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PROCEEDINGS
of the
NINTH PACIFIC SCIENCE CONGRESS
of the
PACIFIC SCIENCE ASSOCIATION
1957
VOLUME 4
BOTANY
Published by the
SECRETARIAT, NINTH PACIFIC SCIENCE CONGRESS
DEPARTMENT OF SCIENCE
BANGKOK, THAILAND
1962
Our 552 7-7-66 10,000
Call No.
Author
OSMANIA UNIVERSITY LIBRARY
flip
Accession No.
This book should be returtfcd on or before the date
last marked below.
Vol
Published by the
SKCRRTARIAT, NINTH PACIFIC SCIENCE CONORLSS
DrPARIMhNT OF SCILNC L
BANGKOK, THAILAND
1962
PUBLICATION COMMIT I'LL
and
EDITORIAL STAI I
Chairman: OK. CHARNC; RAIANA.RAI
\' ice-Chairman- M. u. CIIAKRATONC, IONC.YAI
MR. WiMINCilON BRINK PRO! . NOPAKHl'N lONC.VAI
MR. D V. SASSOON MRS. CAIIMRINI I SIIMN
MR. 1MH R()( IIANAIMJRANANDA OR. TYOIA N \ RANONC
OR PRAPRIT NA NACiARA MR. CIIA1\\AI SANCiRl'JI
MRS. M. ISAF^I (II i ARRAR, Secretary
|'KIN.|ri> A I 1HAI \\A1ANA 1 \NICH ( K l> V >, K\MA I R()A1>, HAN'(.KOK MJ<S HOONPRIN<t F SUW'AN, PRINILR BE 25O5
CONTENTS
1'ldt tor's Note . . .1
Abbreviations . 11
Participants ......... in
Standing Committee Repoits
IOSBI RG. r.R , Report ofthe Chairman of the Standing Committee on Pacific Botany 1
WAI KI R, i II., Concerning the Flora of Japan, Fnghsh Translation 4
nu. SHIN - YiNCj. Status of the Nora of China Project . 5
\\ AI KI R. i H , Report on the Botanical Section ofthe Program for Scientific Investigation ofthe
Ryukyu Klands . 6
si noDiwiim), KDSNOTO, Botany in Indonesia, 1953- 1957 7
MOORI , i r( \ n , Botany in New /ealand, 1953 - 1957 . 10
\\AI MR, i n , Report of the Subcommittee on Bibliography 14
DOIY, MS, Report of the Subcommittee on Algae 17
STI INI R, MONA i ISA, Report ofthe Subcommittee on Common Names of Pacific Plants IS
HANK, i. P, II, Report of the Subcommittee on Fthnobolany . . 2!
Appendix
ROGI RS. DAVID, Summary of Studies on Mtiiuhot r.senlenta 24
i AM, H.J., Report of the Subcommittee on Pacific Plant Areas 2N
SKOI rsiM RG, (".. Report ofthe Subcommittee on Nature Protection . 29
si i YODIWIRYO, KUSNOTO and DH MY, ANWARI, Problems Confronting Botanical Institutions in
the Tropics . ......... . ^9
KOSTI RMANS, A., Problems of Botanical Institutes in the Tropics , . 41
PtJKSiGi ovi , J.w., A Brief History of Botanic Gardens with Special Reference to Singapore . 44
BURKILI , n. M., Some Current Problems ofthe Botanic Gardens, Singapore 46
SUVAFABANDHU, KASiN, Problems Facing the Herbaria in Thailand 52
VAN STI i MS, c.G.r. j , Interaction and Cooperation between 7 ropical and Temperate Herbaria . 54
PROWSI , G.A., Problems in a Spceiali/ed Botanical Department . . 56
VIROI, ROBI RT, The Various Technical Aspects of Botanical Research in New Caledonia 58
WOMI RSII Y, J.s., Notes on the Techniques Employed for the Collection of Botanical Speci-
mens in Papua and New Guinea 61
S\mposium on Vegetation Types of the Pacific
DANSI KI AU, Pii KKi , Vegetation Study and Recording 64
HOSOKAVVA, r., OMURA, M. and NISIUHAKA, Y., Grading and Integration of Epiphyte Commu-
nities ... . ... ... .... 76
KUC'in [R, A.W , Vegetation Mapping m the Pacific Region S3
POKU RIS, R., The Problem of the Origin of the "Savannahs" of the Islands of the Pacilic .... 86
iiiJRi IMANN, J M., The Structure of Some Biocoenoses of New Caledonia . 89
KICKING, RIIOY w , The hog Belt Rain Forest of the Pacific Northwest (U.S.A.) .... ... 95
IAII KAWI, MISAO, 'I he Iconography of the Vegetation of the Natural Forest in Japan ... 99
WANG, rm - vvu. The Development of horest Communities in Eastern Asia . . 103
IOSBIRG, i R , Dynamics of Atoll Vegetation . . . . 114
rosBi RG, i .R , Tropical Pacific Grasslands and Savannas .... 118
KIRA, IAIUO, OGAVVA, iii'SAio and YODA, KYOJI, Some Unsolved Problems m Tropical Forest
Ecology . . . . . 1 24
HORIKAWA, Yosiiiwo. The Occurrence of Tropical Plants in the Japanese Archipelago and its
Phytogeographical Significance . . . .... 135
i Cii i R, [ RANK i , Unifying Concepts in Vegetation Study as Applied to the Pacilic Basin . 1 36
VAN STI i NIS, c G G j , The School-Flora as a Medium for Botanical Education m the Tropics 139
MFIJI R, w.. Botanical Exploration and Education in Present Day Sumatra 141
HOITIUM, R.I ., Problems of Publicity: with Special Reference to the Need for Popular Books
dealing with Focal Plants . . 146
no i Y. MAXWI i i. s., I unctions of the Algae in the Central Pacific 148
JOHNSON, j. HARI AN, Comparison of the Calcareous Algal Eloras of Recent and Fossil Reefs 156
IOSBIRG, I.R., Qualitative Description of the Coral Atoll Ecosystem ... 161
P\LUMBO, RAI PH K, The Relationships between Atolls and Benthic Algae 168
\\OOD, i j mu.t'soN, The Microbiology of Coral Reefs 171
c IIAPMAN, v j , 'I he Marine Biogeographical Provinces of the South Pacific 174
no, PHAM - HOANG, Coastal Marine Plants Around Cauda Harbour (Near Nhatrang) 179
SCAGI i , ROW KI i ., Benthic Algal Productivity in the North Pacific with Particular Reference
to the Coast of British Columbia 181
RYrm:R, JOHN H., On the Efficiency of Primary Production in the Oceans ... 188
sii v\, PAUL c.. Comparison of Algal Floristic Patterns in the Pacific with those in the Atlan-
tic and Indian Oceans, with Special Reference to Codium .... 201
PROWSI , G.A., Distributional Relationships of Malayan Freshwater Algae 217
in DMANN, Ji AN, The Rhodophyta Order Acrochactiales and its Classification ... . ... 219
IOKIDA, JUN and YAIHJ, IIIROSHI, Some Observations on Lammaria Gametophytes and Sporo-
phytcs 222
PAPINIUSS, GLORGI K, Clearing Old Trails in Systematic Phycology . 229
YI i ASQIII /, GRi.GORio i.. On the State of Phycological Knowledge in the Philippines 234
SCAGLI , ROBi RT p., Phylogcnctic Relationships of Certain Dorsivcntral Rhodomelaceae .... 239
Symposium on I'thnohotain of 1 lunland and Contiguous Countries
BANK, IHIODORI P . II, Opening Remarks on Hthnohotany and Ecology . 244
JONIS, \OIMY ii.. The Nature and Status of Fthnobotany . . ... . 246
SMITINAND, ii M, Materials Used for Thatching m Thailand . ... 248
SAMAPUDDIII, k., Some hood Plants in the ForesN of 'I hailand . . . 250
SMIIINAND, TI M, Note Made from Local Knowledge of the Use of Poisonous Plants by the Thai
People . . . . 262
PHDMXUSRI, pRASimu, Tea in Thailand . . . 264
IUGBY, nisi. The Manufacture of Sugar from the Sugar Palm in Upper Mandailing,
Sumatra . . .... 266
BART i ITT, IIAKI i Y HARRIS, Possible Separate Origin and Evolution of the Ladang and Sawah
Types of Tropical Agriculture 270
BARii.Lir, HARI i/v HARRIS, Some Words Used in Connection with Primitive Agriculture in
Southeast Asia . ... 274
WIATHFRWAX, PAUi , Ethno-Ecological Relationships between the Mai/e Plant and Man in
Western Ancient America 276
BANK, IHLODORF P., II, Elhiiobotany of Northern Peoples and the Problem of Cultural Drift . . 279
BANK, THLODORE p., II, Medicinal Plant Lore of the Aleut 281
STHINL R, MONA i ISA, The Problem of Vernacular Names of Plants in the Pacific and its Solution 285
HU, smu - YING, An Enumeration of the Food Plants of China with Vernacular Names 289
TUYAMA, TAKASI, The Problem on the Vernacular Plant Names in Japanese 291
PURSLGLOVF, J.w., Common Vernacular Names of Plants of the Pacific Basin 293
BARRAU, JACQUES, Notes on the Significance of Some Vernacular Names of Food Plants in
the South Pacific Islands 296
CONKLIN, HAROLD c., Ethnobotanical Problems in the Comparative Study of Folk Taxonomy. . 299
AUSTERLITZ, ROBERT, A Linguistic Approach to the Ethnobotany of South-Sahalin 302
DWYHR, R.E.P., Vernacular or Common Names of Plants 304
JOHN, HAROLD ST., Origin of the Sustenance Plants of Polynesia, and Linguistic Evidence for
the Migration Route of the Polynesians into the Pacific (Abstract) 308
BARRAU, JACQUES, Edible Yams of the South Sea Islands, Species Present, Vernacular Names
and Distribution 309
Symposium on Vernacular Names of Plants
COLE, KATiiihLN, Acelo-Carmine Stain in the Cytogemitical Investigation of Some Marine
Algae of the Pacific Coast 313
ELLIOTT, JACK c., The 1956 Typhoon Season on Okinawa (Abstract) 316
TIENG, TRAN-NGOC, The Evolution of the Germinative Power of Rice Seeds 318
DE JONGI , P., Stimulation of Yield in Hevea Brasiliensis 320
MENNINGER, EDWIN A., Flowering Trees that Fail to Flower in Alien Lands 325
EDITOR'S NOTE
At the meeting of the Committee on Publications of the Ninth Pacific Science Congress, it was
agreed that the increasing number of communications and papers presented at each Congress has
become a very difficult problem for the Publication Committee and the editorial staff to cope with and
that too much time is required to complete the publication of the Proceedings; therefore, it was recom-
mended that the following principles governing publication be followed:
a. That invited contributions to the scheduled symposium be published in full;
b. That reports of the Standing Committees be published in full;
c. That other papers submitted to the Congress during its sessions be published in abstract only,
the abstract not to exceed 500 words;
d. That papers which, though listed on the program or included in the pre-Congress abstracts
published in advance but not actually submitted to the Congress at its sessions, should be
disregarded ;
e. That authors be asked to indicate by a definite and early date 1 whether they prefer to publish
their papers in sources other than the Congress Proceedings; that if this is done, the Congress
should be acknowledged;
f. That all proof reading be the responsibility of the editorial committee, and that this committee
shall consider the manuscripts in their hands by a definite date as final; 2
g. That authors be held responsible for submitting their material in good English;
h. That on matters arising during the course of publication and not specifically covered in the
statement of policy the editorial committee is empowered to act.
In accordance with the resolutions of the Committee, the editorial board has edited the reports and
manuscripts where necessary to bring uniformity and consistency to the format. Typographical and
grammatical errors as well as errors in phraseology, spelling, or technical terms have been corrected,
wherever possible, but in cases where the exact meaning of the original copy was not clear, the text has
been left as submitted by the autor.
In order to reduce the cost and bulk of the publication, appendices, illustrations, and exhibits when-
ever considered not vital to the text have been eliminated.
Tf an author requested to publish elsewhere, his paper has been mentioned in the footnote under
the respective titles, but if an author who presented a paper at the Congress failed to submit his manu-
script either iafull or in abstract, his paper and the discussions thereon have been eliminated entirely.
It was also decided that, in order to complete the publication of the Proceedings as soon as possible,
each division be published in a separate volume. Short volumes or the ones that do not require too much
editorial work will be released first. Therefore, among the twenty volumes planned, any volume may
appear first. They will not appear in consecutive order.
The editorial board wishes to thank all authors who were prompt in submitting their revised manu-
scripts in good form and, in particular, members of the Standing and Organizing Committees, too
numerous to be named, who have helped in collecting the manuscripts pertaining to their respective
divisions.
The Board wishes in particular to thank Dr. F. Raymond Fosberg for going over and correcting
the Special Symposium on Climate, Vegetation, and Rational Land Utilization in the Humid Tropics
under UNESCO;
Mr. Saman Buravas of the Royal Mines Department for helping by redrawing charts and maps
in order that they might reproduce clearly when printed;
Mr. J. Alan Tubb of the FAO Regional Office, for his assistance in going over and clarifying some
of the papers in the Fisheries and Oceanography volumes and in translating some of the French papers;
Dr. Pradisth Cheosakul of the Department of Science for editing the Chemistry in the Development
of Natural Resources volume;
Last but not least, the Board wishes to thank the Thai Watana Panich Press for their cooperative ef-
forts, far beyond the requirement of the contract, in devoting all their resources to printing these volumes.
1 January 1, 1958, in the case of the Ninth Congress.
2 March 1, 1958, in the case of the Ninth Congress.
ABBREVIATIONS
APFC
CAA
CSIRO
ECAFE
EQUAPAC
FAO
IACOMS
ICA
ICAO
ICSU
IGY
IPFC
IRC
JCRR
NORPAC
PIHI.CUSA
PIOSA
SFATO
SPC
UN
UNESCO
UNICFF
USDA
USIS
USOM
WHO
WMO
Asia-Pacific Forestry Commission
Civil Air Administration
Commonwealth Scientific and Industrial Research Organization (Australia)
Economic Commission for Asia and the Far East
Equatorial Pacific (occanographic survey)
Food and Agriculture Organization
International Advisory Committee on Marine Sciences
International Cooperation Administration
International Civil Aviation Organization
International Council of Scientific Unions
International Geophysical Year
Indo-Pacific Fishery Commission
International Rice Commission (FAO)
Joint Commission on Rural Reconstruction (Taiwan, China)
North Pacific (occanographic survey)
Philippine Council for United States Aid
Pan-Indian Ocean Scientific Association
South-East Asia Treaty Organization
South Pacific Commission
United Nations
United Nations Educational, Scientific and Cultural Organization
United Nations International Children's Emergency Fund
United States Department of Agriculture
United States Information Service
United States of America Operations Mission
World Health Organization
World Meteorology Organization
PARTICIPANTS*
ASHTON, PETER SHAW, Botanist, State Forest Office, Brunei.
AUBERT DE LA RUE, EDGAR, Geologist, National Centre for Scientific Research, 18 rue Ribera, Paris
XVI", France.
BALASIRI, DAROON, Instructor, School of Pharmacy, University of Medical Science, Bangkok,
Thailand.
BANK, THEODORE FOUL, ii, Executive Director, Institute for Regional Exploration, University Station,
Box 2143, Ann Arbor, Michigan, U.S.A.
BARTLETT, STEPHEN, Head of F&M Department, National Institute for Research in Dairying, Univer-
sity of Reading, Reading, England.
BELARDO, LUZ ouvEROS, Dean, College of Pharmacy, The Philippine Women's University, Manila,
Philippines.
BOONJAVATANA, CHAROEN, Supervisory Unit, Department of Teacher Training, Ministry of Educa-
tion, Bangkok, Thailand.
BRYNE, BRIAN, Teacher, Mt. Hermon, Darjeeling, India.
BURKILL, HUMPHREY MORRISON, Director, Botanic Garden, Singapore.
CALDWELL, ARTHUR FRANCIS, Department of Pharmacy, Faculty of Medicine, University of Malaya,
Singapore.
CAPELI , ARTHUR, Reader in Oceanic Linguistics, University of Sydney, Sydney, N.S.W., Australia.
CHANTARASRIKUL, ANONG, Bangkhen Experimental Station, Department of Agriculture, Ministry of
Agriculture, Bangkok, Thailand.
CHUMDFRMBHADEJSUK, SHALAM, Srisuriyothai School, Ministry of Education, Bangkok, Thailand.
CROCKER, ROBERT LANGDON, Professor of Botany, University of Sydney, Sydney, N.S.W., Australia.
DASANANDA, SALA, Deputy Director-General, Rice Department, Ministry of Agriculture, Bangkok,
Thailand.
DOTY, MAXWELL s., Professor of Botany, University of Hawaii, Honolulu 14, Hawaii.
DOUGLAS, BRYCE, Senior Scientist, Smith Kline and French Laboratories, Philadelphia, c/o Chemistry
Department, University of Malaya, Singapore 10.
EGLKR, FRANK EDWIN, Aton Forest, Norfork, Connecticut, U.S.A.
FELDMANN, JEAN, Professeur & la Faculte de Sciences de Paris, Laboratoire de Biologic Vegetale
Marine, Faculte dc Sciences de Paris, 195 rue St. Jacques, Paris 5 e , France.
FOSBERG, F. RAYMOND, Botanist, U.S. Geological Research Council, Washington 25, D.C., U.S.A.
GAUSSEN, HENRI MARCEL, Professor, University of Toulouse, rue Raymond JV 21, Toulouse, France.
HARRISSON, TOM H., Curator, Sarawak Museum, Brunei.
HEIM, ROGFR, Director of the National Museum of Natural Sciences, Museum National D'Histoire
Naturelle, 57 rue Cuvier, Paris 5 e , France.
HO, PHAM HOANG, Charg de covas a la Faculte, Universite de Saigon, 12 Hong Thap Tei, Saigon,
Vietnam.
HOERNER, GODFREY RICHARD, Chief University Advisor, Kasetsart University/Oregon State College
ICA Contract, Bangkhen, Bangkok, Thailand.
HORMCHONG, TWFE, Instructor, College of Education, Ministry of Education, Bangsaen, Thailand.
HU, SHIU-YING, Botanist, Arnold Arboretum, Harvard University, 22 Divinity Ave., Cambridge,
Mass., U.S.A.
HURLIMANN, JOHANN H., Botanist, Ciba J.A., Friendensgasse 51, Basle, Switzerland.
JALAVICHARANA, KAHN, Chief, Plant Industry Division, Department of Agriculture, Bangkhen,
Bangkok, Thailand.
JIRAWONGSE, VICHIARA A., Lecturer, Department of Pharmacy, Chulalongkorn University, Bangkok,
Thailand.
* Initials or names in italics represent Thai titles.
iii
JOHNSON, J. HARLAN, Professor of Geology, Colorado School of Mines, Golden, Colorado, U.S.A.
KASHEMSANTA, M.C. LAK, Director-General, Department of Agriculture, Ministry of Agriculture,
Bangkok, Thailand.
KETUSINH, OUAY, Professor, Department of Physiology, Siriraj Hospital, Dhonburi, Thailand.
KUCHLER, A.W., Professor of Geography, University of Kansas, Lawrence, Kansas, U.S.A.
LOETSCH, FRITZ, Professor, Hamburg University, Bundes forschungsanstalt Forstwirtschaft, Reinbek
bei, Hamburg, Germany.
MCHALE, THOMAS RiLEY, Economist, 24 Blackstone Blud., Providence, R.I., U.S.A.
MEKAYONK, VICHIT, Suan Sunantha College, Bangkok, Thailand.
NA-THALANG, SOMBOON, Director, Rubber Division, Department of Agriculture, Ministry of Agri-
culture, Bangkok, Thailand.
OGAWA, HUSATO, Researcher, Laboratory of Plant Ecology, Institute of Polytechnics, Osaka City
University, Manami-Ogimachi, Kitaku, Osaka, Japan.
PAPENFUSS, GEORGE F., Professor, Department of Botany, University of California, Berkeley 4, Cali-
fornia, U.S.A.
PAVANARIT, SRIROT, College of Education, Department of Teacher Training, Ministry of Education,
Bangkok, Thailand.
PHUMXUSRI, PRASIDHI, Agriculturist, Mae Malai Farm, Mae Rim, Chiang Mai, Thailand.
PICHAISIRI, SOMSUGDI, Instructor, Department of Teacher Training, Ministry of Education, Bangkok,
Thailand.
PROWSE, GERALD ALBERT, Botanist and Algologist, Fish Culture Research Station, Batu Berendam,
Malacca, Federation of Malaya.
RAYMOND, MARCEL L., Taxonomist, Montreal Botanical Garden, 4101 East, Sherbrooke Street, Mon-
treal, Canada.
RICHARDS, PAUL WESTMACOTT, Professor of Botany, University College of North Wales, Bangor,
Great Britain.
RUHLE, GEORGE CORNELIUS, Park Naturalist, Hawaii National Park, Hawaii, U.S.A.
RYTHER, JOHN HOOD, Marine Biologist, Woods Hole Oceanographic Institution, Woods Hole, Mass.,
U.S.A.
SACHET, MARIE-HELENE, Bibliographer, Pacific National Research Council, Washington 25, D.C.,
U.S.A.
SAMAPUDDIH, KRiT, Royal Forest Department, Bangkok, Thailand.
SANSUKH, SOMSAK, Teacher, College of Education, Department of Teacher Training, Ministry of
Education, Bangkok, Thailand.
SANTOS, JOSE VERA, Head, Department of Botany, University of the Philippines, Quezon City,
Philippines.
SCAGEL, ROBERT FRANCIS, Associate Professor, Department of Biology and Botany, and Institute of
Oceanography, University of British Columbia, Vancouver 8, B.C., Canada.
SEIDENFADEN, GUNNAR, Danish Minister, Danish Legation, Bangkok, Thailand.
SETYODIWIRYO, KUSNOTO, Director, Botanic Gardens of Indonesia, Djl. Bubulak 25, Bogor, Indonesia.
SILVA, PAUL CLAUDE, Associate Professor, Department of Botany, University of Illinois, Urbana,
Illinois, U.S.A.
SMITINAND, TEM, Chief, Section of Botany and Zoology, Forest Products Research Division, Royal
Forest Department, Bangkok, Thailand.
STEINER, MONA LISA, Chairman of sub-committee on Vernacular Names of Plants, 2833 Park Avenue,
Pasay City, Philippines.
SUVAGONDHA, CHAMLONG, Professor and Dean, School of Pharmacy, Bangkok, Thailand.
SUVARNASUDDHI, KHio, Deputy Director-General, Royal Forest Department, Bangkok, Thailand.
SUVATABANDHU, KASIN, Department of Agriculture, Bangkok, Thailand.
THIRAWAT, SUKHUM, Chief, Division of Forest Control, Royal Forest Department, Bangkok,
Thailand.
iv
THONGUMPHAI, KHAJORN, Teacher, Chachoengsao Teacher Training College, Chachoengsao,
Thailand.
TRINANDWAN, SA-KORN, Teacher, College of Education, Ministry of Education, Bangkok, Thailand.
TROLL, CARL TH., Professor, University of Bonn, Universitat, Geographisches Institut, Bonn, Germany.
TUYAMA, TAKASI, Professor, Laboratory of Botany, Faculty of Science, Ochanomizu University,
Otsuka, Bunkyoku, Tokyo, Japan.
VALLARTA, MARIA T., Head, Department of Botany, The Philippine Women's University, Manila,
Philippines.
VAN STEENIS, c.G.G.J., Professor, Flora Malesiana Foundation, Marelaan 55, Oegstgeest, Netherlands.
VESTAL, EDGAR F., T.A. Plant Protection Advisor, USOM, Bangkok, Thailand.
WALKER, EGBERT H., Department of Botany, Smithsonian Institution, Washington 25, D.C., U.S.A.
WEATHERWAX, PAUL, Visiting Professor, College of Education/Indiana University ICA Contract,
Bangkok, Thailand.
WYCHERLEY, PAUL RENODEN, Botanist, Rubber Research Institute of Malaya, P.O. Box 150, Kuala
Lumpur, Malaya.
YEN, DOUGLAS ERNEST, Scientific Officer, Vegetable Station, D.S.I. R., Robinson Road, Otahuhu,
New Zealand.
YODA, KYOJI, Researcher, Laboratory of Plant Ecology, Institute of Polytechnics, Osaka City Uni-
versity, Minami-Ogimachi, Kitaku, Osaka, Japan.
BOTANY
Standing Committee Chairman: F. R. FOSBERG
Organizing Committee Chairman: M.C LAKSHANAKARA KASHEMSANTA
Standing Committee Reports
REPORT OF THE CHAIRMAN OF THE STANDING COMMITTEE ON
PACIFIC BOTANY
F. R. FOSBERG
Chairman, Pacific Vegetation Project, Washington, D.C., U.S.A.
The Standing Committee on Pacific Botany,
established under this title by the Eighth Pacific
Science Congress, succeeding the former Standing
Committee on Pacific Plant Areas, includes the
following members (arranged alphabetically):
Theodore P. Bank II (U.S.A.) (Chairman,
Subcommittee on Pacific Ethnobotany).
Nancy T. Burbidge (Australia).
Pierre Dansereau (Canada) (Chairman, Sub-
committee on Pacific Plant Ecology).
Maxwell S. Doty (Hawaii) (Chairman, Sub-
committee on Pacific Algae).
Ramon Ferreyra (Peru).
F. Raymond Fosberg (U.S.A.) (Chairman,
Standing Committee).
Hiroshi Kara (Japan).
Ir. Kusnoto Setyodiwiryo (Indonesia).
H. J. Lam (Netherlands) (Chairman, Subcom-
mittee on Pacific Plant Areas).
E. Looho (Indonesia) (Chairman, Subcommit-
tee on Medicinal Plants).
Lucy B. Moore (New Zealand) (replacing the
late W.R.B. Oliver).
Eduardo Quisumbing (Philippines).
Carl Skottsberg (Sweden) (Chairman, Subcom-
mittee on Nature Protection).
Mona Lisa Steiner (Philippines) (Chairman of
Subcommittee on Common Names of Pacific
Plants).
Kasin Suvatabandhu (Thailand) (Secretary,
Standing Committee).
Egbert H. Walker (U.S.A.) (Chairman, Subcom-
mittee on Botanical Bibliography).
John Wyatt-Smith (Malaya).
The committee has functioned principally
through its eight subcommittees. The reports of
these subcommittees (which follow) speak for
themselves, and material contained therein will
not be repeated here. Their accomplishments
are amply sufficient to justify continuation of this
method of functioning of the Standing Com-
mittee. It is therefore recommended that the
Botany Division of the Ninth Congress consider
very carefully the lines of effort that it wants the
next Standing Committee on Pacific Botany to
emphasize and that it recommends for each such
line or field the establishment of a special sub-
committee, as was done in 1953.
The present Standing Committee is composed
of the chairmen of the subcommittees and suffi-
cient other members to give it wide geographical
representation. No members were selected from
countries that were represented already by sub-
committee chairmen. When Professor Kusnoto
had to resign the chairmanship of his Subcom-
mittee on Medicinal Plants because of pressure
of other duties, on his recommendation Dr.
Looho was appointed to replace him, but he
continued as a Committee member. We deeply
regret the loss by death of one of our members,
Dr. Oliver, chairman of the predecessor of this
committee. Miss Moore was asked to replace
him.
The principal efforts of the Chairman were
to assist the Organizing Committee of the Ninth
Congress with the planning of the botanical
program and to compile a list of Pacific botanists.
To accomplish the first aim, a visit was made
to Bangkok in April, 1956, when the program was
thoroughly discussed and names were suggested
of people who might agree to organize symposia
and to be chairmen of sessions. Most of these
people have responded to the invitations of the
Organizing Committee, resulting in the rather
1
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
ambitious program of the Botany Division,
and, incidentally, in the excellent representation
of Pacific botanists at the Congress. Where
possible, the activities of the subcommittees have
been tied into the program. If certain fields have
been neglected, it was because the program could
scarcely have been made longer. Cooperation
with other divisions was recommended where it
seemed appropriate.
The second of the tasks was more difficult,
and the product is admittedly imperfect. A
request was sent to all Standing Committee
members and a few others for lists of names from
their several geographic areas. This resulted
in a few lists. To these were added names known
to the Chairman, encountered in the literature,
or found in published address lists. The resulting
list was duplicated and circulated to a geograph-
ically representative group of botanists for
criticism, additions, and corrections. The result
was gratifying and enabled us to get together the
list that is before you and which will be mailed
to all those appearing in it. It is hoped that it
will contribute substantially to the primary aim
of the Standing Committee, to stimulate com-
munication and cooperation between the botanists
interested in the Pacific Basin.
A word is in order about the scope and limita-
tions of this list. It was arbitrarily limited to
those branches of botany that have a definite
regional aspect, and that are not properly included
in agriculture or other strictly practical fields
which may have the resources to compile lists
that will better serve their purposes. Thus many
names were excluded in such fields as plant phy-
siology, genetics, plant chemistry, cytology,
agronomy, etc. Reliance usually had to be
placed on printed sources for such information.
Undoubtedly serious errors were made. We
humbly ask forgiveness and hope that a future list
may rectify all errors and omissions.
We take great pleasure in acknowledging our
gratitude to the Pacific Science Council Secreta-
riat and the National Science Foundation for
financial help that made the production of this
list possible. Copies will be distributed to all
people whose names appear on the list and to
libraries of institutions known to be interested
in Pacific Botany. Persons or institutions not
listed or which do not receive copies may receive
them by requesting them, so long as the copies
last.
We wish also to express our gratitude to the
Pacific Science Board of the National Research
Council for continuously making facilities and
2
space available for the functioning of the Stand-
ing Committee, and for help in numerous ways.
It is customary for the reports of standing
committees to include summaries of work in the
field accomplished or undertaken during the
period of the Committee's activity. We have
asked Committee members for such information
on their areas. A few have responded, and their
contributions are briefly summarized here, along
with some other widely known current activity.
This is only a sampling of the field, and details
and most names are purposely omitted, as it is
impossible to list all of them. Several of the
reports submitted by Committee members seem
sufficiently comprehensive that it is desirable to
append them in full to this report, in which cases
little in the way of summary is included.
By far the most comprehensive and widespread
program in the Pacific Area is that of the Flora
Malesiana, whose able editor is with us and may
care to give us current news of it. Under its
auspices have appeared many preliminary re-
visions of groups as well as a number of the
final treatments for the Flora. The Flora male-
siana Bulletin continues to be the world's most
informative botanical news organ, as well as
containing much valuable bibliographic informa-
tion.
Revision of Cheeseman's Manual of the Flora
of New Zealand is complete, and the book is in
press. For further information on New Zealand,
see Miss Moore's report.
The C.S.I.R.O. in Australia is carrying out
extensive botanical and ecological survey work in
the Northern Territory and in New Guinea,
and botanists in other parts of Australia continue
to be very active. It should be specially noted
that the botanical exploration of the subantarctic
Macquarie Island, recommended by the Second
Pan-Pacific Science Congress, has been carried
out by a group from the C.S.I.R.O.
Along the Pacific side of South America, work
on the flora and vegetation of Peru, centered at
the Universidad de San Marcos, Lima, and that
on Colombia, centered at Bogota, Medellin,
and especially at the U.S. National Herbarium,
is especially active and productive. The Flora of
Panama is being continued by the Missouri
Botanical Garden.
In the U.S., work on a manual of California
plants is going on at the Rancho Santa Ana
Botanical Garden and New York Botanical
Garden. At the several campuses of the Univer-
sity of California, Jepson's Flora of California
is drawing toward completion, and much work in
VOLUME 4
BOTANY
systematic, biosystematic, and ecological fields
is being done. Pomona College and affiliated
institutions also form an active center of taxo-
nomic and ecological work, especially on plants
of arid southwest North America. Work on the
vegetation of Alaska has been carried out under
various auspices, especially in connection with
work on permafrost by several military sponsored
agencies and the U.S. Geological Survey.
In Japan there has been much activity in many
institutions, principally on the vegetation and
flora of Japan. Especially valuable as a basis for
all other such work are Hara's bibliographic
checklist of the flora of Japan and an English
language revision of Ohwi's Flora of Japan, the
latter in cooperation with several U.S. Institu-
tions.
We have no news of the status of recent
botanical work in mainland China, but the Flora
of China Project of the Arnold Arboretum,
U.S.A., is being continued under the direction of
Dr. Shin-ying Hu. (See accompanying report by
Dr. Hu.)
For a report of progress on a flora of the
Ryukyu Islands see the accompanying report by
E.H. Walker.
In Thailand aquatic and economic plants are
under study by our Thai colleagues, and a Danish
expedition has been studying and collecting the
flora.
At the University of Hawaii is a project on
Pacific Pandanus, as well as active work on algal
ecology and ocean productivity.
The Pacific Vegetation Project has continued
its investigations on the vegetation and flora of
Micronesia, as well as some other Pacific areas
and on coral atolls generally, sponsored by the
U.S. Geological Survey. A by-product of this
is the volume, Island Bibliographies, published
by the Pacific Science Board.
On other insular areas, work on the flora of
New Caledonia has continued at Zurich and by
R. Virot at Paris, on that of Fiji at the U.S.
National Herbarium and at Suva, and on the
food plants of the Pacific at the University of
Marseilles and the South Pacific Commission.
Finally, we must mention the completion and
publication of two magnificent volumes on the
vegetation and the origin and affinities of the flora
of Juan Fernandez by our eminent colleague, Carl
Skottsberg of Gothenburg, Sweden.
Mention should be made of the status of the
volume on the Vegetation Provinces of the
Pacific Basin, based on the symposium at the
Eighth Pacific Science Congress. Manuscripts
are in hand for the greater part of the chapters,
but about a third are still not available. They
are mostly reported in various stages of comple-
tion. It is hoped that this work can be promptly
brought to a conclusion, as all indications are
that the volume is much needed and would imme-
diately be widely used. Experience with this
volume tends to discourage attempts at coopera-
tive works of this sort, as it seems to be necessary
to spend time out of all proportion merely to
urge unenthusiastic collaborators to complete
their promised contributions.
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
CONCERNING THE FLORA OF JAPAN, ENGLISH TRANSLATION
E. H. WALKER
Smithsonian Institution, Washington, D.C., U.S.A.
An American serviceman, Leopold Charette,
initiated the translation into English by J. Ohwi
of his Flora of Japan (in Japanese, 1953). Fred
Meyer, formerly at the Missouri Botanical
Garden, and E.H. Walker, Smithsonian Institu-
tion, are editors, and the project is currently
sponsored by the Missouri Botanical Garden.
Funds have been provided by the National
Science Foundation, U.S.A. Dr. Ohwi recently
submitted a treatment of the Pteriodophyta, a
group not included in the original flora, in
English. With the permission of the editors, he
then translated this into Japanese for publica-
tion in Japan. This came from the press in
November, 1957. (A copy was circulated through
the audience.)
VOLUME 4
STATUS OF THE FLO RA~OF~CH1NA7 PROJECT
BOTANY
SHIN-YING HU
Arnold Arboretum, Jamaica Plain, Massachusetts, U.S.A.
HISTORY
This project was initiated in 1953 as a coopera-
tive project between A Continental Development
Foundation and the Arnold Arboretum of
Harvard University. Over thirty thousand
dollars has been invested in it.
ACCOMPLISHMENT
1. A complete index to the species of the
flowering plants has been prepared. This is in
card catalogue form and it consists of approxi-
mately 100 thousand cards each covering names,
synonyms, important literature references, au-
thentic collections, and the distribution.
2. The preparation of a bibliographical enu-
meration of the plants of China has commenced.
The treatment of the Compositae is now complete.
The manuscript consists of about 1,500 typed
sheets, double spaced. That of the Orchidaceae
is almost done.
PUBLISHED FLORA AND MANUSCRIPT
1. A model of the flora, a treatment of the
Malvaceae, was published in 1955 and distributed
in 1956. It shows what can be done.
2. Manuscript for the next issue covering
Palmae, Araceae, Ericauloneceae, Conmelina-
ceae, and Panderderaceae is at hand.
PROBLEMS
The supporting foundation can no longer
provide financial support to the Project. The
undertaking itself is a tremendous task. Now
moral and financial support are required for its
continuation.
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
REPORT ON THE BOTANICAL SECTION OF THE PROGRAM FOR
SCIENTIFIC INVESTIGATION OF THE RYUKYU ISLANDS
E. H. WALKER
Smithsonian Institution, Washington, D.C., U.S.A.
The SIRI program of the Pacific Science Board
and the U.S. Army was designed to provide
basic scientific information for the development
of the southern Ryukyu Islands. As botanist on
this program, I spent the three summer months
of 1951 in Okinawa and the Southern Ryukyus
mainly collecting herbarium material for use in
eventually preparing a Flora of the area.
In 1952, the U.S. Civil Administration and the
Government of the Ryukyus mimeographed an
issue of 200 copies of a Flora of Okinawa,
originally prepared by the Okinawans, Sonohara,
Tawada, and Amano, and edited by me. In 1954,
they issued my book of some 300 pages Important
Trees of the Ryukyu Islands, dealing with 209
species, each illustrated, the text in both Japanese
and English in parallel columns. In 1953, I
attended the Eighth Pacific Science Congress as
delegate from this program and contributed
two papers dealing with Okinawa and the
Southern Ryukyus, and on my return made
further collections in Okinawa. With assistance
from this program, a herbarium was established
at the University of the Ryukyus in 1955 and
encouragement given to the training in America
of an Okinawan botanist to become its curator.
In 1954, a new Flora of Okinawa and the
Southern Ryukyus was started. It is based upon
all available specimens, which have now grown
to occupy some ten cases in the U.S. National
Herbarium in the Smithsonian Institution, and
on critical studies by many contributors. As
now being prepared it will contain keys, brief
characterizations, and critical notes. The thor-
oughness of this work was greatly increased by
the opportunity to select and borrow critical
herbarium specimens from Japanese and Taiwan
herbaria and to make collections in Okinawa
of palms and bamboos while enroute to this
Congress.
Besides the material results of this program,
so useful in the development of the Ryukyus and
of our scientific knowledge of this area, such as
the publications and critically studied scientific
specimens, certain intangibles may be noted.
Especially significant is an increased under-
standing, goodwill, and cooperation between the
botanists and institutions of Japan, Okinawa,
Taiwan, America and some European countries.
Surely the investment in this program is bringing
rewarding returns.
VOLUME 4
BOTANY INTNDONESIA, 1953-T957
KUSNOTO SETYODIWIRYO
Director, Botanic Gardens of Indonesia, Bogor* Indonesia.
BOTANY
This is a brief report of botanical activities in
Indonesia during 1953-1957.
The staff of the Treub Laboratory was engaged
in conducting the following scientific investiga-
tions.
INVESTIGATIONS IN THE FIELD OF GENERAL BOTANY
(a) Research with growth hormones, isolation
of growth promoting compounds of a fern,
application of a new Ageratum test, applica-
tion of commercial growth hormones for the
rooting of cuttings of cultivated plants.
(b) Investigations on the interaction between
epiphytes, micro-organism, and host-plants
in order to clarify a supposed symbiotic ef-
fect. Microscopy of the lutoids in Hevea
latex.
(c) Cytological investigations of the Amorpho-
phallus and Corypha. Description of the
meiotic divisions, the explanation of a new
phenomenon found in the meiotic prophase
in the genus Corypha, in respect of the
function of the nucleolus. Study on the
pollen grain mitosis of Raffles ia arnoldi,
and the establishment of the number of
chromosomes. Study on the sterility of
Monodora myristica. Allium spp. were
used to test the effect of the digitonin on
the nuclear division.
(d) Genetical investigations with Digitalis spp.
carrying out interspecific crossing between
D. lanata and D. lutea. Observation of
different Digitalis spp. cultivated on higher
sea-level.
(e) Anatomical investigations on the leaf and
root structure of Fagraea borneensis and
Smilax sp.
INVESTIGATIONS IN THE FIELD OF BIOCHEMISTRY
(a) Investigations of the chemical composition
of latex in the family of Apocynaceae.
(b) Determination of the rubber hydrocarbon
constituent of Balanophora sp.
(c) Investigations on the cardiac glycosides of
Theyetia peruviana, Antiaris toxicaria and
various other plants of the families of Ascle-
piadaceae, Apocynaceae, Moraceae. Test
for the presence of alkaloids and saponins
in plants collected in Indonesia.
INVESTIGATIONS IN THE FIELD OF MICROBIOLOGY
(a) Isolation of pure cultures of Pseudomonas
sp. collected from diseased Hevea and
Digitalis plants.
(b) Investigation of Xanthomonas sp. isolated
from infected rice plants.
(c) Obtaining of new material for the collection
of fungi and bacteria maintained in the
laboratory for scientific and industrial pur-
poses. Exchange of samples with institutions
abroad.
Lectures. Delivering a course on the cultivation
methods of orchids in 1956. Lectures on Cyto-
logy and Genetics at the Faculty of Agriculture
of the University of Indonesia. The same lec-
tures, and also on Biochemistry, were given at the
Biological Academy, established by the Ministry
of Agriculture in October, 1955, in Bogor.
Excursions. An excursion was carried out to the
islands of Timor, Roti, etc. in order to collect
medicinal plants; another excursion in Java,
Celebes and in the Lesser Sunda Islands to collect
cytological material of monocarpic palms.
Guest research workers in the Treub Laboratory.
Prof. Dr. Koernicke, Germany, Prof. Dr. F.
Fagerlind and G. Wibom, Sweden, Prof. Dr. P.
Buchner, Italy, and Dr. P. Surany, U.S.A., have
visited and worked for varying periods. Dr.
H.P. Bottelier, Holland, worked in the Treub
Laboratory on a Fellowship for six months.
Exhibition. The Treub Laboratory has partici-
pated in the Agricultural and Horticultural
Exhibition at Pasar Minggu near Djakarta in
1954, with a show material on the effect of the
growth hormones and the symbiosis between
epiphytes, microorganism, and host plants.
HERBARIUM BOGORIENSE
In 1953, a combined expedition of Dr. A.
Kostermans of the Forestry Service and Dr. W.
Meyer of the Herbarium went to the island of
Nunukan in North East Kalimantan (Borneo)
7
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
and the adjacent mainland. The collections
amounted to 1,500 numbers of Phanerogams and
600 Cryptogams. Two other members of the
staff, Mr. Groenhart and Mr. Van Borssum
Waalkes, made a trip to West-Central Sumatra
and the Mentawei Islands. The Herbarium
received 4,630 plants in exchange, whereas more
than 14,000 specimens were sent abroad as
duplicates and more than 8,000 sheets were
loaned for study purpose. One part of Volume 2
of Reinwardtia appeared with monographies and
papers on Phanerogams.
In 1954, a guest worker, Dr. Alston from the
British Museum in London, made collecting trips,
sponsored by the Herbarium, to South Borneo,
Sumatra, North Celebes, Ternate, and Ambon.
A trip by Mr. A. Groenhart was made to Banten
in West Java. The collections of the expedition
of Dr. Kostcrmans of the Forestry Service to
Central Kalimantan, amounting to 1,500 field
numbers, were ceded to the Herbarium. More
than 3,000 duplicates were received from abroad,
in exchange for almost 12,000 duplicates, sent
abroad to twenty Institutes. One big volume
appeared of Reinwardtia containing numerous
articles on Phanerogams and Cryptogams.
In 1955, an expedition of Dr. A. Kostermans
went to Central Kalimantan for three and one-
half months. The collections, consisting of about
one thousand field numbers, are incorporated
in the Herbarium collections. Furthermore, an
expedition of Dr. Kostermans to Central Suma-
tra for the purpose of collecting timber for re-
search in cellulose for rayon, brought back more
than 600 field numbers for the Herbarium. Dr.
Kostermans was accompanied during the first
part of this trip by Mr. Anwari Dilmy, Keeper
of the Herbarium. The two gentlemen accom-
panied Prof. Kusnoto, the Director of the Botanic
Gardens to Padang Tinggi, West Coast of
Sumatra, for the official opening ceremonies of
the Branch Garden there. Plans were outlined
as to how to investigate the extensive area of
virgin forest surrounding this new Garden.
Almost 20,000 specimens have been sent abroad,
on loan and as gifts, to thirty Institutes, whereas
3,000 specimens apart from the Kostermans
collections were received in exchange. One
back-volume, containing the plates of Ebenaceae
of Bakhuizen's monography appeared, and one
part of Reinwardtia, containing numerous taxo-
nomical articles. The Herbarium building un-
derwent big repairs and improvements and is
now in a good condition.
In 1956, Mr. Van Borssum Waalkes made an
expedition to Timor. Mr. Jacobs made a
8
combined trip with Dr. Meyer from the Agricul-
tural Faculty at Pajakumbuh to Mount Korintji
in Central Sumatra, whereas one guest-worker,
Mr. Forman from Kew, made trips to East Java
with Mr. Dilmy, and to Sumatra. Dr. Koster-
mans, who made an expedition to the mountain
region in Central Kalimantan, brought back
about 800 field numbers and living plants of these
unexplored regions. The collections are incor-
porated in those of Herbarium Bogoriense. Two
parts of Reinwardtia appeared this year with
articles on Cryptogams and Phanerogams. Mr.
Dilmy and Dr. Kostermans attended the UNESCO
Symposium in Ceylon.
In 1957, Mr. Forman made a trip to North
Celebes. Dr. Kostermans collected in East Java,
especially bamboos. Mr. Dilmy made a trip to
the U.S.A. and Europe, sponsored by the Ford
Foundation and UNESCO. Two volumes of
Reinwardtia appeared. In 1956 more than
10,000 specimens were sent out in exchange,
and about 1,000 specimens were received. In
1957, exchange and loan of material was carried
on as usual and is still increasing. Dr. Koster-
mans worked up the Lauraceae of Madagascar,
which appeared in a series of articles in Bogor
and Brussels. A monograph on Durio by
Dr. Kostermans, assisted by a student of the
Biological Academy, Mr. Sugeng, has been com-
pleted.
The botanical activities of the Bureau of
Horticulture at Pasar Minggu, near Djakarta,
include the following:
(1) Distribution of fruit trees. An attempt
is being made to establish the correlation between
the distribution of fruit trees with ecological
factors, viz. rainfall, soil-type, hours of sunshine,
water-table, temperature. As an example of
these investigations, it has been found in various
grape-growing centres that it is very probable
that a minimum of 70% of sunlight is required
for success.
(2) Cabbage breeding. A scheme on the
breeding of high quality cabbage varieties capable
of seeding in the tropics is being worked out.
Crosses between high quality non-seeding varie-
ties and lower quality seeding varieties have
been carried out. A vernalisation project on
non-seeding high quality varieties is being
planned.
(3) Virus free potato seed stock. In connec-
tion with the problem of keeping a virus-free
seed stock collection of potatoes, insect free
chambers are being built in a glasshouse. Various
plant-indicators are already multiplied for the
VOLUME 4
BOTANY
purpose. Import of wild and cultivated species
of South America is being planned.
(4) Citrus decline. An investigation into the
cause of citrus decline is going on. Indications
are very strong that the decline is not due to
soil, but to other sources. Nematodes have
been isolated from roots of plants. Budding
trials on lime ("djeruk nipis") with scion material
from declined citrus trees shows stem-pitting and
vein clearing in the leaves.
The division of Phytosystematics, -geography,
and -sociology of the Faculty of Agriculture,
University of Indonesia, Bogor, put it as one of
the objectives of botanical investigations by the
senior students of the Faculty, the study of the
main features of phytogeography and -ecology
in Indonesia. Several reconnaissance expeditions
have been undertaken in 1953 to the Lawu Moun-
tain (3,300 meter above sealevel) in East-Central
Java, and Sumatra, in 1954 to Central and East
Java, Banten (West Java), and the islands in the
Bay of Djakarta. We have got the opinion that
in all probability the site factors climate and soil
series, together with the geographical location,
have an important influence on the structure and
floristic composition of our tropical plant com-
munities. After the departure of the Head of the
Division, Prof. Dr. S. Bloembergen, at medio 1 955,
in a very limited scale ecological surveys in trop-
ical rain forests have been carried out in East
Kalimentan and the Mount of Lawu. Further
botanical investigations are planned. Recently,
Mr. G.A. de Weille, staff member of the General
Research Station at Medan (North Sumatra),
has taken his doctor's degree in agriculture
at the Agricultural Faculty of Bogor, on a thesis
entitled "Blister blight control in its connection
with climatic and weather conditions".
Since 1953 the Central Research Stations
Association (C.P.V.) at Bogor has published the
following articles of botanical interest in English:
(1) Schweizer, J., The physiology of latex as
basis for tapping systems (1953).
(2) Wiersum, L.K., Results of some preliminary
experiments on stimulation of latex
yields (1953).
(3) Vollema, J.S. and Lasschuit, J.A., The
mildew resistant clone LCB 870 (1953).
(4) Vink, A.P.A. and Alphen de Veer, E.J. van,
Mechanical and chemical lalang control
(1953).
(5) Zeehuizen, J.J. and Arentzen, A.G.J.,
Divergencies between strain and mo-
dulus determinations (1953).
(6) Zeehuizen, J.J. and Schoon, Th. G. F.,
Preoculation of the yellow fraction by
alkaline salts of weak acids with preser-
vative properties (1953).
(7) Zeehuizen, J.J. and Schoon, Th. G. F.,
Oxalic acid as a coagulant in the pro-
cessing of rubber (1953).
(8) Zeehuizen, J.J. and Schoon, Th. G.F., The
influence of glyoxal on the vulcaniza-
tion characteristics of natural rubber
(1954).
(9) Wiersum, L.K., Observations on the rooting
of 1/cvea cuttings (1955).
(10) Wiersum, L.K., Physiological investigations
into the process of yield stimulation of
latex (1955).
(11) Paardekooper, E.C., Results of testing of
Hevca seedlings obtained by hand pol-
lination during 1927-1944 (1956).
(12) Paardekooper, E.G., The occurrence, pro-
perties and possible application of
juvenile type buddings (1956).
(13) Paardekooper, E.C., Further data on early
selection in Hevea (1956).
(14) Knaap, W.P. van der, Results of tea clone
experiments at Pasir Sarongge (1955).
(15) Knaap, W.P. van der, Notes on disease
incidence of Exobasidium vexans Mas-
see on tea (1955).
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
BOTANY IN NEW ZEALAND, 1953-1957
LUCY B. MOORE
Botany Division, C.S.I.R., Wellington, New Zealand.
PERSONAL NOTES
First must be mentioned the death of the
veteran botanist Dr. W.R.B. Oliver on 16 May
1957. Dr. Oliver published a revision of the
genus Aciphylla in October, 1956, (Trans. Roy.
Soc. N.Z. 84, pp. 1-18), and was active in field
work until the end of that year, making a collect-
ing trip to Norfolk Island in November. He
was able to carry on a limited amount of herba-
rium work until within a week or two of his
death. Another loss to New Zealand botany
was the death, in July, 1957, of Mr, G.O.K.
Sainsbury who, having completed studies on
New Zealand and Tasmania mosses, was working
actively on collections from some Pacific Islands
and from Malaya. The herbaria of both these
collectors are at Dominion Museum.
Distinguished botanical visitors to New Zea-
land during the years under review included
Dr. F.W. Went, Dr. H. Godwin, Professor R. D'O.
Good, Dr. W.M. Hicsey, and Dr. J.W. Gregor,
all for rather short periods. There was a good
representation of Australian botanists for the
A.N.Z.A.A.S. meeting in Dunedin in January,
1957, and the International Grasslands Congress
in November, 1956, and the Empire Forestry
Conference now meeting here have also brought
numbers of plant scientists to our country.
Dr. H.H. Allan was one of the twelve biolo-
gists to receive the honorary degree of Doctor of
Philosophy at the Linnaean Jubilee celebrations in
Uppsala in May, 1957.
With the appointment of Dr. W.R. Philipson
to the recently established Chair of Botany at
Canterbury University College, we have, for the
first time, a full Professor of Botany at each of the
four main University institutions.
Dr. G.H. Cunningham, F.R.S., retired from
the directorship of the Plant Diseases Division
of D.S.I.R. at the end of September, 1957.
The name of his successor has not been an-
nounced.
INSTITUTIONS AND SOCIETIES
Apart from the purely botanical departments
in the four University Colleges and two of the
museums, the Botany, Crop Research, Grass-
10
lands. Plant Chemistry, and Plant Diseases
Divisions of D.S.I.R., and the Forest Research
Institute, increasing amounts of work of direct
botanical import are coming from several other
bodies, some of which employ one or more
professional botanists, e.g., Dominion Physical
Laboratory, Soil Bureau and N.Z. Oceanogra-
phic Institute of D.S.I.R., soil conservation
service under Department of Agriculture, and
certain local bodies known as Catchment Boards.
Several rather recently formed societies meeting
annually discuss botanical problems, e.g., N.Z.
Genetical Society (established in 1949), N.Z.
Ecological Society (established in 1951), N.Z.
Microbiological Society, N.Z. Association of
Agricultural Scientists, and N.Z. Geological
Society. The last three formed since 1953.
In 1953, Botany Division, D.S.I.R., transferred
to Christchurch, though a small Sub-Station
is retained temporarily in part of the old quar-
ters in Wellington.
CRYPTOGAMS
ALGAE
In 1956, Chapman published a systematic
account of the marine Myxophyceae and Chlo-
rophyceae (Jour. Linn. Soc. 55 No. 360, pp. 333-
501). Dellow worked on ecology in Auckland;
Naylor investigated anatomical and life history
problems in brown seaweeds and produced a
check list of marine algae for Dunedin. Mason's
checklist of N.Z. Charophyta appeared in 1956.
E.A. Flint is currently working on soil algae as
part of a team dealing also with fungi, bacteria,
yeasts, amoeboid and ciliate protozoa, and
actinomycctes.
FUNGI
In systematics, there has been a steady flow
of papers from Cunningham and Dingley. G.B.
Cone has continued her collecting of agarics but
has not reached publication. A textbook on
Forest Fungi, prepared by M.E. Lancaster, was
published for Forest Service in 1955. A study of
Australasian Cyttariaceae (Rawlings, T.R.S.N.Z.
84, pp. 19-28) has interests across the Pacific.
VOLUME 4
BOTANY
Mycorrhiza and the rhizosphere generally have
been studied in relation to practical problems.
Here might be mentioned the work on plant
viruses at Plant Diseases Division, including
some very fundamental probing under R.E.F.
Matthews.
LICHENS
In Dunedin, there is great interest in this group,
cheifly by Wm. Martin and the chemist J. Murray.
Unfortunately there has been no N.Z. lichenolo-
gist amongst the scientists with the N.Z. Antarctic
party at McMurdo Sound.
BRYOPHYTES
Mrs. Hodgson's work on Hepaticae continues.
An important landmark was the publication in
1955, of Sainsbury's Handbook of the New
Zealand Mosses, with illustrations by Nancy
M. Adams (Roy. Soc. N.Z. Bull. 5, 490 pp.,
76 pi.).
PTERIDOPHYTES
Cytological work on ferns is in hand both at
Auckland and Canterbury University Colleges.
J.D. Lovis of Leeds spent a year, 1955-1956
here, continuing his investigation of Asplenium
trichomanes and comparing southern hemisphere
forms with those of the north. W.F. Harris'
Manual of the Spores of New Zealand Pterido-
phytes appeared in 1955 (D.S.I.R. Bull. 116.
186 pp., 10 pi. 4 fig.).
PALYNOLOGY
The appearance of Harris' Manual, of Cran-
well's account of pollen of N.Z. monocots in
1953, and of Couper's "Upper Mesozoic and
Cainozoic Spores and Pollen Grains" (N.Z.
Geological Survey Paleont. Bull. 22, 1953,
77 pp. 9 pi.) testify to the interest in this field.
The lack of the dicot pollen atlas promised by
Cranwell is still acutely felt, but, meanwhile,
Hards and Moar continue to publish work on
fresh pollen and that in recent deposits. Atmos-
pheric pollen studies seem to have been discon-
tinued meanwhile. Geologists are using plant
microfossils increasingly for solving chronolo-
gical problems which still concern us here.
Questions of the very recent history of the vege-
tation, e.g., in the last 50,000 years, are also much
to the fore and C 14 dating services are rather
generously available. At this very important
early stage in sketching the outlines of ecological
change, there seems to have been less than full
appreciation of Faegri's advice that only a trained
field botanist will be able to utilise the pollen
analysis technique fully, and some very poorly
substantiated statements have been used as evi-
dence in papers by geologists and physicists.
TAXONOMY OF FLOWERING PLANTS
Revisions of individual genera continue to
appear, but Allan's new Flora, Zotov's account
of the grasses of New Zealand, and Healy's
manual of the adventive plants in New Zealand
are all awaited. Preparation is well ahead at
least with the first of these.
A noteworthy event has been the flowering
and fruiting in cultivation of Tecomanthe speciosa,
plants having been established by cuttings from
the only known wild plant on Great Island of
the Three Kings group.
EXPERIMENTAL TAXONOMY AND
CYTOLOGY
Experimental and cytological studies are
helping to clear up taxonomic problems in
certain groups, e.g., Baylis on Solanum, Brownlie
on ferns, Connor on Agropyron, Brockie on
Epilobium, Rattenbury on several genera, and
Hair on conifers in particular (studies not con-
fined to N.Z. material) and in general on a
long-range project of a chromosome atlas of
New Zealand plants.
GENETICS
This is mostly on plants of economic impor-
tance, e.g., crop plants, pasture grasses, and
timber trees. Bannister's study of a progeny of
Pinus radiata is giving positive results after only
six years. Godley is working on dioecy in indig-
enous plants.
ANATOMY
Lively and varied interest is indicated by these
titles from A.N.Z.A.A.S. meeting: "Aspects of
leaf development in Dicotyledons," B.F. Slade;
"Transformation of vegetative shoot-apex into
the Primordium of a flower," W.R. Philipson;
"Effect of various treatments on the Anatomy
of the stems and roots of apple trees," D.W.
McKenzie. A paper by R. Licitis-Lindbergs on
branch abscission and disintegration of the female
11
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
cones of Agathis australis Salisb, (Phytomor-
phology6, 1956, pp. 151-167) cleared up problems
that have long awaited attention.
PLANT PHYSIOLOGY
A most spectacular change in emphasis in
botanical research here is the increasing interest
in plant physiology. This does not stem at
all obviously from leadership in the University,
but seems to come from practical requirements
in a country where the national economy is
based directly on pasture production. This has
become, for instance, important in the Dominion
Physical Laboratory, an extract from whose
current programme reads (D.S.I.R. Annual
Report, 1957, p. 48) "measurement of mechanical
properties of plants and the investigation of the
relationship between these properties and the
plant's environment; investigation of the moisture
movement through a soil caused by thermal
gradients; setting up plant cabinets to provide
adjustable controlled environments for growing
plants; development of equipment for the
measurement of the amount of heat and water
vapour carried vertically by atmospheric turbu-
lence; measurement of the transpiration rate of
plants under different soil moisture conditions
and in different environments; and, investigation
of how evaporation over a pasture is influenced
by pasture length."
It seems strange that these projects are under-
taken by a laboratory which has no trained bota-
nist on its staff. It does, however, work closely
with Grasslands Division, which is studying
various other aspects of physiology of pasture
plants. Basic work on photosynthesis is coming
from the Plant Chemistry Laboratory.
Isotopes have been used in various investiga-
tions and at A.N.Z.A.A.S., Dr. Cone reported
results using N 15 to support the evidence from
cultural studies that nitrogen fixation occurs
quite generally in foliage of pioneer shrubs and
some weeds and trees.
Walker and his co-workers have demonstrated
the importance of added sulphur for optimum
plant growth in many areas. Jacques' long-term
studies on roots continue, and there is much
activity in seed research in relation to pasture
plants, forest species, and weed control. An
aspect of plant chemistry that has been given some
newspaper publicity is a survey of extracts of
indigenous plants at Auckland University College
for the Cancer Research Project.
12
ECOLOGY
Descriptive ecological surveys continue to
hold an important place. The National Forest
Survey, after ten years of intense activity, has
reached the stage where a full report on "all
essential phases of the volumetric survey of
merchantable-quality indigenous forests" is about
to be released. The same organisation, within
the Forest Service, is now proceeding with three
related projects: (1) Ecological survey to lead to
regional forest accounts accompanied by mile-to-
the-inch forest-type maps; (2) Protection forest
survey covering almost a third of the total land
area of New Zealand, where the vegetation is
non-commercial forest, mostly above 2,000 ft.
altitude, and unoccupied shrublands, grasslands,
and moorlands that lie above forests; here the
prime management objective must be regulation
of the water yield; (3) Exotic-forest survey (F.R.I.
Annual Report 1957, 24).
Out of the Forest Survey arose Holloway's
outstanding paper "Forests and Climates in the
South Island of New Zealand" (T.R.S.N.Z. 82, 2,
1954, 329-410). This put forward the hypothesis
that "the forests, as a whole, are in an unstable
condition consequent on comparatively recent
changes in regional climates and that, as a result,
and active redistribution of species is in progress
with resultant development of a wide range of
new, though by no means stable, forest types."
Evidence is set out to show that the forests of
the whole of the South Island are comprehensible
in terms of this hypothesis which is shown not to
be in conflict with evidence derived through
studies of the soils or through study of Polynesian
traditions. This paper is obviously one of the
highlights of botany in New Zealand in recent
years.
Regional or local ecological surveys include
several on tussock grasslands (Barker, A. P.; An
ecological study of tussock grassland. D.S.I.R.
Bull. 107, 1953. Tussock Grassland Research
Committee: The high-altitude snow-tussock
grassland in the South Island of New Zealand.
N.ZJ. Set. Tech. 36, 1954. Riney, T.A., and
Dunbar, G.A.: Criteria for determining Status
and Trend of High Country grazing lands in the
South Island of New Zealand. Soil Conserv.
and Rivers Control Council, 1956. Wardle, P.,
Mark, A.F.: Vegetation and Climate in the
Dunedin District. T.R.S.N.Z. 84, 1956. Miller,
F.L. et. al.\ Shotover River Survey. Otago
Catchment Board Bull. 1. 1956). In this connec-
tion, Gradwell's studies on frost action in high
VOLUME 4
BOTANY
country are relevant. In the North Island, there
are accounts of forests of Taranaki and West
Taupo, and of a rather different kind, Druce's
detailed account of a single catchment of a
few acres which is to be reserved for long-term
studies of soil formation and change.
The "General Survey of the Soils of North
Island" (1954), the publication of a map showing
the North Island soil-forming ash showers, and a
pamphlet on "Soil Pattern in New Zealand"
will all help plant ecologists.
Turning to ecology of seashore, Chapman's
"Mangrove and Salt Marsh flats of the Auckland
Isthmus" is in the press and intertidal studies
include Dellow's comprehensive account of
the shores of the Hauraki Gulf and those of
G.A. Knox and E.J. Batham in the South Island.
Field work has not been confined to the main
island and there have been expeditions to or
reports on Auckland, Chatham, Three Kings,
Kermadec, and Pitcairn Islands, and New
Caledonia. R. Cooper of Auckland Museum was
working during the winter of 1956 on taro and
kumara in several parts of the Pacific, and it is
understood that he has a generous Rockefeller
grant for similar work further north.
PESTS AND DISEASES
Members of the Plant Diseases Division
produced a manual on "Plant Protection in New
Zealand" (704 pp. 477 illustrations, 1956); and
in 1954, Chamberlain presented a comprehensive
account of plant virus diseases in New Zealand.
At Forest Research Institute, a full-time Forest
Biology Survey was commenced early in 1956,
the overall purpose being "to maintain a contin-
ual and systematic inspection of all forests for
the presence of newly introduced insects and
fungi, and to detect any dangerous increases in
the numbers of existing pathogens." Research
into the incidence and effects of "noxious animals"
especially deer and possums, has been stepped up,
but the teams working on the relations of these
animals to vegetation are not as strong botanically
as could be wished. A spectacular drop in the
number of rabbits (without infection with my-
xomytosis) has coincided with a series of better-
than-average seasons, leading to striking vegeta-
tion changes in some of the more rabbit-ridden
dry districts. These effects too, have been quite
inadequately documented owing to the shortage
of field botanists.
Manuka blight, a disease of Leptospermum
caused by a scale insect, has spread partly by
natural dispersal and partly by deliberate transfer,
until now, ten years after first being noticed,
it occurs throughout New Zealand. Affecting
one of the commonest and most widespread of
shrub species, it will certainly have direct effects
on forest regeneration, though it has been
welcomed by farmers whose poorer pastures have
always been subject to invasion by manuka.
A most dangerous addition to the weed flora
is water hyacinth, which has recently been found
to seed freely in many waterways and has already
entailed an expensive control programme.
On the other side of the picture, plant quaran-
tine provisions have been much improved with
the recent installation of modern methyl bromide
fumigation plants at several main ports.
CONSERVATION
At the time of the setting up of the National
Parks Authority in 1953, there were five National
Parks, variously controlled. To these have been
added three more (Mount Cook, 151, 800 acres;
Nelson Lakes, 139,833 acres; and Urewera
119,614 acres, and it was enlarged in 1957).
Moves are afoot to secure reserves in other places,
especially in Chatham Islands and in the country
west of Lake Taupo, both regions where active
agricultural developments have now been made
possible by use of modern machinery and, in the
volcanic country, by the use of cobalt to keep
stock healthy. In both places, the remnants
of highly characteristic vegetation are dwindling
rapidly.
13
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
REPORT OF THE SUBCOMMITTEE ON BIBLIOGRAPHY
E. H. WALKER
Smithsonian Jstitution, Washington, D.C., U.S.A.
The subcommittee members were selected to
represent as many countries and branches of
botany as possible. When completed the commit-
tee represented continental United States, 6; Ha-
waii, 2; Japan, 1; Free China, 1; New Zealand,
2; Australia, 1; and Holland, 1; a total of 14
members. The majority are taxonomists, but
phytogcography, ecology, economic botany,
history of botany, and library science are repre-
sented.
As there was no previous committee on whose
activities a program could be built, the first activity
was to circularize the members and a few others
for their ideas of the possible aims and activities
of the subcommittee. Following is a summary of
some of the suggestions submitted. Some members
pointed out the need for defining more specifically
boundaries of the area to be included, its present
limits being a bit indefinite. Severval asked that
a survey be made of the areas and fields now
covered by bibliographies, their perspectives and
adequacy, thus revealing where there are gaps to
be filled. In order that such information be made
available to users, it was suggested that a regulary
appearing bibliographical serial be established.
Some pointed out the need for a guide for doing
bibliographical work and the establishment of
certain standards and procedures. For the benefit
of workers without access to adequate libraries,
cooperation between individuals and institutions
for loans and microfilming, as well as certain
bibliographical research were suggested. Jt was
further proposed that editorial assistance and
guidance be established to aid authors to prepare
their bibliographic manuscript, so they will be of
maximum value to researchers and librarians,
the principal users of bibliographic tools. Close
cooperation with established bibliographic enter-
prises, notably the Society for the Bibliography
of Natural History in England and Biological
Abstracts, was suggested. An enlarged new
edition of E. D. Merrill's Bibliography of Pacific
Botany was proposed. Several of these needs
would be provided by establishing a bibliographic
service at some centrally located institution,
closely linked with similar work at other institu-
tions, thus broadening the service. Finally,
careful consideration was urged for the adoption
of modern new techniques in bibliographic pro-
14
cedures so as to broaden their usefulness and to
conserve limited funds.
Two progress reports were issued by the sub-
committee, including summaries of these sugges-
tions and reports on various known activities
of the members of the committee and other perti-
nent endeavors. These were mimeographed and
distributed by the staff of the Pacific Science
Association to the members of the subcommittee
and others, especially to the chairmen of the
other subcommittees and standing committees,
and to certain institutions. The present report
constitutes in effect the third report.
It may be well here to report on various pro-
jects in this field. The chairman aided Mrs. Ida
K. Langman in obtaining a National Science
Foundation grant which enabled her to spend
nearly a year searching in Mexican libraries for
material for her extensive bibliography of Mexican
botany. She is now extending her work by
similar searches in American libraries. The
chairman has rendered some assistance as a
member of the editorial board for the vegetational
bibliographies being prepared by Jack McCormick
at the American Museum of Natural History.
He reports little recent progress due to other
obligations. It was found that the Subcommittee
on Algae is deeply involved in bibliographic work,
recognizing that this is fundamental to the
development of any field of study, especially in
its initial stages. Just how the Subcommittee on
Bibliography can aid in the bibliographic work
of the other subcommittees remains to be worked
out.
For several years Robert Cooper, botanist, and
Enid Evans, librarian, both at the Auckland
Museum, have been preparing a bibliography of
New Zealand botany. Their major problem
seems to be finding someone with access to
literature on that area who has the time, ability,
and resources to run down known-to-exist items
not available to them. Thus the establishing of
the bibliographic service proposed above seems
urgently needed.
Two most important bibliographies have been
issued during the existence of this subcommittee.
First is the bibliography in volume five of the
Flora Malesiana prepared by Mrs. Van Steenis-
Kruseman. It will be of inestimable value to
VOLUME 4
BOTANY
workers over a wide area. Participation in its
subsequent improvement has been solicited
by distributing separately bound interleaved
reprints to key workers. The second important
contribution is Island Bibliographies by M.H.
Sachet and F. R. Fosberg. This is in effect a
cover title for three separate bibliographies,
entitled: (1) Annotated Bibliography of Micro-
nesian Botany, (2) Bibliography of Land Ecology
and Environment of Coral Atolls, and (3) Selected
Bibliography of the Vegetation of the Tropical
Pacific Islands. This was duplicated by the
National Research Council.
R.T. Hoogland of Canberra, Australia, has
revealed that, as opportunity permits in connec-
tion with work on his New Guinea collections, he
is accumulating material for a botanical bibliogra-
phy on that great area. Miss Nancy Burbidge,
also of Canberra, is accumulating bibliographic
material in connection with preparation of a
generic list of Australian plants. Edwin H. Bryan
of the Bishop Museum maintains several valuable
bibliographic files on various subjects in the
Pacific and has submitted two papers for the
bibliography symposium of the Congress.
The chairman of this subcommittee, in cooper-
ation with F.R. Fosberg and M.H. Sachet, and
under grant from UNESCO, has prepared a manu-
script supplement to Merrill's Botanical Bibliogra-
phy of the Islands of the Pacific. Dr. Merrill,
on retiring from his long and highly productive
work on Pacific botany, turned over his materials
accumulated for another edition of his well-
known Pacific bibliographies to E. H. Walker
and F. R. Fosberg. The UNESCO grant was
insufficient and other obligations too great to
permit the preparation of the supplement that
will eventually be needed, but this limited work
will go a long way in the eventual preparation of
that work. It is in effect a token recognition of
Dr. Merrill's legacy to Messrs. Walker and
Fosberg.
Work on the needed supplement to the
Merrill and Walker Bibliography of Eastern
Asiatic Botany may begin in the near future,
because of as yet informal encouragement by
the National Science Foundation and the Ameri-
can Institute of Biological Science, Encouraging
promises of full cooperation by Japanese and
Chinese botanists were received by the chairman
on his recent visit to those countries. The pro-
mises are bright that this supplement will better
meet the needs of the Oriental botanists than did
the original work.
Time prevents reporting here on various
projects that may come within a broader defini-
tion of bibliographic work. The term is not
confined to the preparation of bibliographies,
but embraces all work with books. It would
scarcely be justified to attempt to enumerate here
the botanical works that have been or arc being
prepared in this area or the library developments
that are pertinent to bibliographic work in its
widest sense, but such would indeed be very illu-
minating and profitable.
In June, 1956, the chairman of this subcom-
mittee was asked by the Chairman of the Organiz-
ing Committee on Botany of the Ninth Pacific
Science Congress to convene a symposium on
Bibliographic Problems in Natural History in
the Pacific to be held at this Congress. This
constitutes the culminating activity of this sub-
committee. The objective of this symposium will
be to receive the ideas of its participants through
a limited number of distributed and delivered
papers and through free discussion at both formal
and informal gatherings. The results will then be
summarized. It is hoped that from the various
reports of the Subcommittee on Bibliography and
the coming symposium a more concrete program
for the development of this work in the Pacific
may be formulated and implemented.
DISCUSSION
F.E. EGLER: I would like to draw your attention to a
bibliography project now centered at the American
Museum of Natural History (New York 29). This project
is essentially the responsibility of Dr. Jack McCormick of
that institution. It is, or was until recently, entitled
Regional Vegetation Literature. It is not an all-botany
bibliography, but one concerned specifically with vegeta-
tion (plant community) publications with special emphasis
on identifying the localities at which studies have been
made. The area involved at present is North America and
South America, and this is part of our specific area. I
understand that they are seeking collaborators for different
countries. Publication of the first of a new series of
publications is already far advanced.
F.R. FOSBERG: The supplements to Merrill's Bibliog-
raphy of the Pacific Islands and to Island Bibliographies
were aided by a small financial grant by the UNESCO Humid
Tropics Committee. No provision was made for their
publication, but there is some indication that funds may be
available for their publication. 1 would suggest that this
session could appropriately adopt a recommendation that
15
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
these supplements be published so that they would be of
help to others.
M.S. DOTY: The phycology sub-committee has recog-
nized "bibliography' 1 as perhaps the foremost of the
barriers to phycological research which might be lowered
by efforts of the sub-committee. Thus, after consultation
with Dr. Walker, compilation of bibliographies was begun
on a form adoptable to his extension of E.D. Merrill's
Bibliography of Pacific Botany. This material should be
considered for inclusion in the present "First supplement
... M to that publication.
C.G.G.J. VAN STEENIS: Naturally I want to support
publication of bibliographies. However, it would be very
desirable to have them in printed form, instead of less well
surveyable and more bulky mimeograph. Has Dr.
Fosberg tried to have them published in the Bernice Bishop
Museum Bulletins or Philippine Journal of Science which
have a wide circulation in the Pacific area?
16
VOLUME 4
REPORT OF "THE : SWCOMMITTEE
BOTANY
M. S. DOTY
University of Hawaii ', Honolulu, Hawaii.
Two mimeographed circulars were prepared
and sent to members of the subcommittee. These
evoked considerable interest and demand. We
suggest the practice be continued, but duplication
and distribution should be done by the Pacific
Science Association Secretariat in Honolulu.
Participation in bibliographic matters was
begun. This was after conferences and corre-
spondence with the Bibliography Subcommittee
Chairman and with a local group assembled in
Honolulu to make a report to that subcommittee.
In conformation with the style used by E.H.
Walker of that subcommittee, bibliographic
materials have begun to be assembled in Honolulu
regarding algae in the Pacific. Jt is planned that
this information will be made available to anyone
at cost or, eventually, be duplicated. We do not
anticipate duplicating the various efforts to pub-
lish bibliographic materials but, rather, assist in
their extension and help make them available
to prospective users.
The major phycological problems have been
sought by correspondence with the subcommittee
members and enumerated in the mimeographed
information bulletins distributed to the sub-
committee. These suggestions were to promote
floristic studies, develop bibliographic sources,
facilitate exchange of ideas and materials, and
encourage functional studies of the algae.
It is further suggested the Standing Com-
mittee on Botany of the Pacific pass a resolution
strongly favoring adherence to the International
Botanical Code of Nomenclature and in particular
to those pre-eminent principles the type method
and priority.
17
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
REPORT OF THE SUBCOMMITTEE ON COMMON NAMES OF
PACIFIC PLANTS
MONA LISA STEINER
Pasay City, Philippines.
INTRODUCTION
The complex problem of common names of
plants, their variability, recording, and stability
in some respects, was brought before a session of
the Eighth Pacific Science Congress. Although
standardization was undertaken successfully in
some regions, it was pointed out that standardiza-
tion of common names of plants in the Pacific
Basin is not feasible, because too many language
groups are encountered in this part of the world.
It was consequently agreed that a subcommittee
on Common Names of Economic Plants be estab-
lished, which would work towards the compila-
tion of a list of common names of plants in
active use in the Pacific Basin together with their
botanical equivalent, to be listed in form of a
dictionary.
As such a vast task can not be accomplished
uno acttt, the project was divided into the follow-
ing groups: foodplants, forest trees, weeds and
forage plants, and ornamentals. For the Ninth
Pacific Science Congress, only foodplants were
worked on.
A list of basic foodplants, based on Philippine
material, was prepared and sent to botanists
interested in vernacular names and familiar
with certain regions in the Pacific. Only generally
used foodplants were included, not famine
foodplants or those of local occurrence. Varie-
ties and hybrids were generally not listed, also
not all names of products, as those of the coco-
nut, for instance, could fill a book.
In the beginning only tropical plants were
included, but later on it was apparent that even
in the tropical Philippines many temperate zone
foodplants are being cultivated and that a separa-
tion was impractical. In the first attempt, China,
Japan, and other countries not directly bordering
the Pacific, such as Thailand, were not included;
but as the compilation progressed, it became evi-
dent that such additions might prove very useful,
as few references are available in Roman script
in those languages.
MEMBERS OF THE COMMITTEE
Miss Marie Neal
Bishop Museum, Honolulu 17, Hawaii.
Mr. B.E.V. Parham
Department of Agriculture, Suva, Fiji.
Mr. R.E. Dwyer
Director of Agriculture, Papua, Port Moresby.
Dr. F. Raymond Fosberg
c/o National Research Council, Washington, U.S.A.
Dr. Shiu-ying Hu
Arnold Arboretum, Jamaica Plain, Mass., U.S.A. (since 1957).
Mr. H. Ando
Hiroshima University, Sendamachi, Japan (since 1957).
Mr. Faustino Miranda
Institute de Bilogia, Casa del Lago, Chapultepec.
Miss N. Burbidgc
Canberra, Division of Plant Industry, Australia.
Polynesia.
Fiji.
Papua.
Micronesia.
China.
Japan.
Mexico.
North Australia.
18
VOLUME 4
BOTANY
Mr. H. Keith
has resigned on account of transfer.
(North) Borneo.
Mr. Jacques Barrau
Laboratoire d' Agronomic Tropicale, Marseille, France.
Melanesia.
Ida Langmann
Academy of Science,
Philadelphia, U.S.A.
West Coast of North America.
Dr. Mona Lisa Steiner Philippines.
2833 Park Avenue, Pasay City, Philippines, chairman.
ACCOMPLISHMENTS
As many references as possible were scanned in
Manila for vernacular names of focdplants in
the Pacific Basin and combined with published
and unpublished lists of vernacular names sent by
members of the committee and others as listed in
the bibliography. Sincere gratitude is herewith
expressed here for all the valuable cooperation.
The Pacific Basin was then divided into various
regions, such as Micronesia, Polynesia, Malane-
sia, and the locality or region in which a common
name is being used is then indicated. Sometimes
numerous languages are encountered within one
region, yet frequently one name is of more
general usage, while others are of local occur-
rence only. In the Philippines, for instance, one
language of about 80 has been selected as the
National Language, namely Tagalog, and this
language can be understood all over the country.
The various common names of a region have
been compiled, but the more generally used name
has been underlined. In addition to the various
Malayan, Polynesian languages, etc., the common
names of foodplants in Chinese, Dutch, English,
French, German, Japanese, and Spanish were
added.
Although the chairman is well aware that such
a compilation can never be complete, an attempt
has been made to compile all the gathered infor-
mation in the form of book. With the valuable
assistance of the National Research Council in
the Philippines and the aid of Dr. Patrocino
Valenzuela 200 copies of A PRELIMINARY
COMPILATION OF VERNACULAR NAMES OF
FOODPLANTS IN THE PACIFIC (195 pages)
were mimeographed in February, 1957, and
sent to all members of the committee and
institutions that might aid in this undertaking.
Members of the committee have then been asked
to check the vernacular names given, fill in gaps,
also to underline the most generally used name
in a certain region. The preliminary compila-
tion thus checked and filled in was to be returned
to the chairman before the Ninth Pacific Science
Congress.
Since the compilation was sent out, much new
material was gathered by the chairman who went
on six months leave in Europe. Particularly,
the common names in the major languages were
filled in. Among those who returned a checked
copy was Mr. H. Ando, Hiroshima University
who added more Japanese names. Dr. Shiu-Ying
Hu, Arnold Arboretum, is preparing Chinese
equivalents. Other new additions, too, are waiting
to be included. After the various additions and
revisions are made and incorporated, it is hoped
that the first volume of the compilation will
come out in print.
SUGGESTIONS
1. The arrangement of compiling plants alpha-
betically within families was more practical for
the compiler, but should not be followed in the
final work. The scientific names are to be
arranged alphabetically.
2. No personal acknowledgements have been
made in the preliminary report, but all used
references, printed and unpublished, were included
in the bibliography. The final dictionary, however,
should have special acknowledgements.
3. So far no cross index has been made, but as
soon as all the new information has been
gathered, a cross index should be prepared.
4. A taxonomist of standing should check the
validity of the used scientific names and their
synonyms.
19
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
5. Other workers should be appointed to work printing of the revised and enlarged compilation
on forest trees and medicinal plants while the of common names of foodplants. It should also
work on foodplants is to be completed. be distributed through the Pacific Science Asso-
6. A fund should be made available for the ciation.
20
VOLUME 4
BOTANY
REPORT OF THE SUBCOMMITTEE ON ETHNOBOTANY
T. P. BANK II
Institute for Regional Exploration, University Station, Ann Arbor, Michigan, U.S.A.
The Subcommittee on Ethnobotany of the
Standing Committee on Pacific Botany was organ-
ized in 1954-55 and is composed of twenty-five
members.
The Chairman has circulated a request for
information about significant ethnobotanical
studies currently underway in various parts of
the Pacific area. Replies have been slow, but
those that have been received form the basis of
the following report.
SOUTH PACIFIC
Dr. Toichi Mabuchi of Tokyo reports that the
Japanese Society of Ethnology plans to begin
a field study soon on rice agriculture in Southeast
Asia, notably in Thailand and Cambodia, which
will require three years. He is continuing his own
studies of aboriginal tribes of central Formosa
and has investigated the correlations between
social organization and agriculture. Mr. Kokichi
Segawa, who is now in Brazil for the Japanese
Government, has been collaborating with Gordon
Bowles in the compilation of An Illustrated
Ethnography of the Formosan Aborigines, Vol. I:
The Yami by Segawa and Kano, and this work
includes ethnobotanical illustrations and des-
criptions.
Thor Heyerdahl has continued his field work in
the Pacific searching for new evidence for diffu-
sion. Dr. George F. Carter suggests that the
discussions at Bangkok "might well include some
review of this ... with a call for further re-exami-
nation of the botanical evidence for diffusion."
Carter urges that botanists and anthropologists
collaborate to "get that ethnobotanical dic-
tionary moving, and to include in it meanings or
names, whenever possible, and to put in the plant
usages, and most especially ritual usages."
Dr. Jonathan L. Hartwell, Laboratory of
Chemical Pharmacology at the U.S. National
Institutes of Health, is collaborating with various
colleagues in an extensive survey of the literature,
both from America and abroad, and folklore,
pertaining to medicinal plant uses. The purpose
is primarily pharmacological to open up leads
for experimental work on the isolation of active
substances from plants. He is especially interested
in plants used for the treatment of cancers,
tumors, warts, and corns, and he requests that
botanists and anthropologists who are similarly
interested communicate with him for an exchange
of information.
Dr. Harold St. John, who is a subcommittee
member, will not be at Bangkok but will submit
papers to two sections, on taxonomy and verna-
cular names. He is currently in the field for three
months studying Pandanus in Netherlands New
Guinea, Australian New Guinea, Papua, and
Queensland. From this and his other extensive
work with Pandanus will undoubtedly come future
ethnobotanical reports.
Dr. Robert Ornduff writes from Berke^y that
his current sphere of activity is non-ethnobota-
nical, that he is doing biosystematic work with
Senecio species from New Zealand, as well as
with Californian Composite genera and the genus
Senecio in the western United States. He is also
collaborating with anthropologists at Reed
College in a project on the Warm Springs Indian
Reservation in Oregon.
Jacques Barrau has prepared a paper on "Notes
on the significance of some vernacular names of
food plants in the South Pacific Islands" which is
of considerable interest to ethnobotanists.
R.D. Hoogland writes from Australia that
"Botanists in Australia have paid very little
attention to ethnobotany. Apart from the enor-
mous amount of work still needed in local taxo-
nomy and the pressure on economic aspects, this
can probably partly be accounted for by the small
aboriginal population and the seemingly little
use made of the native flora."
"The latter seems to be the case also in New
Guinea. In comparison with the islands of Indo-
nesia, where the native flora is extensively used
in medicine, very little use is made by the New
Guinea natives of their vast resources in this
field. During my collecting trips in New Guinea,
I have tried to collect material of native drugs
for chemical testing and found it very difficult to
get any at all. Others have noticed the same,
e.g., D'Alberts (1880); Professor Baas Becking
tells me that he has the impression that this
applies to the whole of Melanesia."
Anthropologists have been working in various
areas in New Guinea and have assembled various
21
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
ethnobotanical data, none of which, however, has
been published in recent years. These data apply
in particular to food-plants, both cultivated and
collected in the forest. Others have been working
in Sumatra and Borneo (from Australia ed.)."
Robert Suggs is currently doing archaeological
work in the Marquesas. Previously he indicated
to Professor Bartlett that he would try to collect
as much ethnobotanical information as he could
and would seek native names for the plant spe-
cimens he hoped to bring back.
George Carter adds a further note to the effect
that Heyerdahl's Galapagos expedition "demon-
strated that the Peruvians had sailed to those
islands and used them as fishing stations for
hundreds of years. Since this was presumably
done with balsas, it strengthens his case that the
Peruvians were fully capable of sailing into the
the Pacific and returning... (linguistic similarities
in Polynesia and South America) the total culture
picture suggests extremely long continued con-
tacts, with some of the later ones coming from
America."'
Jonathan Sauer writes from the University of
Wisconsin that he is continuing his work with
grain amaranths, and with Sasuke Nakao has
published further data and interpretations of
Asiatic grain amaranths, crops domesticated by
the American Indians and introduced to Asia by
unknown means (Land and Crops of the Nepal
Himalaya, ed. H, Kihara, pp. 141-146. Kyoto,
1956). '
Dr. Sauer is also continuing to grow specimens
of pot-herb amaranths collected in seed from
various parts of Southeastern Asia, preparatory
to systematic study, and he reports that he is
currently engaged in a monographic study of all
species of Canavalia (sord-bean and jack-bean)
from all parts of the world, with special emphasis
on the aboriginal cultivated varieties. He invites
workers to send him specimens of Amaranthus
or Canavalia and promises to identify them and
give a report to the collectors.
Dr. Tom Harrisson indicates that he has col-
lected considerable information on the eth-
no -ecological aspects of primitive agriculture in
Borneo and that he is willing to exchange infor-
mation. He is reporting on ethnobiological lore
in Borneo at the Congress.
Harold C. Conklin is currently in the Philip-
pines to study tropical forest agriculture and will
present a number of papers at the Bangkok
meetings.
Professor Bartlett, now an emeritus professor
at Michigan, is principally engaged in compiling
22
a gigantic annotated bibliography on the subject
Fire in Relation to Primitive Agriculture and
Grazing in the Tropics. Volume 1 has already
been distributed and is about out of print; volume
2 is out and ready for distribution (A-G, Tropics
in general; H-J, South Asia and Oceania a total
of 873 pages); and he has enough material already
gathered for a third volume of equal proportions.
He is also actively advising ethnobotanists,
archaeologists, and botanists who are in the field
or about to take to the field as to what problems
still need to be attacked. He writes that "sweet
potatoes introduced into the Orient by the Spanish
should be compared in every way, cytologically,
chromatographically, and otherwise, with those
that spread into Polynesia from New Zealand."
He is especially interested in the possibilities of
tracing sweet potato migrations by means of
chromatography. He also wants someone to
study native gardening carefully all along the
Southeast Asia coast.
As an appendix, a brief report is attached
covering studies on Manihot esculenta by Dr.
David Rogers who is continuing this work at the
New York Botanical Garden, in addition to
editing Economic Botany.
NORTH PACIFIC
In Hokkaido, Japan, Dr. Robert Austerlitz
has been working for several seasons with Gilyak
and Ainu informants. His project is primarily
a linguistic study, but he has developed an active
interest in ethnobotany and has compiled a large
annotated list of plants used for foods, medicines,
poisons, and handicrafts. His specimens and
field identifications have been checked by Dr. M.
Tatewaki of the Botanical Institute at the Hok-
kaido University.
T.P. Bank II is continuing studies of Aleut and
Pacific Eskimo plant lore. Recently he spent
a year in Hokkaido to compare this material with
what is known about Ainu plant uses. He is
especially interested in Eskimo- Aleut migrations,
Aleut medical knowledge, and the question of
cultural diffusion across the North Pacific.
Bank is commencing an ethnobotanical and
toxicological study of aconite poison whaling in
the North Pacific, in collaboration with workers
at Hokkaido University. Part of the study will
include an evaluation of the variation of toxicity
in various races of Aconitum and the significance
of this to anthropological questions pertaining to
the use of aconite poison by primitives in the
North Pacific region.
VOLUME 4
BOTANY
Archaeological studies by Bank and others at
Michigan have also utilized ethnobotanical
methods for determining chronology of cul-
tural events in the Aleutians. Radiocarbon
dating of archaeological deposits and pollen
analysis have indicated that the Aleuts arrived in
the Aleutians at least 3,500 years ago at a time
when the climate may have been appreciably
colder. Preliminary results of pollen studies show
that the beginning of amelioration of climate
toward the onset of the thermal maximum in the
Aleutians may have occurred between 4,900
and 4,000 years ago.
Others are doing botanical and ecological work
in the Aleutian area, including Robert Kindschy,
Jr. at the University of Idaho, who collected
ethnobotanical data among the Nikolskian Aleuts
in 1957.
No reports have been received from ethno-
botanists at the University of Washington, but
presumably studies are being continued among
the Indians of that region and in southeastern
Alaska.
23
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS^
APPENDIX
SUMMARY OF STUDIES ON MANIHOT ESCVLENTA
DAVID ROGERS
New York Botanical Garden, New York, U.S.A.
PURPOSE
1. To provide a convenient classification of
the cultivars of Manihot esculenta.
2. To demonstrate relationships among the
varieties.
PROCEDURE
1. Assemble cultivars in museum plots for
ready study of the varieties.
2. Classify, using morphological character-
istics, in a framework of taxonomic categories.
DISCUSSION
Jamaican and Central American cultivars represent the
range of material so far studied. In Jamaica, a collection
of 107 named varieties were brought together from all
parts of the island at Bodies Experimental Farms. After
considerable study, forty-five cultivars were recognized.
In Costa Rica, at the Inter-American Institute of Agri-
cultural Sciences at Turrialba, Dr. Jorge Leon had collected
the locally occurring cultivars. To this collection were
added variants from other Central American countries,
Cuba, a few from the East Indies, and several from Brazil.
In this collection about twenty-five cultivars are found to
be distinct.
The cultivars from both areas fit into a framework of
sub-specific categories of two convariants (equivalent to
subspecies) and several sub-convariants (comparable to
series).
The classic divisions of Manihot esculenta (sweet vs.
bitter) have been set aside in this work. No morpholog-
ical categories are coordinate with such a division.
Work is in progress to publish a key to the cultivars
studied, and proposals for continuation of the studies in
South America are being made.
24
VOLUME 4
SUPPLEMENT "TO^THE REPORT~6F
THE SUBCOMMITTEE ON ETHNOBOTANY
BOTANY
RECOMMENDATIONS
The distinctions between ethnobotany on the
one hand and economic botany on the other are
not always clear. This has been brought out
during the present Congress, where papers repre-
senting both points of view are presented in a
symposium on the ethnobotany of Thailand and
contiguous countries. Indeed, ethnobotanical
reports are distributed throughout various ses-
sions dealing with a number of diverse disciplines,
including anthropology, botany, linguistics,
conservation, and agriculture. This is not a
deadly sin, but it does make for a lack of coordi-
nation and a scattering of thoughts and conclu-
sions, when perhaps the true function of the study
of ethnobotany is to bring them together.
Ethnobotany should deal with verbal traditions
only, except for those traditions which have
recently found their way into the literature,
whereas economic botany is more concerned with
the modern concepts of the science of plants and
of their uses. The word "primitive" has various
connotations among different people, but perhaps
it is still our best key to the content of ethno-
botanical study. Ethnobotany is the study of
the interrelations of primitive man and plants, in
the sense that primitive denotes a lack of any
written language and therefore the preservation
of traditions by verbal means only.
Chemical analysis of useful plants is not usually
considered a part of ethnobotanical work, but
the study of plant debris in archaeological depo-
sits, studies of plant patterns in the vicinity of old
village sites, pollen analysis, and radiocarbon
dating are all related to our main objectives in the
study of ethnobotany.
With this rather brief definition of the content
of pur study, the following specific recommen-
dations are made in the hope that they will stimu-
late a more thorough consideration of how
ethnobotanical research can contribute most to
the solution of problems in the various disciplines
of which ethnobotany is a part.
1. A summary of our state of knowledge of
Southeast Asian ethnobotany is clearly needed.
It is important that we should not leave ethnobo-
tany in its present unorganized state of dispersed
data throughout monographs which are actually
in content anthropological, botanical, geograph-
ical, linguistical, or agricultural. For example,
a great deal of plant lore is included in folklore
and written literature of Southeast Asia but was
nevertheless not put there for the purpose of
transmitting the oral traditions pertaining to
plants; it includes scattered references to plant
uses compiled by casual travelers, biologists,
linguists, and others. Often, this material is not
the sort that is recovered by Occidental scholars
but, if found, is more likely to be assembled by
native scholars. One objective of Thai ethno-
botanists might reasonably be to round out a
regional monograph on ethnobotany that would
stress man's relationship (traditional) to the
environment through trial and error.
2. A matter that is frequently neglected in
making dictionaries is adequate attention to
primitive gardening and agriculture. Although
there are dictionaries of primitive tribes which
have valuable ethnographic content, there are
many languages that have no records of plant
names, implements used in gardening, and so
forth. A comparative study of gardening in
Eastern Asia has not been made, but such a
monograph would be extremely useful.
3. Because of its inter-disciplinary nature,
ethnobotanical study is frequently pursued by
persons with a variety of training, with a result
that many different field methods are employed,
and the data are not presented in any uniform
manner. The effective worker should be enough
of a botanist to make careful and systematic
collections for identification of the plants and
precise meaning of the words. Nothing is more
frequently commented upon by botanists than
that the materials collected for identification by
non-botanists are too often hopelessly inadequate.
Every contributor to ethnobotany, whether he be
a linguist, anthropologist, or geographer, should
become enough of a botanist to discriminate
critically between related plants and to have at
least as discerning an eye as one who is frequently
referred to somewhat condescendingly as a
tribesman. The ethnobotanical worker cannot
always be a linguist, but he should equip himself
with at least the basic knowledge of approved
linguistic field recording techniques and train
himself to distinguish at least the major sound
units of the language with which he is working.
He should of course attempt to fit his botanical
data into a cultural context, which presupposes
at least a basic knowledge of the culture he is
25
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
entering and a strong curiosity about the part
*vhich plants play in this culture.
4. International cooperation is indispensable.
\uthorities are not numerous for most plant
groups; linguistic information for many tribes is
;carce. The field worker is faced with the dis-
:ouraging task of either identifying everything
limself with the help of plant keys, which are
lard to find, or else seeking out the appropriate
jpecialists and requesting their cooperation.
It would be of inestimable service to field workers
f rosters of specialists could be compiled for
^ach major region in the Pacific Basin and for
Southeast Asia. This is a service that could be
Derformed by the ethnobotanists themselves.
5. The program in ethnobotany of the Pacific
Science Congress might best be used as a basis
or indicating what might be done by scholars in
5iam and elsewhere, who themselves may not
lave an adequate means of securing identifications
)f all materials but who because of their linguistic
ibilities with vernacular names are in a strategic
position to cooperate with international agencies.
\s a step in this direction, a list of suggested
esearch, compiled by Professor H.H. Bartlett
U.S.A.), is appended.
6. One phase of cthnobotanical endeavor that
las received but little attention in Southeast Asia
s the securing of chronological correlations with
)lant materials preserved at or near archaeolog-
cal sites. Although pollen analysis might not
)e feasable in some parts of the tropics, because
>f poor preservation of the pollen, it nevertheless
ihould be attempted. Dendrochronology may
ilso prove practical in some areas. Ethnobot-
inists should be encouraged to work with the
dentification of plant materials from archaeolog-
cal sites and seek comparative ethnographic
:orrelations for their material. They can and
hould contribute to the rapidly growing collec-
ions of plant materials for radiocarbon dating
>f cultural and biological events, and especially
hey could be of service in insuring that the proper
>recautions are taken during collection of the
naterials so that contamination from biological
ources is avoided.
APPENDIX
A List of Suggested Ethnobotanical Research
or Southeast Asia Compiled by Professor H.H.
3artlett, University of Michigan, Ann Arbor,
Michigan, U.S.A.
1. Listing of plants mentioned in old Siamese
16
vernacular literature, with notes on identity,
utility, ceremonial importance, etc.
2. Listing of plants mentioned in old Siamese
translations of Indie Buddhist literature, etc.,
with parallel listing of Siamese and Tndic (San-
skrit, Pali, Prakrit) plant names.
3. Notes applying to specific geographic locali-
ties on the utilization by the Siamese of local
plants of all sorts.
4. Descriptions, for specific localities, of
ancient traditional Siamese horticulture, village
gardening, industrial preparation, marketing and
utilization of plant products, especially indigo
and other dye plants, palm sugar, starch, glue,
tannin, etc.
5. Descriptions, with local names, and utili-
zation of the cultivated or wild varieties of any of
the following: bamboo, rattan, upland rice, yam,
other root crops, coconut, pot herbs ("greens"),
spices, narcotic plants (for example, betel nut
palm; betel pepper), fruit trees, etc.
6. Materials or house-building botanical
source, geographic origin, mode of transpor-
tation, etc.
7. Plants traditionally used in medicine, and
the lore concerning them.
8. History of the ancient pharmacopoeia.
9. Plant poisons and the lore concerning them,
with reference to vernacular literature. (Control
of insect pests, rodents, rice birds, predatory
animals, etc.)
10. Descriptions of primitive agriculture of any
localized ethnic or linguistic minority groups, and
of their village gardening, with notes on specific
plants, vernacular names, utilization, etc.
11. Description of sacred trees and groves, and
botanical identification of the species concerned.
12. Ceremonial uses of plants. Use of plants in
religious rituals.
13. Traditional ornamental gardening in specific
localities.
14. Materials used in basketry.
1 5. Textile plants ; history, traditional utilization.
16. Utilization of seaweeds for food or other
purposes.
17. The plant lore of weeds and wild plants.
18. Effect of population impact on remaining
forest areas in Siam; extent of man-made grass-
lands; utilization of same for grazing, etc.
19. History and method of use of lontar palm
leaves as writing material in Thailand, illustrated
by lontar manuscripts.
VOLUME 4
BOTANY
20. Ancient methods of making indigo (and
other dyes) with information on the plants used.
21. The geography, mode of growth, harvesting,
and utilization of floating rice in deep flood-waters
of Siam.
22. The sources and preparation of spices and
perfume used by local tribes.
23. The varieties, cultivation and utilization of
Pandanus in Thailand.
24. The process of manufacturing sugar from
the sugar palm (or other source).
25. The enrichment of abandoned dry rice lands
with useful woody plants by aboriginal tribes.
26. The useful plants of the Semang (or other
people) of Thailand.
27. Sacred forests in Thailand.
28. Sacred trees in Thailand.
29. The use of fungi as food in Thailand.
30. Food plants of the silkworm in Thailand.
31. The identification and use offish-poisoning
plants in Thailand.
32. Native systems of classifying plants.
33. Plants used as sources of drinking water.
34. Plants producing gums, resin, and rubber,
and methods of utilization of the products.
35. Primitive methods of making fermented
drinks.
36. Plants used as emergency and famine foods.
37. Village horticulture.
38. Plants used for thatch.
27
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
REPORT OF THE SUBCOMMITTEE ON PACIFIC PLANT AREAS
Compiled from letter by H.J. Lam of Rijksherbarium, Leiden, Netherlands.
Since the last Pacific Science Congress in 1953,
the chairman of the subcommittee has been
unable to devote a significant amount of time to
the project of preparation of distribution maps
which was the task of this subcommittee.
The Philippine Journal of Science has offered
to publish the maps and accompanying text, if
it were carried to a point where this is indicated.
The main problem outstanding seems to be that
of securing funds to employ a competent bota-
nist to do the actual work of compiling the maps.
Prof. Dr. S. Bloembergen, formerly of Java, is
known to be available for this task, if funds, to
the extent of $1,000 per year for half time, or
$3,500 per year full time, can be made available.
Office space and facilities in the Rijksherba-
rium, as well as the close cooperation of Dr. Lam,
will be made available.
It is urged that efforts be made to find the
necessary funds to carry this project forward.
28
VOLUME 4
BOTANY
REPORT OF THE SUBCOMMITTEE ON NATURE PROTECTION
C. SKOTTSBERG
Gothenberg, Sweden.
During the Seventh Pacific Congress, Auckland,
N.Z., 1949, 1 resigned as Secretary to the "Stand-
ing Committee for the Protection of Nature in
and around the Pacific" and was succeeded by
Dr. F.R. Fosberg. During the Eighth Congress,
Manila, 1953, the Standing Committee for Pacific
Botany was organized with Dr. Fosberg as Chair-
man and Mr. Kasin Suvatabundhu as Secretary,
while I was to take charge of a Subcommittee on
Nature Protection. Of the former local represen-
tatives few could be counted on any longer so
that new members had to be found.
JAPAN
As was pointed out on former occasions, the
Imperial Japanese Government has long been
actively protecting the natural resources of Japan,
and there has been no reason for the Pacific
Congresses to make recommendations. Through
the Japan Nature Protection Society, the general
public is engaged in the movement, and among
the directors and trustees are many representa-
tives of cooperative bodies such as the National
Parks Association, the Ornithological Society,
the Forestry Association, Landscape Society,
Alpine Club, etc.
PHILIPPINE ISLANDS
(DR. QUISUMBING in letter to
PROF. VAN STEENIS, Dec. 10, 1954).
"There are three categories that could be in-
cluded in the matter of conservation, areas or
localities, rare species and interesting species.
The matter... is really a very serious one... . One
of the greatest difficulties that we are encountering
in the Philippines is the matter of kaingin (shifting
agriculture) and also the vast cutting for the lumber
industry. Many of the areas... cut for timber
have been declared agricultural lands, to place
people from congested areas... . Doubtless in
this project many species are liable to be destroyed
and perhaps disappear for good. A national
committee has been appointed by the President of
the Republic to reserve certain forests... and we
have many reservations now. The difficulty ...
is that cutting for kaingin purposes continues and
it is very difficult to police these areas for lack
of forest guards and on account of the fact that
these people in the remote barrios need a place
where they can expand and cultivate crops even
temporarily."
Conservation of areas. "In Mindanao we have
a sizeable area covered by Lansium domesticum.
The Japanese had killed some of these trees during
the occupation and some of the natives in that
particular area went as far as killing trees to
collect the fruits... . I have indicated to our autho-
rities that the area should be preserved. In another
area in the province of Zambales, Vanda Lamella-
ta var. boxalli is very common on trees. I have
seen this place before devastation... with the trees
covered by this orchid in flower. As most of
these areas are private lands we find difficulties in
having them preserved. Even if the government
would expropriate the land, it is difficult to con-
trol the vandals; the orchids are collected in bud
and brought to Manila by peddlers. There is also
a project now of conserving strips of forests in the
central portion of the island of Leyte.
I am going to submit a list of plant species
included in the law for conservation, but as my
files have been destroyed I shall look into the
original law to get the names. One is Lilium
philippinense, which was very common in the
grassy area in the pine belt of mountain Province.
In early days it was indeed a sight to see this
lily when in bloom. The Igorots have been
gathering plants and brought to Baguio for sale.
...Another species in the list for conservation is
Phoenix hanceana var. philippinensis which is
found only on the island of Pabtan. The main
species is reported from Southeastern China,
Thailand, and Formosa."
INDOMALAYA
(according to letters to PROF. VAN STEENIS).
Malaya. J. Wyatt-Smith, Forest Botanist,
Kepong. Oct. 11, 1954.
"The position as I see it is that it is impossible
with the present botanical knowledge of this
region to consider reservation for the preservation
of isolated species or endemics except in very
special circumstances. Future collections, explora-
tion and taxonomic studies may well reveal the
occurrence in abundance of a hitherto "rare"
species and endemics may no longer be endemics
29
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
through being found in other countries or by
being reduced in taxonomic studies. Further,
if I am not committing botanical heresy, is
it so vital to preserve isolated endemic species
in a genus unless there is something very special
about them which is of interest phylogenetically ?
Endemic genera and single representatives in a
country or outliers of a genus by all means."
"I am wholeheartedly in support of preserving
large areas of all forest types throughout the
country, and thereby preserving complete asso-
ciations together with the rest of the plants and
animals, etc., making up the synusia. With this
in mind several Jungle Reserves have already
been constituted in Malaya and many more are
under discussion (see Malayan Forester, Vol.
XIII, 1950, pp. 92-94, and Malayan Annual
Reports), and steps are also being taken to
preserve certain areas such as Jugra Hill which is
the only place in the Federation where Shorea
gratissima is found."
"As regards preservation of 'existing isolated
rare' species..., the Malayan Nature Society is a
fairly influential body and preservation and
conservation has already come under discussion.
I am Secretary of the Selangor branch and closely
connected with the main committee... so I am in
some sort of position to push anything you con-
sider desirable in this connection."
Singapore Island. J.W. Purseglove, Director,
Botanic Gardens, Singapore.
"I am in complete agreement that everything
possible should be done to preserve plant species
which are in danger of extinction and 1 will do
all 1 can to help in this matter. The total area of
Nature Reserves in Singapore is approximately
8,500 acres controlled by a Nature Reserve Board
of Management of which I am ex-officio Chair-
man. The area reserved is a noteworthy achieve-
ment of a small colony of some 224 sq. miles with
a population of over 1 million. T doubt if we
can hope for any further reservation on this
island."
Further information on Singapore Island is
given in a circular "Nature conservation on
Singapore Island and in particular the threat to
the Pandan Nature Reserve", signed by H.
Burkill, Chairman, Nature Reserve Board, Botan-
ic Gardens, Singapore, 1957. The circular gives
the history of the forest reserves, which are
characterized as follows.
"Each of Singapore's Reserves, besides the
general purposes of recreation and education,
was created for a particular biological reason.
30
Rare plants exist in all of them. Bukit Timah
is the classical collecting area for a very large
proportion of Malaya's flora. It is the type
locality for probably more species of plants than
any other area of comparable size in the world.
It is the only remaining piece of primary lowland
forest on Singapore Island. Kranje typifies the
transition from muddy fore-shore to a dry land
vegetation, whereas Pandan is typical tidal
mangrove. These two form a series in the land-
building character of mangrove vegetation. With
the total exploitation of other mangrove areas in
South Johore, Pandan is the last remaining piece
of typical mangrove. It is moreover the most
southerly of continental Asia. Labrador conserves
a primitive species of fern, Dipteris conjugata,
which is found elsewhere in Malaya only above
3,000 ft. altitude. Pandan is the type locality
for many marine animals and all the Reserves
and the Water Catchment Area are sanctuaries
for wild animals."
Of great general interest is Mr. Burkill's answer
to the proposals to utilize Pandan:
"Proposals have been made to clear the whole
of the Pandan Reserve for the establishment
of prawn ponds. This is the second attempt to
remove control of the area from the Nature
Reserves Board for commercial exploitation,
the previous one in 1956 being successfully
resisted. It is not the purpose of these notes to
set out the arguments for these proposals. Suffice
it to say that the Nature Reserves Board is un-
willing to cede the area since it is a legally con-
stituted body with a duty to perform."
The following general remarks regarding
Malaya may also be quoted from Mr. Purse-
glove's letter to Dr. van Steenis (see above).
"With the present state of collecting within
the Malaysian region, are we justified in assum-
ing that a plant is in danger of extinction because
it has only been found in one small area? Until
much more collecting has been done and the
flora and its distribution known more fully than
at present, the possibility of rare and interesting
species or species at present only recorded from
one small locality turning up in an area at present
little collected would seem to be considerable.
1 think that the most we can do at the moment
is to list those species which reputable botanists
consider in danger of extinction and do what
we can to preserve them, the method of preserva-
tion being dependent upon local conditions."
"Unless a sufficiently large area is preserved
ecological conditions will be altered which might
VOLUME 4
BOTANY
result in the loss of species. In secondary com-
munities reservation may result in a change in
vegetation with resulting loss, especially when a
plant may depend for its continuance on biotic
factors. It would be difficult to look after a
large number of small areas and see that they are
adequately patrolled, even if money and staff
are available. Making a small reserve without
adequate patrol and advertising the presence of
the species might lead to its more rapid extinction.
The possibility of preserving the plant in cul-
tivation might be considered."
INDONESIA
In addition to the late Dr. K.W. Dammerman's
beautifully illustrated work, "Preservation of
Wild Life and Nature Reserves in the Nether-
lands-Indies," published for the Fourth Pacific
Science Congress in 1929, comprehensive reports
appear in the Proceedings of the Fifth Congress,
Vol I, 1933, but after that no further report was
received until 1949, when a few remarks by Dr. A.
Hoogerwerf were included in the Proceedings,
Vol. IV, 1953, referring to the work done by the
Netherlands Indies Association for Nature Protec-
tion which issued annual reports.
It would have been of particular interest to
obtain official statements as to what happened
during the Japanese occupation and the following
insurrection which ultimately led to the establish-
ment of the Republic of Indonesia. It is, however,
good to learn from a letter to Dr. van Steenis,
written Oct. 25, 1954 by the Director of Kebun
Raya Indonesia (formerly Buitenzorg Botanical
Garden), Prof. I. Kusnoto, that the efforts made
by his predecessors to convince the government
of the necessity of plant protection by law will
be renewed, and that everything will be done in
order to intensify plant conservation. A restric-
tion on the export of rare species, among them
orchids, is planned, as well as the protection of
the localities for rare species, by setting aside such
areas as reserves. For detailed information the
departmental Head of the Herb. Bogoriense and
the Chief of the Botany Division of the Forest
Research Station should be approached.
Sumatra. Dr. W. Meijer recently published a
paper on "Natuurbescherming in Midden-Su-
matra" (Natuur en Landschap 101 , 1956), contain-
ing an interesting reconnaissance of protected
natural monuments such as Harankloof, a low-
land virgin forest at Pantji conserved since 1930
and locality for Rafflesia Arnoldi, Aneikloof,
type locality for Amorphophallus titanum, etc.
The forest reserve on Mount Sago, 2,000 meters,
is now being investigated scientifically. An area
of 10 hectares, 1,000 m. above sea level, included
150 different kinds of trees, almost equalling
Tjibodas on Java in richness. Nature conser-
vation in Sumatra is an enormous task, but
wholeheartedly supported by the Governor and
the Director of the Forest Service. The immensely
interesting extinct volcano Mt. Singgalang has
recently been proposed as a forest reserve.
BRITISH NORTH BORNEO
(The late G.H.S. WOOD, Forest Botanist,
in letter to DR. VAN STHENIS, Oct. 5, 1954).
"Regarding the preservation of species and
ecological types I am following Malayan practice.
This is primarily the making within forest reserves
of virgin jungle reserves of 50 to 100 acres which
will not be interfered with in any way. Con-
stituted forest reserves at present occupy only
3% of the total land area of North Borneo, but
it is anticipated that large areas now being worked
as concessions will become forest reserves. No
jungle reserves have so far been made but it is
expected that creation of such reserves will be
made obligatory on concession areas and con-
stituted reserves. The object is to keep samples
of ecological and physiognomic types rather
than particular species undisturbed."
"Forest reserves are also being made in forest
stands which are not necessarily economic (or for-
seeablyso); hence numerous species are being
preserved intentionally or not. An example of
this kind is Tabawan Island in Darvel Bay, which
I recently visited with Mr. Wyatt-Smith. This
island supports a peculiar forest type dominated
by Casuarina sumatrana, Calophyllum spp. and
an unidentified tree, known locally as bangkan;
only three scarce Dipterocarpaceae are recorded
so far."
"You will appreciate that in general forests
on the flat land of the east coast of North Borneo
are undisturbed and offer considerable scope for
retention as forest reserves containing selected
virgin jungle reserves. On the west coast, both
flat land and mountainous slopes up to 3,500 ft.
have suffered severely from either permanent or
shifting cultivation. Undoubtedly some species
have already become extinct and more will follow,
and little can be done to protect individual species.
But priority is given to the formation of forest
reserves on the west coast before the remaining
virgin stands, particularly at low altitudes, are
destroyed; hence the great majority of species
31
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
will survive apart from those which are very
local."
"It is expected that it will be easy to reserve
areas over 3,500 ft. on mountains on the west
coast. A Land Utilization Committee is planned
for North Borneo and I shall propose that the
whole of Mt. Kinabalu-Mt. Tambuyukon area
and the Mt. Trus Madi area should be reserved
as national parks (game and forest reserves) quite
apart from the formation of other forest reserves
in the mountains. Proposals have already been
made for a national park in the Ulu Segama area.
On the other hand little can be done on the
flat coastal fringe of the west coast which is now
relatively densely populated. In this area few
untouched samples of lowland forest remain on
well-drained ground, and existing swamp forest
(either in forest reserves or outside) is threatened
with conversion to padi land. Generally speak-
ing we have so little information on the distribu-
tion of rare or local species that even if it were
economically feasible to make reserves for them
one would not know where to make them. In
the case of tree species it is easier, for there is
justification in preserving them as part of a partic-
ular forest type and very often this means that
the rest of the distinctive associated synusium of
lianes, under-storey trees, and shrubs is also pre-
served. In this connection you may be interested
to know that most limestone hills on the east
coast (with their economic importance as pro-
ducers of birds' nests, which are controlled by the
Forest Department) are being included within
forest reserves. As we become acquainted with
other ecological types I earnestly hope that they
can be similarly treated, but no one can deny
that large areas at present under forest in North
Borneo must ultimately be lost to permanent
agricultural land as the population increases.
Only careful choice of forest reserves can preserve
most of the species."
(August 3, 1955.) "It has occurred to me that
the project suggested in your letter of March
8, 1955, might be easily extended by enlisting the
aid of either FAO or UNESCO, the former
being particularly active in Malaysia. I know the
idea is not new but some sort of census or survey
of existing or potential areas of purely scientific
value would be of great and permanent value.
To start with, some idea of the extent of such
areas might be obtained, followed by detailed
surveys with estimation of animal and plant
populations. Being of primarily scientific turn
of mind, I am particularly interested in such a
project and I am sure the aid of many scientific
32
bodies could be brought to bear on the preserva-
tion of species which would otherwise be liable
to extinction."
A few days later (August 9, 1955), Mr. Wood
sent a circular letter to seven timber trading com-
panies of North Borneo.
"Gentlemen,
Reservatjon of virgin junglej>lots.
This matter was raised at a meeting with repre-
sentatives of the main timber firms in Sandakan
on 6.8.55. A number of Forest Reserves have
been constituted, which help the preservation of
much of the native flora and fauna of North
Borneo, but it is desirable to make as many of
these as possible in widely scattered areas even
if they are small. In concessions concessionaires
are asked to provide details of any areas which
they do not wish to exploit (e.g. inaccessible
areas on hills, swampy areas, etc.). These in-
accessible areas are often the home of rare plants.
On his recent tour Professor H.G. Champion
said that such virgin jungle reserves should not
be less than 50 acres, with a substantial surround;
further details can be found in Chapter XIV
of the North Borneo Forest Manual. Naturally
reserved areas of this sort are just as desirable in
forest which is commercial and provide a yard-
stick for regeneration procedure in adjacent
exploited areas. The reservation of virgin
forest in North Borneo will allow the preservation
of most native species, especially in the national
parks on Mt. Kinabalu and in Ulu Segama, but
additional widely scattered small areas of virgin
jungle left in concession are of great value."
"Please send lists of any areas of this sort which
you wish to abandon under the terms of your
agreement."
(November 26, 1955.) "Further to my letter of
3.8.55... another point has occurred to me re-
garding the conservation of rare species and veg-
etation types. You may be aware that present day
silvicultural treatment in forest reserves through-
out the tropical belt of the Commonwealth are
tending more and more to the elimination of
weed trees by poisoning. From a scientific point
of view this is serious, because many of the weed
trees include rare species which one wishes to
protect. If forest is logged but not silviculturally
treated, there seems to be little reason why such
species should not persist, but systematic poison-
ing may eliminate them in the long run."
VOLUME 4
BOTANY
"I have a feeling that large climbers would be
more affected than under-storey trees, for not
only are they broken when large commercial
trees are felled, but they have to be ruthlessly
poisoned if they are not to inhibit the young
commercial saplings of the new crop. If this is
the case one can foresee a time when forest
reserves (other than protection and ecological
reserves) would be largely composed of com-
mercial trees alone."
"I may be wrong in my assessment of the recu-
perative powers of the under-storey trees, but
quite frequently one meets examples of species
which are so rare in the restricted localities in
which they do occur that I hesitate to predict
their future in the event of intensive silvicultural
treatment."
"It strikes me that this problem is much worse
in Malaysia than in Africa, for here we have so
many endemic species of limited range."
SARAWAK
(MISS WINIFRED BROOKE,
c/o The Museum, Kuching, in letter to
DR. VAN STEENIS, October 13, 1954).
"1 have consulted Mr. F.G. Brown, B. Sc.,
Conservator of Forests here, and he has given me
information regarding reserves."
"At present very little is known regarding
species and their localities in this country and
there is little danger of exterminating any of them.
Rafflesia can only be protected by keeping the
exact localities concealed and by education regard-
ing the imaginary character of its advantages as
a medicine in childbirth (suppose they are imag-
inary). It grows in a forest reserve. About
one-fifth of this country consists of forest reserves
where normally no trees can be felled although
the department may destroy useless species. It
could no doubt give and enforce, to a reasonable
extent, orders for the protection of any species
of tree or plant found peculiar to a reserve and
not as in the case of Rafflesia of a special value
to the local people."
"Several species of Conifers grow on the sum-
mit of Mt. Poi and according to Beccari on that
of Matang, and he mentions other interesting
species on Matang. Mt. Poi is a forest reserve
and Matang is kept very largely in its natural
state by the Water Board as Kuching obtains
her supplies from there."
"Hunting and the collection of forest products,
gums, fruits, etc. is permitted in the reserves; and
the fauna, especially the birds in them and else-
where, are in considerable need of protection."
"The Governor of Sarawak, Sir Anthony Abell,
is encouraging a project to make Santubong
Peninsula a national park, also a small lake in
the north of the country. The matter is, I believe,
well under way."
"When and if we have real information regard-
ing a rare species with a limited range I consider
that now is the time to arrange to protect it
because at present it would be no hardship to
anyone to avoid felling, digging up or building
on a particular strip of ground; others equally
suitable abound except possibly for air-strips;
later on, if the country develops, it might seriously
interfere with work or planning to protect some
small locality near an industrial or agricultural
project."
NETHERLANDS NEW GUINEA
(F. RAPPARD, Head of Forestry Service,
Hollandia-Haven, in letter to DR. VAN STEENIS,
October 26, 1954). Summary in English.
"1 have discussed your letter on 'protection of
rare plants in Netherlands New Guinea' with (my
assistant) Versteegh and (Professor) Lam. We
haven't at present the slightest idea what are
rare species, nor shall we know for a couple of
decades, and if we try to do something it will
remain a useless project. The only way to get
some results will be to create some reserves
where lumbering is forbidden, but such areas
will be in the high country. Proposals have
already been made, also for establishing one
coastal reserve."
TERRITORY OF PAPUA AND
NEW GUINEA
(j.s. WOMERSLEY, Chief of Division of
Botany, Dept. of Forests, Lae, in letter to
DR. VAN STEhNIS, August 28, 1956).
"Firstly, I would agree with you that the funda-
mental definition of 'rare' in terms of plant species,
at least as far as New Guinea is concerned, will
prove exceedingly difficult to determine for the
practical work of this Committee (C. of Pacific
Botany). In fact, 1 would hesitate to say that
anyone species in New Guinea today is so rare
that a real possibility of its extinction exists.
As an example of this I could quote Dendrobium
johnsonii, which immediately after the war was
believed to be an exceedingly rare orchid which
was faced with extinction due to considerable
export of plants made immediately prior to 1940.
33
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
However, further investigation has shown that
this orchid is as common as any almost through-
out the whole island within its own altitudinal
range."
"I am more concerned at this stage, at least
for New Guinea, in the establishment of national
parks of considerable size which could provide
refuges against the pressing tide of agricultural
development for the unique flora and fauna.
The same problem has concerned the Animal
Ecologist, Mr. K. Slater, in considering how the
rare species of Birds of paradise may be protected
against agricultural development of the land.
Of course the very terrain of New Guinea ensures
the protection of the flora and fauna although
serious disturbances can be caused even to high
mountain forest if agricultural development
interferes with water-sheds and streams. "
'There is also, at least in the Trust Territory,
the problem of land ownership which is quite
clearly vested with the indigenous people unless
land is (a) purchased by the Administration, or
(b) declared waste and ownerless. Virtually all
land purchases to date have been designed to
provide agricultural or residential land and only
very few areas have been proclaimed waste and
ownerless. This position appears to me to ensure,
at least for several generations, that much of the
existing forested areas quite unsuitcd to native
agriculture will remain more or less in their
present state."
"However, I cannot concede that this removes
the necessity to find some means of proclaiming
flora and fauna reserves to protect the wild life
of the Territory."
"In answering your specific questions I feel the
project is very real particularly in those countries
of Malaysia where the population density is so
great that there is real danger of rare species
becoming extinct."
"To your second question I shall be happy to
join you as a collaborator with the proviso that
at this stage and with our relatively limited knowl-
edge of the distribution of plants in New Guinea,
I would not be prepared to assert that any partic-
ular species was so rare that it needed protection.
We already have embodied in our Customs Ordi-
nance adequate power to limit the commercial
exploitation of our indigenous flora, e.g. by collect-
ing of live orchids. Specimens of flora, fauna and
minerals are prohibited exports except by consent
of the administrator or his nominated delegate."
34
WESTERN AUSTRALIA
(MR. c. A. GARDNER, in letter to DR. COSTIN.)
Regarding plants threatened with extinction.
"1 think that we can safely say that a number
of our species are threatened with extinction in
the near future, and that a number are already
extinct. Such arc plants endemic to a small
area. I very much doubt that the following will
be seen again:
Philotheca ericoides (Harv.) F. Muell
Darwin ia carnea C.A. Gardn.
Cryptandra eriantha Diels.
Leschenaultia hirsuta F. Muell
Calcadenia Drummondii Benth.
Species threatened with extinction in the near
future include
Marianthus ringens (Drum & Harv.) F.M.
Casuarina fibrosa C.A. Gardn.
Boronia capitata Benth.
Hemigenia viscida S. Moore
Asterolasia grandiflora (Hook.) Benth.
squamuligera (Hook.) Benth.
Dryandra vestita (Ripp.) Meisn.
speciosa Meisn.
The above are a few that come to mind. I
will keep your request in mind, and add to this
from time to time. With large scale operations
in agricultural development, it is inevitable that
many of our localized endemics must disappear."
EASTERN AUSTRALIA
(J.H. WILLIS.)
The following lignose species may be in danger
of disappearing:
Acacia peuce F. Muell
Choristemon humilis H.B. Williamson
Eucalyptus serrulata Blakely and Beuzeville, a
unique type in the genus Eucalyptus
NEW ZEALAND
(MISS L.B. MOORE.)
Three Kings Islands.
List of species in danger, according to obser-
vations in 1951 by Mr. Baylis:
Alectryon grandis Cheesem. (Sapindaceae, 3 or
4 adult trees)
VOLUME 4
BOTANY
Braehyglottis arboresccns W.R.B. Oliver (Com-
positae)
Elingamita Johnsonii Baylis (perhaps a dozen
trees)
Hebe insularis (Cheesem.) Ckn. & Allan (Scro-
phulariaceae, recorded as spreading)
Paratrophis Smithii Cheesem. (Moraceae, re-
corded as spreading)
Pittosporum Fairchildii Cheesem. (recorded as
spreading)
Pleetomirtha baylisiana W.R.B. Oliver (Anacar-
diaceae, only the type tree known)
Rapanea dentata W.R.B. Oliver (Myrsinaceae,
perhaps a dozen trees)
Tecomantha speciosa W.R.B. Oliver (Bigno-
niaceae, only the type plant known)
The reason for recovery was the removal of the
goats.
POLYNESIA AND MICRONESIA
(from F. R. FOSBfcRG.)
Island floras are peculiarly vulnerable to des-
tructive forces of many sorts, as the areas occu-
pied by species are frequently very restricted and
populations are often very small. As has been
many times reiterated, most endemic island
species of plants are in danger of extinction if
present tendencies toward disturbance and des-
truction of island habitats are not checked.
In general the actual situation seems to con-
tinue to deteriorate throughout the insular world,
judging by what personal observations we have
been able to make and from reports of others,
mostly from small indications that would be
useless to detail here. There are a few brighter
spots in the general scene. Major gains and losses
have been listed in a report on the conservation
situation for Oceania included in the report of
the Standing Committee for Pacific Conservation,
given elsewhere in the Congress and need not
be repeated here.
Work progresses slowly on the preparation of
a file of information on the most immediately
threatened species in Oceania which could serve
as a guide for immediate action to save some of
these. Reliable and up-to-date information is
not easy to obtain, however. Contributions of
such information are earnestly solicited.
CALIFORNIA
(J. TH. HOWELL.)
"The distribution of the plants considered in
this report is that given by Jepson in his Manual
of the Flowering Plants of California. As a
by-product of the analysis of that work for the
present report, I published a Tabulation of Cali-
fornia endemics' (Leaflets West. Bot. 7, 1955), in
which I discuss matters relating to this important
and distinctive feature of the California flora.
Genera endemic in California.
(a) Known to occur in a State or National
Park:
m Scquoiadendron Buchholz (Taxodiaceae)
Sedella Britt. & Rose (Crassulaceac)
m Draperia Torrey (Hydrophyllaceae)
m Phalaeroseris Gray (Compositae)
m Orochaenadis Cov. (Compositae)
m Gilmania Cov. (Polygonaceae)
Oreonana Jepson (Umbelliferae)
m Whitneya Gray (Compositae)
m Holozonia Greene (Compositae)
To these will perhaps have to be added the
following which probably can be found in a pre-
serve :
m Heterogaura Rothrock (Onagraceae)
Pseudohahia Rydb. (Compositae)
(b) Not known in a Park or other Preserve,
"National Forests" not considered:
m Etosperma Swallen (Gramineae)
m Odontostomum Torr. (Liliaceae)
m Carpenteria Torr. (Saxifragaceae)
Acanthomintha Gray (Labiatae)
m Holocarpha Greene (Compositae)
m Crockeria Greene (Compositae)
m Tracyina Blake (Compositae)
m Neostapfia Davy (Gramineae)
m Hollisteria S. Wats. (Polygonaceae)
m Lyonothamnus Gray (Rosaceae)
Monolopia DC. (Compositae)
m Blepharozonia Greene (Compositae)
m Eastwoodia Brandagee (Compositae)
Genera preceded by an m are monotypical.
The following trees, endemic in California,
are found in a State or National Park:
Abies magnified Murr
Cupressus macrocarpa Hartw.
Sargentii Jepson
Pinus balfouriana Jeffrey
,, monophylla Torrey
muricata Don.
ponder osa Dougl. var. Jeffreyi Vasey
sabiniana Dougl.
35
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Finns torreyana Parry
tuberculata Gord.
Scquoiadendron gigantewn (Ldl.) Buchholz
Torreya californica Torr.
Lithocarpus densiflora (Hook.&Arn.) Rehd.
Quercus chrysolepis Liebm.
Douglasii Hook. & Arn.
lobata Nee
morehus Kellogg
Platanus racemosa Nutt.
Aesculus californica (Spach) Nutt.
Fraxinus dipclala Hook. & Arn.
These are not found in Parks or other
Preserves :
Abies bracteata (Dougl.) Nutt.
Cupressus Forbesii Jepson
go vcniana G ord .
inacnahiana M u r r .
var. Baker i Jepson
nevadensis Abrams
pygmaea Sargent
Sargent ii var. Duttonii Jeps.
Pinm contorta Dougl. v. Bolanderi Vasey
,, Coulteri Don.
Pseudotsuga macrocarpa Mayr.
Taxus rev if olid Nutt.
Quercus dumosa Nutt. var. Macdonaldii Jeps.
Wislizenii A. DC.
Jug Ian s californica Wats.
Hinds ii Jepson
Lyonothamnus calif or nicus Gray
Acer negundo L. var. californicum Sargent
The California Floral Province, extending from
southern Oregon to northern Baja California,
with exclusion of considerble areas in the eastern
part of the State related floristically to the Colum-
bia Plateau, Great Basin or Sonoran Desert, was
proposed by me in 1957 (Leaflets West. Bot. 8:5).
The number of endemic genera is 65."
MAINLAND OF CHILE
National Park of Talinay, Prov. of Coquimbo.
Visited on April 30, 1955, by Dr. and Mrs.
Skottsberg, accompanied by Mr. Carlos Jiles,
a resident of Ovalle. The forest was found in
primeval condition and in no way threatened ; it is,
in fact, in better condition than the nearby quite
similar Frai Jorge forest north of the mouth of
Limari River and has been much less visited than
this. The latter has been well surveyed by Messrs.
Munoz and Pisano, and before by Dr. and Mrs.
Skottsberg (publication in Acta Horti Gotoburg
36
18, 1950). A corresponding survey of Talinay
ought to be undertaken. The composition is the
same in the two places, but Cerro Talinay is about
30 m. higher (700 m.) and makes an impression
of being even more humid; both are enveloped
in fog and the vegetation drips with water. Nei-
ther is of difficult access; in the case of Talinay
a motor road leads to the foot of the mountain,
and from there the ascent is made on foot through
dense chaparral, where, in places, Puya chilensis
is dominant. The forests are famous as consist-
ing of a highly hygrophilous community, sur-
rounded on all sides by xerophytic vegetation and
separated from the south Chilean rain forests
by many degrees of latitude. The great Pan-
american Highway will pass one km. from Tali-
nay, and it is expected that the peculiar forest will
be of great touristic value. It should be remem-
bered that the soil cover of delicate ombrophilous
herbs and bryophytes, where the foot sinks
deep, cannot bear any traffic at all without
getting damaged and that the extreme rain forest
type with lianas and epiphytes covers only a
narrow strip. The distance from the Highway
to Frai Jorge is about 20 km.
THE JUAN FERNANDEZ ISLANDS
In my Report for 1933-38 to the Sixth Pacific
Congress I told of the establishment, in 1935, of
the Juan Fernandez National Park, the first of
its kind in Chile. Even if, as little was done to
enforce the regulations, an efficient conservation
of the native vegetation still remaining wasn't
in any way ensured, nothing like the devastation
that has taken place in the last twenty years could
be expected. That much would have happened
between 1917, when I surveyed the islands, and
1935, could be foreseen; the intentional introduc-
tion to Masatierra of one of the dangerous pests
from the mainland, a European species of Rubus
(so far determined ulmifolius) had been reported,
and the renewal of Masafuera as a convict settle-
ment in 1927 ought to have left its marks. In
order to find out what had happened and to
make a comparison between 1917 and now, I
returned to the islands in December, 1954, and
spent three months there. Also this time I was
accompanied by Mrs. Skottsberg.
Masatierra. The settlement remains a fishing
village, but it had grown considerably, from
about 200 to about 600 people, which is too much,
as too many persons are not engaged in the
fishing industry. As before, the population
concentrates around Cumberland Bay, the only
VOLUME 4
BOTANY
harbor. The lack of suitable land and the exceed-
ingly broken topography forbids agriculture if
not on a miniature scale, and horticulture showed
little increase, even if small cultivations had
extended into the valleys east of the Colonial
Valley, but this had little to do with the destruc-
tion during the thirty-eight years that had passed
since our previous survey. 1 do not think that
the number of cattle, about 250, was much
greater than before; the difference was that there
was almost no pasture left, and we shall see the
reason. Sheep had been introduced and allowed
to increase without any restriction at all. The
whole island lay open to them; they had invaded
the drier forest type and the brushwood of the
high ridges; through the action of a number of
enterprising individuals the national park had
been turned into a sheep farm. The official
figures stated about 3,750 sheep, of which 3,100
grazed the lower treeless western part and the
remainder the denuded slopes of the easternmost
valley of Frances, but in addition numerous
others were observed also in other valleys and
on the high ridges so that the total number must
have exceeded 4,000. Certain areas had been
turned into a desert, and the valley systems of
Frances, Ingles, and Villagra offered a horrible
sight. Once the soil formed by the easily disin-
tegrating basalt, had been deprived of its plant
cover, native (as in Villagra) or not, erosion set in
with full force. Horses are of slight importance
as communication between the coves is so much
easier by motor boat; the old trails to Villagra
and Frances are seriously damaged and almost
dangerous; the lower part of Villagra, in 1917
covered by native grassland, had changed beyond
recognition, and the zig-zag trail across Salsi-
puedes to Ingles valley, worn deep into the rock,
had been completely effaced.
Goats, introduced in the 16th century, had
become naturalized and must have been very
plentiful once, but in the 19th century their
number had become so much reduced that they
were protected for sentimental reasons because
they descended from the goats of Alexander
Selkirk, with whom Defoe's Robinson Crusoe was
identified. In 1916-17 few were shot because
they had retreated to places of extremely difficult
access, and in 1955 they were scarce and not
seen by us. But there was good help for this;
under the pretext of providing goat's milk a
small herd of angora goats had been imported
to the colony, where they took to the mountains
at once, multiplied and spread. One day 1 counted
sixty on one of the steep ridges above the settle-
ment; being pure white, they are easily discern-
able even from a large distance, and some were
seen later on the most inaccessible precipices.
They are private property so that hunting them
is illegal. Two more additions to the fauna should
be mentioned, Nasua rufa, introduced to help
combat the big rats, and rabbits. Nasua prefers
the dense forest and is said to be scarce; it is an
excellent climber and accused of liking birds'
eggs.
It is hardly probable that, in old times, much
damage was done in the native forest after the
wild pigs had been exterminated, except where
logging was practised, until the arrival of the
terrible Chilean pest, the shrub Aristotelia
chilemis (maqui) some time before 1854. In
1917, it filled all the valley bottoms and lower
slopes above the everywhere denuded coastal
belt, penetrating the native forest and prevent-
ing, by its deep shadow, the endemic trees all
native trees are endemic from germinating, so
that their disappearance from the lower forest
belt was foreseen already by Johow in the 90's.
The maqui had continued its progress and ad-
vanced into the upper forest belt, reaching the
ridges in 500-600 m. in many places; the fruits
are dispersed by the thrush. Another Chilean
shrub, Ugni Molinae, locally known as murtilla,
introduced on purpose because of the aromatic
berries during the latter half of the 19th century,
was, in 1917, practically confined to the barren
slopes above the Colony. It had increased tre-
mendously and invaded the native shrub along
the steep ridges, the home of many of the rare
endemic genera and species of plants, some of
them now on the verge of extinction. All our
good old collecting grounds were more or less
ruined. A special list of rare and vanishing
species has been prepared.
One of the most precious and scientifically
interesting forest products, the endemic and
monotypical palm Juania australis, since long
protected by law, has almost entirely disappeared
from the accessible parts of the forest belt, but
freshly manufactured curios of it were still
offered for sale.
The Pangal Canyon, the only typical canyon
in Masatierra and a gem, will serve as an example
illustrating the changes since 1917. Toward the
sea was a fine stand of Boehmeria excelsa along
the stream, and the interior had fine native
forest with stately tree-ferns right up to the ter-
minus, a verical cliff wall with a cascade. Of all
this vegetation, nothing remains; the outer slopes
are naked, full of weeds and grazed by cattle;
the interior, a jungle of maqui; the waterfall is
reached using the tunnels made by the cows.
37
PROCrr.DINGS OI THE NINTH PACIFIC SCIENCE CONGRESS
The bramble-berry, introduced in the 20's
by a colonist living in the Anson valley, to be
used as a living fence around his orchard, had
destroyed it and spread like wild-fire up the valley;
the small open place at the foot of Mt. Yunque,
often visited by us and surrounded by fine forest,
was now changed into an impenetrable thicket.
From here it spreads by man and birds the fruit
being edible in all directions and pioneer speci-
mens florish on the high ridges above 500 m. If
nothing is done to stop it, it will invade the island;
the people complains of the "zarzamora," but
nothing has been done. It may be too late now
and will, in any case, cost a lot of money and labor,
a matter for the high authorities on the mainland.
It goes without saying that the changes also
affect the fauna. The endemic land birds are
decreasing, and the peculiar pacific land shells
will not, I presume, find the maqui a suitable
biotope. The insect life, mostly small incon-
spicuous forms, is still surprisingly rich, at least
on the higher forested ridges; numerous genera
and perhaps some 90",, of the species are endemic.
Whether fires have played any important role
I cannot tell, but I was told that in 1930, a fire
swept Ingles valley, and very likely this is true.
It is, anyhow, ruined; the pasture is gone, and
sheep and cattle left to starve to death. They
cannot get out.
Santa Clara Island, a satellite of Masatierra, has,
as far as we know, never borne any forest growth,
but was once covered by grassland sprinkled with
a few dwarf rosette trees belonging to the endemic
genera Rea and Dendroseris. All this is gone with
the exception of some specimens growing in inac-
cessible gorges. The islet is overstocked with sheep
and looks like a desert; in 1955 the sheep num-
bered about 500, but we saw no goats, once said
to be numerous "Goats Island" is an old name
for the place. The detached rock Morro del
Spartan, separated from the islet by a narrow
channel not crossed by sheep or goats, gives some
idea of what the scenery must have been in bygone
time. This morro is the type and only locality for
Chenopodium Sanctae Clarae, of which half a
dozen specimens were observed.
All visitors agree that Santa Clara lacks fresh-
water, but rain falls during the winter months
when some water must be found, otherwise
sheep could not exist. According to recent
information the owner of the sheep was going to
evacuate the island.
Masafuera. The law says that no permanent
habitation shall be set up on this island ; and when,
38
in 1930, the convict station was abandoned,
Masafuera was left uninhabited. However, in
1940, a fishing colony was established at the
entrance to Casas valley and took possession of
the government buildings. Langust fishing,
the principal industry of Juan Fernandez, is
profitable around this island, and the colony,
though living under rather primitive conditions,
was prosperous. No cultivations of any kind
exist now, there were only about thirty head of
tame cattle, a few horses, and twenty sheep, so
that the damage done was negligible. In 1917,
the marks of the first penal settlement were very
conspicuous; the accessible forest patches had
been logged and grazed; huts had been built in
the uplands; potatoes and vegetables were
grown, but the high land south of the Vacas
valley, crowned by the summit, 1,570 m. above
sea level and extremely wet, was covered by an
impenetrable fern forest and showed almost
no sign of human activity. In 1917, good forest
groves were found in the upper gorges of the
valleys all along the east side of the island the
western side is a precipice between 300 and
700 m. above sea level, the maqui was rare, and
the grassland higher up, with its extensive fields of
big ferns and bands of Gunnera along the streams
and scattered stout specimens of Dicksonia,
made the impression of a peculiar fern savanna
above 900 m. Goats were very plentiful, and many
plant species confined to places they could not
reach. And now, in 1955, this wonderful island
offered a very sad sight. The forests had been
logged and were full of maqui; the first bram-
bleberry shrubs had made their appearance. We
had difficulty in recognizing places we used to
know so well and now found utterly ruined, a
result of a series of devastating grass and bush
fires. The dates given were 1939, 1942 or 1943,
and 1944, and one of them, which swept the In-
ocentcs ridge to the highest summit of the island,
was not accidental. Another fire had run over
the Barril ridge across to the gorges on the
western side. The fern savanna was gone;
charred Dicksonia trunks told the story. Forest
groves of small size in the hanging gorges cut
into the west wall of the island are still unharmed.
The only large patch of closed wood is at the
northern end. This was visited. It is of a dry
type on the flat ridge between two valleys. In
this part, natural Stipa grassland, very monoto-
nous, covers extensive areas; otherwise the up-
land is overrun by Anthoxanthum odoratum and
Rumex acetosella, just as it was in 1917, and the
native Alpine species are scarce except close to
the edge of the west wall in 1,100-1,400 m.
VOLUME 4 BOTANY
PROBLEMS CONFRONTING BOTANICAL INSTITUTIONS
IN THE TROPICS
KUSNOTO SHTYODIWIRYO and ANWARI DILMY
Botanic Gardens, Bogor, Indonesia.
If we may compare botany to a tree, we can say
that it has its roots in Europe but that it is coming
into flower in the tropics. Most of its flowers,
however, are still in bud. To cause these buds
to open is, in brief, the task of botanical institutes
in the tropics. We mean: to realise the splendid
possibilities for scientific research provided by
the wealth of tropical forms.
With respect to the taxonomic study of a trop-
ical flora, the task of an institute in the tropics
is more complicated than that of an institute in
a temperate climate. The latter can be merely
a "working" herbarium where botanists deal with
the material that comes to their hands. But a
tropical institute not only has to elaborate
material for the purpose of compiling taxonomic
publications, it also must supply the material
itself, by collecting it on expeditions in the field,
an undertaking that may be too expensive or too
complicated for an institute in a temperate
climate.
In the course of history, every botanical insti-
tution has had its ups and downs, but this must
not divert our attention from distinguishing the
two kinds of herbaria: the "working" herbaria
and the "supplying" herbaria. An example of
a working herbarium is the Kew Herbarium.
Here botanists are preparing and publishing
revisions, but their material must come from
somewhere else. An example of a supplying
herbarium is the one at Sandakan. Here no
proper taxonomic work is performed, but material
is collected for the Kew botanists and others.
The Bogor Herbarium has always both produced
taxonomic studies and supplied material ; the same
is true for the herbaria at Singapore and Manila.
A modern taxonomic publication, especially
if the author aims at a high standard, is the result
of the cooperation of many people. Specialists
are spread all over the world; the type material
of Malaysian plants is scattered over at least
a dozen herbaria, and for good results so much
collaboration is necessary that we sometimes
wonder whether the task of the Keeper of a
modern herbarium is anything else than shipping
and inserting material and sending botanists
abroad. In any case, if we wish not to botanize
alone but to contribute to international botany,
our best policy is to show how we can be most
useful to others. This can be done by utilizing
the specific advantages of the site of our institute.
We need people who are able to collect critically
and to make notes in English to go with the
material, thus giving important information to
the herbarium taxonomist. The collector must
be prepared to take at least three duplicates a
dozen or more is still better. Further, it is ex-
tremely useful for him to have so much knowledge
of the existing forms that he can pre-identify the
material collected. At Bogor, Mr. Nedi and
Mr. Noerta perform this work. Although lacking
advanced schooling, they have accumulated exten-
sive knowledge during field work with botanists
and continued training themselves in recognizing
families and genera. They are invaluable because
in this respect they have achieved what a pro-
fessional botanist achieves only after long, long
experience. Therefore, if a man is found who
shows talent in recognizing plants, he should be
taught and encouraged, no matter what his status
is. Even a common labourer could thus become
one of the treasures of the institute.
Finally, duplicates of the material collected
must be distributed. This is by no means easy.
The material must be divided into adequate
portions and labels must be copied carefully.
All this material must be packed and shipped.
The dispatch of duplicates is not one of the minor
occupations of the Bogor Herbarium staff; we
need mention only that in 1956 we shipped a
total of seventy-four packing cases of duplicate
specimens.
A herbarium connected with an important
tropical garden has another special function to
fulfill, namely to collect and depict garden
material and to give information asked for. A
tropical garden containing many "wild" species
can provide more direct knowledge than a garden
in a temperate climate.
Because the functions of a tropical garden are,
as we have seen, quite varied, it can easily be
understood that we must prepare ourselves not
to lose our heads when we have to face difficulties
resulting from lack of funds or lack of trained
staff. The most important thing is to keep what
we have. We therefore consider the alcohol and
39
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
the corrosive sublimate that preserve our speci-
mens to be the life blood of the tropical herba-
rium. Almost equally important are the tins and
the paper that protect them against breakage
and dust.
It is similarly important to improve our
material. When material required for a revision
is sent out on loan and later returned and inserted,
the new identifications by specialists will help us
in turn to identify new specimens by comparison.
As apparent from the above, a well-trained
technical stafT is of the utmost importance. A
herbarium with only a botanist is nothing. A
herbarium with good technical staff is at least
something. This contradiction holds good
especially in the tropics.
The work of a botanist will stimulate other
activities: collecting and distributing duplicates
and gaining information from living material.
Of course he should work and publish, but he
must remain aware of the fact that an excellently
prepared and annotated plant specimen, widely
distributed, is better than a second-rate publi-
cation.
With respect to the creation of new local
institutions, it is well to recall the function peculiar
to the tropical herbarium, namely to supply
material to the great international body of
science. A library as extensive as that at Bogor
is necessary to operate a "working" herbarium,
and it must be kept up to date. But a properly
functioning "supplying" herbarium requires only
an active botanist with a few willing local men,
the necessary collecting and drying equipment,
and facilities for shipping the material.
The tree of botany has become too large to be
tended by one institute only. For efficient growth,
proper flowering, and plentiful fruit, many
gardeners are necessary, each endeavouring to be
the right man in the right place.
DISCUSSION
C.G.G.J. VAN STEENIS: I congratulate Prof. Kusnoto
on his excellent exposition of the functions of tropical
herbaria. I am glad to learn that the Bogor institution
intends to combine supply of material to other herbaria
with work on the flora, as indeed all tropical institutes
should.
S.Y. HU: One must be careful to collect and distribute
enough duplicates of each specimen.
p.s. ASHION: May not too many specimens for identi-
fication or as representatives of local collections embarrass
the major herbaria and strain their facilities?
r.G.G.j. VAN SIEENIS: Well annotated, fertile material
will always be gratefully received by the herbaria in the
temperate regions.
H.M. BURKILL: The large herbaria still need such
material, even of common species. Duplicate specimens
are the best currency available to botanical institutions.
40
VOLUME 4 BOTANY
PROBLEMS OF BOTANICAL INSTITUTES IN THE TROPICSt
A. KOSTERMANS
Botanist of the Forest Research Institute* Bogor, Indonesia.
The problems discussed here deal with Botanic
Gardens and Herbaria. Those problems may be
divided into three categories: Time, Climate,
and Man.
Botanical investigation in its initial state was
taxonomical and started outside the tropics,
although its aim was the study of tropical plants.
For centuries, materials, which served as bases for
descriptions, floras, etc., have been heaped up,
first in European and later also in American
herbaria. Tropical institutes, like the Botanic
Gardens in Bogor, the Calcutta Botanical Garden
(in a lesser way), the Rio dc Janeiro Garden,
were more or less considered transit-institutes to
facilitate collections to be sent abroad. As soon
as foreign botanists withdrew from such institutes,
the work came usually more or less to a standstill.
The reasons are obvious: tropical institutes are
usually situated in countries where science is in
its initial stage or the local population has not yet
reached the stage of development for pure scien-
tific research. These comparatively young nations
have other, more important problems, which
confront them. The repercussions of a weak
economy were felt several times during the English
and Dutch Colonial period; in poor times, the
Bogor Institute was deprived of its staff, and work
came to a standstill, while the collections were
neglected.
One of the main difficulties, confronting the
Herbaria of young countries, is that most of the
type material is abroad and not represented in
the country itself. This is a big handicap, as it is
a time and money consuming work to study such
specimens abroad, as several European Herbaria
are unwilling to send material on loan for fear of
damage and loss (which is quite understandable).
On the other hand, we should never forget, that
because collections were kept outside the tropics,
many of the older collections are still in good
condition. Without these collections, progress
in taxonomy would be much hampered.
The young tropical countries, which will take
up now the completion of taxonomic and floristic
work are confronted with almost insurmountable
difficulties. We will mention only a few. Taxo-
nomy in Europe and America is not very much
appreciated by the "man in the street," who
t Presented by Kusnoto Setyodiwiryo.
considers it a waste of time and money and hence
these governments pay poor salaries for too few
jobs, and funds for working arc scarce. As these
conditions prevail in European countries, we can
imagine, that they are far worse in tropical
countries.
There is only one remedy: education. The
wealth of the vegetation in the tropics can never
be made useful to man without thorough investi-
gation, which must be carried out in botanical
institutes and must start with inventory, which
means taxonomic work.
Besides lack of funds, there is a lack of qualified
botanists. The jobs arc not attractive, as payment
is poor and the study long and difficult. For a
taxonomic botanist, knowledge of four or five
foreign languages is an absolute must, which
means that a student in Indonesia, who is taught
only one foreign language in secondary school,
has to spend much time and energy to make up
for the other languages, and this is often a hope-
less task.
Then comes problem of maintaining existant
herbaria and collections of new material.
It is certainly not true that in the humid tropics
the dried material deteriorates very quickly. That
depends on how the material has been preserved
and how it is stored. Attack by insects is suc-
cessfully checked by poisoning the specimens
with an alcoholic solution of corrosive sublimate.
Bulky parts of specimens, like fruit, where the
sublimate does not penetrate, are likely to be
attacked and part of this material should be
conserved in spirits. The glueing of the specimen
on the mounting sheet by means of a poisoned
glue prevents further attack of insects. We could
observe that the material in the Singapore Herba-
ria was far less attacked than that in Bogor; in
Bogor the material was not glued on (nowadays
the method of glueing has been adopted too).
The glueing does not interfere much with taxo-
nomic investigation, as tepid water easily loosens
the parts, which can be taken off for investigation.
On the other hand, the advantages of glueing are
enormous. Not only is the deterioration of the
plant checked, not only is insect damage less, but
the damage caused by handling is reduced to a
41
PROCEEDINGS OF THH NINTH PACIFIC SCIENCH CONGRESS
minimum. The damage done by handling and
dispatching the Bogor material has done so much
damage that the material must be considered
poor in comparison with that of Singapore.
As herbaria in the tropics are usually open
buildings, infection is more likely than in colder
climates. This may be prevented by storing in
tin boxes with tight fitting lids; an insect repellant
might help to chase insects.
There is, however, a great danger for specimens
on loan from abroad which are often not
poisoned. To young workers this is often not
known, and the stored material may be finished
by insects in a couple of weeks. Storage of such
material is only possible in hermetically sealed
boxes, provided with an insecticide. As the
material, however, is handled, this is not much
good. The safest way, which we have adopted
in Bogor, is to poison the loaned specimens in
a pure alcoholic corrosive sublimate solution,
wherein the mounted sheet is immersed for a short
period. The sheet remains clean, and the poison-
ing is invisible.
Upkeep of a collection provided the funds
are there docs not add any problems, as this
routine work is within reach of a lower-paid
official. In Bogor, however, we have trouble in
filing the material properly. Personnel doing this
have practically no inkling of scientific names,
and mistakes are numerous. Again, this is a
problem which could easily be eliminated by a
better paid official with a higher education.
Next comes the collecting of new material.
If the herbarium in a tropical country wants to
stand on its own legs, it should collect as much
material as possible to replace type specimens as
far as possible. A condition, as prevailing in
Bogor, where only half or less of the known and
described plants are represented is an unsatis-
factory condition.
Collecting, however, is possible only by means
of a team of qualified botanists with ample funds.
So long as these two requisites are not fulfilled,
the work should be postponed, as we have learned
from experience in the Forest Research Institute
that indiscriminate collecting is a waste of time
and money.
A "Botanical Survey/' as set up by the Indian
Government to initiate and guide an over-all
survey, has much to commend itself, provided it
has enough funds and qualified personnel (which
is for the time being not the case in Indonesia).
As matters stand in Indonesia, where no flora
is extant and most of the type material is not
available in the country, the only alternative was
to continue the taxonomic survey in the way it
has been done for almost 200 years. A body has
been created, the Flora Malesiana Foundation,
initiated and under the able directorship of
C.G.G.J. Van Steenis in Leiden and Kusnoto
Setyodiwiryo in Bogor, to publish the flora of
not only the area of Indonesia, but also the Phi-
lippines and Malaya. The bulk of the scientific
work is done in Leiden, where a botanical staff
has at its disposition an adequate library and
necessary herbarium material. The funds for
this important work are provided by the Indone-
sian Government. Moreover, the Bogor Botanic
Gardens take an active part by providing herbar-
ium material.
It is in this way only, that botanical investiga-
tion may be carried on in Indonesia. In the
future, Indonesian taxonomists should take an
active part in this large scale important work.
For the time being, Indonesia may contribute by
increasing its activity in collecting material all
over the area and in providing the means (funds
and help) to support the foreign taxonomists,
who work on this flora on a non-profit base.
Again it is proved here that more than any
other science taxonomy is international, and the
whole world benefits from the scientific cooper-
ation of nations.
The two main problems of the Botanic Gardens
in Bogor are lack of funds and (as is the case
with numerous, also extra-tropical gardens) a
difficulty in getting the collections properly
named. For a large garden as that of Bogor
which consists mainly of woody plants (amongst
which are several undescribed ones), a qualified
taxonomist is urgently needed. If a professional
botanist is added to the Garden Staff, the danger
that the importance of the collection will decline
is eliminated. With a non-botanical staff, the
trend is more to raise ornamentals, and the scien-
tific collection may be neglected.
Continuous collecting of seeds and seedlings
of botanical, more than of ornamental or business
interest, is an absolute must. This is, even under
prevailing circumstances, always possible, pro-
vided the staff is convinced of the scientific im-
portance of the Garden.
DISCUSSION
C.G.G.J. VAN STLENIS: I consider the material at Singa-
pore and Bogor to be of similar quality. Glueing specimens
42
to sheets was developed to reduce accidental loss (or theft)
of portions of the specimens, but this practice is most
VOLUME 4 BOTANY
troublesome to research botanists. Despite the greater o. SEIDENFADAN: Taxonomic research is the essential
cost in paper and space, wrapping each specimen in a basis for economic development of natural resources;
separate cover is the best protection against friction damage governments must realize this and give adequate financial
and loss ot detached fragments. support. Institutes must correct the opinion that such
^^^
P.R. wvrntRLhY: 8-hydroxy-quinoline-potassium sul- r.c,.j. VAN SIEENIS: We should all endorse His
phate may be used as a substitute for, or in addition to, bxcellency's statement, may we record this? (There were
mercuric chloride. Paradichlorobenzene is a valuable insec- no dissenting voices, and the meeting appeared to be in
ticide in herbarium cabinets. general agreement.)
43
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
A BRIEF HISTORY OF "BOTANIC GARDENS WITH
SPECIAL REFERENCE TO SINGAPOREt
J.W. PURSEGLOVE
Professor oj Botany, Impci ial College of Tropical Agriculture, Trinidad. '\
This paper is a precis of PURSEGLOVE:
History and Functions of Botanic Gardens with
special reference to Singapore: Tropical Agri-
culture 34(3), July 1957.
Botanic Gardens are places at which to study
plants. The earliest were herb gardens, the first
being that at Pisa founded in 1543. From then
through the Linnean era to the present day,
gardens have developed on scientific concepts,
accumulating not only living plants but herba-
rium material and literature, some on a world-
wide basis, some on a local basis.
Though many gardens were established in the
16th and 17th centuries in Europe, the earliest
botanic garden in the tropics is thought to have
been that at Pamplemousses in Mauritius, es-
tablished in 1735. In that century and the next,
many tropical gardens were set up, chiefly with
the object of studying plants of economic impor-
tance either local or introduced. In Asia, the
gardens at Calcutta were founded in 1786; at
Penang (the first of three), about 1796; Buiten-
zorg (Bogor), 1817; Peradeniya, 1821; and at
Singapore (the first of three), in 1822. These
gardens and the men who have served in them
have laid the foundation for the scientific study of
Asian botany and of much of the forest and agri-
cultural economy of the tropics. One has only
to consider the tremendous value to the world
the following tropical agricultural plants have
and peculiarly that most of them were removed
from their natural habitat to alien lands: Hevea,
cacao, quinine, coffee, cloves, nutmegs, sugar,
bananas, limes, vanilla, cassava, sweet potatoes,
maize, and many others.
In Malaya, the first Penang Gardens were
founded soon after the establishment of a trading
station by the East India Company in 1786.
Similarly the first gardens in Singapore were
founded but three years after the trading post
was set up. The attention of both was given to
the growing of nutmegs, cloves, and other crops
of commercial value. Of the surviving successors
to these Gardens, that at Penang is now a pleasure
gardens; and only that at Singapore is a botanic
garden in a true sense. This latter came into being
t Presented by H.M. Burkill.
t Formerly Director of Botanic Garden, Singapore.
44
in 1859 as the garden of an Agri-horticultural
Association. Within a few years, the government
had taken over its administration; and its work
expanded to include a study of the flora of the
Malay Peninsula, as well as work on crops of
possible economic value, such as Cinchona,
coffee, eucalyptus, ipecacuanha, tea, maize,
sugar-cane, colanuts, mahogany, oil palm, cacao,
and, of course, the classic pioneering of Hevea.
The Singapore Gardens made the greatest
progress in laying the foundation for agriculture
and forestry in Malaya under Ridley in the two
decades following 1888 when he arrived as
Director. Ridley also conducted the most exten-
sive research into the flora of the country. He
founded The Agricultural Bulletin of (he Malay
Peninsula in 1891 to be superceded by The Agri-
cultural Bulletin of the Straits and Federated
Malay States in 1901.
Through the researches of the Botanic Gardens,
the Departments of Forestry and of Agriculture
were set up to extend the work of their respective
lines. This did not mean that the Botanic Gar-
dens ceased their interest in these matters. The
work of all generations of members of the Depart-
ment has had a bearing on the evaluation of
Malaya's inherent wealth in its vegetation and
in its agri-horticultural potential which is now
made known through its serial publication, The
Gardens' Bulletin, and The Revised Flora of
Malaya. Even during the Japanese interregnum
in Malaya, work at the Gardens continued under
the direction of Japanese botanists. Though this
period was fraught with difficulty, the herbarium
and library suffered no loss, but the living col-
lection of plants was sadly reduced. In the post-
war years, the Gardens quickly regained their
former good condition, but these latter years have
been characterised by lack of trained staff.
The work of a botanic garden is botanical,
horticultural, and educational. For the first
purpose, the garden must be a museum of living
plants to compliment the herbarium, a museum
of dried plants. In the Singapore Gardens, some
3,000 perennial species grow, not counting hy-
brids. The herbarium contains about 400,000
sheets, mostly of Malaysian origin, whose quality
and quantity are constantly being increased by
loans for revision and by exchanges of duplicate
material. Collecting is done in Malaya and in
neighbouring countries where the vegetation has
botanical affinities. This work leads to taxonomic
studies which are necessary preliminaries to the
preparation of floras. With the complexity of
Malaysian botany, this work is a tremendous
undertaking. Over the past one and a half
centuries that Malaysian botany has been studied,
an immense amount of information has been
gathered. In 1921-24, Ridley published the first
Flora of the Malay Peninsula. So much more
data are now available that this work is in urgent
need of being superceded by a revised Flora, a
work which has already been begun. Similarly
the Singapore Gardens is throwing in its lot
with the immensely more difficult task in the pre-
paration of a regional flora, The Flora Malesiana.
It is only from such undertakings that man can
live, for, to quote Linneus "Science, and in the
first place botany, is the only reliable basis of
private as well as national economy."
On the horticultural aspect, the exchange of
seed and the acquisition of species of possible
value from other countries are equally important
activities. It is the natural sequel to the botanical
research just mentioned. From it has arisen,
for example, the immense and vitally important
Hevea planting industry. Besides Hevea, there
are numerous other plants, of less apparently
over-riding importance but nevertheless of value
commercially (e.g. oil palm, cacao), medically
(e.g. Rauwolfia spp.), and aesthetically (e.g.
Mucuna, cacti, orchids). The work on agricul-
tural plants is now the responsibility of the
Department of Agriculture, but nevertheless the
Gardens continues to play an important part.
Horticulturally Malaya's relatively seasonless
climate is a difficulty for many spectacular orna-
VOLU_ME_4 BOTANY
mental do not receive the required stimulus to
flower. Breeding work has been done on some
groups of plants to select responsive varieties,
and the main breeding work at present being
done is on orchids.
Educationally the Gardens help to increase
people's, in particular children's, interest in plants.
School parties regularly visit, and the Gardens
supplies material to schools and the University
for teaching and planting. A considerable
amount of advisory work is also done and active
encouragement is given to such bodies as the
Singapore Gardening Society, the Malayan
Orchid Society, and the Malayan Agri-Horticul-
tural Association.
In addition to the foregoing, the Gardens has
the responsibility of running the Nature Reserves
on Singapore Island in which all wild life is pro-
tected. The botanical and educational aspects
of the Gardens' work are represented in this
activity.
The future, with political maturity of Malaya
and Singapore, holds two cogent problems.
Firstly, there is the question of trained staff, for
much will depend on the new recruits who will
have to take over the duties heretofore exercised
by overseas men, and who it is to be hoped will
carry on the traditions of competent research
built up over the past century. Secondly, there
is the question of finance, a satisfactory solution
to which rests on the ability of the staff to demon-
strate to politicians and the public the worthwhile
role the Gardens plays in the economy and pres-
tige of the country. Two and a half centuries ago
the following lines were written about botany:
"It is certain, however, that this Science, like all
Sciences, flourishes sometimes more and some-
times less, all in accord with the inclination of
Rulers and the Favour of Government."
(Commelijn) How true it is still today!
DISCUSSION
p.w. RICHARDS: The collections of living plants in the
tropical gardens must not be neglected, nor limited to
economic or ornamental species. Visiting botanists find
them invaluable for a wide range of studies.
J.H. HURUMANN: Private institutions are faced with
the same difficulties as official organizations, and so their
utility is limited.
45
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
SOME CURRENT PROBLEMS OF THE BOTANIC GARDENS,
SINGAPORE
H.M. BURKILL
Botanic Gardens, Singapore.
ADMINISTRATION
In a period of transition of administrative
authority such as is going on in Singapore at
present, it is an advantage to maintain a sense of
humour. Many odd things crop up, like the
zealous Treasury official who, at an application
for funds to buy some more herbarium cabinets,
quoted Government's General Orders which
require a Director of a department to review his
department's records when they become five years
old with a view to destroying those of no further
import. Strictly correct, of course, but when do
botanical records (herbarium specimens) reach
an age for destruction ? Or again, another official
dealing with equipment for the Gardens wanted
to know how many persons normally sat on a
garden seat. He had plainly yet to learn the facts
of life that in Singapore's predominantly ado-
lescent population, "two's company; three's
none!"
These points will serve to show that in the
growing political responsibilities of the new
nations of Southeast Asia there is now a cadre of
administrative officers with everything to learn
in the shortest of time. Rome was riot built in a
day. An intelligent inquisitivcness is a healthy
sign amongst this new band of administrators,
but the simplest way of learning is by coming to
see and personal discussion, and one could
wish that this, disrupting though it may at times
be to normal routine, was more often practised.
Though it may be adequate to copy one's pre-
decessors, it is desirable that those who hold the
purse-strings and those who run the department
maintain a close liasion. Thus one may hope to
avoid the unpredictable response to a normal
contingency and gain the advantage of both
parties speaking and thinking alike.
PERSONNEL
Every branch of scientific endeavour through-
out the world is short of trained personnel. The
Botanic Gardens, Singapore, has not been un-
affected. The normal compliment of qualified
staff is four botanists, i.e. a Director, an Assistant
Director, a Keeper of the Herbarium, and a
46
Botanist, and two horticultural Curators. From
1946 to the end of 1954, except for a short period
in 1948-49, there was no Assistant Director.
From 1952 to 1956, the post of botanist was
vacant. From 1946 to 1954, the horticultural
branch was virtually down to one officer, as one
Curator was seconded for duty as Agricultural
Officer. These shortages have created much
difficulty in carrying out a full programme of
research and have thrown additional burdens on
the existing staff. This Department is not in the
process of "empire-building"; its full establish-
ment of botanical officers has remained un-
changed for 33 years, and its compliment of
Curators has been reduced to one since the
Penang Botanic Gardens were handed over to
the Department of Agriculture in 1946.
For the first time since 1946, a full compliment
of senior staff was at work from December, 1954,
though one post was vacant but held temporarily
by a re-engaged pensioner in the absence of any
qualified younger man suitable for permanent
appointment. For two years this position was
maintained and even the vacant post was filled.
Then in 1957, the Singapore Government intro-
duced a policy of accelerated Malayanisation of
the public service. This resulted in the loss of one
botanical and one horticultural officer, of which
only the latter has been replaced. Further losses
are expected. It is not intended to discuss politics,
but it is essential if the situation is to be under-
stood. Briefly, it is that local men are coming
forward in greater numbers for higher training.
Facilities are now available at the University of
Malaya and abroad by means of awards to qual-
ify local men for appointment to posts hitherto
held by officers from overseas. For the expatriate
officer, the element of uncertainty, the offer of
compensation for loss of career, the knowledge
that sooner or later he will have to retire, and the
younger he goes the better chance he will have of
starting again in new employment are all cogent
reasons for the loss of trained man power in
government service in Singapore. Many ex-
patriate officers have and are leaving before local
men are ready to fill their posts. Some or all of
these reasons apply to the Botanic Gardens
VOLUME 4
BOTANY
though it is apparent there will be no claim for
appointment from local persons for some years.
Now most biological work is essentially long
term particularly that of the botanist. Plant
collections and written notes and papers are only
a part of the botanical archives an institution
possesses. Equally important are the personal
experiences, the stored-up impressions and ideas
of its officers. Much of this information does not
get onto paper and published till towards the end
of a man's career. Let me take two examples:
The Flora of Malaya (Ridley, 1922-25) and the
Revised Flora of Malaya (Holttum, Vol. I 1953,
Vol. II 1955) could never have been written by
any other authors in as complete and competent
a form from the available collections and liter-
ature alone simply because such works are the
better for the authors having had first hand per-
sonal experience. Ridley was in Malaya for 24
years; Holttum, for 31 years.
The point I come to then is that frequent change
of staff where it involves a complete break of
environment in research workers' careers is bad
for the institution. It may be bad also for the
individual, but that is outside the present issue.
The changes which are now taking place in the
Singapore Botanic Gardens through political
circumstances are in danger of seriously putting
the clock back. It can never, of course, go com-
pletely back, but it does mean that over the next
ten years or so there will be an unfortunate break
in that intangible quality, personal experience,
which is such an important asset to any institution.
I have no solution to offer. Advice has been
tendered. Recommendations have been made.
The government has realised the situation.
Decisions have been taken. Events are running
their course.
How is the loss to be made good ? The physical
loss of bodies and the abstract loss of minds will
no doubt eventually be recovered. For the former
we depend on the University of Malaya and
overseas universities where Malayans are training,
but for the present, prospects are not good for
only a few students are coming forward to take
botany. For the latter, there can be no progress
till the former is made good, and then only time
will tell.
This shortage of trained personnel has caused
the introduction of a new concept of recruitment.
Whereas it was previously accepted that persons
arranged their own training and, except for a few
fortunates who could obtain scholarships, paid
for it privately, this liability now devolves to a
considerable extent on Government and on
public funds. Further, whereas it was usual to
appoint a man already trained, it is now accepted
that a promising but untrained or partly trained
man is appointed to a post (or is anyhow bespoke
for the post) and then trained for it. This is no
doubt the easiest way of creating a corps of quali-
fied man in the shortest possible time and is in
accordance with Government's increasing con-
cern with scientific matters.
In the subordinate cadres, the position has been
much easier except where specialised knowledge
is required. Librarians appear to be hard to come
by, and the Departmental library has been with-
out adequate management for a long time. This
throws out of gear all the normal services that
one expects of a librarian such as receipts, loans,
and exchanged; referencing and cataloguing;
binding; pest control and prophylaxis; rack
arrangement; and the assistance expected by
botanical officers in locating literature. The
solution to the shortage of librarians lies in train-
ing, but Malaya at the present time does not offer
any facilities and few students are going overseas
to study for librarianships.
One point on which this meeting might raise
discussion is on the nature of the basic training
required by a librarian of a botanical library.
To run such a library, in my opinion, the librarian
must have had some botanical training so that
he may know the difference between, for example,
agriculture, horticulture, and botany in their
general meaning or between a phanerogam and
a cryptogam in a more technical sense. With
such knowledge he can without troubling the
botanical staff carry out the functions of a
librarian of a botanical library. I would therefore
expect in the position I now find myself in the
matter of recruiting a librarian to start with some-
one with some knowledge of botany and to train
him in library techniques. In this way I would
ensure having a librarian capable of handling
botanical literature.
On the other hand, there is a body of opinion
amongst librarians that a librarian should be a
librarian and nothing else. I am assured that the
botany around him would look after itself. But
in a specialised library it surely calls for
specialised knowledge to classify competently
the constant inflow of literature, particularly if
one considers that it is part of a librarian's duty
to maintain a subject card index. The librarians'
opinion to which I have just referred indicates
a severe limitation of usefulness, but it is a posi-
tion which now holds in Singapore and perhaps
in general elsewhere. It is a matter on which I
would greatly welcome comments.
47
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
With regard to other subordinate grades we are
in no particular difficulty for the departmental
organisation provides the means of training
whereby a youngster can earn and learn his way
to positions of responsibility. In fact the Botanic
Gardens of Singapore in some respects are rather
like an old family concern; out of the twenty-six
botanical and horticultural posts on the staff, no
less than nine are filled by fathers or sons of
present or past employees of the Department.
I myself take pride in being in this category.
COLLECTING
The Botanic Gardens, Singapore, is the only
botanical institution in the British Territories of
Southeast Asia. It is true there are other herba-
ria, but they tend to be specialised, with the result
that the sphere of activity of the Department has
been spread over the Malay Peninsula and Bor-
neo. Recently occasional visits have been pos-
sible to Borneo, but till 1955 these were restricted
by shortage of staff. In Malaya, with ease of
access, quite a different problem has arisen. For
the past nine years there has been armed insur-
rection in the country by a small proportion of
the population which has taken to the forests.
The trouble was incipient from the end of the war,
but gained official cognisance in June, 1948, with
the euphemistic title of "Emergency." For some
years it reached serious proportions and even now
is the cause of difficulties and restrictions in
several parts of Malaya.
This state of affairs has had a very important
effect on collecting and botanical exploration, as
it has been impossible to penetrate the forests.
At times it has been forbidden, or unwise, to stop
one's car on the main roads to have a look at the
roadside vegetation. To wander into the forest
would have been an invitation of trouble with
either the police or military on patrol or from the
insurrectionists. In either case, one would have
been at the receiving end. Planters, miners,
foresters, and surveyors have had to work accom-
panied by armed escorts. 1 know only too well
from personal experience the physical and mental
limitation such a condition imposes.
The consequence has been that collecting has
been severely restricted to those areas one could
safely visit and these on the whole are the areas
best known botanically, i.e. the vicinity of towns
and hill stations. The less accessible centre of
the Malay Peninsula is where exploration is most
wanted, for example, Gunong Kerbau, Gunong
Tahan, Ulu Kelantan, and theTrengganu plateau.
48
Armed conflict such as Malaya is experiencing
has been or is a feature in most of the countries
of Southeast Asia during the past decade. It is
something with which some at this meeting may
have had close personal experience. It is some-
thing which all of us can deplore, though, I fear
there is nothing this meeting can do to alleviate it.
It has been the biggest obstacle in the past ten
years to sociological, industrial, and research
progress.
The situation improves in Malaya, and botan-
ical exploration into the more remote parts of
the country may soon be possible again, though
the pre-war style of expedition lasting a month
or more with men and equipment to match is
likely to be little more than a pipe-dream except
on rare occasions, for even some dreams come
true! But in general we will have to work on the
principle of light excursion lasting not more than
a few days or on single-day collecting trips.
An aspect of some importance is the need for
botanists to have some familiarity with the types
of vegetation of neighbouring countries. A
foreign correspondent writes that he cannot plan
a trip to Malaya as he is unable to obtain the
necessary currency exchange. A Malayan cor-
respondent says his proposed trip to another
country is limited by the same curbs. I would
like to suggest that exchange of botanists between
institutions of neighbouring countries should be
encouraged in much the same way as students
and university lecturers are exchanged. The free
provision of herbarium amenities is naturally
simple, but the principle may well be expanded
to the offer on a reciprocal basis of facilities for
field work, transport, subordinate assistance,
and lodging.
VISITING RESEARCH WORKERS
Many scientists call at Singapore on their way
to other countries. Few stay to work. If tropical
botany is to be better understood by botanists
from institutions in temperate countries, it is very
desirable that some botanists come to work for
short periods at tropical herbaria. They thus
acquaint themselves with some of the problems
of tropical botany, carry their impressions back
to their home institutions, and interest their
colleagues. In the last few years, we (Botanic
Gardens and the University of Malaya) have been
fortunate in obtaining grants to permit two young
botanists to come to Malaya for a year each.
It is hoped that this may be repeated regularly
to the advantage of all concerned. There are
VOLUME 4
BOTANY
certainly numberless lines of investigation ready
for working on. The grants have come from the
British Government Colonial Welfare and
Development Scheme, but there are International
Agencies which could or should furnish some
assistance. Claims for assistance are more than
their funds can stretch to, and botanical matters
do not usually rank high in priority. Plants are
fundamental to life, and their study is man's first
science. It is our duty to raise the sociological
status of botany in the minds of the world's
administrators.
DISTRIBUTION AND EXCHANGE
Botanical security can be achieved only by
ensuring permanent accessibility to collected
material, which are the basic botanical archives.
Essentially, the problem is one of availability and
adequate distribution.
The development of botany in the countries of
Southeast Asia at the present time is still very
much interwoven with herbaria of the temperate
world. The connections of the Malaysian flora
are especially strong with Kew, British Museum,
Leiden, and the Arnold Arboretum. Many other
links exist between tropical Asia, tropical Africa,
and temperate countries.
For our part, our system of distribution of
duplicates may help to perpetuate this depend-
ence on Europe, but it is a system which cannot
be changed if we are to continue to make the best
uses of our material and if we are to follow the
principle of distribution for security sake. That
our top duplicates go to those institutions where
experts are working on the groups of plants con-
cerned is a general policy. For us it usually means
to Kew, Leiden, and the British Museum. The
longer we send to them, the more important their
Malayan collections become; the more important
their Malayan collections are, the more reason we
have for adding to them and keeping them up-
to-date. This is a sort of cleft stick in which we find
ourselves, albeit not an unpleasant one, nor an
unprofitable one. This is the position which we
at Singapore hope to maintain at least for the
foreseeable future till there is a capacity and talent
amongst the local population for things botanical.
Thereafter one would wish that such distribution
be maintained as a feature of cooperation for
mutual benefit. The extent of distribution of
duplicates of the higher plants after sending to
these museums takes a wide range. All is done
on free exchange, but it is limited to those her-
baria of Europe, N. America, and Australia with
Malaysian affiliations and to herbaria of the
Indo-Pacific region, with whom we are in ex-
change. There is, I believe, a growing opinion
that a regional flora cannot be considered as
peculiar to a water-tight compartment. The time
is coming when the floras of the tropics may have
to be taken as a whole. I am thinking, therefore,
that our system of distribution is inadequate and
that we may have to exchange our material for
tropical African and tropical American material
also, if the expanding requirements of taxonomic
study are to be readily met. This will impose
greater demands on storage space and the limited
numbers of duplicates available for distribution.
It is perhaps worth mentioning that during the
past three years for every number collected by
our staff and laid in our herbarium there has been
an average of 4^ duplicates available for dis-
tribution. We could without difficulty distribute
double that, and we would do so were it not for
the difficulty of handling so many duplicates in
the field. Any increase in the number of exchang-
ing institutions has obvious drawbacks.
It is, however, not so much the normal run of
current collections which give rise to the greatest
concern, but the type specimens, the classical
botanical archives. The Singapore Herbarium
has about 1,000 type sheets, many are holotypes,
others are iso-types. There may or may not be
duplicates elsewhere; but of the older ones, it is
likely that distribution was much more limited
than one would now-a-days deem desirable.
But whatever their distribution elsewhere, we have
an obligation to ensure their availability to present
and future botanists. Several herbaria with
valuable collections suffered losses during the
late war. The Singapore Herbarium was fortunate
to have had no damage, for which the actions of
Professors Tanakadate and Kwan Koriba, Dr.
Holttum and Mr. Corner must be recognised.
But preservation whether from war damage or
from accident is a problem concerning all her-
baria housing valuable material. For ourselves,
we have recently had most of our type sheets
microfilmed and propose to distribute positive
copies to certain herbaria for their reference as
well as for safe custody. Except for an adequate
distribution, this is perhaps the nearest approach
to a satisfactory solution that one could have,
expedient though it is.
LOANS
Free exchange and free loan are, in general,
principles to be commended. Loans from the
Singapore Herbarium since 1946 have involved
49
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
39,000 sheets or about 10% of the total herbar-
ium collection. These loans, totalling nearly
100 lots, are looked on with favour as most of
them are for revision purposes, and the sheets
thus come back annotated with the current
taxonomic name.
The increasing complexity of taxonomic revi-
sion is reflected in the length of time these loans
are out; for those sent out and returned, the
average has been twenty-two months. Seven
were out over four years; two over five years;
and one over six years. Of the material currently
out on loans, there are 11,000 sheets issued in
thirty-two different consignments covering genera
or sections or, for a few, the whole material of
orders. Three have been out over nine and one-
half years; two over eight years; and two over
seven years. I cannot help but think that some-
thing has gone wrong with these! What of the
fervent instructions of loaning herbaria that
the loan is only for three months?
The accusing finger can no doubt be pointed
at the Singapore Herbarium too. It is a common
problem, but the point I would make is that the
extensiveness of loans now out is apt to cause
considerable embarrassment in handling current
collections, as there is much which cannot be
matched. It is not necessarily a temporary prob-
lem, but is likely to grow with the expansion of
botanical study.
BOTANIC GARDEN OR PUBLIC PARK
Most botanical institutions have attached to
them some extent of land, garden, estate experi-
ment station, or whatever the appropriate title
might be, where plants may be grown under
observation. A botanic garden usually attracts
many visitors, though basically it is a research
station rather than a pleasure garden. The
Singapore Gardens were laid out originally as
the latter on formal lines of landscaping, in all
eighty-six acres, when in 1882 the Government
took them over as part of a botanical institution.
The design has remained unchanged for almost
100 years.
Problems inherent in the setup of the Singapore
Gardens are these:
(1) Space;
(2) Pressure of the public.
With regard to the first, the Gardens are but
four miles from the centre of the city and are in a
built-up residential district. Expansion is impos-
sible. With the old Economic Garden which was
attached to the Botanic Gardens, the area was 188
50
acres which was close enough to 200 acres,
often considered an optimum size. In 1925, the
Economic Garden of 102 acres was excised for
a college football field. The Frenchman's charac-
terisation of the British "Toujours le Sport" is
justly applied. But the result is that chiefly in
regard to tree species, planting is greatly res-
tricted. If the lawns of the gardens were planted
over, more desirable introductions could be tried,
but concession has to be paid to appearance and
to horticultural considerations. The need for an
arboretum is paramount at the present time.
The acquisition of additional land has been con-
sidered off and on for the past thirty years since
the Economic Garden was lost and is now com-
ing to a head again with further demands on the
Gardens.
The other problem has many facets. The
Gardens are one of the few open spaces within
the City Area. In fine weather on week-ends or
public holidays, many thousands of people visit
them. In general the public's behaviour is per-
fectly satisfactory, but in their train is the time-
old trouble of the few anti-social individuals
against whom restrictions have to be imposed
which affect all visitors to the Gardens: the bicy-
clists, who as a class, are pests; the poachers who
try to catch fish in the Gardens lake; the pilferers
who break cuttings ofT plants so that the plants
have to be removed; the orchid fanciers who
remove orchid pollenia for fertilising their own
blooms and thus cause our plants on display to
fade quickly. These are the troublemakers; their
kind is common throughout the world.
Of our own domestic horticultural problems,
the biggest is monkeys. They are utterly des-
tructive. All horticultural practices arc fair game
for interference. Anything special has to be
established under a wire cage or behind an elec-
trified fence. Any attempt to reduce their numbers
is a signal for outcry ; ones office is beset with holy
matrons and true citizens defending their birth
right, for to the public the Botanic Gardens are
a place to come to feed the monkeys.
The amenities which the public has in the
Gardens are undoubtedly appreciated, and it
seems to me that the best possible balance is
struck between botanical/horticultural require-
ments and the public's usage as a park. With
more children learning biology in the schools,
it is gratifying to have orderly parties conducted
by teachers. This is a good augury, for there are
many too many people who look upon plants as
something to cut down or pull up according to
size, and it is a function of a botanic garden,
VOLUME 4
BOTANY
rather than a park, to foster a love of plants in
the predominately destructive human mind. In
the following section I refer to a very closely
related subject.
NATURE CONSERVATION
This subject is dealt with in another sympo-
sium, but as it is a problem confronting the Botan-
ic Gardens, Singapore, it is correctly mentioned
here, if but briefly.
In Singapore, the preservation of small sample
areas of vegetation of the indigenous flora (and
fauna) is a major problem. There are five areas
covering about 8,000 acres constituted nature
reserves by law under a board of trustees with
the Director of Botanic Gardens as Chairman.
Though this legislation was enacted as recently
as 1951 and even more recent land utilisation
planning has respected the areas as reserves, it is
proving difficult to maintain them intact against
catch-penny commercialism, demands for
resources they contain (granite), and expanded
development plans hitherto unforeseen.
Two of the reserves are type localities for species
of plants and marine animals. All are essential
for education and research, in addition to their
obvious attractions as places for recreation and
open air enjoyment. Their very accessibility
from the city adds greatly to the amenities they
offer, but this too adds to the covetousness with
which certain persons would seek to destroy them
for commercial exploitation. Though there is
a growing land hunger in Singapore, there are
yet other similar areas of unreserved land else-
where on the Island awaiting rational utilisation.
SUMMARY
None of the problems described here are in-
superable. Some indeed are axiomatic in running
an organisation. Some, given goodwill and
patience, will resolve themselves. Others are due
to force majcur. Solutions to the first two kinds
lie in our own hands. For the third, we must
adapt ourselves as good citizens and good
biologists.
D/SCUSS/ON
G.A. PROWSE : Men of long proven, practical experience
are being ousted by others with better academic records
but less knowledge of and training in the particular sub-
ject. This occurs where inexperienced administrators
fail to appreciate the relative merits of experience and
proper qualifications. Better mutual understanding
between institutions and the authorities they serve is
required.
P.R. WYCHERLEY: All papers have referred to the
shortage of trained staff. In Southeast Asia especially,
too few students are attracted to botany; many graduates
seek employment elsewhere, although they are sorely
needed in botanical, agricultural, and forest research
stations.
K. SETYODiwiRYo: Botany is a poorly paid profession
in Indonesia, hence the few young scientists entering the
field.
H.M. BURKILL: Many Singapore graduates become
teachers. We must raise the sociological status of botanists.
E.H. WALKER: We need better advertisement of the
importance of botanical work which should promote
more recognition and reward for those engaged in plant
science. These difficulties are very general.
C.G.G.J. VAN STEENIS: The exchange of botanists be-
tween different tropical institutes and between them and
the temperate region herbaria, as suggested by Mr. Burkill,
will greatly benefit the development of tropical botany.
Tropical institutes must not be left to develop in isolation;
personal contacts are invaluable to all. Bogor and Singa-
pore have laid a good foundation in accommodating visit-
ing specialists; this successful enterprise must be extended.
Specimens are sometimes out on loan for many years,
because the monographers are often university teachers who
cannot devote all their time to taxonomic revision. Plan-
ning when the specimens are required by the borrower
will help, and herbaria should remind borrowers of the
outstanding loan periodically after a reasonable time for
study has been allowed.
R. HEIM: Young botanists must be encouraged to take
up taxonomy. Botanists must aid the establishment of
reserves and parks to retain living specimens of the flora,
as well as planting formal botanical gardens. The wide
distribution of duplicate specimens cannot be emphasized
too strongly.
E.H. WALKER: I endorse the need for new taxonomic
studies. Some of the problems of loaned specimens and
exchanges of specialists could be resolved if there were
regular conferences of the directors of botanical institutes
to advance cooperation.
p.s. ASHTON: The working taxonomist and collector
must be able to visit not only herbaria but also the different
source areas in the region, irrespective of political bounda-
ries. A pooling of resources is needed so that botanists in
poor countries may reap the benefit of facilities established
by their richer neighbors.
E.H. WALKER: There is a lack of curators in the her-
baria I have recently visited in the Pacific area. Funds
have been cut, and although the senior men may remain,
whole groups of essential assistants have been unwisely
dispensed with. Individual institutes probably cannot
work out their own salvation. General guidance, coordi-
nation of plans, and assistance in bringing their needs to
the notice of the authorities arc all necessary.
M.S. DOTY: I would remind you that we can pass reso-
lutions for the attention of the Pacific Science Association
or of UNESCO, whose support will add more weight to
our conclusions.
51
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
PROBLEMS FACING THE HERBARIA IN THAILAND
KASIN SUVATABANDHU
Department of Agriculture, Ministry of Agriculture, Bangkok, Thailand.
Anybody may realize that Thailand, land of
freedom and land of smiles in Southeast Asia,
is an agricultural country. She is very rich in
natural resources. Her soil is fertile and suitable
for various kinds of cultivation. Rice and teak
are famous and recognized by many customers.
The population is too little in comparison
with the vast area. Her economy is dependent
on plants and plant products.
In Thailand, as in many other "ropical countries,
botanical knowledge which is of basic impor-
tance is still in its initial state. Speaking in general,
the Thai people have a great interest in plants,
their economic value, and even the ecological
relations. Courses in botany are offered in the
university, but these are only for the scientific
background of medical, pharmaceutical, biolo-
gical, and agricultural students. The teaching
for the advancement of Botany is still in the
future. The Thai flora still receive scanty atten-
tion. No botanic gardens exist in the kingdom,
but there are a few reserved parts of the forest.
As a matter of fact, botanical works such as a
survey and systematic study have been made,
but for the use of other institutions which are
considered the main arteries of the country's
economy, i.e., Royal Forest Department and
Department of Agriculture.
In 1906, the Royal Forest Department started
the systematic study of plants, but these were
mainly timbers of economic importance and
some other forest products. A herbarium was
also built up. In 1920, the Botanical Section of
the then Ministry of Commerce was formed
with the object, in the first place, of making a
survey throughout the kingdom of plants yielding
or likely to yield, economic products. This sur-
vey was to include an enquiry into the properties,
quantities, and accessibility of such plants.
Again the herbarium was built up. This Section
is now under the Division of Plant Science, Depart-
ment of Agriculture. A great deal of work has
been done on the flowering plants of Thailand,
but it is still incomplete. Studies of ferns and
mosses have been made to a lesser extent, but
the lower plants, fungi, and algae are almost
unknown.
One might question why botanical studies and
work have not developed further in Thailand.
Shortages of trained scientific personnel and
scientific literature as well as inadequate funds are
all responsible.
The most important of these is the shortage of
personnel. In the two mentioned botanical
sections, there are only a few members of the
official staff, and work progressed very slowly.
Collections which are necessary and important
are made on a small scale. There are too few
duplicates to offer to other institutes. The main
factor is the difficulty of finding persons interested
in making a career of this purely scientific work.
There is a demand for persons in various fields
of applied science such as medicine, engineering,
architecture, agriculture, etc. Pure science has
thus far not attracted sufficient students and
knowledge of the country's vegetation is still very
scant. The shortage of trained personnel creates
other problems in developing this botanical
knowledge such as limited advanced study and
little coordination with other institutions.
At present official persons in various fields
have begun to realize the importance of botanical
knowledge. We hope that in the future this kind
of purely scientific work will have the full support
of the government, as a part of the country's
economic development.
DISCUSSION
C.G.O.J. VAN STEENIS: The lack of a botanic garden on
a scientific basis in Thailand is amazing in view of the
agricultural nature of the economy. Indonesia and Malaya
are examples of how botanic gardens, with their herbaria
and libraries, have aided agricultural development by
knowledge of the flora. The establishment of a botanical
garden requires careful long-range planning, for slow
but sure growth is needed. This is a government responsi-
bility. I propose that this section endorse any resolution
52
supporting the establishment of a botanic garden by the
Thai Government. (There was general agreement that a
botanic garden is an urgent necessity for any tropical
country, including Thailand, which lacks such an institute
for the botanical study and development of its agricultural
and forest resources.)
M.L. STEINER: Our attempts to rehabilitate the gardens
in Manila have failed. The trees and shrubs we planted
are destroyed. The promises of help from the authorities
VOLUME 4 BOTANY
have come to nothing, how can we press the point with tion is needed. The government may stand in need of
them? education in this matter.
E ' H - WALKER: Vigorous action is needed on all fronts.
interest larger organizations in obtaining effective support
C.G.G.J. VAN STFTNIS: Government, not private, ac- from the authorities.
53
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
INTERACTION AND COOPERATION BETWEEN TROPICAL
AND TEMPERATE HERBARIA
C.G.G.J. VAN STEENIS
Flora Malesiana Foundation* Oegstgeest* Netherlands.
Not long ago I was struck by a plea to mono-
graphers by Brother Alain (Havana) who com-
plained of the fact that monographers frequently
study less material than desirable and often
confine their work to the revision of the material
of some large, European herbaria (Taxon 6,
1957: 46-47). He pointed to the deficiency in
this respect of Radlkofcr's monograph of the
Sapindaceac in the Pflanzenreich.
It is true that in many monographs of this
series only a very limited amount of material
was borrowed from institutes outside Berlin-
Dahlem, and that for essential material only.
Two decades ago when working at Bogor,
Java, I had a similar complaint about Malaysian
material, and I wrote to Berlin to ask if we could
receive timely notice as to which families or
genera were under revision in order to loan our
material for examination by the monographer.
The answer was unsatisfactory, as it was argued
that the monographers could generally not
master large loans from all herbaria abroad and
did not solicit to ticket all specimens, as their
facilities and specially their time were restricted.
Nearly all of the monographs therefore suffer,
in varying degree, from incompleteness. And
the same holds for various other revisions general-
ly. The reference to Radlkofer's work is not
a particularly good example as it was essentially
finished before 1910 and remained, as far as I
know, in Radlkofer's desk for decades. Radl-
kofer did not add much to it in later years, and
the final posthumous editing was performed by
the redaction of the Pflanzenreich.
This situation looks lamentable and dangerous
to the progress of international taxonomical work.
It is partly due to the fact that tropical collections
are accumulating so fast that many large herbaria
are overcrowded. Furthermore, the number of
herbaria is still increasing.
On the other hand, monographers are still
scarce or monographic work is not sufficiently
envisaged to be the major source of creative work
and the available time of each monographer,
in contrast with the dynamism of collections, re-
mains static, i.e. twenty-four hours in a day and
night.
54
A long experience with the Flora Malesiana
revisions, which cover only a part of the tropics,
has taught that completeness in ticketing all sheets
which is generally a multiple of all collecting
numbers of all herbaria in the world containing
Malaysian plants is an impossibility. And from
this experience I infer that such a procedure
for a monograph of even only a medium-sized
family would lead to insurmountable personal,
technical, and financial difficulties; it requires
an immense amount of time from the side of
the monographer to cope with the routine work
involved.
To give a slight idea of what colossal numbers
of sheets must be revised for a monograph, in a
study of Cyperus only for Malaysia and from a
limited number of herbaria, about 17,000 sheets
passed through the hands of Mr. Kern for
comparison or identification and ticketing! For
Fimbristylis, the number of sheets was about
8,000.
The historical development of plant taxonomy
and distribution of collections has led to the
present situation of rather few large world her-
baria in which collections are presented and
worked on from all parts of the world, and rather
few very large herbaria which are more or less
concentrated on certain major regions of the
World (Africa, New World, Old World, Austra-
lia, etc.). Whether we like it or not, this situa-
tion has to be accepted.
Most of these institutes have less scientific
staff members than would be desirable; some are
even ridiculously understaffed. Generally staff
members must, by necessity, perform official
and routine work and are left little time for mono-
graphic work.
Combining this need of time on the side of the
monographers with the just mentioned immense
increase in sheets, there is an absolute necessity
for them to use their time as efficiently as possible,
resulting in limiting their work primarily to
essential material and to that which is immediate-
ly available, rather than to strive towards com-
pleteness in ticketing all specimens of their group
they can lay hands on. After all, a revision or
monograph derives its value from its usefulness,
VOLUME 4
BOTANY
from keys that work, from the degree of accuracy
by which genera and species are delimited and
described, and not from immense lists and
enumerations of specimens from this or that
herbarium.
I have touched on this subject in my prepara-
tion of the Flora Malesiana (ser. I, vol. 1, p.
Ixv) in a chapter on the policy of distributing
duplicates as advocated and applied by the late
Dr. Merrill in the Philippines who was in a simi-
lar position as we were at Bogor and Brother
Alain in Cuba. Dr. Merrill has recommended
that the tropical collectors and institutes take the
initiative and distribute duplicates to the large
world herbaria, specially to those from which it
can be expected that they will name them immedi-
ately or in future, and to those where already
large, authentic collections are represented from
the region where the new collections are made.
Of course the tropical institutions cannot expect
to come on terms of an exactly balanced ex-
change, as the world herbaria have generally
little to offer in this respect. Dr. Merrill did this
always on what he called a "free exchange basis,"
that is: both parties give what they are able to
produce. As far as material is concerned, the
tropical institutions will therefore be largely the
donating party.
But their effort is materially rewarded in several
respects: firstly, the collections, provided they
have been preliminarily identified to families and
preferably to genera, come into "scientific cir-
culation," they arc "available" and will come more
easily under the eyes of monographers, interested
staff members, or visitors of the large institutes.
Secondly, wide distribution of duplicates safe-
guards collections against calamities. Thirdly,
the reward to the donating party will exist in the
ultimate naming of its collections by specialists.
Fourthly, its initiative and diligence provides the
donating institution with favourable contacts
abroad and with specialists; it will stir up interest
in tropical plants. In passing I may give the
practical hint of printing at the bottom of each
duplicate label a request that any new identifi-
cation of the sheet be kindly brought to the
notice of the donating institution.
The history of the exploration of the Philippine
flora, through the impetus of Dr. Merrill, proves
the wisdom of the policy just mentioned. Even
though the calamity of war came over Manila
and destroyed what had been built during two
decades of extremely hard, intelligent, and dili-
gent work, the liberal way in which duplicates
had been distributed still guarantees the possibil-
ity of writing a complete Flora of the Philippines.
The rich collections assembled in tropical
centres must be brought to the notice of the work-
ers and visitors of the big world herbaria. This
is the duty of tropical assembling places, and
besides a duty it is a great privilege, because trop-
ical botany will, for at least half a century to come,
not only be mostly dependent on, but will also
benefit from contributions of the great scientific
centres in the temperate regions of the globe.
I believe that Brother Alain has touched on a
subject worthy to explain during this Congress
which serves to cultivate goodwill, understanding,
tolerance, and cooperation. On one hand I be-
lieve he underestimates the labour involved in
monographical work; on the other hand, he un-
derestimates the extreme value of tropical herb-
aria taking the initiative in their own hands. Trop-
ical herbaria should recognize the extreme value
of a very liberal policy of distributing duplicate
specimens of good material of native plants to the
world herbaria. Specialists, on the other hand,
are invited to give timely notice, preferably in
Taxon, of the subject of their intended monograph
or revision. In this way a satisfactory interaction
and cooperation can be realized.
DISCUSSION
E.H. WALKER: The collection at Manila was destroyed
by fire during the war, fortunately many duplicates are in
the Smithsonian Institute. This illustrates the need for
wide distribution of duplicates. Breakdowns in the loan
of this material call for more cooperation between directors.
55
PROCEEDINGS OF THEJNINTH PACIFIC SCIENCE CONGRESS
PROBLEMS "IN" A SPECIALIZED BOTANICAL "DEPARTMENT
G.A. PROWSE
Fish Culture Research Station, Batu Berendam, Malacca, Federation of Malaya.
While a botanical department of a freshwater
fish culture research station is not strictly speaking
the same thing as a botanical institution, many
of the problems are similar, while others will be
peculiar to the station itself. For that reason it
seems worthwhile bringing these problems before
a meeting of this kind, with the hopes that the
ensuing discussion may result in their solution,
or at least suggestions of their solution.
Taxonomy. Limitations of staff make it neces-
sary that a botanist working in such a station as
ours undertakes taxonomic studies on the various
plant groups, although unlike other botanical
institutions, he is at least limited to those plants
in an aquatic environment. The botanist here,
however, cannot afford to confine himself to one
particular group and must spread his efforts,
although inevitably he finds himself taking a
greater interest in one of the groups. Because
of this he finds himself dependent on the goodwill
and cooperation of others elsewhere, and various
problems necessarily arise.
LITERATURE
In common with other institutions, a newly-
opened iish culture research station always finds
itself short of the relevant reference works. This
is as true in the botanical department as elsewhere.
Many essential books are scarce or out of print,
while others are so expensive as to be beyond
the department budget. It is vitally necessary,
particularly when describing a new species, that
the botanist should have all the available reference
works at hand. The lack of this is seen in the
appalling number of synonyms for many organ-
isms. Photo copying seems to be the answer to
this lack of relevant literature, and it can take
several forms. Photostats are large and easily
read, but they are comparatively expensive and
are better bound for storing. Microfilms are
cheaper but necessitate a special reader and are
troublesome to store, and it is not always easy to
refer to a page and illustrations at the same time.
Microcards, where they have been printed, are
much better, and are ideally used with a special
reader but are quite easy to read with a dissecting
microscope. Furthermore they are black print
on white background, and are easy to store in
56
filing cabinets, being the same size as a filing card.
In cases where microcards have not been made,
microtape can be made from an existing micro-
film and gummed on to a filing card to make what
is in effect a microcard. The extra cost is very
little. All these methods may be beyond the
budgetary limit when numerous large works have
to be copied.
COLLABORATION OF
WORLD AUTHORITIES
Lack of literature at hand means that either
we must borrow, or else send material elsewhere
for identification. With the macrophytes, par-
ticularly the flowering plants, good herbarium
specimens should present little difficulty. With
the microscopic plants, however, especially the
algae, numerous special problems arise. Drawings
of the organism, in as many aspects as possible,
or good photographs are often valuable, but
there is always the danger that the artist's inter-
pretation is not strictly accurate, and a photo-
graph always lacks depth of focus. Nevertheless,
in the absence of the actual organisms themselves,
these can be a very good second best.
The despatch of actual material is a little more
difficult and entails special methods. The usual
method of preserving algae is in 5% formalin,
and for things like desmids or diatoms, where
the structure of the cell wall is important, or for
the Chlorococcales, it seems perfectly adequate.
For cell contents such as in Spirogyra, it often
causes distortion, while the more delicate flagel-
lates may become almost unrecognisable, and
colonial forms may break up. In such cases
Picric acid, or one of the compound preservatives
might be better and it would be necessary to
experiment. The flagellates in particular are a
difficult problem as they so easily become un-
recognisable.
Permanent slides suggest themselves as an
answer, and they are particularly important for
the preservation of holotypes. The mounting of
the frustules of diatoms is too well known to cause
much concern. With other algal groups it is not
so easy. The serial dehydration necessary for
many mountant fluids often leads to the distortion
or complete loss of the organism concerned.
VOLUME 4
BOTANY
A direct mountant is indicated, and in this respect
I have found Polyvinyl alcohol very useful. It
both fixes and mounts the living organism and can
be diluted with water to the required strength to
prevent plasmolysis. It will set hard, although in
the wet Malayan climate I have found it better to
encourage its drying-out in a desiccator. Its
main disadvantage is that objects become so
transparent in it, and it may be necessary to add
a dye. Glycerine mounts, while easily prepared
and useful, are too fragile to send by post. More
experiments are obviously needed to find the best
permanent mountants for the algae.
The ideal way would be to send living material,
for then it could be examined in the living state
and even cultured. Naturally this presupposes
despatch by air, and if it is possible to send the
material direct from airport to airport without
intervening postal delays, so much the better.
The problem is to keep the organisms alive and
in reasonable condition throughout the journey.
Drastic variations in temperature are nearly
always harmful to algae, and these are just what
they seem to be subjected to when carried from
country to country. To overcome that, it would
be necessary to insulate the container as much as
possible, or even enclose it in a thermos flask,
with consequent increase in air-charges. As
containers for the water samples, 1 have found
polythene bags very useful, as there is little danger
of them breaking, and several of them could be
packed inside a single larger container. Another
problem is bacterial action, which can become
quite serious in the time taken for the sample to
reach its destination. Field collections of water
samples inevitably contain some bacteria, and
these are certain to multiply. Perhaps the careful
use of one of the antibiotics may overcome this
difficulty. A further serious difficulty is oxygen
deficiency in the water. It must be remembered
that the algal specimens will be travelling in the
dark, and therefore any oxygen used up in their
respiration cannot be replaced by that due to
photosynthesis. Furthermore, in natural field
collections there are bound to be a number of
small animals which will further deplete the
oxygen supply. The tubes or polythene bags
should not be more than half full, and any dense
suspensions should be diluted with well oxy-
genated water. Living samples have been suc-
cessfully sent over long distances, but they have
usually been pure cultures with very few organisms
in the container. Thus if a pure culture, partic-
ularly bacteria- free, could be maintained, it
would be no difficulty to put a minute drop in
\ tube of sterile culture solution. The estab-
lishment of such cultures, while essentially part
of the work of a fresh-water fish culture research
station, is outside the scope of this paper.
The foregoing should give some idea of the
problems which occur in such a specialized
botanical department as ours at Malacca, and
I have no doubt there are many other such de-
partments with similar problems. I have indi-
cated some of the possible ways of solving these
problems, but I am hoping to gain much informa-
tion and advice from the discussion which will
follow.
DISCUSSION
E.H. WALKER: Some of the problems are bibliograph-
ical, and I hope, as I have mentioned from time to time,
you will support the meeting on this subject.
M.S. DOTY: Polythene bags are permeable to air,
although not to water. The amount of water may be
profitably reduced until it forms only a thin film which can
absorb the maximum oxygen.
G.A. PROWSE: Desmids and flagellates are the most
difficult.
M.S. DOTY: Some fixatives are of value.
E.H. WALKER: Cooperation between institutes and
airlines are needed to improve transit facilities.
H.M. BURKILL: The use of the diplomatic bag has been
advantageous in difficult cases.
E.H. WALKER : We need scientific attaches.
J.H. HORLIMANN: They could help in the numerous
quarantine difficulties.
p. WEATHERWAX: Soil samples are often unacceptable
to the health authorities, but they can be sent through with
cooperation. What are conditions like in the freight holds
of aircrafts?
M.S. DOTY: Specimens must go in the pressurized part
of the plane; the freight holds are freezing. Packages
should be clearly labelled to this effect, and, if possible,
be put in the hands of the second officer of the aircraft,
who can also reduce delays or unfavorable storage on the
ground at intermediate stops.
57
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
THE VARIOUS TECHNICAL ASPECTS
OF BOTANICAL RESEARCH IN NEW CALEDONIAt
ROBERT VIROT
Museum National (VHisloire Naturellc, Paris, France.
This note aims essentially at suggesting a
few measures which would enable the conditions
of botanical research within our territory of the
Southwest Pacific Islands to be improved.
SYSTEMATICS
In this field, efforts should be directed, above
all, towards enabling the investigator who is not
always an expert to have the means of making
a rapid and exact determination of the specimens
he has gathered or, inversely, towards providing
him with a precise representation of the morphol-
ogy of the species he wishes to obtain.
In this connection, it should first of all be made
clear that no adequate reference herbarium
exists on the spot at present. The material which
can be utilized is limited, in this case, to fragmen-
tary collections provided exclusively by metro-
politan and foreign botanists. The investigators
thus find that they arc almost always obliged to
have the necessary determinations made, or
confirmed, by the competent services of the
Natural History Museum in Paris, this leading to
a considerable loss of time. As a palliative to
this difficulty, the following measures would
appear to be highly recommendable:
a) the drawing up of several basic herbaria,
as complete as possible, centralised in Nou-
men at the seat of the scientific and adminis-
trative bodies who are the most directly
interested: Institut Frangais d'Oceanie,
Commission du Pacifique Sud, Service des
Eaux et Forets, Service de T Agriculture,
Mus<e no-caledonien. Botanists who are
collectors "de passage" should always under-
take to enrich these herbaria by the deposit
of duplicates; and where only one specimen
of a new species is gathered, this should be
provisionally represented by a full-sized
photographic reproduction;
b) the opening, for each species, of an identical
file, the number of these corresponding to
the basic herbarium and which will contain,
in addition to the classical elements, a very
detailed morphological description, a syn-
t Presented by J.H. Hiirlimann.
58
onymy, vernacular names, general, chorolo-
gy, list of the known neo-caledonian locali-
ties, ecology, etc. A photographic documen-
tation carried out in the natural state and
referring not only to the leguminous aspect
as a whole (port), but also to details: leaves,
flowers, fruits.
Another problem, of primary importance, is
that of a practical bibliography. The only flora
published up to the present- that of A. Guillau-
min (1948) was intended by the author as a
preliminary work only, destined to meet imme-
diate needs. Consisting of synoptic tables only
and totally deprived of illustrations, it is accessi-
ble only to investigators who are already well-
informed. In addition, the rigid framework of its
presentation does not lend itself easily to am-
plifications, a disadvantage which is particularly
inconvenient when one realises that there still
remains a great deal to be done before the floristic
inventory of New Caledonia is complete. In our
opinion, the ideal formula to adopt is to be
found in the drawing up of a flora carried out on
the principles which inspired O. Degener for the
publication of his New Illustrated Flora of the
Hawaiian Islands: a perforated sheet movable
for the description of each species, the recto
being reserved for the text (diagnosis, ecology,
chorology, etc.) and the verso for several sketches
of their features. It is clear that such an arrange-
ment would allow for the easy insertion of addi-
tional sheets concerning new species in the same
way as it would allow for the replacement of
obsolete keys of determination by up-to-date
documentation.
RESEARCHES ON THE TERRAIN
Cartography. For the moment, the topograph-
ical Services of New Caledonia put two types of
terrestrial maps at the disposal of the public:
1:40,000 maps and 1:10,000 maps. Utilized
specially for the delimination of mining and
agricultural concessions, the 1 :40,000 maps cover
only a fraction of the area of the Grande Terre.
Further, their planimetric representation is
VOLUME 4
BOTANY
simplified in the extreme. Indeed, we find only
the contours of the coast and the waterways and
roads, with a few summary particulars with regard
to swamps, the European settlements, the large
forest areas, etc. As regards the relief, dotted
lines indicate the position of the principal ridges
and the peaks mentioned are mostly defined, not
by their altitude, but by their order number with-
in the triangulation framework. Although the
1:10,000 maps in curves are of course more
exact, they exist only in scattered sheets and
cannot therefore serve for purposes of general
study.
However, all these gaps are shortly to be satis-
factorily solved, thanks to the publication of a
new map under the auspices of the Institute
Geographique National. Drawn up from aerial
surveys, this map, the final scale of which is not
yet fixed (1 :50,000 or 1 : 1,000,000), will cover not
only the Grande Terre as a whole but also the
islands geographically dependant on it: Pine
island, the Loyalty group, etc.
We would also mention the excellent geological
map 1:100,000 in 10 sheets (Grande Terre and
Pine Island) drawn up following on the work of
the mission led by P. Routhier from 1946 to
1949. Already partially on sale, within the
programme of publications of the Office de la
Recherche Scientifique et Technique Outre-Mer,
it represents, a considerable advance compared
with our previous knowledge in the subject and
will be of inestimable service to the ecologist.
Ecology. While the equipment of the pedolog-
ical section of the French Institute of the Pacific
Islands now enables very complete analyses of
soils to be carried out, nothing positive has as
yet been accomplished in the field of micro-cli-
matological researches. The local National
Meteorological Service whose observation sta-
tions are, incidentally, almost all placed in the low-
lying regions near the shores directs its activities
mainly towards objectives that are primarily
practical: definitions of the characteristics of the
general climate, weather forecasts with a view to
the improvement of the conditions of air and
maritime traffic, etc.
The accelerated developments of the ecological
discipline requiring data which are more and more
exact, from now on the serious study of the
dynamics of the leguminous groups of New
Caledonia, cannot be envisaged without the
support of more solid arguments than the spora-
dic measurements carried out up to now by na-
turalists in the course of their movements. In
the United States, for instance, the use of auto-
matic and portable micro-climatological stations
placed within the various strata of given groups
has already given very encouraging results.
Noted every week, the tapes of the registering
instruments thus constitute homogeneous series
of continued observations. This technique, it
seems to us, should find a perfect application
within the limits of our Southwest Pacific terri-
tory, more particularly within the upper zones
of the mountainous regions where it is not always
easy to remain for sufficiently long periods.
Transport. At the present time the roads ac-
cessible to motor traffic are limited to two arterial
roads running along the East and West Coasts
and communicating with one another by three
or four transversal roads. In very many cases this
arrangement hinders, in particular, the rapid
approach to areas of work situated in the moun-
tain groups. Very often, long and fatiguing
distances over extremely varied ground have to
be covered on foot in order to reach the goal
which has been set. During the ascension, the
investigator, obliged to mobilise all his physical
forces in overcoming natural obstacles, can devote
only a small part of his intellectual faculties to
detailed scientific observation. Heavy rains or
intense sunshine often add to this depressing
picture.
In the absence of suitable landing ground,
the light airplane would not be suitable to radical-
ly overcome these difficulties, but the helicopter,
to which the problem of a large landing ground
does not apply, certainly represents the solution
for the future. Incidentally, one of these helicop-
ters used experimentally by a local industrial com-
pany for mining prospection has, up to the
present, given entire satisfaction. Thus put down
in the area of work without any previous fatigue,
the investigator finds himself able to give of his
very best during the time he remains on the spot
and, afterwards, also during the descent towards
the lower lying regions.
PHYTO-SOCIOLOGY AND A MAP OF
LEGUMINOUS GROUPS
The transport difficulties of which we have
just spoken have very considerably retarded the
inventory of the neo-Caledonian vegetation.
Indeed, great progress still remains to be made in
the knowledge of nature and the distribution of
the leguminous groups. In order to attain this
end, it will be necessary to call in the assistance of
aerial photography on a large scale and in colour.
Personally, in the course of flights at low altitude,
59
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
we noted the clearness with which certain masses
physiognomically very individualised, such as
the Mangrove, the herbaceous belts of the sandy
beaches, and the photophileous forest of Melaleuca
Leucadendron, were to be distinguished, thanks
to their morphology and their colouration. Of
course, a general map, made up from an assem-
bly of aerial cliches, does not in any case do away
with the classic analytical surveys made on the
spot; but, judiciously interpreted, it possesses the
enormous advantage of enabling us to give the
precise localisation of the large laguminous for-
mations and, as a consequence, to rationally
coordinate the researches on the ground.
DISCUSSION
M.S. DOTY: I fear helicopters may prove out of the
question; it costs US$400 to fly one for an hour.
J.H. HURLIMANN: I admit we may not be able to realize
all our hopes, but the possibility is noted.
60
VOLUME 4
BOTANY
NOTES ON THE TECHNIQUES EMPLOYED FOR THE COLLECTION
OF BOTANICAL SPECIMENS IN PAPUA AND NEW GUINEAt
J.S. WOMERSLEY
Division of Botany, Department of Forests, LAE.
The collection of botanical specimens in
tropical countries has always presented a number
of problems not faced by collectors in temperate
latitudes. The normally high humidity combined
with temperatures usually exceeding a minimum
of 75 degrees Fahrenheit bring with them the
problem of rapid fungal and bacterial growth.
A large number of tropical plants, particularly
those of the rain forests, develop abscission tissue
with extreme rapidity in the leaf petioles and
petiolules. An inspection in any herbaria of the
specimens of tropical plants will reveal how
widespread this is.
Various botanists in the tropical woHd have
met these problems in a variety of ways. Eleven
years experience in Papua and New Guinea has
produced the techniques and procedures which
will be described below, but before doing so it is
necessary to give some background regarding the
nature of the country, its people, and tran-
sportation.
In New Guinea, botanical collecting has been
and still largely is associated with geographical
exploration. As yet little has been done toward
the exhaustive compilation of the flora of a
limited area. The botanist and his equipment
must therefore be mobile. Road transport is
available to a limited but rapidly increasing
extent. Air transport particularly by light aircraft
carrying loads up to 1,200 pounds weight is
widely available. Water transport descending
in size from coastal vessels of about 300 tons to
canoes is generally available in coastal areas.
Canoes and small launches may be used to some
extent on most lowland rivers. A few areas may
be reached by float plane, landing on lakes.
Using the most appropriate transport, the
botanist gets himself into the general area he
plans to visit. Further progress is on foot with
human porterage as transport for the collecting
equipment. Fortunately, in most areas the local
inhabitants are prepared to assist parties by
carrying equipment and supplies. The rate of
pay varies rather widely throughout New Guinea.
The maximum load is about 40 pounds per adult,
or where a load is carried jointly by two men,
t Presented by H.M. Burkill.
a total of 80 pounds weight. Equipment must
therefore be of a size and weight which allows it
to be made up into suitable sized bundles for
carrying.
The actual equipment used differs somewhat
from that of a botanist in temperate countries.
Most of the flora is woody so that an axe is
indispensable if justice is to be done with this
great group of plants. One is led to suspect that
at least as far as New Guinea is concerned the
difficulty of falling large trees has sometimes
turned the attention of botanists away from the
tree storey. Lane-Poole, when collecting in New
Guinea in the 1920's and being almost exclusively
concerned with the forest flora, used a rifle
successfully to bring down branches of forest
trees. A shot gun can also be used to good
advantage at times. Secateurs are extremely
useful for cutting the actual specimens, while
heavy gloves or a machete facilitate the collection
of thorny palms and trees.
As the specimen is finally prepared, usually
represents but a fragment of the original plant,
adequate descriptive field notes are essential.
The value of the collections is greatly enhanced
and frequently identification facilitated if wood
samples are collected. General practice in New
Guinea is to obtain a wood sample of all plants
having a woody stem. If this exceeds 6 inches in
diameter, the sample is usually trimmed to 15
inches long with a cross section of 4 inches by
4 inches. Bark is retained wherever possible.
Where weight is a serious consideration, the wood
sample may be reduced in size.
Turning to the specimens of foliage, every effort
has to be made to collect flowering and fruiting
material of each species. Most tropical herbaria
have a considerable quantity of sterile or inade-
quately fertile material which can be only partially
identified and must await the recollection of more
adequate material. This does not mean that
under certain circumstances such as a collector
in a "new" country or where a plant appears to
have some special interest, specimens lacking
flowers or fruit should not be collected. I can
repeat, though, that a wood sample in such cases
61
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
can be of immense value in identifying the
collection.
The technique adopted for preparing the
botanical specimens in Papua and New Guinea
is based on the formalin technique of Schultes,
used successfully by him in South America.
However certain modifications have been made.
At the base camp are required newsprint, folded,
but preferably clipped to 18 inches by 12 inches;
frames, 18 inches by 12 inches constructed as a
lattice from wood 1 inch by inch thick; a
rectangular tank, the inside dimensions of which
are 18-^ by 12} inches by 12 inches deep,
fitted with a lid; strong string; sisalkraft; (a water-
proof bituminous paper); and concentrated
formalin.
Actual collecting requires newspapers, frames
(or wire press), strong cord (leather straps
deteriorate too quickly), and labels. The speci-
mens, suitably pruned to display the essential
parts to best advantage, folded in the case of large
or compound leaves, are numbered by attaching
a number bearing manilla tag, the number on
the tag corresponding of course to the relevant
number in the field note book. Soft pencil or
crayon is most suitable for numbering on the tag.
Do not use ordinary ink. A pad of folded news-
paper is opened so that all but the lowest sheet is
folded on the left. The lowest sheet is reversed
so that the fold is on the right. The numbered
specimens of each collection are now placed
singly between the sheets. Wherever possible,
ten or more replicates of each collection should
be obtained. With this number it is convenient
to confine a single packet of paper to each col-
lection. The reversed bottom sheet is finally
folded around the packet thereby providing
closure at both edges. The packets are then
placed between a pair of wooden frames and
pressure exerted by means of several turns of
strong string at each end. A bundle not exceeding
twelve inches in thickness is desirable. These
bundles are then transported to the base camp.
The next step consists of preserving the speci-
mens from fungal and bacterial action by immers-
ing them in formaldehyde solution of 4% con-
centration by weight, specific gravity about 1.014
at 75 degrees fahrenheit. The formalin can be
prepared by diluting a concentrated formaldehyde
solution. As normally received in New Guinea,
this rarely exceeds 35% formaldehyde by weight
and should be broken down in the proportion of
one part of concentrate to eight parts of water.
An alternative procedure has recently been
adopted with success using the solubility of
62
paraformaldehyde in hot water made alkaline
with hexamine. This produces a formaldehyde
solution, the specific gravity of which is adjusted
to 1.014 after cooling. Solution can readily be
effected by adding about one ounce of hexamine
to three gallons of water which is heated to the
boiling point. After removing from the fire,
slowly stir in three pounds of powdered para-
formaldehyde. Due to the formaldehyde vapour
produced, this can be most unpleasant. An
alternative which works equally well is to place
the paraformaldehyde powder at the rate of one
pound per gallon in a one gallon, wide mounthed,
plastic, screw top bottle. The hot water, which
contains hexamine at the rate of one ounce to
three gallons of water, is then added to each
bottle, the lid screwed, on and the bottles shaken
intermittently until solution is complete. After
cooling, the specific gravity is adjusted as before.
The use of paraformaldehyde has obvious
advantages over concentrated formaline where
weight is a factor to be considered.
Having prepared the formalin solution, the
rectangular tank, the inside of which has been
coated with bitumastic paint, is half filled with
4% formaldehyde. The prepared bundles of
specimens referred to above are immersed, a
stone may be used to stop the bundle floating,
the lid put on and the whole lot left for 18-24
hours. The following day the bundle of speci-
mens is removed; the free liquid is drained off;
and it is securely wrapped in waterproof sisalkraft
paper. In this condition, specimens have been
kept without deterioration for three months.
The bundles of formalin preserved specimens
are sent by whatever transport is available to the
divisional headquarters at Lae. Here the bundles
are opened, the specimens placed between drying
pads which are interspersed with corrugates for
ventilation, and dried in cabinets heated with 150
watt globes. Drying is complete in 24-36 hours.
Where succulents or palms having a waxy epider-
mis are being collected, the addition of a liquid
detergent to the formalin at the rate of about one
tablespoonful to each gallon is advantageous in
accelerating the penetration of the leaf tissues by
the formalin. Probably no harm would accrue
from the general use of a little detergent with the
formalin.
The main attraction of this method as used
in New Guinea is that the collector does not have
to spend time while in the field in tending drying
ovens or racks. For the successful use of the
method, though, access to some central drying
facility is essential.
VOLUME 4
DISCUSSION
BOTANY
C.G.G.J. VAN STEENIS: JFor herbarium work, it is essen-
tial that the morphological characters of the specimen
should be well preserved. Dehiscence of leaves, leaflets, and
floral members must be prevented by rapid killing. Alcohol
and/or formalin are excellent, but whatever facilities are
available such as heat may be used if circumstances dictate.
I agree with Drs. Prowse and Walker, field characters must
be recorded at collection. Otherwise such valuable infor-
mation as scent, color, and glossiness will be denied to the
herbarium botanists who compile the floras for practical
use. It is essential that every duplicate specimen should
have a full copy of the original field label.
E.H. WALKER: The original field labels on the Manila
specimens were destroyed in the fire; the duplicates lack
this information, which is now permanently lost. Hence
the need, despite the labor involved, of providing every
duplicate specimen with a copy of the field notes.
63
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Symposium: Vegetation Types of the Pacific.
VEGETATION STUDY AND RECORDING!
PIERRE DANSEREAU
Institut Botanique, Universite de Montreal, Montreal, Canada.
Many methods are used for describing vege-
tation, although much of the available literature
is appallingly unmethodical in the sense that the
standards are poorly defined. No valid criticism
of method or approach, on the other hand, can
be made without an honest effort to evaluate
purpose and scope which are an ultimate measure
of adequacy.
This symposium on vegetation types of the
Pacific does not hope to provide a complete
inventory of all synecological work, even less an
accurate description of all known plant com-
munities. Research on vegetation has been very
widely scattered and many bioclimatically im-
portant areas have not been studied at all. Nor
was it possible to request a uniform approach
of all the participants in this joint effort. It has
seemed more useful to pick contributors who had
a good deal of field knowledge and who had
already made a contribution to the description
and interpretation of Pacific landscapes and to
give them a free rein in their own course. The
resulting papers arc sure to be somewhat hetero-
geneous, and their divergence in method will no
doubt be pointed out by Frank E. Egler.
It seems to me, however, that some reflections
on structure variation are an appropriate preface
to this presentation and that examples from
various parts of the Pacific can be usefully quoted
in view of subsequent correlation with the geolog-
ical, historical, climatic, dynamic and other
interpretations which are forthcoming. At pre-
vious congresses (New Zealand, 1949, Philippines,
1953), I have made similar contributions which by
now have been amply discussed with many
colleagues. I feel that a further step can be
undertaken at this time, and I will therefore
offer some elaborations upon the system which
I have previously proposed (1951, 1952, 1953,
1957a, 1957b).
THE FEATURES OF STRUCTURE
Spatial distribution of the biomass is a function
of life-form, size and coverage, which in turn
may vary seasonally (relative deciduousness) and
t Presented by F.E. Egler.
64
which may be caused by diverse assemblages of
leaf-type and texture classes. Table 1 repeats
my earlier attempts (1951, 1952, 1957a)to provide
a simple, practical scale for grading actual stands
of vegetation or for abstracting therefrom a
number of well-individualized types.
1 have previously discussed in some detail the
choice of the alternates presented under each of
the six criteria that appear in Table 1. But I now
find that systematic application by myself and by
others gives me cause to introduce certain modi-
fications or to suggest some alternatives. There-
fore Table 2 is offered as a new key to the system,
and Fig. 1 shows how the symbols (now simpli-
fied) can be plotted on squared paper (8 squares
high, 25 squares long).
These modifications improve the system in
a number of ways.
Life-form. Inasmuch as the former "trees"
and "shrubs" did not necessarily refer to plants
at the adult stage of their development but only
to woody individuals of a certain height, at a
certain time, it is best to designate by a single
symbol all such individuals. This leaves category 1
with only 5 instead of six alternatives. It will
also be noted that the symbols now proposed
for lianas (L) and epiphytes (E) are much easier
to draw.
Stratification. In substituting this heading for
"size," a more objective picture is conveyed,
and the sliding scale principle is better applied.
Thus, all symbols followed by the same figure
(1 to 7) will situate that life-form at the same
level (e.g. in the same layer) as all other life-forms
coupled to the same figure. This was not so
previously: e.g., Tt and Ht were 23 metres apart.
Now, the synusiae are well separated, and the
layers are united!
Figure 2 shows some types of vegetation that
had been previously plotted with the 1951 key.
Although this may be an improvement in practical
procedure, it still does not allow to show a certain
number of features (see 7, Table 8), which, for
certain purposes, it may be desired to illustrated.
Coverage. Therefore, some of these features
VOLUME 4
BOTANY
are added here and shown in Figs. 3, 4, and 5.
The two diagrams previously published (7, Figure
1 3) and now re-plotted show in their upper layer
a certain total crown coverage that could very
well be the result of different stem growths.
Inasmuch as the system so far has made little
provision for emphasizing the closeness of stems,
a new means is now offered to show this. Figs.
3, 4, and 5 outline two different approaches: one
by direct drawing of independent stems and
crowns (and modification of the crown symbols
from circles to ellipses) and the other by drawing
lines that show how many times closer the stems
are than the normal coverage symbol allows for.
A further refinement can be introduced to
represent the principal outlines of crowns as
indicated in Fig. 6. There would be no particular
merit in extending such categories to herbs, lianas,
epiphytes or bryoids, inasmuch as leaf type
pretty well determines outline anyway.
Of course there always remains another alter-
native, which would consist in prolonging the
strip many times: 50, 75, 100 or more squares,
instead of 25, would allow an almost literal repre-
sentation of the principal life-form types along this
linear transect.
Function. Relative deciduousness (or leafing
periodicity) is of course quite important for it
affects volume at different seasons. But there is
another item of great importance to vegetation
analysis, the dispersal function. This has been
discussed at some length by Dansereau and Lems
(6) . It is very remarkable that the different layers
show such marked contrasts in the distribution
of, for instance, wind- and animal-dispersed
species.
In a recent paper, Keay (8) analyzes a second-
growth forest in Nigeria where he finds that wind-
disseminated tree species are dominant. On the
contrary, in the Canary Island laurel forest,
bird-dissemination is the rule at the tree level.
In the temperate forests of the Pacific, a very large
number of trees are dispersed by wind: from
Alaska to northern California, the tallest members
of the needle-leaved forest (Picea, Tsuga, Sequoia,
Pinus) have very light diaspores, whereas a greater
number of the shrubs and herbs have fleshy
fruits. The Mediterranean area of California,
on the other hand, has a vast array of oaks and
some fleshy-fruited trees like Arbutus. A similar
situation obtains in New Zealand, with the
northern forest rich in fleshy fruits, the southern
rich in nuts and small winged diaspores.
CORRELATIONS OF STRUCTURE
Although this system was devised essentially,
in fact exclusively, in order to show the features
of vegetation and later and otherwise relate them
to environmental elements (causal or not), in
actual recording it is found useful to plot a certain
number of the latter, for instance site conditions.
Table 3 offers a small repertory of the usual
alternatives: climate, relief, qualities of soil,
land use.
THE MAPPING OF STRUCTURE
The problems of mapping are very numerous
and each scale presents its own difficulties. No
easy rules can be established (and surely none are
explicitly recognized) when it comes to stripping
a large-scale map (say 1 inch = 1 mile) of its
detail if the information is to be relayed to a
smaller scale (for instance, 1 inch = 8 miles). As
a matter of fact, it is rarely a matter of omitting
detail; it is more likely a matter of interpreting
the units of the large-scale map so as to lump
them into units of a greater order of magnitude.
This involves a judgment, something which is not
strictly a matter of record or observation, of
compilation of features that already lie on the
map itself, but something that is essentially classi-
ficatory. An example of this is provided by the
excellent maps of the Toulouse group, for instance
Gaussen's (J) Perpignan sheet. The colours
represent each a "series" (we would say a sere):
a number of plant communities, usually of in-
creasingly complex structure, which lead to a
terminal stage. 1 The intensity of the colour, on
the other hand, does indicate changes of structure.
This procedure appears to me worthy of standard-
ization and lends itself to a very useful procedure.
Thus, for instance, the kauri forest area of New
Zealand or the highland pine region of the Island
of Luzon (Philippines) in their developmental
stages from grassland or scrub to mature forest
could be represented by a series of structure
diagrams, and on a map by the corresponding
colours.
Schmid (9) had already recognized the desir-
ability of producing structure diagrams in the
margin of vegetation maps in order to show the
range of variation which must of all necessity be
involved in any one category of vegetation type
[except on a very large-scale map which really
attempts to show the exact status of the plant-
l This is generally the climatic climax although occasionally an edaphic or topographic climax. See ( 3) for a fuller discussion
of this topic.
65
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
cover (Pflanzendecke) at one time!].
It is hoped that the present schemes lend them-
selves to fairly wide application and can be used
to reveal at a glance the many-sided aspects of
vegetation types that play an important role in
the landscape. It may well remain to demonstrate
to the biologist and, to the paleontologist what
the historical and physiological conditioning of
these types really is. It is granted, of course, that
such considerations are fundamental, but the fact
that they are virtually disregarded in the present
instance, should not disguise the urgency of a
better comparative understanding of the
functional aspect and distribution of vegetation
types as they are to be observed in the living
landscape at present.
REFERENCES
(1) Dansereau, Pierre, 1951, Description and
recording of vegetation upon a structural
basis, Ecology, 32 (2): 172-229.
(2) Dansereau, Pierre, 1953, Structural units
of vegetation in tropical and temperate
climates with special reference to Pacific
areas, Proc. Seventh Pac. Sci. Congr.,
5:100-112.
(3) Dansereau, Pierre, 1956, Le regime clima-
tique regional de la vegetation et les
controlcs edaphiques, Rev. Canad. Bio/.,
(4) Dansereau, Pierre, 1957a, Biogcography : An
ecological perspective. The Ronald
Press Co., New York, xii + 394 pp.
(5) Dansereau, Pierre, 1957b, A preliminary
note on the structure variations of
temperate rain forest, Proc. Eighth Pac.
Sci. Congr., 4 (Botany): 407-436.
(6) Dansereau, Pierre and Lems, Kornelius,
1957, The grading of dispersal types in
plant communities and their ecological
significance, Contrib. Jnst. Rot. Univ.
Montreal, 71 (in press).
(7) Gaussen, H., 1946, Feuille No. 78: Perpig-
nan. In: Carte de la Vegetation de la
France (1 :250,000). Centre National de
la Recherche Scientifique, Paris.
(8) Keay, R.W.J., 1957, Wind dispersed species
in a Nigerian forest, Journ. EcoL, 45(2):
471-478.
(9) Schmid, Emil, 1954, Anleitung zu Veget-
ationsaufnahmen. Vierteljahrsschrift
der Naturforschenden Gesellschaft in
Zurich, 1C (1954), 1,37pp.
66
Six categories of criteria to be applied
1. LIFE-FORM
T
9
trees
F
9
shrubs
H
V
herbs
M
CD
bryoids
E
epiphytes
L
^
lianas
VOLUME 4
Table 1.
BOTANY
2. SIZE
t tall (T: minimum 25 m)
(F: 2-8 m)
(H: minimum 2 m)
m medium (T: 10-25 m)
(F,H: 0.5-2 m)
(M: minimum 10 cm)
1 low (T: 8-10 m)
(F,H: maximum 50 cm)
(M: maximum 10 cm)
3. COVERAGE
b barren or very sparse
i discontinuous
p in tufts or groups
c continuous
4. FUNCTION
I I deciduous
1 1 1 1 semideciduous
"|| 1 1 evergreen
evergreen-succulent ;
or evergreen-leafless
5. LEAF SHAPE AND SIZE
n <^> needle or spine
g A graminoid
medium or smal
broad
compound
V
P O thalloid
6. LEAF TEXTURE
sclerophyll
succulent; or fungoid
67
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Table 2.
A revised scheme of the six categories of criteria to be applied to a structural description ol
vegetation types.
1. LIFE- FORM
4. FUNCTION
w
L
E
H
M
9
a
^
V
Q
erect woody plants
climbing or decumbent woody s 1
i
plants
e :
N
epiphytes
j j
&
herbs
bryoides
2. STRATIFICATION
1 more than 25 metres
2 10-25 metres
3 8- 10 metres
4 2-8 metres
5 0.50-2 metres
6 0.10-0.50 metres
7 0.0-0. 10 metres
3. COVERAGE
b barren or very sparse
i interrupted, discontinuous
p in patches, tufts, clumps
c continuous
68
deciduous
semideciduous
evergreen
evergreen- succulent ;
or evergreen-leafless
5. LEAF SHAPE AND SIZE
o
v V
P o
needle or spine
graminoid
medium or small
broad
compound
thalloid
6. LEAF TEXTURE
x
k
filmy
| | membranous
sclerophyll
succulent; or fungoid
a. UTILIZATION
virgin
pastured
lumbered
harvested
ploughed
burned
VOLUME 4
Table 3.
Categories and symbols for recording site conditions.
d. DRAINAGE
BOTANY
lUUUl
L/VNAl
b. SOIL STRUCTURE
soft
medium
hard
c. SOIL TEXTURE
(A, B, C horizons)
bedrock | V V y"
boulders
pebbles
sand
silt
clay
organic
mmm
excessive
good
deficient
e. RELIEF
flat
depressed
rolling
abrupt
f. EXPOSURE
use symbols of compass,
N, S, E, W
e.g.: NW
g. CLIMATE
use Koppen symbols
e.g.: Dfc
69
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
v^
r' C
f C
p
c
p
? ? c
2
3
4
5
6
7
Wl W2 W3
W4
W5 W6 W7
LI
AE,
L2
a
L3
CD
L4
A E3
E4
CD
L5 CD
L6
E5
A E6
L7
E7
n
n
H4 H5 H6 H7 M6 M7
Fig. 1. A graphic representation of all the symbols combining criteria 1 and 2 of Table 2. This is a new scheme, super-
seding the one proposed in 1951.
70
VOLUME 4
BOTANY
Aceretum rubri Wldhze(ozb) WZdhzi W3dhzp W4dhzi
WGdhzi(azb) HGdhzb W7dazb H7dazb
M7enxp
Betuletum populifoliae W2dazi W3do(h)zb W4dhzb
W6dazb H6dvzb H7dg(o,v)zp
M7enxp
Fig. 2. A red maple stand and a wire-birch stand plotted according to the new scheme (Table 2 and Fig. I). (Compare
with Figs. 13 and 14 of the 1951 version.)
71
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
B
Fig 3. The upper la>cr of the red maple stand (see Fig. 2), showing different distribution of stem and crown spaces cor-
responding to identical coverage and illustrating two ways in which the stand can be plotted.
72
VOLUME 4
BOTANY
B
000000 000000 00 00
Fig. 4. The upper layer of the wire-birch stand of Fig. 2, illustrating variations in stem within an identical total crown
coverage and two ways in which these differences can be plotted.
73
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
B
000X00
Fig. 5. Another variant of crown-stem distribution in a wire-birch stand (see Figs. 2 and 4).
74
VOLUME 4
BOTANY
Fig. 6. A series of crown outlines for tall woody types (W 1, 2, 3), which can fit the perimeter of the symbols in Fig. 1.
75
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
GRADING AND INTEGRATION OF EPIPHYTE COMMUNITIES!
T. HOSOKAWA, M. OMURA, and Y. NISHIHARA
Department of Biology, Kyushu University ', Fukuoka> Japan.
INTRODUCTION
It is possible to say that there may be consider-
able subjectivity in classifying vegetation, even if
we adopt the abstract method of Ziirich-Montpel-
lier's school, attaching great importance to
characteristic species, which is commonly em-
ployed in continental Europe. We have a strong
urge to find some more objective method of
classifying vegetation. Fortunately, our attention
was called to Gpodall's objective method (I)
for the classification of vegetation. This method
is based on positive interspecific correlation of
major component species by making use of the
values of their frequency. We modified his
method to some degree (I), and used it and
Sorensen's method (5) for classifying epiphyte
vegetation growing on beech trees in Mt. Hiko,
Southwest Japan.
METHODS
The concept on which the modified method of
classifying vegetation is founded is that if there is
neither species of positive nor of negative inter-
Specific correlation of significance level in any
given plant group, such a group is conceived of
as being of homogeneous construction. Then,
the epiphyte vegetation is classified objectively
by four procedures according to this method of
consideration, practically almost the same as
Goodall's methods (I) of separating groups.
(a) Procedure I. The species of higher fre-
quency, as well as of positive interspecific correla-
tion of P< 0.001 level of significance were used
for separating groups by excluding all quadrats
in which the single correlated species did not
occur, until this particular species was no longer
present in a given group. One of the final groups
of residue was considered in its possible combina-
tion with another group, if there was not recog-
nized any species of positive interspecific correla-
tion which reached a high significance level in a
newly combined group. We expect, however,
that there may exist some species of negative
interspecific correlation of P< 0.001 or 0.0 1<
P< 0.001 level of significance even in such a final
group conceived to be certainly homogeneous as
judged by positive interspecific correlations of
significance level.
(b) Procedure II. The species of lower fre-
quency and simultaneously of positive inter-
specific correlation of a high significance level
which is quite the same as Procedure I, were used
for separating groups by excluding all quadrats in
which the single correlated species occurred, as
well as by excluding those in which it did not
occur (Procedure I). Recombination of the final
groups of residue was made by the same method
as Procedure I.
(c) Procedure III. As a substitute for positive
interspecific correlation, the species of higher
frequency and of negative interspecific correla-
tion of high significance level were used for sepa-
rating groups like Procedure I. In this case too,
we may expect some species of positive inter-
specific correlation of the same level of signific-
ance remaining in the final groups.
(d) Procedure IV. As a substitute for the
species of higher frequency, we used those of
lower ones for separating groups. The treatment
is quite the same as Procedure II.
Procedure I is clearly the best one for classifying
the vegetation of epiphytes, as shown in Table 1 ,
notwithstanding the difference that exists among
the three series of methods different in quadrat
size and number found in random sampling on
five beech trees: one standing at the peak, Kita-
dake, of 1,150 m alt.; two at the ridges of 925 m
and 930 m alt. ; and the other two at the moun-
tain-slopes of 910 m and 960 m alt.
The smaller the quadrat size, the more numer-
ous the groups become. Fig. 1 shows the simi-
larity of correlation among all the groups recog-
nized in the three series of groupings by means of
the sampling data of the different kinds of quadrat
size. Each group is designated by the names of
two important species which bear the highest and
the next best figures of positive indicator value.
However, if there are only a very few or not any
species having positive indicator values of signi-
ficance level in a group, we indicate it by enclosing
the name of the species not reaching significance
t Presented by F.E. Egler.
76
Table 1.
Success of different procedures in dividing data,
which were obtained from different sampling
sizes of quadrats, into homogeneous groups.
f
CO
S'
r
O
No. of interspecific
correlation
I
1
{.
ce
i
with P*
Q.
2
1
s
J
o.oi-aooi
< 0.001
I
7
1
-*
01
n
7
2
2
O
i
4
24
IB
3
IV
S
7
2
i
9
3
ho
O
n
7
2
1
00
O
s>
in
6
10
8
*
TV
7
4
2
i
10
3
2
^
n
1 f
2
6
Nd
CM
J5
m
8
5
8
r>
<
w
6
4
6
* The table shows the number of Positive (in Procedures III & IV)
and negative (in Procedures I & 1J) interspecific correlation of sig-
nificance level in the final groups.
level in parenthesis within a rectangular frame of
broken line. The degree of independence of every
epiphyte-group classified in these ways is ex-
pressed by the species number of positive (in
Roman type) and negative (in italics) indicator
values (1, 2) of significance level (P < 0.01),
which are shown beneath the rectangular frame
(Fig. 1) in which is written the group name.
The similarity of correlation among the groups,
depending on the three simultaneous series, is
VOLUME 4 BOTANY
expressed by using the "Quotient of Similarity"
(=QS, 5). In Fig. 1, the highest value of this
Quotient between one group of one series and a
group of the other series is shown near the con-
nected line between groups, and the correlations
between them are shown by various forms of
lines. Only the major connection-lines of simi-
larity, excluding minor ones, are shown. In
comparing groups between two series with each
other, to which both groups belong, the thick line
indicates the closest correlation where the highest
similarity is recognized between the two groups.
The thick broken~lines having a triangular arrow-
head indicate that a given group from which the
arrow-headed broken line started has the highest
value of QS correlating with the group indicated
within those of the other series. According to QS,
especially the groups centering around those
having the highest values of QS, as shown in
Fig. 1, we may regroup the epiphyte groups of
each series into four group-types.
Of every group in the series (Series 1) which
resulted from the Procedure I of arranging the
data obtained from the smallest size of quadrat
(500 sq cm each), Table 2 shows the ecological
distribution concerning vertical position and expo-
sure on trees and topographical positions of host
trees. The state of ecological distribution of
every group in the other two series, both Series 2
and 3 obtained from Procedure I, which resulted
from the data of larger quadrat (1,000 sq cm
Series 2, and 1,500 sq cm Series 3) is not so
different from that of the smallest sampling series
(Series 1) shown in Table 2.
As shown in Fig. 1, in each series, several
somewhat homogeneous groups were recognized
as the result of objective classification of epiphyte
vegetation in terms of interspecific correlation,
while there were some groups which had few
species of positive indicator value of significance
level. In order to make clear the correlational
similarity of floristic composition among the
independent groups of each separate series,
Sorensen's grouping method (5) based on the
value of QS was performed. An outline of this
method is shown in Fig. 2 and Table 3. For
example, we will explain the method of integra-
tion of those epiphyte groups in the Series 1 de-
rived from sampling data of the smallest quadrat
(See Table 3 and Fig. 2). The order of arrange-
ment of the epiphyte groups was the same as that
(Series 1 ) of Fig. 1 . At the level of QS = 40 (Table
3 and Fig. 2), the Homalia japonica Homalio-
dendron scalpellifolium group (Group 1) and the
Thuidium cymbifolium - Homaliodendron scalpel-
77
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Series 5
Series 2
Scries I
Series 3
~ ~~ ^ * - "~ "*
_ 3 3 -^-\
Fig. 1. Classificatory schema of epiphyte communities, depending on Procedure I on the basis of the data obtained from
the three kinds of sampling sizes of quadrats (Series 1 means the grouping resulted from 423 quadrats of 500 sq cm; Series 2,
that of 282 quadrats of 1,000 sq cm; and Series 3, that of 150 quadrats of 1,500 sq cm). The numerical figure written in
Roman type beneath every rectangular frame in which is shown the group name, indicates the number of species which
has the positive indicator value of significance level in that corresponding group, and that of italic type is of negative indicator
value. The numerical figures situated near a connected line indicate the value of Quotient of Similarity between the groups
connected with the line.
lifolium group (Group 5) were unified (QS = 42)
into one group (Group 2), and at the same time
Groups 3 and 4 (QS = 41) were combined into one
group (Group c) ; at the QS = 30 level, the Group b
and e (QS = 32) into the Group B, and the Group c
and d (QS = 33) into the Group C; and so on.
The number of groups differs to some degree at
78
different levels. Noting Fig. 1 and Table 3, we may
say that the group which has only a few species
bearing indicator value of significance level among
the component species, correlates with the other
groups by the lower value of QS. Though the
number of groups differ much in the different
series which differ from each other by the size of
VOLUME 4
Table 2.
BOTANY
Showing the state of the ecological distribution of epiphyte groups by the numbers of quadrats.
Vertical distribution on trees: trunk-bases (Tb), trunks (T), lower parts of crowns (Cb), interior parts
of crowns (C), and top-most parts of crowns (Ct). The distributed side on trees: open side (O),
intermediate side (M), and sheltered side (S). Distribution related to topography, viz., topographical
distribution of host trees on which the epiphytes grow: a beech tree standing at the peak, Kitadake,
of 1,150 m (Fj), at the ridge of 930 m (F 2 ) and 920 m (F 3 ), and at the slope of 910 m (F 4 ) and 960 m (F 5 )
alt. The figures having a thick underline indicate the predominant position of a given group.
O
i
T3
f
r&
^xHdbitat
Group ^x.
Vertical distribution
on trees
Distributed
side
on trers
Uistnbutioa
related to
topography
Tb
T
Cb
c
a
M
S
Fi
F 2
F3
F4
Fs
I
Parmelia loeirior
Uhtd uispula
2
9
13
20
1
3
5
7
5
3
4
Diiranolpmd tlagiliformc
MetzQerid conjugate
2
3
A
12
7
9
8
1 1
13
8
2
4
1
Graphic sp.
Cetrdsia colldta
7
3
6
8
1
12
7
6
6
17
1
1
n
Bouldyd mitteii
An z id. japonic a
6
32
29
4
34
24
13
15
16
20
9
1 1
Ortiwdicranum kaktodeose
Bouldyd mite nii
1
7
7
7
8
8
6
18
30
3
1
Ptirmelid homogcries
Frulldiud, monilidta
8
13
33
10
1
25
23
17
13
10
5
9
i
AnomotLon girdkUi
DolifomHm cymbffolid
4
57
17
S
4
23
36
3
10
12
27
1 1
forsst/vcmid ( ryptuxvides
Porclld irentisia
6
MVMM
14
48
6
17
33
23
15
15
15
18
23
i
3
i
fhuidium. {y/nbifol/w*
HomdiodtAomn siidpeJIijolium
6
2
12
2
7
8
homalid idpo/uca
Hoindliodendnon <4djpcV(folium
26
8
9
12
13
1
2
19
92
12
HHIMWMB
75
Total no. of quadrat
68
131
lie
80
26
160
131
132
90
92
74
quadrat for drawing data from the epiphyte
vegetation, the larger the size of the quadrat, the
fewer the number of groups becomes. Notwith-
standing the difference of series, however, in any
case the epiphyte vegetation is classified roughly
into two groups (e.g., Fig. 2): one is of epiphytes
growing on trunks and trunk-bases, and the other
those of the interior parts of crowns and the top-
most parts of crowns.
At a certain level where we can determine the
value of QS, as we hoped, as a given criterion of
integration of epiphyte groups, we can unify
those groups into the higher rank of group at a
given level, as well as we can understand the
degree of similarity of floristic composition among
those groups. Moreover, it is certainly note-
worthy that the order of arrangement of those
groups (Figs. 1 and 2) which are naturally
arranged in the case of classifying vegetation,
such as their ordering positions from the top-
most parts of crowns down to the trunk-bases,
is quite in accord with the environmental gradient.
79
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Group QQ
farnel/a
U/ot<* crisp.
Dicntnolom* f. -
Metzgeri* nnj.
Grttphif //>. -
Cetrttrk* coi.
40
30
Boulaya. mitt -
AnziA
Btuya mitt.
k -
m*tcry.-
c JhuMium cymk -
i Horn* lie* Jc
e
20
ffl
10
8
Fig. 2. Series 1 of Sorensen's grouping method, in which the integration was performed at the foundation of final groups
which were recognized in terms of classifying the epiphyte vegetation through Procedure 1 on the basis of the data obtained
from 423 quadrats of 500 sq cm.
COMPARISON BETWEEN EPILIAS AND
THE GROUPS OF THE PRESENT
TREATMENT
In order to compare those groups resulting from
such methods of treatment with the five epilias
(3) of epiphyte communities which we have
studied and designated to the same epiphyte
vegetation in the beech forests of Mt. Hiko (4),
and moreover to make clear the correlation be-
tween those five epilias and the groups based on
the present treatment, we selected six correspond-
ing epiphyte-groups (the Group A, B, C, D, E,
80
and F in Figs. 2 and 3) at the level of QS = 32
(Fig. 2) in the series of sampling with the smallest
size of quadrat (Series 1). Comparison was made
between them by using the values of QS of each
one (Fig. 3). Here the correlational similarities
among those groups are clearly recognized.
Indication of the similarity between an epilia and
a group, which is shown by the lines and the
numerical figures of QS, was done in the same
way as in Fig. 1. What is written beneath each
epilia-name surrounded with a rectangular frame
is the number of the characteristic species, and
similarly that of the epiphyte-group name means
VOLUME^ BOTANY
Table 3~~~~
Showing the values of QS between the epiphyte groups at four different levels in Series 1, which
is sh
10.
8.
9.
4.
5.
6.
7.
2.
5.
1.
own
N
in Fig. 2.
d. b. e. d. c. g. f. h.
a,
b.
e.
d.
c.
3
f.
h.
i\
\
24
31
9
5
6
2
6
22
\\
32
12
12
14
5
6
12
21
\
\
\
14
13
10
4
7
18
24
26
\
33
21
11
II
22
28
23
41
\\
26
26
20
n
II
21
50
35
\\
21
12
7
4
to
12
15
14
\\
22
6
5
14
12
II
12
32
\
S
\
7
4
6
6
10
12
36
27
3
0.07
5
2
5
6
25
21
42]
10. 8. 8. 4. 3. 6. V. 2. 6. 1.
A.
F. D. E C. B. A.
the species number of the positive (in roman type)
and negative (italic) indicator value of significance
level. From these figures and the connected line,
the correlational correspondence between each
epilia and group is comprehensible to us. It is
possible that the epilia or group which has corre-
spondingly fewer characteristic species or fewer
species bearing significant indicator value among
the component species, in either case, does not
necessarily show any correlational correspondence
between such epilia and group.
Accordingly, because of the success of our
several years of varied studies of corticolous
vegetation in the beech forests of Mt. Hiko, South-
west Japan, we would draw the conclusion that the
following four distinct communities of epiphytes,
which are considerably developed there can be
recognized: the Thuidium cymbifolium-Homalio-
dendron scalpellifolium epilia (3) or Thuidium
cymbifolium-Homaliodendron scalpellifolium group
(at the lower part of tree trunks), the Pterobryum
arbuscula - Anomodon giraldii epilia or Forsstroe-
mia cryphaeoides - Anomodon giraldii group (on
trunks), the Cetraria collata f. nuda - Boulaya
mittenii epilia or Boulaya mittenii - Frullania
moniliata group (at the upper part of trunks and
on boughs in crowns), and the Ulota crispula-
Pertusaria sp 2 epilia or Parmclia laevior - Ulota
crispula group (at the top-most part of the crowns).
It will be probable that the other three remains
shown in Fig. 3, viz., the Graphis sp t - Pertusaria
spj epilia, Graphis sp. -Cetraria collata group, and
Dicranoloma fragiliforme - Metzgeria conjugata
group, are all unlikely to be developed into
independent social units or groups.
CONCLUSION AND SUMMARY
In cases where it is difficult to distinguish plant
community individuality, as in corticolous com-
munities consisting chiefly of bryophytes and
lichens, it seems to us to be possible to make a
classification of such vegetation and to systema-
tize the communities objectively. This would be
81
Epilia
e U/ota crispu/a -
* Pcrtusctria sp.2
PROCEEDINGS OF THE NINTH_ PACIFIC SCIENCE CONGRESS
45
Epiphyte group
Cctmria
* Bou/dydL mittenii
15
/
'
Perl us Aria sp. i
Ptervbryum drbuscula -
Anomoaon gimldii
8
r\ Pcirmelici laevior -
U/ota crisputa
** fragl/iforme -
Cetntria.
sp. -
.'y p Bou/ayd jnittenii -
' Frvllajrii* noni/idtd
14
D Forsstroenu'ci cryphdeoides -
u> Anomodon girctldii
L Uuidium cymtifolium - Y ^\\ ~n' !f * l " m rvmtifnlium -
' homaliodendmn scalptHifolium \ 54 541 ' flomaliot/ein/ron stxtlpellifolium
5 (9
Fig. 3 Classificatory schema of epiphyte communities, showing so as to make comparison between each epilia and group
by means of the value of QS. Indication of correlational similarity between them is the same to Fig. 1.
done on the basis of floristic composition, by
classifying the communities into some homoge-
neous groups by making use of the value of inter-
specific correlation based on the frequency of
major component species and in the light of the
indicator value of significance level (I, 2). These
groups would then be integrated into an appro-
priate number of groups of higher rank at a
wanted level in accordance with the value of
"Quotient of Similarity" (-QS, 5) which is based
on the frequency of component species.
REFERENCES
(1) Goodall, D.W., 1953, Objective methods for
the classification of vegetation, Austral.
Jour. Bot., 1: 39-63, and 434-456.
(2) Hosokawa, T., 1955-56, An introduction of
82
2x2 table methods into the studies of the
structure of plant communities, Jap.
J. Ecol., 5: 58-62 (1955), 93-100 (1956),
and 150- 153 (1956).
(3) Hosokawa, T., Omura, M. and Nishihara,
Y., 1954, Social units of epiphyte com-
munities in forests, VIHe Congr. Intern.
Bot. Paris, 1954. Rapports comm. Sect.
7: 11-16.
(4) Omura, M., Nishihara, Y. and Hosokawa,
T., 1955, On the epiphyte communities
in beech forests of Mt. Hiko in Japan.
Rev. Bryol. Lichen. 24: 59-68.
(5) Sorensen, Th., 1948, A method of establish-
ing groups of equal amplitude in plant
sociology based on similarity of species
content. Dei Kong. Dansk. Vid. Sel.
Biol.Skr.5(4): 1-34.
VOLUME 4 BOTANY
VEGETATION MAPPING IN THE PACIFIC REGION
A.W. KUCHLER
University of Kansas, Lawrence \ Kansas, U.S.A.
Let me begin with a brief report on what has
been accomplished so far in mapping the vege-
tation of the Pacific Region. As you can imagine,
the accomplishments are not at all uniform, but
they are nevertheless very promising. There are,
of course, numerous vegetation maps of very
small areas. I shall ignore these here and report
to you only on maps of countries.
In North America, a vegetation map of British
Columbia was published recently, and maps of
Mexico and Guatemala have been available for
some time. A map of Alaska is now in press, and
there has been a small map of the United States
for a long time; a larger one is now in preparation.
Most of the Central American states remain to
be done. This is a relatively favourable report,
especially if we include the work now in progress.
In South America, conditions are much less
satisfactory, and Peru is the only country of which
a reasonably detailed vegetation map exists.
The most favourable report comes from Austra-
lia. That continent has now been mapped, and
the new vegetation map by Williams has been
published recently. New Zealand has also been
mapped, as well as New Guinea. On the other
hand, the various Pacific islands have been
mapped very unevenly, if at all.
In Asia, progress varies from one country to
the next. The new Soviet map covers a large part
of the Asian continent. Indonesia, the Philip-
pines, and Taiwan have vegetation maps that need
some further work. There are good but small
maps of Korea, Manchuria, Thailand, and Burma.
The greatest gaps are China and Japan, Malaya,
and North and South Vietnam.
Actually, vegetation maps are now available
for the entire Pacific Region, and the International
Bibliography of Vegetation Maps which I hope
to publish some time soon contains over 200
references to countries bordering on the Pacific
Ocean. The scale and quality of these maps vary
within wide limits, and many of the vegetation
maps are too small or too generalized to be of
much use. But modern vegetation maps can be
useful instruments in a variety of scientific investi-
gations, as well as in planning and managing the
use of our lands. This has been demonstrated
many times.
If, therefore, it is evident that vegetation maps
are useful, I should like you to consider the possi-
bility of preparing a vegetation map of your
home land if a good one at a large scale does not
already exist. If you are not in the position to
prepare such a vegetation map, you can perhaps
encourage one of your colleagues to do so.
The maps should be useful to as many people
as possible and not just to those of one country.
For this reason, a certain degree of coordination
is valuable. In addition to suggesting that new
and large scale vegetation maps be made of every
country of the Pacific Region, I should therefore
like to propose that the vegetation be mapped in
its major physiognomic and structural features,
and that these broader types be refined according
to their florist ic character by listing the dominant
genera or species wherever that is feasible.
Structure and floristic composition are the two
basic features which all vegetation types on earth
have in common. They should therefore be given
priority on vegetation maps. If other features
are to be added, such as ecological aspects of the
habitat, they might well be in the nature of a
supplement rather than primary information.
Such supplementary information may be given
wherever this is feasible and useful, but I think
it should be only in addition to the two basic
features of physiognomy and floristic com-
positon, and not instead of them.
Of course, there are many ways to make a
vegetation map, and it is quite possible that
authors may wish to adapt their maps to the
circumstances of their respective countries. The
possibilities of variation are greater than many
authors realize; and while a vegetation map
should not be cluttered, there is, nevertheless, no
need to ignore any chances to make the map
content more meaningful.
Those who wish to make large scale maps will
find interesting and stimulating models in Carl
Troll's vegetation map of the Nanga Parbat area
in the western Himalaya, or in Heinz Ellenberg's
vegetation-site map of Leonberg, Germany; both
maps are at the scale of i : 50,000. Larger scale
phytosociological maps, as published under the
direction of Louis Emberger or Reinhpld Tiixen,
are perhaps not yet feasible for the Pacific Region
83
PROCEEDINGS OF THh NINTH PACIEIC SCIENCE CONGRESS
as they require more detailed information than
is generally available. As far as I know, only
some Japanese scientists have successfully at-
tempted to prepare such maps, and only in very
limited numbers.
Inspiration for small scale maps may be gained
by studying Kurt Hueck's map of Germany
or Henri Gaussen's map of France, both at 1 :
1,000,000. More recently, our Soviet colleagues
have published vegetation maps of their country
at 1 :4,000,000, and Williams published his new
map of Australia at 1 :6,000,000. My own map of
the United States is at 1:14,000,000, and this
small scale is justified only because the map was
included in an atlas.
The preparation of a new vegetation map of
the United States at a larger scale is now in pro-
gress, but I do not expect to complete the manu-
script for another three or four years. For the
large countries of Australia, Canada, China, the
Soviet Union, and the United States, scales of
1 :4,000,000 or 1 :6,000,000 can be justified, but
for smaller nations a scale of 1 : 1,000,000 should
be the minimum.
1 mentioned that many vegetation maps of
the Pacific Region are of too small a scale or too
generalized to be very useful. This need not be
discouraging because the field of vegetation map-
ping is developing only now. The scale can always
be enlarged and more details can be added. Far
more significant are the map content, its organi-
zation, and representation. A consideration of
these features is fundamental and, indeed, im-
perative before preparing a vegetation map.
Today I should like to present to you two
problems for discussion. The first one is that of
the map content. Some authors argue that a
vegetation map should contain nothing but
information on vegetation; others want to show
more, especially ecological features and also land-
use data. A strong case can be made for both
sides. I do not propose here that a decision should
be taken in support of one approach to the ex-
clusion of the other. Both approaches are valu-
able and justifiable. But I do feel that the matter
should be discussed now so that those who
return home to prepare or promote a vegetation
map of their area will see more clearly what
problems they will have to face.
The argument for showing only vegetation on
a vegetation map is strong and logical. Vegetation
is so complex that the map can always be filled
with as much detail as the scale permits. If the
over-all character of the vegetation is simple,
then further detail can be shown by introducing
84
various structural aspects of the vegetation,
differences in the lowest synusia, floristic varia-
tions, transitions associations, sub-associations,
variants, sub-variants, and other features. The
map is then filled to capacity, and the entire
information refers only to vegetation. By exclud-
ing all other kinds of information, the vegetation
can be described in the greatest possible detail.
But there are others who feel that some detail
of the vegetation can or should be sacrificed in
order to introduce ecological information. There
is, of course, no doubt that such information is
revealing and valuable, and therefore should be
given.
However, the introduction of non-vegetational
data creates a conflict: what part of the vegeta-
tional information is to be sacrificed for the sake
of ecological data, and also, how much ecological
information may be introduced without under-
mining the character of the map as a vegetation
map? A further problem is the kind of ecological
information to be shown in a vegetation map.
Should it be any kind, whatever the author
happens to favor at a particular time and place,
or should it be so organized that a logical system
can be applied? If so, what specific ecological
data are to be selected as it is impossible to in-
dicate the environment comprehensively?
Usually, vegetation maps with ecological
information lack unity, but they are nevertheless
very numerous. This shows that many authors
place much value on this approach. On the other
hand, there are only very few vegetation maps
that are based on a strictly organized system
which includes both vegetational and environ-
mental features. I am thinking of the works by
Heinz Ellenberg and Henri Gaussen. There are
also vegetation maps which are strictly vegeta-
tional but which have strong and definite ecolog-
ical implications. The maps by Reinhold Tiixen
and his collaborators belong in this class. On the
maps by Hueck, or by Lavrenko and Sochava,
ecological information is sometimes given, some-
times implied, and sometimes omitted. Carl
Troll fits the vegetation into the landscape, so
that his vegetation map, too, is ecological in
character; his method is most enlightening but
is most readily applied to mountainous terrain
on large scale maps.
The fact that the most interesting or the most
valuable vegetation maps were not made in the
Pacific Region is not at all a cause of concern.
What really counts is the basic ideas on which a
vegetation map rests. Once grasped, these ideas
can be applied in the Pacific Region as well as
VOLUME 4
BOTANY
anywhere else. The problem of the map content
is well worth discussing because the character
and the usefulness of the maps depend on it.
The other feature I wish to present to you for
discussion is the use of color. Several authors
have shown that the judicious application of
color and color patterns can greatly enrich the
map content. Many years ago, Riibel in Swit-
zerland proposed a color scale for Swiss vegeta-
tion maps. This scheme turned out to be too
limited and was not often applied. Much later,
Emil Schmid, also of Switzerland, was careful to
select harmonizing colors to facilitate map reading
but refrained from proposing definite colors for
given vegetation types as Riibel had done.
Schmid has made more of a contribution than it
seems because many vegetation maps have dis-
turbing color schemes. Clashing colors make
map reading difficult and easily suppress im-
portant features on the map.
Sochava of the Soviet Union briefly discusses
the use of color on vegetation maps and proposes
that the color scale used on the Soviet map be
employed elsewhere, too. But he failed to indicate
just how he arrived at his color selections, and
therefore it is not possible to use his ideas sys-
tematically in other countries.
The only person who so far has presented a
complete and logical color system for use on
vegetation maps is Henri Gaussen of France.
He systematically uses the sequence of colors in
the spectrum and different patterns within each
color to express vegctational and ecological
information. The system has been criticized by
Sochava, and indeed, some modifications may be
desirable. But the fact remains that so far, and
as a method of using colors on vegetation maps,
Gaussen's work is unique.
If you agree with me that vegetation maps
should be made of the entire Pacific Region, then
we should ask ourselves whether or not colors
should be used uniformly throughout the region
in order to tie the area together and integrate the
individual maps into a larger whole. Question 1
is therefore: should a color scheme be prepared
for the vegetation of the Pacific Region, or should
the use of color be left to the discretion of each
author? If a more or less uniform system seems
desirable, then there follows question 2: should
the use of colors be restricted to vegctational
information, or may colors refer to ecological
data as well? The more systematic the use of
color, the more readily can the reader distinguish
the major groups of features at a glance.
Finally, there is the use of symbols which may
be employed to enrich the vegetational and/or
ecological information. And again, as in the case
of colors, should symbols be used indiscrimi-
nately or should they be restricted to any one
type of features? Uniformity of approach cer-
tainly facilitates the appreciation of vegetation
maps, but whether such a form of coordination
can or should be achieved is a matter for dis-
cussion here and now.
Certainly the present state of vegetation map-
ping in the Pacific Region leaves much to be
desired. Certainly, enough vegetation maps
have been published all over the world that the
basic ideas can be grasped and developed.
I want to urge you to clarify the thinking of all
of us who arc interested in preparing vegetation
maps of the Pacific Region.
85
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
THE PROBLEM OF THE ORIGIN OF THE "SAVANNAHS 7 '
OF THE ISLANDS OF THE PACIFIC!
R. PORTERES
Museum National d'Histoire Naturelle, Paris, France.
People who deal with the subject of the vege-
tation of the islands of the Pacific always em-
phasize the youthful nature and the ubiquitous
and more or less ruderal flora of the savannahs
of these islands.
The "savannahs" are qualified as "secondary"
and as being the outcome, in fairly recent times,
of the clearing of forests for cultivation and the
subsequent abandonment of the cultivated areas.
The floristic lists given are convincing in this
respect, apart from some reservations which
should be made.
Virot (1957) considers that the "savannahs" of
New Caledonia are of secondary origin and the
direct outcome of the deforestation of the island.
The first explorers already pointed out the pre-
sence of vast "savannahs." However, they
represented only "a degraded and secondary
aspect of the vegetation and . . . they are made up
almost entirely of the species introduced, parti-
cularly the graminaceous species." (Virot, p.
62). "The secondary "savannahs," existing at the
time when France took possession of New Cale-
donia, were, from the beginning, used in part as
natural pasture land. Generally speaking, they
consisted of stretches of dry grasses, among
which the ubiquitous pan-tropical plant pre-
dominated, the graminaceous species for the
most part, of which the principal ones are Hetero-
pogon contortus Roem., Aristida pilosa Labill.,
Eleusine indica Gaertn., Dactyloctenium aegyp-
tiacum Willd., etc..." (Virot, pp. 65 and 66).
"First of all, it is all but certain that almost all
the grassy stretches of New Caledonia have no
other origin than that of the destruction of virgin
forest by fire." (Virot, p. 68). "Among the
graminaceous species may be mentioned, among
others: Heteropogon contortus Roem. and Schult;
pan-tropical pyrophytic plant, typical of "se-
condary savannahs", and Themeda triandra
Forsk; which are, further, widely prevalent in
the interior herbaceous formations of a large
part of Africa and tropical Asia." (Virot, pp.
128-129).
The examination of the lists of species reveals,
in addition, a deficiency in leguminous plants.
t Presented by R. Heim.
86
Papy ( 1954) also points out the absence of prairial
leguminous plants, endemic species as well as
those introduced in ancient times, in the group
known as the Society Islands.
Thus, what the authors here call "secondary
savannahs" would be nothing but anthropic
herbaceous clearings.
Are there no "primary savannahs" in existence
in the Pacific islands? Have they, incidentally,
ever existed ?
We will try to give here both an analysis of
this question and a possible interpretation with
the help of elements of isolated value.
Two hemicryptophytic graminaceous plants to
be found to a certain extent everywhere in the
Pacific Region should have our attention : Themeda
triandra Forsk and Heteropogon contortus Roem.
and Sch., two andropogoneous plants.
Themeda triandra is exclusively paleo-tropical
in its distribution; Heteropogon contortus is pan-
tropical, present in the old and the new tropical
world.
J. Lebrun (1947) proposed the setting-apart of
a superior group in the phyto-sociologicous
African hierarchy, the Themedetalia triandrae
order. This order, of paleo-tropical value, would
in East Africa include an alliance, the Themedion
triandrae afro-orientale. (Lebrun). Among the
vegetable associations included herein, Lebrun
has studied and defined an association with
Themeda triandra Forsk. and Heteropogon con-
tortus^ distinguished as Themedeto-Heteropogone-
tum, in accordance with the appellative rules of
nomenclature in phyto-sociologicous subjects.
The extension of the genus Themeda in the
whole of the inter-tropical and sub-tropical zone
of the old world contrasts with its total absence
in America.
Very generally speaking, all the species of
Themeda are orophileous or sub-orophileous, at
least on approach to the equator. In relatively
high latitudes, they become "planitiary" (habitat
in plains).
The "prairies" of Themeda triandra or of
T. quadrivahis Kunth characterize a prairial
VOLUME 4
BOTANY
pyrophytic type of humid estival tropical climate.
This type, as we shall see later, seems to be very
ancient in the world. The discontinuity in the
area of distribution of two of these species is a
partial proof of this antiquity.
The existence of a Themedion triandrae afro-
orientate does not however presume the existence
of an exclusively West African alliance, at least
in the sense of a Themedion triandrae afro-occi-
dentale. Lebrun (1947) gave an explanation on
this subject, suggesting for this latter region a
vicarious order in relation to Themedetalia and
based on another Andropogon Hyparrhenia
diplandra Stapf.
We have shown (R. Porteres 1951) that, if
traces could be found in West Africa of the
Themedetalia order, of the Themedion alliance
and even of a Themedeto-heteropogonetum asso-
ciation, this has now all collapsed and is rclictual.
The Hyparrhenietalia partially took its place,
not as a floristic vicariant, but as an eco-vicariant.
We have clearly indicated the ecological contrast
between these two orders and have demonstrated
the relictual state of the Themedetalia in West
Africa.
Taken generally, the prairial formations of
Th. triandra can be defined as "pyro-climatic"
and those of Hyparrhenia diplandra as "anthropo-
pyro-edaphic" (R. Porteres, 1951).
In East Africa, the floristic individuality of the
Th. triandra and //. contortus grouping had al-
ready been recognised by R. Staples (1926) in his
pastoral experiments on the cultivation of the
South African Veld. For Staples and Lebrun,
the grouping tends towards a seasonal contrast
which results in a marked periodicity of the
vegetation of this prairial type and thus favours
its maintenance. The periodical spreading natural
fires facilitate the extension of the establishment
of Th. triandra and maintain, in particular, the
association with //. contortus.
At Dahomey it is said that a residue of this
association exists (Porteres, 1951).
Certain differences in the ecological behaviour
of the two species could dislocate the association.
Th. triandra tends towards lighter and better
porous soils than H. contortus. The latter species
at present pan-tropical, appears to be more
aggressive, probably on account of its facilities for
the scattering of the seeds by man and animals
(spikelets which catch on with their awns).
Th. triandra is less aggressive, and this perhaps
explains its absence in America.
In certain countries of the Old World, Th.
triandra is replaced by T. australis (South Africa),
T. quadrivalvis (Madagascar, part of India). Life
associated with Heteropogon contortus is also
to be noted.
The presence in the Pacific lands, either of
//. contortus alone, or of Th. Triandra alone,
or of the two species together, raises in our
opinion the problem of the existence or the non-
existence of primeval herbaceous groups, of
"primary savannahs."
If these species are of recent introduction, the
problem does not arise; they would indicate a
tendency towards the regression of the potential
vegetation of the "present secondary savannahs"
which would thus be approaching a pyrophile
stage.
If these species are present since very remote
ages, this means that pyrophytic prairies existed
and formed part of the natural landscape of the
Pacific islands.
The fact that certain isolated islands or groups
of islands in the Pacific, such as the Society
Islands, possess only Heteropogon contortus and
do not possess Th. triandra give us the impression
that the pyrophytic conditions of these islands
are recent and that primary savannahs have never
existed there.
However, the establishment of the vegetation
of all these islands is of recent origin, by the
covering of uplifts (or volcanic lava stretches)
with vegetation, dating from the Pliocene age,
or even the Pleistocene or the Holocene age.
It is thus impossible to deduce anything much,
except perhaps that it was man who destroyed
forest vegetation and who favoured the her-
baceous clearings which fires were beginning
to ravage.
Quite different is the problem of New Caledonia
where we ascertain the presence, together with
H. contortus, of Th. triandra. Other species of
Themeda exist in New Caledonia: Th. ciliata
which is known to be annual and of recent intro-
duction; Th. gigantea, an annual para-littoral
species which does not interest us here and which
was probably brought in anthropically from India
over Melanesia.
Is the Th. triandra in New Caledonia a species
which is the outcome of natural ancient distri-
bution? Was it part of the normal surface of
this island in archaic times ?
The genus Themeda covers a total paleo-tropical
area reaching from one extremity of Africa over
India and Australia and into New Zealand (?).
Themeda triandra, we may say, is to be found in
the whole of the area covered by the genus. The
species includes numerous variations, certain of
87
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
which have been established as independent
species, and the surviving varieties of which are
apparently well isolated morphologically; this
isolation is also, correlatively, of a geographic
order and reaches as far as the low systematic
levels, so that it has been possible to use it for
various interpretations in Africa (Stapf 1919,
Porteres 1951).
Incidentally, the genus Themeda (and in parti-
cular the species Th. triandra in the wide sense)
particularly merits a very thorough study at
various levels: systematic, geographic, caryologic,
cytogenetic, physiological, ecological, sociological,
and pastoral.
CONCLUSIONS
It is not possible to know whether the Pacific
lands carried balanced herbaceous formations of
primeval origin in ancient times. One now finds
herbaceous clearings which have been introduced
historically. None the less, the existence of the
pan-tropical species Ileteropogon contortus Roem
and of the paleo-tropical species Themeda triandra
Forsk in these meadow lands permit us to assume
either a prairial advance towards a pyrophytic
stage or a relictual state of pyrophytic prairial
conditions.
To know whether primary prairies ever existed
in the Pacific lands is a problem which is far from
being solved.
In the case of the Central Pacific islands which
are geologically young, one should perhaps put
forward the view that they have never existed;
but the case of certain outer, geologically ancient,
chains of islands which belong to the Tertiary Age
of Indo-mclanesian Asia or of the Australian
continent, or of an antarctic continent, may be
different.
BIBLIOGRAPHY
Staples, R.R., 1926, Experiments in Veld manage-
ment, Union S. Afr. Dept. Agric. Sc.
Bull., 49.
Lebrun, Jean, 1948, La Vegetation alluviale au
Sud du lac Edouard, Bruxellcs, Jl:
566-598.
Papy, H. Rene, La Vegetation dcs lies de la
Societe et de Makatea, Vol. I, 386 pp. (in
Geographic forest, du Monde, Tome V,
Sect. 2, VoL 1, Art III, 26me Partie).
Guillaumin, A., 1948, Flora analytique et synop-
tique de la Nouvelle-Cale"donie-Phanero-
games. Vol. I, 369 pp., Paris.
Guillaumin, A., et Virot R., Contributions a la
Flora de la Nouvelle-Caledonie. CII
Plants recoltees par M.R. Virot. Mem.
du Museum National d^Histoire Naturelle.
88
VOLUME 4 BOTANY
THE STRUCTURE OF SOME BIOCOENOSES OF NEW CALEDONIA
J.H. HURLIMANN
Ciha* Basle, Switzerland.
Attempts to describe thoroughly and classify
the New Caledonian vegetation were started only
about 30 years ago. Last year, an important
study by R. Virot was published, giving for the
first time a synopsis of the vegetation groups of
the whole island. Together with the studies by
Daeniker, Sarlin and other authors, we now
have substantial data regarding the composition
of several groups of plant species. However,
precise observations of documentary nature are
partly lacking with regards to frequency, arrange-
ment and development of species within the bio-
coenoses. The works of Dansercau, Richards
and Schmid set out the lines along which such
studies should be conducted, which are of funda-
mental importance to the knowledge of the ecolo-
gy of the species and its application in forestry,
agriculture and livestock.
By adopting the method described by Schmid,
we endeavoured to get an idea of the structure of
parts of some biococnoses. Unfortunately, due
to limited time, all generalized study was impos-
sible. On another occasion, we described this
method, making certain changes that appeared
useful, for the determination of the structural
features and their illustration.
The purpose of this paper is to give an idea of
the structural facts which are recognizable through
these biocoenological surveys.
Sarlin mentions the classification of the New
Caledonian vegetation into 7 distinct formations
established by previous authors:
1. Coastal zone with Oceanian flora;
2. from to 3-400 m the Niaouli Savannah
( Melaleuca leucadendron) ;
3. from 400 to 1,000m medium altitude forest;
4. from 1,000 to 1,500m dry coniferous forest;
5. the forest galleries;
6. lower serpentine scrub;
7. summit scrub.
Those who know New Caledonia well realize
that this classification is superficial, incomplete
and even erroneous (for example the definition
of the formation under item 4 as "dry forest."
Virot has chosen the following system to give
a comprehensive view:
1. Halophilous formations, including:
a) mangrove,
b) herbaceous vegetation of beaches,
c) belt of so-called "Seaside" trees;
2. Non-halophilous formations, divided into:
A forest series, including:
a) coastal climatic forests (coastal hillside
wood),
b) hydrophilous forests of the river banks,
c) photophilous forests of the marshes,
d) mesophilous forests of the valleys,
e) shade-tolerant forests (insufficiently speci-
fied),
f ) oro-nephepiphilous forests,
g) photo-xerophilous forests;
A series of shrubs, including:
h) edaphic sclerophilous-xerophilous scrub,
i) orophilous scrub;
A herbaceous series, including only:
k) paludal groups.
Of course, this classification will only be provi-
sional since it does not take into account the mixed
floristic features which constitute the vegetation
of different regions of the island. Despite this
defect, Virot's classification shows considerable
progress when compared with previous classifi-
cations.
Our structural studies were carried out mostly
in the formations which Virot calls vallicol-meso-
philous and oro-nepheliphilous, which are
obviously very mixed formations. By way of
comparison, we will also discuss a survey made in
a secondary coastal forest. On the other hand
we will not talk about shrub vegetation of ser-
pentine areas which we have not been able to
study in detail.
The forests in which we were most interested
because of their aspect are those with a predom-
inance of Neo-caledonean Fagaceae, of Trisyn-
gyne spp. They are distinct from other types of
forests by the fact that only one species forms,
nearly exclusively, the dominant tree stratum.
Our surveys Nos. 6, 9 and 10 were made in
Trisyngyne codonandra Baill. forests; survey No.
7 in a 7>. Balansae Baill. forest. All the fragments
studied are located on peridotitic mountains
quite near to one another: the farthest apart are
separated by less than 30 km. In spite of this,
relatively important differences are apparent,
even between the three Tr. codonandra biocoe-
noses.
Due to lack of time, it will not be possible to
reproduce here the complete lists of species.
89
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Their study shows that the floristic composition
is very variable in Tr. codonandra forests. Among
all the species found in the 3 surveys, there are
only two species, Tr. codonandra itself and
Symplocos cf. Pancheri, found in all three surveys.
Five species are common to surveys 6 and 9,
eight to surveys 9 and 10, while surveys 6 and 10
have no particular species in common.
It is interesting to see that survey No. 7, in
spite of the dominance of another species of
Trisyngyne, seems to connect the other fragments
of vegetation. In fact, surveys No. 7, 9 and 10
have 3 species in common, and surveys 6, 7 and
9 two species. The total of the plants approxi-
mately identified in the 4 fragments studied is
about 150 phanerogamic species. The coincidences
found are therefore rather poor, and it will be
admitted that the floristic-statistics method is
not of very great use in these circumstances.
Now let us proceed to the information received
on biocoenological structure. By grouping the
plants together, forming provisional ecological
types, the following frequency of the different
types in the biocoenoses fragments occurs :
Table 1.
Frequency of provisional ecological types in the vegetation according to the surveys
(in per cent of small squares of the surveys):
a
b
c
e
8 =
i
k =
n
o
pst
pep
VE
VB
w
X
y
trees more than 1 2 m high (number)
shade produced per crown
trees more than 8 to 12 m high (number)
shade produced per crown
trees less than 8 m high
trees with tufts of leaves, 2 to 8 m high,
few branchings
trees with tufts of leaves, less than 2 m high,
few branchings
prostrate trees
shrubs more than 2 m high
shrubs less than 2 m high
woody lianes or lianes with stiff stems
prostrate shrubs
erect herbs
herbs forming dense clumps
herbs forming rosettes
herbs forming stems
epiphytic herbs
bryophytes on the ground
bryophytes on wood (dead or living)
lichens
saprotic mushrooms
algae
3
no data
8
no data
9
18
4
5
6
1
13
22
10
60
21
9
47
45
1
7
5
6
4
50
30
7
4
38
1
85
85
9
3
10
~2
96
3
23
37
1
5
11
10
2
9
6
1
73
77
1
4
67
8
32
43
1
1
2
13
1
2
46
15
11
53
57
3
1
To these figures must be added those of young plants which, according to their state of growth, parti-
cipate in the composition of the various biocoenoses strata:
Young plants more than 1 m high
Young plants, 20 -100 cm high
Young plants less than 20 cm high
21
71
54
9
42
72
45
10
4
48
17
74
95
The most noticeable differences are found in the density of class c trees (including class e and
the young trees over 1 meter high) and in that of non-epiphytic undergrowth (including young plants
90
of less than 20 cm high),
are as follows:
Class c etc.
Class p etc.
VOLUME 4 BOTANY
The total percentage of participation of these 2 categories in the 4 surveys
6
48
67
10
97
136
43
49
63
156
Therefore, surveys Nos. 7 and 9 present a much more developed undergrowth than Nos. 6 and 10.
This fact is also a result of the number of small squares entirely without any vascular plants (in % of
the total of squares):
6 9 10 7
10
1
17
These differences are due to the growth of the
dominant tree strata, producing more or less
intense shade according to the density of the
foliage.
The frequency of the shrubs follows the same
lines, but their absolute number is very small,
so that they only play a minor part in the structure
of the vegetation as a whole.
The proportion of lianes is almost similar in
the 3 surveys Nos. 6, 9 and 10, but there is an
abundant presence of young creepers (Frcycinctia
sp., Smilax sp. and Alyxia sp .) in survey No. 6.
Jn the participation of young plants, it is ad-
visable to compare the number of adult and young
subjects of various species. These observations
give valuable information on the probable
development of the biocoenoses in question. The
lack of young plants of dominant species can
induce a change in the structure of the whole
vegetation, whilst an abundance of young plants
is a certain guarantee for the maintenance of
population. Below are the most striking examples
of both statements, found in our surveys:
a) Very abundant young plants, compared to adult
plants:
young adult
Survey No. 6
Trisyngyne codonandra \ 39 3
Tree No. 40 22 1
Survey No. 9
Araucaria muelleri 108
Palm tree No. 24 311
Survey No. 10
Trisyngyne codonandra 46 3
Rubiaceae No. 29 23 1
Survey No. 7
Trisyngyne balansae 652 8
SapindaceaeNo.il 42 10
Guttiferae No. 32 33
b) Abundant adult plants, few young plants:
young adult
Survey No. 9
Trisyngyne codonandra 4 7
Cunoniaceae No. 9 2 14
Myrtaceae No. 3019 3 17
Survey No. JO
Sapindaceae No. 34 08
Survev No. 7
Hibbertia sp. No. 14 8
Montrouziera sp. No. 4 117
Leucopogon dammarifolius 6 14
In surveys Nos. 6, 10 and 7, seed reproduction
of the dominant species is well favoured. On
the other hand, its future is uncertain in survey
No. 9. Here, the young Araucaria muelleri plants
occupy a very important place in the vegetation
of the lower strata although there are no adult
trees of this species in the square of the survey or
in its immediate neighbourhood.
However, the abundant seed production and
germination are still not sufficient to guarantee
the stability of the stock of plants. Furthermore,
development must also take place without any
particular hindrance, whilst allowing for a succes-
sive elimination of young plants by competition.
In surveys Nos. 6 and 10, this phenomenon is
apparently present since there are young plants
of all sizes. The case of survey No. 7 is different :
beside 8 adult trees, we find 543 young plants of
10 cm or less, 105 of 11-20 cm, only 4 plants
between 21-30 cm and no taller plants. Beside
an irregular seed production from year to year
which seems likely, it is probable that further
growth of the young plants is hindered when they
have reached a certain age, although we cannot
indicate the factors responsible for this.
We should like to mention here a study made by
Mr. Letouzey, Conservateur des Eaux et Forets
91
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
in French Camerons, on the Lophira alata forest
of the coastal zone of the Camerons. There,
the Lophira alata has the same symptoms of com-
petitive insufficiency in its youth, and the author
presumes that without protected artificial regen-
eration, the forest composition would change.
The interpretation of survey No. 9 is rather
delicate. The four young plants of the dominant
species are 20, 30, 40 and 40 cm high respectively.
An adult plant of 6 meters only has not yet
attained its optimal growth. However, the massive
growth of young Araucarias between 20 to 160
cm high seems to show that this forest will be
changed as and when the old Trisyngync die. This
transformation is apparently favoured by a rather
weak shade produced by the adult Trisyngync, as
their crowns are not contiguous, and by the fact
that other species participate in the formation of
the dominant stratum (A gat his ovata, Salauopris
sparsiflora and Rhodamnia andromcdoides). In
fact, at this high altitude, Trisyngyne does not
seem to be at its best, and its position seems
to be endangered by the strong competition of
other plants. Changes of secondary order in
ecological conditions may consequently easily
provoke structural changes and accelerate the
dynamic tendencies of the biocoenose.
One last structural component on which we
would like to draw attention is the distribution
of the species in the area surveyed. Beside
plants like Phclline lucida and Myodocarpus
fraxinifolius of survey No. 6, or Leucopogon dam-
marifolius of survey No. 7, which are all quite
regularly distributed over the whole area, there
are others such as Rubiaceae No. 29 of survey
No. 10, or Lophoschocnus montisfontium of sur-
vey No. 9, the former located in one corner only,
the latter separated in three distinct groups.
These irregularities can be caused by two com-
pletely independent factors: 1. the character-
istics of the biological system of dispersion of
fruits or seeds, as well as the possibilities of a
vegetative reproduction, or, 2. conditions of the
biocoenological medium allowing the develop-
ment of plants in one area, and hindering it in
another.
The first factor is apparently responsible for the
grouping of the plants of Rubiaceae No. 29 men-
tioned: all the young plants are grouped around
an adult tree. To a certain extent, tree No. 40
of survey No. 6 and Spiraeanthemum sp. No.
2 of survey 10 show similar phenomena. On the
contrary, with regards to Lophoschoenus montis-
fontium, their relation to the medium seems more
important: it is apparent that it is found mostly
92
under the spaces left open by the crowns of domi-
nant trees. The contrary could be said regarding
Phelline lucida of the same survey (No. 9) which
seems to follow the shade of the trees. Therefore,
this species shows that the group forming tenden-
cy is subject to the influence of the vegetation
itself, having a homogenous distribution in a
forest of regular shade, forming groups in a
forest of irregular shade.
The presence of groups of certain species, in
relation to other plants, is very interesting for
the classification of vegetations. However, care
should be exercised in order to avoid overesti-
mation of the value of the groups. Daeniker has
always insisted on the necessity of considering
groups within a wider framework, to look at them
as fragments of a mosaic. Temporary dominant
or subdominant conditions caused or suffered
during certain states of individual growth of a
species can be very important, without justifying
however a classification of biocoenoses into
smaller defined units. In this respect, we recall
the lively discussions which took place during
the last International Congress of Botany in
Paris concerning the existence of tropical asso-
ciations.
Taking our structural analysis into account,
we will not hesitate to put surveys 6 and 10
together as having both the same type of biocoe-
noses, despite the absence of common species in
the under-growth and herbaceous stratum. This
type is characterized by one dominant stratum of
trees of equal height, reaching 12-15 m. The
lower strata are not very developed. There
seem to be minor differences only in the compo-
sition of the stratum of small trees (presence of
many trees of provisional ecological type e) as
well as in the frequency of young plants.
Survey No. 9 is clearly different in its structure.
Dominant adult trees do not reach the dimensions
of the Trisyngyne of the two afore-mentioned
surveys. The crowns do not form a contiguous
cover and consequently, the undergrowth and the
herbaceous stratum are much more developed.
This structure is emphasized by the abundance of
young plants. Analysis of the fragments of
neighbouring vegetations will doubtless show
that this is a mosaic stone forming a transition to
distinct biocoenoses of higher altitudes.
Survey No. 7 belongs to a third type of biocoe-
noses. The dominant trees reach 12-15 m in
height, but without forming a complete cover at
crown level. The foliage of TV. balansae produces
less shade than that of Tr. codonandra. Therefore
the undergrowth and the herbaceous stratum are
VOLUME 4
BOTANY
well represented and heliophilous shrubs such as
Leucopogon, Dracophyllum and Hibbertia, as
well as the heliophilous orchid Eriaxis rigida,
can be found. Contrarily to what we find in
survey No. 9, there is a lot of herbs with a tendency
to form aerial stems without leaves (Lophos-
choenus spj, while rosette herbs are lacking.
The common characteristics existing between
the four surveys cannot fail to be noticed: the
dominant stratum reaching a certain height above
the ground and formed essentially by one species
alone, with, at most, 2 to 4 species reaching the
same level, but isolated between the Trisyngyne.
The most frequent families by number of individ-
uals are the following: Fagaceae, Polypodiaceae,
Orchideae, Cyperaceae, Sapindaceae, Rutaceae,
Aquifoliaceae and Guttiferae (thus, three families
comprising mostly herbs).
Now let us compare briefly another survey
with the four fragments discussed: No. 8, situated
near No. 7, on a steep slope of a pcridotite moun-
tain. Even at a superficial glance a complete
difference from the afore-mentioned biocoenoscs
is apparent.
Above a tree stratum forming a continuous
dome but unequal as to its level above the ground
(between 10-15 m in general) a few isolated trees
grown up to a height of 20-40 m (Agathis lanceo-
lata, Araucaria hernicri, Alhizza sp. and other
species). Their ecological influence is unim-
portant compared to the trees forming the dome.
Despite the dense shade produced by the latter,
the stratum of trees of less than 8 m high (types c
and e) is represented in abundance, and small
shrubs (type k) are more frequent than in any
Trisyngyne forest surveys. Herbs are also found
in greater abundance than in fragments of shaded
biocoenose of Tr. codonandra Nos. 6 and 10.
Finally the very high number of young plants
(33 % of the square are young plants higher than
1 m, 91% young plants of 20 to 100 cm, 77%
young plants smaller than 20 cm) indicates that
this biocoenose is well balanced and that its
participants are well adapted to the given condi-
tions.
This is explained by the very different floristic
composition of this forest, compared to previous
surveys. The most represented families by number
of individuals are in this case: Rubiaceae, Pan-
danaceae, Arabiaceae, Guttiferae, Euphobiaceae,
Myrtaceae, Laureaceae and Myrsinaceae, i.e.,
families of trees found in all dense tropical forests
of the world. The undergrowth includes a rather
large number of sciaphilous species, specially
a Phyllanthus and two Psychotria. The Freyci-
netta genus is also well represented, specially
among young plants. Amongst the genera en-
countered in the Trisyngyne biocoenoses, only
Phelline and Myodocarpus fraxinifolius are found
in greater number, hence, trees which are also
found under shade in survey No. 6.
Finally, on 100m 2 , we have found 14-16 species
in the strata of trees of over 8 m high, and it is
certain that this number would increase if the area
observed was extended.
Therefore the structure of the biocoenose frag-
ment of survey No. 8 is that of a tropical forest.
By its complexity and the great amount of species
it contains, it is clearly distinct from Trisyngyne
forests. Of course, it does not correspond to
dense forests of the type of equatorial rain forest
of the big continents, but it is noticeable that the
general idea, if I may call it so, is similar. Besides,
we have seen very similar forests from the struc-
tural point of view in Central America, in East
Cuba and on the Eastern slope of the Drakens-
berg, South Africa.
Trisyngyne forests, with their simpler structure
and predominance of one species only in the
higher tree strata, are more reminiscent of forests
of temperate regions. However, it would be
unwise to use this term in defining any vegetation
of New Caledonia. In fact, the geological evolu-
tion of this island, the phylogeny and epi-ontology
of its vegetation are so different from what is found
in temperate countries that we would prefer to
create a new expression: "paratropical forest"
next to tropical vegetation properly speaking.
To add another type of structure, we would
like to mention briefly the study of a fragment
of a secondary forest, greatly influenced by man,
located on the slopes of Ouen Toro near Noumea.
The ground is of sedimentary origin, and the
vegetation varies from very impoverished grass-
lands to regenerated forests, as described by
Sarlin in his work.
. In the fragment studied, we again find a domi-
nant tree stratum, only reaching 8 m in general,
composed nearly exclusively of Acacia spirorbis.
A few trees of other species are mixed therein:
Casuarina cunninghamiana and Melaleuca leuc
adendron. Under the shade of these trees,
the crowns of which are close enough together
without producing however a too intense shade,
-the small trees and shrubs are few. On the con-
trary, the herbaceous stratum is very developed
(specially Bidens pilosus, Plectranthus parviflorus,
Rynchelytrum roseum and other Graminaceae and
Composites),and the ground is abundantly covered
by young Passiflora suberosa plants, Observing
93
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
the distribution of herbs throughout the survey,
it is apparent that the structure is not firmly
established, because the herbs are generally lim-
ited into groups, with small interference of species.
Therefore, we are dealing with initial stages rather
than with a normal vegetation competition.
As I said in the beginning, this summary of our
biocoenological studies of New Caledonia is not
sufficiently complete to give you a general view of
the types of structures found on this island. We
have spoken neither of swamp forests on sedi-
mentary grounds which are distinct from those
found on peridotitic grounds, nor of the Niaouli
savannah (Melaleuca leucadendron ) , which has
its own many difficult problems as to its original
genesis, problems which perhaps only a detailed
analysis of the structure could solve. We have
not discussed the following points: the shrubby
"bush" and the serpentine vegetations, with their
various strata and types; to what extent are they
natural, are they caused by fire or by cutting?
We were unable to compare the vegetation of
the peridotitic, schistose and gneissic mountains,
and we have not established the characteristics
of various altitude zones. All this would require
much more time for field work than we had, and
the only thing we seek to accomplish by this paper
is to show the Wciy in which one can attempt to
solve these questions.
94
VOLUME 4
BOTANY
THE FOG BELT RAIN FOREST OF THE PACIFIC NORTHWEST (U.S.A.)t
RUDY W. BECKING
The Pennsylvania State University, University Park, Pennsylvania, U.S.A.
Coastal forest types in western Washington and
Oregon resembling tropical forests by their
luxuriant epiphytic growth on trees, have been
called "rain forests." Generally these rain forest
types are confined to the Fog Belt Subregion.
PHYSIOGRAPHY
The unique physiography of the Fog Belt Sub-
region consists of high mountain ranges directly
parallel to the Pacific Coast blocking the path
for southwesterly ocean winds. These mountain
masses (Coast Range and the Olympics) form a
barrier of 3,000 to 8,000 feet elevation forcing
these moisture-laden storms to rise and condense,
resulting in the highest orographic precipitation
for the whole North American continent. Several
large rivers have cut through the Coast Range
creating wide valleys open to these ocean winds.
Glaciers advancing down from the Olympics have
scoured other deep, steep-sided, U-shapped valleys,
in places sometimes more than a mile wide.
These glacial valleys are essentially level with
a gradient of about 1 %. The highest precipita-
tion occurs at the end of the 2-30 mile long and
broad valleys around 1,500-2,000 feet elevation,
sometimes resulting in an annual precipitation of
more than 300 inches.
CLIMATE
The proximity of 2-20 airline miles to the
Pacific Ocean creates an ideal marine climate in
this narrow Fog Belt Subregion, extending some
500 miles along the Pacific Coast, from California
into Alaska.
General characteristics of this marine climate
are:
(1) A very narrow fluctuation of temperature
extremes hardly exceeding 20F annually with
an average annual temperature of 49 - 50F.
(2) The longest recorded frost free period
extending well over 200 days, ranging from 180-
310 days which is exceptional for the 44-49 paral-
lel northern latitude.
(3) Consequently, it has no snowfall directly
near the ocean. Six miles further inland in the
valley bottoms snow depth increases from a few
inches to two feet and at a distance of 15
miles inland from 2 to 4 feet. Above 1,000 feet
elevation, annual precipitation in the form of
snow can even amount to 300 inches at high
elevations.
(4) No appreciable drought period in the
summer. During the early morning hours incom-
ing ocean fog will create considerable fog drip
within the rain forest preventing these forest types
to dry out. Forest fires which commonly ravage
more inland forest types have never been able to
penetrate into these rain forest types.
VEGETATION PATTERN
Rain forest types cover river flats and the foot
hills up to about, 1,000 feet elevation constituting
of two, possibly more, alliances of coniferous
forest types dominated by the Sitka spruce and
western hemlock, and of deciduous forest types
dominated by the big -leaf maple and red alder.
The American rain forest types are character-
ized by:
(1) Unusual tree heights and sizes. The largest
and highest individuals of Douglas fir, Sitka spruce,
red cedar, and hemlock are all found within the
rain forest. Tree heights of 300 feet and more are
not uncommon in virgin stands of sheltered
valleys with tree ages of about 2,500 years.
(2) Buttressed tree bases and collonades.
Sitka spruce and hemlock regeneration usually
starts upon rotting logs and stumps. Seedling
growth is very slow on these substates until its
roots strike mineral soil. Roots become heavier
and fuse in order to support the increasing tree
size. Gradually, the nurse-log decomposes, leaving
a collonade of trees with buttressed bases and
exposed root collars.
(3) Luxuriant epiphytic growth on stems and
branches of trees. In temperate forests, epiphytic
vegetation is rarely encountered. Low tempera-
ture and occasional dessication in the summer
have limited the epiphytic flora to a few crypto-
gams, lichens, bryophytes, and pteridophytes.
No profusion of vascular epiphytic plants, so
t Presented by F.E. Egler.
95
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
typical of tropical rain forest, exist. Trees may
become sometimes so heavily laden with an
upholstering of rain-soaked bryophytes that they
are susceptible for windthrow.
The coniferous forest alliances can be divided
into four main forest types:
The Sitka Spruce - Hemlock forest type. This
is the most common rain forest type covering
river flats, slopes, and ridges. Its tree layer is
dominated by Picea sitcheniss and Tsuga hetero-
phylla usually in an intensive mixture, however
locally pure stands of one of these dominant trees
are also found. Its shrub layer is dominated by
Acer circinatum or Rubus spectabilis particularly
in river flats and on lower slopes. Its herb layer
is characterized by the dominance of Polystichum
munitum, Oxalis oregana, and occasionally
Tiarella trifoliata.
The characteristic species composition includes:
the group of character species of this forest
type:
Arceuthobium tsugen- Moneses uniflora
se
C V/V? ton la un (flora
Cornus canadensis
Disporum smith! i
Mcnziesia ferruginea
Picea sitchensis
Rubus pedatus
Tsuga heterophylla
Vaccinium alaskaense
Vaccinium ovalifolium
Particularly the stands on river flats, lower and
middle slopes are characterized by the common
moisture indicators:
Acer circinatum
Athyrium felix-femina
Blechum spicant
Disporum oreganum
Dryopteris dilatata
Fesfuca subulifiora
Luzula parviflora
Maianthemum
dilatatum
Melica subulata
Montia sibirica
Oxalis oregana
Polystichum munitum
Rubus spectabilis
Sambucus callicarpa
Stachvs cilia ta
Tiarella trifoliata
Stands on upper slopes are generally character-
ized by the lack of moisture indicators and by the
abundance of:
Gaultheria shallon Linnaea borealis
The Coastal Sitka Spruce forest type. This
forest type is limited to bluffs and west slopes
directly facing the Pacific Ocean. Dominant
trees of Sitka spruce, hemlock, and red alder
are stunted by wind. Its shrub layer is usually
dominated by Gualtheria shallon, sometimes
96
on the lower slopes by Rubus spectabilis.
The characteristic species composition consists
of.
Alnus ruhra
Arceuthobium tsugense
Lonicera involucrata
Picea sitchensis
Thuja plicata
Tsuga heterophylla
Vaccinium ova turn
together with most of the common moisture
indicators.
The Sitka Spruce -Lodgepole Pine forest type.
This forest type is confined to areas with sandy
soils and dune formations in the vicinity.
Its characteristic species composition consists
of:
Gaultheria shallon
Lonicera involucrata
Mains rivularis
Picea sitchensis
Pinus contorta
Rhododendron
californicum
Thuja plicata
Tsuga heterophylla
Umbel la ria calif or nica
Vaccinium ova turn
with hardly any of the common moisture
indicators.
The Sitka Spruce Swamp forest type. Meander-
ing rivers create extensive swamp forest areas in
their deltas. Trees are situated on humps within
the swamp. Its tree layer is rather open, charac-
terized by the dominance of Sitka spruce and red
alder mixed with an occasional hemlock; its
shrub layer is dominated by Rubus spectabilis.
Its characteristic species composition consists of:
Alnus rubra
Cardamine angulata
Car ex obnupta
Chrysoplenium glecho-
maefolium
Lonicera involucrata
Lysichitum camtschat-
cence
Oenanthe sarmentosa
together with the character species of the
Sitka Spruce - Hemlock forest type:
Menziesia ferruginea
Moneses uniflora
Vaccinium alaskaense
Vaccinium ovalifolium
with the common moisture indicators:
Athyrium felix-femina
Blechnum spicant
Dryopteris dilatata
Luzula parviflora
Maianthemum
dilatatum
Polystichum munitum
Rubus spectabilis
Sambucus callicarpa
Stachys ciliata
Tiarella trifoliata
VOLUME 4
In the deciduous rain forests, the Riverflat
Maple and the Red Alder forest types are disting-
uished. The Riverflat Maple forest type is noted
for its luxuriant epiphytic cover, being considered
usually the veritable rain forest. In virgin stands
the dominant bigleaf maple is mixed with Sitka
spruce, red alder, cottonwood, hemlock, and
grand fir, but in second growth stands usually
only the maple is present. A luxuriant cover of
bryophytes and vascular plants cover the soil,
rotting logs and tree trunks while a dense herb and
shrub layer will add to its jungle-like appearance.
However, on the Olympic Peninsula these
Riverflat Maple forests have a rather open and
park-like character, lacking chiefly the dense layer
of high herbs and shrubs. The reason for this is
the presence of some 4,000 Roosevelt elk (Ccrvus
canadensisvar: roosevelti) wintering in these rain-
forest valleys inside the Olympic National Park.
As the Roosevelt elk is gregarious, frequently
banding in herds from 20 to 100 animals, the
effects of elk herds' browsing and trampling are
significant. The lack of high herbs and shrubs and
the introduction and abundance of grasses and
grazing indicators have given this forest type an
overbrowsed appearance. Since 1935, elk cxclo-
sure plots have been established within the
Olympic National Park. At the present the vege-
tation inside these fenced plots is characterized
by a tall herb layer and a dense shrub layer of
Acer circinatum, Sambucus callicarpa^ and
Rubus spectabilis indicating its natural appear-
ance without the overbrowsing effects of the
Roosevelt elk. In other parts of the Fog Belt
Subregion where no Roosevelt elks are present,
deer will frequently graze in the Riverflat Maple
and Red Alder forest types, so that grazing
indicators and grasses form a characteristic com-
ponent of these deciduous rain forest types.
The Riverflat Maple forest type. The charac-
teristic species composition of this forest type is
comprised of:
in its tree layer:
Pseudotsuga taxi folia
Selaginella oregana
(epiphytic)
Thuja plicata
Tsuga heterophylla
Abies grandis
Acer macro phyllum
(dom.)
A Inns rubra
Picea sitchensis
Populus trichocapra
in its shrub layer :
Acer circinatum (dom.) Holodiscus discolor
Corylus californica Oplopanax horridus
JBOTANY
Samhucus callicarpa
Tax us brevi folia
Osmaronia ccrasiforis
Rubus spectabilis
in its herb layer:
the group of common moisture indicators:
Acer circinatum
A thyrium feliz-femina
Blechnum s pi cant
Dent aria tenella
Disporum oreganum
Dryopteris dilatata
Festuca subuliflora
Luzula par vi flora
Mciianthemum
dilatatum
Melica subulata
Montia sibirica
Oxaiis oregana
Rubus spectabilis
Sambucus callicarpa
Stachys ciliata
Tierella trifoliata
Trientalis latifolia
the group of riverflat moisture indicators:
Asarum caiulatum
Circaea pad fie a
Dicentra Jormosa
H vdrophyllum tenuipes
Mitella caulescens
Nemophila par vi flora
Rosa nut k ana
Symphoricarpos
albus
Tell in HI grandiflora
Tolmiea menziesii
Urlica Ivallii
Viola glabclla
the group of grazing indicators:
Adenocaulon hi color Calium tri/idum
Agrostisstoloniferajexarala Osmorhiza nuda
Bromus sitchensisjvulgaris Poa kelloggii
Cardamine oligosperma
Carex leptopoda
Deschampsia elongate/
Galium aparine
the group of Riverflat Maple forest character
species:
Ranunculus bong-
ardii
Stellaria crispa
Trisetum cernuum
Carex hendersoni
Car ex mertensii
Elymus glaucus
Galium oreganum
Epiphytes on trees are comprised of a large
group of bryophytes and lichens of which the
most common ones are :
Antitrichia curtipendula
Camptothecium lutescens
Claopodium crispijolium
Eurhynchium oreganum
Frullania nisquallensis
Homalothecium nuttallii
Hylocomium proliferum
Hypnum subimponens
Lobaria oregona
Metzgeria conjugata
Metzgeria pubescens
Mnium menziesii
Mnium punctatum
Mnium venustum
Neckera douglasii
Neckera menziesii
Pore I la navicularis
Pseudoiso thecium
sWloniferum
Rhytidiadelphus
loreus
Rhytidiadelphus
triquetrus
97
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
A few plants are also characteristic epiphytes:
Montia heterophylla Selaginella oregana
Polypodium vulgare
The Red Alder forest type. It occupies similar
habitats as the former forest type and is usually
in a successional transition towards it. Charac-
teristic for this forest type is a dense, even-aged
stand of Alnus rubra, the almost complete lack
of any shrub layer in its typical form and the lack
of a dense epiphytic upholstering of the tree
trunks. Mostly, the red alder is densely covered
by crustaceous and foliosc lichens with some
bryophytes, comprised of:
the group of common epiphytes:
Antitrichia curtipendula
Cetraria glauca
Frullania nisquallensis
Ne( '/< cm do u% las ii
Parmefia enteromorpha
Pannelia physodes
Pcrtusaria multipuncta
P or el la navicularis
Pseudoiso thecium stol-
onijerum
and the group of epiphytes typical for red alder
trees:
Cetraria scutata
Dimino\veL\ia cirrhata
Ever n ia prunastri
Graphis scripta
Lecanora subfusca
Ochrolechia tartarea
Orthotrichum consi-
tnile
Orthotrichum Ivellii
Parmclia pertusa
Par me I ia sax at His
Radula bolanderi
Radula complanata
The characteristic species composition of this
Red Alder forest type is comprised of:
the group common moisture indicators:
Acer circinatum
A thy Hum felix-femina
Dryopteris dilatata
Luzula par vi ft or a
Oxalls oregana
Polvstichum munitum
Rhamnus purshiana
Ruhus spectabilis
Samhucus callicarpa
Stachys ciliata
TiareUa trifoliata
the group of riverflat moisture indicators:
Circaea pacifica Osmaronia cerasifor-
Dicentra formosa mis
Hydrophyllwn tenttipes To/mica menziesii
Mitella caulescens Viola glabella
the group of swamp indicators:
Cardamine angulata Lysichitum camtschat-
Chrysoplenium glech- cense
omaefolium Oenanthe sarmentosa
Corydalis scouleri
the group of grazing indicators with the follow-
ing additional grazing indicators typical for this
forest type:
Cirsium lanceolatum
Equisetum arvense
Rumex acetosella
Rum ex obtusifolia
Stellaria graminea
and the group of character species of the Red
Alder forest type:
Epilohium adenocaulon Pleuropogon refractus
Glycerin elata
Mimulus guttatus
Mite (la ova Us
Pctasites spcciosus
Ribes petiolare
Senecio triangularis
Yaleriana sitchensis
The deciduous rain forest types are strictly
confined to valley bottoms and coves along
streamlets below 1,000 feet elevation. They are not
completely restricted to the Fog Belt Subregion
but may be found locally in the foot hills of the
Cascades where the annual precipitation is
unusually high. Both forest types occur on
alluvial deposits close to the river. The open
river bars are first colonized by red alder and
cottonwood.
Successional trends of the Red Alder forest type
towards the climax Riverflat Maple forest type can
be frequently encountered, but no observations
have led to the belief that eventually the deciduous
Riverflat Maple forest will evolve into a conifer-
ous rain forest type. When the river alters its
course, it may disrupt forest vegetation and occa-
sionally Douglas fir may get established in pure
even-aged stand patches. The forest communities
in such Douglas fir stands are closely allied to the
Sword Fern-Douglas Fir alliance. As fire is not
the tool for opening up the forest canopy for
future Douglas fir regeneration, only clearing of
forested land by river erosion and redeposition
of new land will perpetuate the Douglas fir forest
community. Otherwise, it will evolve into the
climax of the Riverflat Maple forest type.
98
VOLUME 4
BOTANY
THE ICONOGRAPHY OF THE VEGETATION OF
THE NATURAL FOREST IN JAPANt
MISAO TATEWAKI
Hokkaido University, Sapporo, Japan.
The present project is a research into the forest
vegetation of Japan. The writer wished to obtain
a phytosociological description of the various
forest vegetations, once the tremendous treasure
of Japan. The primeval forests in Japan have
been largely destroyed, and this destruction will
continue until public pressure halts the abuse.
The writer selected carefully the experimental
plots represented by the fine forests in the present
Japan where the excellent physiognomies of the
natural forest have been fortunately preserved.
To analyse the construction of the forest
vegetation, the writer used "the sociation" as the
synecological unit. He treated it as the species
in the systematic botany. This idea results from
the field surveys of the complicated forest veget-
ation of Japan in these twenty-five years. For
the description of the sociation, the bisect method
accompanied by the map, the sketch, the photo-
graph, and the analytical table was used. It is
the shortest and the wisest method to express the
composition of the forest vegetation in the natural
stand. The writer has attempted to describe
the vegetation accurately and to analyse precisely
and carefully the construction of the forest layers.
Generally speaking, Japan ranges from the
zone of southern temperate forest to that of the
northern temperate, each represented by the
evergreen broad-leaved forest and the summer-
green one. The former includes the subtropical
rain forests in its southern points and the latter
sometimes accompanied by the subarctic forests
in the northern part. The latter in Hokkaido
except the southwestern part shows the inter-
mediate feature between the temperate and the
subarctic forest, expressing the stronger influence
of the temperate character.
From the viewpoint of the vertical vegetation,
the pure thicket of the Siberian dwarf pine is well
developed in the alpine belt of the mountain
ranges in the north from Central HonshQ (the
main island) through the Be tula Ermani forest.
The subarctic or needle-leaved forest represented
by Abies and Picea follows the Ermarfs birch
forest or directly contact the dwarf pine thickets.
But it is very characteristic to render the forest
t Presented by F.E. Egler.
vegetation in the Japan Sea side by the absence
of the Picea- Abies forest.
The climax forests are as follows in the different
forest zones:
Subtropical rain
forest
Salt marsh
Coast
Warm temperate
forest
Sandy shore
Coastal district
Mountain dis-
trict
Ficus Wightiana
Kandela Camlet
Livistona subglobosa
Shii-Kashi
Pinus Thunbergii
Machilus Thunbergii
Abies firma-Tsuga Sieboldii
Cold temperate forest
Seashore Quercus dentata
Light soil Quercus crispula
Valley Pterocarya rhoifolia-Aescu-
lus twbina- Cercidiphyllum
Japonicum
Needle-leaved
forest
Hokkaido
Picea- Abies
Picea jezoensis- Abies sach-
alinensis
Bog Picea Glchni
Honshu Picea je 20 ens is va r . hondoen-
s is- Abies Veitchii
Northern mixed forest
Sandy shore Quercus dentata
Plain Ulmus propinqua-A cer mono
S wany A Inus japonica
Hill Acer nwno-Tilia japonica
Valley Cercidiphyllum japonicum-
Fraxinus mandshurica-Jug-
lans ailanthifolia
Up to the present, the following three volumes
(Mil) have been published. The last two volumes
(IV- V) are already prepared and will be published
in June of 1958.
I. THE CENTRAL SANYU DISTRICT,
HONSHO
By M. TATEWAKI and T. TSUJll
Res. Bull. Coll. Exp. For. Coll. Agr.
Hokkaido Univ. 18-1. 1 54. (1956)
99
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Hiroshima, famous following the first dropping
of the atomic bomb, was the center of the
research. It is facing the Inland Sea (Setonaikai)
and is situated 3424'N., 13227'E. We studied
a belt which extended from the seaside at Hiro-
shima to the inland mountains that form the
backbone of the Chugoku Region. The distance
is about 42 km, and the rise in vertical elevation
from sea level to the mountains is 1,346 m. Along
the transects under consideration, five experi-
mental localities were studied as follows:
Name of forest
Castanopsis cuspi-
data
Cinamomum Cam-
phora
Castanopsis caspi-
data
A hies firma-Tsuga
Sieboldii
Fagus crenata
Quercus crispula
Tsuga Sieboldii
Name of
Transect
Eleva-
locality
number
tion (m)
Ujina
l.a
20
^
l.b
40
iwakuni
2.a
60
2.b
80
Minochi
3.a
1,000
Nakanoko
4. a
1,000
>
4.b
950
Miyajima
5.a
460
, ,
5.b
500
5.c
470
^
5.d
460
5.e
460
5.f
420
,,
5-g
200
91
5.h
40
,,
5.i
50
Abies fir ma
11. ISLAND OF YAKUSH1MA
By M. TATtWAKI
ibid. 18-2. 53 148. (1957)
Geobotanically, the Island of Yakushima is
one of the most important localities in Japan.
It is extremely interesting, with many endemic
and rare elements. Yakushima is roughly orbi-
cular in shape, about 27.1 km long and 26.7 km
wide, with an area of approximately 544 sq km,
lying between 13023'-40' East Longitude and
30 13'- 28' North Latitute. In the central part
of the island, there are high mountains, such as
Mt. Miyanoura (1,935 m), Mt. Nagata (1,890m)
and Mt. Kuromi (1,836 m). All rivers extend
toward the sea from these central highlands,
and enroute they often form deep gorges.
In 1916, the American botanist Dr. H. Wilson
visited Yakushima and spoke admiringly of the
beautiful primeval forests ofCryptomeriajaponica.
100
Both before and after him, many Japanese bota-
nists have turned their attention to floristic and
vegetational studies of the region, among them,
Profs. G. Koizumi, G. Masamune, S. Hatsushima,
Drs. M. Kawada, K. Imanishi, Mr. Z. Tashiro
and so forth. The results of their important
researches were cited briefly in the paper. The
studies of Prof. G. Masamune are especially
important, for he described the flora in excellent
and precise terms.
The two islands Yakushima and Tanegashima
form a special phytogeographical district of the
East Asiatic Warm Temperate Zone. Tokara
Strait, lying just to the south of Yakushima
island, acts as a line of demarcation between this
zone and the elements at the northern limit of their
distribution, which perhaps is only natural, con-
sidering the unique geographical position of the
islands.
But it is quite remarkable that certain Japanese
elements, which make up a prominent part of the
flora also have their northern limit of distribution
in this region, namely: Cephalotaxus drupacea,
Torreya nucifera, Abies fir ma, Chamaecyparis
obtusa, Castanca crenata, etc.
The causes of this peculiar and somewhat
confined distribution have been attributed to the
presence of high mountains on Yakushima and
deep sea separating the islands from the southern
areas. These geographical barriers may be con-
sidered to be the primary ecological factors
operating against a more widespread occurrence
of the Japanese elements named above. Along
the transects under consideration, nine localities
were studied as follows:
Name of
locality
Kurio
Transect Eleva-
number tion (m)
l.a
l.b
Name of forest
Kandelia Candel
l.c
Mugio
Nabeyama 3. a
Near
Kosugidani 4.a
2.a
40
3.a
20
3.b
20
4.a
800
5.a
1,000
5.b
5.c
6.a
1,000
1,000
1,100
Ficus Wightiana-
Ficus retusa
Machlus Thunder-
&H
Quercus Wrightii
Distylium racem-
osum
Cryptomeria ja-
ponica
6.b 1,100
VOLUME 4
BOTANY
Name of Transect Eleva- Name of forest
locality number tion (m)
Name of Transect Eleva- Name of forest
Near
Kosugidani 7.a 960
7.b 960
7.c 960
Mt. Ishizuka 8.a 1,200
8.b 1,440
Mt.
Miyanoura 9. a 1,760
Abies firma
Tsuga Sieboldii-
Cryptomeria ja-
ponica
Tsuga Siebohlii
Tsuga Sieboldii
Cryptomeria ja-
ponica - Trochoden-
dron aralioides
Cryptomeria ja-
ponica
III. FOREST VEGETATION OF
SOUTHERN KYOSHO
By M. TATFWAKI and T. MISUMI
ibid. 18-2. 149 208. (1957)
In some limited localities in Southern Kytishu,
the forests are characterized by subtropical
vegetation. Pure forest of Kandelia Candel, Cycas
revoluta, and Livistona subglohosa, respectively,
are representative. The latter two forests, occur-
ring in rather small areas, are found at the points
of the peninsula protruding into the Pacific Ocean
washed by warm currents or on the islets sur-
rounded by warm current. The forest vegetation
in the low lands of the district under consideration
predominantly consists of evergreen broad-leaved
trees. The present study was carried out from the
seaside to the mountain to heights of about
1,000 m., ranging from the subtropical forests
to the temperate needle-leaved forest through the
evergreen broad-leaved forests. Along the tran-
sects laid out for study, eight experimental local-
ities were situated as follows:
Name of
Transect
Eleva-
Name of forest
locality
number
tion (m)
Kiire
l.a
Kandelia Candel
Koyama
2.a
520
Distylium racem-
osum
11
2.b
540
11 *1
Bir6-jima
3.a
20
Livistona subglob-
osa
11
3.b
20
11 11
Hosaki
4.a
20
Cycas revoluta
11
4.b
20
11
Toisaki
5.a
20
11 11
Magaya
6.a
140
Cinnamomum Cam-
phora
locality
Aoshima
number
7.a
Mt.
Kirishima 8. a
8.b
8.c
tion (m)
5 Livistona subglob-
osa
990 Abies firma-Tsuga
Sieboldii
1 ,050 Tsuga Sieboldii
960 Abies firma
IV. THE SOUTHERN
SHIKOKU DISTRICT
By M. TAIL.WAKI and T. TSUJH
ibid. 18-2. 149208. (1957)
The dominant feature of the forest vegetation
of Southern Shikoku is represented by "pluvii-
silvae." In several places of the district under
consideration, the natural physiognomy of the
forests has fortunately been kept and poses an
important problem for geobotany in spite of the
comparatively small areas in question.
The general arrangement of the forests will be
explained first. Along the shore, the Pinus Thun-
bergii forest is commonly developed on sandy
beaches. On flat terraces, the Pittosporwn Tobira
forest mixed with Eurya emarginata forms the
front zone against the sea. It attains a height
ranging 1-3-5 m influenced by the wind. The
Camellia ja ponica forest succeeds it and is then
followed by the Machilus Thunhergii forest
inward.
In the Machilus forest belt, Distylum racetnosum,
Podocarpus Nagi\ and Elaeocarpus sylvestris are
sometimes found, forming groves in some areas.
The pure Quercus phillyraeoides forests are gen-
erally found on the rocky slope along the shore.
Nephrolepis cordi folia and Pyrosia Lingua are com-
monly found in the undcrlayer. The Castanopsis
cuspidata forest is also found inland, but it has
lost its primeval physiognomy. Only one place in
Ikku, near Kochi, a rather young natural forest,
was selected for ecological analysis.
The elements of the subtropical forests are well
represented by Fieus Wightiana and Livistona
subglobosa. Their areas are very limited. The
former forest is found in Cape Muroto and Cape
Ashizuri. The latter is found only in Ashizuri,
but the question whether it is an invader from
the south or a remnant is still unsettled. We
consider it as a vestige.
Along the transects under consideration, six
experimental localities were situated as follows:
101
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Name of Transect
locality number
Cape
Eleva- Name of forest
tion (m)
Ficus Wightiana
Quercus phillyrae-
oides
Distylium racemo-
sum
Castanopsis cuspi-
data
Livistona subglob-
osa
Pittosporum Tob-
ira
99
Camellia japonica
Machilus Thunber-
8"
Quercus phillyrae-
oides
Livistona subglob-
osa
V. GEOBOTAN1CAL STUDY ON
THE FAGUS CRENATA FOREST IN
THE DISTRICT OF ITS NORTHERN LIMIT
By M. TATEWAKI, etc.
The Japanese beech, Fagus crenata, is the
representive tree of the cold temperate forests in
Japan. Up to the present, the precise study of
its distribution within the northern limit was
carried out by Tatewaki in 1948. The geobotani-
cal study of the Japanese beech forest of the
district under consideration is a most important
and interesting problem not only from the view-
point of geobotany, but also from that of forestry.
During these ten years, especially during the last
five years, Tatewaki accompanied by the members
of his institute, T. Misumi, T. Igarashi,
S. Watanabe, and S. Kawano, have devoted
themselves to the study of this question under
the auspices of the Hakodate Regional Forest
Office. Along the transects under consideration,
seven localities were studied, as follows:
Muroto
l.a
5
11
l.b
20
99
l.c
100
Tosa Ikku
2.a
80
Kashima
3.a
5
Cape
Ashizuri
4.a
20
4.b
40
4.c
40
99
4.d
100
4.e
100
99
4.f
40
,,
4.g
40
4.h
40
,,
4.i
40
,, ,,
4-j
80
91
4.k
80
99
4.1
130
Yotate-yama
5.a
30
99 19
5.b
30
Misaki
sample plot
20
Name of Transect
Eleva-
Name of forest
locality number
tion (m)
Utasai
.a
60
Fagus crenata
99
.b
100
99 99
Tsubame-
no-sawa
.c
560
M )
99 99
.d
560
91 99
Shirai-gawa
.e
280
99 99
Yamato-
no-sawa
l.f
180
91 19
Mt, Kariba
2.a
740
Betula Ermani
99 99
2.b
680
Fagus crenata
99 99
2.c
540
99 99
99 99
2.d
420
99 99
99 99
2.e
420
>
Mt.
Oshamanbe
3.a
860
Betula Ermani
99 99
3.b
700
99 99
19 99
3.c
720
Fagus crenata
-Betula ermani
99 99
3.d
640
Betula Ermani
-Fagus crenata
Mt.
Oshamanbe
3.e
620
Fagus crenata
99
3.f
620
91 99
3.g
660
11 99
3.h
580
Mt. Ohira
4 a
800
11 99
4.b
340
11 99
4.c
340
19
4.d
320
11 91
4.e
360
11 99
4.f
280
,, ,,
4.g
220
11 91
4.h
280
,,
4.i
100
91
4 j
200
99
4.k
540
99
4.1
360
99
4.m
920
99
4.n
260
V
4.o
320
99 99
4.p
500
99 99
4.q
700
99 99
4.r
600
19 99
4.s
460
4.t
420
99 99
4.u
300
91 99
4.v
700
99 99
4.z
240
102
BOTANY
THE DEVELOPMENT OF FORESf~COMMUNlTIES IN EASTERN ASIAt
CHI-WU WANG
University of Florida y Gainesville, Florida ; U.S.A.
INTRODUCTION
Over the land mass of continental eastern Asia,
forest communities once flourished in an almost
unbroken expanse of over 30 degrees of latitude,
from the tropics to eastern Siberia. Throughout
the immense geographic range there is a gradual
displacement of forest components, with resultant
intergradations and segregations of forest types.
The forest vegetation of continental eastern
Asia comprises the following main types 1 :
(1) The montane coniferous forest formation:
spruce-fir forest, larch forest;
(2) The broad-leaved deciduous forest forma-
tion : the mixed mesophytic forest, the mixed
northern hardwood forest, the deciduous
oak forest, birch forest;
(3) The broad-leaved evergreen forest forma-
tion: the evergreen oak forest, the rain
forest, the littoral forest.
The main types of forest vegetation are arranged
in approximately the latitudinal and altitudinal
sequence as presented in the vegetation map.
Throughout the long geologic past, since the
advent of angiospermous plants in the late
Mesozoic Era, there were always areal and com-
positional differentiations in the vegetation. The
author assumes that all the natural phenomena in
geologic time were governed by laws that govern
the Universe today. The plants of the existing
vegetation, as were plants throughout the ages,
are the direct descendants of pre-existing ones
with which they are genetically most closely
related. Upon this premise, the developmental
concept of plant communities used in this paper
is based. The phylogenetic relationships of the
components of the plant communities offer the
most reliable clue in interpretations of develop-
ment, differentiation and relationship among the
different forest communities, their present pattern
of geographic distribution, and their vicissitudes
during geologic time.
Evidence afforded by the existing vegetation
and that of the geologic past suggests that the
existing forest communities originated from a
type that resembled the modern rain forest. The
differentiated, usually simpler and specialized
types of forest communities, different as they are,
arc linked with the ancestral type represented in
the modern rain forest by a continuum of varia-
tion among phyletic stocks. The modern rain-
forest has been changed through contraction and
spcciation among its components, as have other
forest communities. It still retains the multitude
of phyletic stocks of which the other forest com-
munities are composed and to which they are
linked by a close phylogenetic bond.
The ancestral community, as exemplified by
the existing rain forest, is generally represented
by primitive forms and usually by far more sec-
tions, genera, and even sub-families. In other
words, the rain forest is composed of descendants
from a higher level of plant evolution, and hence
an older one in time sequence, while the other
forest communities are composed of later-differ-
entiated and often specialized forms of the same
phyletic stocks.
The compositional and arcal differentiations of
the forest communities are primarily the results
of two basic processes: historic contraction and
regional speciation. They are the manifestation
of the innate potentialities of all the phyletic
stocks of forest trees ever evolved on the earth's
surface. The extreme paucity of components in
certain simple forests of wide extent, e.g., birch
forest, larch forest, and the multitude of mono-
typic groups and ogliotopic genera in the poly-
phyletic communities, e.g., mixed mesophytic
forest, furnish evidence of the extent of contrac-
tion. On the other hand, the clusters of numerous
closely related species, in both the polyphyletic
and the monophyletic communities, and the geo-
graphic varieties, races, and forms recognized
within population of extensive range bear witness
to the active process of regional speciation.
Through contraction, which widened the gaps
of discontinuity both in character and in range,
distinct species (such as Populus tremuloides)
and subgenera (such as the black oak group,
Erythrobalanus, of North America) and genera
(such as the evergreen beech group, Nothofagus,
of the Southern Hemisphere) likewise have
t Presented by F.E. Egler.
1 The Forest Types of Continental Eastern Asia I, II. 8th Pacific Science Congress, Manila, Philippines.
103
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
resulted from the species clusters. The differen-
tiation of subfamilies and families is only the
prolongation of the process, which in turn trans-
forms simple types of forest communities into
polyphyletic types in time to come.
In the present treatise the segregation and
development of two main forest types, the
mixed mesophytic forest and the mixed northern
hardwood forest, are traced. The closest living
representatives of these two forest types are found
in eastern North America. Frequent references
are made to the forests of eastern North America.
The author believes that the segregation and devel-
opment of forest communities, as visualized
above, is an universal phenomenon and has been
contemporaneous in time sequence throughout
the world. But it is more fully expressed in the
existing vegetation of eastern Asia which extends
from the tropics to the Arctic and which is the
greatest repository of Tertiary relics. The main
types of forest vegetation are not only spatially
and compositionally differentiated entities, but
each of them reflects a common level of plant
evolution. They are linked by a strong bond of
descent upon which the interpretation proposed
in this paper is based.
THE MIXED MESOPHYTIC FOREST
COMPOSITION
The mixed mesophytic forest of eastern Asia is
lloristically the richest in composition of all the
deciduous broad-leaved forests, and is surpassed
in complexity only by the rain forest. Not only
does it include a multitude of tree species, but the
trees represent a large number of remotely related
phyletic stocks. It includes also numerous relics
of an "arcto-Tertiary flora" whose most closely
related modern species are found in the mixed
mesophytic forest of North America. The tree
components of the mixed mesophytic forest of
eastern Asia include more than 10 genera of
conifers and over 50 genera of hardwoods ^ever-
green broad-leaved trees):
Conifers
Cephahtaxus
Cryptomeria
Cunn ingham ia
Cupressus
Nothotaxus
Pinus
Pscudotsuga
Pseudolarix
Taxus
Tor re y a
Tsuga
Broad-leaved trees
A canlhopanax A cer
104
Aphananthe Fagus
Nyssa
Betula Fraxinus
Paulownia
Camptotheca Gymnocladus
Phellodendron
Carya Halesia
*Photinia
*Castanea Hovenia
Pistacia
* Ccistanopsis Idesia
Platycarya
Celt is *IHicium
Prunus
Cercidiphyllum Juglans
Pterocarya
Cladrastis Kalopanax
Pterostyrax
Daphniphy 'Hum Koelreuteria
* Quercus
Davidia Liquidatnbar
Sassafras
Diospyros Liriodendron
Sorbus
Ehretia *Lithocarpus
Tetracentron
Elaeocarpm Maackia
Tilia
Emmenopteris Magnolia
Trema
Eucommia *MangIietia
Ulmus
Euptelea Meliosma
Zelkova
Evodia Moms
The "mixed mesophytic forest climax" of
eastern North America, as defined by Braun (5,
40-41), includes 25 species and varieties of dom-
inant trees in the aboreal layer. They represent
the following genera:
Acer
Fraxinus
Prunus
Aesculus
Halesia
Quercus
Betula
Juglans
Tilia
Carya
Liriodendron
Tsuga
Castanea
Magnolia
Fagus
Nyssa
The following genera, which sometimes appear
in the climax stands, can be added to the above
list:
Cehis
Ckidrastis
Cornus
Gvmnocladus
Ilex
Liquidatnbar
Morus
Oxydendron
Pinus
Plalanus
Robinia
Sassafras
Taxus
Wmus
Aesculus
All these 30 genera are represented in the mixed
mesophytic forest of the Yangtze Valley, except
for Oxydendron, Plalanus, and Robinia.
Braun (4* 5) maintains that the mixed meso-
phytic association of eastern North America is the
"lineal descendant" and persisting remnant of the
"undifferentiated forest of the Tertiary," and the
most complex and oldest association of the
deciduous forest formation. It occupies a central
position in the deciduous forest as a whole; and
from it or its ancestral progenitor (the mixed
Tertiary forest) all other climaxes of the decidu-
ous forest have arisen.
How satisfactory this center of radiation theory
for the mixed mesophytic forest is, will be verified
VOLUME 4
BOTANY
by further studies of my American colleagues.
It is known, however, that the Tertiary forest of
the middle latitudes of eastern North America was
far richer in tree components than the existing
mixed mesophytic forest. It included the follow-
ing genera that do not exist in the present forest
(Wilcox flora, Eocene) :
* Acacia
*Comhretum
*Pistia
*Anona
*Cordia
*Pithecolohium
Ar Wear pus
*Cryptocurya
Psycho tria
*Avicennia
* Dalbergia
* Sap Indus
*Berchemia
*Diospyros
Schefflera
*Buttneria
* Engelhardtia
*Sideroxylon
*Caesalpinia
* Eugenia
*Staphyleu
*Canavalia
*Ficus
*Sterculia
*Canna
Glvptostrobus
*Terniina/ia
* Cassis
*Oreodaphne
Zygophvllum
*Cedrela
*Osmanthus
*Cinnamomum
Paliurus
The fossils assigned to " Dryophyllwn" (3) are,
according to Sharp (18), more closely related to
modern Mexican oaks. Representatives of these
genera now exist in the broad-leaved evergreen
forest region of eastern Asia, and the majority of
them (*) still can be found in the southern part of
eastern North America south of the mixed meso-
phytic forest region, mostly in Florida and
Mexico (18).
The present mixed mesophytic forest of eastern
North America is primarily a deciduous commu-
nity. If it is the lineal descendant of the Tertiary
forest, as suggested by Braun, the evidence indi-
cates that the existing forest has been changed
considerably from its progenitor.
HISTORIC CONTRACTION
The most obvious change in the composition
of the mixed mesophytic forest has been the
nearly total (as in eastern North America) or
partial (as in eastern Asia) elimination of ever-
green broad-leaved trees. This change resulted
in a transformation into an essentially deciduous
community, such as the mixed mesophytic forest
of today, from a forest community that was
dominated by, or at least, included a considerable
portion of evergreen broad-leaved trees in the
arboreal layer. The modern equivalents of these
Tertiary evergreen broad-leaved trees are mostly
associated with the rain forest of today.
This trend, as indicated by fossil evidence, is
also suggested by the components of the existing
forest. The fact that the greatest concentration of
epibiotics is in the mixed mesophytic forest is of
special significance as a clue to the historic
contraction of this community.
Most of the epibiotics of the mixed mesophy-
tic community are represented by monotypic gen-
era, or by monotypic family, such as:
Amentotaxus
Bretschneidcra
Cephalotaxus
Ccrcidiphyllum
Davidia
Decaisnea
Dipt crania
Euptelca
Emmenopterys
Halesia
Huodendron
McUiodendron
Metasequoia
Nothotaxus
Pseudolarix
Rhoiptelea
Sargentodoxa
Sinofranchetia
Sinojackia
Sinowilsonia
Taiwania
Tetracentron
Torricellia
The above-mentioned plants include only those
epibiotics of regional distribution which charac-
terize the geographic intergradations within the
mixed mesophytic community. The epibiotics
arc not all necessarily restricted in range, however.
The following include those that are distributed
in the entire region of the mixed mesophytic forest
and beyond. They are either monotypic groups
or oligotypic "polytopic genera," i.e., small genera
composed of only a few geographically disjunc-
tive species, usually isolated in the remote parts of
the world.
(1) Monotypic families: In addition to Amento-
taxaceae, Bretschneideraccae, Cercidiphyllaceae,
Metasequoiaceae, and Rhoipteleaceae mentioned
above, are: Ginkgoaceae (Ginkgo) , endemic to the
mixed mesophytic region of the Yangtze Valley,
and Eucommiaceae ( Eucommia), Yangtze Valley
and the transitional zone of the Northern Pro-
vinces.
(2) Monotypic genera: In addition to those
mentioned in the last section (Davidia, Nothot-
axus, Pseudolarix, Taiwania, Tetracentron), are
the following:
Bishchojia
Camptotheca
Cryptomeria
Eusc aphis
Fokienia
Fortunearia
Idesia
Platycarya
Poliothyrsis
Pteroceltis
Tapiscia
(3) Poly topic genera: The following include those
polytopic genera that now occur in more or less
the entire range of the mixed mesophytic commu-
nity. Those genera that have persisted only in
certain parts of this region were discussed in the
preceding section.
105
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Aphananthe Eastern Asia and Australia 3-4
species, China 1
Buckleya Continental eastern Asia 3, east-
ern North America 1
Chionanthus Continental eastern Asia 1, east-
ern North America 1
Cladrastis Continental eastern Asia 2, Japan
1 , eastern North America 1
Cunninghamia Yangtze Valley and farther
south 1, Taiwan 1
Gymnocladus Continental eastern Asia 1, east-
ern North America 1
Liquidamhar Eastern Asia 1, western Asia 1,
North and Central America 1
Liriodendron Continental eastern Asia 1, east-
ern North America 1
Nyssa Continental eastern Asia 1, east-
ern North America 4, Himalaya
to Java 1
Pseudotsuga Continental eastern Asia 2, west-
ern North America 3-4
Sassafras Continental eastern Asia 1 , Tai-
wan 1, eastern North America 1
The multitude of relics in the mixed mesophy-
tic forest is in sharp contrast to their extreme
paucity in the areas just to the north, which
are occupied by plant communities developed
over a new land surface and in a new habitat
resulting from climatic changes in relatively recent
geologic history. This fact seems to indicate
that the mixed mesophytic forest is situated at the
periphery of the selective pressure which has
been effective enough to decimate part of the
population, and yet favorable enough for the
survival of the rest- a fact which possibly explains
the preservation of the numerous ancient relics
and the extremely rich iloristic composition of
the mixed mesophytic forest.
Braun (5) aptly observed that the comparable
community of North America is "the most
complex and the oldest association of the Deci-
duous Forest Formation." The complexity of
this type of forest, however, is not due simply
to the multiplicity of tree species. Strikingly
indeed, it is composed of many t4 one-of-a-kind"
elements. The 60 species of crown trees that are
found throughout the entire range of the mixed
mesophytic community represent no less than
50 genera. This fact and the presence of the
numerous monotypic groups and polytopic genera
suggest that the mixed mesophytic type, complex
as it seems to be, is but an impoverished relic of a
floristically much richer community.
106
TROPICAL ORIGIN
The original composition before the impover-
ishment included more genera that are related
to the trees of the rain forest, as suggested by the
history of the mixed mesophytic forest of eastern
North America. In the present forest of eastern
Asia the rare occurrence of such genera as
Podocarpus, Torrcya, lllicium, Phoebe, Manglietia,
and evergreen oaks (Quercus, Castanopsis, Pas-
ania) suggests similar conditions.
The comparative morphology of angiosperms
(19, 1 ) suggests that the deciduous trees in general
arc derived from evergreen stocks which are close
to the more primitive forms. The major compo-
nents of the mixed mesophytic forest, which are
naturally all deciduous trees, are nearly all repre-
sented in the rain forest by evergreen trees of
closely related genera or species. In fact, most of
the trees of the mixed mesophytic forest as listed
below are deciduous representatives of phyletic
stocks which are predominantly evergreen, viz:
( Diospyros)
( Elaeocarpus )
( Daphniph vllum, Mal/otus)
(Poliothyrsis, Idesia)
(Llquidambar , Forlunearia,
Sinowilsonia)
(Sassafras, Under a, Lit sea)
( 1. iriodendron , Telracen iron )
( Morus)
( Da vi did)
(Chionanthus, Fraxinus)
( Emmenopterys)
(Evodia, Phvllodendron )
(Buckleya) '
( Koelreuteria)
(Tapis c\a, Euscaplus)
(Halesia, Sinojackia, Huo-
dendron, Rehderodendron,
AlniphyUum)
Two species of the genus Castanea in the mixed
mesophytic forest are evergreen. Even Acer,
which is represented by over 50 species of decidu-
ous trees in eastern Asia, and the Boraginaceae,
which is essentially a family of herbaceous plants
of the arid regions, are both represented by decidu-
ous trees (deciduous Acer and Ehretia) in the
mixed mesophytic forest, but by evergreen trees
in the tropics (section of entire leaved maples,
Cordia Ehretia, and Tournefortia). The woody
boraginaceous plants represent the primitive
section of the family (9) .
The closest kin of the deciduous trees of the
mixed mesophytic forest can nearly all be traced
Ebenaceae
Elaeocarpaceae
Euphorbiaceae
Flacourtiaceae
Hamamelidaceae
Lauraceae
Magnoliaceae
Moraceae
Nyssaceae
Olcaceac
Rubiaceae
Rutaceae
Santalaceae
Sapindaccae
Staphyleaceae
Styracaceae
VOLUME 4
BOTANY
to the rain forests of the tropics, which usually
include more tribes, genera, and sections of the
phyletic stocks to which the deciduous trees of
the mixed mesophytic forest are related in a
continuum of variation. The weight of evidence
indicates that the mixed mesophytic forest has
emerged from the northern fringe of a type of
evergreen broad-leaved forest similar to the
present rain forest. The evergreen components
were gradually eliminated, probably in late
Tertiary. Through further contraction and
speciation the multitude of relics and numerous
closely allied species of certain large genera of
deciduous trees which characterize the present
mixed mesophytic forest were differentiated.
THE MIXED NORTHERN
HARDWOOD FOREST
The mixed northern hardwoods forest, in conti-
nental eastern Asia is situated between the boreal
coniferous forest farther to the north and the
deciduous oak forest of the relatively arid North-
ern Provinces, which in turn separates the mixed
northern hardwoods from the mixed mesophy-
tic forest of the Yangtze Valley. Despite the
discontinuity, the mixed northern hardwood
forest is closely related to the mixed mesophytic
forest in composition. It appears to be a severely
depauperized form of the mixed mesophytic
forest.
DEPAUPERIZED MIXED MESOPHYTIC FOREST
The primary components of the mixed northern
hardwoods are Acer (8 species), Tilia (5 species),
and Betuht (9 species), intermixed with consider-
able proportions of white pine (Pinus koraiensis),
Quercus, Fraxinus, Juglans* Maackia, Phelloden-
dron, and Ulnms. This forest also includes, as
minor components, trees of the following genera:
Alnus Mains Pyrus
Carpinus Moms Salix
Celtis Populus Sorbus
Kalopanax Prunus Zelkova
Taxus, Magnolia, and shrubby Cornus and Lindera
also occur as rare accessories.
In comparison with all the other types of deci-
duous broad-leaved forests, the mixed northern
hardwoods is, next to the mixed mesophytic, the
richest in composition. All the above mentioned
26 genera are common to both types. However,
the specific representative of these genera usually
are not the same. Furthermore, there are many
more genera (over 40) of evergreen and deciduous
trees among the primary and minor components
of the mixed mesophytic forest that are not repre-
sented now in any other type of deciduous forest.
But some of them are known to have occurred
in the northern regions in the Upper Tertiary.
This fact shows the gradual retraction of the range
of certain "southern" genera and the depaupe-
rization in components of the northern lobe of a
more extensive mixed mesophytic forest which
once flourished in the northern region. This
process resulted in the simpler type of forest
community of deciduous broad-leaved trees now
designated as the mixed northern hardwood forest.
All the genera of the forest components, without
exception, are included in the present mixed
mesophytic forest, and some of the eliminated
components are preserved in the Tertiary fossils
of this region.
With the exception of Tsuga and Fagus, all the
genera of the primary forest components of this
community in North America are represented in
the mixed northern hardwood forest of eastern
Asia, which is somewhat richer in composition
than its North American counterpart. Common
forest components such as Phellodendron, Maac-
kia, and Zelkova and a common climber, Actinidia,
are not known in North America. Schizandra
is represented in North America only in the South.
Recent interspecific hybridization (8) between the
white pine of the Northeastern Provinces of
China (Pinus koraiensis) and the eastern white
pine of North America (Pinus strobus), both of
which are important components of the respec-
tive mixed northern hardwoods, gives almost
100% fertility. The striking compatibility of the
two vicarious species furnishes genetic evidence
for the close affinity of the two similar but widely
disjunct forest communities.
POST-PLEISTOCENE EXPANSION
An entirely different interpretation of the
formation and the present distribution has been
proposed for the mixed northern hardwood forest
of eastern North America. According to Braun
(5, p. 533), the vegetation of the mixed northern
hardwoods is "the result of post- Wisconsin migra-
tions, which brought about an expansion from the
unglaciated Allegheny Plateau and northern
Allegheny mountains (physiographic sections of
Appalachian Plateaus) without pronounced modi-
fication of type. The climax elements are almost
entirely a result of these expanding migrations."
In the interpretation of the present distribution
of the deciduous forests of eastern North America,
great pains were taken to correlate the physio-
graphic history with forest development, particu-
larly the Pleistocene glacial boundary. The ice
107
PROCFEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
advanced across the Appalachian Plateau, the late
Tertiary haven of the mixed forest. From fossil
evidence, Braun (5, pp. 512-513) deduced that
"nowhere far beyond the glacial boundary was
climate during the glacial stages sufficiently severe
to displace occupying vegetation," and this
ensured continuous occupancy by the late Tertiary
mixed forest, the progenitor of the present mixed
mesophytic forest, on the unglaciated part of the
Appalachian Plateau and a small isolated ungla-
ciated area (the Reading Prong and the New
Jersey Highlands) farther east. Braun (5, p. 529)
maintains "Continuity of occupation since early
Tertiary time accounts for the antiquity of its
vegetation ; lack of extreme changes, for the con-
ti nuance of one climax type; and a consistently
humid climate (with rare lapse to subhumid)
for the prevalence of this climax/' The explana-
tion of the present forest distribution as the result
of post-Pleistocene expansion and segregation of
the vegetation from the unglaciated areas to the
glaciated areas where the former vegetation was
removed by the advancing ice, is plausible enough.
MIGRATION AND SURVIVAL
However, it does not necessitate the continuous
occupancy of the late Tertiary mixed forest not
far from the front of the continental ice sheet, an
assumption that is still open to debate. Pcrigla-
cial phenomena such as block-fields and thick
surficial deposits moved by soli-fluction probably
of Wisconsin age are found in the higher parts of
the Appalachian Highlands as far south as the
Great Smoky Mountains (6,600 ft, circ. Lat.
35 N.) where the cove hardwoods type (circ.
3,000 - 3,500 ft) is considered as the most typical
of the mixed mesophytic communities. At the
present time, areas in the subarctic or in high
mountains where similar periglacial phenomena
are actively forming are all essentially treeless.
By analogy, Denny reasons that the Appalachian
Highlands were likewise essentially devoid of
forest when such periglacial phenomena were in
action ( 1 ).
This inference does not entirely preclude the
continuous occupancy of the late Tertiary mixed
forest in the unglaciated areas still farther away
from the glacial boundaries. Tt does indicate,
however, that the Tertiary mixed forest of eastern
North America, like the similar type of eastern
Asia, has undergone severe displacement, espe-
cially in regions close to glaciations. It also
suggests that to a considerable extent the present
mixed mesophytic forest is the result of post-
108
Pleistocene migrations from areas farther to the
south.
There are indisputable evidences of Quater-
nary glaciations in the mixed mesophytic forest
region of eastern Asia (13, 14, 15, 2, 21) . Glaci-
ated areas include: Lushan (1,480 m, Lat.
2930' N.), Chiuhuashan (900 m, Lat. 3032' N.),
Tienmushan (Tienmongshan, 1,547 m, Lat.
3025' N.), and Huangshan (1,700 m, Lat. 30
10'N.). These mountains are situated in the
southern part of the Lower Yangtze Valley, not
too far from the coast.
The Lower Yangtze Valley, which is now occu-
pied by mixed mesophytic forest, was glaciated
not once but three times. In the Poyang glacia-
tion (the oldest and also the largest) and the next,
the Taku glaciation, piedmont glaciers were
formed. The third, the Lushan glaciation, pro-
duced only small glaciers in the higher mountains.
These glaciations were separated by interglacial
periods of genial or even subtropical climate,
resulting in the high oxidation of the ferruginous
soil and incipient lateritization of the boulder-clay-
like deposit and the Red Loam (76, chapter on
Pleistocene climate).
Despite the successive glaciations, the existing
vegetation of the above-mentioned glaciated
mountains, as evidenced by the preserved natural
forests, is of the mixed mesophytic type which
includes not only nearly all the tree genera of the
mixed mesophytic forest of the unglaciated Appa-
lachian Plateau, but also survivals from those
genera that are known to have occurred in the
late Tertiary mixed forest of eastern North
America and to have perished subsequently.
There are distinctions between forest types and
forest regions. The mixed forest of the Miocene
whose components were almost identical with
those of the present mixed mesophytic forest but
which occurred more than 500 kilometers north
of the northern limit of the present type, has been
replaced by the present deciduous oak forest.
It is then reasonable to infer that in the Quater-
nary glaciations the areas that are now occupied
by mixed mesophytic forest, but were at that time
glaciated or at least exposed to periglacial condi-
tions, could not possibly have been occupied by a
type that was similar to the present mixed meso-
phytic forest. The presence of the mixed meso-
phytic forest, no matter how complex or ancient
the flora may be, does not imply that the area has
never been glaciated nor does it necessitate the
continuous occupancy of this type in that parti-
cular area.
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BOTANY
PRE- PLEISTOCENE CONTINUITY AND REGIONAL
SPECIATION
Nor is it necessary to assume the total elimina-
tion of forest communities and to preclude the
possible persistence of a depauperized form of the
late Tertiary mixed forest farther to the north,
particularly in the maritime regions where the
influence of warm currents prevailed. The mixed
northern hardwoods forest, Braun maintains, was
almost entirely a result of post-Wisconsin migra-
tion so far as the climax elements are concerned.
To her disadvantage, the forest flora of the
deciduous forests of eastern North America with
which she was dealing, rich as it is, is relatively
simple in comparison with that of eastern Asia.
The geographic intergradations in the whole range
of the American deciduous forests are not well
expressed by the regional speciation of their
components. Of the 25 species and varieties listed
as "dominant trees of the arboreal layer" of the
mixed mesophytic forest of eastern North Ame-
rica, all are represented in other "climaxes of
deciduous forest." Gymnocladus, Halesia, Liqui-
dambar, and Oxydendron are perhaps the only
tree genera of the mixed mesophytic forest that are
not represented in the deciduous forests of the
North. Not only are most of the genera in com-
mon, but their representatives in the primary
components of both the northern hardwoods
and the mixed mesophytic forest are of the same
species, which extends in a continuous range. This
fact seems to have led Braun to the conclusion
that all the other deciduous forest climaxes have
arisen from the centrally located "undifferentiated
mixed mesophytic forest," supposedly the oldest
association of the deciduous forest formation
preserved in the Appalachian Plateau.
In a forest flora like this, evidence of geo-
graphic differentiations could be found at the
subspecific level, e.g., the local races and eco-
types. Recent studies of the American beech
(6), ash (22, 23) walnut (24), maple (10), hem-
lock (17) demonstrate that forest trees are made
up of a number of more or less distinct types with
well-differentiated ranges.
In the deciduous forest communities of eastern
Asia, on the other hand, geographic differentia-
tions are well expressed on the species level. The
mixed mesophytic forest and the mixed northern
hardwoods in particular, are not only richer in
components, but the tree genera of the mixed
northern hardwoods that are in common with the
mixed mesophytic forest are represented mostly
by different species. In other words, distinct re-
gional speciation has taken place along with the
differentiation of forest types. The interpreta-
tion advanced by Braun of the mixed northern
hardwoods of eastern North America cannot
satisfactorily explain this phenomenon. The
mixed northern hardwoods forest of eastern Asia
cannot possibly be entirely the result of post-
Pleistocene expanding migrations from un-
glaciated areas south of the ice front :
(1) There are only a few species of trees in the
mixed northern hardwoods that are connected
with the mixed mesophytic forest in a more or less
continuous range. Most of the other non-endemic
species of the primary components are generally
restricted to the north and rarely occur south of
the deciduous oak forest region. These few wide-
ranging trees occur both in the mixed mesophytic
forest of the south (MM), the deciduous oak
forest of the northern Provinces (N), and the
mixed northern hardwoods of the Northeastern
Provinces (NE). Some of them extend as far as
Japan (J), Korea (K), eastern Siberia (S), and
Sakhalin (SK). This group of wide-ranging trees
includes the following:
Acer mono (A. pic turn var. parviflorum) (MM,
N, NE, K, J)
Betula japonica (B. mandshuricd) (MM, N, K,
J, S, SK)
Carpinm cordata (MM, N, NE, K, J, S)
Fr ax inns chinensis (MM, N, NE)
Populus iremula var. davidiana (MM, N, NE)
Ulmus japonica (MM, N, NE, K,J)
V.pumila (MM, N, NE, S)
A number of lesser plants can be added to this
list, e.g., Schizandra chinensis (MM, N, NE, K, J,
S) and Actinidia arguta (MM, N, NE, K, J, S).
The true mistletoe, Viscum album, unlike the
American form, Phoradendron Flavescans, extends
from the mesophytic forest to the boreal regions
(Siberia) with a total disregard for glacial bound-
aries.
(2) The majority of the primary components
of the mixed northern hardwoods are tree species
of a northern range which rarely if ever appear in
the mixed mesophytic forest region. A number of
these northern trees are endemic to the Northeast-
ern Provinces and the adjacent part of Korea
where this type of forest is extensive, or restricted
to the general region of the Northeastern Pro-
vinces (NE), Korea (K), Japan (J), and eastern
Siberia (S). Some of them extend even to
Sakhalin (SK). This group of trees includes the
following:
Acer barbinerve (NE, K)
A. Mandshurica (NE, K)
A. pseudo-sieboldianum (NE, K)
109
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
A. te%mentosum (NE, K)
A. triflorum (NE, K)
Alnus hirsuta (NE, K, J, S)
Be tula cos tat a (NE, K, S)
B. er/wfl/i/ii (NE, K, J, S)
B. platvphvlla (NE, K, S)
B. Schmidtii (NE, K, J, S)
Fraxinus mandshurica (NE, K, J)
Maackia amurensis (NE, K, J)
Phellodendron amurensis (NE, K, S, SK)
Sorbus amurensis (NE, K, S)
7Y//0 amurensis (NE, K, S)
7. megaphylla (NE)
Pinus koraiensi\ (NE, K, J)
Ttfjotf cuspidata (NE, K, J, S, SK)
Magnolia sieboldii (NE, J, K, S), Cornus alba
(NE, K, S), K/Vw amurensis (NE, K, S), and a
number of shrubs and lesser plants can be added
to this list. None of them extends to the mixed
mesophytic forest region from which the mixed
northern hardwoods is separated by the decidu-
ous oak forest region.
(3) The discontinuity of the two allied forest
types, the mixed northern hardwoods and the
mixed mesophytic, is paralleled by the similar
disjunct occurrence of the following genera:
Alnus (MM, NE, K, J, S)
Lindera(MM, NE, K, J)
Maackia (MM, NE, K, J)
Phellodendron (MM, NE, K, J)
Taxus(MM, NE, K, J, S)
The following plants occur in both the mixed
northern hardwoods and the mixed mesophytic
forest, but not in the intervening area:
Acer caudatum (Himalaya, MM)
A. Caudatum var. ukwunduense (A. ukurun-
duense) (NE, K, J)
A. ginnala (MM, NE, K, J)
Actinidia kohmicta (MM, NE, J, K, S)
A. polygama (MM, NE, K, J)
Lindera mollis (MM, NE)
Kalopanax pic turn (MM, NE, K, J)
Phellodendron sachalinense (MM, K, J, SK)
Samp locos paniculata (MM, NE, J)
The distribution patterns of these three groups
indicate progressive contraction in an extensive
mixed mesophytic type of forest which once
extended in a continuous range to the present
mixed northern hardwoods region of the North-
eastern Provinces and the adjacent maritime areas
of eastern Siberia. This disjunction of some of
110
the components probably is due to the increased
continentality in the intervening area. Tertiary
fossil deposits show the actual existence of the
pre-Quaternary continuity of this type of mixed
forest. Trees almost identical with living ones
in the present mixed northern hardwoods and the
mixed mesophytic forest once occupied the inter-
vening area.
In addition to the compositional affinities, the
pre-Pleistocene continuity of a mixed hardwood
forest community, and the present disjunct
range of those components that are common to
both the mixed northern hardwoods and the mixed
mesophytic forest, favor the explanation that the
present mixed northern hardwood forest is a
depauperized form of a pre-Pleistocene mixed
hardwood forest community that was similar
to the present mixed mesophytic forest. The
prc-Pleistocenc community extended over a
continuous wide range with an essentially homo-
geneous composition but with well-developed
geographic differentiations of tree species. Region-
al speciation perhaps resulted in the numerous
endemic species in the subsequently disconnected
and depauperized forest communities, although
all the genera of these endemic species and, in
fact, all the genera, without exception, of the
primary components of the present mixed north-
ern hardwoods are represented in the present
mixed mesophytic forest.
The presence of the endemic species, parti-
cularly those that are restricted to this region
and the adjacent Korea- Japan-eastern Siberia-
Sakhalin areas, precludes a post-Pleistocene
origin of the mixed northern hardwood forest.
If the present mixed northern hardwoods forest
were the result of post-Pleistocene expanding
migration from unglaciated areas, as proposed by
Braun for the mixed northern hardwood forest
of eastern North America, then the amelioration
of climate should be such that the entire region,
including the intervening area, would be suffi-
ciently favorable for the continuous extension of
a type of mixed hardwood community that
connected both the mixed northern hardwoods
and the mixed mesophytic forest in gradual
intergradations.
The establishment of a forest community
including such disjunct plants as Lindera, Symplo-
cos, Taxus, Maackia, Phellodendron, and a few
species of maples in the intervening area a
condition which the theory of post-Pleistocene
expanding migration requires would necessitate
a substantial increase in precipitation and a
milder climate than the prevailing one.
VOLUME 4
BOTANY
If it is assumed that the mixed northern hard-
wood forest of the entire range is of post-Pleisto-
cene origin, then the implication is that the com-
ponents common to this region have not only
migrated to Siberia but have also crossed a land-
bridge to Japan and Sakhalin. A direct land
connection between the mainland and Japan and
Sakhalin Island was possible at the height of the
Ice Age when the sea level was sufficiently
lowered. However, such a land connection could
not have been in existence when the climate
became sufficiently warm and humid to effect
the supposedly post-Pleistocene expanding mi-
gration.
On the other hand, if it is assumed that only the
mixed northern hardwood forest of the mainland,
i.e., the Northeastern Provinces (Manchuria)
and eastern Siberia, is the result of post-Pleis-
tocene expanding migration, and that similar
forests in Japan and Sakhalin are relics of the
Tertiary, then the persistence of these forests in
Japan, and particularly in Sakhalin which is
situated farther north and is more rigorus in
climate, serves only to prove the possible survival
of the mixed northern hardwood forest. This
forest includes a number of the same species of
trees in at least the maritime regions of the
continent just across the sea.
Pleistocene fossils also indicate that the present
mixed northern hardwood forest is the depauper-
ized form of a Tertiary mixed hardwood forest
rather than that it is of post-Pleistocene origin.
Zelkova (MM, N, NE, K, J), now extinct in Sibe-
ria, was growing with Ginkgo in the early Pleis-
tocene on the Amur River, the frontier between
the Northeastern Provinces and Siberia (11).
Juglans mandshurica (N, NE, K, S) now occurs
on the mainland only, but was found in the
Pleistocene deposits of Japan (12). The fossil
record shows that a progressive reduction in
components and shrinkage in their range had
taken place. The continuous occurrence of a
mixed deciduous hardward community which
was slightly richer in composition and which had
a more northerly limit than the present mixed
northern hardwood forest is evidenced by fossil
remains to have occurred as late as the early part
of Quaternary.
COMPLEMENTARY EVIDENCE IN LOWER PLANTS
So far, the evidence used in the discussion of
forest development has been limited as much as
possible to the primary forest constituents, with
only occasional excursions into the lesser growth,
so that the main thesis can be presented with
clarity and effect, without unnecessary reference to
long lists of plants which have already been a
burden to the reader. In the mixed northern
hardwoods forest now under discussion, certain
lower plants offer complementary evidence for
the continuous persistence of the type during
the Pleistocene in this general region. The rare
occurrence of Hymenophyllum and Trichomanes
in the present mixed northern hardwoods and
coniferous forests on the frontier between Siberia
and the Northeastern Provinces is of special
significance.
Hymenophyllum and Trichomanes, are two
genera (sen. Lat.) of Hymenophyllaceac which
include 400-500 species of filmy ferns, mostly in
shaded habitats of the humid parts of the tropics.
In North America, one species (Trichomanes
hoschianum) reaches as far north as Kentucky
and Illinois. Three species occur in Europe.
The numerous species of Hymenophyllum and
Trichomanes of eastern Asia are concentrated in
the evergreen forest region. They occur only
occasionally in the mixed mesophytic forest, and
have been observed in the exceedingly wet valleys
of western Yunnan in Abies-Picea- deciduous
hardwood forests. Over the great distance
between the Yangtze Valley and the Siberian
frontier, the Hymenophyllaceac have never been
observed nor are they likely to be found. They
sometimes reach high latitudes in both hemi-
spheres, but only under the conditions of maritime
climate, e.g., Hymenophyllum tunbridgense on
the Faroe Islands (Lat. 63' N.) in the North
Atlantic.
Kryshtofovich (II, 12) and Vorobiov (20)
reported the discovery of Trichomanes parvulum
and Hymenophyllum \\rightii in the Suchan River
basin of eastern Siberia (Ussuri), close to the
Northeastern Provinces, a considerable distance
from the coast. Hymenophyllum wrightii was
previously known from Sakhalin and Japan (as
far north as Hokkaido), but not from Korea
proper except on Quclpart Island.
In eastern Siberia, Hymenophyllum and Tricho-
manes grow together on a cliff within a virgin
forest of Picea ajanesis, Abies nephorlepis, and
Taxus cuspidata. A rare fern, Pleurosoriopsis
makinoi, is found nearby. In this general region,
Vitis amurensis, Schizandra chinensis, Actinidia
spp., Phellodendron amurense, and Juglans mand-
shurica may still be found. Kryshtofovich
concluded that they were survivals of the Tertiary
forest.
The forest around the growth of Hymenophyl-
lum on Sakhalin consists mainly of Picea ajanensis,
111
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
and Abies sachalinensis, with admixtures of Betula
japonica, Taxus cuspidata, and Euonymus sachali-
nensis. The soil is thickly covered with Osmunda
cinnamomea and Athyrium pterorhachis. In
places, Ilex rugosa covers the soils with a thick
layer.
The forests of western Yunnan in which
hymenophyllaceous ferns occur are thousands
of kilometers away from the Siberian coast;
nevertheless, the composition is essentially the
same, including Abies delavayi, Picea complanata,
Taxus chinensis, Betula caudate, B. albosinensis,
Sorbus spp., and Magnolia globosa, A variety of
Betula japonica (var. szechuanica) is known to
this region. The consistency of the composition
of the kind of forest in which hymenophyllaceous
ferns are associated in such widely disjunct areas
suggests the probable conditions of the relic forest
that survived the Pleistocene Epoch in the mari-
time regions of eastern Siberia and adjacent
regions.
In the above discussions on the segregation and
development of forest communities in continental
eastern Asia, frequent references have been made
to the similar type of forests in eastern North
America. This comparison deserves critical
examination because two diametrically different
conclusions were reached from observations of
two forests that are exceedingly alike in com-
position, developed under essentially similar
habitat conditions, and situated in approximately
the same latitudinal ranges.
The differences in opinion do not imply the
author's approval or disapproval of the theory
advanced by his American colleague. The author
realizes that the forest communities of the two
land masses are similar but not the same. How-
ever, the author does believe that the basic pro-
cesses that shaped the existing vegetation, as
revealed by available evidence and presented in
this treatise, are universal and contemporaneous
in the general time sequence, but the extent of
contraction and speciation has involved regional
differences.
DISTRIBUTION or THE MAIN TYPES OF NATURAL
VEGETATION OF EASTERN ASIA
112
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BOTANY
REFERENCES
(1) Bailey, I.W. and E.W. Sinnott, 1916, The
Climatic Distribution of Certain Types
of Angiosperm Leaves, Am. J. Dot., 3:
23-39.
(2) Harbour, G.B., 1934, Analysis of the Lushan
Glaciation Problem, GeoL Soc. China
Bull., 13: 647-656.
(3) Berry, E.W., 1930, Revision of Lower
Eocene Wilcox Flora, U.S.G.S. Prof.
Paper 156: 1-196.
(4) Braun, E.L., 1947, Development of the Deci-
duous Forests of Eastern North Ameri-
ca, Ecol Monograph, 17: 211-219.
(5) Braun, E.L., 1950, Deciduous Forests of
Eastern North America, Blackisten Co.,
Philadelphia.
(6) Camp, W.H., 1951, A Biogeographic and
Paragenetic Analysis of the American
Beech (Fagus), Yearbook, Am. Phislo-
sophical Soc., Philadelphia 1950: 166-
169.
(7) Denny, C.S., 1951, Pleistocene Frost Action
near the Border of the Wisconsin
Drift in Pennsylvania, Ohio J. ScL, 51
(3): 116-125.
(8) Johnson, A. G., 1952, Personal Communica-
tion.
(9) Johnston, T.M., 1951, Studies in the Bora-
ginaceae XX. Representatives of Three
Subfamilies in Eastern Asia (Cordioi-
deae, Ehretioideae, and Heliotropioi-
deae), /. Am. Arb., 32 (1): 1-26.
(10) Kriebel, H.B., 1956, Patterns of Genetic
Variation in Sugar Maple (unpublished
Doctoral Dissertation).
(11) Kryshtofovich, A.N., 1 930, Hemenophyllum
wrightii V.D.B. on the Sakhalin Isl.,
Bull, dn Jard. Bot. Principal de L'UESS
f., XXIX, L/vr., 3-4: 1412.
(12) Kryshtofovich, A.N., 1935, Hemenophyllum
and Trichomanes the Ussuriland of the
USSR. Sunyatsenia, 3: 22-25.
(13) Lee, J.S., 1933, Quaternary Glaciation in
the Yangtze Valley, GeoL Soc. China
Bull., 8: 15-44.
(14) Lee, J.S., 1934, Data relating to the study
of the Problem of Glaciation in the lower
Yangtze Valley, GeoL Soc. China Bull.,
8:395-422.
(15) Lee, J.S., 1936, Confirmatory evidence of
Pleistocene Glaciation from the Huang-
shan, Southern Anhui, GeoL Soc.
China Bull., 15:279-290.
(16) Lee, J.S., 1939, The Geology of China, Lon-
don.
(17) Olson, J.S. and H., Hiensteadt, 1957, Pho-
toperiod and chilling control Growth of
Hemlock, Sciences, 125 (3246): 492-494.
(18) Sharp, A.J., 1951, The Relation of the
Eocene Wilcox Flora to Some Modern
Floras, Evolution, 5(1): 1-5.
(19) Sinnott, E.W. and Bailey, I.W., 1915, Inves-
tigations on the Phylogeny of the Augio-
sperms 5: Foliar Evidence as to the
Ancestry and Early Climatic Environ-
ment of the Augiosperms, Am. J. BoL,
2: 1-22.
(20) Vorobiov, P.P., 1933, A Discovery of
Hemenophyllum wrightii V.D.B. in the
Suchnan Region of the District, Bull.
Far East Branch, A cad. Set. ISSR, 1-2-2:
141.
(21) Wissmann, H. Von., 1937, The Pleistocene
Glaciation in China, GeoL Soc. China
Bull., 17: 145-168.
(22) Wright, J.A., 1944 A, Genotypic Variation
in White Ash, J. Forestry, 42 (7):
489-495.
(23) Wright, J.A., 1944B, Ecotypic Differentia-
tion in Red Ash, J. Forestry, 42 (8):
591-597.
(24) Wright, J.A., 1954, Preliminary Report on a
Study of Races in Black Walnut, J.
Forestry, 52 (9): 673-675.
113
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
DYNAMICS OF ATOLL VEGETATION
F.R. FOSBERG
Pacific Vegetation Project, Care of National Research Council, Washington, D.C., U.S.A.
Coral atolls are flat islands and reefs of lime-
stone of organic origin lying only slightly above
sea level and not in immediate proximity to higher
land. They are in the form of groups of small
islands or islets on reefs that enclose shallow
bodies of sea water called lagoons. Atolls are
very common in tropical seas, especially the
central and western Pacific, central and western
Indian Ocean, and in the Caribbean region.
They are almost lacking in the eastern Pacific, the
Atlantic, and the eastern Indian Ocean.
Because of the relative uniformity of their sub-
stratum, topography, and temperature, their
vegetation is rather simple, though by no means
uniform. Their wide diversity of rainfall and
geographical position contributes to a degree of
variation in vegetation that would perhaps not
otherwise be expected.
The atoll habitat seems to be a very young one,
geologically speaking, though it may well have
existed in something like its present form many
times in the past. The occurrence after the last
glaciation of a warm dry period or "climatic op-
timum" would logically have brought about
a restriction of the glaciers and ice-caps to less than
their present extent and a corresponding rise in
sea level. Geological evidence suggests that this
rise may have been in the neighborhood of either
2 or 3.5 m above present level. At that time,
most of the atolls would have been submerged
and have existed only as reefs awash with little
or no dry land. With the gradual fall of sea level
that started perhaps 3,500 years ago, platforms
of reef limestone were exposed to erosional pro-
cesses and to the deposition of the loose sedi-
ments produced by this erosion. The resulting land
habitats were a mosaic of area of sand and gravel
bars, beaches, dunes, and flats, as well as reef
breccia platforms variously eroded by action of
rainwater. The material of this diverse substra-
tum was limestone, pure except for possible minute
amounts of drifted pumice.
Weathering of this substratum by the solvent
action of rainwater commenced, of course, imme-
diately when it was exposed above the waves. This
process was augmented by the development of
vegetation even in its earliest stages, when very
soon after coral limestone is exposed to air it is
colonized by microscopic blue-green algae. Some
114
forms that grow on solid surfaces have the ability
to bore into the limestone. Others inhabit the
interstices in the surface layers of sand-textured
deposits, causing the particles to cohere and form
a crust that is friable when dry and gelatinous
when moist. The boring types speed the weather-
ing of the limestone by breaking down the outer
layers. Both types are suspected of contributing
fixed nitrogen to the resulting soil, though this
has not yet been demonstrated for the species
involved. Certainly some humus is added as the
plants die and decompose.
The soil that forms in such situations is at best
a poor one, fit only to support plants of the most
absolute pioneer types. With the exception of
calcium, the essential elements for plant nutrition
are either scarce or chemically almost unavailable.
Especially characteristic of these soils are high
pH and deficiencies of nitrogen and iron, as well
as of certain trace elements such as manganese and
zinc. Phosphorus is initially low but may be
increased very early by excrement from sea-and
shore-birds. Nitrogen may be added in the
same way. Extremely high salinity makes these
soils even more unsuitable for most plants, as
does the almost complete absence of organic
matter.
As noted above, the development of microscop-
ic blue-green algal vegetation, indicated by a
darkening of the white or pinkish limestone,
may tend to ameliorate the deficiency of organic
matter and nitrogen. Even so, the number of
plants fitted to colonize such a habitat is at best
very low. This number is further restricted by
the difficulties of dispersal over wide expanses of
sea. The natural dispersal agents are practically
limited to water, wind, and sea-birds.
Almost the only members of the available floras
that are adapted to colonize such essentially
strand habitats are strand species, found every-
where along the shores of islands and continents.
These species are mostly adapted to dispersal
by water, through various floating mechanisms,
and by birds, through sticky or burr-like fruits
that adhere to feathers or feet or through fleshy
fruits that may be eaten and carried for some
distance in birds' digestive tracts. A few species,
such as some grasses, with very small fruits, and
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BOTANY
some plants with winged fruits, may be carried
by winds, especially typhoon or hurricane winds.
Recorded observations on the original compo-
sition of the vascular vegetation on new coral
atoll habitats are few. Pemphis acidula seedlings
are known to become established in tiny pockets
of sand caught on otherwise bare limestone
rock. On coral sand and gravel banks, bars,
or flats Portulaca lutea, Lepturus repens, Boer-
havia spp., Guettarda speciosa, Tournefortia argen-
tea, Scaevola sericea, Suriana maritima, Heliotro-
pium anomalum, Jpomoea pes-caprae, Vigtia man-
na, Pandanus tectorius, Cocos nucifera, Barringto-
nia asiatica, and Thuarea involuta seedlings have
been observed in absolutely unprotected appa-
rently unmodified habitats. Of these, Cocos,
Ipomoea, and Barringtonia have been seen in
such circumstances only very rarely. Certain
other species, though not observed as primary
colonists on new areas, undoubtedly are able
to fulfill this function.
The simplest types of atoll vegetation are es-
sentially aggregations of these pioneer species.
In the driest atolls as well as in a few very remote
ones that few species have reached, succession
seems not to have progressed much beyond this
stage where the original composition still charac-
terizes the vegetation; the actual composition
and local variations may result from chance or
from local differences in substratum, position in
relation to prevailing wind, distance from sea,
or altitude. Characteristic types on such atolls
are Scaevola scrub, scrub forest of Tournefortia,
Guettarda and Scaevola; Lepturus grassland or
Lepturus-Tournefortia savanna; mosaics of thicket
and grassland; or dwarf scrub of Sida fallax or
of Sida with Heliotropium. It is uncertain whether
the Pisonia grandis forest, the Pisonia-Cordia
forests, and the Pandanus forests in some of these
extreme habitats represent pioneer stages or
have developed after some slight alteration of
the habitat and resulting succession. Mention
should be made here of extreme halophytic types,
such as mats of Sesuvium portulacastrum on
lagoon margins and in saline depressions, stands
of Pemphis where the roots are flooded by sea
water at high tide, and scrub of Scaevola or
Suriana bathed continuously by salt spray.
The establishment of vegetation of any kind
tends to bring about change and amelioration of
the habitat in a number of ways. The simplest
of these, of course, is shading. Even a slight
protection from the heat of the sun, from the
drying effect of the wind, and from salt spray may
make possible the establishment of additional
species or create more favorable conditions for
some already present. Addition of humus to the
soil tends to lower its alkalinity and to increase
its water-holding and base exchange capacities.
Phosphorus and nitrogen are increased by accu-
mulation of bird excrement which, however,
tends to be washed down through the porous soil.
Nitrogen may also be contributed by the activity
of bacteria on the roots of Vigna. of Azotohacter,
and probably of blue-green algae. Nostoc is at
times abundant on semi-shaded soil. A further
effect may be the concentration of minor nutrient
elements in humus. Rain tends to reduce salinity
as well as to leach out nutrients, and to build
up a body of fresh ground water at a slight depth
in the soil.
These processes are, of course, continuous and
simultaneous, and usually make possible an
increase in the flora and in the complexity of the
vegetation. In submesic atolls, those with less
than 2.5 m of precipitation, a mixed forest
results which, in the Northern Marshalls, for
example, may have all or any combination of
about 16 species of trees and shrubs: Guettarda
speciosa, Tournefortia argentea, Tenninalia samo-
ensis, Morinda citrifolia, Pandanus tectorius,
Pisonia grandis, Soulamea amara, Cordia subcor-
data, Pemphis acidula, Ochrosia oppositifolia,
Allophylus timorensis, Hernandia sonora, fntsia
bijuga, Pipturus argentea, Scaevola sericea, and
Suriana maritima. These may occur in any
arrangement or combination with chance, dis-
tance from the sea, and wetness of climate the
principal factors involved. Such forests tend to
be more luxuriant with greater rainfall. Substra-
tum may be a factor in the pattern, though the
solution is not clear except in a few cases such as
where a pure or almost pure stand of Pemphis
is found on bare rock. Many of the species listed
may be dependent on shade for the establishment
of their seedlings.
In this mixed forest, ferns appear and mosses
become more prominent. The ferns are principally
terrestrial, and very few species occur. Certain
successional phenomena may be observed, as the
gradual dropping out of Tournefortia from the
more dense forests, the assuming of dominance,
in places, by single species, such as Pisonia,
Ochrosia, Cordia, Intsia, or Pandanus. Forests
made up exclusively of Pisonia arc especially
frequent and occur under a wide range of mois-
ture conditions. Almost no other species are
found established under a thick stand of Pisonia.
The factors involved in these changes are not
usually obvious, and the matter needs much more
115
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
careful study. With fairly abundant moisture,
Ochrosia seems able to succeed any of the other
species, even Pisonia, once it gains a foothold.
Under Ochrosia its own seedlings usually form
a veritable carpet, growing not more than a few
cm high unless a break occurs in the dense cano-
py, when they shoot up rapidly to fill it.
It is of interest that the most stable, or most
nearly "climax" types here seem to be composed
of single species.
In these submesic atolls, also, man has been
able to establish an apparently permanent foot-
hold. He has brought great alteration to much of
the vegetation within his influence. Native
forests have been cleared and replaced by coconut
groves and plantations, and locally by mixed
coconut and breadfruit, or by Calophyllum. In
wet places he has planted taro, or rarely, man-
groves. He has excavated pits down to ground
water for raising taro and other crops. Later
he has abandoned most of these in the submesic
atolls. Increasing wetness of climate is reflected
strikingly in greater luxuriance of coconut plan-
tations and greater abundance of breadfruit trees.
On the wetter atolls, the limitation on the de-
velopment of vegetation seems to be much more
the lack of available flora than the severity of the
environment. The far greater richness of the
vegetation of atolls in the western Carolines and
especially in and near the Malaysian Archipelago,
where a large flora is available, seems to support
this idea.
It is not clear whether the much greater pro-
minence of mangroves in wet atolls is a function
of climate or of their proximity to the rich
mangrove swamps of Malaysia.
Succession to a truly mesophytic vegetation,
even to what might be termed rain forest, is in
these wet atolls unimpeded either by dryness
or by salinity.
Studies of the vegetation of reasonably undis-
turbed atolls of this character are as yet lacking.
Those that have been accessible for investigation
have been profoundly altered by man. The
anthropogenic vegetation is correspondingly
richer there than in drier atolls. In fact it is
difficult at times to tell what species have been
brought by man. The most striking major
aspects are solid, dense forests of giant bread-
fruit trees and extensive taro-pits or marsh gar-
dens in which many food plants in addition to
taro are grown. The weedy vegetation and the
undergrowth in coconut plantations are both
more luxuriant and richer in species.
116
Generally speaking, after the earlier stages of
colonization and development of vegetation have
been passed, the differences in atoll vegetation
in the Pacific may be interpreted in terms of
three major gradients that from dry and seasonal
to wet climates, that from saline to fresh sites,
and that from the east, with a very small available
flora, to the west and southwest with the rich
Malaysian and Melanesian floras.
All three of these gradients, as expressed,
exhibit increasing richness of composition of
vegetation. Toward their extreme ends of
dryness, salinity, and remoteness, the tendency
to complete dominance by single species is marked.
Such types dominated by single species, are rare
in mesic atolls excepting the forests of Pisonia
grandis.
The elTects of decreasing salinity are reflected
in the increased luxuriance and richer composi-
tion of vegetation in the interior of fairly wide
islets, and, very strikingly, by the great increase
in number of species on even slightly elevated
atolls over those lying at sea level. The increase
on the latter type, however, may also be inter-
preted as a result of greater age of the land surface
in raised atolls.
From dry to wet atolls, both luxuriance and
number of species increase in a striking fashion,
whether we are considering only native vegetation
and flora or including that established by man.
The two driest atolls of the northern Marshalls
have vascular floras of nine species each, while
some of the wet southern atolls of the same group
have as many as a hundred or more species.
Accurate figures to show the effect of remote-
ness are not available for atolls at the ends of the
gradient; but it is possible that Clipperton
Atoll had no native plants, and only two are
recorded from Ducie Atoll. Vostok and John-
ston Islands, not so far to the east, but very
remote, had respectively two and three species
of native vascular plants. However, the floras
of other atolls in southeastern and central Poly-
nesia are more diverse than these, but nothing
to compare with the large numbers on various
East Indian Atolls, or even the Caroline ones.
Accurate floristic figures are much to be desired
for atolls in the Malaysian area.
It is thought that in very few atolls has vegeta-
tion developed or succession reached its maxi-
mum possible development under present climatic
conditions, as the soils are very immature and
oversea migration of plants is sporadic and
VOLUME 4 BOTANY
uncertain. Where atolls have been left undis- owing to the disturbances caused by hurricanes
turbed, though, the climatic conditions have and by man's activities, most atoll vegetation
tended to arrest development at different points, may be regarded as being in a more or less active
corresponding to degrees of wetness. However, dynamic status.
117
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
TROPICAL PACIFIC GRASSLANDS AND SAVANNAS
F.R. FOSBERG
Pacific Vegetation Project, Care of National Research Council, Washington, D.C., U.S.A.
Grassland vegetation, a collective term for
plant communities in which grasses and grasslike
plants are the dominant growth form, is fairly
widely distributed in the tropical Pacific Basin
but in aggregate makes up only a small part of
the vegetation. Natural, as compared with man-
made, grassland covers an even smaller area.
In some instances it is difficult to be certain of the
history of a given area.
The classification of grassland may be based
on any of several features composition, physiog-
nomy, structure, habitat, or economic considera-
tions. Because no adequate study has been made
of the region as a whole on any of these bases, and
because the required information is not available
on any of these features for all Pacific grasslands,
it seems logical to base a tentative arrangement
on such data as are available on all features.
Many tropical grasslands, including some Pacific
ones, contain scattered trees and are then gen-
erally called "savanna." Every stage exists
between grassland without trees through savanna
and woodland to forest, depending on the spacing
of the trees. A savanna may be arbitrarily defined
as grassland with trees that are spaced, on the
average, farther apart than twice the diameter
of their crowns, or with trees covering less than
20 per cent of the ground. The significance of
trees in grassland varies, depending on the kinds
of trees. In some cases they may represent
invasion leading to re-establishment of forest;
in others they may belong to species that are
able to resist burning when well grown, but
which fire will kill when young. Still other
savanna trees, usually small ones, belong to
species that are found nowhere else than scat-
tered in grassland. Considerable work has been
done on savannas in other parts of the tropics
but rather little on those of the Pacific. Here they
will be treated mostly as variants of grassland
types and discussed individually in the above
framework.
The Pacific grasslands may be conveniently
separated into those of high altitudes and those
of medium to low altitudes. Fixed altitude limits
may not easily be assigned to these two categories,
but the high-altitude types either are mostly
natural or are moderate expansions of natural
118
grassland, while many, if not most, of the middle
and low altitude types seem to result from or to
have been vastly extended by the activities of man.
Montane grasslands are extensively developed
in the Pacific Basin principally in the Andes
where they occupy the area between timber line
and perpetual snow. Timber line ranges from
about 10,500 to 1 1 ,000 feet in Colombia to nearly
13,000 feet in Bolivia, and there is a correspond-
ing variation in snow line from perhaps 15,000 to
18,000 feet. The Andean montane grassland is
a bunch-grass or tussock formation with Stipa,
Calamagrostis, Poa, Agrostis, and some other
genera as the principal grasses and with abundant
cushion plants and a rich herb and shrub flora.
This grassland belt stretches from Patagonia to
northernmost Colombia and, in poorly developed
form, to Costa Rica. It is continuous in the
south, but is broken into isolated patches in the
lower northern Andes. Still, even in Colombia,
the aggregate area is large (3, 4, 5).
In addition to much local variation, correlated
with altitude, topography, hydrology, exposure,
and human interference, there is a large-scale
regional variation from north to south, related
to climate. This variation expresses itself in three
major grassland regions or formations, designated
conveniently by the local names "pramo,"
"jalca," and "puna." They occur in that order
from north to south and are characterized by
increasing aridity southward.
The paramo is the most remarkable of the three,
marked by features that might well lead to the
argument that it is not a grassland at all. The
most striking feature is the remarkable composite
genus Espeletia. The grotesque gray rosettes of
this group, called "frailejones" by the local people,
are an almost constant feature of the landscape
above timber line from the Venezuelan Andes to
the Paramo del Angel in northern Ecuador, where
they may be seen in great abundance, but where
their occurrence stops abruptly. South of the
Rio de los Charcos they are completely unknown.
Although usually forming only a small part of
the cover, Espelatia gives a unique aspect to the
landscape that amply warrants separation of the
pdramo from the jalca to the south. In addition,
there is a very considerable flora of herbaceous
VOLUME 4
BOTANY
and shrubby plants with striking local endemism
which sets off this wetter northern paramo region.
Giant bromeliads (Puya spp.), Rumex tolimensis,
Lupinus alopecuroides, Aragoa spp., Blechnum
arboreum, and other strange plants add to the
unreality of this landscape.
On level or more gently sloping ground, the
soil is usually black and highly organic. Rainfall
is high, fog is frequent, and mosses, ferns, and
other hygrophytes are in places very abundant.
The arrangement of the mountains somewhat
parallel to the direction of the trade winds lessens
possible rain-shadow effects.
The paramo vegetation thins out upward to
the level of perpetual snow. The upper part,
or super-paramo is more strictly grassland, though
sparse. Downward, the shrubby sub-paramo
vegetation gradually replaces the grassland in a
transition to the "sotobosque" or "elfin forest"
and the mossy forest.
Disturbance by man in the region just below
the paramo has had the interesting effect of caus-
ing at least some elements of the paramo vegeta-
tion to spread downward in places, producing,
what might be called quasi-paramo, very similar
to the paramo in aspect down to elevations as low
as 3,000 m. But many paramo species are lacking,
and there are many introduced weedy species,
including grasses, not normally found in the
paramo. Active human utilization of the pdramo
does not extend much above 3,500 m.
From the Charcos River in northern Ecuador
to north-central Peru, the jalca (often also
called paramo or even "wet puna") occupies a
broadening belt, similarly placed with regard to
the forest belts below and perpetual snow above.
It is a grassland of bunch-grass or tussock species,
with, again, many non-graminoid herbs and
shrubs, but lacking the Espeletia and thus present-
ing a far more conventional mountain meadow
appearance. This is varied, however, by such
oddities as Puya, Azorella, and Distichium.
The jalca, though mostly somewhat drier than
the paramo, is still a mesophytic grassland, and
the ground cover is complete. Fog is still an
important factor in the climate. Human occupa-
tion extends locally to 4,000 m elevation. A
limiting factor seems to be the cold, resulting
from the high degree of cloudiness.
Southward, where the climate becomes signi-
ficantly drier, the grass clumps become more
discrete; Stipa becomes the dominant genus;
and bare ground becomes apparent, at least locally
between the clumps. This is the beginning of the
"puna" or high altitude "steppe."
Here timber line, on the east slopes, is higher.
Woodland of Polylepis reaches 4,200 m. The
associated herbs and shrubs in the grassland are
mostly more xeromorphic. Such genera as Te-
traglochin, Pycnophyllum, Lepidophyllum, and
Opuntia are found. Large woody species of the
cushion plant, Azorella, are common and locally,
along with the shrub Lepidophyllum, provide fuel
for the inhabitants.
Here human activity normally extends to 5,000
m, and a road crosses a pass in Bolivia at over
6,100 m. This increase in the altitude limits of
effective human occupation seems due to the
higher insolation and lower degree of cloudiness.
Enormous areas of puna occupy the broad part
of the high Andes, the "altiplano" of southern
Peru, Bolivia, and Chile. On the east slope it
merges with the "ceja de la montana" or elfin
forest of the Yungas, on the west into the desert
western slopes of the Andes in Peru and Atacama,
Chile. Southward it extends, at lower and lower
altitudes, to the south temperate zone and even,
in modified form, to Patagonia.
Elsewhere in the tropical Pacific the only equi-
valent of Andean high-altitude grassland is a
weakly developed belt in Hawaii. This is charac-
terized by sparse stands of densely tufted species,
up to 1 m tall, mainly Trisetum glomeratum,
Agrostis sandwicensis, and Air a nubigena, at timber
line on the high volcanoes above 2,600 m. The
belt merges downward with the upper forest and
upward into bare lava, cinder, and clinker slopes.
In the saddle between Mauna Loa and Mauna
Kea on Hawaii, similar grassland is dominated
by tufted perennial species of Eragrostis. Exces-
sive grazing by sheep and wild goats has damaged
and altered these grasslands.
Mention may also be made here of extensive
pastures existing at middle altitudes on the north-
ern part of the island of Hawaii. These are
artificial and are made up of numerous species of
forage grasses deliberately introduced for the
raising of beef cattle.
Alpine grassland, in the mountains of Papua,
dominated by Aulacolepis, Poa, and Danthonia,
are mentioned by Brass (2, p. 177), but no des-
cription has been found.
At low and middle altitudes in the Pacific
tropics, there are of course large expanses of such
cultivated grasslands as sugar cane and rice fields.
Various minor grassy communities, such as
marshes, old fields, salt-grass flats and other strand
types, and pastures are of reasonably general
distribution. Worthy of special mention are the
Lepturus repens grassland of coral atolls and other
119
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
strand situations and the reed marshes composed
of Phragmites karka. Stands of Lepturus are
widespread in small patches on dry coral sands
and cover substantial areas on Christmas Island
and other dry atolls of the Central Pacific and on
Pokak Atoll, the driest of the Marshalls, but are
strangely missing in Hawaii.
Lepturus repens, in spite of its name, is a bunch
grass, and occurs in discrete rather small tufts of
fine but hard culms, and narrow but harsh leaves.
Frequently these tufts will send out long wiry
runners which root at the nodes and send up
smaller tufts. Eventually a loose mat may be
formed. Usually one or more of several broad-
leafed herbs and dwarf shrubs occur mixed with
the grass, in varying proportions. These are
Portulaca lutea, Boerhavia sp., Sida fallax, and
Heliotropium anomalum. The last is lacking in
the Marshalls but found in Wake and the Line
islands. Scattered low, rounded trees of Tourne-
fortia argentea may be found, making a savanna
of very characteristic appearance.
Reed marshes are widely distributed on higher
islands, usually, but by no means always,
near sea level just back of the coasts. Fairly
extensive marshes are found on Saipan, Guam,
and Truk islands. They are usually almost pure
stands of the large tropical reed, Phragmites karka.
It forms hollow canes about 1 to 1.5 cm thick,
with broad leaves above, and with panicles
bronze-colored at flowering. Pandanus trees and
Hibiscus tiliaceus may occur scattered through
these marshes to form savannas.
Much more extensive are several upland types
of grassland or savanna. Most widely distributed
of these, though possibly not largest in area, is
that dominated by the genus Miscanthus, prin-
cipally by Miscanthus floridulus, commonly called
"sword-grass" because of the scabrous cutting
edges of its leaves.
In its simplest form, Miscanthus grassland is a
pure stand of sword-grass, crowded large tufts of
reed-like culms with harsh leaves, up to 3 and
even 4 m tall. It is found thus on relatively
fresh ash slopes, as on the volcanoes of the
northern Marianas. Reasonably pure stands are
also widespread in denuded areas on the older
volcanic islands of southeastern Polynesia, es-
pecially Mangareva. Generally, however, there
is a varied accompanying flora of sedges, other
grasses, and several broad-leafed species of herbs
and shrubs. Most of these grow on erosion scars
and other bare spots, which are an almost con-
stant feature of the habitat of Miscanthus. In
Guam, the Miscanthus grassland forms a mosaic
120
with a low, soft grassland composed mainly of
Dimeria chloridiformis. In this mosaic, Dimeria
occupies principally the more level or gently
sloping situations, fine clay soil that dries out
fairly well in dry weather. The steeper, more
rocky sites, as well as those with more permanent
moisture, are covered predominantly by Miscan-
thus.
Fire is a frequent feature of Miscanthus grass-
land and the associated Dimeria. In burned areas,
an aggregation of weeds appears, to be gradually
crowded out by the grasses. If erosion starts,
an erosion scar community of ferns, small shrubs,
and various herbs tends to establish itself on
the bare soil. As time goes on, this community
is replaced by the grasses. Most of the plants
peculiar to these grasslands are members of this
erosion scar community. Members of the genera
Myrtella, Melastoma^ Geniostoma^ Hedyotis, 77-
monius, Dianella, Euphorbia, Phyllanthus, Glochi-
dion, Machaerina, Rhynchospora, Fimbristylis,
Cantharospermum, Chrysopogon, and Ischaemum,
as well as the ferns Gleichenia, Sphenomeris^
Lindsaya, and Cheilanthes, plus Lycopodium,
make up the community. They are mostly either
very wide-ranging species or very restricted
endemics.
The bottoms of permanently or usually wet
ravines in the mosaic are occupied by brakes of
reeds (Phragmites karka) which show up as a
deeper green or, at flowering time, a rich bronze
against the pale green or buff of the Miscanthus.
These reeds are very tall, often filling ravines up to
the level of the grass tops on the terraces along
the ravine margins. Only the difference in color
betrays the existence of the ravine to the casual
observer.
In this mosaic, seedlings of Casuarina equiseti-
folia tend to appear; and an area not burned for
a few years may assume the character of a savan-
na, or even of a sparse forest. Pandanus also form
limited savanna.
Westward in the Pacific, other grasslands tend
to replace Miscanthus as the important types.
In Yap and Palau, savannas of ferns, Nepenthes,
and sedges with some grasses and many secondary
herbs and scattered trees and shrubs of various
kinds occupy the places where Miscanthus would
be expected.
In the Philippines and northward, Miscanthus
floridulus tends to be replaced in certain situations
by Miscanthus sinensis. The exact environmental
relations of these two species are not always
clear, nor is it easy to distinguish them under
some conditions. In the Ryukyus where Miscan-
VOLUME 4
BOTANY
thus may be unusually abundant on limestone,
in contrast to its behavior farther east, the species
involved may often be Miscanthus sinensis though
this has not been determined with certainty. No
information is available on the Miscanthus (?)
grasslands found in western Papua between 1,000
and 2,000 meters of elevation and mentioned
by Brass (2, p. 176).
In places in the Philippines, Caroline Islands,
and probably elsewhere, Saccharwn spontaneum,
a large bunch grass with the aspect of Miscanthus,
may form considerable stands. Not too much is
known of the distribution of this type of grassland.
Since the war, Saccharum spontaneum has ap-
peared in Saipan and spread with great vigor.
It is suspected that the plants involved may really
be seedlings from cultivated sugar cane. True
Saccharum spontaneum^ course, may well have
been introduced accidentally with war supplies,
or perhaps after the war. The species seems na-
tive and not excessively abundant in the high
Caroline Islands from Kusaie to Palau, where it
is frequently mistaken for Miscanthus. The
latter is known with certainty in the Caroline
Islands only from Ponape.
In the Fly River area in New Guinea and west-
ward are large areas of savanna (2), dominated
by Ophiurus exaltatus, a coarse grass reaching 2
m in height, but also characterized by Themcda
triandria, Imperata cylindrica^ and Ischacmum sp.,
and scattered trees of Eucalyptus, Melaleuca,
and Tristania. Tristania dominates or shares low,
wet places with Eriocaulon and many other herbs.
In places, Ophiurus, Imperata, and Sorghum
nitidum form a grassland on well-drained soils.
Elsewhere, Germainia ca pi tat a forms savannas
with Banksia. Low, ill-drained soils have a
ground cover of sedges with numerous other
flowering herbs and small trees of Melaleuca and
Banksia. These savannas fade imperceptibly into
"savanna forests" or woodlands. Brass considers
these savannas to be natural and to be extensions
into New Guinea of the "open forests" of Aus-
tralia.
From Guadalcanal to the Philippines, and
undoubtedly westward in Indonesia in such
islands as Celebes, Timor, and the lesser Sunda
group, occurs a rather coarse grassland domi-
nated by Themeda triandria, the "kunai" grass of
New Guinea. In Guadalcanal, Florida, and
probably Buka in the Solomons, an almost pure
stand of Themeda, with wet places dominated by
Phragmites karka (9, pp. 90-91) occurs on the
coastal plain and runs up onto low mountain
ridges several miles inland, where it changes to
savanna and gradually to forest upward. Theme-
da triandria is a stiffish grass 1 to 2 meters in
height, with drooping flower clusters.
In the Philippines, the Themeda type of grass-
land is the matrix of many of the pine savannas
of Benguet, Luzon. Themeda triandria, Themeda
gigantea, Chrysopogon aciculatum, species of
Ischaemum, Panicum, Andropogon, and Fimbris-
tylis, as well as various other grasses, sedges, and
broad-leafed plants make up this grass cover.
Merrill (8) mentions large areas of Themeda-
dominated grassland throughout the Archipelago
and lists a large number of species as common in
open grasslands. There are obviously many
variants of this type as well as of the Imperata
grassland to be described next.
Although Themcda and its associates are main-
tained by burning, if the burning is carried to an
extreme, this type of grassland may give way to
Imperata cylindrica and associated species.
Imperata grassland covers vast acreages in the
New Guinea-Indonesia-Southeast Asia region,
and is usually, if not always, associated with man.
Its altitudinal range is enormous from sea level
to as much as 2,500 m (in New Guinea, according
to 6). Imperata is the "lalang" of Malaya, the
"alang-alang" of Indo China, the "cogon" of the
Philippines. It may be thought of as the true
"climax" of the man -shifting-cultivation-burning
complex in this part of the world. Imperata, with
the tough matted wiry buried rhizomes, is about
as near as nature comes to producing a really
fire-proof plant. When the shoots are burned
off, the rhizomes remain uninjured and immediate-
ly respond by sending up flowering culms, fol-
lowed by a new crop of leafy shoots.
Though Imperata is a harsh, unpleasant grass
in texture, it is not large. It forms a tough hard
sod, is not bunchy, and seldom exceeds 1 meter
in height. Its bright green color gives a deceptive
appearance of succulence, but animals do not
care much for it. It is commonly associated with
a rather characteristic lot of widespread plants
belonging to such genera as Melastoma, Pteridium,
Lygodium, Eurya, Gleichenia, Rhynchospora,
Fimbristylis, and Andropogon.
Although this type of grassland is easily invaded
by woody species, such as pine, such invasion
seldom leads to anything, as the invaders are
generally susceptible to fire. And fire, the factor
that brings about dominance by Imperata, is
seldom long absent where Imperata has achieved
such dominance.
Much study has been devoted to the origins of
the grasslands and savannas of tropical America
121
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
and Africa. A paper by Beard ( 1 ) provides a
convincing set of generalizations about at least
the tropical American savannas. The grasslands
of the western tropical Pacific have received little
attention, as it has generally been assumed that
they arc of secondary origin, due to man's activ-
ities. At least one student (9) has vigorously
contested this view as applied to the grassy areas
of Guadalcanal, withal not too convincingly.
His conclusions are largely based on low rainfall,
as shown by a very short period of record; on
absence of evidence of burning at the present
time; on the existence of strips of forest along
streams; on the existence of two peculiar animals,
an endemic subspecies of button quail an obli-
gate grassland bird and a fish adapted to inter-
mittent streams; and especially on a curious
argument that, because the climate cannot support
rain forest the grassland must be "climatic." No
mention is made of the possibility of other types
of tropical forest. From the data he presents,
there is no doubt whatever that these grasslands
depend for their existence on the rain shadow
cast by the high Guadalcanal mountains. Whether
they are climatic in origin is another matter.
They may very well be so, but it is not established
to complete satisfaction in his paper that man in
earlier times did not have a hand in their origin
or spread. The fact that he describes them as an
almost pure stand of Themeda triandria and men-
tions almost no accompanying flora makes their
natural origin at least open to question.
Similar views concerning the Miscanthus grass-
lands of Fiji and Guam have been expressed oral-
ly by a number of casual observers. In the case of
Fiji, these views were based on the low rainfall,
and in that of Guam on edaphic considerations.
Some study has been devoted to the origin of
the Guam grasslands. Here, on historical evi-
dence, there was much more forest even within
historical times. The introduction of deer and
of the custom of driving them by fire caused a
great reduction of this forest during Spanish times.
Many of the plants characteristic of the present
grassland were clearly introduced from America
and elsewhere during the Spanish period. These
facts favor the idea that this grassland is of
secondary nature. However, as in tropical
America, there are some native, even endemic
species, restricted to the grasslands. Such are
Phyllanthus saffordii, Ischaemum longisetum,
Machaerina aromatica, Hedyotis grandiflora, Wik-
stroemia elliptica, and Spaihoglottis micronesiaca.
Glochidion marianum is found on both Guam and
Ponape; Myrtella benningseniana, Timonius albus,
122
Hedyotis fruticulosa on Guam and Yap; and
Dimeria chloridiformis on Guam and Palau.
In addition there are certain widespread plants
of similar habitats that are not obviously intro-
duced, such as Miscanthus floridulus and Dianella
ensifolia (both known as fossils from Pagan,
northern Marianas), Curculigo orchioides, Fim-
bristylis tristachya, Fimbristylis annua, Melasto-
ma malabathricum, Lindsaya ensifolia, Cheilanthes
tenuifolia, and Lygodium seandens.
This assemblage of plants could not likely
have developed or lived in forest, as it is made up
of shade-intolerant species which do not now
inhabit nearby forest. The obvious possible
habitats were considered. Fresh ash slopes,
such as those covered by Miscanthus in the
northern Marianas, were ruled out, as such have
apparently never existed in Guam. Open ridge
crests do not seem likely, as in a climate such as
that of Guam and at such low altitudes the ridges
are likely to have been wooded. Also many of
the species concerned do not elsewhere inhabit
such ridge crests. Landslide and erosion scars
would be a possibility for some of the species, if
there ever had been an abundance of landslides,
but this does not seem a sufficient possibility, in
the absence of humans and grazing animals to
cause large numbers of landslides and aggravated
erosion. The late Josaiah Bridge suggested a
correlation with bauxite deposits. This would
be a possibility, as many of the same plants are
associated with bauxite elsewhere. However,
extensive deposits of bauxite are lacking in the
areas concerned. In any event, even with bauxite,
open habitats would have been required.
In attacking the same problem in the tropical
American savannas, Beard (I) noted a constant
association of what he regarded as natural savan-
na with areas of mature topography flat or gently
undulating country, on plains, terraces or plat-
eaus, having soil with impeded drainage, generally
a permeable layer lying on an impermeable one.
It seemed pertinent to look for such an asso-
ciation of savanna and topography on Guam,
past or present. When asked about this, J.I.
Tracey suggested that certain flat-topped erosion
remnants scattered over southern Guam might
represent the remains of an ancient flat erosion
surface. These "mesitas" present precisely the
flat surface and permeable surface layer overlying
a clay subsoil that are required by Beard's theory.
The general geological reconstruction of southern
Guam indicates that this flat surface may have
been in existence for a long period, possibly since
some time in the Miocene. If this is so, a proper
VOLUME 4
habitat would have been available for the develop-
ment of the native part of the floristic assemblage
now characteristic of the Guam grasslands.
It is suggested that the history of these grass-
lands is about as follows :
1. The gradual appearance of grasslands and
their flora, and their encroachment on the forest
as the topography matured during the erosion
cycle following the most recent emergence of
most of the area from the middle Miocene Alifan
submergence.
2. The gradual restriction of these grasslands
and encroachment of forest as the flat surface was
dissected during elevation and rejuvenation.
3. The persistence of grassland only in small
patches, mostly on the "mesitas" or erosion rem-
nants during the time of the Chamorro inhabi-
tants of Guam.
4. The rapid and extensive expansion of the
grasslands onto the denuded eroded hills, with
addition of numerous introduced species from
the time of the Spanish occupation to the present
day.
Similarity in situation on other Micronesian
islands, such as Palau and Yap, suggests that
possibilities of a similar history might be inves-
tigated when the necessary geological information
is available. Until floristic, geologic, and soils
data can be assembled for other Pacific grasslands,
little more can be said about their origin beyond
the statement that in most regions they have at
least been greatly extended, if not actually brought
into being, by human agencies.
Merrill (7, 8) categorically considers the
Philippine grasslands to be the result of human
activity. The fact that some of the plants found
in these grasslands are endemic to the Philippines
suggests that this may be an extreme interpretation
and needs more study. However, it is easy
enough to see that the pine savannas of Luzon are
increasing their area very conspicuously at the
present time. Certainly it is safe to assume that at
least most of the Philippine grasslands are secon-
dary.
Descriptions of the New Guinea savannas (2)
suggest that at least some of them may be natural,
with the same characteristics of alternate extreme
desiccation and waterlogging described by Beard
for tropical America. These areas would amply
repay further study.
REFERENCES
(1) Beard, J., 1953, Ecol. Monogr. 23: 149-215.
(2) Brass, L., 1938, Jour. Arnold Arb., 19:
174-190.
(3) Cuatrecasas, J., 1934, Trab. Mus. Nac.
Cicnc. Nat. Ser. Bot., 27: 1-144,
(4) ..... ___ ., 1956, Suelos Ecuatoriales 1:
13-30.
(5) Fosberg, F.R., 1944, Jour. N. Y. Bot. Card.
45: 226-234.
(6) Lam, H.J., 1945, Sargentia 5: 153, 168.
(7) Merrill, E.D., 1912, Phil. Jour. Sci. Bot. 7:
145-208.
(8) __ _ , 1923-1926, An enumeration of
Philippine flowering Plants. 4 vols.,
Manila.
(9) Pendleton, R.E., 1949, Ecol. Monogr. 19:
75-93.
123
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS __
SOME UNSOLVED PROBLEMS IN TROPICAl7 FOREST ECOLOGY^
TATUO KIRA, HUSATO OGAWA, and KYOJI YODA
Laboratory of Plant Ecology, Institute of Polytechnics, Osaka City University, Osaka, Japan.
SITUATION OF TROPICAL FORESTS
IN THE WORLD VEGETATION
In the ordinary sequence of climatic zones on
the earth, the moist climates of temperate and
tropical regions are separated from each other by
the intervening dry climate which prevails over
middle latitudes. So that the two major forest
areas on the continents, temperate and tropical,
rarely come into contact in their geographical
distribution. This geographical isolation has made
plant ccologists overestimate the contrast between
temperate and tropical forest vegetation in phy-
siognomy, community structure and floristic
diversity. Indeed the luxuriant growth of huge
trees as well as the immense richness of plant
species in tropical forests is a wonder to those who
are acquainted with temperate vegetation, but
nevertheless it is not right to lay too much stress
upon this first impression.
Along the western fringe of the Pacific Basin,
a long chain of islands stretches from north to
south along the continent, starting from Sakhalin
through Japanese Archipelago, the Ryukyus and
Formosa to the Philippines, where both summer
and winter monsoon bring abundant rainfall
throughout the year. It should be emphasized
that this part of the western Pacific is perhaps the
sole example in the world that represents the un-
interrupted series of moist forest climates from
the subarctic to the tropical zone. According to
the researches of Japanese ecologists (1,2,3), no
less than six forest zones are discriminated along
the western coast of the Pacific. The parallel
nomenclature of climatic and vegetation zones
are as follows.
CLIMATIC ZONE
0. Polar zone
1. Arctic zone
2. Subarctic zone
APPROXIMATE RANGE VEGETATION ZONE DOMINANT OR CHARAC-
OF WARMTH INDEX* TERISTIC TREES
month-degrees Barrens, snow and ice
0-15 Tundra
15-45 A 55 Evergreen conifer forest Abies, Picea, Larix, Betu-
la, etc.
Deciduous broad-leaf Fagus, Quercus, Acer, etc.
forest
Lucidophyllous or Cyclobalanopsis, Shiia,
laurel forest Machllus, etc.
,, Castanopsis, Lithocarpus,
Cinnamomum, etc.
Subtropical rain forest Ficus, Machilus, Lager-
stroemia, and many other
genera
240 - Tropical rain forest Floristic composition
extremely diverse. Genera
of Dipterocarpaceae are
most remarkable
* Warmth index is a convenient geographical index of thermal climate proposed by Kira (1). It is given by summing
the mean monthly temperature (t) above 5C or by I (/ - 5). Winter months in which / < 5C are excluded.
124
3. Cool temperate zone 45 A 55 - 85
4. Temperate zone 85 - 140
41. Warm temperate zone 140-180
5. Subtropical zone 180-240
6. Tropical zone
VOLUME 4
BOTANY
These systems also proved applicable to the
altitudinal zones on high mountains of Formosa,
continental China, Himalaya (4,5), and Malay-
sia (2), with slight modifications. The zones
presented here seem more in number than those
in any other systems hitherto proposed. This is
not strange because the differentiation of vege-
tation zones along thermal gradient is expected
to be most complete under moist climates. Where
the subhumid or arid climate interferes with the
normal sequence, the thermal zones may reduce
in number or even be discontinued as is usually
the case in other parts of the world, on which
most other systems of zonal classification have
been based.
These zones are more or less physiognomical.
They do not represent the discrete units of vege-
tation, but are no more than the vague patterns
joined together by gradual transition. As stressed
by Imanishi since 1937 (6,7,8) and recently by
Curtis (9 JO) and Whittaker (11J2), one zone
gradually and continuously gives way to another,
when the vegetation is analysed in terms of the
distribution of each component species along an
environmental gradient, e.g., thermal, moisture,
etc. Fig. 1 illustrates an example of the distri-
bution of some twenty conifer species in central
Japan along the thermal gradient, including both
horizontal and altitudinal. As seen in the figure,
all distribution curves are bell-shaped and neither
the ends nor the modes of curves tend to con-
centrate into definite groups, although the area
Warmth index (month -degrees)
Fig. 1. Distribution of 24 conifer species along thermal gradient in the Central District, Japan. 1 :Pinus pumila.
2 : Pinus koraiensis. 3 : Abies Mariesii. 4 : Abies Veitchii. 5 : Picea jezoensis var. hondoensis. 6 : Larix Kaempferi.
1 : Tsuga diver sifolia. 8 : Picea bicolor. 9: Tax us cuspidal a. 10 : Abies homolepis. 11 : Pinus pentaphylla. 12 : Thuja
Standishii. 13 : Chamaecyparis pisifera. 14 : Picea polita. 15 : Chamaecyparis obtusa. 16 : Thujopsis dolabrata. 17 :
Tsuga Sieboldii. 18 : Sciadopitys verticillata. 19 : Abies firma. 20 : Cryptomeria japonica. 21 : Pinus Himekomatsu.
22 : Pinus densiflora. 23 : Torreya nucifera. 24 : Pinus Thunbergii.
125
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
surveyed includes four zones from arctic (alpine)
to temperate.
In the similar way, as we travel southwards
from the subarctic conifer forest of Sakhalin and
Hokkaido, the nature of forest community gradu-
ally but steadily changes towards the tropical
rain forest of the Philippines. No discontinuous,
sudden change is found except where the flora is
impoverished due to the small size of the islet as
on the Tokara Group at the southern end of
Japan, or where the thermal gradient is extra-
ordinary sharp as in the narrow straight between
Formosa and Botel Tobago. Thus we are inclined
to consider that the aforementioned difference
between temperate and tropical forest is nothing
but the contrast between both ends of a con-
tinuous series.
As an instance, the floristic diversity of the
tropical rain forest, so often spoken of in ecolog-
ical literatures, will be considered from this view-
point. In recent years, plant and animal ecolog-
ists have recognized that the number of species
and the number of individuals per species in a
community are subject to certain statistical regu-
larities. Let us now adopt Fisher's law of
logarithmic series ( 13) . Denoting by s the number
of species which is represented by n individuals
in the sample randomly collected from a certain
community, Fisher's law is given by
s = a
x n
where x (0 < x < 1) is the value that represents
the completeness of sample and the more com-
plete is the sample the nearer it approaches unity.
a, on the other hand, is independent to sample
size and means the number of species represented
by a single individual when the community is
completely sampled. The latter value or the
index of diversity offers an exact measure of esti-
mating the floristic diversity of a community.
As shown in Figs. 2 and 3, the law is fairly well
satisfied by forest samples of utterly different
nature and from different climatic zones. Es-
pecially noteworthy is the fact that a single domi-
nant forest in the temperate zone (Fig. 2) and a
tropical rain forest that contains nearly 100 tree
species per hectare (Fig. 3) equally well fit the
law. Apparent correlation is found when the
values of a obtained from different climatic zones
are plotted against thermal gradient (Fig. 4). These
evidences suggest the continuous decline of
floristic diversity from tropics to high latitudes
and do not support the view that the tropical forest
is essentially different in its character from the
temperate forest.
126
In his monograph on the tropical rain forest,
P.W. Richards (4) has recently stated that the
tropical rain forest is not a highly specialized plant
community as usually believed, but represents
the most original type from which more impov-
erished vegetation of temperate regions have
been derived through evolutional history. With
regard to floristic diversity, he found that the
diversity was most pronounced in the mixed forest
on mesic sites, and that, even under the same
rain forest climate, forests on perhumid or sub-
humid sites tended to be poorer in tree species
and to have a few more or less apparent domi-
nants. A similar principle might also be applied
to the changes due to the fall in temperature.
PRODUCTIVITY OF
TROPICAL FOREST ECOSYSTEM
In view of the continuous concept of the world's
vegetation as stated above, the study of tropical
forests, especially of the mixed rain forests, is
doubtlessly one of the most important subjects
for plant ecologists, as it may justly be called the
starting point of all kinds of vegetation study.
Owing to the unfavorable climate and inaccessi-
bility from civillized areas, however, researches
on tropical forest ecology in the past were mostly
confined to community floristics and morphology.
Among the problems left for future study, we are
especially interested in the productivity of forest
ecosystems under rain forest climate.
As early as in 1930, Vageler (15) estimated the
annual organic matter production by tropical
rain forest to be 100-200 tons per hectare. There
is little doubt about the widespread conception
that tropical rain forest is by far the most pro-
ductive of all terrestrial ecosystems. Vast accumu-
lations or standing crop of living plant body on
unit forest area is mainly responsible for the con-
ception. But the amount of standing crop is not
always proportional to the true productivity or
the velocity of organic matter production, usually
given by the amount produced per year and per
unit area. As a considerable portion of annual
production is accumulated as wood year after
year, and leaves as well as small branches and
roots are continuously shed off day by day, the
productivity of a forest can not be estimated from
the standing crop alone.
The difficulty in determining the true productiv-
ity caused considerable variation in the estimates
by different authors even for temperate forests.
Filzer (16), for instance, estimated the dry matter
production of German forests to be ca. 4.6 t/ha/
year. This figure was largely common, he stated,
to pine, spruce, and beech forest, and also to the
productivity of pastures. This seems, however,
somewhat underestimated. From the growth
analysis of a 12 year old stock of ash in Denmark,
Boysen Jensen (17) estimated the annual produc-
tion of dry matter per hectare as follows:
1. Wood 4.09 tons
2. Leaf 2.72
3. Branches shed off 0.57
4. Respiration loss 3.10
5. Total 10.48
Underground parts were excluded from the
estimation. Assuming the Top/Root ratio to be
5.0, annual net production (1+2 + 3) and gross
production (5) may amount to 8.9 and 12.6 t/ha
respectively. Miller's comprehensive work (18)
clarified the age-productivity relationship in
Danish beech forest. The net productivity reaches
VOLUME 4 BOTANY
its maximum (ca. 16 t/ha/year) at 30-40 years,
whereas the maximum gross production (ca. 28
t/ha/year) is found at 60-90 years when increased
ratio of respiration loss results in the decrease of
net production by 2-4 t/ha.
In 1956, Satoo (19) published the result of his
growth analysis on a regenerated stand of aspen
in central Hokkaido about 40 years in age, and
estimated the net productivity of aerial organs to
be 8.7 t/ha/year. With the assumption that
ca. 30% of gross production is lost by respiration
and T/R ratio is equal to 5.0, the estimated annual
gross production becomes 14.9 t/ha. If the under-
growth of bamboo grass is included in the cal-
culation, it would rise to as much as 20 t/ha.
Both Japanese and Danish estimates are on
the same order of magnitude. Because Denmark
and Hokkaido belong to the cool temperate zone
20
No. of individuals per species
30
Fig. "Z. Species number-individual number relation in a lucidophyllous forest of central Japan, dominated by Shiia
cuspidata. The curve indicates Fisher's logarithmic series fitted to the data. Sampled area 0.16 ha. Data after Miki, 1932.
127
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
S
50-1
40
o
30
20-
10-
n
"so"
"ioo iso
No. of individuals per species
Fig. 3. Species number-individual number relation in a tropical rain forest, Para, Brazil. Sampled area 3.5 ha. Data
after Pires et ai, 1953.
128
VOLUME 4
BOTANY
(warmth index approximately 50 and 65 month-
degrees respectively), we may tentatively conclude
that cool temperate forests produce 15-25 tons
of dry organic matter per hectare and per year
under moderate edaphic conditions.
No exact estimates arc now available for
warmer climates, but the production of some
temperate grasslands may be worth mentioning.
Although the standing crop of living plant of
forest ecosystem greatly exceeds that of grassland,
there is little evidence that the productivity of the
latter is much lower than that of the former when
they thrive under the same climate. In hilly
pastures of Japan belonging to the temperate
zone, where the natural grasslands, of Miscanihus
sinensis are maintained for thatching materials,
60 r
40
20
a standing crop of summer foliage of more than
60 tons per hectare (fresh weight) is common.
The approximate equivalent of this in total dry
matter including root is 27 tons. As the annual
increment of living rhizomes seems negligibly
small in a well-developed grassland, this amount
of summer crop may roughly equal the net pro-
duction of the year. Thus the gross productivity
would amount to 39 t/ha/year.
In this connection, the relation of agricultural
productivity to thermal gradient is very suggestive.
Kawakita (20) compared the agricultural pro-
ductivity of eight districts of the Japanese Archi-
pelago in terms of the average yields in calories
of edible parts of main food crops. When
the productivity is correlated with the average
O O
/
/
/
/
/
/
O
100 200
Warmth index (month -degrees)
Fig. 4. Fisher's index of diversity (a) increases with temperature.
300
129
PROCEFDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
10
20C
20xl0 3
s
X
15
-o
o
Ui
cx
"03
10
Mean annual temp.
Warmth index
100
200 month -deg.
Fig. 5. The relation of agricultural productivity to thermal climate. 1: Sakhalin. 2: Hokkaido. 3: Northeastern
Hondo. 4: Central and southwestern Hondo. 5: Sikoku. 6: Kyusyu. 7: Ryukyus. 8: Formosa. Data after Kawakita
1949.
130
thermal data, approximately linear regression is
obtained (Fig. 5). It is to be noted that the
warmth index appears more linearly correlated
to the productivity than the mean annual tem-
perature. If similar linear regression be expected
between the warmth index and the productivity
of natural ecosystems and the gradient of the
regression line remains as it is, we could estimate
the productivity of tropical forests by extrapola-
tion. According to the regression of Fig. 5, the
productivity at 260 month-degrees of warmth
index which is the standard value for humid
equatorial climate is about 5.26 times as great as
that at 55 month-degrees. Assuming the gross
productivity of a forest at 55 month-degrees to be
15-25 t/ha/year, the expected value in the tropics
would be 79-132 t/ha/year, falling on the same
order with that of Vageler's estimation. It is
also of interest to note that the gross productivity
of marine algae in a coral reef community of
tropical Pacific water was 24g/cm 2 /day in glucose
(21). Equivalent dry matter of this as ordinary
plant materials is about 80 t/ha/year.
A most reasonable representation of productiv-
ity is found in the photosynthetic energy effi-
ciency or the ratio of radient energy utilized for
the organic matter synthesis to the total radient
energy available. The efficiency or Lindeman's
ratio has been studied mostly with aquatic
ecosystems, and it is now widely recognized that
eutrophic lakes and ocean waters in the temperate
zone having phytoplankton as principle producer
have the annual efficiency of 0.3-0.4%. Higher
efficiencies were recently observed by Odum (21,
22) in the tropical coral reef and the water-grass
community of a freshwater thermostatic spring in
Florida, the ratio to the total available radiation
at the water surface being 2.9% and 2.6% res-
pectively. One author (23) has stated that the
energy efficiency in natural ecosystems hardly
exceeds 1 % and that forests are in general even
lower than aquatic ecosystems in their efficiency.
However, our calculations on the aspen forest
of Hokkaido mentioned above resulted in a con-
siderably higher percentage. Annual gross pro-
duction of 20 t/ha in this forest corresponds to
the annual efficiency of 0.8%; and, if only the
radiation during 5.5 months' growing period is
considered, the ratio increases to 1.4%. As for
the more productive beech forest of Denmark,
where the available radiation becomes less owing
to higher latitude (56N.), a still higher percentage
is to be expected. For Miscanthus grassland of
central Japan, the estimated efficiency is 1.9%.
We might therefore conclude from these evidences
that the forest ecosystem could reach a much
VOLUME 4 BOTANY
higher level of energy efficiency than formerly
believed, presumably as high as 2.5%.
Average annual amount of solar radiation on
the equator is 0.339 cal/cm 2 /min according to
Simpson (24). This is equivalent to 1782 x 10 10
cal/ha/year. Similar estimates by Fukui (25)
give almost the same value. If the tropical forest
is assumed to produce photosynthetic assimilates
with the efficiency of 2.5%, 44.5 x 10 l calories
are expected to be utilized for organic matter
synthesis, or 117 tons of glucose are produced
per hectare and per year. One hundred and seven-
teen tons of glucose roughly correspond to 106
tons of dry plant materials. This estimate again
agrees well with the result obtained by the ex-
trapolation of the productivity-temperature re-
gression in Fig. 5.
FALL IN PRODUCTIVITY
CAUSED BY FOREST DESTRUCTION
One of the greatest difficulties confronting
tropical agriculture is the catastrophic decline of
productivity that occurs when the forests have
been cleared and converted into such artificial
ecosystems as farmland or plantation. This is
especially the case with humid rain forest areas.
In general the agricultural exploitation of these
areas is as yet far from success. Only the shifting
agriculture on upland slopes and the paddy rice
cultivation on irrigated plains, both of which
have presumably originated among the inhabi-
tants of humid tropics, are well adapted to this
climate.
In temperate regions, the agricultural pro-
duction is no less efficient than the production of
natural ecosystems. Transeau (26) estimated, for
example, the energy efficiency of gross organic
matter production for a heavy crop of maize in
Ohio to be 1.6%. Similar efficiency for average
rice crop in Japan (ca. 4.6 tons of unhulled grains
per hectare in air-dry weight) is about 1.1%. The
highest yield of rice hitherto recorded from Japan
amounts to 15.3 t/ha, and the gross productivity
equivalent to this grain yield means the energy
efficiency of about 3.3 %. Such high productivity
of temperate agriculture is of course maintained
by the addition of chemical and organic ferti-
lizers, which not only compensates the amount
of minerals removed from the soil as crops and
through leaching, but also enriches the soil beyond
the level of mineral content under natural condi-
tions. Natural ecosystems, especially the forest
with its stratified structure, seem more efficient
in utilizing the available space as well as the
131
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
radiant energy that falls upon it, as compared
with singled-layered, widely spaced crop com-
munities. It is perhaps the effect of enriched soil
minerals that enables agricultural ecosystems to
overcome this handicap.
In the tropics, however, the return of minerals
to the soil is so large in amount that to replace it
with manuring under cultivated conditions is
a hard task. On the other hand, high temperature
accelerates the decomposition of litter to such an
extent and the leaching of soil minerals is so rapid
owing to heavy rainfall that the soil of cleared
forest land is quickly devastated. We have seen
in Fig. 5 that the agricultural productivity is
nearly proportional to the warmth index through-
out the long stretch of Japanese Archipelago,
from Sakhalin to the Ryukyus, where the way of
agricultural land utilization is essentially the same.
Turning to Formosa, however, the productivity
suddenly falls below the expected value. Al-
though the change in the traditional system of
land utilization may be responsible for the fact
to a certain extent, this appears to suggest an
unavoidable limitation imposed upon the tem-
perate way of agriculture under tropical climates.
Extremely low mineral content of tropical
forest soils must be mentioned here. In regions
of nearly equal climatic humidity, the organic
matter or the nitrogen content in the soil can be
represented as the function of temperature,
decreasing exponentially with rising temperature
(27). In contrast to temperate forest soils con-
taining an average of 0.1-0.3% total nitrogen,
the percentage for tropical forest soils hardly
exceeds 0.05%. Similar relations may well be
expected for other mineral nutrients. The layer
of undercomposed organic materials is also
poorly developed on the floor of tropical rain
forest. But the minor standing crop of minerals
stored in the soil does not always indicate the
lower productivity of a whole ecosystem, as the
standing crop is nothing but the balance in the
budget of minerals which at every moment are
set free from the litter and absorbed by the roots
again. Several authors have reached the same
conclusion (14) that there is an almost closed
cycle of plant nutrients in the tropical rain forest
ecosystem and that the minerals are at once
reabsorbed by plants as soon as they are released
from the decomposing plant materials.
The dynamics of soil mineral budget is quite
different in temperate forests. To cite an example
from the study of Ovington (28), average dis-
tribution of nitrogen in several pure stands of
different tree species, artificially established at
132
Abbotswood, England, were as follows:
In canopy 274 kg/ha
Tree boles 132
Ground flora 32
Forest floor 235
Mineral soil 7,041
(Underground parts are omitted)
Considering the age of stands (mostly 40-45
years), the amount of nitrogen in the annual
increment of living plant body as canopy, boles,
and ground vegetation may be of the order of
200-250 kg/ha. When compared with the amount
present in soil and litter (235 + 7,041 - 7,276
kg/ha), the turnover of soil nitrogen would be
only 1/30-1/35 time/year, if the addition due to
rainfall and nitrogen-fixers be compensated by
the loss through leaching.
As to tropical forests, judging from the pro-
ductivity and the average nitrogen content of soil,
the annual turnover of soil nitrogen might be not
far from 1. Namely, the speed of circulation of
plant nutrients is 30-35 times more rapid in trop-
ical than in temperate forest. The speed of
circulation is most probably the function of
temperature, and to establish the turnover-tem-
perature relation as known functions may provide
a possible approach to understand the dynamics
of the tropical rain forest ecosystem.
What we have said above is little more than
common ecological knowledge. Our intention
is to point out the importance of the study of
ecosystem metabolism in the rain forest and its
implications to tropical agriculture. Intensive
study in forest areas where shifting cultivation is
prevalent may throw new light on the problem.
SUMMARY AND CONCLUSION
1. It is pointed out that the tropical rain forest
occupies a terminal situation on a long continuous
series of forest vegetation from subarctic to tro-
pical climate. Along the scries or the latitudinal
thermal gradient on the earth, the floristic diversi-
ty, productivity, and other characters of forest
ecosystem continuously changes, ending in the
tropical rain forest with its extreme richness in
flora and organic matter production. The im-
portance of the western Pacific area is stressed
which represents the sole example in the world of
uninterrupted continuation of humid forest
climate from high latitudes to the equator.
2. By the extrapolation of a temperature-
productivity curve obtained from temperate
regions, the annual gross productivity of tropical
rain forest is estimated to be 105 25 t/ha in dry
matter. This agrees well with another estimate
based on the energy efficiency of organic matter
production by forest combined with the amount
of total radiant energy available in the tropics.
3. The cause of the failure of rain forest area
as arable land is discussed as related to the eco-
system metabolism. It is suggested that the very
rapid circulation of plant nutrients between plant
and soil is mainly responsible for the sudden
decline of productivity that ensue from the clear-
ance of rain forest. A rough estimation has
shown that the turnover of soil minerals may be
some 30 times greater in tropical forest than in
temperate forest.
4. These results are only tentative, but they
may serve as the starting point for intensive field
researches on tropical rain forest which are
earnestly desired in the near future.
REFERENCES
(/)* Kira, T., 1945, A new system of climatic
classification in eastern Asia as the basis
for agricultural geography, Kyoto.
(2)* Kira, T., 1945, New classification of cli-
mates in southeastern Asia and western
Pacific, Kyoto.
(3)* Suzuki, T., 1952, The East-Asiatic forest
climaxes, Tokyo.
(4) * Imanishi, K., 1953, Nature of Nepal Hima-
laya. Kagaku 23:406-416, 464-468.
(5) Kawakita, J., 1956, Vegetation. In 'Land
and crops of Nepal Himalaya', Scientific
results of the Japanese expeditions to
Nepal Himalaya, 1952-1953, edited by
H. Kihara, Tokyo, pp. 1-65.
(6)* Imanishi, K., 1937, Community analysis
and community classification. Geogr.
Rev. Jap., 13:725-736.
(7)* Imanishi, K., 1937, The altitudinal regions
of the Northern Japanese, Alps. Jour.
Jap. A/pine Club, 31:269-364.
(8)* Imanishi, K., 1949, Principles of bio-
sociology, Osaka.
(9) Curtis, J.T. and Mclntosh R.P., 1950, An
upland forest continuum in the prairie-
forest border region of Wisconsin,
Ecology, 32:476-496.
(10) Brown, R.T. and Curtis, J.T., 1952, The
upland conifer-hardwood forests of
* In Japanese.
** In Japanese with English summary.
BOTANY
northern Wisconsin, EcoL Monogr.,
22:217-234.
(11) Whittaker, R.H., 1952, A study of summer
foliage insect communities in the Great
Smoky Mountains, Ecol. Monogr., 22:
1-44.
(12) Whittaker, R.H., 1956, Vegetation of the
Great Smoky Mountains. EcoL Monogr.,
26: 1-80.
(13)
Fisher, R.A., Corbet, A.S. and Williams,
C.B., 1943, The relation between the
number of species and the number of
individuals in a random sample of an
animal population, Jour. Animal
12:42-58.
(14)
(15)
(16)
(17)
(18)
Richards, P.W., 1952, The tropical rain
forest, Cambridge.
Vegeler, P., 1930, Grundrissdertropischen
und subtropischen Bodenkunde, Berlin.
Filzer, P., 1951, Die naturlichen Grund-
lagen des Pflanzenertrages in Mittel-
curopa, Stuttgart.
Boyscn Jensen, P., 1932, Die Stoflproduk-
tion der Pflanzen, Jena.
Boysen Jensen, P., 1949, Causal plant geo-
graphy, Del Kgl. Danske Videnskah.
Selskab, Biol. MeJd., 21 (3): 1-19.
(79j**Satoo, T., Kunugi R. and Kumekawa, A.,
1956, Amount of leaves and production
of wood in an aspen (Popitttts Davidiana)
second growth in Hokkaido, Bull. Tokyo
Univ. Forests, 52: 33-51.
(20)* Kawakita, J., 1949, A quantitative repre-
sentation of land productivity in terms
of calory yield. Geography for Social
Life, 19: 6-10.
(21) Odum, H.T. and Odum, E.P., 1955, Troph-
ic structure and productivity of a
windward coral reef community on
Eniwetok Atoll, EcoL Monogr., 25:
291-320.
(22) Odum, H.T., 1957, Trophic structure and
productivity of Silver Springs, Florida.
EcoL Monogr., 27:55-112.
(23) Park, O., 1949, Community metabolism.
In Allee, W.C. ct al. : Principles of
animal ecology, New York. pp. 495-528.
(24)* Fukui, E., 1941, Climatology. Revised
Edition, Tokyo.
(25)** Fukui, E., 1953, Climatology of radiation;
133
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the meridional distribution of insolation (27) Jenny, H., 1941, Factors of soil formation,
over the earth's surface. II., Geogr. Rev. New York.
Jap., 26: 573-585. ^ Ovington, J.D., 1957, The volatile matter,
(26) Transeau, E.N., 1926, The accumulation organic carbon and nitrogen contents
of energy by plants, Ohio Jour. Sci., of tree species grown in close stands.
26: 1-10. New Phytologist, 56: 1-11.
134
VOLUME 4
BOTANY
THE OCCURRENCE OF TROPICAL PLANTS IN THE JAPANESE
ARCHIPELAGO AND ITS PHYTOGEOGRAPHICAL SIGNIFICANCE
YOSHIWO HORIKAWA
Botanical Institute, Hiroshima University, Hiroshima, Japan.
(Abstract)
As one of the most characteristic features of
the Japanese flora, it is cited that several plants of
the tropical origin are growing wild there. This
fact was already clarified by me with respect to
bryophytes. As to the vascular plants, the follow-
ing is known about the distribution of some
species. According to their northernmost limit
of habitat in the Japanese Archipelago, the species
are divided into ten types:
Type 1 . ranging from Indo-Malaysia north
to 44 L.N. in Japan.
Trichomanes parvulum Poir.
Type 2. to 41 L.N.
Vitex rotundifolia Linn, fil., Albiz-
zia julibrissin Durazz.
Type 3. .__to 38 L.N.
Dicranopteris glauca Underw., Hy-
menophyllum bar bat um Baker, Dro-
sera spathulata LaBill.
Type 4. to 37 L.N.
Psiloswn nudum Beauv.
Type 5. _to 36 L.N.
Ceratopteris thalictroides Brong.,
Mecodium poly ant has Copel., Ly-
copodium cernuum Linn., Vittaria
flexuosa F6e. Anodendron affine
(Hook, et Arn.) Druce.
to 35 L.N.
Nephrolepis cordifolia Presl., Ipo-
moea indica (Burm.) Merr.
Type 6. -
Type 7. __
_to34L.N.
Histiopteris incisa J. Smith, Humata
repens Diels, Cocculus laurifolius
DC, Solarium aculeatissimum Jacq.,
Type 8.
Dichondra repens Forst, Ipomoea
pes-caprae (Linn.) Sweet, Senecio
scandens Hamilt. ex D. Don,
Dianella ensifolia (Linn.) DC.
...to 33 L.N.
rnacrorrhiza (Linn.)
Alocasia
Schott.
Type 9. . ... . to 31 L.N.
Ficus retusa Linn., Kandelia Candel
(Linn) Merr., Messerschmidia ar-
gentea (Linn., fil.) Johnston.
Type 10. to 30 L.N.
Blechnopsis oriental is Presl., Bias-
tus cochinchinensis Lour., Melas-
toma candidum D. Don., Spinifex
Uttoreus (Burm. fil.) Merr.
(The horizontal and vertical ranges of each
species are to be discussed with the frequency
maps.)
At the present stage of our knowledge, it is
hardly possible to form a safe conclusion about
the phytogeographical explanation of the occur-
rence of these tropical plants in the Japanese Ar-
chipelago. It is more likely that these species
were once distributed much more widely in tem-
perate regions of East Asia, such as China, Korea,
and Japan. Jn later times, through climatic
change, they were unable to compete with more
aggressive species and have been reduced to their
minor role in the vegetation. The coast of the
islands of Japan has been warmer under the
influence of the warm sea-current of "Kuro-
shio," and they may have survived here as living
fossils.
135
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
UNIFYING CONCEPTS IN VEGETATION STUDY AS
APPLIED TO THE PACIFIC BASIN
FRANK E. EGLER
A ton Forest, Norfolk, Connecticut, U.S.A.
My role in summarizing this symposium on
Vegetation Types of the Pacific Basin is fraught
both with customary and uncustomary difficulties.
J should state first that my position as Chairman
of this program is only as a latter-day substitute
for Dr. Pierre Dansereau, to whom should go full
credit for organization, and who regrettably was
unable to come to Bangkok despite earlier plans
to the contrary.
During the course of this symposium we have
been presented with several succinct and admira-
ble contributions. At first glance, it may appear
that these are totally disconnected and unrelated
"islands" in the "sea" of vegetation (plant com-
munity) science. Perhaps we should face one fact
at once: the terrestrial vegetation of the Pacific
Basin, as distinct from that of the rest of the
world, has but one element that binds it all
together a vast mass of water called the Pacific
Ocean which is not a substratum for terrestrial
vegetation, and acts as the best possible agent to
prevent the unification of that vegetation. Thus
the Pacific Basin is a geographic entity, united
more by the common interests of its human popu-
lations than by any fundamental vegetational
relationships.
The papers presented in this symposium serve to
place the spotlight on a few elements of the whole
problem, much as if a spotlight were to pass over
a crowded stage, temporarily lighting a few per-
sonalities, but leaving all the rest in darkness.
This momentary lighting however is very im-
portant. It serves to add a definite increment to
our knowledge of Pacific Basin vegetation, even
as does each succeeding Pacific Science Congress.
My purpose at this time is no more than to place
these papers in the larger framework of such a
stage.
There are several "viewpoints" from which
vegetation may be approached. No one view-
point has precedence over any other. In the
rest of botany, for example, the morphologist
does not vie with the physiologist as to which is
most "important," or which comes "first." So it
should be with the students of plant communi-
ties. The floristic composition of vegetation is
one such viewpoint. Within the Pacific Basin,
136
the floras segregate into at least three majoi
groups. There is a circum-Arctic flora on the
mountains near the Alaska coast, out across the
Aleutians, and down the high elevations of the
Kamchatka Peninsula. This flora, giving rise tc
tundra types of vegetation, has not been repre-
sented on this symposium. Progressing south-
wards, we find temperate floras that have more in
common with themselves than with the arctic 01
tropical floras that bound them on each side.
Becking, Tatewaki, and Chi-Wu Wang have each
contributed to our knowledge of these areas,
The tropics, by comparison, is a relatively unified
region, characterized in its vegetation by the
relative absence of damaging cold weather (al-
though how that damage is described in purel}
meteorologic terms is still an ecologic puzzler),
Porleres, Hiirlimann, Kira, Horikawa, and Fos-
berg have given us interesting information or
vegetations developed from these floras. Of these.
Fosberg has handled the atoll problem, where the
flora is attenuated to point of poverty, thus
greatly simplifying some aspects of vegetation
study. The temperate floras of the southern he-
misphere, of Australia, New Zealand, and South
America are not represented on this symposium,
except for Fosberg's comments on the montane
grasslands of the Andes.
Passing on to a second viewpoint, interest in
the form and structure of vegetation is a major
preoccupation of some vegetation scientists,
Becking, in handling northwestern United States,
is concerned largely with the make-up of vegeta-
tion in terms of the plant communities, according
to the ideas of Braun-Blanquet. Hurlimann is
concerned with structural details of a tropical
forest. Thirdly, Dansereau has presented us
with an expression of his system of shorthand
symbols for recording the structure of communi-
ties. Like most systems of shorthand, it is be-
wildering at first look, particularly this lollipop-
oriented one. (Freudian overtones specificall>
not implied!) Furthermore, Dansereau is ver>
much alive, and I fear there may be revisions tc
the revisions before the gentleman passes on tc
a stable end-stage of heavenly climax, revisions
that make it difficult for ones of limited mentalit)
like myself to keep up with him. It was ques-
tioned from the audience as to whether this
system had been or could be used in the floristical-
ly more complex tropics. It is my personal
opinion that it is in areas of just such complexity
where this system might prove most serviceable,
in that it describes structure unrelated to details
of floristic composition, which details for most
tropical regions are not adequately known.
One micro-facet of the morphology of plant
communities is that of epiphyte communities, of
those aggregations of plants which utilize another
plant as a substratum, and which can be studied
as a separate element in the total phytosociologic
picture. Hosokawa and his colleagues have car-
ried on a great many studies with epiphyte com-
munities, both in Japan and on Pacific islands.
Their present contribution is one more in a
notable series.
Another aspect of the form and structure of
vegetation, related not to the different communi-
ties involved, or to the structure of those commu-
nities, but to whole mosaics of communities, is
the grassland-savanna-forest problem. These
types of vegetation are found in close association
with each other in many parts of the tropics.
Their interpretation as being "natural," or
induced by primitive tribes and persisting into the
present as "relicts," or induced by contemporary
agricultural and pyric factors, is a phase of scienti-
fic investigation discussed by Porteres and Fos-
berg. It is a fascinating field of inquiry not only
for the academic mind, but for the Vegetation
Manager who wants to manipulate and convert
vegetation, sometimes into types markedly differ-
ent from what now exists.
In the third "viewpoint," that of functions and
processes, it is quite interesting that not a single
paper is developed along the traditional lines of
"plant succession" to "climax," with an orderly
diagram showing the origins of the various
"seres," culminating by means of arrows, in one
final end-stage. Nevertheless, all the speakers
have considered adequately and worthily the
various dynamic functions that are involved in the
life-activities of their various vegetations.
The "ecologic" (environmental) approach, a
fourth viewpoint, is coincidentally absent as
the major approach in any one contribution to
this symposium. This approach, dominant in the
thinking of possibly the major part of vegetation
research, at least in America, is not omitted from
this symposium by intent. Although we have no
paper entitled "The influence of climate on the
distribution of . . ." or "Soil chemistry as related
_yoLUME_4 BOTANY
to the vegetation pattern in . . .," there has been
no slight, with any speaker, as to an awareness
of the importance of the environment in the
behavior and phenomena of vegetation.
The geographical distribution of vegetation
types is a fifth viewpoint, and cartography (map-
ping) is one of its most important aspects. In this
branch of knowledge, Kuchler has distinguished
himself, particularly in accumulating a world
bibliography of vegetation maps. His presenta-
tion and views serve excellently to round out
this symposium. Aerial photography has enor-
mously enhanced our facilities for vegetation
mapping in recent years. It should be pointed out,
however, that the vegetation types which are dis-
tinguishable from the air are generally the phy-
siognomic types, that is, types distinguished as
forest, grassland, savanna, and desert. Equally
distinguishable are types dominated by an aer-
ially visible species, such as pine forests in a
mixed hardwood region. There are dangers as
well as advantages to this facility, for it leads us
to overlook the fact that there may be other and
even more significant kinds of vegetation types
which are not related to these aerially visible
differences. Therefore one cannot overemplasize
the need for accessory studies on the ground by
competent botanists.
A sixth and final viewpoint, that of distribution
in time, or history, has also been underplayed,
if we view its importance in an ideal vegetation
science. Both Porteres and Fosberg have
touched the subject in their inquiry on the
possibly-human origins of grasslands and savan-
nas, and Chi-Wu Wang has considered the history
of the Chinese forests as it extends back into
geologic time.
As a postlude in one sense, and as an open door
to larger vistas in another sense, the spotlight
should be played again on Fosberg's treatment
of atoll vegetation. The idea implicit here is
developed more extensively by him in another
paper delivered elsewhere at this Congress, "...
Description of the Coral Atoll Ecosystem."
Now in all other papers of this Symposium, we
have considered "vegetation" as our subject of
study. For many years, however, scientists have
realized that vegetation is but part of a larger
integrated whole, called by such names as eco-
system, microcosm, landscape, and "organism"
(not the biologic organism). The obvious intellec-
tual advantages of this approach are often offset
by the very complexity of the natural phenomena
involved, and the fact that adequate scientific
knowledge from many disciplines is seldom
137
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
encountered in one man. The atoll, by its very
nature, is a distinctive microcosm, existing at the
interface of sea and air, with energy absorbed
from and radiated to its environment, with a
sort of feed-back mechanism that keeps the
system in a relatively steady state. From this
view, the vegetation all but loses its separate
identity in the structure and behavior of the larger
"whole." It is not to be implied that the same
conceptual approach is immediately applicable to
continental areas, with their enormously greater
complexities, and less clearly defined boundaries
in space. Fosberg's researches greatly advance
our ideas in these conceptual realms, and indicate
that the views are both feasible and practicable.
In closing this symposium, which had been
planned by Dansereau as Chairman of the
subcommittee concerned with Vegetation (plant
communities), I wish to say that in my opinion
this subcommittee, acting within the framework
of these Congresses of the Pacific Science Associa-
tion, has a brilliant future ahead of it. A unified
vegetation science is just beginning to emerge
from the various nationalistic disciplines which
originated mainly in North America and Europe.
There is now no other area in the world than this
Pacific Basin where scientists of so many nations
can cooperate so advantageously, with the vigor
that comes from the interplay and interchange
of ideas, of problems, and of research accom-
plished. I look forward to seeing this vigor
translated into a distinct world contribution
towards the description and understanding of
the world's vegetation.
138
BOTANY
THE SCHOOL-FLORA AS A MEDIUM FOR
BOTANICAL EDUCATION IN THE TROPICS
C.G.G.J. VAN STEENIS
Flora Maletiana Foundation, Oegstgeest, Netherlands.
A proper understanding of plant structure is
an absolute necessity for botanical education,
without which pupils of the secondary schools
the source from which university students will be
recruited will not be able to understand plant
function, plant ecology, flower biology, forestry,
and agriculture.
The identification of a living plant by means of
keys and a description is still the best means to
gain an insight in its structure and its contrasting
features with other plants. Plant atlasses may
seem to lead to the same result along a more easy
way. But this is educationally basically wrong,
it leads only to a vague notion of the general
habit of a plant and to its name, not to any knowl-
edge of its structure.
Most Floras have a dualistic character, they
tend to give a complete survey of all the species
of a country or a more restricted area and must
serve simultaneously for education.
In tropical countries this dualism is untenable
by the wealth of the floras, and would necessitate
large and expensive works in which the educative
purpose is entirely lost. Such Floras would con-
tain descriptions of plant forms from remote
mountains and forests which could never serve
for regular class demonstration and which no
teacher could be expected to know himself, let
alone have available in the living state in sufficient
quantity for class teaching.
Complete Floras may be desirable for botanists
and advanced amateurs, but for school-children
they represent the 'Tables of the Law', they have
no educative value and are deterrent rather than
awakening interest in plants.
It is compulsory that in the class each pupil
has in hand a living specimen and the Flora, with
at most a pin or needle as a simple utensil; lens
and forceps will generally be not available and
are unnecessary for the purpose.
The idea is that the identification of a plant is
performed in the class, step by step, and that
under guidance of the teacher each pupil can
follow the way along which the ultimate goal,
the name, is reached with subsequent checking
of the description. This simple botanical detec-
tive work must give satisfaction.
Such a School-flora must be adapted to the
following points:
(a) A selection of plants, hence no complete-
ness whatsoever.
(b) Availability of at least a number of species
incorporated in it, as the teacher must be able to
collect for one lesson enough material to give a
specimen to each pupil. Consequently the choice
of selection is restricted to plants available in or
near towns and cities.
(c) Extensive use should be made of cultivated
plants from school-gardens or nurseries, town
parks, roadsides, etc., resulting in inserting in the
Flora numerous cultivated and ornamental plants.
(d) To bring pupils into contact with biotopes
in his immediate vicinity brings along the insert-
ing in the Flora of some typical plants of beach,
mangrove, rice-field, swamps or pools, dry-
farming fields, estates, waste ground, secondary
growths, etc.
(e) The absence of a lens and forceps and
knowledge how to use these necessitates the
omission of plants with small flowers which
escape easy observation with the naked eye.
Besides, there is no educative advantage in dis-
secting small flowers against using large ones.
(f) There must be no obscurity in the identifi-
cation keys to the family, genus or species, and
when the name of the latter is reached there
should be a clear not too elaborate description for
checking the identity. Great care should be given
to attain a faultless text.
(g) Interest in the plant should be raised in
adding where desirable some brief additional
notes on uses for mankind, biological topics, or
other noteworthy data.
(h) In absence of adequate pre-knowledge
technical terms should be avoided and the keys
and descriptions should be as much as possible
worded in plain language ; a plate showing schema-
tic pictures to elucidate necessary technical terms
may be handy.
(i) Representatives of the most important
tropical families should be inserted by native,
introduced or cultivated species.
139
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
With such a flora only a select number of species
can be identified; it serves merely for educative
purposes', the idea of completeness is entirely
abandoned.
If we imagine that during the courses in the
secondary school not more than about 25-50
species will be identified in the class, the School-
flora should contain not more than about 300-
400 species in all.
With a few collaborators I have realized such
an octavo School-flora for Indonesia, with about
400 species belonging to about 300 genera and
120 families. It covers about 400 printed pages 1 .
It is not the intention that pupils will know or
learn all these 400 plant species; no more than
10% can be treated in the class. But the number
of 400 leaves room to the teacher to make his
choice in accordance with available plants in
sufficient quantity in his vicinity. Representative
species of both the everwet and seasonal regions
should be inserted. It further allows pupils who
are interested to use the book for their own
pleasure.
School-floras, like Alston's The Kandy Flora'
(1938) and Merrill's 'Flora of Manila' (1912)
follow a scheme which is less adapted to practical
use as our School-flora for Indonesia. Both
contain generic diagnoses which I feel are un-
necessary for secondary schools.
My main objection is, however, that they re-
present a complete flora of the vicinity of a town,
Kandy and Manila respectively, without restric-
tion to easily available plants for class teaching
and selection on size of flowers. They both still
contain a dualistic element (serving education and
botany) and are not entirely devoted to the goal
of being merely a tool for educative purposes.
In the School-flora for Indonesia we could not
avoid mention of such important groups as
grasses, sedges, and ferns, and have been com-
pelled to add some simple pictures for a few re-
presentatives of these groups.
The insertion of common and widely distributed
ornamentals, weeds, fruit trees, wayside plants,
etc. is convenient to use the flora in the whole of
Indonesia in cities and towns below 1,000 m
altitude, the site of most training centres. And
I am rather certain that it could for the same
reason be used in most parts of tropical southeast
Asia, Micronesia, and Melanesia.
I believe that the model of a School-flora of
which the principles have been just outlined, is
a valuable tool for education purposes and the
wakening of the interest of pupils.
I find it desirable to bring these principles to
the general notice of teachers and hope it will be
a stimulant to the writing of similarly adapted
School-floras in other tropical countries.
DISCUSSION
F.R. FOSBERG: E.H. Bryan, of the Bishop Museum is
writing the flora of Guam which fills Van Stccnis require-
ment for School-flora. The common species are empha-
sized so that the ordinary people of Guam can learn. Such
books acquaint young students with the nucleus of botany
before college time.
C.G.GJ. VAN STEENIS: In Java, education in schools is
faced with the problem that knowledge of Java's flora
(4,500 species) is impossible. Therefore it is a necessity
to have easy tools to instruct people in the morphology of
certain plants. One disadvantage is that the teacher must
know plants and names himself.
M.L. STEINER i I do not agree with the exclusion of small
species of flowers.
C.G.G.J. VAN STEENIS: For secondary classes it is suffi-
cient to study only large flowers.
M.H. SACHET: I disagree.
F.R. FOSBERG: People may learn botany, but not of
plants. America has a tendency not toward School-flora
but toward picture books which form bad habits.
p. WEATHER WAX: A written examination would find
out if students can use the key.
V.A. JIRAWONGSE: Is there an advantage in using just
vernacular names?
C.G.GJ. VAN STEFNIS: Vernacular names are different
in different parts of the world, and Latin names are used.
"Flora voor de scholen in Indonesia". Noordhoff-Kolff, Djakarta, 1st ed. 1949, 2nd ed. 1951.
140
VOLUME 4
BOTANICAL EXPLORAffON^AND EDUCATION IN
PRESENT DAY SUMATRAt
BOTANY
W. MEIJER
Faculty of Agriculture, Pajakumbuh, West Sumatra,
SCENERY AND BOTANICAL RICHNESS
IN SUMATRA
Sumatra, as compared with Java, is a land of
promise for botanical exploration. On the
densely populated island of Java, most of the
lowland flora is destroyed, and almost the whole
country is divided among agriculture, forestry,
cattle-breeding, and fishery. Botanical explora-
tion has already been so thorough in Java that
only rarely are new plants found. Bogor is still an
important botanical centre on Java. In Sumatra
we find a quite different situation though not
everywhere. It is not only a land of promise but
also of contrasts. Regions like Djambi and
Indragiri are still largely covered by primary
forests in which are found elephants, rhinos, and
tigers. Only 2-5 plant specimens have been
collected per 100 sq km in these regions. I read
in the newspaper that the tribes of the primitive
Kubu-people are still living in these forests. It is
told that they walk undressed and are forbidden
by their own law to plant.
Quite different situations are found in other
places: the Deli-region in the Northeast has been
widely cultivated and is still the area of big estates
of oil palm and tobacco and rubber. Minangka-
bau on the West Coast is to be seen from the air
like an enclave between the jungles of South
Tapanuli, Indragiri, Djambi, and Bengkulu. It
was long ago colonized by people who made this
a region one of mixed garden agriculture and who
still preserve their special customs. The land-
scape has much the same features as Java, but
agriculture is far less intensive than in Java
and the flora of the kampongs also shows many
differences. Minangkabau people possess, for
example, their own races of rice (hundreds!);
their own names for and assortment of bamboos ;
their own names for at least twenty-five different
kinds of cultivated bananas; their own way of
planting tobacco, cinnamon, sugarcane, clove
(tjenkeh), and gambir; their own way of burning
grassfields and secondary forests, of plundering
the forests, taking no care of silviculture, land
and soil protection. It is very difficult to change
cultivation customs which cause destruction of
t Presented by J.V. Santos.
the landscape and impede further development.
Sumatran students entering the new University
Andalas are not selected among the Kubus or
among kampongs far from the town; they are
largely the sons and daughters of businessmen
and government officials urbanized people who
have lost contact with their rural surroundings.
They have been strongly influenced by the
blessings of modern culture from America
and Europe, but are still in search of their own
style.
Pajakumbuh, is an ideal spot for botanical
exploration and education. Here we find in a
region of 10 sq miles (about 14 km square),
a flat basin with mixed gardening and marshes
(Pajakumbuh means the plain of marshes),
surrounded by hills on which tobacco and gam-
bir are planted and forest clad mountains com-
posed of three different types of rocks: sandstone,
limestone, and the old volcano Mt. Sago. The
lowland forests, rich in Dipterocarpaceae, of the
large East Sumatra coastal region are on a dis-
tance of about 1 5 miles. The whole flora of the
Pajakumbuh region probably has at least 2,000
species of flowering plants. For Mt. Sago
alone, I could list more than 800 species. The
whole phanerogamic flora of Sumatra may be
estimated as about 10,000 species including, for
example, 800 orchids.
When we read in Flora Malcsiana that the
collecting density in W. Sumatra per 100 sq km
is 38 (332 in W. Java), we can understand why
our recent collections from the Pajakumbuh
region contain numerous novelties new to
science, or plants hitherto known only in Malaya,
or collected only in South or North Sumatra.
Several of these plants are really common in
special types of vegetation. It is becoming clear
now that forest types are strongly correlated with
soil types, the so-called climax theory has no
scientific basis! The forest vegetation on lime-
stone hills is quite different from that on sand-
stone hills. Analogous differences are seen in the
weed vegetation and the cultivated races of rice.
Limestone regions of the plain are very different
from sandstone regions. The latter appeared to
me in need of phosphate manure which could be
141
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
found in caves in the limestone hills! We are
drawing the attention of the agriculture service
to this situation.
We have already more or less compared the
flora of Mt. Sago with several other mountains
in Central Sumatra. Impressions of the lowland
forests were collected in West-Indragiri. The
sandstone region was the subject of the most
recent expedition. More thorough studies of
the limestone flora and of marsh forests remain
for the future. We are presently making a
thorough inventory of economic plants in the
Pajakumbuh region devoting attention to all kinds
of fruit trees, vegetables, legumes, races of rice,
weeds, and secondary vegetation. We are able
to advise and help the population in the rural
reconstruction area served by our Faculty to
use their waste lands more efficiently and to
improve their mixed gardens. The culture region
is far less productive as compared with Java.
Our hope is that investigation will precede
long-run planning in Central Sumatra and that
natural resources will be better preserved than at
present. Reforestation should become more
than incidental planting of Pinus, Acacia auri-
culaeformis, and some Toona sinensis. Good
measures against burning and for forest and soil-
protection should be taken. The farmers ought
to be educated in this direction.
We are ourselves trying to do this in our recon-
struction area. We have succeeded in getting
the road improved and have gathered all kinds of
informations about economic botany, started a
reforestation project and have built a nursery-
garden. We are now engaged in project to plant
the waste hills which are burned every year and
show heavy signs of erosion. Our exploration task
cannot be separated from our informations and
education. The local population needs much
information and education. People are no
longer content with their standard of life. Indone-
sian society is in transition! Descriptive botany
and botanical ecology are very useful and practi-
cal in tropical agriculture !
THE ROLE OF BOTANICAL EDUCATION
IN SUMATRA TODAY
There will be no practical application of our
studies in forestry, cattle-breeding, fishery and
agriculture because they have no role in the
botanical education of the country. Technical
assistance for this requires more than three years.
The Dean of our Faculty estimates that ten years
will be necessary to get Sumatra's own people
142
sufficiently interested and educated in scientific
botany.
Are the facilities for such an education already
provided at the new University of Andalas ? No ;
after two years of struggle with many kinds of
difficulties, a botanical institute is being built, but
because of a lack of equipment, it is still impossible
for the departments of plantphysiology, micro-
biology, and cytogenetics to do their work on
a modern level. Our Faculty is still without any
laboratories for physics, chemistry, and agro-
nomy. For social-economic science we have
nobody! In Indonesia it often takes two years
before ordered instruments arrive. The education
of skilled personnel takes much patience and
good-will. Minangkabau people have another
discipline of work than Europeans or Americans.
Foreign medical doctors feel the same difficulties,
but fortunately self-education and self-criticism
are also known here. The non-experimental
parts of botany can start from the first days of
arrival of a field botanist. Old newspapers for
drying plants are to be bought on the local
market; bamboo sasaks are easily made; and the
sun is the great drying oven: Kampong people
may be found who know the roads into the forests
and many local tree names. The botanist must
be able to get the authorities interested in his work ;
he needs transport, funds, and even soil for expe-
rimental gardens. Knowledge of the Indonesian
language to educate the students in botany is
very helpful. A good exploration of the nearby
kampongs and mountains makes it possible to
teach botany outside the lecture room in its
natural surrounding. Then you have the simple
task of showing the new students for the first time
in their life what cotyledons really are (the word
Dicotyledons they know) ; opening their eyes to the
special structure of flowers of coconut palm,
banana, and papaya; teaching them to distinguish
fruit trees, legumes, vegetables, curbits, green-
manures, and ornamentel plants; teaching them
to observe plant forms, to describe them, and to
learn how to make use of keys for identification.
It is astonishing how little is known about this;
a lot of goodwill, patience and perseverance is
asked from the teacher. As soon as microscopes
Arrive in my case after one year and the funda-
mentals of plantmorphology and knowledge of
plants in the surroundings are understood, you
may start with laboratory courses in plant anato-
my and cryptogams. Please do not look for Rhoea
discolor to demonstrate plasmolysis; use the red
leaved Cannas one meter from your door. As to
anatomy, don't look only for the classical exam-
ples, but take Portulacca, Sida, and Amaranthus
VOLUME 4
BOTANY
growing along the door. The classical systems
of vascular bundles without secondary growth
seem to be an exception among tropical plants.
Show the students who know only red and white
rice, the hundreds of varieties growing in the
surroundings. They will be astonished, enjoying
the wide range of variation like every taxonomist
enjoys seeing how nature plays with forms.
Make the students aware of the fact that their
own country should be the real centre for cultiva-
tion and breeding of bananas, not Central or
South America. Eradicate the theory that bana-
nas were imported from America into Indonesia;
eradicate the idea that the clove came from
Zanzibar, that the local Rafflesia the biggest
flower of the world is insectivorous. Demon-
strate the wild species of Musa growing on
Mt. Sago how you describe them and document
your finds with photographs, how you further
investigate the many cultivated varieties. Don't
teach them a mass of facts, but methods and
synthetic views. What a difference between an
ordinary paddy-field seen as a basin with rice or
seen as a dynamic whole, a habitat in and
around which phanerogams, weeds, algae, bac-
teria, soil composition, and the origin and
breeding of rice-varieties all play a role ! What a
difference between a kampong as an ordinary
collection of fruit trees and vegetables and the
same seen through the vision of a botanist who
points out the origin of some elements in the
forests which grew here during former times;
the influence from India, from America; the
influence of Dutch agriculture; the richness in
varieties, the flower and seed biology and
embryology of Manggas, Djeruk, Djambu, and
Bananas; the role played by algae and water-
plants in small fish ponds fertilised from the
primitive houselets above them.
The most clever and interested students may
take a part in expeditions to the highest mountain
summit of Sumatra (Kerinchi, 3,800 m) or in the
lowland forest of Indragiri. They learn the most
common forest plants; they learn how to organ-
ize an expedition, how to collect plants, how to
documentate the collection. They become
teachers in secondary schools, discuss with you
during evenings in the bivaques the best system
for teaching botany there and arrange for you to
give popular lectures to the local population,
pupils of secondary schools, and their teachers.
It might be necessary then to borrow clean trou-
sers from one of your students, but you will find
your own improvised lecture in the newspapers,
and the radio will broadcast that according to
you botanical study is very important for daily
life of the people.
When you tell people living along the coast that
the brown algae like Sargassum has in principle a
method of reproduction analogous to that of
human beings, a clever boy may ask you to tell
the how and why of this and he hears something
about sperms and egg cells. A school teacher
living at 1 km distance from the coastal mangrove
asks you after your lecture to repeat for him what
mangrove forest is and where he can find it.
Local authorities want to hear from you about
the possibilities of agar industry. Don't fall into
this pit.
KAMPONG PEOPLE TEACHING A BOTANIST
Is a botanist in tropical regions only explorer
and educator? No; he himself learns very much
from the simple illiterate kampong people.
They know their varieties. Some of them know
trees in the forest, only from making a cut into the
bark. Why not learn that? Some people may tell
you about plants to be used as medicines. You
may find drugs which are useful but unknown to
the academic doctors, Indonesian or foreign, and
you may find a good basis for botanical education
of students in medicine and pharmacy.
COORDINATION BETWEEN BOTANICAL WORK AND
LOCAL SERVICES AND AUTHORITIES
Especially growing institutes need a lot of local
support and local understanding. At the start in
Pajakumbuh, we were without housing and often
without our own transport for excursions and
expeditions. The behaviour of a botanist looking
everywhere in gardens, in paddy fields, and in
forests for plants was badly understood by the
population. What do you look for? What is the
purpose of your work ? I was asked these ques-
tions everywhere. I made it the subject of a pub-
lic lecture on the first anniversary of the Faculty.
After that popular lecture, I got spontaneous
help from the head of the local civil service to
build an emergency herbarium, a building of
8x14 meters from which several thousand dupli-
cates are already distributed to London (Br.
Museum, Nat. History), Kew, Leiden, Amster-
dam, Geneva, and Singapore. Officers of the
forest service joined several of my expeditions.
The trip to the summit of Mt. Kerinchi was simul-
taneously done by the head of Forestry with
several of his officers. Many problems of forest
botany, reforestation, and rural reconstruction
are often discussed with my forestry friends,
semi-academic Indonesians. All projects for
143
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
nature protection are jointly made with the forest-
ry service.
The head of the service for agriculture is still
like a father of the Faculty. He started it locally,
and he supports our work in every way. He is
as much disappointed as we when we cannot get
the money and equipment from the Central
government. With the head of the Veterinary
service, we started our project for local recon-
struction at the base of Mt. Sago. It is not neces-
sary for us to tell the leaders of the Forestry
service that their officers rarely enter the forests,
to tell the agriculture service that its apparatus
has very little practical use for the population, to
tell the veterinary doctor that the grasslands on
the experimental station are badly treated by
burning, etc. They all know this, but they are
waiting for our students who shall have to im-
prove the present situation. The country waits
for a new, better educated generation. We are
clearly confronted with problems in our desa
pertjobaan, rural reconstruction area, two kam-
pongs with mixed gardens, surrounded by paddy
fields, hills and with above them the forest of
Mt. Sago. How are we able to advise, encourage,
and educate the population here? Botani-
cal inventories, socio-economic research by the
students, practical projects for a cooperative
movement including evening courses to be given
by students are started here. Thus our work
finds roots before modern laboratories and equip-
ment are provided. Of course the illiterate popu-
lation is not awakened within one year from its
apathy, but there is more movement than we
anticipated a year ago when this project was
started.
INTERNATIONAL CONTACTS AND HELP
When agriculture students after three years of
study receive only laboratory courses in botany,
not any practical education in physics, chemistry
and agriculture we can speak about mismanage-
ment among the authorities who started this
Faculty and who have the responsibility for its
welfare. We must realize, however, that Indo-
nesia is a land in transition with many financial
and economic troubles. As long as these troubles
are not resolved, reconstruction of the country
remains difficult. It is, of course, disappointing
to educate students in agriculture in botanical
subjects while other subjects like chemistry and
agriculture itself are largely postponed by lack of
teachers, equipment, and working rooms. It is
very irritating that improvement of this unsatis-
factory situation is so long postponed. Visits of
144
several foreign agencies UNESCO, Ford Foun-
dation, 1CA, always came at the most inapprop-
riate time. Now only some local oil companies are
giving support (for chemistry). The best help we
received was for the library, especially from Ame-
rica, after the Botanical & Gardens of Bogor
gave it a good start.
As to botanical work, it is, of course, almost
impossible to collect at new institutes within some
years all the old books and journals which are
necessary for revision work and monographs.
This is another reason to concentrate on explora-
tion, documentation in the field, and to distribute
as many duplicates as possible of the valuable
collections. The returning namings will help in
the work of ecology and plant geography.
The amount of work to be done (Stencilled
lecture notes must be prepared for the students as
no books in their own language are available,
and they cannot read the Dutch language), is so
over-whelming that almost no time is left for
preparing scientific papers. Work of years ago
still remains unpublished. Then we get requests
from abroad to send material living or in liquid.
We have to go to Java to get the chemicals for
fixation; we are not able to obtain glasswork
with stops and often cannot fulfill the wishes of
foreign colleagues because of lack of time and
provisions. The British Museum (Natural
History) got a living specimen of Rafflesia by
aeroplane. Nobody understands how many
bureaucratic barriers had to be broken down for
that within some days. I had first to get all kinds
of permits from Djakarta and, was not allowed to
pay until a Dutch firm took over the responsibil-
ity. The Faculty waited two years before the first
foreign journals entered the library. A large
botanical institution in America, formerly special-
ising in the flora of Malaysia, writes sour and
formal letters to me in reply to enthusiastic pro-
posals which are for mutual benefit. Other
institutes like Amsterdam, Leiden, Cambridge,
British Museum and Singapore are largely
cooperating and stimulating.
THE FUTURE EAST AND WEST
The work of botanical exploration and educa-
tion in tropical regions needs the help and support
from temperate regions of the globe. There are
the books, the old publications, the old centres of
botany, the treasures gathered by Blume, Miquel,
Hooker, Ridley, De Candolles, and Merrill.
There are still many more botanists working than
here. Here we have the richest part of flora of
the world still to be studied.
VOLUME 4
BOTANY
What will be the future of the tropical forests?
Will they be destroyed before scientific interest is
raised among the people of these countries?
Must all foreign botanists go away from here?
Will it be possible for a botanist from Europe
or America to devote his whole life to the richest
flora of the world, or will political boundaries and
nationalistic movements force him to restrict
himself to the scanty flora in his own country or
to shift his attention to parts of the tropics which
are still under colonical rule? History teaches
us that the fate and function of botany in our
world cannot be separated from international
cooperation.
In my opinion, this is understood by the grow-
ing generation of Indonesian scientists, the best
of them being further educated in Europe and
America. The joint use of natural resources of
the world will have its natural equivalent in joint
exploration and research.
CONCLUSION
This lecture started like the talk of a circus-
artist and ended like a sermon. I hope that you
understood that botanical exploration and educa-
tion has found a good start in Central Sumatra
itself, but that this young plant has still many
difficulties in growth. The climate is often frus-
trating and disappointing for it. Much skill,
perseverance, and a sense of humour are needed
for its growth. Have we to laugh about this
undertaking or to pray for it?
145
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
PROBLEMS OF PUBLICITY: WITH SPECIAL REFERENCE TO
THE NEED FOR POPULAR BOOKS DEALING WITH LOCAL PLANTSt
R.E. HOLTTUM
Kew Gardens, Surrey, England.
In the modern world we are all increasingly
dependent on the applications of science, and
education for the young must include an intro-
duction to both physical and biological science
if they arc to have an intelligent understanding
of their environment. And it is not only the
young who need to have access to such knowl-
edge; there is need for popular publications for
adults, to keep them up to date with the changing
world.
All life is dependent on the green plant, and
though modern technology has greatly changed
our ways of living, it has not altered that fact.
But the modern habit of living in towns and cities
tends to prevent a great many people from
realizing the fact, and therefore education in
biology is of even greater importance than it was
when the majority of people lived in close depend-
ence on their natural surroundings.
Modern political developments have added a
new urgency to all aspects of education. Demo-
cratic self-government, to be effective, depends on
an educated electorate.
Thus the first essential of education in biology
is an understanding of the significance of plants
as the basis of all life and as a basic natural
heritage at the disposal of mankind, and education
of this nature must be presented in terms of local
plants in their relation to the local climatic
environment. Knowledge of this kind is being
gradually acquired at the various tropical bota-
nical institutions. Such institutions thus have a
duty to contribute knowledge about local plants
which may be used in this necessary education in
biology. And such institutions will in the future
depend for their very existence on a general
understanding of their importance by the public
and by the politicians elected to power by the
public. Publications of this kind become neces-
sary publicity from the point of view of the
institutions themselves.
Biological science is not like physical science,
the important data of which are independent of
environment. Every different tropical country
has different plants and animals, and there are
many differences of climate which control and
f Presented by J.V. Santos.
146
limit the activity of those plants and animals,
and also of man in exploiting them for his own
use. Thus there is need for different presentations
of these subjects in different countries; one general
text book, or a series of publications from a
single source, will not meet the situation.
The great difficulty in preparing such educa-
tional and popular presentations of a biological
nature lies in the extreme complexity of the sub-
ject matter and also in the fact that there has been
far less biological study in the tropics than in
temperate regions; and studies made in tem-
perate regions are only of limited help because
they deal with a different set of organisms in a
different environment.
Before anyone can write a simple introduction
to a subject for the help of beginners, he must
himself be master of the subject, or at least have
a wide enough knowledge to be able to select for
the beginner those aspects of it which will be most
helpful. In order to obtain the necessary wide
outlook, one must spend a great deal of time in
study; and this involves not merely the study
of books already written, but also the local
plants themselves, as existing works cover only a
fraction of what one needs to know. Original
observation and the correlation of such observa-
tion are creative processes quite different from the
assimilation of knowledge already recorded in
books. The ability to carry out such creative
work is something that comes only from long
practice.
As an example which has come to my personal
attention, I will refer to the question of bamboos
in Malaya. Bamboos are of great importance to
the countryman who uses them every day in
many ways, and any student of biology in Malaya
should learn something about them. But there
is nothing in print about bamboos in Malaya
which is of any use to a person who wishes to
know how to begin to study the subject, nor
even any work which is of direct value to the
expert; the only usable taxonomic works have
been written in India and in Java, and they leave
untouched much of what is in MaUiya. Further-
more, nobody has given critical thought to the
VOLUME 4
BOTANY
general classification of this group of plants as
a whole (quite apart from Malaya) for nearly
a century. Some twenty years ago I began to
take an interest in the subject, and after a good
deal of casual enquiry I sat down to write a sys-
tematic account of what information was avail-
able, based on specimens and data accumulated
in Singapore. This showed me how inadequate
this information was, and during subsequent
years I made several journeys with the object of
adding to it. Having left Malaya, I cannot add
further original observations, and I have written
an account of the subject, as far as is possible on
present information, so that others may start
where I leave off; in particular, I have tried to
present an introductory statement which will
help a new observer to begin his work.
This work on bamboos is only a small aspect
of the study of Malayan plants, and it may be
matched in other families of plants and in other
parts of the tropics. But unless this kind of work
is done, the peoples of the world will not be able
to know the scope of their natural heritage of
plant life, and they may well, in ignorance, destroy
a large part of that heritage before it has been
studied. Such primary study of native plants
can be accomplished only by experienced workers ;
it is not something that can be undertaken by a
beginner. This is often not understood by ad-
ministrators and politicians, and even scientists
who are not acquainted with the problems of
taxonomy sometimes fail to appreciate the great
amount of such work which remains to be done
in the tropics.
Of course there is a great amount of informa-
tion already recorded about tropical plants.
Much of this information is in taxonomic mono-
graphs and formal Floras which can be under-
stood only by a specialist. There is need, as above
noted, for specialists to select from this mass of
data such parts as are useful as an introduction
to the subject. The university student, or the
school teacher in training, needs to have such an
introductory presentation. When the student
in turn becomes a teacher in a school, he will
find yet another problem that of introducing
children to the same kind of knowledge. The
teachers in their turn must thus review the subject
and write books which will serve their particular
needs.
From my present standpoint, the work of the
tropical botanical institution is, therefore, two-
fold: the continued study of natural vegetation
and the recording of the results of that study in
formal scientific publications and the interpre-
tation of that knowledge in terms useful to the
student (especially the student who will be a
teacher), and to the adult who wishes to develop
an interest in such matters.
The institutions will be able to do tins only if
there is a sufficient reserve of natural vegetation
and if they have sufficient trained and experienced
stalT to carry out both the fundamental investi-
gation and its interpretation for the student and
the common man. In these days, the institutions
will not be provided with the staff, equipment,
and means of publication unless the general
public is aware of the importance of these things.
The right kind of publicity is needed, and this
can come only from the institutions themselves.
It should be a function of this Congress to help
to impress on governments and on peoples that
the work of tropical botanical institutions, and
with it the preservation of natural reserves of
plant life, is a matter of fundamental and urgent
practical importance.
DISCUSSION
j.v. SANTOS: Mr. Holttum appears to support Van
Steenis' idea.
C.G.G.J. VAN SIEENIS: Dr. Holttum in his paper in-
tended that there should be more simple, popular knowl-
edge of plants. Dr. Holttum has written papers not
intended for secondary schools. Structure must start first.
F.R. FOSBERG: 1 am pleased to hear this discussion
about starting education in secondary school.
147
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
FUNCTIONS OF THE ALGAE IN THE CENTRAL PACIFICt
MAXWELL S. DOTY
Department of Botany, University of Hawaii, Honolulu 14, Hawaii.
The algal roles in nature at sea are those of
being the primary producers: primary producers
of food and materials in the euphotic zone. The
rates of algal production and perhaps the sizes
of algal standing crops are a function of the con-
centrations and rates of addition of otherwise
limiting chemicals to this euphotic zone and to
light intensity. To an extent these quantitative
population features are a function of the abilities
of the organisms themselves. The major roles of
the algae are related to whether they are plank-
tonic or benthic in nature.
In the open sea the algae are plankters, which
in at least some cases must lead a heterotrophic
life. As an example, the Coccolithophores domi-
nant in euphotic Mediterranean waters (3) have
also been found to be abundant at depths of 1,000
to 2,500 meters, far below the euphotic zone.
Such organisms may get there initially through
sinking or by vertical mixing. In our own work,
we have upon occasion found the most chloro-
phyll-rich waters at the very bottom of the eupho-
tic zone, if not below it. Returned to the surface
through upwelling or vertical mixing or rising
through changes in density, such organisms may
act as the "seed" for normal populations.
The food producers in the sea, the phyto-
plankton algae, make that 73 per cent of the
earth's surface covered by ocean as productive as
the land on the average per unit of area. This
production may be estimated variously (e.g., 20,
22, 9) between 0.8 and 15 x 10 l tons of carbon
per year.
The rate of primary production varies in dif-
ferent parts of the ocean. At the poleward limits
of the temperate seas with seasonal change and
the vertical mixing of various kinds that occurs
there, the substances normally limiting to phyto-
plankton production become more readily avail-
able. Measurements of phytoplankton product-
ivity have been found (15) to increase three-fold
as one moved into this productive water from the
temperate region. In the divergent current
regions of the Central Pacific along the equator,
water from the dysphotic zone upwells. In such
regions the productivity has been measured (16)
to be almost ten times that several degrees to the
north or south. In these two instances, the rate
increases are correlated with areas where the
inorganic materials normally limiting to algal
growth are being brought upwards into the eupho-
tic zone from below. These areas of major size
and consistent presence may be correlated with
high fish production: albacore tuna in the first
instance cited and yellow-fin tuna in the second.
Phytoplankton productivity rates increase as
land is approached. This has been shown (12)
to be an increase of two or more orders of mag-
nitude. The phenomenon is apparently due to
two processes leading to high concentrations of
essential minerals near shore. They are (a) con-
tributions from the land brought by water, and
(b) substances accumulated in the area from the
sea by benthic forms.
General algal productivity in the non-enriched
areas of the sea may be supported by recycling of
materials derived from the sea or initially from
emergent land. Such areas may gradually become
poorer in the elements essential for primary
productivity through the movement of materials
to layers of water below the euphotic zone.
Fixed nitrogen may be exceptional in that
supplies of this element in combined form may
be derived from rainfall or may dissolve in the sea
and be combined there by microorganisms,
particularly the blue-green algae. In fact on
atolls and in the sea, nitrogen fixation by blue-
green algae is expected to be a major as yet non-
assessed factor. If the major premises of this
and the previous paragraphs are valid, then a
relative increase in the ratio of nitrogen to, e.g.,
phosphorus might be measured in going away
from areas where there is enrichment from shore
or the depths.
It is of considerable interest to note the trend,
even in the study of benthic algae, away from
more descriptive aspects and toward more func-
tional aspects. Thus in past years we had oceanic
t The conclusions presented herein without citation were drawn from studies financed largely by contracts between the
Botany Department of the University of Hawaii and the U.S. Atomic Energy Commission (AT-(04-3)-15) or the U.S.
National Science Foundation (NSF G 1992 and NSF G 3833) or with the U.S. Office of Naval Research (NR 388-001)
with the U.S. National Research Council's Pacific Science Board, and by funds from the Graduate Research Committee
of the University of Hawaii.
148
VOLUME 4
BOTANY
phytoplankton studies concentrating on observing
the kinds and numbers of algae. The outstanding
example of this is the magnificent series of studies
by Lohman (17). Today we have concentration
on studies of oceanic phytoplankton such as
those of Steemann-Nielsen and Aabye-Jensen
(23), using isotopic carbon, and those of Ar-
rhenius (this Congress) on bottom deposits where-
in only the algal function or results of algal
functions are observed and measured. But very
few, an example being the fresh-water lake studies
of Rodhe (21) have been able to link both obser-
vational and functional studies. Marine algal
studies of this kind should be encouraged.
The benthic algae are more familiar and are
the forms thought of as the marine algae first,
even by phycologists themselves. These are the
algae normally living attached to the substratum
and commonly ranging in size and form from
half-ton kelps to solitary diatom cells. In com-
parison to the plankton algae they are very well
known taxonomically and morphologically. For
reasons of economic interest, species of some
genera, such as Chondrus, Macrocystis, Laminaria,
Gelidiwn, Eucheuma, Gratilaria, and Porphyra
are rather well known, even in regards to their
production (e.g., 28 on Gelidiuni). Their con-
tributions otherwise are conjectured but little
assayed as yet. Two roles can be expanded upon
here as conspicuous in the more central Pacific:
these are essential mineral concentration near
shores and material construction of shores and
atolls.
The mangrove areas in the Hawaiian Islands
are conspicuously clean and devoid of the char-
acteristic faunal and floral populants. In the few
years mangroves have been planted there, the well
known algal combination of Caloglossa, Bostry-
chia, and Catenella apparently has not appeared.
Surprisingly enough some of the mangrove areas
studies on Mindoro (east of Calapan), the Philip-
pines, and in the Thousand Islands (northwest
of Djakarta), Indonesia, were similarly devoid of
these macroscopic algae. But then, too, these
localities were devoid of the intertidal mud char-
acteristic of the areas heavily populated (e.g.,
Pine River shores in Queensland, Australia) by
the above algal triumverate. The theory of
causality rears its head at this place: mangroves
develop on reef flats and elsewhere where no mud
is present, but do the algae appear where there
is mud, or, do they cause the mud to be deposited ?
In the Pandan Nature Reserve in Singapore
where there is a magnificent development of
these algae on both mud and mangrove roots,
the algae were seen to be muddy, covered with
a seemingly inseparable coating of mud, while
the roots on which they were growing were,
otherwise, quite devoid of any regular deposit
of mud. In fact the roots except for the algae
were as clean as those in Hawaii or the roots of
"non-algalated" mangroves in Indonesia or the
Philippines.
In Hawaii, in Singapore, and elsewhere,
Vtntcheria forms dense turfs of erect filaments on
flat mud areas of deposit, at the edges of drainage
channels. It appears, from their mud-choked
flat nature, that these turfs act as mud-trapping
agents. The presence of this phenomenon far
outside the mangrove area (e.g., at Coos Bay,
in Oregon, U.S.A. 5) inclines one to exclude this
community from the mangrove community,
though it is strongly developed in some mangrove
areas.
Reefs and non-igneous islands as we know
them in the Pacific are of biological origin and
largely rather directly the results of algal activity.
Reefs as we are using the term are solid aggre-
gations largely of carbonate, which has usually
been deposited by Rhodophyta, Foraminifera,
Coelentcrata, or Chlorophyta. The order of
abundance in a reef is often listed in order as
these groups appear above.
This definition of reefs excludes wave cut
benches and solution benches, though it is postu-
lated here that some of their common physiog-
nomy is caused by the same effects of algal
growth. Both the benches and reefs often have
an outer raised margin (rampart or algal ridge)
and a higher algae-covered area of reef flat just
inshore of the outer margin. Wentworth (26)
attributes this form in the case of the benches to
an algal function: that of reducing erosion. On
reef edges such as found in the Tuamotu Archi-
pelago (11) in addition to building the reef margin
upwards, the algae seem to play the same pro-
tective role seaward of the channel area (back-
ridge trough). Thus a common role of the algae,
mitigating the results of erosion, is postulated
here as a major cause of this similarity in form
often seen on all three shore types.
In different places and perhaps at different
geological times, the relative importance of the
above groups of organisms giving rise to coral
has changed. As an example it may very well be
that among Chlorophyta, formerly the Dasycla-
dales were more important than now. In atolls
seaward reef edges are usually predominantly
red algal (Corallinaceae) in composition while
149
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
those facing lagoons are predominantly coelen-
terate in composition. Paleontological study of
reefs has been of academic interest until recently
when certain ore and oil associations with fossil
reefs have brought economic reasons to bear on
a better understanding of reefs.
Coral islands are common in the Western
Atlantic and elsewhere. Here this term is used to
denote islands the detrital material of which is of
biological origin but islands which are not based
on or surrounded by a currently active reef flat.
Perhaps those low islands, such as the Bahamas
in the Caribbean or the Thousand Islands in In-
donesia, are of this general sort. Often they are
largely of algal detritus.
The coral islands arc of several types including
those which are perhaps relics of the normal
reef island situation to be discussed below. John-
ston and Sand Island at what may be called here
Johnston Atoll (south of Hawaii) and some of
the smaller islands of the Hawaiian group (e.g.,
Laysan) are perhaps of this sort. The composition
is generally thought of as predominantly algal
but is often line or for other reasons hard to assay
as to origin. Some of the Bahama group and
Bermuda have been described as consisting largely
of Halimeda detritus. These are of easily recog-
nizable rather uniform fragmental composition.
Coral islands of animal skeletons, in addition
to the two algal types above, are found. The
Thousand Islands near Djakarta seem to be at
least in part of this sort. It may be that such
islands arc more frequent in the Eastern Indo-
Pacilic and, having been investigated first, gave
rise to the seemingly false idea of the dominance
of coelenterate corals in reefs and islands of bio-
logical origin. Such islands, as typified by those
near Djakarta, when small, slope gently from at
least ten meter depths (as judged by gradual shift
in water color) to low tide level and, then, termi-
nate upwards in low (one meter above high tide
line) Hat island fragments. Larger islands may
have reef flats that appear to slope somewhat
toward the island. In such cases mangroves,
instead of covering the intertidal shore or reef
flat, are at times separated from shore by a body
of water, which might be termed a lagoon. If
one terms the shallowest water at the seaward
margin of such a lagoon, a reef, it must be kept
in mind that it is different from the reefs domi-
nated by red algae in at least two respects aside
from the actual organisms present. First, such
reefs seem to be gently rounded in contour, in
their shallowest areas or slope steadily into the
sea rather than being quite plane or even with a
raised margin as in the case of algal reefs.
150
Secondly, it appears that they do not extend
upward to such a level that they are exposed to
the air as much as are the red algal reefs. It
appears that a majority of Central Philippine
shore reefs are of this latter type.
Though structurally of coelenterates, this last
type of coral island, or reef, is inseparable from
the zooxanthellae and filamentous algae which
pervade it. Though far from satisfactorily under-
stood, it appears that hermotypic corals, the large
and only corals contributing significantly to reef
and island substance, are dependent upon these
algae. It may be debated whether this dependence
is nutritional or functions more in removal of
waste products. Recent work (1, 13) showing
the loss of a large part of the photosynthate from
unicells would lead us to expect the nutrient was
available, in addition to the older arguments that
these endophytes were nutritive; e.g., hermotypic
corals develop only in the euphotic region. Un-
doubtedly the algae untilize the mineral waste
products that diffuse away from the coelenterate
cells, but it seems less likely that this is essential
to the livelihood of the animals. If the hermo-
typic coelenterates are dependent on the algae,
then in turn we can say that even the animal
dominated reef structures are quite directly related
to algal activity.
A note at this point is to be entered as to the
effects of algae on coral island (19) intertidally
and above high tide line. Their growth on almost
all the surfaces of these low oceanic islands is
a phenomenon associated with the marine en-
vironment. The gastropods in rasping them away
as food remove consolidated materials. Their
growth on, and in, coral rock must effect its
solution. These two phenomena are blue-green
algal phenomena and as the major of these
algae, Entophysalis, is also abundant on much of
the non-regularly inundated part of the island,
the general phenomenon of nitrogen fixation by
such algae should not be overlooked. On these
low oceanic islands, the usual nitrogen fixing
organisms, such as the Rhizobium of legume
roots, are essentially absent. In wetter areas,
such as Majuro in the Southern Marshall Islands,
the island surfaces, where devoid of larger vege-
tation, may become covered by Nostoc commune.
Such algae have been shown (27) to fix up to 90
per cent of the nitrogen they use even when sup-
plied with what would appear to be adequate
supplies of fixed nitrogen.
Coral reefs as found in the Central Pacific and
the islands on their reef flats seem to be main-
tained at the sea surface by algal activity. Char-
acteristically, atolls are formed. There is, as
VOLUME 4
BOTANY
often presented, a variety of reefs in relation to
igneous islands or underwater bases. The theore-
tical origins of atolls and the accumulation of
islands on atoll reefs have been elaborated on at
length by many authors. Other than to note
that these authors have concerned themselves
with the formation of "the classical atoll in full
bloom" from its beginnings and have given little
attention to what may be called their old age or
decline, we will restrict ourselves to the postulated
algal roles not discussed above.
Let us first consider in brief the distribution of
algae across an atoll. As an example let us con-
sider Raroia in the Eastern Tuamotus(7,//,/#J or
Eniwetok in the Northern Marshall Islands
(25, 9) i as the algae can be found in an investi-
gation leading in over the upwind seaward reef,
moving downwind, across the lagoon and thence
traversing the leeward atoll rim. Upon approach-
ing the atoll coming downwind, the water
becomes shallower as the bottom slopes gently
toward the surface. Rather abruptly at depths
of about six to eight meters, the hermotypic corals
become covered or arc replaced by coralline
algae. In shallow water, there is little else in
sight. In deeper water, coclenterate corals arc
scattered irregularly over the gently undulate
surface radiating away from the atoll. In shallower
water, radiating rows of individual coelenterate
coral organisms give way inshore to grooves and
spurs covered with a smooth coating of algal
coral.
Spurs extending toward the atoll, at the sea
surface, may extend upwards to such levels that
the seaward margin of the reef protrudes perhaps
a meter between waves even at high tide. The
spurs tend to be especially well developed oppo-
site islands and may be fused laterally into a
ridge, the famous algal ridge of the literature.
Grooves extending toward the atoll may run
through the algal ridge or terminate outside of it.
They may extend through the ridge covered by
the laterally fused ridge material and reef surface
inshore of the algal ridge. They may be open
channels throughout or closed over so that the
innermost open end is a blow-hole, sometimes
enclosed in a low dome, on the inner part of
the reef.
Inshore of the algal ridge, the surface is usually
plane, horizontal and at low tide level. This
surface may be as smooth as a concrete sidewalk.
Sometimes it is covered with a turf of jointed
coralline or other algae. As Wentworth sug-
gested (26) for solution and wave-cut benches the
algae here also probably play a role in lowering
erosion, as discussed above.
Still further inshore, the reef surface becomes
lower and tends to slope so that water coming
in over the algal ridge flows to the right or left
and back out over the reef margin through
grooves or the so-called (11) excurrent areas.
This area has been termed a back-ridgc-trough
and termed elsewhere a channel area. This area
appears to be one where corrosion and corrasion
are dominant over deposition. Indeed from the
algal pavement of the outer reef flat to the lagoon,
the physical features appear to be largely regulated
by mechanical forces though the materials may
be predominantly of algal origin, and these latter-
may in turn regulate the mechanical forces.
The sea reef, phycologically, can be divided
into four major zones horizontally in so far as
the populations are concerned: (a) Porolithon
onkodes dominates the surfaces from depths of
six to eight meters up onto the outer reef flat;
(b) the outer and higher reef flats which may be
covered by an algal turf; (c) the channel or reef-
pool area; and (d) the shore intertidal and supra-
tidal areas. In designating these we purposely
avoid terms used in discussions of vertical dis-
tribution.
Porolithon onkodes above its maximum depth
comes quickly to cover almost all of the surface.
It produces the algal ridge as a scries of overlaying
overlapping pustules of several centimeters to
a few decimeters in lateral extent and from a few
millimeters to a centimeter or two thick. Within
the cavities resulting reside a host of animal sorts,
of which annelids, fishes, crabs, and foraminifera
are conspicuous. On the surface patches of small
algae, e.g. Centroceras, Microdictyon, and blue-
greens, are scattered about as are such animals
as limpets, barnacles, and Cypraeids. Poroli-
thon onkodes is a most peculiar coralline alga in
that it will live on the most brilliantly illumi-
nated shore areas and stand some exposure to
air. This is in contrast to other genera of crustose
coralline algae such as Lithothamnion^ which
never grows extensively under such circumstances.
The algal turf on higher parts of the reef flat
occurs in at least two major forms: as relatively
pure stands of jointed corallines on outer reef
flats (7) underlain by Porolithon onkodes ; and as
mixed stands, on inner reef flats (24), of Jan fa.
Amphiroa, and relatively small fleshy algae such
as Caulerpa ambigua, Polysiphonia, Herposiphonia
1 Perhaps the confusion regarding Lithothamnion and the algal ridge arises from the fact that Porolithon onkodes was
originally described by Heydrich in the genus Lithothamnion.
151
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
secunda, Laurencia, and Gelidium intermixed often
with large proportions of blue-green algae. Larger
fleshy algae often occur in patches or isolated
clumps or sandwiched in crevices or the holes
opened by boring animals, i.e., where freed from
the browsing of fishes. Two functions must be
considered for this turf: first, in providing a direct
or indirect source offish food; second, in reducing
erosion of the reef flat.
The channel or pool area is often sandy or
floored by shallow pot holes. Algae are few
except under the edges of reef boulders, e.g.,
Diciyosphaeria. Often the surface is pinkish
from what seems to be incipient growths of
PoroUthon onkodcs. Larger algae, such as Liagora,
Tur binaries and Padina sometimes occur. The
aspect is that of a shallow but open tidepool.
At one time (February), the reef pool area of the
eastern end of Ujcland in the Marshalls was
strongly pink from the air. Investigation on the
ground revealed this to be due to an extensive
growth of Liagora.
The intertidal and supratidal areas are usually
covered by, below, a brown and, above, sun-
blackened coatings of Entophysalis. As one pro-
gresses away from shore and salt, Anacystis and
finally Scytonema or, if very wet, Nostoc similarly
tends to cover stable surfaces. The roles of these
algae in nitrogen fixation and shore processes are
taken up above.
Upwind lagoon shores are usually sloping sand
beaches. There is little of macroscopic biological
nature on such sandy beaches. When not sandy
beaches, they are representative of shore types
elsewhere in the lagoon. The color of the water
changes as one moves out into the lagoon from
the shore. As seen from the air, the color is that
of the sand beneath, usually a very light buff
color, darkening until depths of six to eight feet
arc reached when the color is definitely green
tinged. The water appears greenish and gradually
darkening until at perhaps depths of twenty feet
when it can be said to be blue. When shallower
places in the lagoon are approached, a reverse
color sequence shows.
The most shallow places in lagoons are the
brown, sometimes pink-edged or light buff-
centered reef patches, reaching low tide level.
Chunks of rock protruding above low tide level
are often blackened with blue-green algal growths.
If islands are present, they are likely to be similar
to the sandier reef islands. Reef patches have
various forms (4) which may be arranged in an
evolutionary sequence (ll)\ either in the lagoon
changing in relation to the bottom, or just growing
152
upwards, or in relation to changes in the shor
location with time.
A typical reef patch reaching the low tide sui
face of the lagoon is coated on its sides wit
Caulerpa and, especially on less precipitous area
and near the base, with Halimeda. The Caulerp
is reputed by the Tuamotuan natives to be a mos
important turtle food. Small fleshy algae occui
especially on the uppermost six to eight feet c
the rocky column, and serve as fish food. Th
classical relationship between the coelenterate
and the algae perhaps enables the hermotypi
corals to grow and play their role in the produc
tion of the reef patches. The brown coloratio
of the coelenterate corals on top of the reef pate
is probably largely due to the endozoic algs
pigments. Occasionally the margins of rcc
patches will be pink from growths of PoroUthon
especially on their upwind sides, or they ma
bear a fur of Rhodomelaceous algae. The center
of reef patches are often depressed and san<
floored with scattered coral fragments. Ther
above the sand such algae as Udotea and th
turf-forming species of shore areas are oftei
abundant.
Diving and careful dredging in lagoons hav
contributed (9J4) considerably to our knowledg
of the algae on lagoon bottoms. With the ex
ception of Halimeda, little in the way of corallini
or calcium-depositing algae is found. Contrar
to popular opinion, Halimeda has not been foun<
( 11 J 4) to occur in dense meadows on the bottom
Instead, though lagoon bottom sediments ar<
often predominantly of Halimeda segments, th
living thalli are almost always clustered abou
the talus bases or occur on the sides of ree
patches. Despite this, Halimeda is perhaps th<
agent most responsible for the filling of lagoons.
Reefs are usually found on the downwinc
lagoon shores. These have been termed f/7
shore-bound reef patches for both their anatonr
and components are very similar to what is founc
on the lagoon reef patches. The roles of the alga<
seem to be the same as on lagoon reef patches.
Islands on the downwind reefs are, phycologi
cally, very similar to those on windward reefs
The islands which occasionally occur on ree
patches are also phycologically similar.
Seemingly downwind or lea sea reefs are phy
cologically like those of the windward shores anc
the algal roles are the same. In some cases, e.g.
at Arno Atoll in the Southern Marshall Islands
the downwind reef edge may have many mon
coelenterate corals on it than the sea reef edges
VOLUME 4
BOTANY
elsewhere, and also the algal ridges are often less
conspicuous.
As described above, the atoll system seems to
persist in equilibrium with the sea as long as
changes in the earth's crust are slow. If the crust
shifts upward, the reef can be eroded back toward
sea level; or if the rate of elevation is sufficiently
fast, a raised atoll can result. On the other hand,
should the atoll sink, upward growth of the
organisms would keep the atoll in contact with
the sea's surface, unless the sinking rate were
so fast the organisms could not do so. Apparent-
ly a sinking rate of over one or two centimeters
per year would result in a sunken atoll, at least
that is the current estimate for the maximum rate
of atoll upgrowth. Examples of atolls where
such phenomena have taken place arc numerous.
The atoll around Johnston Island, south of
Hawaii, seems to have sunken on only one side;
so a long crescent rim is left at the sea's surface.
The atoll of Anaa, east of Tahiti, is an example of
an atoll where one edge has been raised, in this
case about five meters, while the rest is at normal
atoll elevations in respect to the sea surface.
Prospective algal roles in both the erosion and up-
building have been taken up above.
Finally the benthic algae seem to play a most
important role in the concentration of the che-
mical elements essential to life processes. As a
result of this process, as one nears shores the
biological standing crops as well as the productiv-
ity rates increase enormously. Let us consider
the events that would lead to this end around a
newly created igneous island such as a volcano
suddenly rising above the sea's surface. Essen-
tially this condition prevails in Hawaii when new
lava flows reach the sea. From a study there of
lava surfaces appearing in the sea in 1955 we know
it is the algae which appear first and before the
animal populations characteristic of older shores
in the same general area.
In the open ocean we would assume that when
a new igneous mass appeared there would at first
be no concentrations around its shores. Leached
materials from the newly cooled lava would be
diluted and washed away in the passing ocean
stream. The first macroscopic organisms to
appear would be expected to be such as Enter -o-
morpha, Ectocarpus, and Polysiphonia, as they are
in Hawaii, if the new island were in similar waters.
These pioneers appear in that order of importance
and time in the Central Pacific. Other populants
follow. The benthic standing crop increases as
the algae convert inorganic materials from the
water into organic materials or deposit them as
inorganic compounds such as calcium carbonate.
This must apply as well to phosphorus and
nitrogen and carbon and the other elements
essential to protoplasm. Because these popula-
tions are benthic forms they do not wash away.
Following the algae, there appear the animals
which feed upon the algae: the littorines, limpets,
and blennies. A certain amount of the increase
in standing crop shifts over to these herbivorous
forms and then, theoretically, to carnivores. The
detritus and fecal material are undoubtedly de-
graded to a mineral state, and thus the water
near shore becomes enriched in inorganic nu-
trients. The observed near-shore increases in
plankton must be the results of this enrichment.
It is observed that the calcareous organisms
appear conspicuously a year or more after the
non-calcareous algae arc well established. With
the very long-term evolution of an atoll, the
binding of materials becomes predominantly an
activity of the calcareous forms in so far as mass
is concerned. This implies a shift to such herbi-
vores as the parrot fish, some species of which
live on calcareous algae. The detritus and fecal
material they produce contribute to the sediments
of the lagoon or island shore as well as to the
enrichment of the waters about the high island
or atoll.
It is expected that the supply of fertilizer
elements and the standing crop of organic material
would increase until a new steady state is achieved.
Previous to this time, i.e., while the standing
crops are increasing, one might expect the water
approaching the young atoll to contain more of
the fertilizer elements than the water leaving the
atoll. This is without taking into account sub-
stances from a high island around which the atoll
might be developing. In later stages, the amount
would be expected to be equal. Perhaps in old
age the amount leaving (at least leaving the
euphotic zone) would exceed the amounts arriving
and being bound by the primary productivity of
the algae. It may be that plankton algal pro-
ductivity near shores, as well as the size of the
standing crops of phytoplankton, may serve as
an index of the relative age of an atoll.
SUMMARY
The phytoplankton algae are described as
having major roles affecting fertilizer concentra-
tions or in primary productivity resulting from
variations in fertilizer salt concentrations found
(a) in transition or shear zones enriched by ver-
tical mixing or eddies, (b) in divergent current
153
PROCEKDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
areas by upwelling, (c) in wind disturbed areas
by vertical mixing, and (d) near land by fertilizer
increases arising both through concentration
by benthic organisms and through addition by
fresh-water. The benthic algae are described as
having major roles in primary productivity of
food, in the fertility of the soil, and in construction
and destruction of shores, islands, and reefs,
particularly those of atolls.
REFERENCES
(1) Allen, M.B., 1956, Excretion of Organic
Compounds by Chlamydomonas, Arkiv
F. Mikrobiol., 24: 163-168.
(2) Arrhenius, Gustav, (Mss.) Climatic Records
on the Ocean Floor. (Paper presented
to symposium on marine upwelling at
9th Pacific Science Congress, Bangkok,
Thailand.)
(3) Bernard, F., (Mss.) Donnecs R6centes sur la
Fertilite Elementaire en Mediterrance.
(Paper presented to the 1957 I.E.S.
annual meeting in Bergen, Norway.)
(4) Denielsson, Bengt., 1954, Native Topograph-
ical Terms in Raroia, Tuamotus, Atoll
Research Bulletin, 32: 92-96, 3 figures.
(5) Doty, Maxwell, S., 1947, The Marine Algae
of Oregon, I: Farlowiu, 3: 1-65.
(6) , 1 954a,* Current Status of Carbon-
fourteen Method of Assaying Productiv-
ity of the Ocean (As of April, 1954),
9 pp., 6 appendices, University of Hawaii.
(7) , 1954b, Floristics and Plant Ecol-
ogy of Raroia Atoll, Tuamotus, Atoll
Research Bulletin, 33: 1-41, 1 figure.
(8) , 1955,* Current Status of Carbon-
fourteen Method of Assaying Productiv-
ity of the Ocean (As of February,
1955), 52 pp., 4 appendices, University
of Hawaii.
(9) , 1956,* Current Status of Carbon-
fourtecn Method of Assaying Productiv-
ity of the Ocean (As of April, 1956),
51 pp., 6 appendices, University of
Hawaii.
(10) , 1957,* Current Status of Carbon-
fourteen Method of Assaying Productiv-
ity of the Ocean (As of July, 1957),
41 pp., 2 appendices, University of
Hawaii.
(11) Doty, Maxwell S., and Morrison, J.P.E.,
1954, Interrelationships of the Organisms
on Raroia aside from Man, Atoll Re-
search Bulletin, 35: 1-61, 8 figures.
(12) Doty, M.S., and Oguri, M., 1956, The Island
Mass Effect, Journal du Conseil pour
V Exploration de la Mer, 22: (1) 33-37.
(13) Fogg, G.E., (Mss.) Extracellular Products of
Phytoplankton and the Estimation of
Primary Production. (Paper presented
to the 1957 I.C.E.S. annual meeting in
Bergen, Norway.)
(14) Gilmartin, MalvernJr., 1956, The Ecological
Distribution of Certain Central Pacific
Atoll Benthic Algae, iii and 78 pp., 8
ligures, and a map. M.S. Thesis, Univer-
sity of Hawaii, Honolulu (Un-
published).
(15) Guillard, R.R.L., Doty, M.S., and Oguri,
M., 1955, The Productivity of Waters
North of Hawaii as Determined by
Carbon-fourteen Uptake. Abstract from
the Proceedings of the Hawaiian Acade-
my of Science, 1955-1956, p. 10.
(16) King, J.E., Austin, T.S., and Doty, M.S.,
1957, Preliminary Report on Expedition
"Eastropic". U.S. Fish and Wildlife
Service, Special Scientific Report
Fisheries 201, 155 pp.
(17) Lohmann, H., 1908, Untersuchungen zur
Feststellung des vollstandigen Gehaltes
des Meeres an Plankton, Komm. z.
Wiss. Unters. D. Deutch. Meere in Kiel
und d. Biol. Anst. Helgoland. Wiss.
Meeresunters, N.F. Abt. Kiel. 10:
131-370.
(18) Newell, N.D., 1954, Reefs and Sedimentary
Processes of Raroia, Atoll Research
Bulletin, 36: 1-35.
(19) Newhouse, Jan., 1954, Ecological and Flor-
istic Notes on the Myxophyta of Raroia,
Atoll Research Bulletin, 33:42-54, 2
figures.
(20) Riley, Gordon, 1953, Letter to the Editor.
Journal du Conseil pour r Exploration
dela Mer, 19(1): 85-89.
(21) Rodhe, Wilhelm, (Mss.) The Primary Pro-
duction in Lakes: Some Results and
Restriction of the C 14 Method.
* These are annual reports to the U.S. Atomic Energy Commission^ Division of Biology and Medicine, of work accom-
plished under contract AT-(04-3)-15 between the U.S. Atomic Energy Commission and the University of Hawaii.
They have not been published and are not generally available but are listed here for convenience in completing the list
of information sources.
154
VOLUME 4
BOTANY
(22) Steemann-Nielsen, E., 1952, The Use of
Radio-active Carbon (C 14 ) for Measur-
ing Organic Production in the Sea,
Journal du Council pour V Exploration de
la Mer, 18(2): 1 17-140, 7 figures.
(23) Steemann-Nielsen, E., and Aabye-Jensen,
E., 1957, Primary Oceanic Production.
The autotrophic production of organic
matter in the oceans, Galathea Report,
1:49-136, 41 figures.
(24) Taylor, W.R., 1950, Plants of Bikini and
other Northern Marshall Islands. Uni-
versity of Michigan Studies, Scientific
Series, 18, 15 and 227 pp. Plates 1-79
and frontispieces.
(25) Tracey, J.l.Jr., Ladd, H.S., and Hoflrneistcr,
J.E., 1948, Reefs of Bikini, Marshall
Islands, Journal of the Geological So-
ciety of America, 59:861-878, Plates
1-1 1 and 8 figures.
(26) Wentworth, Chester, 1939, Marine Bench-
forming Processes, II : Solution benching.
Journal of Geomorphologv, 2: 3-25, 12
figures.
(27) Williams, A.L:., and Burris, R.H., 1952, Ni-
trogen Fixation by Blue-green Algae and
Their Nitrogenous Composition, Ameri-
can Journal of Botany, 39: 340-342.
(28) Yanegawa, T., and Tanh, K., 1957, Studies
on Agar-agar in Japan, Proc. Eighth Pac.
Sci. Cong. (Philippines), 3: 215-223.
DISCUSSION
R.C. MURPHY: Will the mere shallowing of water, as
an ocean current approaches an island, cause a concen-
tration of plankton life, and therefore increase the food
resources for fish and birds?
M.S. DOTY: In Hawaii, in respect to the circulation in
the North Pacific, there is no upwelling along the islands.
G.L. CLARKE: As the contained life is an integral part
of the water volume, no such vertical concentration of
plankton can occur. I attribute the acknowledged enrich-
ment of life in the coastal waters to more favorable con-
ditions for nutriment and reproduction. If the animals
are brought by currents, they will also be carried away by
currents that pass around the island unless some mecha-
nism to concentrate them exists. This might be accom-
plished by the animal's own swimming reactions.
G.F. PAPENFUSS: It is known that in temperate seas the
larger algae are very important in providing protection,
or a home as it were, for many kinds of animals and thus
contribute directly to the concentration of life in shallow
waters. Is this also true in tropical regions where the algae
are much smaller?
M.S. DOTY: This is true also in the tropics and parti-
cularly among the coralline algae which provide many
niches and serve for protection of small animals.
H.M. HURKHILL: What species of the mangrove com-
plex has been introduced to Hawaii for purposes of land
formation ?
M.S. DOTY: It is only the Rhizophora spp.
H.M. DURKIUIL: The primary pioneer mangrove in
Malaya is considered to be Avicennia spp., but in the man-
grove areas of Singapore a Dictysta sp. looks as if it takes
an active initiation in the binding of mud found in the
intertidal zone and fiats below the mangrove limit. With-
in the tidal prawn ponds of the mangrove belt, Vaucheria
appears to be an important mud-binder which may criti-
cally limit the commercial life of the ponds.
G.F. PAPLNFUSS: Can it be that the gradual sloping
of the reef on the windward side of an atoll is owing to the
scouring action of the waves on that side as compared with
the conditions on the leeward side?
F.R. FOSBERG: It should be pointed out that the erosion
on the slopes of atoll reefs is of two sorts - that by ordinary
waves and that by typhoon or storm waves. The latter
may be much more important ; they come from the south,
predominantly, and produce different effects than ordinary
waves. The function of the other encrusting organisms is
to cement the relatively fragile growth-lattice of the reef
into a rock capable of resisting ordinary wind waves.
155
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
COMPARISON OF THE CALCAREOUS ALGAL FLORAS OF
RECENT AND FOSSIL REEFS
J. HARLAN JOHNSON
Department oj Geology ', Colorado School of Mines, Golden, Colorado, U.S.A.
INTRODUCTION
Reefs ranging in age from the Eocene to the
Recent are known from a number of islands of
the Western Pacific, and they have been encoun-
tered in borings made in several of the islands.
It has been the author's good fortune to have
spent much of the last ten years studying the fossil
algae from these reefs. The fossils consist only
of calcareous algae: crustose corallines, articu-
lated corallines, Halimcda, and a few Dasycla-
daceae. The non-calcareous forms have not been
preserved, so any comparisons between Recent
and ancient floras will have to be limited to cal-
careous algae.
Unfortunately Recent calcareous algae have
received relatively little study. They are probably
the least known of any of the algal groups.
Possibly this is because they are more difficult to
collect, being commonly firmly attached, and
because of the necessity of decalcification, section-
ing, and study with a microscope.
THE RECENT ALGAL FLORA
The Recent Calcareous Algae found in the
Tropical Pacific appears to form a surprisingly
uniform widespread flora consisting of a relative-
ly small number of genera and species. Common-
ly at a given locality there may be great numbers of
individuals of a few species. The magnificent
algal ridge at Bikini has been built largely by two
species: Porolithon gardineri and Porolithon
onkodes, while the great Lithothamnion bank of
Haingsisi near the Southwest point of Timor,
so vividly described by Weber Van Bosse (77,
pp. 4-5), is composed largely of rounded masses of
Lithothamnium eruhescens. Most of the common
species are surprisingly widespread, extending
from the East Indies over the tropical Pacific
and westward across the Indian Ocean. Several
species extend from the Red Sea to the Marshall
Islands and even to Hawaii. Dawson records
some species from both the coasts of Vietnam
and western Mexico.
Very few comprehensive listings of the species
found in a given area are known to the writer.
156
However, Table 1 will give an idea of the general
complexion of the Recent Flora.
FOSSIL FLORAS
The oldest reefs of the Western Pacific for which
we have much information are of Eocene age.
The author has made detailed studies of large
collections of Eocene material from Saipan (10)
and the Eniwetok cores ( 11), and is now studying
a large collection from Guam. He has also stud-
ied small collections from the East Indies and
from Ishigaki in the southern Ryukyu Islands.
Ishijima has described Eocene species from
Taiwan, and adjoining areas.
The Eocene flora is another relatively small
homogeneous flora with a wide distribution. A
number of species, or very closely related forms,
have been found from the western Mediterranean
to the Mariannas and even to the Marshall Is-
lands. Data showing the general composition
of the Eocene flora of crustose coralline algae is
shown on Chart 2.
The known Miocene flora is considerably
larger than the Eocene flora, with a greater
representation of species belonging to the genera
Lithophyllum and Mesophylluw, fewer species of
Archaeolithothamnium, and about the same num-
ber of Lithothamnium. This flora also has a
number of links with the Western Mediterranean
and even some with the West Indies.
The Pleistocene Flora is not well known.
Much more collecting and study is needed.
However, it appears to be essentially the same as
the Recent. The only difference noted so far is
that in some localities it may contain more
species than the Recent. This may merely mean
incomplete knowledge of the Recent. Most of
the Pleaistocene species are still found growing
along the present coastlines. The Pleistocene
flora is characterized by the presence of large
quantities of Porolithon, Goniolithon, and Amphi-
roa. In some regions (Palau, Saipan, Bikini, and
Eniwetok) there is also an abundance of Halimeda
debris. The Lithothamnium and Lithophyllum
present usually belong to thin encrusting species.
Archaeolithothamnium occurs, but it is represented
by only a few rarely found species.
VOLUME 4
BOTANY
Too little is known about the Pliocene Flora
to merit a discussion here. Charts showing
details of the composition and distribution of
these floras will be published in the author's
forthcoming report on the fossil algae obtained
from the deep drillings at Eniwetok, Funafuti,
and Kita-Daito-Jima (11), which will probably
appear during 1958.
REEFS OLDER THAN TERTIARY
No reefs older than Eocene arc known in the
Western Pacific region. Reef building corals and
rudistids have been dredged from the tops of
Guyots in the areas around the Marshall Islands
(1), but no algae were described. Permian reef
building algae have been found in Timor and
Table 1.
Composition, Recent Floras of Calcareous Algae.
The numbers indicate the number
of species of each genus.
Genus
Porolithon
Goniolithon
Lithophyllum
Lithothamnium
Archaeolithothamniuni
FoslleUu
Lithoporellu
Heteroderma
Jan ici
Ainplnroa
Cora/Una
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Halimeda
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157
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Japan. However, it is necessary to go to distant
continents to learn the character of the algal
floras of the Mesozoic and Paleozoic Reefs. The
Mesozoic reef floras are almost unknown. The
Paleozoic floras have an entirely different com-
plexion consisting largely of calcareous green
algae. Calcareous red algae appear to have been
rare before the Jurassic. During that period they
were represented mainly by members of the ex-
tinct family of the Solenoporaceae. The Coralline
Table 2.
Geographical Distribution of Eocene Coralline Algae found in the Western Pacific.
Genus and Species
Archaeolithothamnium
A. cf. A. chamorrosum Johnson
A. dalloni Lemoinc
A. cf. A. hemchamlri Rao
A. numniuliticum (Giimbcl) Rothplet/
A. oulianovi Pfcnder
A. a IT. A. saipancnsum Johnson
A. cf. A. sociahile Lemoinc
LAthophyUum
L. cf. lingusticum Airoldi
Lithotluuunium
L. cf. L. uhraidi Lemoine
L. crisputhallum Johnson
L. kumhecrustwu Johnson
L. cf. nwreti Lemoine
L. tcipachawn Johnson
Mesophyllwn
M. robustus Johnson
M. vaughanii (Howe) Lemoine
Corallina
C. prisca Johnson
Eniwetok |
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X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
158
VOLUME 4
algae (Family Corallinaceae) are not common
before the beginning of the Tertiary. Most of the
genera of today extend back into the Eocene,
only a few into the Cretaceous. No represen-
BOTANY
tatives of Recent genera have yet been found in
rocks older than middle Jurassic. The known
geologic range of the common genera of the
Corallinaceae is shown in Table 3.
Table 3.
Known Geologic Range of Common Recent Genera of Coralline Algae.
Porolithon
Goniolithon
Lithophyllwn
Mesophyllwn
LithothumniwH
Archaeolithothamnium
Fosliella
Lithoporellti
Jania
Amphiroa
Coralline!
REFERENCES
(1) Barton, E.S., 1901, The Genus Halimeda:
Siboga Expedition Monograph, 60: 1-32,
pbs. 1-4, Leyden, Holland, Brill.
(2) Dawson, E. Yale, 1954, Marine Plants in
the Vicinity of Nha Trang, Viet Nam,
Pacific Science, 8:373-469, 63 figs.
(3) , 1956, Some Marine Algae of
the Southern Marshall Islands, Pacific
Science, 10:25-66, 66 figs.
(4) , 1957, An Annotated List of Ma-
*J
c
v
u
0>
CCS
Pleistocene
Pliocene
t*
8.
Q-
5
0>
c
V
u
.2
Ui
4>
|
4>
C
V
o
.2
Oligocene
Eocene
Cretaceous
Jurassic
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
?
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
?
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
?
X
X
X
X
X
X
X
X
rine Algae from Eniwetok Atoll, Pacific-
Science ; 11:92-132, 31 figs.
(5) Foslie, M., 1900, Calcareous Algae from
Funafuti, Del. Kgl. Norske Via 1 . Selsk.,
Skrifter, 1900, 1:3-12.
(6) Foslie, 'M.H. and Printz, H., 1929, Contri-
butions to a Monograph of the Litho-
thamnia, Det. Kongl. Norske Vid. Selsk,
Museet, Monograph, 60 pp. 75 pis.
(7) Hamilton, E.L., 1956, Sunken Islands of the
Mid-Pacific Mountains, Geol. Soc.
America Mem., 64, 97 pp., 12 figs., 13 pis.
159
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
(8) Ishijima, W., 1954, Cenozoic Algae from
the Western Pacific, Tokyo ^ Yuhodo
Company, 87 pp., 69 pis.
(9) Johnson, J. Harlan, 1954, Fossil Calcareous
Algae from Bikini Atoll, U.S. Ceol.
Survev Prof. Paper 260-M, P. 537-545,
pis. 188-197.
(10) _, 1957, Fossil Calcareous Algae
from Saipan, U.S. Geol. Survey Prof.
Paper 280, Part E, pi. 25 (in press).
(11) _., 1958, Fossil Calcareous Algae
from the Eniwetok, Funafuti, and Kita-
Daito-Jima Drill Holes, U.S. Ceo*.
Survey Prof. Paper 260-W (in press).
(12) Johnson, J. Harlan and Ferris, B.J., 1949,
Tertiary Coralline Algae from the
Dutch East Indies, Jour. Paleontology,
23, (2): 193-198, pis. 37-39, (March).
(13) Lemoine, Mme. P., 1939, Lcs Algucs Cal-
caires Fossiles de L'Algerie, Mat.
Carte Geol. de L'Algerie Series, 9,
128 pp., 3 pis., 80 figs.
(14) Lignac-Grutterink, L.H., 1943, Some Ter-
tiary Corallinaceae of the Malaysian
Archipelago: Verk. Geologisch-Mijn-
bouw Kundig Genootschap veer Neder-
land en Kolonien, Geol. Ser., Deel, 13:
283-297, 2 pis., (Dec.).
(15) Sripada, Rao K., 1943, Fossil Algae from
Assam: 1 The Corallinaceae, Proc. Nat.
UAcad. Sciences, India, 13, part 5:265-
299.
(16) Taylor, W.R., 1950, Plants of Bikini and
other Northern Marshall Islands: Uni-
versity Michigan Press, 227 pp., 79 pis.
(17) Weber van Bosse, A., and Foslie, M.H.,
1904, The Corallinaceae of the Siboga
Expedition: Siboga, Exp. Monograph
LXI, 1 10 pp., 16 pis., Leyden, Holland.
DISCUSS ION
M.S. DOTY: Can Halimcda be recognized in fossil depo-
sits?
j.i i. JOHNSON: Halimcda can be recognized to genus
quite readily, but it is not as easy to compare with species
because parts used by the taxonomist dealing with modern
species may not be preserve in the fossil record. They are
quite abundant and contribute a great deal to the shallow
water deposits back as far as the Cictaceous.
j.v. SANTOS: Which plays the greater role in the forma-
tion of an atoll, the coelentcrate corals or the coralline
algae?
J.H. JOHNSON: It may vary to some extent, but in
general the formation of the atoll is made up basically of
coral and then the coralline algae come in, with diatoms
and foraminifera finally filling in the spaces.
160
VOLUME 4 BOTANY
QUALITATIVE DESCRIPTION OF THE CORAL ATOLL ECOSYSTEM
F.R. FOSBERG
Pacific Vegetation Project, Care of National Research Council, Washington* D.C., U.S.A.
A coral atoll may be described, in the briefest
terms, as a cap of limestone of organic origin on
a mountain on the floor of the ocean, rising to or
only slightly above sea level. Some lie on shallow
banks or continental shelves. The cap is a ring-
like ridge or reef surrounding a body of water
termed a lagoon. Some parts of this reef may
emerge above high tide level as islets. These may
either be remnants of former higher reef levels or
detrital accumulations. Much or all of the reef
surface below mean low-tide level and down to
depths where sun-light penetration is very atten-
uated is composed of communities of living
plants and animals. In bulk, at least, these are
mostly organisms that secrete limy skeletons.
Accumulations of these skeletons make up,
almost exclusively, the reefs and upper parts of the
mountain down to the volcanic (or other) base-
ment rock on which the reefs originally started
to grow. The depth of this limestone is known
for only a few atolls and may vary from at least
1 ,400 meters to, probably, very much less.
The concept of the ecosystem, first proposed by
Tansley (9), is that of an interacting system com-
posed of an environment and all of the organisms
involved with it. It is normally an open system
because there is a continuous, though variable,
influx and loss of energy and material. Such a
system is, of course, an abstraction constructed
to facilitate understanding of the complex pro-
cesses involved in a segment or class of segments
of the biosphere. As such, its extent is limited
only by selection and definition of the segment
or segments under study. Thus it may be varied,
in different examples, from the smallest observa-
ble unit of environment in which organisms live
to the entire world's biosphere as a whole with its
total environment (4). As the ecosystem is
limited only by the extent of the effective environ-
ment, the maximum could be, theoretically, the
entire universe. Practically, however, the defini-
tion will not ordinarily extend to the ultimate
sources of energy, or even of material. It will be
restricted to such extent as will best facilitate
observation and understanding of the portion
of nature under immediate study. This concept,
of obvious and increasing utility but not too easy
to handle, and never, apparently, used by its
creator, has been, in recent years, adopted by a
number of ecologists (e.g., 6, /, 5, 4, 2, 3). No
two have defined or formulated their ecosystems
in exactly similar terms, nor is there any critical
need, at this stage, to do so.
In this paper, the coral atoll ecosystem will be
described in terms of processes involving transfer
or transformation of energy and material, with
only incidental reference to the actual organisms
involved in the system or to the physical struc-
tures found in the environment. It is recognized
that in a complete account of such a system these
aspects, also, would be described fully. For
present purposes it may suffice to say that the
biotic component of the system is composed of
phytoplankton and zooplankton; free-living but
bottom-dwelling animals and other hctcrotrophic
organisms of many sorts; an enormous aggrega-
tion of sessile or fixed organisms representing
most classes of algae, a few seed plants, and prac-
tically all phyla of invertebrate animals; and a
diverse assortment of land animals and plants.
Many of the marine organisms secrete skeletons
of CaCO 3 which arc added to the material of the
substratum. This process forms a lattice-work
of limestone in which free skeletons or loose
fragments lodge. By several processes these
may become bonded in such a way as to form a
rather hard and resistant rock. This is built up in
the form of a ridge or reef usually enclosing a
shallow body of water or lagoon and rising
variously to somewhat below, near, or just above
high-tide level. This reef is ordinarily rather
flat-topped, of varying width, with irregularities
or islets extending above high-tide level. Waves
commonly break on the outer margin and water
flows from the sea to the lagoon and back to the
sea over the flat or gently sloping reef surface or
through gaps in it. The flow may be in and out
with the tides, or in over the windward and out
over the leeward sides. The islets are commonly
composed in part or wholly of loose porous
limestone debris and are mostly covered by
vegetation that includes representatives of all
major groups of land plants. The land faunas
are made up of a large number of species of
insects and other arthropods, worms, land
molluscs, a few reptiles, some birds, and a few
mammals, including rats and man. Larger islets
may contain within their porous structures a body
161
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
of fresh ground water floating on the underlying
salt water and retarded by friction from free
diffusion with it.
These atoll structures are found in most regions
of the tropical and, more rarely, subtropical seas.
Temperatures range generally between 75 and
85F or in full sun on land, higher, decreasing
of course with increasing depth in the sea.
General climates range from relatively dry,
perhaps 600 mm precipitation, to wet, 5,000 mm
or more. The atolls are in trade wind, doldrum,
and monsoon belts. Insolation is generally high,
cloudiness low to moderate.
Most atolls are inhabited by human beings,
some by relatively large populations. These
exert a generally appreciable, often profound,
influence on the functioning of the ecosystem.
Of specific importance in this connection are the
economic activities of planting coconuts, harvest-
ing, drying, and exporting coconut meat, and
importing in exchange various foods and other
materials.
With this description of the general physical
and biological situation, we may proceed to
describe in more formal terms, the abstraction
called the coral atoll ecosystem. This may be
outlined as follows in 12 sections, lettered A to L.
A. Media.
B. Nourishment or inflow.
C. Production.
D. Transformation.
E. Decomposition.
F. Excretion.
G. Accumulation.
H. Turnover.
I. Miscellaneous other elTects and processes.
J. Losses.
K. Balance.
L. Trends.
A. Media: The media in which the system
exists are two a layer of sea water and a super-
imposed layer of air. These, by the nature of
the earth-system itself, are constantly changed by
sea and air circulation. Through them or by
means of them, all exchange, gain and loss, of
matter and energy takes place. These two media
are the most universal and pervasive components
of the ecosystem and at the same time its environ-
ment, influencing in some measure everything
in the system.
B. Nourishment or inflow: As the atoll is an
open system, there is a continuous addition of
matter and energy in many forms. Fundamental,
of course, is the daily increment of solar energy
162
without which the system, in anything like its
present form, could not exist. Its functioning is
in almost every respect dependent on either photo-
synthetic or thermodynamic processes, which are
dependent on constant addition of energy from
the sun. The nourishment of organisms and the
circulation of both air and water are important
examples.
The energy of wind, mostly indirectly a form of
solar energy, also exerts its force in various ways
in the system. Most of this energy is received
elsewhere and transported to the atoll.
The gravitational energy of both sun and moon
contributes importantly to sea-water circulation
in the form of tides. The movement of ground
water in atoll islets also is influenced by tidal
fluctuation.
Essential components of the media, such as
O 2 , N 2 , H 2 O, CO 2 , as well as dissolved salts and
suspended organic matter, and even living pro-
pagules and disseminules of organisms are con-
tinually renewed or carried into the system by air
and ocean currents. Relative concentrations of
the various components of the media are main-
tained at a rather constant level in this manner.
The replenishment of the ground-water bodies in
islets is dependent on incoming fresh water,
mostly evaporated elsewhere and deposited as
rain on the islets from wind-borne clouds.
Surface currents, upwellings, tradewinds, mon-
soons, and cyclonic storms are important aspects
of the circulation patterns involved. The intro-
duction into the system of phosphorus, on which
organic activity is completely dependent, is
believed to be controlled to a considerable extent
by upwellings of deep-sea water. One important
route of transport of phosphorus from areas of
upwelling to the atolls is by means of fish-eating
birds and their young which deposit phosphates
in their excreta within the area of the system.
Essential mineral elements, nitrogen, and organic
carbon are also brought in by the birds at the same
time. Organic matter, in the form of drifting
plankton, driftwood, and dead organisms, is
brought to the atolls by currents. Currents also
bring small amounts of mineral elements in the
form of pumice as well as in solution. Volcanic
ash arrives by way of winds, especially high alti-
tude winds.
Finally, with changing patterns of human
activity on atolls, increasing amounts of imported
foods and other materials as well as alien organ-
isms are introduced into the system. These
introductions are effecting various rather pro-
found changes in the equilibria and altering
greatly the physical appearance of the atolls.
VOLUME 4
BOTANY
C. Production: The elaboration by plants of
basic organic materials from elements and simple
inorganic compounds is termed production, in
an ecological sense. Such elaboration provides
the fuel and building materials for all other life
processes. The effective capacity of a system for
production is called its productivity. (1) The
most obvious productive process is photo-
synthesis, by which carbohydrates arc elaborated.
Algae and green plants utilize CO 2 , H 2 O, and
energy from sunlight for this purpose. Such
algae occur as components of plankton, within
the cells of corals and other coelenterates, fixed
on the reef surfaces, terrestrially on soil and rocks,
and epiphytically on tree trunks. Mosses are
found in many terrestrial situations on the islets,
as well as on tree trunks, especially where they
are shaded. Ferns and psilopsids are common
growing on land and epiphytic on trees. Seed
plants grow principally on land, but some are
epiphytic and a few are marine, growing in shal-
low lagoon situations on sandy bottoms. All of
these groups, together, account for the origin of
most of the carbohydrates used in the system.
(2) The other essential type of production is the
fixation of atmospheric nitrogen its oxidation
and elaboration into simple compounds. It is,
well known that this fixation is accomplished by
bacteria in the soil and in nodules on the roots
of certain leguminous plants. Less well known,
but possibly more important in the atoll system,
is fixation of nitrogen by blue-green algae. This
occurs on the soil surface and possibly in fresh and
salt water. Much of the nitrogen available to
atoll organisms is probably fixed within the system,
but important quantities enter the system by way
of birds, rain, and ocean currents.
D. Transformation: The alteration oj primary
and fabrication of secondary organic compounds:
This function may be viewed as a succession of
processes, mostly involving a break-down of
organic compounds and their re-elaboration into
more complex ones. Some of these are of an
enormous order of complexity (e.g., nucleo-
proteins).
(1) Autotrophic plants, in the nourishment of
their own protoplasm and elaboration of stored
material, cellulose, lignin, and other materials,
carry out the first major step in a series of
turnovers of the products of photosynthesis. Of
course, additional inorganic materials are incorpo-
rated by this process, and many of the elaborated
compounds are infinitely more complex than the
original carbohydrates produced by photosyn-
thesis.
(2) Heterotrophic (parasitic and saprophytic)
plants carry this process a step farther in utilizing
already elaborated complex substances, as well as
simpler materials derived from their hosts and
organic substrata. Here may be mentioned the
utilization of dissolved organic matter in the
media by facultatively heterotrophic planktonic
algae as discussed by Saunders (8).
(3) Animals, feeding on plants in various ways,
convert plant organic matter into animal organic
matter. The principal classes of processes by
which this is accomplished are as follows:
a. Eating of phytoplankton by zooplankton.
b. Utilization of material produced by zooxan-
thellae, by their coelenterate hosts.
c. Reef grazing and boring.
d. Eating of land plants by animals.
e. Eating of dead plant parts by animals.
f. Parasitism of plants by animals.
(4) Secondary conversion of animal matter to
animal matter is accomplished as a result of
three well-known classes of processes, namely:
(a) Predation
(b) Parasitism
(c) Scavenging.
These are carried on in a great number of different
ways by a large number of animals. Reef grazing
and boring are important here, too.
(5) Reconversion of animal matter to plant
matter is not as conspicuous a process, but is
important nevertheless. There seem to be no
insectivorous plants on atolls, so this reconver-
sion is principally accomplished by bacteria and
fungi living mostly on dead, and occasionally on
living organic matter. It is an interesting question
whether zooxanthellae utilize in any way the
materials of their hosts' tissues.
E. Decomposition (usually but unfortunately
termed "reduction"): The destruction of the
elaborated organic compounds and reconversion
back to simple inorganic compounds and relative-
ly inert organic residues: Two main categories
of processes are involved here. (1) Physiological
oxidation (inappropriately termed respiration by
many plant physiologists), which is the oxida-
tion of carbohydrate materials within living cells
releasing the energy required for other life pro-
cesses. This process goes on constantly in all living
things. (2) Non-biological oxidation, both by
burning and by the slow oxidation of dead materials
that normally takes place on exposure to atmos-
pheric oxygen, aided or not by hydrolytic and
catalytic action.
163
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
The principal products of both sorts of processes
are CO 2 and H 2 O, with, of course, inorganic
and inert organic residues. Chemical energy is
released and converted into other forms.
F. Excretion (within the system): The release
of waste products and residues by organisms into
the media:
(1) In water, CO 2 and O 2 are released, as well
as excreta and soluble metabolic wastes. Calcare-
ous and siliceous skeletons and oily material
remain after disintegration of organisms.
(2) On land, likewise CO 2 , O 2 , and metabolic
wastes arc released in solution in air or water.
Guano and other excreta, as well as deciduous,
caducous, or severed plant parts are deposited
on the surface of the ground to decompose.
G. Accumulation- Storage of materials in
unchanging or very slowly changing form, i.e.,
temporary withdrawal of material (and energy)
from free circulation in the system.
(1) In bulk the limestone from calcareous
skeletons of plants and animals represents the
greatest and most important accumulation, the
principal component of the atoll itself.
(2) Phosphatic residues, mainly calcium phos-
phates, are present as phosphate rock, compo-
nents of soils, guano, and at least in some atolls
(e.g., Washington Island) as a phosphatic mud
or sludge on the lagoon bottom.
(3) Humus, both as raw humus in Pisonia
forests, and as more stable humic residues in Aj
horizons of soils, plays an important part in the
functioning of the system. The acid raw humus
contributes to the formation of phosphate rock,
and the soil humus helps to maintain the soil
in condition to support growth or larger plants
and micro-organisms. Humus, though relatively
inert, is continually undergoing a slow oxidation.
(4) Slight accumulations of charcoal, metal
oxides, silica, and silicates occur where human
activity is significant. Silica from sponge,
radiolarian, and diatom skeletons also occurs in
minute amounts, as well as small quantities of
silicates from floating pumice.
(5) Finally, fresh water, in the ground-water
lens, as well as in the several states of soil water,
may be regarded as a temporary accumulation.
The organic matter and other substances in
living organisms represent a large total quantity
but, as they are in a constant state of turnover,
should probably not be classed as an accumula-
tion.
These accumulations, along with the materials
in solution in the media, may be regarded as the
164
reservoirs of materials on which the other com-
ponents may draw for nourishment.
H. Turnover of materials and energy: Cate-
gories D, E, and F, above are to be classed as
turnover. In addition several more processes
may be so regarded.
1. Re-use of CO 2 released by oxidation.
2. Re-use of O 2 released by photosynthesis.
3. Re-use of H 2 O released by metabolic and
external chemical processes.
4. Re-use of fixed nitrogen, both from meta-
bolic wastes and from primary biological oxida-
tion.
5. Re-use of mineral nutrients released by
excretion and breakdown of organic materials.
6. Withdrawal from and return of various
materials to media.
7. Withdrawal from and return to accumula-
tions.
I. Miscellaneous other effects and processes
taking place within the system.
1. Inhibition by salt (NaCl). The organic activ-
ity in terrestrial situations seems subject to a
considerable inhibition by the salinity of the sea-
water medium. This inhibition results from
difference in osmotic pressure, the chemical
effects of absorption of excess sodium and chlo-
rine ions and consequent inhibition of absorp-
tion of others. The number of land organisms
completely adapted to the normal salt concen-
tration of the sea is limited. Hence establishment
of immigrant organisms is severely limited, and
many of those that become established function
at below their optimum levels. Salt water enters
the land environment as wind-borne spray, as
storm waves, and by diffusion through the
ground. The conspicuous nature of the limiting
effects of salinity may be a reflection either of the
small extent of the land habitat and consequently
great exposure to salt or of its probable geolog-
ically recent origin that has allowed little time
as yet for evolution of a special atoll biota.
2. Effects of sea-air interface: Category 1 is
really only one of the consequences of the fact
that the land portion of this ecosystem is a thin
lens inserted in the general sea-air interface.
The distribution of many organisms, marine as
well as land, is influenced by the character of this
interface. Aeration, principal release of energy
from insolation, frequently an abrupt break in
temperature gradient, local high salt concentra-
tions resulting from evaporation, solution and
other forms of erosion of limestone, and the
VOLUME 4
BOTANY
shaping of the contours of vegetation and con-
trol of its composition are all consequences of
the nature of this interface. Many more could
be enumerated.
3. Shelter effects. One of the reasons for the
diversity of animal life in such an apparently
simple environmental complex may be the
variety of habitats resulting from the surface
irregularity of the several substrata. The vegeta-
tion, the deeply pitted rock, the porous soil, and
the intricate nature of the reef lattice provide
shelter of various types for a large number of
species of animals (and plants, too) that have
widely differing requirements.
4. Burrowing and turning over of soil by crabs
is an important factor in the process of incor-
porating organic matter into the soil. Crab
burrows are very common on many atolls, and
fresh mineral soil is often piled or scattered
around their entrances. The mechanical tilling of
the soil in this manner has been compared to that
accomplished by earthworms in other habitats.
It doubtless is a process of great importance,
though no careful assessment of its extent or
effects has been made.
J. Losses (or excretion from the system):
a. Of the principal substances lost from the
system the first three listed below are present in
such constant proportions in the media outside
the ecosystem that the losses may be considered
as balanced almost exactly by inflow. The others
are fluctuating quantities, and there is no exact
relation between inflow and loss.
1. CO 2 , carried away by winds and currents.
2. O 2 , carried away by winds and currents.
3. N 2 , carried away by winds and currents.
4. Fresh water dispersed into media and carried
away by winds and currents.
5. Nitrates and organic N, carried away by
currents.
6. Phosphates, carried away by currents in
solution and suspension.
7. Other dissolved mineral substances, carried
away by currents.
8. CaCO 3 , carried away by currents in solu-
tion and suspension.
9. Plankton, carried away by currents.
10. Dead animals and plants and detached living
fixed organisms, carried away by currents and
storms.
11. Birds which migrate.
12. Export of copra.
13. Export of pearl shell, etc.
b. Energy losses:
1. Light, by reflection.
2. Heat, by radiation and convection and
carried by winds and currents.
3. Chemical energy lost with elaborated mate-
rials.
K. Balance: The resultant of all of the factors
at work on the segment of the universe (or of
nature) occupied by the atoll ecosystem is the
atoll itself. It may be thought of as a system
in a state of dynamic equilibrium with a positive
offset represented by the physical mass of the
atoll with its associated biota and its total of
organic and mineral matter over and above that
of the normal media air and sea water that
otherwise would occupy the space. All the
characteristics described serve to set the system
off from the surrounding undifferentiated media.
L. Trends: With such complexity, it is hard to
estimate trends, though it may be easy to discern
them. Over very long periods, the trend is obvi-
ously toward greater accumulation of material
and probably toward increasing complexity.
This trend usually seems directly related to slow
subsidence of substratum on which the atoll is
built, and may be expected to continue. On a
shorter time scale, it is possible to suggest that
during periods of general or eustatic rise in sea
level mass will increase, by addition of calcareous
material in layers. Biotic complexity may at the
same time decrease somewhat with tendency
toward loss of land habitats. With fall in sea level,
the trends may be the opposite loss of mass by
erosion and gain in biota with appearance of land
habitats, increased activity of sea birds, and
especially the results of occupation by man.
Presumably for about the last 3,500 years the
latter trend has been generally maintained.
Whether or not the last few decades have wit-
nessed a change in this trend is uncertain.
It seems clear that these major trends are
controlled by factors external to the system. The
ultimate control of sea level is as yet by no means
clear. The variation in CO 2 content of the air has
been suggested (7) as a factor that determines,
or at least influences, world temperatures, eva-
poration of sea water, accumulation of ice, and
consequent effects on sea level. It has been sug-
gested that the recent apparent reversal of the fall
of sea level may be due to the vastly accelerated
industrial activity which pours great quantities of
CO 2 into the atmosphere. If this is a valid
assumption it seems reasonable to think that the
165
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
present rise will continue, probably at an increas-
ing rate. Thus a prediction might be made that
the presently observed loss of land above water
by erosion may be accelerated by a rise in sea
level and consequent submergence of much or all
of the land area of atolls. Such predictions,
however, rest on very insecure bases at present.
On a still shorter time scale are the effects
produced by the occupancy of the atolls by man,
and especially modern man. These effects tend
to be drastic as far as the land portions of the
ecosystem are concerned, but trends arc as yet
hard to isolate. Certainly the replacement of the
native vegetation by coconut plantations and the
rise of the export of copra are notable and
probably involve a complex of related or depen-
dent effects. This change will probably continue
but certainly at a decelerated rate, as land for
expansion of plantations is becoming scarce.
Augmentation of the land biota will probably
continue as man's effect on the land environment
continues. Pollution of lagoons will undoubt-
edly increase, with resulting encouragement to
some organisms and ill effects on others. Fishing
activities have tended to decrease with contact
with civilization, but this trend may probably be
reversed and with use of such effective methods as
dynamiting and poisoning the marine biotas may
undergo considerable change. There has as yet
been little attempt to measure the results of such
factors so that here, as in other aspects of the
system, estimation of trends is highly speculative.
If such prediction is of interest, attention should
be directed toward critical study of the details
of the working of the system outlined above, to
clarify it and fill in the parts that are at present
inferential. It seems possible that if a firm under-
standing of this ecosystem is achieved it may be
used as a model in terms of which to study other
ecosystems.
SUMMARY
The general physical and "physiological"
framework of the coral atoll ecosystem has been
outlined in terms of the media in which the system
exists, nine categories of processes taking place
within the system, the balance or dynamic equi-
librium in the resultant structure brought about
by these processes, and suggested trends in the
state of this equilibrium. This highly generalized
picture rests on a vast accumulation of facts and
upon inferences drawn from them and from
pertinent facts derived from study of related or
analogous situations in other systems. It is hoped
that this description may serve, until a better
conception is devised, as a framework around
which an understanding of this segment of nature
may be built and as a guide for future research
designed to clarify our knowledge and apprecia-
tion of coral atolls.
REFERENCES
(1) Billings, W.D., and Bliss, L.C., 1955, An
Alpine Snowbank Ecosystem in the Medi-
cine Bow Mountains of Wyoming,
Bull. Ecol Soc. Amer., 36: 76 (abstract
only).
(2) Cain, S.A., 1956, The Expansion of the
Human Ecosystem, 1-10 (duplicated,
distributed at Symposium: Values in
human ecology, New York, Dec. 28,
1956).
(3) Dansereau, P., 1957, Biogeography, An Eco-
logical Approach, 1-394, New York.
(4) Evans, F.C., 1956, Ecosystem as the Basic
Unit in Ecology, Science, 123 (3208):
1127-1128.
(5) Costing, H.J., 1956, The Study of Plant
Communities, 1-440, San Francisco.
(6) Pitelka, F.A., 1955, High Arctic Tundras as
an Ecosystem, Bull. Ecol. Soc. Amer.,
36: 97 (title only, paper read at East
Lansing meeting of Ecological Society
of America, Sept. 1955).
(7) Plass, G.N., 1956, Carbon dioxide and the
Climate, Amer. Scientist, 44: 302-316.
(8) Saunders, G.W., 1957, Interrelations of Dis-
solved Organic Matter and Phytoplank-
ton, Bot. Rev., 23: 389-409.
(9) Tansley, A.G., 1935, The Use and Abuse of
Vegetational Concepts and Terms, Eco-
logy, 16: 284-307.
DISCUSSION
F.E. EGLfcR*. The approach used by the author and its
potential as an approach to land utilization is most valu-
able. The concept of the ecosystem has been slow to gain
recognition, possibly because it is based on the investiga-
tions of many fields of knowledge, seldom as the result of
166
one individual's effort. The plant ecologist often comes
from the field of taxonomy and may not be fully equipped
to handle phytosociological problems. The integration of
scientific knowledge by an ecologist requires an unusual
breadth of understanding.
VOLUME 4 BOTANY
M.S. DOTY : It is necessary to distinguish between poten- regarding when it stops being an atoll and when it becomes
tial and net productivity. a lagoon. With reference to the energy relationships, the
F.R. FOSBERG: Although potential productivity is some- constant blowing of wind corning from outside may contri-
thing to think about and discuss, the actual productivity bute to the system, but the chief source of energy is from the
is the only tangible aspect of it that could be evaluated. sun. The wind may build up sand and accumulate energy
G.F. PAPENFUSS: Are there atolls that lack a lagoon? in the system, but it may also do the opposite and release
F.R. FOSBERG i There is some question of interpretation potential energy from the system.
167
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
THE RELATIONSHIPS~BETWEEN ATOLLS AND BENTHIC ALGAEt
RALPH F. PALUMBO
School of Fisheries, University of Washington, Seattle, Washington, U.S.A.
In the U.S. Pacific Northwest, one is accus-
tomed to seeing large fronds of such algae as
Macrocystis and Ncrcocystis floating around in
the cold water of Puget Sound. In the Marshall
Islands of the warm Central Pacific Ocean, on
the other hand, where the species of algae dis-
cussed in this paper were collected, the absence
of these or any of the large forms of algae is
very apparent. There only colonies or individual
specimens of tiny filamentous algae or branched
species rarely exceeding a few inches in length
were found. The striking size difference in the
marine flora of the two regions brings up the
question as to the nature of the conditions caus-
ing this difference, and, at the same time, poses
the question as to reasons for the differences in
distribution of the algae in the atoll itself. This
paper will be confined to the latter problem.
A coral atoll is a relatively small outcropping
in the middle of an ocean and contains a diversi-
fied environment in which the growth of various
organisms is possible. The open ocean surround-
ing an atoll is a source of the minerals used in
the production of organic matter and thus acts
as a nutrient solution for the activities of the
various photosynthetic organisms present. The
principal members of the atoll community engag-
ing in photosynthetic activity, of course, are the
benthic and the planktonic algae. The distribution
of planktonic algae of the atoll lagoon is primarily
dependent on the local currents, and con-
sequently, the problem of this paper is to deter-
mine the differences in the environment which
might account for the distribution of the benthic
algae about the atoll.
Recent studies by Johnson (5) and others have
confirmed earlier suggestions that the calcareous
algae play an extremely vital part in the building
of coral atolls. Transects across the reef at
Bikini Atoll indicate that 40-90 % of the reef is
comprised of algal deposits, consisting primarily
of the remains of species of the green coralline
alga Halimcda and of the red coralline algae such
as PorolithoH, Cioniolithon, Mesophyllutn, and
Lithothamnion. Other organisms, such as corals
and foraminifera, have contributed to the reef
building process, but to a lesser extent. These
deposits extend to a depth of over 2,500 feet and
t Presented in abstract by M.S. Doty.
168
date back to the Miocene epoch. They serve as
pointers to the equilibrium or relatively stable
conditions that exist between the atoll inhabitants
and their environment. This theory has also
been promulgated by Odum and Odum (8) who
stated that the productivity of a reef just balances
the respiration, suggesting that the reef commun-
ity is a true ecological climax or open steady state
system.
The conditions that have existed, apparently
for millions of years, making possible this reef
building process, revolve around the activity of
these algae and their ability to flourish in this
type of environment. The warm temperatures
and the abundant radiant energy bathing the
exposed reefs combined with the rich supply of
oxygen and carbon dioxide form ideal conditions
for photosynthesis. The ability of these algae
to secrete calcium carbonate as well as other
minerals caused these particular species to sur-
vive, and their cementing action caused the
consolidation of algal and other debris in the
formation of the atoll mass.
The calcareous algae mentioned above are
found primarily in the regions of pounding surf,
where turbulence and oxygen rich waters are at
a maximum. Some of the non-encrusting cal-
careous algae, such as Codium and Udotea,
however, are found in the more protected habi-
tats of the tide pools and along the western
leeward reefs where the surf action is less severe.
These species apparently cannot survive the
rigorous action of the waves impinging upon the
northern reefs, but they do require similar condi-
tions of light, temperature, and mineral compo-
sition optimum for the growth of the more hardy
corallines.
The calcareous algae, however, are not the
only algae contributing to the mass of the reef.
Odum and Odum (8) have shown that the fila-
mentous green algae interwoven in the skeletal
material of the coral reef make up a large por-
tion of the mass and consequently increase the
overall productivity of the reef.
Of the conditions influencing algal growth in
an atoll, temperature is probably the least im-
portant. Studies that showed insignificant
VOLUME 4
BOTANY
temperature differences in the shallow waters
on the different sides of the coral reef were made
at Rongelap Atoll by Sargent and Austin (9).
The temperatures across the three hundred-
meter reef varied from 28.3C on the seaward
side to a maximum of 29.4'C on the lagoon side,
not sufficient to account for the differences in
productivity per unit area on the reef and in the
adjacent waters. Munk, Ewing, and Revelle (7)
found very slight temperature differences with
depth in the lagoon at Bikini Atoll. At three
feet below the surface, the temperature varied in
24 hours between 27.2 l 'C and 27.T J C and at
165 feet between 27.05 and 27.08C, indicating
a thorough mixing of the top and bottom layers
of water. This idea was partially supported by
radiological determinations made after the Baker
atom test in 1946, by these investigators as well
as by W.L. Ford (3). The temperatures in the
atoll environment, then, vary only slightly and
probably not enough to account for distribution-
al differences of the organisms present. The most
significant part that temperature plays undoubt-
edly occurs during low tides when the tide pools
become very warm and the beach rock areas are
exposed to the direct sunlight for hours at a
time. Here the only algae that survive are the
blue-green algae such as Anacystis and Schizo-
thrix, which possess simple structures and gela-
tinous sheaths.
Of the factors limiting the growth of photosyn-
thetic organisms, light is often the controlling
one. In the murky waters of many northern
seashores, the stratification of algal species is
well defined. This zonation can be explained
at least in part by the amount and quality of the
incident light and its effects upon the photosyn-
thetic activity of the various algae. For exam-
ple, the Chlorophyta are found only in shallow
water, as the red light from which they obtain
energy for photosynthesis cannot penetrate more
than a few feet. Some of the Rhodophyta, on the
other hand, may be found growing at depths up
to 300 feet because they can utilize the blue wave
lengths which penetrate to this depth. The Phaeo-
phyta use both the red and the deeper penetrating
orange-yellow light and thus frequent the zone
extending from the low tide line to a depth
of approximately 50 feet. Whether such an
explanation can be used to account for the dis-
tribution of the algae in an atoll environment is
open to question. Wells (11) reported that
"light or radiant energy is the principal factor
controlling the depth of growth of mutually
interdependent reef corals and symbiotic zooxan-
thellae... temperature is the principal factor con-
trolling their geographic distribution." On the
other "hand, Sargent and Austin (9) stated in
Rongelap lagoon, even though the light intensity
at 15 meters was only 20 " of that just below
the surface of the water, it was still adequate for
maximum photosynthesis in free-living plants. In
the unusually transparent water of Eniwetok
lagoon at depths up to 45 meters, the present
author and others of this laboratory using self-
contained breathing apparatus saw luxuriant
"fields" of green algae, especially Halimeda
nionilc, H. gigcis, and Caulerpa raccmosa, attached
to the white bottom sand. Gilmartin (4) has also
observed these and other green algae at depths
of 60 meters. Others have also noted the pre-
ponderance of green algae at depths usually
restricted to the deeper living red algae (10).
This can signify only that sufficient light does
penetrate the clear waters of these lagoons to
permit maximum growth of the "greens" and
possibly to discourage growth of the "reds." It
was also noted that some of the same species
inhabiting the deeper, relatively calm water were
also growing vigorously in shallow turbulent
water on the edge of the seaward reefs. Good
examples of this type of algae are the green
Halimeda opimtia and the red Asparagopsis taxi-
fonnis. In these instances at least, light does
not appear to be the controlling factor.
Another factor important in the growth of
algae is the mineral content of the medium.
Few quantitative studies have been made of the
mineral content of the waters in an atoll environ-
ment. Sargent and Austin determined the
amount of dissolved phosphorous in the water
at several localities on one of the reefs at Rongelap
Atoll and found that only small differences existed,
in fact, the values were so low that the measure-
ments were felt to be uncertain. The total dissolved
phosphorous over the reef at a depth of about
forty centimeters varied from 0.31 to 0.66 mg
atoms/liter. Measurements of Mg++, Ca++, Cr,
and total hardness made by Cloud (1) at Onotoa
Atoll in the Gilbert Islands showed that even
though there were diurnal variations in the
values for some of these factors, there were no
significant differences in the levels in the shallow
water at the four localities studied. Radiological
studies to determine the relative uptake of trace
minerals by marine organisms have been con-
ducted by this laboratory. These studies have
shown that minute amounts of manganese,
zirconium, zinc, cobalt, cesium, and iron can
be detected in amounts too small to be detected
by ordinary chemical techniques (6). The con-
centration of these minerals by sedentary marine
169
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
organisms over a long period of time could be
useful in the measurement of the different minerals
present and could also be useful in evaluating
their distribution about the atoll. Until further
information of a similar nature encompassing
several ecological situations is obtained, no finite
conclusions can be drawn regarding the effects
of varying mineral composition on algal growth
in the environment of an atoll.
A factor which must be considered in the
distribution of an alga is its habit or life form.
Several classifications of the life forms of marine
algae have been outlined, and it has been sug-
gested (2) that such classifications could be
useful in determining whether certain ecological
situations are characterized by the same types of
algae. As such an approach has been used with
partial success, it might well be expanded to include
a wider range of conditions and geographical
areas in order to evaluate its dependability.
Unfortunately the information relative to the
chemical and physical factors in an atoll environ-
ment is far from complete. The information we
do have indicates that the quantitative differences
in these factors are probably too small to ac-
count for distributional patterns among the
algae and other organisms present. Radiological
studies to aid in determining the relative amounts
of minerals about the atoll should be made, and
these, in conjunction with mineral nutrition
studies using radioactive tracers, might help to
clarify the role of minerals in the atoll environ-
ment. It has been proposed that the life form of
an alga may be the prime factor in determining
its distribution, and some studies indicate this
to be the case. However, sufficient discrepancies
exist to warrant further study of the problem.
In summary it can be stated that the occur-
rence of an alga in a particular ecological niche
is dependent upon many different factors, among
which the following must be included: the life
form of the alga, the chemical and physical factors
of the environment, the dynamic factors of wave
action and immersion, and the biological factors
imposed by other organisms in the vicinity.
REFERENCES
(1) Cloud, P.E., Jr., 1952, Preliminary report on
geology and marine environment of
Onotoa Atoll, Gilbert Islands, Nat. Res.
Counc., Pacific Science Board, Atoll
Res. Bull., 12:6-73.
(2) Feldmann, J., 1951, Ecology of Marine
Algae. In G.M. Smith, Manual of Phy-
cology, Chronica Botanica Company,
Waltham, Mass. pp. 312-334.
(3) Ford, W.L., 1949, Radiological and Salinity
Relationships in the Water at Bikini
Atoll, Trans. Amer. Geophvs. Union,
30(1): 46-54.
(4) Gilmartin, M., 1956, Personal communica-
tion.
(5) Johnson, J.H., 1954, Fossil Calcareous Algae
from Bikini Atoll. Bikini and Nearby
Atolls, Part 4, Paleontology, Geol.
Survey Prof. Paper 260-M, pp. 537-543.
(6) Lowman, F.G., Palumbo, R.F., and South,
D.J., 1957, The Occurrence and Dis-
tribution of Radioactive Non-fission
Products in Plants and Animals of the
Pacific Proving Grounds. Applied
Fisheries Laboratory, Univ. of Washing-
ton., U.S. Atomic Energy Commission
Report UWFL-5L
(7) Munk, W.H., Ewing, G.C., and Revelle,
R.R., 1949, Diffusion in Bikini Lagoon.
Trans. Amer. Geophys. Union, 30 (1):
59-66.
(8) Odum, H.T., and Odum, E.P., 1 955, Trophic
Structure and Productivity of a Wind-
ward Coral Reef on Eniwetok Atoll,
Ecol Monogr., 25(3): 291-320.
(9) Sargent, M.C., and Austin, T.S., 1954, Biolo-
gic Economy of Coral Reefs. Bikini and
Nearby Atolls, Part 2, Oceanography
(Biologic), Geol. Survey Prof. Paper
260-E, pp. 293-300.
(10) Taylor, W.R., 1950, Plants of Bikini. Univ.
of Michigan Press, Ann Arbor, Michi-
gan, 218 pp.
(11) Wells, J.W., 1954, Recent Corals of the
Marshall Islands. Bikini and Nearby
Atolls, Part 2, Oceanography (Biolo-
gic), Geol. Survey Prof. Paper 260-1,
pp. 385-486.
DISCUSSION
F.R. FOSBLRO: One must keep in mind that in the Pleis-
tocene there have been some violent fluctuations with great
changes in sea level and that the changes are of such an
170
extent that one would not think in the same terms. The
interpretation of a steady state existing since the Miocene
cannot be rigidly applied.
VOLUME 4
BOTANY
THE MICROBIOLOGY OF CORAL REEFSt
E.J. FERGUSON WOOD
Division of Fisheries and Oceanography, Cronulla, Australia.
The microbiology of coral atolls is a sadly
neglected field of marine research. We have little
information, apart from the studies on the
bacterial precipitation of calcium carbonate made
by Drew (6), Lipman (7), and Bavendamm (5) ;
some notes on blue-green algae associated with
coral reefs by Baas Becking (1 ) ; and some con-
clusions that may be drawn by inference from the
work of Baas Becking and Wood (3) and Baas
Becking, Wood, and Kaplan (4).
Drew came to the conclusion that large-scale
precipitation of calcium carbonate was caused in
tropical waters by a marine bacterium which he
called Bacterium calcis. He showed that this
bacterium could reduce nitrates to ammonia,
which reacted with the bicarbonate in sea water
to cuase the precipitation of calcium carbonate,
the calcium being derived from calcium sulphate.
This is, of course, theoretically possible in a
region where the calcium and carbonate equilibria
are in delicate balance near the saturation point
for calcium carbonate. Lipman questioned
Drew's hypothesis and considered the production
of ammonia from organic matter was the impor-
tant factor in calcium precipitation by marine
bacteria. He also pointed out that there were
so few bacteria in the open sea that extensive
precipitation was unlikely. Bavendamm showed
that there were enough bacteria in bottom depo-
sits to cause extensive calcium carbonate preci-
pitation, but concluded that there were no spec-
cific bacteria concerned.
Zobell (9), reviewing the situation, states that
fewer than 5 per cent of bacterial species in the
sea are endowed with the ability to liberate free
nitrogen from nitrate or nitrite in the presence
of abundant organic matter. My own observa-
tions show that few marine bacteria can liberate
either nitrogen or ammonia from nitrates, and
1 doubt whether this process is important in
calcium carbonate precipitation.
Possibly of greater significance in microbial
calcium carbonate precipitation is the utilization
of carbon dioxide by autotrophic bacteria and
photosynthetic microorganisms. There is no
information on the occurrence or abundance of
such organisms in the coral reef biocoenosis.
t Presented in abstract by R.F. Scagel.
The environmental factors which may affect
lime deposition include temperature, pressure,
hydrogen ion concentration, and rcdox potential.
In fresh water, salt concentrations also have an
effect, but in the sea, these do not vary sufficiently
to be important.
The effect of temperature on calcium carbonate
precipitation is stressed by Sverdrup ct al. (8),
who show that, because of the rapid decrease of
solubility of carbon dioxide with rise of tempera-
ture, extensive deposition is possible only in
warmer waters. These authors suggest, too,
that, as pressure increases the solubility of carbon
dioxide, lime deposition will be confined to rela-
tively shallow waters.
It should be remembered that the carbonate
equilibrium is very easily reversible, and that
precipitation presupposes re-solution with a
slight change of conditions in the right direction.
Thus, organisms removing carbonate by photo-
synthesis or photoreduction may, and do, con-
tinue to produce it in the dark by respiration.
Baas Becking and Wood, and Baas Becking,
Wood and Kaplan have considered the relation
between pH/Eh, along with the microorganisms
and the environment for a number of biocoen-
oses. In the first paper, the limitations of a
number of autotrophs by the environment was
studied; in the second, the chemical equilibria
imposing possible limitations were delineated,
and this has been carried still further by Baas
Becking (in press). It is unfortunate, but unavoid-
able, that no observations were made in coral
reef areas, so we can get only an approximate and
theoretical picture, of these regions. In the
estuaries of temperate regions, the pH of the
water where measured ranged from 7.0 to 9.4,
with a mean of 8.25, while the muds ranged from
near 5.0 to near 9.5, with a mean of 7.4.
The estuarine waters had a mean Eh of +362
mV with a range from + 150 to + 500 mV, while
the muds ranged from + 600 to - 350 mV with
two means, one at about + 325, and the other at
about - 75 mV.
Biologically speaking, all autotrophic processes
except sulphate reduction and photoreduction
171
PROCFFDINGS OF THE NINTH PACIFIC SCIFNCh CONGRESS
could occur in cstuarine waters, while sulphate
reduction and photorcduction of carbon dioxide,
as well as the other autotrophic reactions are
possible, and can be shown to occur in the estua-
rine muds. Baas Becking and Wood consider
that the sulphur cycle dominates the muds, and
that these in turn dominate the water in shallow
estuaries. The lower mean Eh of the muds
reflects this dominance.
In fresh- water limestone regions studied by
these authors, the pH ranged from 5.5 to 9.0, with
a mean of 7. 19, while the Eh ranged from -+ 100
to +500 mV with a mean at 333 mV. The
pH shows that bicarbonate ion dominated the
waters, even those with high concentrations of
calcium (at 7.2, there will be little COj ion). In
sea water, at a pH over 8, the equilibrium of the
CO 2 , HCO 3 , CO 3 would move towards the
carbonate. At that pH there will be no free
CO 2 , and 4 parts of bicarbonate to one of
carbonate ion. A change in the pH to 9.4 will
reverse the bicarbonate-carbonate ratios to 1 part
of bicarbonate to about 3 parts of carbonate ion,
i.e., carbonate will become the limiting factor in
photosynthesis.
In temperate estuaries, a pH of 9.3 - 9.4 is
reached on shallow flats with a large plant popula-
tion, e.g., of Zostent, in the middle of the day.
Although, theoretically, calcium carbonate
should actively precipitate under these conditions,
the greater acidity of the muds will limit the
amount actually deposited. Another limit is
imposed by the short duration of these high pH
values, and the counter-effect of respiration at
night.
Coral reefs differ from the estuaries studied by
Baas Becking and his colleagues in having, on the
whole, less photosynthctic plants, and conse-
quently less organic detrital matter in the muds.
This should lead to a lower activity of the organ-
isms of the sulphur cycle, which depend on
heterotrophic bacteria to produce the low Eh
required for starting sulphate reduction. The
green and purple sulphur bacteria also require
low Eh values to initiate the oxidation of sulphy-
dryl. These regions will also differ from the
fresh-water limestone regions in having a con-
siderable amount of calcium sulphate which was
absent in the areas studied by Baas Becking,
Wood and Kaplan.
It can be safely predicted that the alkaline limit
of the coral reef environment will be controlled
by carbonate, and the lower Eh limit by the
amount of organic matter, i.e., reducing material.
172
The other limits cannot be estimated except by
actual observation.
In temperate estuaries, and in mangrove
swamps, phosphate is largely controlled by the
sulphur cycle in the mud (2). The insolubility of
the sulphides of iron cause the release of phos-
phoric acid from ferric phosphate in the presence
of hydrogen sulphide. An alternative method for
the collection and storage of phosphate by
microorganisms is the chelation of phosphate in
the pcctic sheaths, also described by Baas Beck-
ing and Mackay. I have noticed that encrusting
growths of blue-green algae usually have a zone
of sulphate reduction immediately below them,
and could thus use the sulphur cycle to obtain
their phosphate. However, one would expect
the pectic chelation mechanism to be of greater
importance in the phosphorus cycle on coral
reefs. Nitrogen fixation may also be an important
function of the Nostocaceae on coral reefs.
It will be realized from the account that 1 have
given that we are largely in a field of pure specula-
tion when we consider the microbiology of coral
reefs, but that we have a priori some means of
approach to the problem. This approach should
include (1) the study of the microbial compo-
nents of the coral ecosystem, (2) their interaction
and cumulative effect on the environment, and
(3) the actual biological mechanism of lime
deposition especially as skeletal material.
REFERENCES
(1) Baas Becking, L.G.M., 1951, Notes on some
Cyanophyceae of the Pacific region,
Proc. Kon. Ned. Akad. Wet., 54: 2.
(2) Baas Becking, L.G.M., and Mackey, Mar-
garet, 1956, Biological processes in the
estuarine environment, V: The in-
fluence of Enteromorpha on the environ-
ment, Proc. Kon. Ned. Akad. Wet.,
B59: 109-123.
(3) Baas Becking, L.G.M., and Wood, E.J.F.,
1955, Biological processes in the estua-
rine environment, I &1I: Ecology of
the sulphur cycle, Proc. Kon. Ned.
Akad. Wet., B58: 160-181.
(4) Baas Becking, L.G.M., Wood, E.J.F., and
Kaplan, I.R., 1957, Biological processes
in the estuarine environment, X: The
place of the estuarine environment
within the aqueous milieu, Proc. Kon.
Ned. Akad. Wet., B60: 88-102.
VOLUMH 4 BOTANY
(5) Bavendamm, W., 1932, Die mikrobiolo- (7) Lipman, C.B., 1924, A critical and experi-
gische Kalkfallung in der tropischen mental study of Drew's bacterial
See, Arch. Mikrobiol., 3: 205-276. hypothesis in CaCO 3 precipitation in
the sea, Carnegie lust. Wash. Dept.
(6) Drew, C.H., 1914, On the precipitation of Mar. #/W., 19: 179-191.
calcium carbonate in the sea by marine (8) Sverdrup, H.U., Johnson, M.W., and Flem-
bacteria, and on the action of denitrify- ing, R.H., 1942, The Oceans. Prentice
ing bacteria in tropical and temperate Hall, N.Y.
seas, Pap. Tortugas Lab. Wash., 5: (V) Zobell, C.E., 1946, Marine Microbiology.
7-45. Chronica Botanica Co. Waltham, Mass.
173
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
THE MARINE BIOGEOGRAPHICAL PROVINCES OF
THE SOUTH PACIFIC?
V.J. CHAPMAN
Auckland University College, Auckland, New Zealand.
Our present knowledge of the biogeography of
the South Pacific stems from contributions spread
over a number of years dealing with algal zona-
tion and ecology in South America (23,24,25,26),
Australia (9,1,12,20,31,32), and New Zealand
(21,8,4,5,2,3,10,11,15). The subantarctic islands
have also been examined, the first records being
those of Hooker (14) and Harvey (13), but there
are recent ones for the New Zealand subantarctic
islands (6), for the Cro/et Is. (17) and for the
subantarctic islands generally (18). Another
island flora that is relevant to any discussion
on biogeographical provinces is that of Juan
Fernandez (16).
The present author believes that proper appre-
ciation and understanding of the provinces is
most readily secured by using as a basis the major
categories recognized by Stephenson (27,28.29)
in his work on the marine fauna and flora of
South Africa. The three major categories were:
(a) A warm water component, predominant
at Durban and gradually decreasing to Port
Nolloth. This can be regarded as essentially a
subtropical province.
(b) A cold water component associated with
the cold upwclling on the S.W. African coast,
and gradually disappearing towards Cape Town.
This can be regarded as forming essentially a
cool temperate province.
(c) A southern intermediate flora of the Cape
district which can be regarded as a warm tem-
perate province.
These major groups were correlated with
changes in mean sea temperature, and Stephenson
(29) related them to the main southern littoral
faunas as described by Eckman based upon
intervals of 10C. These are as follows:
( 1 ) Tropical and sub-tropical faunas : minimum
temperature not below 20C.
(2) Warm temperate faunas: minimum tem-
perature 10-20C.
(3) Cool temperate faunas: minimum tem-
perature 0- 10C.
(4) Antarctic faunas: minimum temperature
0C or lower.
t Presented in abstract by M.S. Doty.
174
A similar ten degree temperature barrier has
been proposed by Setchell (22) as operative for
marine algal zones. Recent work by Bennett
and Pope (1) on Australian faunas suggests that
the minimum sea temperature boundary of 10C
between warm and cold temperature faunas may
need to be raised to 11.5C. In Australia, the
principal marine faunal and floral provinces
were first proposed by Hedley but they have been
amended since then. At the present time the
following can be recognized:
(a) The Banksian from Cape York southwards.
(b) The Solanderian in the Coral Sea area.
(c) The Peronian in Southern Queensland and
Northern New South Wales.
(d) The Maugean in Bass St. and around
Tasmania.
(e) The Flindersian in Victoria and South
Australia.
(f) The Dampenan in Western Australia and
up to the Northern Territories and the
Gulf of Carpentaria.
Recent work on the littoral faunas and floras
has served to substantiate these provinces (1),
and they have also been shown to be valid for
the pelagic Dinoflagellates (33). Of these Austra-
lian provinces, the Maugean must be regarded
as cool temperate, being associated with cold
antarctic waters. The Peronian and Flindersian
provinces are both warm temperate, whilst the
Solanderian and Damperian are tropical. Fur-
ther work on those parts of Australia just north
of the Peronian and Flindersian provinces will
probably show (hat there are two small subtrop-
ical provinces characterized by a mixture of
tropical and warm temperate species.
In New Zealand, Moore (19) has summarized
the distribution of some 200 marine algae, and
as a result she recognized seven distinct provinces
which can be compared with those of Powell
for the marine fauna and of Cockayne for the
terrestrial plants. The present writer, whilst
agreeing generally with the provinces of Moore
and Powell is of the opinion that some modi-
fication is desirable on the basis that has been
outlined above. The Kermadec algal flora are
VOLUME 4
BOTANY
based on old lists (1 ) 9 and a recent collection
clearly contains tropical, e.g., Caulerpa racemosa,
C. wekbiana, Pocockiella nigrescens, and warm
temperate species, and can therefore be regarded
as a sub-tropical province. Such information
as is at present available indicates that Norfolk
Is. falls into the same province. Off the shores
of South America, the flora of Juan Fernandez
suggests affinities with Australia and New
Zealand, but there is a distinct warm water
element (e.g., Hydroclathrus, Padina, Micro-
dictyon japonicum, Chaetomorpha antennind)
which justifies treating it as subtropical. The
Aupourian or Auckland province is definitely
warm temperate (11), the proportion of warm
temperate species gradually decreasing towards
Cook Strait. The subtropical convergence crosses
New Zealand in Cook Strait and, as the mini-
mum sea temperature of the convergence is
around 10 C C, it forms the logical boundary
between the warm temperate province of the
northern island arid the cool temperate province
of the South Island. The Chatham Islands lie
very close to the sub-tropical convergence, and
they possess a flora with warm and cool temperate
elements. There appears, however, to be a
slightly higher proportion of warm temperate
elements, and it may therefore be placed in the
warm temperate Auckland province, or as a
sub-province of the Auckland province.
The marine algal flora of most of the South
Island of New Zealand must be regarded as cool
temperate, and it would seem to have a relation-
ship with the Maugean province of Bass Strait
and Tasmania. It is to be noted that although
Tasmania lies just north of the sub-tropical
convergence nevertheless the waters of the cold
antarctic current maintain a cool temperate
flora and fauna. The present writer considers
that the central and intermediate provinces of
New Zealand should be merged into a single
province, the Zealandian.
The Forsterian (Stewart Is.) and Antipodean
(= Rossian of Moore) provinces of New Zealand
comprise the subantarctic islands. Both provinces
contain a large proportion of southern species,
so much so that it seems desirable to recognize
a sub-antarctic province which would also include
Kerguelen, Crozet, and Heard Islands. In this
case, therefore, the minimum sea temperature
boundary would be less than 10C. It may be
suggested that further work in and around Mag-
ellan Straits, Tierra del Fuego, and the Faukland
Islands may show that they also represent a sub-
antarctic province with a flora differing from that
of the cool temperate province to the north and
the extreme antarctic province of the Antarctic
continent and adjacent Islands.
There remain the coral reefs and atolls of the
South Pacific. Our knowledge of the marine
algal floras of these islands is at present very
meagre, but it can be suggested that they repre-
sent essentially a tropical province which can con-
veniently be called the Central Pacific province.
It remains to be seen whether New Caledonia,
the New Hebrides, and the Solomon Islands
group, now placed in the Solandcrian province
have a flora sufficiently similar to that of the
Central Pacific to justify the extension eastwards
of the Solanderian province.
Some further confirmation of the conclusions
that have been reached above can be derived from
a comparative study of the basic zonations in
the South Pacific, especially those of Australia
and New Zealand. Guiler (12) has compared
(Table 1) the basic (i.e., not greatly exposed nor
greatly protected) zonation of Tasmania with
that of New Zealand, Chile, Victoria, and S.W.
Africa; and he considers that ecologically the
zonation in Tasmania indicates a greater affinity
with that found in New Zealand than with that
of southern Australia. In making the compara-
tive study, the basis used is that proposed by
the Stephensons (30).
In Table 2 a comparison is given of the basic
zonation of exposed and protected shores in the
Auckland province of New Zealand with the
zonation reported from the Zealandian province
of the S. Island (Banks Peninsula).
Finally a comparison may be made of the basic
zonation from three of the Australian provinces
(Table 3).
The distinction between the tropical flora and
the other two is very evident. The similarity
between the Peronian and the Aucklandian
provinces is also brought out. In New Zealand,
Pyura is more frequent in the South Island,
and it evidently is a representative of the cool
temperate zone. A fuller comparison of the
varied zonations that have been described from
the different provinces is really essential to
bring out the finer points that demonstrate the
differences.
Whilst our knowledge of the algae and the
algal zonations in many parts of the South
Pacific is as yet lacking or very inadequate,
nevertheless from the data available it would
appear that the picture that has been drawn
represents the situation as it is at present. It
may also serve to highlight those areas from which
further information would be very valuable.
175
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Table 1.
Tasmania
New Zealand
Victoria
S.W. Africa
Chile
Supra-
littoral
fringe
Littorinids
Littorinids
Myxophyceae
Littorinids
Littorinids
Littorinids
Chamaesipho-
Chthalamus
Chamaesipho-
Chthalamus
Chamaesipho-
Chthalamus
Bolanus
Chamaesipho-
Bolanus
Mid-
littoral
Galeolaria
Hermella
Volsella
Elminius or
Saxostrea
Algae or
Galeolaria
Pomatoceros
Serpulid
worms
Corallines
Corallines-
Hormosira or
Mytilus
Pyura
Corallines
Corallines
Sub-
littoral
fringe
Lessonia
Xiphophora
or
Durvillea
Ecklonia Cystophora
Carpophyllum Durvillea
Lessonia or
Durvillea
Laminaria
Ecklonia i
Macrocystis
Lessonia
Durvillea
Table 2.
Hauraki Gulf : Piha 1 Poor Knights Is. Banks Peninsula
(Protected) (Exposed) (Exposed) (Exposed)
Littorinids Melaraphe Melaraphe Mclaraphe
Supra-littoral fringe (Mclaraphe) |
Myxophyceae Bostrychia Myxophyceae Bostrychia
i Chamaesipho spp. Chamacsipho Chthalamus Chamaesipho
Elminius Volsella i Modiolus Elminius Serpulid worm
Saxostrea Gigartina alveata Apophloea Elminius
Mid-littoral Corallina- Pachymenia Corallina- Mediolus
Hormosira Mytilus i Novastoa Mytilus
Corallina-
Halopteris
Sublittoral fringe Carpophyllum Durvillea (1 sp.) Xiphophora Durvillea (2 sp.)
Ecklonia Carpophyllum Carpophyllum
Supra-littoral fringe
Mid-littoral
Sub-littoral fringe
176
S. Australia
(Flindersian)
Melaraphe
Myxophyceae
Barnacles
Galeolaria
Corallines
Ecklonia
Cystophora
Table 3.
I New South Wales
(Peronian)
Melaraphe
Tetraclita (exposed)
Barnacles
(Chthalamus-Tctraclita)
Chamaesipho
Galeolaria
Pyura
Corallines
Hormosira
Pyura
Ecklonia
Phyllospora
Queensland
(Solanderian)
Tectarius
Melaraphe
Barnacles
(Chthalamus -Tetraclita)
Ostrea
Coral
Sargassum
VOLUME 4
BOTANY
REFERENCES
(1) Bennett, I., and Pope, E.G., 1953, Inter-
tidal Zonation of the Exposed Rocky
Shores of Victoria, together with a
re-arrangement of the biogeographical
provinces of temperate Australian
shores, Aust. Journ. Mar. and F. W. Res.,
4(1): 105.
(2) Beveridge, W.A., and Chapman, V.J., 1950,
The Zonation of Marine Algae at Piha,
New Zealand, in relation to the tidal
factor (Studies in Inter-tidal Zonation
2), Pac. Sc/., 4(3): 188.
(3) Carnahan, J.A., 1952, Inter-tidal Zonation
at Rangitoto Island, New Zealand
(Studies in Inter-tidal Zonation 4), Pac.
Sci., 7(1): 35.
(4) Chapman, V.J., 1950, The Marine Algal
Communities of Stanmore Bay, New
Zealand (Studies in Inter-tidal Zonation
1), Pac. SW., 4(1): 63.
(5) 1956a, A Revision of the
Marine Algae of New Zealand, Pt,
I: Myxophyceae and Chlorophyceae,
Journ. Linn. Soc. Bot., 55: 333.
(6) , 1956b, Marine Algal
Provinces and Zones in New Zealand:
their place in a World System, Proc.
1st Geog. Conf., N.Z. Geog. Soc., p. 27.
(7) Cotton, A.D., 1911, Marine Algae from
the North of New Zealand and the
Kermadecs, Kew Bull., 61: 253.
(8) Cranwell, L.M., and Moore, L.B., 1938,
Inter-tidal communities of the Poor
Knights Islands, N.Z., Trans. Rov.
Soc. N.Z., 67: 375.
(9) Dahin, W.J., Bennett, I., and Pope, E.C.,
1948, A Study of Certain Aspects of the
Ecology of the Inter-tidal Zone of the
New South Wales Coast, Aust. Journ.
Sci. Res. Ser. B., 1 (2): 176.
(W) Dellow, U.V., 1950, Inter-tidal Ecology at
Narrow Neck Reef, New Zealand (Stu-
dies in Inter-tidal Zonation 3), Pac.
Sci., 4(4): 355.
(11) , 1955, Marine Algal Ecol-
ogy of the Hauraki Gulf, New Zealand,
Trans. Roy. Soc. N.Z., 83 (1): 1.
(12) Guiler, E.R., 1952, The Nature of Inter-
tidal Zonation in Tasmania, Pap. Proc.
Roy. Soc. Tasm., 86: 31. Handbook of
the New Zealand Flora, London.
(13) Harvey, W.H., 1855, Algae in J.D. Hooker,
Flora Novae Zelandiae, 2:211, London.
(14) Hooker, J.D., 1847, Flora Antarctica, 2,
London.
(15) Knox, G.A., 1953, The Inter-tidal Ecology
of Taylor's Mistake, Banks Peninsula,
Trans. Roy. Soc. N.Z., 81 (2): 189.
(16) Levring, T., 1941, Die Meeresalgen der
Juan Fernandez-Inseln, Nat. Hist.
Juan Fern\ and Easter Is., Ed. C. Skott-
sberg,2: 601.
(17) ___ , 1944, Meeresalgen von
den Crozet Inseln und Kerguelen, Ark.
Dot., 31 (8): 1.
(18) _______ - , 1945, Marine Algae
from some Antarctic and Subantarctic
Islands, Lunds. Univ. Arss. N.F. Avd. 2,
(19) Moore, L.B., 1949, The Marine Algal Pro-
vinces of New Zealand, Rov. Soc. N.Z.
Proc. 6th Sci. Congr., p. 187.
(20) Newton, L., and Cribb, A.B., 1951, Some
Aspects of Algal Ecology in Britain and
Australia, Research, 4: 449.
(21) Oliver, W.R.B., 1923, Marine Littoral Plant
and Animal Communities in New Zea-
land, Trans, and Proc, N.Z. Inst., 54: 496.
(22) Setchell, W.A., 1920, Stenothermy and
Zone Invasion, Amer. Nat., 54: 385.
(23) Skottsberg, C., 1907, Zur Kenntnis der Sub-
artanktischen und Antarktischen Meere-
salgen I. Phaeophyceen, Wiss. Ergebn.
d. Schwed. Sildpolar Expd. 1901-03,
4, Stockholm.
(24) . ________ ..,1921, Marine Algae I:
Phaeophyceae (Botanische Ergebnisse
der Schwediochen Expedition nuch
Patagonien unde der Feuerland, 1907-
09), K. Svenska Vet-Akad. Hand, 61,
Stockholm.
(25) __ ._. _, 1923, Marine Algae II:
Rhodophyceae, Ibid., 63, Stockholm.
(26) , 1950, Sodra Ishavets
Algflora, K. Svenska Vetenskap-Akad-
Arsbok, p. 367.
(27) Stephenson, T.A., 1939, The Constitution
of the Inter-tidal Fauna and Flora of
South Africa, Part I, Journ. Linn. Soc.
Zoo I., 40: 487.
(28) , 1944, Ibid., Part II, Ann.
Natal Mus., 10: 261.
(29) , 1948, Ibid., Part III, Ann.
Natal Mm., 11 (2): 207.
177
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
(30) Stephenson, T.A. and Stephenson, A., 1949, of the algal ecology, Trans. Roy. Soc.
The Universal Features of Zonation be- S. Aust., 71 (2): 228-229.
tween Tide Marks on Rocky Coasts, (32) , 1948, Ibid., II: ThePen-
Journ. EcoL, 37 (2): 289. nington Bay Region, Ibid., 72 (1): 144.
(33) Wood, E.J.F., 1954, Dinoflagellates in the
(31) WomersIey,H.B.S., 1947, The Marine Algae Australian Region, Aust. Journ. Mar.
of Kangaroo Is. I: A general account and F. W. Res., 5 (2): 171.
178
VOLUME 4
BOTANY
COASTAL MARINE PLANTS AROUND CAUDA HARBOUR
(NEAR NHATRANG)t
PHAM-HOANG-HO
Deputy Director, VInstitut Oceanographique, Nhatrang, Vietnam.
The Nhatrang bay, and the sea coast of Central
Vietnam in general have a physiognomy which
distinguishes them from the rest of the whole
coast from the ecological point of view.
The "Annamitic Cordillera" ends there with
rocky outgrowths delimiting sandy bays while
alluvial deposits brought in by large rivers con-
tinue to build elsewhere muddy and sandy coasts.
Climatologically, the coast of Central Vietnam
owes the Cordillera a climate diametrically op-
posed to that of the rest of the country. When
in the North and the South the summer monsoon
brings in the rain, the dry season (from February
to August) prevails in the Centre. The yearly
average of rains amounts to 1,441 mm against
1,979 in Saigon.
At Nhatrang, the thermic regime is also the
same as in Saigon, with a small temperature
difference (4) between the warmest and the
coolest month.
Finally, oceanographically the central coast is
distinct. There, in fact, diurnal and semi-diurnal
tides manifest themselves on an equal basis. The
tide regime is a mixed one as compared with
a semi-diurnal regime in the South and a diurnal
regime in the North. (We are awaiting details
about the harbour establishment along the Viet-
nam coast).
Thus the physical, climatological, and oceanog-
raphical data put emphasis on the interest of
a comparative study of the ecology of the central
sea-coast and that of the rest of Vietnam.
We began with the study of the submarine
vegetation at Nhatrang, mainly on rocky spots.
We took, for example, the rhyolitic point of
Qiuda. We could distinguish the three general
bionomical zones of Stephenson (1949), viz.
a Littorine zone, a Cirripede zone, and the sub-
littoral fringe. But the zones which can be easily
evidenced are from top to bottom:
(a) the Calothrix pilosa Born, and Flash girdle,
30 cm wide, watered by high sea spray (embruns)
and submerged under the biggest spring tides
(lunar tropic). This is a discontinued girdle,
confined to rough substrata.
Upward accessories: Littorina scabra Lin.,
t Presented by M.S. Doty.
Tectarus gramdaris Gray, Lygia sp., ubiquists:
Grapsus grapsus.
(b) the Brachytrichia maculans Com. girdle,
covering 80% to 90% of the area. Another typical
growth, Littorina scraba Lin., has a maximum
thickness which changes according to the tide.
In the anfractuosities are the Isognomon sp.
and Pollicipes mittellae ("mode battu"). And at
the lower limit are the patella: Cellana amussilata
Reev., Acmaea saccharina Linn. A. granostriata
Sch Patella aster Reev., etc.
(c) the Ostrea cucullata girdle, situated at the
mid-tide level with Ectocarpus braeviarticulatus
J.Ag., Chnoospora pacifica J.Ag., Ge Helium pusil-
lum Le Jolis ("mode battif), Acrocystis nana
Zan. On the walls, eurytopics colonies of Paly-
thao sp.
(d) the Gelicliella acerosa girdle at the low water
levels is the large Phaophyccae zone : Sargasswn
ssp. Turbinaria ornata and Turbinaria sp. ("mode
battu") Padina commersoni Bory., Chnoospora
implexa J. Ag. ("mode peu battu a calme"). The
great phaeophyceae shelter abundant epiphytic
flora. On padina commersoni Bory, for example,
we found Griffithsia metcalfii Tseng, G. tennis
C. Ag.,Acrochaetumgmci/e, Centrocerasclavatum
Mont., Ceramium mazatlanense Daws., Poly-
siphonia sp. Great acorn-shells (Balanus tintin-
nabulum) were covered with Melobeseae or Ralfsia
sp. at the beaten spots; on the less beaten rocks
were Gastrochaena cymbium Sprengl. On the
scattered rocks you find Enteromorpha intestinalis
Link. On the small grounds where the waves
are not sharp are the tufts of Ectocarpus mitchellae
mixed with Enteromorpha clatharata J. Ag. and
E. Kyllini Blid.
Inside the beach rock blocks we found
Lithodomus lima Jouss., L. Malaccarnus Reev.,
Pholadidea far rot i Jouss., Area turella, several
crustaceous: Gonodactylus, Alpheideae, Isopoda;
Spongiaires and Polychaeta.
(e) the infra-littoral level begins with Thalassia
Hemprichii Asch., Halophila ovalis Hook, Diplau-
thera Uninervis Asch. that is found only at the low
spring waters. You find there Colpomenia sinuosa
Derb & Sol., Padina commersoni, Gracilaria
179
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
crassa Harv., Bornetclla oligospora, Neomeris dcma setosa and other sea-urchins, Mulleria sp.,
annulata Dick, Halimeda gracilis Harv., Struvea the Crinoids. It is also the coral zone whose
anastomasans Pice., Galaxaura fastigiata Dec., study is being conducted by M.M.G. Ranson
etc At the lower part, you see between rocks: and Nguyen-Thanh-Tri.
Ceratodictyon spongiosum Zan., Halymenia dila- We do not yet have data about the deep lower
tata Zan., Liagora ceranoides Lam., Liagora infra-littoral zone. Dawson (1954) found Gala-
farinosa Lam., Q. Jilamentosa Chou, etc..., Dia- xaura Vietnamiensis Daws, at 30 m deep.
DISCUSSION
M.S. DOT Y: Is the 4 C. temperature difference referred P.H. HO: The temperatures given are for air.
to for air or water?
180
VOLUME 4 BOTANY
BENTHIC ALGAL PRObljCTIviTY ~IN THE NORTH PACIFIC WITH""
PARTICULAR REFERENCE TO THE COAST OF BRITISH COLUMBIA
ROBERT F. SCAGEL
University of British Columbia , Vancouver, Canada,
In comparison with the progress in our knowl-
edge of most groups of plants especially con-
cerning their life-histories and distributions
the advances made in marine phycology have
been relatively slow. The limited access to living
material or to the facilities to maintain the larger
marine algae in the living condition for a pro-
longed period of time, the difliculties of col-
lection particularly in the subtidal zone and
the lack of any extensive direct economic value
until recent years have all contributed to this
slow progress. However, in spite of these dif-
ficulties, there has been a considerable amount
of interest in the marine algae, including many
studies of their ecology. Although this interest
has been fairly widespread in a number of
countries of the world, until recently there has
been little activity in the field of marine benthic
algal ecology on the Pacific Coast of North
America and nothing of a comprehensive nature
has been published for this area. It is an ana-
chronism that this should be so in a region which
received such prominence some fifty years ago
through the efforts of a pioneer in the field, the
late William Albert Setchell.
Although the knowledge of the effect of tem-
perature on the world-wide distribution of plants
both horizontally and vertically had developed
gradually over a period of many years, it was
only during the last hundred years that the atten-
tion of phycologists was brought to a consider-
ation of the reasons for the observed distributions
of the marine algae. The historical development
of this trend of thought and investigation during
this early period has been reviewed by Setchell
(30). Starting over fifty years ago and extending
through a series of papers from 1893 to 1935,
Setchell made a noteworthy attempt to explain
the world-wide distribution of marine algae, and
especially members of the Laminariales on the
Pacific Coast of North America, on the basis of
latitudinal and seasonal temperature distributions.
The physical data available during this early
period were limited, but many of the principles
set forth by Setchell concerning the distributions
of marine algae are as sound now as when they
were first proposed. Except for more precise
knowledge of the physical and chemical factors
of the environment and the distributions of the
algae concerned, much of SetchelFs ecological
work can still be used as a good foundation for
further study. Although it was largely a two-
dimensional approach to the marine environment,
Setchell's work made a significant contribution
to the development of marine algal ecology.
Lamouroux (23, 24) had suggested the possibi-
lity that temperature stratification in the sea
might account for the vertical distribution of the
marine algae and had considered the effect of
tides on intertidal zonations, but this trend to
analyze the vertical distribution of the marine
algae was not taken up in detail until much later.
Coleman (12) was one of the first to emphasize
the use of tide levels to account for the vertical
distribution of the marine algae in the intertidal
zone. In a study in Oregon, on the Pacific Coast
of the United States, Doty (15) has given further
evidence for the relationship between the verti-
cal distributions of marine algae and critical tide
levels.
There have been a number of lists of marine
algae published, and attempts have been made
not only to relate the floras of one area to another,
such as that by Okamura (25, 26) in the North
Pacific, but also to account in a general way for
distributions on the basis of ocean currents, such
as that by Isaac (22) in the area around South
Africa. However, there soon followed a decided
shift to intertidal studies of regional areas, such
as that by Feldmann (17) in the Mediterranean
and Chapman (11) and his students in New Zea-
land. Some attempts have also been made to
describe universal features of intertidal zonation
throughout the world (32). At the same time,
there has been a tendency to place greater em-
phasis on the interrelationships between the
various organisms.
Many of these intertidal studies have been of
great value as an initial descriptive stage of in-
vestigation, and there is a need for further de-
scriptive studies of this type in new and unde-
scribed regions. However, the variety of systems
of nomenclature and terms that have been pro-
posed by marine ecologists to describe zonation,
associations, and other ecological concepts have
181
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
frequently only complicated the descriptive study
rather than succeeded in explaining the observed
phenomena. This has led to some confusion in
terminology. It is a debatable point whether there
can be such a thing as a universal system of classi-
fication beyond a generalized scheme, such as
that proposed by Ekman (16), and it is question-
able whether some of the systems proposed can
contribute further to progress in marine algal
ecology even in regional studies without simpli-
fication or clarification.
Although the shift in emphasis to the inter-
relationship of organisms was an important one,
in some instances this approach has been respon-
sible for excluding adequate concurrent studies
of the physical and chemical aspects of the en-
vironment. It is for this reason that a case may
be made for reassessing our position in marine
algal ecology, and a critical evaluation of the next
steps to be taken to further our progress is timely.
Perhaps what may be called a three-dimensional
or an oceanographic approach can be used to
analyze more precisely various factors in the
marine environment and the relationship of these
factors to bcnthic algal productivity. A step in
this direction has been considered recently with
some measure of success by Dawson (13, 14) in
Baja California, Doty (15) in Oregon and Wo-
mersley (34) in Australia.
To the oceanographer, the most complicated
physical or chemical situation to explain may be
the smallest unit of the environment with which
he is faced. This is partly a problem of instru-
mentation. However, it is usually much easier
not only to recognize significant discontinuity in
properties, such as temperature, salinity even
plankton distributions over extensive areas of
the ocean than in restricted or local regions, but
also to use such information in describing dyna-
mic processes. Hence, it is suggested that more
attention should be given to studies of the general
distribution of various physical and chemical
properties in the marine environment in an
attempt to set up some workable hypotheses to
account for observed distributions of marine
algae. In this way we may hope to explain and
account for biological phenomena rather than be
satisfied by a description of the phenomena or by
terms to describe them which do nothing more
than give names to dynamic aspects of marine
ecology much in need of logical explanation.
With increased activity recently in oceanography
in the Pacific, we may now hope for more abun-
dant and usable data on some of the more general
oceanographic properties of the North Pacific.
In specific cases, and particularly in more
182
restricted areas, the ecologist will be forced to
turn more attention to obtaining in situ physical
and chemical data before further progress can
be made.
One can arbitrarily start by summarizing all
the factors in the marine environment as geolog-
ical, physical, chemical, and biological. The
way in which these are considered may be some-
what a matter of interpretation. Salinity, for
example, may be considered directly, as a chemical
factor, or indirectly as a physical factor respon-
sible for changes in density and thus contributing
to the pattern of circulation. Likewise, the nature
of the substratum may be considered indirectly as
a geological factor, or directly as a physical or
mechanical factor restricting or permitting estab-
lishment of benthonic organisms because of
particle size. There has been much written on
many of these aspects of ecological study in
special cases, but it is suggested that, in a general
overall reassessment of the environment, an
attempt be made to proceed from this more
general position to the particulate. This approach
may initially lead only to the erection of further
hypotheses, as the indirect or direct nature of the
action of any particular factor in the environment
may ultimately be established only by further
experimental work either in the field or in the
laboratory. This is the approach that is being
promoted in the studies on marine algae being
carried out on the Coast of British Columbia,
Canada, some aspects of which will be mentioned
briefly in this review.
Although on occasion in the past the position
of the taxonomist has been questioned in the
scientific field, it is apparent that the need for
fundamentally sound taxonomic studies of the
marine algae is almost as great now as it has ever
been, particularly since the recent progress in
knowledge of the fields of biochemistry, cytology,
and genetics. However, in the sense implied here,
the work of taxonomist has reached a logical
conclusion only when it is applied, and it is this
application that is frequently left for the ecolo-
gist. As a step in the direction of increasing our
knowledge of the taxa which comprise the tools of
the algal ecologist and of completing this de-
scriptive phase of the study of the marine ben-
thonic algae, an annotated check list has just been
completed for the Coast of British Columbia and
Northern Washington (28). Based on this list,
further studies are now in progress to augment
the existing data on the marine flora of British
Columbia, not only insofar as distributions are
concerned, but also relating to life-histories,
growth, reproduction, and seasonal aspects.
These problems are now being tackled in marine
laboratories which have running seawater avail-
able, as well as in the controlled-environment
tanks and rooms on the campus of the University
of British Columbia. These fundamental studies
are basic to all other aspects of ecological
research, and more especially when an attempt
is made to use specific organisms as indicators
of oceanographic conditions.
Not only does this descriptive phase require
an adequate consideration of the taxonomic
aspects, but a more complete description of the
other factors in the environment is needed. With
increased activity recently and currently in general
oceanographic studies of the North Pacific by
a number of institutions on the Pacific Coast of
Canada, in Japan, and in the United States, there
has resulted a considerable body of knowledge,
but it is still far from adequate for the ecologist,
particularly for one interested in coastal dyna-
mics. Circulation problems, temperature, sali-
nity, oxygen, and in some areas, phosphate and
nitrate distributions are fairly well known in a
broad and general sense, but at the present time
very few restricted areas are known oceanograph-
ically in sufficient detail. More precise physical
and chemical data, even in these more restricted
areas, are also inadequate. The distribution of
other chemical constituents, and to a large extent
even a knowledge of the plankton composition,
distribution, and activity, is almost completely
lacking. It is obvious that the more data that
become available the more clearly one can tackle
problems relating to specific distributions and
set up field and laboratory studies to test hy-
potheses.
It is already apparent that much can be done
experimentally both in the field and in the labora-
tory with benthonic marine algae. Many of the
problems encountered by the ecologist dealing
with the larger marine algae present unique
culture problems and require an entirely different
approach, both in the field as well as in the labo-
ratory. Some studies on growth and reproduction,
particularly of some of the larger Laminariales,
have been done in this region, both in the field
(27) as well as in the laboratory. Although the
size of many of the cold-water marine algae add
new problems, at least some of the stages can be
carried out to the point where transplants can be
made from the laboratory into the sea for further
study. Transplant experiments of natural popu-
lations of these larger marine algae, even in the
case of Macrocystis, are quite feasible.
The successful use of the experimental approach
in the laboratory is primarily dependent on having
VOLUME 4 BOTANY
facilities for maintaining temperature and light
control, although the size of plants may again
present certain special problems. Cultures of
Laminariales have been maintained in con-
trolled-environment tanks at the University of
British Columbia for as long as a year, during
which the complete sexual generations were
cultured and the young sporophytes reached a
length of fourteen inches, well past the stage
where secondary morphological characteristics
had developed. These studies have permitted
indisputable identification of the sporophytes to
genus and in some cases to species. The study of
cultures in this group suggests that much of the
early work on gametophytes in the Laminariales,
and in fact even on the early sporophytes, may be
in some question. In most of the early studies,
reported plants were not grown long enough to
establish beyond doubt the characteristic
secondary morphological features of the sporo-
phytes of the genera from which zoospores were
initially obtained. In the presence of contaminat-
ing zoospores of other species, there is not other
way of establishing that the same species or even
the same genus in the Laminariales was obtained
in the sporophyte generation succeeding the
gametophyte generations in culture.
It would be remiss not to mention much of the
worthwhile physiological work that has been
done on marine algae. However, there is a need
for a great deal more physiological work, parti-
cularly of the type done by Gail (18, 19,20) in an
attempt to related physiological processes more
specifically and directly to the environment and
ecological problems encountered in the field.
In physiological studies, there is frequently a
tendency to proceed more and more deeply into
special aspects of the physiological behaviour or
biochemistry of an organism under artificial
conditions, rather than to project back to the
field and attempt to explain behaviour under the
conditions existing in the natural environment.
Almost all of the quantitative aspects of benthic
algal productivity in this area have related to
species of economic interest (27,21). These studies
have dealt largely with quantities and distribu-
tion and have contributed little to an evaluation
or an explanation, in terms of oceanographic
factors, of the causes for this production.
Obviously the ideal of a functional interpre-
tation is dependent on an adequate and balanced
knowledge of all of the foregoing aspects of
the qualitative and quantitative features of both
the organisms and the environment. Many,
much-needed, data are still lacking. An attempt
to follow this line of investigation has been started
183
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
on the Coast of British Columbia, and in a some-
what more restricted area at the north end of
Vancouver Island in Queen Charlotte Strait.
Further detailed work is in progress in the Strait
of Juan de Fuca at the south end of Vancouver
Island between Vancouver Island and Northern
Washington.
The Pacific Coast of Canada is ideally suited to
a study of benthonic organisms and the effect of
oceanographic factors on their distribution both
in the intertidal and the subtidal zones. Although
the coast of British Columbia is only about 600
miles long, proceeding directly from the Strait
of Juan de Fuca to Dixon Entrance, if all its
various ramifications are included, there is a
coastline estimated at about 25,000 miles in
length. The tidal amplitude in this region is
great, ranging from about 1 1 feet at the southern
boundary to nearly 26 feet at the northern boun-
dary. As a result of thorough mixing in the
coastal region, the upper zone in this area,
except for a few local anomalies, is characteristi-
cally rather uniform in temperature at any one
period and fluctuations occur within narrow
limits. The annual range in temperature of the
sea water near the surface is from about 6 degrees
C to 18 degrees C. On the other hand, because
of the excessive run-off from large rivers,
especially through the long mainland inlets,
there are conditions ranging from practically
fresh water at one extreme to full ocean salinity
of about 34% at the other extreme. The oceanog-
raphic conditions characteristic of the Coast
thus provide a particularly ideal area in which
to study the distribution of marine benthonic
organisms in relation to salinity over a rather
extensive geographic area.
Throughout the Coast, the physical nature of
the substratum ranging from mud and sand
at one extreme to solid rock at the other deter-
mines to a large extent the organisms which are
found in a specific area. However, a comparison
of the flora and fauna on various types of bottom
is possible in a number of regions which are
otherwise oceanographically rather similar. This
permits a correlation of the distribution of a wide
variety of plants and animals with other physical
and chemical factors of the environment.
Biological observations, extending over the
whole length of coastline, indicate that there is
a high degree of uniformity in the populations of
many benthonic plants and animals extending
from the Strait of Juan de Fuca to Dixon
Entrance. This would be expected under the
relatively uniform conditions of temperature
indicated. In attempting to correlate the dis-
tribution of some of these organisms with salinity
characteristics, as well as other oceanographic
factors, there are several areas on the Coast
which could be used for purposes of this study.
Although some supporting observations have
been made in the Strait of Juan de Fuca and
Dixon Entrance, and further work is being done
in the former area at the present time, the present
review is restricted largely to a consideration of
the vicinity of Queen Charlotte Strait near the
north end of Vancouver Island.
Although soundings are still somewhat incom-
plete for the area, a general study of the bottom
topography in Queen Charlotte Strait indicates
extensive shallows, particularly along the Van-
couver Island side of the Strait and around Mal-
colm Island. In this region an abundant and
varied intertidal and subtidal flora and fauna
are supported. In the central part of the Strait,
and between Nigei and Vancouver Islands, there
are deeper channels exceeding 100 fathoms in
depth. These channels are not continuous,
however, with the deeper waters of the mainland
inlets and Johnstone Strait, and exhibit physical
and chemical properties quite distinct from the
latter.
The salinity distribution near the surface in
Queen Charlotte Strait indicates a general cir-
culation in a counter-clockwise fashion. The
run-off from the mainland inlets along the north
shore and at the east end of the Strait, particu-
larly from Knight Inlet at the east end of the
Strait, contributes large volumes of fresh water
which tends to move seaward at the surface mix-
ing as it progresses along the north shore into
Queen Charlotte Sound with the deeper more
saline water below. The more saline water from
the open ocean and Queen Charlotte Sound
moves into the Strait centrally as well as along the
north side of Malcolm Island. The intrusion of
high salinity water along the deep channels in
the central part of the Strait is also apparent.
This general pattern of salinity distribution, with
fluctuations to varying degrees near the surface,
is pronounced in the upper zone to a depth of
at least 20 meters.
An analysis of surface salinity data over a
ten-year period from Pine Island Lighthouse
(/ through 10), which is near the entrance to
Queen Charlotte Strait, indicates a salinity
maximum of about 34% and a minimum of about
28% with an annual mean of 31.75%. Although
this station is not fully characteristic of the whole
of Queen Charlotte Strait, the data available give
VOLUME 4
BOTANY
a close approximation of the annual salinity
fluctuations in the more oceanic part of the
Strait. Insufficient data are available from
Pulteney Point on Malcolm Island as yet to
analyze the seasonal fluctuations in the central
region of the Strait, but, for the period available,
a range of about 28% to 32% with an annual
mean of 28.95% is indicated.
A comparison of the temperature-salinity
characteristics of various parts of the Strait and
the connecting bodies of water by means of T-S
diagrams indicates the discreteness of the water
masses typical of Johnstone Strait, Knight Inlet,
and Queen Charlotte Sound. The T-S diagrams
for Queen Charlotte Strait indicate a character-
istically intermediate condition between these
extremes in properties of temperature and salinity
for the greater part of the Strait.
An analysis of surface temperature data over
a ten-year period from Pine Island Lighthouse
(7 through JO) indicates a temperature maximum
of about 12 degrees C and a minimum of about
5 degrees C, with an annual mean of 8.6 degrees
C. Although this station, as already indicated,
is not characteristic of the Strait in all regions,
it probably gives a reasonable approximation of
the annual temperature fluctuations for the outer
part of the Strait. As is true in regard to salinity,
insufficient data are available from Pulteney
Point on Malcolm Island as yet to analyze the
seasonal fluctuations in the central region of the
Strait, but for the period available a range of
from about 5 degrees C to 1 1 degrees C with an
annual mean of 8.25 degrees C is indicated.
The tidal amplitude in this area is about 17
feet, so that there is exposed an extensive inter-
tidal flora and fauna during lowtide periods.
Although no attempt will be made to analyze the
vertical distribution of the organisms of the
intertidal zone here, it may be pointed out that
many of them can be related to certain tide levels.
Biological observations have been made
throughout Queen Charlotte Strait, although a
more intensive study has been undertaken at
Hope, Deer, and Malcolm Islands. The observa-
tions have been restricted so far to the more
conspicuous algae and invertebrate animals in
the intertidal and immediately accessible subtidal
zones. These localities present a transition from
Hope Island, where the highest salinities are
encountered, to the north and east sides of the
Strait where the lowest salinities are found. Deer
Island and Malcolm Island are intermediate
between these extremes.
Some organisms in the area are more cos-
mopolitan in their distribution, particularly in
their tolerance to extreme dilution. Extending
throughout the Strait are forms such as Alaria
tenuifolia Setchell f. tenuifolia, Cymathere tripli-
cata (P. and R.) J.Ag., Costaria costata (Turn.)
Saunders, Costaria mertensii J.Ag., Laminaria
saccharina (Linnaeus) Lamouroux f. saccharina,
Nereocystis luetkeana P. and R., Porphyra per-
forata}. Ag. f. perforata, Rhodomela larix (Turn.)
C. Ag., Odonthaliafloccosa (Esper) Falk., Mytilus
edulis Linnaeus, Littorina planaxis Philippi and
Strongylocentrotus drobachiensis (Miiller).
Restricted to the region of highest salinity, as
at Hope Island, are Postclsia palmaeformis Rupr.,
Lessoniopsis littoralis (Farl. and Setch.) Reinke,
Laminaria setchellii Silva, Pelvetiopsis limitata
(Setchell) Gardner f. limitata, Dilsea californica
(J.Ag.) O. Kuntze, Erythrophyllum delesserioides
J.Ag., Jridaca Uneare (S. and G.) Kylin, Hymenena
setchellii Gardner, Ptilota asplenioides (Esper)
C.Ag., Ptilota californica Rupr., Ptilota hypnoides
Harvey, Mitel/a polymerus (Sowerby) and
Flustrella corniculata (Smitt). Pleurophycus
gardneri Setch. and Saund., Pterygophora califor-
nica Rupr., two forms of Hedophyllum sessile
(C.Ag.) Setch. and Styela montereyensis Dall are
also present in the regions of highest salinity but
are somewhat less restricted in their distribution
and extend into the Strait almost as far as Deer
Island.
A few organisms extend still further into the
Strait but only slightly beyond Deer Island.
Among these are Alaria nana Schrader and Alaria
marginata P. and R. Still others extend from
Hope Island past Deer Island down into the
Strait as far as Malcolm Island, but not as far as
the east and north sides of the Strait. Among
these are Macrocystis integrifolia Bory, Egregia
menziesii (Turner) Aresch. subsp. menziesii,
Alaria valida (Kjell. and Setch.) f. valida, the
bullate form of Hedophyllum sessile (C.Ag.)
Setch., Constantinea simplex Setch., Mytilus
californianus Conrad, Strongylocentrotus purpu-
ratus (Stimpson) and Haliotis kamtchatica Jones.
Isolated populations of the abalone (Haliotis
kamtchatica), which have been noted further
eastward in Johnstone Strait, and which may be
related to local oceanographic features, such as
upwelling, present somewhat of an anomaly to
the general distribution. A further exception to
the distribution described in this group is that of
Macrocystis integrifolia. Although Macrocystis
occurs in regions of high or relatively high sali-
nities and extends as far down the Strait as
185
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Malcolm Island and over to Numas Island, it
does not occur in the most exposed areas on the
open coast where Postelsia and Mitella are en-
countered.
There is a distribution of organisms here which
follows closely the pattern of salinity distribution
in the Strait, which in turn reflects the circulation
within the area. It would be premature to say that
salinity is directly responsible for the observed
distributions of all the organisms encountered.
But one may say that the distribution reflects the
dependence on waters characteristic of the open
ocean and exposure. In some instances, it may be
directly salinity that is a causal factor. The open
coast, on the other hand, has organisms associated
with surf conditions. It has been suggested that
the high oxygen requirement of certain organisms
is met only in such an exposed environment.
However, the measurement of oxygen values in
the sea in this area, does not directly support
this argument, as the oxygen content of the waters
within the sheltered Strait is as high or higher
than in the surf in the exposed regions. This is
particularly true in the central part of the Strait
when there is a heavy bloom of phytoplankton
in the area, at which time the water may be super-
saturated to as much as 175 per cent. Likewise,
although it is known that many marine algae
have a high inorganic phosphate requirement,
there is no evidence that this nutrient is ever
limiting in this area within the zone occupied by
the benthonic marine algae. There is a great
need for further knowledge of the presence, dis-
tribution, amounts, and availability of many
more dissolved inorganic and particulate organic
substances. The evidence so far points also to
the need for a study of quantitative removal and
the rate of removal of such substances and precise
requirements for growth and reproduction of the
benthonic marine algae.
The restriction of certain organisms to surf
conditions suggests that constant movement' of
water is required to provide nutrients and gases
which may be rapidly exhausted from the im-
mediate micro-environment of the individual
alga, or in the case of the sessile marine inverte-
brate, such as Mitella polymerus, to provide
particulate food. It may be that lowering the
concentration or removal by dilution or water
movement of some substances which accumulate
above a certain level of concentration in the
micro-environment is just as significant as the
availability of others.
In summary, a detailed study of the distribution
of marine benthonic organisms in Queen
186
Charlotte Strait has been limited so far to the
more conspicuous algae and invertebrates en-
countered. The relationship of these distributions
to the salinity distribution indicates that more
intensive study of the flora and fauna in this area,
as well as elsewhere on the Coast, will provide
further supporting evidence of the effect of ocean-
ographic variables on the distribution of marine
benthonic organisms and the possibility of using
such organisms as indicators of oceanographic
conditions both in time and space. It is hoped
that this oceanographic approach both quali-
tatively and quantitatively may then lead not
only to a clearer understanding and explanation
of the fundamental relationships between the
organisms and their environment, but also to an
understanding of the interrelationships and in-
teraction among the organisms themselves.
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(1) Anon, 1944, Observations of Seawater Tem-
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Coast of Canada, Fisheries Research
Board of Canada, Nanaimo; Pacific
Oceanographic Group, Vol. 5, 1942
and 1943.
(2) - - , 1946, Ibid., 6, 1944 and 1945.
(3) , 1948, Ibid., 7, 1946 and 1947.
(4) , 1949, Ibid., 8,1948.
(5) , 1950, Ibid., 9, 1949.
(6) , 1951, Ibid., 10, 1950.
(7) ., 1952, Ibid., 11, 1951.
(8) , 1953, Ibid., 12, 1952.
(9) - , 1955, /AW., 14, 1954.
(10) , 1956, Ibid., 15, 1955.
(11) Chapman, V.J., 1950, The Marine Algal
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Zealand. Studies in Intertidal Zonation 1,
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(12) Coleman, J., 1933, The Nature of Inter-
tidal Zonation of Plants and Animals,
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15 figs., 6 tables.
(13) Dawson, E.Y., 1945, Marine Algae Asso-
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western Coast of Baja California, Mexico,
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(14) , 1951, A Further Study of
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VOLUME 4
BOTANY
(15) Doty, M.S., 1946, Critical Tide Factors
that are Correlated with the Vertical
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Organisms along the Pacific Coast,
Ecology, 27: 315-328, 6 figs.
(16) Ekman,S., 1935,TiergeographiedesMeeres.
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(17) Feldmann, J., 1937, Recherches sur la
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(23) Lamouroux, J.V.F., 1825, Distribution
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(25) Okamura, K., 1926, On the Distribution
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588-592.
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(29) Setchcll, W.A., 1893, On the Classification
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(30) .. _- _, 1917, Geographical Distri-
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(31) _, 1935, Geographic Elements
of the Marine Flora of the North Pacific
Ocean, Amer. Nat., 69: 560-577.
(32) Stephenson, T.A., and Stephenson, A.,
1949, The Universal Features of Zonation
between Tide-marks on Rocky Coasts,
Jour. Ecol., 37 (2): 289-305, pi. 8, 4 figs.
(33) Tokida, J., 1954, The Marine Algae of
Southern Saghalien, Me. Fac. Fish,
Hokkaido Univ., 2 (1): 1-264, 15 pis.,
4 figs., 5 tables.
(34) Womersley, H.B.S., 1956, The Marine Algae
of Kangaroo Island, IV: The Algal
Ecology of American River Inlet, Austra-
lian Jour. Mar. and Freshwater Res.,
7(1): 64-87, 7 pis., 4 figs.
DISCUSSION
G.F. PAPENFUSS: Is anything known concerning growth
rates of the larger brown algae in the North Pacific?
F.R. SCAGEL: In Nereoaplis during July and August,
the blades can increase in length by 2 to 3 inches per day.
Small sporophytcs a few inches in height in April can de-
velop into mature, reproducing plants of over 100 feet
in length by late June of the same year.
187
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
ON THE EFFICIENCY OF PRIMARY PRODUCTION IN THE OCEANS
JOHN H. RYTHER
Woods Hole Oceanographic Institution, Woods Hole, Massachusetts, U.S.A.
INTRODUCTION
What is the maximum possible rate of organic
production in nature? Is this maximum ever
attained in the ocean? What are the limiting
factors of the marine environment which normally
prevent this potential from being realized or
even approached?
These fundamental questions may be attacked
in either of two ways. First, through extensive
and repeated oceanographic surveys we may
accumulate a body of data pertaining to the
magnitude of primary production in the sea.
Eventually these data should include the appro-
ximate range of this process, and through correla-
tion with chemical and physical characteristics
of the water, one should be able to extrapolate
or predict production values for similar oceanic
areas.
Second, we may consider the efficiency of
utilization of radient energy in the photosyn-
thetic process and in the organic yeild of mass
algal cultures, the environmental conditions which
influence and control this efficiency, and the exis-
tence and relative importance of these factors in
the sea. In this way, too, it should be possible
to predict the potential and, if sufficient informa-
tion is available, the actual magnitude of primary
production in the various parts of the ocean as a
function of its physical and chemical characteris-
tics.
Both of these methods have obvious disad-
vantages and limitations, so an attempt will be
made to use a combination of the two in the
following discussion. First the magnitude and
efficiency of photosynthesis will be reviewed
throughout its entire measured range starting
with carefully controlled quantum yield experi-
ments in the laboratory, through mass algal cul-
ture experiments, both indoors and out, to meas-
urements of primary production in the sea,
from the highest known values to the lowest.
This will be followed by a consideration of the
factors which affect photosynthetic efficiencies,
treating these quantitatively as far as is possible.
Finally an attempt will be made to evaluate the
relative importance of these factors in the various
parts of the ocean and to relate them to the
observed values of primary production.
188
THE QUANTUM REQUIREMENT
OF PHOTOSYNTHESIS
The quantum requirement or quantum effi-
ciency of photosynthesis (the reciprocal of the
quantum yield) may be defined as the number of
quanta of light energy required to reduce one
mole of CO 2 to CH 2 O. Although a minority of
investigators, mainly Warburg and Burk (28),
have reported extremely low quantum require-
ments (4 or less approaching 100% efficiency
of utilization of the absorbed light), it is the opi-
nion of most plant physiologists that a maximum
sustained photosyntetic yield requires 8-12
quanta of red light for the reduction of one
mole of CO 2 . (See 3, 30, 5, 6, 2.) Values within
this range have been observed in a wide variety of
plant species, the literature on which is extensive-
ly reviewed by Rabinowitch (77, chapter 29)
and Daniels (2) .
As a quantum of red light contains 41 k cal
and one CH 2 O group has a heat of combustion
of 112 k cal, an average quantum requirement of
10 corresponds to an efficiency of
(112x 100)/(lOx41) = 27%.
Efficiencies of this general order are obtained
only at very low intensities of light and when the
physiological condition of the plants and their
external environment are considered to be optimal
for photosynthesis. Presumably, then, this (appro-
ximate) value represents the maximum biological
potential of plants in converting light into chemi-
cal energy, a maximum which is imposed by
the limitations of the physical-biochemical pro-
cess of photosynthesis.
THE EFFICIENCY OF MASS ALGAL
CULTURE YIELDS
Experiments in mass algal culture by the
Solar Energy Research Group at Agricultural
College, Wageningen, Netherlands, have demon-
strated that the maximum sustained efficiency
observed in photosynthesis experiments is closely
approached in terms of the organic yield of rela-
tively long-term growth experiments. (See
11, 30, 29, 27.) Mass cultures of Chlorella
were grown by this group under artificial illumi-
nation with yield efficiencies, calculated on the
VOLUME 4
BOTANY
basis of the heat of combustions of the cells
produced, frequently ranging from 10-15% and
occasionally greater than 20%. As the organic
yield in such experiments does not take into
consideration respiratory losses which are auto-
matically corrected for in manometric photo-
synthesis measurements, the proximity of some of
their efficiency values to that observed in quantum
yield studies is truly remarkable.
However, when these cultures were grown out
of doors under natural illumination, efficiencies
declined markedly to values of 1-3% under full
sunlight, as high as 6% in shaded cultures (22%
in sunlight) 1 .
Somewhat greater yields were obtained by a
group of Japanese workers (13) who grew Chlo-
rella (and other unicellular algae) throughout
the year in outdoor bubbling cultures. Their
yields, in grams dry weight of organic matter
produced per square meter per day, averaged
16.5 in summer, about 2.4 in winter when
temperatures were apparently unfavorable. If
we may assume that their Chlorella were similar
in composition to the "normal" cells of Ket-
chum and Redfield (9) its heat of combustion
calculated by indirect colorimetry would be 5.4 k
cal/gram, which is comparable to the values
reported by Kok (11). Using this figure, their
yield data, and the average radiation values
for each experiment, a wide range of efficiencies
were obtained from 0.06% to 14.0%, averaging
about 4%. To the author's knowledge, the
maximum yields obtained by this group represent
the most efficient photosynthetic utilization of
natural illumination which has been reported.
THE MAGNITUDE AND EFFICIENCY
OF PRIMARY PRODUCTION IN
THE OCEAN
We may now turn to the ocean and present a
brief and partial review of primary production
values in various coastal and deep sea areas. No
attempt will be made to review the literature
completely, and all earlier values of uncertain
validity are omitted. In general an attempt has
been made to represent as many different types of
marine environments and as wide a range of
production values as possible. Many of the data
are as yet unpublished.
Several different methods have been used to
measure or calculate production by the various
authors. These will not be described here, but
the reader may refer to the cited references for
this infomation. Odum (14), Odum and Odum
(15), and Riley (19) have based their values on
measurements of in situ oxygen changes in the
water, using quite different techniques in each
case. Yentsch used the familiar "light and dark
bottle" method of measuring oxygen production
in his East Sound, Washington, studies. Stee-
mann Nielsen (25 ) and, in part, the present
author employed the C 14 method (24). The
remainder of the values were calculated from
radiation, light penetration, and chlorophyll data
using the method recently described by Ryther
and Yentsch (23).
Radiation values, except where specifically
measured with a standard type radiation meter,
were taken from Kimball (JO) who gives mean
radiation values at sea level on a world-wide and
seasonal basis corrected for average cloud cover.
The use of Kimball's tables many be somewhat
unsatisfactory in estimating efficiencies of pro-
duction measured by the C 14 method as the latter
were obtained on specific days when the radiation
may have deviated considerably from the mean.
This objection does not apply to values obtained
by the chlorophyll-light method which themselves
were calculated by using Kimball's tables. All
other efficiencies were based on direct measure-
ments of radiation except for Odum (14) and
Odum and Odum (15) who estimated radiation
from the equation given by Kennedy (8).
The organic yield (g carbon/m 2 /day) has been
calculated from oxygen production using an
assimilatory quotient of 1.25 (see 22) which
represents organic carbon or approximately
50% of the ash-free dry weight of plant material.
This may be converted into comparable energy
units by multiplying by 2 x 5.4, assuming that
the heat of combustion of the organic yield is the
same as that of normal Chlorella cells or 5.4 k
cal/g (see above). 2 Efficiency was then calculated
by dividing the daily organic production by 50%
of the incident radiation, both expressed as gram
calories/cm 2 /day.
Table 1 gives the seven highest single values for
primary production known to the author for the
marine environment. Of these, the highest values
1 Photosynthetic efficiencies of the utilization of solar energy are calculated here and elsewhere in this paper on the basis
of the visible spectrum only (400-700 mu) or roughly 50% of the total incident radiation.
2 If production is calculated as glucose-carbon (assimilatory quotient = 1), a higher value for carbon assimilation is
obtained. This may be multiplied by 2.5 (as glucose is 40% carbon) x 3.7 (the heat of combustion of glucose) giving
nearly the same energy content as of calculated on the basis of the actual composition of the plants.
189
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
were found in the benthic populations studied by
Odum (14) and Odum and Odum (IS), 10.3 g
carbon/m 2 /day for the turtle grass community
and 9.8 g carbon /m 2 /day for the coral reef, cor-
responding to efficiencies in the utilization of the
radiation reaching the surface of the water of
4.0% and 3.1% respectively.
The other production values in Table 1 range
from 2.8 to 5.4 g carbon/m 2 /day with less than
two-fold variation. The implications of the
striking similarity of these values, obtained
from widely differing types of environments, will
be discussed in the final section of this paper.
In contrast to the high values reported in
Table 1, Table 2 shows what may be considered
as average or normal production rates for various
estuarine, inshore, coastal, and oceanic areas.
Again the most striking feature of these is their
similarity. The shallow water areas have pro-
duction values ranging from about 0.1 to 1.5 g
carbon/m 2 /day, and it is interesting to note that
the same range was observed seasonally in the
two regions where production was followed
throughout the year, Long Island Sound and the
continental shelf off New York.
It is perhaps surprising that the productivity
of a polluted embayment, Great South Bay,
averaged only 0.24 g carbon/m 2 /day in midsum-
mer despite the fact that chlorophyll concentra-
tions in the euphotic layer averaged about
10 mg/m 3 or some 10-20 times that of unpolluted
coastal areas. This is probably due to the
shallow depth of Great South Bay and the high
turbidity of its waters, factors which will be dis-
cussed in more detail later.
Production measurements in Allen Bay, in the
Canadian Arctic made by Mr. Spencer Apollonio
almost daily during the brief open-water season
(July-August) showed no indication of an initial
flowering of any magnitude following the breakup
of the ice, though values did decrease throughout
the period from 0.69 to 0.02 g carbon/m 2 /day,
averaging 0.19. One possible explanation for the
low production of this area is that much of the
euphotic zone was diluted with melt water low in
nutrients concentration.
The oceanic values for the North Atlantic were
obtained during a passage from Woods Hole,
Mass, to Plymouth, England by HMS Discovery
III in April, 1957. They are probably higher
than average for the year as the cruise apparently
coincided with the spring diatom flowering (see
the deep water continental shelf values for April
in comparison with the rest of the year). The
four high values from the Grand Banks were
obtained during the same cruise.
However, five measurements from the Sargasso
Sea made by Atlantis during the same period
showed no evidence of a spring maximum in those
Table 1.
Some maximal values and efficiencies of primary production in coastal and oceanic waters.
Environment
Location
Reference
Date
Radia-
tion
Prod.
Method
g C/m 2 /
day
Eff.
(%)
Turtle grass
Coral reef
Long Key, Fla.
Japton Reef,
Eniwetok Atoll
Odum (1957)
Odum & Odum
(1955)
Aug., 1955
July, 1954
560*
660*
In situ
In situ
10.3
9.8
4.0
3.1
Spring flowering
(upwelling?)
Grand Banks
4440'N.,
4857'W.
Ryther & Yentsch
(unpub.)
Apr., 1957
380*** Chi
5.4
3.1
Polluted estuary
Forge River,
Moriches Bay,
L. I., N. Y.
Ryther & Yentsch
(unpub.)
Aug., 1956
400***
Chi
5.1
2.7
Upwelling
East Sound,
Washington
Yentsch
(unpub.)
July, 1954
700**
Chi
Oxy
5.1
4.8
1.6
1.5
Upwelling
Walvis Bay,
Angola
Steemann Nielsen
(1954)
Dec., 1950
430***
C"
3.8
1.9
Spring flowering
Cont. Shelf
off N. Y.
Ryther & Yentsch
(unpub.)
Apr., 1957
402***
Chi
C14
3.1
2.8
1.7
1.5
* From Kennedy (1949); ** measured; *** From Kimball (1928).
190
VOLUME 4 BOTANY
Table 2.
Magnitude and efficiency of primary production in selected coastal and oceanic waters.
_ j _
Mean
1
Situation
Location
No.
sta.
Reference Date
total
radiation
(g cal/
Prod,
method
Prod, (g
carbon/
m 2 /day)
Eff.
%
cm 2 /day)
Polluted
Great
8
Ryther &Yentsch
Aug., 1956
400
Chi
0.26
0.14
embayment
South
(unpub.)
Bay, L. I.,
N.Y.
Unpolluted
Long Is.
8
Riley (1956)
Mar.-May,1952
400
In situ 1.06
0.57
embayment
Sd., N.Y.
May-Aug., 1952
544
1.73
0.69
Aug.-Nov., 1952
310
i 1.22
0.85
jNov.-Feb., 1952-3
139
0.22
0.35
Feb.-Mar., 1953
252
1.33
1.12
Arctic
Allen Bay,
1
Apollonio,
July- Aug., 1956
302
Chi
0.19
0.14
embayment
Cornwallis
Ryther &
Is., N.W.T.
Yentsch
(unpub.)
Cont. Shelf
Off N.Y.
11
Ryther &
Sep., 1956
317
Chi
0.17
0.12
(25 m depth)
Yentsch
Dec., 1956
120
0.61
1.10
(unpub.)
Feb., 1957
223
0.60
0.58
Mar., 1957
327
0.96
0.64
Apr., 1957
402
1.25
0.68
Cont. Shelf
Off N.Y.
5
Ryther & Yentsch
Sep., 1956
317
Chi
0.14
0.10
(500 m depth)
(unpub.)
Dec., 1956
120
0.17
0.30
Feb., 1957
223
0.09
0.09
Mar., 1957
327
0.53
0.35
Apr., 1957
402
1.10
0.59
Cont. Shelf
N.Atlantic
4
Ryther & Yentsch
Apr., 1957
380
Chi
4.63
2.65
(unpub.)
S. Atlantic
2
Steemann Nielsen
Dec., 1950
476 O4
1.37
0.63
(Benguella
(1954)
Current)
i i
Oceanic
N.Atlantic
36
Ryther & Apr., 1957
380 Chi 1 0.47
0.27
i
Yentsch
,
(unpub.) :
Sargasso
5
Turner, Ryther
Apr., 1957
477 Chi
0.044
0.02
Sea
& Yentsch
i
(unpub.)
Sargasso
1
Steemann Nielsen
June, 1952
440
C14
0.048
0.02
Sea
(1954)
i
Caribbean
3
Ryther (unpub.)
Feb., 1955
387
Ci4 0.43
0.24
Pacific
{
(1105'S.,
1
Steemann Nielsen
Mar., 1952
454
O4 0.15
0.07
17l07'W.)
(1954)
(630'S.,
1
>
410
C" | 0.14
0.07
1642'W.)
(1957'N.,
1
^
460
C14
0.10
0.04
15810'W.)
(423'N.,
1
> *>
| 410
CU
0.26
0.14
16444'W.)
!
1
191
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
waters, the mean of 0.044 g carbon/m 2 /day
agreeing almost exactly with Steemann Nielsen's
values for the Sargasso Sea (0.048) obtained by
the C 14 value during June, 1952. Actually these
values varied by almost ten-fold from a minimum
of 0.009 to a maximum of 0.081.
The remaining oceanic values, from Ryther and
Steemann Nielsen, were obtained by the C 14
method and hence are probably somewhat lower
than would have been given by other methods.
In spite of this, these data indicate (as do the
majority of Steemann Nielsen's values not
included here) that oceanic production is consider-
ably lower than that of coastal and inshore
waters. However, this cannot be considered as a
final conclusion until substantiated by one or
more studies of the annual cycle of primary
production in a truly oceanic area.
Efficiencies of primary production of inshore
waters range from about 0.1 to 1.0 per cent,
again showing this range seasonally as well as
regionally. Efficiencies for oceanic production
are less certain. They appear to be lower, but
also vary by at least an order of magnitude,
ranging from 0.02 to 0.27% in the few data
presented here.
We may now summarize the rough averages of
photosynthetic efficiencies from all sources dis-
cussed above, from quantum-yield laboratory
experiments to the Sargasso Sea. Listed in
decreasing order of magnitude, they appear as
follows:
Quantum yield of photosynthesis 27.0
Indoor algal culture yields 15.0
Outdoor algal culture yields 3.0
Maximal marine values 2.2
Average coastal and inshore
waters 0.5
Average oceanic waters 0. 1
In the following discussion an attempt will be
made to review and evaluate the environmental
factors responsible for the decline of efficiency
from the top to the bottom of this list.
FACTORS AFFECTING PRIMARY
PRODUCTION AND ITS EFFICIENCY
RESPIRATORY Loss
In quantum efficiency experiments, photosyn-
thesis is corrected for the effects of respiration.
However, in the mass algal culture work described
above, the yield of organic matter represents the
net effect of both photosynthesis and respiration.
Similarly the C 14 method for estimating primary
production in the sea appears to measure photo-
192
synthesis minus respiration (see 20). The C 14
values given in Tables 1 and 2 which were ob-
tained by the author have not been corrected
for respiration while those of Steemann Nielsen
(25) have been increased by 4% to allow for res-
piratory loss. The other values reported above
were obtained by methods which correct for res-
piration, and this section does not apply to them.
Respiration is probably never less than 5-10%
of maximum sustained photosynthesis with light
and all other factors optimal. In an earlier
paper, the author showed the relationship be-
tween total daily photosynthesis within the
entire euphotic layer and the incident radiation
falling on the surface of the water (21). This
curve is reproduced in Fig. 1. If one assumes
that respiration is 7.5% of optimal photosyn-
thesis (taking the middle of the range suggested
above), the curve for total daily respiration
within the euphotic layer relative to photosynthe-
sis may be shown as a straight line (i.e., indepen-
dent of radiation), as in Fig. 1 . The ratio of res-
piration to photosynthesis, given as a percentage,
then represents the respiratory loss.
It may be seen that with an incident total
radiation of 100 g cal/cm 2 /day or less the plants
are at or below compensation. With the highest
radiation values reaching the surface of the earth,
respiratory losses are barely less than 30%, and
within the range normally encountered over most
of the earh (250-500 g cal/cm 2 /day), the respira-
tory loss lies between about 30% and 50%.
These losses apply only to plants within the
euphotic layer (the depth of penetration of
approximately 1 % of full sunlight). If part of
the plant population under consideration (wheth-
er an algal culture, a phytoplankton population,
or a bed or attached algae of rooted plants)
extends below this depth, these plants will impose
an additional respiratory loss to the system.
Thus in an unstable water column with a phyto-
plankton population which is wind-mixed to twice
the depth of the euphotic zone, net growth of the
population cannot occur if the incident radiation
is less than about 275 g cal/cm 2 /day.
Similarly a dense culture of Chlorella, although
brightly illuminated at its surface and supplied
with an excess of nutrients and CO 2 , may cease to
grow not because photosynthesis is stopped, but
because the light cannot penetrate to sufficient
depth to permit photosynthesis to compensate
for the respiratory loss of the culture as a whole.
The relationship between photosynthesis and
respiration shown in Fig. 1 is based upon natural
daylight conditions and therefore does not apply
VOLUME 4
BOTANY
200 JOO 400 500 600 TOO
TOTAi INCIDENT RADIATION (fl col / em f doy )
Fig. 1 . Total daily photosynthesis (P), respiration (R), and
the ratio of respiration to photosynthesis ( % respiratory
loss) of phytoplankton within the euphotic zone as a func-
tion of total daily incident radiation.
to cultures constantly illuminated by artificial
lights. Nevertheless it is clear that a large por-
tion of the cells of these cultures are always in
sub-optimal light if not in darkness, and it seems
reasonable to assume that the discrepancy
between the efficiency of yields of algal cultures
grown under artificial illumination (15%) and the
quantum efficiency of photosynthesis (27%) may
be due to this factor alone.
Similarly the C 14 values reported by the
author are only 1/2 to 3/4 as high as the true
values for total photosynthesis, and Steemann
Nielsen's values including a 4% correction for
respiration are probably also low.
REFLECTION Loss
In quantum yield experiments, care is taken to
consider the utilization of only that light which
is actually absorbed by the plants. This cannot be
easily determined in culture yield experiments or
in measurements of natural photosynthetic
rates, and it has sufficed for most workers to base
efficiencies on the utilization of the light falling
on a square unit of water surface. This introduces
a rather small but significant error in that a frac-
tion of the incident radiation is lost through
reflection from the surface or from back scatter-
ing out of the water.
Powell and Clarke (16) measured these losses
from the sea surface. At solar angles greater
than 30 they found that about 4% of the incident
light was lost through the combined effects of
reflection and back scattering on clear days,
about 6.5% on cloudy days. This appears to be
independent of the sea state, from flat calm to
winds strong enough to produce white caps.
Reflection losses increase greatly at solar angles
below 30, as predicted by theory, but as relatively
little absolute radiation reaches the earth's surface
at such low solar altitudes, these higher reflection
values may be discounted. It would appear,
therefore, that 5% is a reasonable average value
to assign to this loss.
Fig. 2.
A) The quantum requirement of photosynthesis as a func-
tion of wave length for Chlorella (after Emerson and
Lewis, 1943) and for Nitzschia (after Tanada 1951).
B) Energy per mole Quantum of light as a function of
wave length.
C) The efficiency of photosynthesis as a function of wave
length.
193
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
WAVE LENGTH EFFECT
It was pointed out that sustained photosyn-
thesis under carefully controlled laboratory
conditions requires 8-12 quanta of red light to
reduce one mole of CO 2 , corresponding to a
mean efficiency of 27%. These experiments
are usually carried out at or near a wave length of
680 mji, the maximum absorption peak for
chlorophyll.
Emerson and Lewis (4) and Tanada (26) have
studied the quantum requirement of photosyn-
thesis at different wave lengths using Chlorella
and Nitzschia respectively. Their curves are
reproduced in Fig. 2A. The differences between
the two species may not be significant as tech-
niques may have differed somewhat, but in each
case the minimum requirement falls within the
range given above, remains relatively constant
between about 680 and 550 m^i, increases to a
maximum at 490 mu, and then partially recovers
between 490 and 400 m|i. Above 685 mjj, the
quantum requirement increases markedly for
both species.
Figure 2B shows the energy per quantum of
light between 400 and 700 mji illustrating the
fact that this drops from a maximum of 71 k cal
per mole of quanta at the blue end to a minimum
of 41 k cal at the red end of the visible spectrum.
As red light is most efficient in terms of its
energy content per quantum and as the quantum
requirement of photosynthesis increases in blue
light, the efficiency of photosynthesis decreases
from its maximum (25-30%) in red light to a
minimum of 12.5% in blue light, as shown in
Fig. 2C.
The spectral distribution of daylight varies with
solar altitude and with the water vapor, car-
bon dioxide, and dust content of the atmos-
phere. Fig. 3 shows the spectral distribution of
daylight under average atmospheric conditions
with an air mass of 2 (solar angle=60) as given
by Moon (12). If the curves for the two organ-
isms shown in Fig. 2C are averaged and the mean
efficiency calculated for the entire visible spec-
trum, weighted for the spectral distribution of
sunlight as given in Fig. 3, this value turns out
to be 18.4%. Thus average solar radiation is
only about 68 % as efficient for photosynthesis as
red light of 680 mu.
In extremely turbid waters, particularly those
containing dissolved pigmented organic matter
(the so-called "yellow substance" of Kalle, 7)
blue and green light may be selectively absorbed
which would result in somewhat higher efficiencies
194
LENGTH ,
Fig. 3. The spectral distribution of daylight under average
atmospheric conditions with air mass-2 (solar angle=60).
6 8 10
INTENSITY (POOT CANDLES XlO* S )
Fig. 4.
A) Photosynthesis of marine phytoplankton as a func-
tion of light intensity (after Ryther, 1956a). Broken line
is the extrapolation of the linear portion of the solid
line, representing hypothetical sustained maximum pho-
tosynthetic efficiency.
B) Efficiency of photosynthesis as a function of light
intensity, as calculated from Figure 4 A, assuming a max-
imum efficiency of 100%.
VOLUME 4
BOTANY
in the utilization of the light which penetrates to
greater depths. However, in normal, clear
coastal and oceanic waters, red light is selectively
absorbed and scattered by the water, and the
blue and green penetrate to the greatest depths
where they would be used still less efficiently than
the incident daylight discussed above.
THE INTENSITY EFFECT
The exact mathematical relationship between
light intensity and photosynthesis is rather contro-
versial (see 27), but an examination of many
intensity-photosynthesis curves (as in 77, chapter
28) leaves little doubt that the relationship closely
approximates linearity up to the saturation inten-
sity. Above this, photosynthesis does not
increase further, and at intensities 1/4 to 1/3 of
full sunlight and above it may become severely
depressed (see 21).
As soon as the intensity-photosynthesis curve
departs from linearity, efficiency begins to
decrease. This may occur at intensities as low
as 1/20 of full sunlight.
The laboratory experiments described above
giving an 8-12 quantum requirement were all
carried out at low intensities on the linear portion
of the curve. If this linear portion is extrapo-
lated, one may then calculate efficiencies at any
intensity by comparing the extrapolated line with
the actual photosynthesis curve. This has been
done in Fig. 4A using the photosynthesis-intensity
curve for marine phytoplankton given by Ryther
(21). The broken line obtained by extrapolating
the linear portion of the curve, represents the
hypothetical sustained maximum rate of photo-
synthesis. The efficiency, obtained from the
00 300 400 500 00
TOTAL INCIDENT NAOIATION ( col/cm'/ay)
Fig. 5. Daily efficiency of photosynthesis by phytoplank-
ton within the euphotic zone as a function of total daily
radiation, assuming a maximum efficiency of 100%.
Fig. 6. The fraction of light absorbed by water as a func-
tion of the depth of the euphotic zone, defined as the
depth of penetration of 1 % of maximum sunlight incident
to the surface.
ratio of corresponding points on the two curves,
is shown in Fig. 4B, with the maximum efficiency
arbitrarily given a value of 100%.
From Fig. 4B and instantaneous values for
total incident radiation (corrected for reflection
loss), one may calculate photosynthetic efficien-
cies throughout the day, and these may be aver-
aged to give the mean daily efficiency. This,
however, will characterize only those organisms
at the surface of the water. The deeper, shaded
plants will operate at higher efficiencies. But as
light is absorbed exponentially, the greater part
of the incident radiation is absorbed at the
higher, less efficient intensities. To evaluate this
quantitatively it is necessary to calculate mean
daily efficiencies at several depths where the surface
intensity is decreased by known amounts, weight
each depth interval according to the amount of
radiation absorbed within that interval, and
integrate over the entire illuminated water
column.
This was done for eight days, each character-
ized by a different incident radiation ranging from
about 30 to 800 g cal/cm 2 /day. The resulting
curve showing the relationship between total
incident radiation and daily photosynthetic
efficiency for the antire phytoplankton popula-
tion is given in Fig. 5, assuming a maximum
efficiency of 100%.
According to these calculations, photosynthe-
tic efficiencies begin to decrease when the incident
radiation exceeds about 50 g cal/cm 2 /day, and
falls to 40% of its maximum on days of highest
radiation. Under normal conditions in tropical
and temperate regions, where average radiation
195
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
values fall between 250 and 500 g cal/cm 2 /day,
efficiencies are reduced to 70-50% of their
biological potential from the effects of light
intensity. Although these calculations are based
upon the photosynthesis-light intensity relation-
ships of marine phytoplankton, they probably
apply reasonably well to marine plants in general.
NUTRIENT DEFICIENCY
The availability of plant nutrients has a pro-
nounced effect upon the rate and efficiency of
photosynthesis, a fact with which plant physiolo-
gists and ecologists are equally well acquainted.
A single example is perhaps sufficient to illustrate
this point. In an earlier paper (20), an experi-
ment was described in which the marine flagellate,
Dunaliella euchlora, was grown in a culture
medium with a limiting supply of nitrogen and
phosphorus, the two elements which are probably
most frequently limiting to photosynthesis in the
sea. The results of this experiment are repro-
duced in Table 3. Exponential growth of the
flagellate occurred for the first five days after
which essentially the same cell concentration
persisted for the next twenty three days. Photo-
synthesis per cell decreased twenty-fold from its
maximum on the second day to its minimum on
the twenty-eighth day. Respiration decreased to
one-half its maximum during the same period.
The two processes were equal and the culture at
compensation level by the end of the experiments.
In nutrient deficient plants, particularly those
limited by nitrogen, the chlorophyll content of the
cells decreases markedly. If these cells absorbed
less light and the latter could thereby penetrate
to greater depths, photosynthetic efficiency per
Table 3.
Photosynthesis (P) and respiration (R)
in a nutrient-deficient pure culture of Dunaliella
euchlora expressed in ml O 2 /24 hrs.
(From Ryther, 1954.)
Days
Mean cell count
(10^ cells/liter)
IT
26
1- 2
61
2- 3
135
3- 4
290
4- 5
390
5- 6
470
6- 7
427
14-15
410
28-29 | 415
P/cell
113.0
121.0
18.3
10.5
10.1
9.5
7.5
6.0
R/cell
(xlO 9 )
13.4
10.0
8.6
5.9
5.7
6.3
5.9
6.1
unit of surface area would not be reduced to
the same extent as when calculated per unit
volume. The following section will have some
bearing on this question. Actually, however,
this is not the case because the concentration of
the carotenoid pigments decreases very slowly
in starved cells and their light absorptive charac-
teristics do not differ greatly from that of healthy
plants. This was determined by measuring the
light absorption of a suspension of Dunaliella
in the 10 cm light path of a Beckman DU
Spectrophotometer, averaging the percent trans-
mission at all wave lengths between 400 and 700
n\[i. This varied from 84% in a healthy, growing
culture to 88% in a nitrogen-starved culture
which had not grown for fourteen days.
Thus under the conditions of this experiment,
the efficiency of photosynthesis decreased to 5 %
of its maximum from the effects of nutrient defi-
ciency. If photosynthesis were measured by a
method which does not correct for respiration,
this effect would be still more striking as the
organisms at the end of the experiment, which
were at compensation level, would indicate no
photosynthesis whatever.
While it is perhaps doubtful that plants in this
extremely impoverished condition normally exist
in the ocean, nutrient deficiency is still probably
the most important single factor limiting the
efficiency of photosynthesis over most of the
marine environment as will be discussed later.
ABSORPTION OF LIGHT BY
WATER AND PARTICULATE MATTER
OTHER THAN PLANTS
Plants living in an aqueous medium are at
some disadvantage over terrestrial forms as some
of the incident radiation must be absorbed by the
water itself. According to Jerlov, the extinction
coefficient of pure sea water is .033. This loss is
negligible if the plants are located near the surface
and sufficiently concentrated so that the light is
all absorbed within a few meters. In a sparse
plankton population, however, as is typical of the
open sea, light may penetrate to much greater
depths and a considerable fraction of it is then
absorbed by the water rather than the plants.
Fig. 6 shows the percentage of the incident light
which is absorbed by the water as a function of
its transparency. From this curve it is obvious
that in such a place as the Sargasso Sea, where the
euphotic zone may exceed 100 meters, over 80%
of the light is absorbed by the water, and the
efficiency of the photosynthetic utilization of the
196
radiation incident to the sea surface is corres-
pondingly lowered.
This, however, cannot be considered as a direct
cause of decreased photosynthetic efficiency,
as a sparse plankton population itself merely
reflects the decrease or absence of photosynthesis
and plant growth which in turn must ultimately
result from a deficiency of plant nutrients or
light. It is mentioned here to point out the fallacy
in the concept that production per unit of surface
area should be comparable in dense and sparse
plankton populations, an idea based on the
assumption that the lower rate of photosynthesis
per unit volume in the latter case may be com-
pensated for by a deeper euphotic zone. This
obviously does not take into consideration ab-
sorption by the water as Steemann Nielsen (25)
has pointed out.
However, there is some justification for this
concept because in coastal or inshore waters with
a shallow euphotic layer, a large fraction of the
light may be absorbed by non-planktonic parti-
culate matter, either organic or inorganic in
nature and probably often associated with the
bottoms. Thus Riley, (19) taking the relationship
between the extinction coefficient and the chloro-
phyll content of offshore waters and plankton
blooms where light absorption by non-planktonic
particulate matter was assumed to be minimal,
calculated that an average of only 1/3 of the light
incident to Long Island Sound is absorbed by
plants, the remainder being attributed to absorp-
tion by other particulate matter.
Additional light may be absorbed in coastal
waters by dissolved pigmented material (i.e.,
the "yellow substance" of Kalle, 7) which prob-
ably consists of organic compounds of terrige-
nous origin and are of relatively little significance
on the open sea. Finally in extremely shallow
areas, such as Great South Bay with a mean depth
of about 2 meters, light may penetrate to and be
absorbed by the bottom.
These various factors probably explain why
production and efficiency values are so similar
in Great South Bay, Long Island Sound, and the
continental shelf area, regions where light absorp-
tion by the water, non-planktonic particulate and
dissolved matter, and the bottom, although of
different relative importance in each case, may
add up to nearly the same total effect.
While light absorption by water was mentioned
above as an indirect cause of lowered efficiency,
symptomatic of other deficiencies, such is not the
case in the absorption of light by particulate and
dissolved matter which are quite independent
_VOLUME 4 BOTANY
of and in competition with the plants for the
available light.
DISCUSSION
This discussion is based upon the premise,
described in the first section of this paper,
that the maximum sustained efficiency of photo-
synthesis is represented by the conversion of
about 27% of the light absorbed by the plants,
this light being at or near 680 mji and at an
intensity well below the saturation level. Turning
from these ideal laboratory conditions to those
existing in nature, and considering the utilization
of sunlight falling on the sea surface by the
underlying plants, we have seen that wave lengths
and intensities are no longer optimal, that light is
lost by reflection and back scattering, by absorp-
tion by the water itself, by dissolved or particu-
late matter other than plants, or by the bottom.
Some of these factors are relatively constant,
others may vary greatly, but all combine to lower
efficiencies of natural photosynthesis to a small
fraction of the "normal quantum yield" as it is
known to the plant physiologist.
Let us consider, for instance, the maximum
possible efficiency in nature by marine plants.
Under the best of conditions, there is still a 5%
loss of light by reflection and back scattering, a
wave length effect resulting in efficiencies some
32% below that observed in red light, and an
intensity effect, depending upon the length and
brightness of the day, reducing efficiencies 30-
50% below that possible in sub-saturation inten-
sities. The accumulated effect of these factors is
to decrease efficiencies 67-87% of the maximum
potential of 27% or to a level ranging from
3.5% to 9.0%.
Are these low values reasonable? They may
be checked by re-examining the organic yields of
carefully conducted outdoor mass algal culture
experiments. Here efficiencies were found to
average about 2% by the Netherlands group,
about 4% by the Japaness investigators. These
yields, however, were not corrected for respira-
tory losses which, we find, could hardly be less
than 30% and may reach 100% in extremely
dense cultures. If we take 50% as an average
respiratory loss, the efficiency of photosynthesis
in these mass cultures would be 4-6%, or roughly
the same as that calculated above.
Are these efficiencies encountered in the sea?
The data presented in Table 1 shows that they may
be attained, or at least closely approached in a
wide variety of different marine environments.
197
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
These, however, all have one feature in common,
a sufficiency of nutrients. The flowing-water,
benthic population appear to live under the most
optimal conditions, and the reasons for this are
obvious -they live at the bottom of a shallow,
clear column of water which removes very little
of the light, and they are bathed in a constantly
replenished medium from which the nutrients
can never become exhausted.
Similarly polluted regions are characterized by
the constant addition of nutrients from an
external source. Here, however, the existence of
dissolved or particulate substances competing
with the plants for light is almost a certainty,
as pollution by definition is of terrestrial origin
and highly productive polluted areas are virtually
confined to harbors, estuaries, etc., which nor-
mally contain considerable amounts of such
substances. Furthermore, most sources of
pollution also contribute substantial quantities of
light absorbing material (i.e., organic detritus,
bacteria, silt, dyes, etc.) to the water in addition
to the essential plant nutrients. Therefore it is
unlikely that such an environment would sustain
production at efficiencies as high as those possible
in the flowing-water benthic communities.
More favorable conditions for photosynthesis
may exist in coastal and offshore areas where the
nutrients originate through upwelling or mixing
processes from the rich, deep ocean waters which
are relatively free of light absorbing substances.
However, high production rates under such
circumstances arc, for the most part, brief and
transitory, either in time or space. Although high
production occurs at or near an area of upwelling,
it quickly diminishes as the upwelled water
spreads over the ocean surface. Production may
remain high at a given geographical area, but it
is not sustained in a given volume of water or by
a given population of plants. Similarly, most
plankton blooms or flowerings are temporary,
usually representing the culmination of mixing
or regeneration processes during a period when
plant growth can not occur (e.g., during temperate
or polar winters). With the return of favorable
conditions for photosynthesis, the temporarily
enriched waters may support a brief and dramatic
period of extremely high production.
According to Red field ( 18), the relative propor-
tion of carbon, nitrogen, and phosphorus in
plankton is 100:15:1 by atoms. A production
rate of 5 grams of carbon per day within a 10
meter water column (such high production
rates result in and are therefore restricted to an
extremely shallow euphotic zone) would therefore
198
require 6 ugA of nitrogen and 0.4 ugA of phos-
phorus per liter per day. There is no deep water
in the ocean rich enough to meet this demand
for more than a few days.
Thus efficiencies of 3-9%, representing the
maximum photosynthetic potential under natural
illumination, are rarely encountered in the ocean.
The average efficiencies reported in Table 2 for
coastal and offshore waters are, for the most
part, one to two orders of magnitude below this.
Two variable environmental factors have been
discussed, both of which decrease efficiencies
below this maximum potential. These are (1)
light absorption by dissolved and particulate
matter other than plants, and (2) nutrient deficien-
cies. The former undoubtedly plays an impor-
tant part in reducing efficiencies in coastal waters.
As mentioned earlier, Riley (19) has estimated
that only one-third of the light entering Long
Island Sound is absorbed by plants. Similar
calculations by the author have indicated that
anywhere from 27% to 66% of the light pene-
trating the waters of the continental shelf is uti-
lized by the phytoplankton, the remainder being
about equally divided between other particulate
matter and the water itself.
It seems doubtful, however, that this factor
can be important in the clear waters of the open
sea, and the conclusion seems inevitable that the
greater part of the oceans is normally nutrients
deficient. Unfortunately relatively little is known
about the concentrations of all the essential
plant nutrients in the surface water of the ocean;
their rates of uptake by the phytoplankton,
limiting concentrations for photosynthesis, and
rates of regeneration by mixing and decomposi-
tion. Consequently the relative importance of this
factor cannot be quantitatively assessed. How-
ever, the conclusion that nutrient deficiencies are
responsible for the low efficiencies (compared to
the potential efficiency of 3% or more) normally
found in the sea is based upon the following bits
of indirect evidence or reasoning: (1) such
nutrient concentrations as have been measured
in the surface waters of the open sea, mainly N
and P compounds, are normally extremely low,
often undetectable. (2) Sustained maximum
efficiencies such as are possible under natural
illumination would require the constant replen-
ishment of nutrients whose only source is the
ocean depths. This water cannot be mixed
upward into the surface layers without, at the
same time, carrying the plants down out of the
euphotic zone. In most of the oceans, thermal
stratification is an effective barrier against such
mixing. (3) The only marine areas where this
maximum potential efficiency is approached,
temporarily or permanently, are those situations
in which some mechanism exists for the enrich-
ment or continued replenishment of nutrients.
(4) No other environmental factor appears to
exist which could decrease photosynthetic efficien-
cies to a comparable degree.
REFERENCES
(1) Clarke, G. L., and James, H.R., 1939, Labo-
ratory Analysis of the Selective Absorp-
tion of Light by Sea Water, J. Optical
Soc.Amer., 29: 43-55.
(2) Daniels, F., 1956, Energy Efficiency in Pho-
tosynthesis, Chapt. 4 in Hollaender,
A., Radiation Biology, Volume III:
Visible and Near Visible Light, McGraw-
Hill, New York.
(3) Emerson, R., and Lewis, C.M., 1941, Carbon
dioxide Exchange and the Measurement
of the Quantum Yield of Photosynthesis,
Am.J. Bot., 28:789-804.
(4) _ _. , 1943, The
Dependence of the Quantum Yield of
Chlorella Photosynthesis on Wave
Length of Light, Ibid., 30: 165-178.
(5) Gaffron, H., 1954, Mechanism of Photosyn-
thesis, in Autotrophic Microorganisms,
4th Symp. Soc. Gen. Microbiol., The
University Press, Cambridge.
(6) Hill, R., and Whittingham, C. P., 1955, Pho-
tosynthesis, Methuen and Co., London.
(7) Kalle, K., 1938, Zum Probleme der Meeres-
wasserforbe, Ann, Hydrog. u. mar.
Meteor., 66.
(8) Kennedy, R. E., 1949, Computation of
Daily Insolation Energy, Am. Meteor.
Soc. Bull, 30: 208-213.
(9) Ketchum, B.H., and Redfield, A. C., 1949,
Some Physical and Chemical Character-
istics of Algae Grown in Mass Culture,
J. Cell, and Comp. Physiol., 33: 281-300.
(JO) Kimball, H. H., 1928, Amount of Solar
Radiation that Reaches the Surface of
the Earth on the Land and on the Sea,
and Methods by which It is Measured,
Mon. Weath. Rev. Wash., 56: 393-398.
(11) Kok, B., 1952, On the Efficiency ofChhrella
Growth, Acta Botanica Neerl, 1: 445-
467.
VOLUME 4 BOTANY
(12) Moon, P., 1940, Proposed Standard Solar-
Radiation Curves for Engineering Use,
J. Franklin lnst. 9 230: 583-617.
(13) Morimura, Y., Nikei, T., and Sasa, T., 1955,
Outdoor Bubbling Culture of Some
Unicellular Algae, /. Gen. Appl. Micro-
Mo/., 1: 173-189.
(14) Odum, H. T., 1957, Primary Production
Measurements in Eleven Florida Springs
and a Marine Turtle-Grass Community,
Limnol. and Oceanogr., 2: 85-97.
(15) Odum, H.T., and Odum, E.P., 1955, Tro-
phic Structure and Productivity of a
Windward Coral Reef Community on
Eniwetok Atoll, Ecol. Monogr., 25: 291-
320.
(16) Powell, W.M., and Clarke, G.L., 1936, The
Reflection and Absorption of Daylight
at the Surface of the Ocean, /. Optical
Soc. Am., 26: 1-23.
(17) Rabinowitch, E. L, 1951, Photosynthesis
and Related Processes, Interscience
Publ., II, Part 1, New York.
(18) Redfield, A. C., 1934, On the Proportions of
Organic Derivatives in Sea water and
Their Relation to the Composition of
Plankton, James Johns tone Memorial
Vol., Univ. of Liver pool, 176-192.
(19) Riley, G. A., 1956, Oceanography of Long
Island Sound, 1952-1954, II: Physical
Oceanography, Bull. Bing. Oceanogr.
Coll., 15: 15-46.
(20) Ryther,J.H., 1954, The Ratio of Photosyn-
thesis to Respiration in Marine Plankton
Algae and Its Effect upon the Measure-
ment of Productivity, Deep sea Res., 2:
134-139.
(21) , 1956a, Photosynthesis in the
Ocean as a Function of Light Intensity,
Limnol. and Oceanogr., 1:61-70.
(22) - - , 1956b, The Measurement of
Primary Production, Ibid., 72-84.
(23) Ryther, J.H., and Yentsch, C.S., 1957, The
Estimation of Phytoplankton Produc-
tion in the Ocean from Chlorophyll and
Light Data, Limnol. and Oceanogr., 2.
(24) Steemann Nielsen, E., 1952, The Use of
Radio-active Carbon (C 14 ) for Measur-
ing Organic Production in the Sea, J.
Cons. Internat. Explor. Mer. 9 18: 1 17-140.
(25) _ _ - , 1 954, On Organic Pro-
duction in the Oceans, Ibid., 19: 309-328.
199
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
(26) Tanada, T., 1951, The Photosynthetic Effi-
ciency of Carotenoid Pigments in Navi-
cula minima, Am. J. Bot., 39: 276-283.
(27 ) Van Oorschot, J.L.P., 1955, Conversion of
Light Energy in Algal Culture, Mededel-
ingen van de Landbouwhogerschool te
Waginingen, Nederland, 55: 225-276.
(28) Warburg, O., and Burk, D., 1950, The
Maximum Efficiency of Photosynthesis,
Arch. Biochem., 25: 410-443.
(29) Wassink, E. C, 1954, Problems in the Mass
Cultivation of Photo-autotrophic Micro-
organisms, in, Autotrophic micro-
organisms, 4th Symp. Soc. Gen. Micro-
bioi, The University Press, Cambridge.
(30) Wassink, E.C., Kok, B., and Van Oorschot,
J.L.P., 1953, The Efficiency of Light
Energy Conversion in Chlorella Cultures
as Compared with Higher Plants, in,
Burlew, J. S. : Algal Culture, from
Laboratory to Pilot Plant, Carnegie Inst.
Wash., PubL, 600.
200
_ VOLUME 4 BOTANY
COMPARISON OF ALGAL FLORISTIC PATTERNS IN
THE PACIFIC WITH THOSE IN THE ATLANTIC AND
INDIAN OCEANS, WITH SPECIAL REFERENCE TO CODIUM
PAUL C. SILVA
Department of Botany, University of Illinois, Urbana, Illinois, U.S.A.
Codiwn (order Codiales or Siphonales of the
green algae) is a relatively large genus (that is,
compared with other genera of marine algae),
comprising about eighty species. It is exceed-
ingly widely distributed, growing on all tropical
and temperate shores with the notable exception
of the Atlantic coast of North America north of
Beaufort, North Carolina 1 . With the cooperation
of scores of workers throughout the world,
during the past eight years, I have been able to
assemble about 2,000 fixed and dried collections.
In attempting to work out a taxonomic treatment
of Codium, I began to perceive certain geographic-
al distributional patterns. It then became im-
portant to determine whether a standard of
reasonableness could be established against which
to judge geographical distributions as a test
of taxonomic soundness. It was of interest to
see whether there was justification for being
suspicious about the alleged occurrence of the
same species, for example, in the Irish Sea and
on the California coast. The first step was to
become aware of previously described distribu-
tional patterns for seaweeds; the second step was
to compare these patterns with those that are
exhibited by Codiwn.
The field of phytogeography is vast, the litera-
ture inexhaustible, its data capable of a variety
of interpretation, extraordinary manipulation,
and distortion. It has the fascination of a chess
game. It is a valid, though treacherous, field of
investigation. While making no claim to com-
petency as a phytogeographer, 1 believe that
information divulged by my studies of Codium
warrants evaluation in general phytogeographic
terms.
CURRENT PATTERNS
It has long been recognized that along most
coasts temperature is probably the most important
factor determining the latitudinal distribution
of marine algae a view particularly expressed
by Setchell (2, 3, 4), who believed that the
effect of temperature on reproductive processes
was critical. Ideally, ocean surface temperatures
could be expected to vary linearly from the
equator toward the poles; however, it is well
known that the major oceanic currents profound-
ly alter this gradient. Zones of algal distribu-
tion thus do not coincide with latitudinal zones.
Figure 1 is a schematic representation (modi-
fied Mercator projection) of an idealized current
system, wherein it is assumed that both sides of
the ocean are delimited by continuous land
masses bearing due north/south. The pattern of
currents coincides in many regions with the
pattern of prevailing winds, but it depends not
only upon the prevailing winds but upon the
rotation of the earth and density differences in
the surface layers. The pattern in the northern
hemisphere has a mirror image in the southern
hemisphere. The following features may be
noted: northern and southern equatorial currents,
equatorial countercurrent, northern and southern
high-latitude currents, northern and southern
high-latitude minor gyrals. Whether a current as
it bathes a particular shore should be designated
"warm" or "cold" depends upon the temperature
that might be expected in the absence of a
current system. Under the idealized conditions
of Fig. 1, the western shores of the ocean would
be warmed by the equatorial currents while the
eastern shores would be cooled by the returning
high-latitude currents. Actually, this idealized
pattern is significantly altered by at least three
factors: (1) the configuration of major land
masses, affecting the major current system;
(2) the configuration of minor land masses, affect-
ing local gyrals; and (3) upwelling, causing local
temperature disturbances.
Figure 2 shows the patterns of currents in the
Pacific. Because of the funnel formed between
Asia and North America, the North Equatorial
Current warms the Japanese coast (as the Kuro-
shio Current) and its high-latitude extension
retains enough warmth, despite the admixture
of cold subarctic water south of the Aleutian
Islands, to be regarded as a warm current when it
1 Bouck and Morgan (J) report the discovery at East Marion, Long Island, New York, of Codium fragile &$$. tomentosoides
(van Goor) Silva, a weed presumably introduced from Europe, where it is becoming increasingly widespread.
201
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
'' '
202
'/>'' L'^??^ 1 * repre f ntati / on of idea . lized current s V stem (modified Mercator projection), a, a' t equatorial currents;
, b , high-latitude currents ; c, c', equatorial countercurrent ; d t </', high-latitude minor gyrals
VOLUME 4
BOTANY
reaches the coast of North America. The north-
ward moving minor branch, the Alaska Current,
is definitely warming; the southward moving
major branch, the California Current, largely
because of the upwelling of cold water soon
reaches a point where it no longer has the effect
of warming, but of cooling. To the northeast
of Japan, temperatures are depressed by the
western part of the Bering Sea gyral (the Oyashio
Current). In the southern hemisphere, the cir-
cumpolar current of the antarctic region sets the
stage for a very different pattern. The continent
of South America extends far south and deflects
a part of the eastward moving cold subantarctic
circumpolar current northward. The resulting
Peru (or Humboldt) Current is further affected
by upwelling, so that it has a chilling effect nearly
to the equator. In the western part of the South
Pacific, there is no current system as well defined
as the Kuroshio system. This may be related
to the great distance separating South America
and^Australia. According to Sverdrup, Johnson,
and Fleming (6, p. 706), "in the western South
Pacific annual variations are so great that in
many regions the direction of flow becomes
reversed, as is the case off the east coast of
Australia."
Figure 3 shows the current patterns in the
Atlantic. In the North Atlantic the Gulf Stream
system, a counterpart of the Kuroshio system in
the Pacific, is well known for its role in warming
the shores of both North America (the south-
eastern part of the United States) and Europe
(the British Isles and Scandinavia). Among the
major terminal branches of this system, the
Irminger Current flows westward to the south of
Iceland while the Norwegian Current can be
traced through two further branches into the
Polar Sea. To the east of Labrador and New-
foundland, the western part of a counter-clock-
wise gyral, the cold Labrador Current, is a counter-
part of the Oyashio Current in the Pacific. Part
of the North Atlantic Current returns southward
off the Iberian and North African coasts and
60 80 100 120 140 160 180 160 140 120 100 80 60 40 20
ENDEMIC SPP.
PACIFIC OCEAN
SCALE Of MILES
<F ~ sb6~"~fooo
80 100 utud 120^ frrt of 140 0****h 160 160 160 tanjtt*. 140 **< of 12Q o^>w<h 100
60 <0
NvtTnoM Scmu or OCM MA
L J NVtTMM A CO.
Fig. 2. Current system and centers of distribution of endemic species of Codium in Pacific Ocean.! Each circle represents
one species.
203
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
A. J NYSTMOM & Co . CHICAGO
NVSTROM SCRIES or DESK MAPS NQ. DD 20
Fig. 3. Current system and centers of distribution of endemic species of Codiwn in Atlantic Ocean. Each circle repre-
sents one species.
904
contributes in part to the North Equatorial
Current and in part to the Guinea Current,
which flows along the coast of Africa as far as
the equator. In the southern hemisphere, the
shape and position of South America profoundly
affects the current pattern: the northeastern
shoulder of Brazil deflects part of the South
Equatorial Current northward across the equator
where it flows northwest along the northern
coast of South America and ultimately contri-
butes to the Florida Current (the first part of the
Gulf Stream system). In the Atlantic, unlike the
Pacific, the surface circulation of the two hemis-
pheres is thus interconnected. The southern tip
of South America deflects the cold subantarctic
circumpolar current not only along the west
coast, but also along the east coast. The resulting
Falkland Current meets the southward moving
branch of the South Equatorial Current (the
VOLUME 4 BOTANY
warm Brazil Current) in the latitude of Uruguay,
where a sharp temperature gradient is thus estab-
lished. Along the west coast of South Africa,
the Benguela Current flows northward; and
being affected by upwelling, it depresses inshore
temperatures until it gradually leaves the coast
and continues westward as the South Equatorial
Current.
Much less is known about the current patterns
of the Indian Ocean (Fig. 4) than of the Pacific
or Atlantic. A major counter-clockwise gyral
in the southern part of the Indian Ocean prevails,
subject to considerable annual variation. The
South Equatorial Current flows southward around
Madagascar and along the cast coast of South
Africa as the warm Agulhas Current. There is a
well-defined eastward moving current between
South Africa and Australia, representing the
returning high-latitude component of the gyral.
ENDEMIC SPP.
INDIAN OCEAN
No 00 22
Fig. 4.--Current system and centers of distribution of endemic species of Codium in Indian Ocean. Each circle represents
one species.
205
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
FLORIST1C PATTERNS
On the basis of considerations that temperature
controls the establishment of populations of
particular genotypes and that currents provide a
means of dispersal, we can make predictions con-
cerning the expected distribution of marine algae.
In seeking explanations of existing floristic pat-
terns, however, we must also consider the historical
factor, despite the difficulty of assessing its role.
A more recent origin is suggested by taxa whose
distribution can be explained without recourse to
the historical factor, while such problems as
bipolar distribution imply greater antiquity.
In the Pacific, on the basis of the Kuroshio
system, we can predict similarities between the
flora of Japan and that of California. In the At-
lantic, on the basis of the Gulf Stream system,
we can predict similarities between the flora of
the northeastern United States and Atlantic
Europe. Considering the interconnected surface
circulation between the northern and southern
hemispheres of the Atlantic, we might also predict
similarities between the flora of Brazil and that
of the Caribbean region. On the basis of the
Antarctic Circumpolar Current we can predict
the existence of a circumpolar antarctic or sub-
antarctic marine algal province. As we shall see,
each of the four predictions is borne out by data
from marine algae in general.
The temperate flora of Japan, although it
comprises endemic species for the greater part
and includes a sizable arctic element, is clearly
related to the California flora. This relationship
is expressed in similarity of general appearance
and in the sharing of many genera and even some
species (including Heterochordaria abietina, End-
arachne binghamiae, Costaria costata, Cymathaere
triplicata, Pikea calif arnica, Baylesia plumosa,
Rhodymenia pertusa, Coeloseira pacifica, Bing-
hamia californica and Pterosiphonia dipinnatd).
The similarities between the flora of the north-
eastern United States and that of Atlantic Europe
have long been recognized. Among common
species shared by the two floras, the following
may be cited: Ulothrix flacca, Prasiola stipitata,
Cladophora albida, C. flexilis and C. gracilis,
Bryopsis plumosa, Ectocarpus confer voides, E.
fasciculatus and E. tomentosus, Giffordia granulosa
Cladostephus verticillatus, Sphacelaria drrhosa,
Elachistafucicola, Chordariaflagelliformis, Eudes-
me virescens, Sphaerotrichia divaricata, Stilo-
phora rhizodes, Arthrocladia villosa, Desmarestia
aculeata and D. viridis, Stictyosiphon tortilis,
Asperococcus echinatus, Desmotrichum undulatum,
206
Punctaria plantaginea, Dictyosiphon foeniculaceus,
Alarm esculent a, Chorda filum and C. tomentosa,
Laminaria digitata and L. saccharina, Ascophyl-
lum nodosum, Fucus serratus, F. spiralis and F.
vesiculosus, Nemalion multifidum, Gelidium crinale,
Dumontia incrassata, Polyides caprinus, Corallina
officinalis, Gloiosiphonia capillaris, Cystoclonium
purpureum, Furcellaria fastigiata, Gymnogongrus
griffithsiae and G. norvegicus, Phyllophora brodiaei
and P. membrani folia, Chondrus crispus, Gigartina
stellata, Rhodymenia palmata, Champia parvula,
Antithamnion cruciatum, Callithamnion byssoides,
C. corymbosum and C. roseum, Ceramium diapha-
num, C. rub rum and C. strictum, Plumaria elegans,
Ptilota plumosa, Seirospora griffithsiana, Sper-
mothamnion turneri, Spyridia filamentosa, Mem-
branoptera alata, Pantoneura angustissima, Phy-
codrys rubens, Dasya pedicellata, Chondria dasy-
phylla, Odonthalia dentata, Polysiphonia elongata,
P. lanosa, P. nigrescens and P. urceolata, Rhodo-
mela confervoides and R. lycopodioides.
In searching for similarities between the flora
of Brazil and that of the Caribbean region, we
encounter a distributional pattern that is remark-
able in that it encompasses portions of both
northern and southern hemispheres. This vast,
rather homogeneous algal province extends from
the Caribbean (with some northward extensions
to Florida, the Carolinas, and Bermuda) to
southern Brazil and shows certain affinities with
the flora of the Cape Verde Islands and the
adjacent African coast and weaker affinities with
the flora of the Canary Islands and of the western
Mediterranean. Taylor (8) in a study of 317
species common in the Caribbean region, found
that 28.4 % were restricted to the Caribbean, 1 1 .0 %
extended northward at least to Beaufort, North
Carolina, 33.1% extended southward along the
coast of Brazil, and 27.5% extended both north-
ward and southward. Taylor concludes that
"the Caribbean flora deserves this name only
because the Caribbean Sea is the area of its great-
est known luxuriance and diversity .... [It] is
actually a west or American Atlantic tropical
flora which, in spite of the Brazil current and the
North Equatorial current, extends down the
Brazilian coast to Rio de Janeiro, with few
replacements." While the direction of the North
Equatorial Current would indeed seem to prevent
or at least to impede the southward spread of a
Caribbean flora, if the possibility of an eastern
Atlantic origin be granted, the existence of this
vast algal province would not be in disagreement
with hydrographic facts.
The circumpolar marine algal province en-
compassing the Antarctic Continent and the
Subantarctic region (including the southern tip of
South America and such islands as the South
Orkneys, South Georgia, South Sandwich, Ker-
guelen, Heard, Macquarie, Auckland, Campbell,
and Stewart) has been summarized by Skottsberg
(5). Characteristic species include Cladophora
pacifica, Adenocystis utricularis, Caepidium antarc-
ticum, Desmarestia rossii and D. willii, Durvillea
antarctica, Geminocarpus geminatus, Halopteris
funicularis, Macrocystis pynfera, Scytothamnus
australis and S.fasciculatus, Utriculidium durvillei,
Balha callitricha, Callophyllis tenera, Chaetangium
fastigiatum, Delisea pukhra, Lithothamnium an-
tarcticum, Lophurella hookeriana, Phycodrys quer-
cifolia, Plocamium hookeri and P. secundatum, and
Polysiphonia microcarpa.
DISTRIBUTIONAL PATTERNS
OF CODIUM
To what extent does the distribution of Codium
agree with previously recognized floristic patterns ?
VOLUME 4 BOTANY
First, it should be pointed out that most species
of Codium are restricted to continuous shores or
closely spaced islands; only a few species occur
on two or more widely separated land masses.
Figs. 2, 3, and 4, indicate the centers of distri-
bution of endemic species in each of the three
oceans. The Pacific with 30 endemic species
leads the Indian with 19 and the Atlantic with 8.
These numbers undoubtedly will be adjusted
upward with further study of certain perplexing
material, as yet unidentified.
Let us look more closely at the Pacific Ocean.
Fig. 5 shows the distribution of four closely
related adherent species. The occurrence of
C. hubbsii Dawson both in Japan and in California,
although separate subspecies probably are in-
volved, should be noted. Codium convolutum
(Dellow) Silva 2 comb. nov. is an example of a
local endemic. On the other hand, C. lucasii
Setchell occurs in South Africa as well as in Austra-
lia. Codium arabicum Kuetzing is interpreted here
as a complex series of microspecies or subspecies
60 80 " 100~ 120 140 160
180 160 140 120 100
COOIUM ARABICUM
C LUCASI!
C. CONVOLUTUM
O C HUBBSII
80 100 lonptudt 12Q fU o( 140 Cwenw.ch 160 180 160 Longitude 14Q Wst of l^Q Gwn^h 1UU HO
J NTSII.OM * Co CM.CAOO
Fig. 5. Distribution of four closely related adherent species of Codium.
2 Codium adhaerens C.Ag. var. convulotum Dellow, Trans. R. Soc. N.Z. 80: 122. 1952.
207
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
extensively distributed throughout the Indo-Paci-
fic region, reaching its northeastern limit in the
Hawaiian Islands. Several interesting patterns
are shown in Fig. 6. Codium spongiosum occurs
in South Africa and Mauritius in addition to the
localities indicated in Fig. 6 and is thus Indo-
Pacific in distribution. The distribution of each
of the three members of the highly specialized
section Digitaliformia is shown: Codium john-
stonei Silva occurs in California and adjacent
Mexico, but lacks a Japanese counterpart; C.
pomoides J. Ag. is restricted to Australia; C.
dimorphum Svedelius occurs in New Zealand and
southern Chile. The distribution of each of five
species of the section Mamilhsa is also shown:
C. minus (Schmidt) Silva 3 comb. nov. occurs in
Japan; its nearest relative, C. mamillosum Har-
vey, occurs in Australia and Hawaii; C. rittcri
S. and G. occurs in Alaska; the remaining two
species are distinct from the other members of
the section and from each other; yet are very
closely related and are extremely local in distri-
bution, C. globosum Lucas occurring in Queen-
sland and C. cranwelliae Setchell in New Zealand.
Fig. 7 shows the distribution of several dicho-
tomous species. Codium contractum Kjellman in
Japan and its very close relative C. macdougalii
Dawson in the Gulf of California, Mexico, are
excellent examples of vicarious species. The
distribution of the three members of the section
Lata is interesting: Codium latum Suringar,
described from Japan, is represented by a sub-
species, C. latum ssp. palmer 7 (Dawson) Silva 4
comb, nov., on Guadalupe Island, Mexico; C.
laminar ioides Harvey is endemic to Western
Australia; while C. platylobium J.E. Areschoug
(not shown in Fig. 7) is endemic to South Africa.
Fig. 8 shows the distribution of various repent
Codium, representing several species or complexes
of microspecies and subspecies. This group of
taxa together with the populations of C. arabicum
indicated in Fig. 5 and C. intertextum Collins and
60 80 100 120 140 IfaO 180 160 140 120 100 80 60 40 20
A CODIUM SPONGIOSUM
0D S DIGITALIFORMIA
O@ S MAMILLOSA
120 . East of 140 Grttnwtch 160
160 Longitude 140 Wtst of 120 Gnnw.ch 100
__ Fig. 6. Distribution of Codium spongiosum, of Section Digitaliformia, and of Section Mamilhsa.
3 Codium mamillosum Harvey var. minus O.C. Schmidt, Bibl. Bot., 23(91): 37, 1923.
4 Codium palmeri Dawson, Bull. So. Calif. Acad. Sc. % 44: 23, 1945.
208
VOLUME 4
BOTANY
Hervey indicated in Fig. 9 form an integral part of
the biocoenosis of coral reefs. The anatomical
variability of these plants from reef to reef is
perplexing.
Despite the high degree of endemism exhibited
by Codlum, a study of the distribution of closely
related species is thus seen to support the recog-
nition of both the Japanese-Californian floristic
pattern and the circumpolar subantarctic pattern.
Moreover, data are provided which support the
recognition of an Indo-Pacific province, the
existence of which has already been indicated by
studies of other algae. This floristic pattern,
while not strongly suggested by current pat-
terns, is not unreasonable if migration through
the Indonesian region is considered possible.
Finally, an affinity between the floras of eastern
South Africa and southwestern Australia is
suggested.
Now let us turn to the Atlantic Ocean. Fig.
9 shows the distribution of two closely related
adherent species, C. adhaerens C. Ag. and C.
intertextum Coll. and Herv. The populations of
C. intertextum in the Canary Islands probably
should be accorded subspecific recognition. Fig.
10 shows the distribution of another pair of close-
ly related adherent species, C. effusum (Raf.)
Delle Chiaje and an undescribed species from
Tierra d*el Fuego and the Falkland Islands (C.
antarcticum Silva). This remarkable distribution
suggests the divergence of a widespread ancestral
stock into a warm-water and a cold-water species.
The absence of a closely related species in the
Caribbean region is notable. Fig. 11 shows the
distribution of a pair of very closely related
globose species, C. bursa (L.) C. Ag. and C.
elisabethae Schmidt, known only from the
Azores. Again it is interesting to speculate on
the absence of C. bursa or a closely related species
in the Caribbean flora. Fig. 12 shows the distri-
bution of three fairly closely related dichotomous
species, C. vermilara (Olivi) Delle Chiaje, C
isthmocladum Vickers, and an undescribed species
from the Guinea Coast (already in literature as
C. guineense Silva). The last two species are seen
to occupy mutually exclusive parts of the tropical
60 80 100 120'" 140 ~ 160 180 160 140 120 100 80 " "60 40
CODIUM LATUM
C. LAMINARIOIDES
C CONTRACTUM
C MACOOUGALII
80 100 Longnud* 1?Q En o< 140 &w>w.<h 160 180 160 lon t .tuO 140 Weu_* 120 G'tn.>. C h 100
/
60 40
" No DO 21
Fig. 7. Distribution of two pairs of closely related dichotomous species of Codium.
209
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Atlantic province. The bipolar distribution of
C. vermilara is unique and suggests a more
extensive distribution in former times. Its absence
in North America is notable. Fig. 13 shows the
distribution of three distantly related dichoto-
mous species. Both C. taylori Silva and C.
decorticatum (Woodw.) Howe occur throughout
the tropical Atlantic province, although the former
is absent in the western Mediterranean. Lastly,
C. tomentosum Stackh., a name at one time applied
to almost any dichotomous Codium, is seen to
have a highly restricted distribution.
It thus can be seen that the temperate North
Atlantic province is not supported by data from
Codium because of the absence of this genus on
the northeastern coast of North America. On
the other hand, full support is given to the recog-
nition of the tropical Atlantic province. The
bipolar distribution of C. vermilara is notable.
CONCLUSIONS
In comparing algal floristic patterns in the
Pacific with those in the Atlantic and Indian
oceans, we find that the Japanese-Californian
alliance is the counterpart of the North Atlantic
province, but the degree of similarity between
the Japanese and Californian floras does not
warrant their recognition as a province. The
Japanese-Californian alliance is supported by
data from Codium, the North Atlantic province,
while lacking such support, is nonetheless well
documented by distributional data from other
algae. The tropical Atlantic province has as its
Pacific counterpart the Indo-Pacific region, which
not only spans the equator but extends over two
oceans. Unlike its Atlantic counterpart, it does not
have affinities with eastern shores; its boundaries
seem to coincide with the limits of coral reefs, and
its flora may be an integral part of the coral reef
biocoenosis. While the components of the flora
of this vast region are not known sufficiently to
warrant its designation as a province, it seems
likely that even after various subregions with
clusters of endemic species have been removed,
there will remain a large area with a fairly homo-
geneous flora. Recognition of both the tropical
80 100 1ZQ 140 160
60 140 120 100 60 60 40 20
No 00 21
Fig. 8. Distribution of repent Codium.
A ) NviTHOM A CO CMICAO
210
VOLUME 4
BOTANY
CODIUM ADHAERENS
C. INTERTEXTUM
No. DD 20
Fig. 9. Distribution of two closely related adherent species of Codium.
211
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
CODIUM EFFUSUM
C. ANTARCTICUM
NVITPOM States or DESK
No. DO 20
A J NYSTKOM 4 Co . CHICAOO
212
Fig. 10. Distribution of two closely related adherent species of Codium.
VOLUME 4
BOTANY
CODIUM BURSA
O C. EL1SABETHAE
No. DO 20
A J NrsritoM A Co CHICAGO
Fig. 1 1 . Distribution of two very closely related globose species of Codiutn.
213
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
CODIUM VERMILARA
C. ISTHMOCLADUM
O C. GUINEENSE.
OF DCS* MAPS N O DD 20 A J NVSTROM A CO.. CMICAOO
Fig. 12. Distribution of three fairly closely related dichotomous species of Codium.
VOLUME 4
BOTANY
O CODIUM TAYLORI
C. DECORTICATUM
C. TOMENTOSUM
N O QQ 20
Fig. 13. Distribution of three distantly related dichotomous species of Codiitm.
215
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Atlantic province and the Indo Pacific region is
strongly supported by distributional data from
Codium. There are no southern hemisphere
counterparts of the North Atlantic province
and the Japanese-Californian alliance. The
circumpolar subantarctic province is well docu-
mented, including support from Codium. Its
northern counterpart, the arctic province, al-
though fully documented (7) has not been consi-
dered in this paper because of the absence of
Codium in that part of the world and because
the existence of this province is dependent upon
climatic and geographic isolation of a mediter-
ranean-type sea rather than upon general oceanic
circulation.
REFERENCE
(1) Bouck,G. B.,and Morgan, E., 1957, The Oc-
currence of Codium in Long Island Water,
Bull. Torrey Bot. Club, 84: 384-387.
(2) Setchell, W.A., 1893, On the Classification
and Geographical Distribution of the
Laminariaceae, Trans. Conn. Acad. Arts
Sr., 9: 333-375.
(3)
(4)
__. , 19 15, The Law of Temperature
Connected with the Distribution of the
Marine Algae, Ann. Missouri Bot.
Card., 2 :287-305.
, 1920, The Temperature Int-
erval in the Geographical Distribution of
Marine Algae, Science. 52 : 187-190.
(5) Skottsberg, C, 1941, Communities of Marine
Algae in Subantarctic and Antarctic
Waters, K. Svenska Vetensk-Akad.
Handl., ser. 3, 19(4), 92 pp.
(6) Sverdrup,H.U., Johnson, M.W., and Fleming,
R.H., 1942, The Oceans: Their Physics,
Chemistry, and General Biology. New
York, Prentice-Hall, 1087 pp.
(7) Taylor, W.R., 1954, Cryptogamic Flora of the
Arctic, II. Algae: Non-planktonic, Bot.
/fry., 20: 363-399.
(8) - - ,1 955, Marine Algal Flora of the
Caribbean Sea and Its Extension into
Neighboring Seas, in Essays in the Natu-
ral Sciences in Honor of Captain Allan
Hancock. Los Angeles, Univ. So.. Calif.
Press, pp. 259-270.
DISCUSS/ON
M.S. DOTY: Does Codium arabicum occur in southern
Japan?
p.c. SILVA: Quite possibly.
G.F. PAPENFUSS: Very few algae are common to North-
west Africa and the Caribbean. West Indian species are
more likely to be found on the east coast of Africa.
p.c. SILVA: There arc marked temperature differences
between the coast of Northwest Africa and the Caribbean.
F.R. FOSBERG: Does Dr. Silva think of these distribu-
tional patterns of algae as related to currents? Do they
indicate historical patterns of plant geography, or are they
related to physiological requirements of the plants?
p.c. SILVA: The historical factor is very important but
difficult to assess. A large part of the genus Codium
seems to be actively speciating although many species arc
widely separated morphologically and would appear to be
relicts. The distribution of recent species at least can be
related to present-day current patterns.
216
VOLUME 4
BOTANY
DKTWBjTIONAL RELATIONSHPS OF
MALAYAN FRESHWATER ALGAE
G.A. PROWSE
Fish Culture Research Station, Batu Berendam, Malacca, Federation of Malaya.
One of the difficulties encountered when read-
ing past papers on the algae of this part of the
world is that the areas concerned are not clearly
defined. Political boundaries rarely coincide with
biological ones, and the term Malaya has been
used to cover anything from the whole of the
Indonesian islands to the Federation of Malaya
itself. I propose to use it to cover the Federation
and Singapore, not through any wish to ignore
the surrounding areas, but because the very
location of the research station makes it necessary
that I should confine my main attentions within
those boundaries.
Looking through the literature on fresh-water
algae, one finds that while there are many papers
dealing with Malay territories generally, very few
of them deal with the Federation of Malaya and
Singapore. We have papers by West and West,
Bernard, Lemmermann, Biswas, and Ruth
Patrick, but very little else. This is partly because
in the past botanists in this part of the world
have been more concerned with the flowering
plants and ferns, and it is only recently that any
serious phycological studies have been undertaken.
With the active interest taken in fresh-water
biology by departments of the University of
Malaya, and with the opening of the Fish Culture
Research Station in Malacca, phycological knowl-
edge in this area has been considerably advanced,
although still very inadequate. It is noteworthy
that the present Acting-Director of the Botanic
Gardens, Singapore is deeply interested in phy-
cology, admittedly marine, but still very encourag-
ing to fresh-water phycologists.
In a survey of the fresh-water algae of Malaya,
one might expect certain things, simply by virtue
of its geography. Firstly it would occasion no
surprise to find those species which are world-
wide in distribution, ranging from arctic to tropic
climates. Superimposed on these distributional
elements would be found those species which are
strictly tropical in range, but still circumglobal
in distribution. Of greater interest, however,
are those species of much more restricted range.
The Malay peninsula forms an integral part of a
geographical system which stretches from Burma
and Thailand in the north right down through
the Indonesian islands to Australia. This Indo-
Malaysian-Australian system was evidently
almost a continuous land mass at one time. One
would expect algal species common to the whole
area, with perhaps minor geographical variations.
In addition, in Malaya one would expect species
having greater affinity with those from countries
in the north, such as India, Burma, and Thailand,
and other species showing greater similarity to
those from Java and Sumatra. Finally it is likely
that a few species may be even more restricted in
range, being confined almost entirely to Malaya.
Such distributional patterns will inevitably be
complicated by the nature of the terrian of the
country concerned, for the various habitats may
differ widely in character. Much of the lowland
of Malaya is lateritic, with swamps and padi
fields. The water is decidedly acid and desmids
abound, whilst the common flagellates are various
species of Chrysophyceae. In many places fish
ponds have been made, with consequent liming
and heavy manuring. Euglenineae are dominant,
or with less heavy manuring, Chlorococcales and
sometimes Microcystis aeruginosa. On the other
hand, in the fast flowing streams of the high
mountains, such as the Cameron Highlands,
diatoms predominate. The limestone peaks of
the north have not yet been explored phycolog-
ically, and should prove to have an interesting
algal flora. In addition, Malaya is divided cen-
trally by the main mountain range, and it would
be interesting to see if those algal species on the
west side differ in any way from those on the east.
Despite these distributional complications,
it should be possible to trace the broad lines due
to geological and evolutionary history, particular-
ly if individual groups are studied. Taking the
very characteristic desmid genus Micrasterias,
we find those species of known world-wide dis-
tribution A/, pinnatifida, americana (only once
recorded in Malaya so far), alata, crux-melitensis,
thoinasicma var. notata and jenneri. Typical
circumglobal tropical species or varieties are M.
mahabuleshwarensis and M. foliacea var. ornata.
In the Indo-Malaysian-Australian group we have
M. ceylanica and M. moebii, although the Mala-
yan form of the latter species is intermediate
217
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
between those from Thailand and Java. M.
thomasiana var. evoluta and M. apiculata var.
tjitjeroekensis show greater affinity for forms
from Java, while M. mahabuleshwarensis var.
bengalica, M. lux and M. anomala are similar to
those from India and Burma. So far, M. torreyi
var. crameri and M. torreyi var. doveri appear
to be confined to Malaya itself. Further inves-
tigations may show that some of these species
are more widespread than at first apparent, but
the list illustrates the fact that Malaya partakes
of elements from both northwards and south-
wards, as well as possessing tropical and world-
wide elements. Other desmid genera show a
similar picture, often just as clearly.
The diatoms of Malaya are not fully worked
out, and we have insufficient representative
collections. For that reason, the picture is less
clear, particularly as so many diatoms are world-
wide in distribution. Examples of these latter to
be found in Malaya are Cyclotella meneghiniana,
Melosira granulata, Synedra ulna, Eunotia pecti-
nalis, Frustulia vulgar is and Pinnularia braunil.
So far I have found none which I would call
purely tropical species, but of much more limited
distribution are Stenopterobia pelagica and Ach-
nanthes crenulata, found also in Indonesia, and
Navicula voigtii occurring also in Thailand and
S. China. Ruth Patrick mentions one or two
species which she found only in Malaya, and I
have found some as yet unnamed ones which
apparently have not been reported elsewhere,
but on such limited collections it would be unwise
to suggest that they are confined to Malaya.
When we turn to the various flagellate groups,
we find a wealth of forms of world-wide distri-
bution also found in Malaya, or else species which
are recorded for Malaya and for places like
Germany and France, which suggests they are
really more widespread. Of the purely tropical
forms, Phacus pyrum is the most typical, but I
would not like to be certain about this species,
for with the flagellates it is often easier to say
which are the species which will not tolerate
tropical conditions rather than vice versa. Of
more limited distribution are Chlamydomonas
lismorensis, Trachelomonas lismorensis, Trachelo-
monas volzii var. cylindracea, and Peridinium
gatunense var. zonatum, all found in both Malaya
and Australia, while Stromhomonas australica is
reported from Australia and S. China as well
as Malaya. It is to be expected that these species
will be reported from Indonesia as well, Trachelo-
monas similis var. hyalina, a very characteristic
species was described from Burma and has proved
quite common in Malaya. No doubt as more
work on the flagellates is done in the various
countries, the picture will become clearer. There
are several species reported from Indonesia and
Venezuela, and now from Malaya, and it seems
possible that they may occur in other tropical
fresh waters.
It will be noticed that most of the examples
I have chosen have well-marked external features,
so that any differences become obvious. No
doubt if such groups as the Myxophyceae and
the Chlorococcales were examined closely, it
would be found that they show similar distribu-
tional patterns.
To sum up, the algal species of Malaya exhibit
an interaction of worldwide distributional range,
circumglobal tropical distribution, and a much
more limited distribution within the Indo-Malay-
sian-Australian area.
DISCUSSION
R.L. CROCKER: You have mentioned geological and
historical evolutionary trends. Have you any theories
on this?
G.A. PROWSE: With the limited amount of information
available, it would be unwise to formulate any theories.
O.F. PAPENFUSS: What literary sources are used in
your work and would Skufa's paper on Burmese algae be
useful to you ?
O.A. PROWSE: I have consulted the various libraries
in Britain and have had photostats made. Good papers
are available on Indonesia, Burma, and elsewhere, but
very little on Malaya itself. I have a photostats copy of
Skufa's paper.
G.F. pAPfcNFUss: Looking at canals near Bangkok, 1
would suggest that the Volvocales are common. Is that
so in Malaya?
O.A. PROWSE: So far they have been comparatively
scarce, but possibly may be more frequent in the limestone
218
VOLUME 4
BOTANY
THE RHODOPHYTA ORDER ACROCHAETIALES AND
ITS CLASSIFICATION
JEAN FELDMANN
Laboratoire de Biologie Vegdtale Marine, Facttlte de Sciences de Paris, Paris, France.
The Acrochaetium-Rhodochorton complex, as
Papenfuss has termed it ( 11 ), groups many little,
mostly epiphytic red algae, distributed in all the
oceans. These algae are generally included in
the order Nemalionales, among which they show
a more simple structure than the typical forms
such as Nemalion and Liagora. The vegetative
fronds consist only of branched filaments with
heterotrichous habit.
As I pointed out some years ago (3), this com-
plex may be distinguished from all other Florideae
(except perhaps from Gelidiales) because it Jacks
a true carpogonial branch. In all the Acrochae-
tiaceae where sexual organs are known, the
carpogonium is borne directly on the side of a
normal cell of the filament or at the top of a few-
celled lateral vegetative branch; sometimes, as
in Crania, the carpogonium may be intercalary by
differentiation of a vegetative cell from the erect
filaments.
The lack of a carpogonial branch and the ex-
treme simplicity of the vegetative frond seem to
me sufficient reasons to put these algae in a
separate order, the Acrochaetiales. Should this
order be considered as the more primitive of the
Florideae or as a reduced group ? 1 am unable to
settle this definitely.
The generic classification of the Acrochaetiales
has for a long time puzzled the phycologists. The
late Mrs. Drew-Baker (I) united all the species
in only one genus, the genus Rhodochorton. More
recently Papenfuss (11 ) proposed a new and much
more natural classification. He chose, as the
main distinctive character, the structure of the
chromatophore and its situation in the cell.
Unfortunately, shortly before the paper of Papen-
fuss appeared, Kylin (5) had proposed another
classification of the Acrochaetiaceae based upon
morphological characters not as good for a sys-
tematic classification as those used by Papenfuss.
Kylin proposed, namely, to place into the genus
Kylinia, created by Rosen vinge (13) for a micro-
scopic species epiphytic on Sporochnus in Den-
mark, all the Acrochaetium species with a single
basal cell. This character does not seem to me
a good one. It would be best, perhaps, to take
into consideration the heterotrichy of these
plants and, if one would use morphological
characters for the distinction of genera, to separate
plants truly heterotrichous from those with only
erect system originating from a single basal cell
and those with only creeping filaments. But
I agree with Papenfuss in considering the type
of chromatophore as a more important criterion
for separating genera.
Kylinia rosulata, type of genus, was described
by Kylin as having a stellate chromatophore with
a central pyrcnoid, consequently Papenfuss (12)
put into this genus all the species previously placed
by him in his genus Chromastrum characterised
by its stellate chromatophore. In fact, the plant
studied by Kylin as Kylinia rosulata is quite
different from the species described by Rosen-
vinge. Both species exist on the coast of France
in the Channel. The species of Kylin is an
Acrochaetium (Acrochaetium ky Unhides nov.
sp.) with stellate chromatophore and with only
asexual monospores. The true Kylinia rosulata pos-
sesses a parietal chromatophore without pyren-
oid and its sexual organs are quite different from
those of Acrochaetium. The male cells (sperma-
tocysta) are borne at the top of a very long
hyaline cell that looks like a hair; and Kylin
thought that Rosenvinge made a mistake, inter-
preting a normal hair with accidentally attached
spermatium at the top for the male organs of
Kylinia. In fact Rosenvinge was right, as I have
ascertained by the study of living material.
The carpogonium of Kylinia arises generally
directly from the spore; and it is, like the vege-
tative cells, appressed on the filament of the host,
but the trichogyne is bent upwards. The plant is
monoecious; and after the fecondation, the
carpogonium produces directly three carpospores
at its top. Kylinia rosulata is thus the simplest
and probably the most reduced type of Florideae
one can imagine.
It is particularly noteworthy to recall the
observations of Levring (7) about a new Kylinia
he described from Australia, the structure and
reproduction of which closely recall those of the
European species.
The carposporophyte of the other Acrochae-
tiales is not so reduced, and there are some dif-
219
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
ferences in its development according to the
different genera. In Acrochaethium (A. borneti,
A. subtilissimum), the carpogonium is divided,
after the fecondation, into three or four cells;
each of them produces lateral branches bearing
terminal solitary carpospores. In Crania (G.
effhrescens), the development is the same but the
carpospores are in chains.
On the contrary, in Audouinella, as studied by
Drew (2), the carpogonium does not divide after
the fecondation but produces directly lateral
filaments with the carpospores terminal. By
this character, the fresh-water genus Audouinella
can be distinguished from the marine genus
Crania.
As to the characters of the chromatophores,
we can distinguish different types among Acro-
chaetiales:
(1) Only one chromatophore per cell; with a
central pyrenoid. The shape and the
situation of the chromatophore may be
variable.
a) The chromatophore may be axial in
the cell with the branches of the chro-
matophore radiating towards the inner
surface of the cell in all directions (stel-
late chromatophore of the genus Chro-
mastrurn of Papenfuss).
b) The chromatophore may be parietal
against the cell wall (genus Acrochae-
tium s. str. of Papenfuss). This parietal
chromatophore may be rounded or
irregularly lobed, sometimes stellate,
but only in one plane.
In fact, there arc intermediate forms
between (a) and (b), and it is difficult,
I think, to use this character for a
generic distinction. There will be some
species, the place of which among
Acrochaetium or Chromastrum of Papen-
fuss, will be difficult to settle.
In some species of this group with
only parietal chromatophore per cell,
the pyrenoid is lacking as in Kylinia
rosulata.
(2) More than one (few to many) parietal
chromatophores in each cell.
a) Chromatophores stellate with pyrenoid.
This is the case for Rhodochorton flori-
dulum but also for plants up to now
included in the genus Acrochaetium.
(A. caespitosum^ A.codii, etc.)
b) Chromatophores without pyrenoid,
disc-shaped or more or less spirally
twisted. This is the case for Rhodo-
chorton rothii (=R. purpureum) and for
the genera Audouinella and Crania.
Considering the mode of development of the
gonimoblast and the characters of the chroma-
tophores as well as the probably haplobiontic or
diplobiontic nature of the life-cycle, one can
divide the Acrochaetiales into two families and
eight genera according to the following key:
ACROCHAETIACEAE
Haplobiontic. Only one chromatophore with
a pyrenoid in each cell. In some species the
pyrenoid is lacking; in others, there are more than
one chromatophore per cell, but each is provided
with a pyrenoid.
A. Only one chromatophore (central or parietal)
per cell.
a) Spermatocysta borne on vegetative un-
differentiated cells, gonimoblast with spo-
rogenous filaments bearing terminal car-
pospores.
(1) Carpogonium transversally divided
after fertilization. . . . Acrochaetium
Naeg. (incl. Colaconema Batt.)
(2) Carpogonium longitudinally divided
after fertilization .... Liagoraphila
Yamada.
b) Spermatocysta borne at the top of dif-
ferentiated elongated and hyaline cells.
(1) Carpogonium transversally divided
after fertilization and producing spor-
ogenous filaments with terminal car-
pospores .... Balbiania Sirodot.
(2) Carpogonium undivided after ferti-
lization directly producing few car-
pospores .... Kylinia Rosenvinge.
B. More than one parietal chromatophore per
cell, each provided with a pyrenoid ....
Rhodothamniella nov. gen.
AUDOUINELLACEAE nov.fam.
Probably diplobiontic (or without sexual re-
production). More than one chromatophore,
always without pyrenoid, in each cell.
A. Sexual organs present. Chromatophores
ribbon-shaped and more or less spirally
twisted,
a) Marine Algae. Carpogonium tranversely
divided after fertilization. Carpospores
in chains .... Crania Kylin.
VOLUME 4
BOTANY
b) Fresh-water algae. Carpogonium un-
divided after the fertilization. Terminal
carpospores .... Audoulnella Bory.
B. Sexual organs unknown. Chromatophores
disc-shaped .... Rhodochorton Naeg. char.
mut.
From a morphological point of view, the two
families I propose to distinguish among Acrochae-
tiales, are very alike as to their vegetative
structure. But they may not be so closely allied
as they seem. Cytologically, Acrochaetiaceae
have a less evolved (more primitive) type of
chromatophore than the Audouinellaceae. Some
species of this family (Audouinella) recall the
Chantransia-stagQ of Batrachospermutn or Lema-
nea and may be reduced forms.
Most of our knowledge about Acrochaetiales
rests upon researches performed on the coasts
of the Atlantic Ocean. In the Pacific Ocean,
important work was done upon the species from
North America by K.M. Drew (1) and upon the
species from Japan by Nakamura (9,10). More
recently, some very interesting new species were
described from Australia and New Zealand by
Levring (7,8), but Acrochaetiales of large areas
still remain unknown.
It is hoped that in the future, accurate studies
on the morphology as well as the cytology, re-
production, and life-cycle of Pacific species will
afford new light upon this most interesting group
of red algae.
REFERENCES
(1) Drew, K.M., 1928, A Revision of the
Genera Chantransia, Rhodochorton and
Acrochaetium, etc. Univ. Calif. Publ. Bot.,
14: 139-124.
(2) _ , 1935, The Life-History of
Rhodochorton violaceum (Klitz.), comb.
nov. (Chantransia violacea Ktitz.), Ann.
of Dot., 49:439-450.
(3) Feldmann,!., 1953, devolution desOrganes
Femelles Chez les Florid^es, Proceed.
Int. Seaweed Symposium 1952, Edin-
burgh, 11-12.
(4) Hamel, G., 1927,Recherches sur les Genres
Acrochaetium Naeg. et Rhodochorton
Naeg., Saint-L6.
(5) Kylin, H., 1944, Die Rhodophyceen der
Schwedischen Westkiiste, Lunds Univ.
Arsskr. N.F., Avd 2, Bd. 40, Nr 2.
(6) ^ , 1956, Die Gattungen der Rho-
dophyceen, Lund.
(7) Levring, T., 1953, The Marine Algae of Aus-
tralia, I. Rhodophyta: Goniotrichales,
Bangiales and Nemalionales, Arkiv for
Bot.. Sen 2, Bd 2, Nr 6.
(8) -
,1955, Contributions to the Ma-
rine Algae of New Zealand, I, Arkiv for
Bot., Ser. 2, Bd 3, Nr 11.
(9) Nakamura, Y., 1941, The Species of Rhodo-
chorton from Japan, I, Sc. papers Inst.
Algol. Research, 2:273-291.
(10) - ._ _, 1944, The Species Rhodo-
chorton from Japan, II, ibidem, 3 :99-l\9.
(11) PapenfussG.F., 1945, Review of Acrochae-
tium- Rhodochorton Complex of the Red
Algae, Univ. of Calif. Publ. Bot., 18:229-
334.
(12) ._ _... _. ,1947, Further Contributions
toward an Understanding of the Aero-
chaetium- Rhodochorton Complex of the
Red Algae, Univ. of Calif. Publ. Bot.,
18:433-447.
(13) Rosenvinge, L.K., 1909, The Marine Algae
of Denmark, I: Rhodophyceae, D. Kgl.
Danske Vidensk. Sehk. Skrifter. Nat.
ogMath. Afd., 7, Raekkel(\)\\-\5\.
221
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
SOME OBSERVATIONS~ON~""~
LAMINARIA GAMETOPHYTES AND SPOROPHYTESt
JUN TOKIDA and HIROSHI YABU
Faculty of Fisheries, Hokkaido University, Japan.
The northwestern part of the Pacific Ocean,
especially around the northernmost island of the
Japan Archipelago, or Hokkaido, and the Kurile
Islands, has been known to be remarkably rich
in number of genera and species of Laminariales
since Dr. K. Miyabe, teacher of Tokida, pub-
lished in 1902 a comprehensive enumeration of
the kelps in Hokkaido (3). An English edition
of this paper has recently been prepared by To-
kida and published by Hokkaido University as
one volume of the Journal of Faculty of Agricul-
ture, Hokkaido University (4). In this paper,
Dr. Miyabe enumerated eight genera, of which
the genus Laminaria was the largest containing
as many as twelve species. However, these
species are not always considered clearly distin-
guishable from one another by some researchers
such as Dr. K. Okamura and Dr. S. Ueda, who
have sometimes expressed their views on the
basis of their field observations. Nevertheless,
Miyabe's species have an important practical
meaning in Hokkaido where most Laminaria
species are very highly estimated for their econo-
mic value. Their market value and the scope of
their use differ remarkably from one another,
each species representing usually one specific
locality or habitat. Their local distinctions are
well reflected in Miyabe's species.
In recent years, an attempt to transplant a
Laminaria of superior quality from its home
locality to other places has been practiced ex-
perimentally in Hokkaido. In connection with
this, there arose a question whether such a tran-
splanted Laminaria was really quite distinct or
not from other related species of inferior quality.
If they were nothing but local or ecological forms
of one and the same species, the transplanted
Laminaria would not be expected to retain its
superior quality long enough. On the contrary,
if it was really a distinct species peculiar to one
district, its characters would be expected to per-
sist to some extent. This question may possibly
be solved not only by the transplanting experiment
itself, but also by a crossing experiment between
certain two species.
Since 1952, cultural experiments of five species
of Laminaria, viz., L.japonica Aresch., L. ochoten-
sis Miyabe, L. rellgiosa Miyabe, L. diabolica
Miyabe, and L. angustata Kjellm., and one species
of Alaria, viz., A. crassifolia Kjellm., have been
carried out as follows by us with the object of
knowing the results of crossing in either couple
of these species, following the method and
principle of the study reported by Schreiber about
twenty years ago, in 1930 (5).
Zoospores were liberated in autumn or early
winter. A dilute zoospore suspension was
pipetted into small Petri dishes containing fil-
tered sea water. These dishes were placed on the
table in the room of our laboratory which was
heated in the daytime by steam from the middle
of November through the middle of April. The
gametophytes developed from those zoospores
remained sterile in such a condition and grew
within one to two months to minute filamentous
thalli visible to the naked eye. The growth of
gametophytes was more rapid in those floating
on the water surface than in those attached to the
bottom of the dish.
Such a filamentous thallus was then picked up
with a fine platinum needle or a pointed pincette,
and, after determining under the microscope its
sex by the size of cells, it was placed in a small
dish or tube containing Schreiber's culture
solution.
A preliminary cultural experiment of Laminaria
religiosa under various conditions of temperature
was carried out from early December, 1952 to
early February, 1953. The cultural tubes were
placed in the flasks filled with water and the
water of each flask was kept respectively at
8C, 12C, 16C, and 20C constantly all day
long, and at 20C in the daytime only. Other
tubes were placed on the table of the room heated
in the daytime as mentioned above. The growth
of gametophytes was best of all in the tubes
kept at 20C only in the daytime and in those at
room temperature, whereas it was worst in those
at constant 20C. The isolated gametophytes
cultured in the tubes at room temperature have
survived already more than three and one-half
years and have proved invariably sterile. They
usually have a minute spherical thallus consisting
t Presented in abstract by G.F. Parpenfuss.
222
VOLUME 4
BOTANY
of monosiphonous branched filaments. A piece of
filament taken from these thalli can be used at
any time as material for a culture experiment,
and it can grow to a new spherical thallus. On
the other hand, the gametophytes became fertile
only when cultured at 8C and 12C, bearing
sexual organs about 25 days and 30 days after
the start of culture, respectively, (cf. Fig. 1).
The eggs thus formed on isolated female
gametophytes of all the species studied by us
could germinate and develop parthenogenetically.
The development of those parthenosporophytes
was in general more or less irregular and resulted
in the production of variously shaped malformed
thalli as illustrated in Figs. 2 and 3. Malformed
sporophytes of similar structure were once re-
ported by Kanda (2) to occur in a mixed culture
of Lamlnaria ochotensis Miyabe (2, Fig. 4, 9-10),
L. cichorioides Miyabe (Fig. 7, 2-1 1) and L. sacha-
linensis Miyabe (Fig. 10, 4). In view of the very
common occurrence of parthenogenesis in our
culture experiments, resulting in the production
of malformed thalli and the occasional produc-
tion of malformed sporophytes in Kanda's mixed
culture, the presence in nature of parthenoge-
netically developed malformed haploid embryos
of kelps are supposed to be not improbable.
However, malformed adult thalli of kelps, such as
lately reported from Hokkaido and vicinity (cf.
/, 6, 7, 8) are not only of very rare occurrence
but also are considered not to have been derived
from such embryos as mentioned above. Such
haploid embryos of kelps may possibly be of a
limited life span.
Crossing experiments were carried out by
putting in a tube a female or male gametophyte
of one species together with a male or female of
another which had been cultured in isolation.
From twenty to twenty-two tubes were prepared
Fig. 1 . Growth rate of gametophytes of Laminaria religiosa as indicated by the number of body cells and time of the first
appearance of reproductive organs (denoted by x) in relation to the water temperature.
223
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Fig. 2. Showing various forms of malformed parthenosporophytes of Laminar ia religiosa in various stages of development.
(Magnification: x 200, x 170, x 95, and x 20.)
Z24
VOLUME 4
BOTANY
Fig. 3. Showing various forms of malformed parthenosporophytes of the following four species of Laminaria. 1-5,
Lamina riajaponica;6-S,L. ochotensis;9, L. angustata\ 10-11, L.diabolica (Magnification: x 200, x 170, x 95, and x 20).
225
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
226
VOLUME 4
BOTANY
Fig. 5. Microphotographs of embryonal Laminar/a sporophytes produced in the following six crossing experiments.
A, Laminaria japonica 9 X L. ochotensis c?, x 50; B, Laminaria religiosa 9 X L.japonica <, x 50; C, Laminaria ochotensis 9
X L. religiosa cT, x 80; D, Laminaria religiosa 9 X L. ochotensis <J, x 80; E, Laminaria angustata 9 X L. religiosa #, x 50;
F, Laminaria ochotensis 9 X Alaria crassifolia <j, x 120.
Fig. 4. Microphotographs of Laminaria gametophytes in various stages of development. A-H, J-N, R, U, Laminaria
religiosa', A, one month old gametophytes cultured at constant 20C; B, twenty-five days old female gametophytes bearing
eggs in a culture kept at constant 8C; C, sixty days old gametophytes in a culture kept at 20C only in the daytime; D,
seventy-seven days old female gametophyte bearing a parthenosporophyte, floating on the surface of cultural solution; E,
thirty days old gametophytes cultured at constant 16C; F, seventy-seven days old female gametophyte floating on the
surface; G, seventy-seven days old female gametophyte attached to the bottom of the cultural dish; H, seven months old
male gametophyte floating on the surface; J-N, five stages of development of one and the same piece of filament which was
first taken at the stage shown in J from an immature female gametophyte floating on the surface; K, two months old; L,
two and one half months old; M, three months old; N, four and one half months old; R, an immature female gametophyte
cultured in isolation, 3.7 months old; U, a mature female gametophyte grown from a thallus similar to that which is illus-
trated in R after placed in a newly prepared Schreiber's solution three weeks before. I, Laminaria japonica, three months old
gametophyte floating on the surface. O-P, Laminaria angustata; O, female and male gametophytes in a nine days old
culture, at this stage of growth, isolation of gametophytes was done; P, a female gametophyte isolated 121 days before.
Q,T, Laminaria diabolical Q, a female gametophyte bearing parthenosporophytes, isolated 78 days before; T, a female
gametophyte bearing parthenosporophytes, 68 days after isolation. S, Laminaria ochotensis, a female gametophyte 3.5
months old, bearing abundant parthenosporophytes. (Magnification: A, E-H, J-M, O-R, U x 80; B x 320; C-D, I, N x 40;
S-Tx 120).
227
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
for each crossing pair. The result of the experi-
ments are summarized in Table 1.
Table 1.
Result of crossing experiments, showing the
combination of species crossed and the structure,
whether normal or abnormal, of sporophytes
produced.
Couple of species crossed
Structur
phytes
Normal
eofsporo-
produced
Abnormal
L. japonica & L. religiosa
L. japonica & L. ochotensis
L. religiosa & L. ochotensis
L. religiosa & L. angustata
L. ochotensis & L. angustata
L. japonica & A. crass if olia
L. religiosa & A. crass if olia
L. ochotensis & A. crass if olia
+
+
+
+
4-
+
-4-
-f
The mark + in the Column "Normal" of Table
1 denotes that a larger number of the sporophytes
produced in the corresponding couple of species
are normal in shape. The parent species of these
normal sporophytes are supposed to have achieved
interspecific fertilization. Such a positive result
of a crossing experiment in Laminaria is believed
to be new to science, provided that the three
species concerned are all valid. Or, this result
may be taken rather as an evidence of specific
identity of the three species. However, in our
present state of knowledge, any conclusive remark
is to be reserved until more decisive data, cytolo-
gical and ecological, are acquired.
REFERENCES
(I) Hasegawa, Y. et #/., 1956, Notes on the Ab-
normal Laminaria Fronds Found in Hok-
kaido and Aomori Pref. (In Japanese),
Monthly Rept. Hokkaido Fish. Exp.
St., 13: 69-70.
(2) Kanda, T., 1946, Culture Studies of Lamina-
riaceous Plants in Hokkaido, (In Ja-
panese), Jour. Inst. Res. Sc. Fish.,
Hakodate Coll. Fish., No. 1, 1-44.
(3) Miyabe, K., 1902, The Laminariaceae of
Hokkaido, (In Japanese), Rept. Invest.
Mar. Resour., Hokkaido, 3: 1-60.
(4) , 1957, The Laminariaceae of
Hokkaido (1902). An English edition,
translated by J. Tokida, Jour. Fac. Agr.
Hokk. Univ., 1, (In the press).
(5) Schreibcr, E., 1930, Untersuchungen iiber
Parthenogenesis, Geschlechtsbestim-
mung und Bastardierungsvermogen bei
Laminarien, Planta, 12 : 331-353.
(6) Tokida, J. et al, 1956, On Two Rare Abnormal
Forms of Laminaria japonica Aresch.
(In Japanese), Collecting and Breeding,
18: 118-119.
(7) , 1956a, On a Malformed La-
minaria with a Spirally Twisted Lamina,
(In Japanese), Monthly Rept. Hokk.
Fish. Exp. St., 13 : 408-411.
(8) , 1957, On Several Examples
of Malformed Kelps in Hokkaido and
Vicinity, (MS read before the 10th
Hokkaido Regional Conference of the
Botanical Society of Japan, 29-30 June
1957).
(9) , 1957a,AChimaeraof,4/tfr/a
and Laminaria Found in Nature, (MS
sent to the editors of "Nature").
DISCUSSION
G.F. PAPENFUSS apologized for summarizing the paper
in the author's absence, and regretted that the illustrations
had not arrived.
R.F. SCAGEL: How were the cultures started?
G.F. PAPENFUSS (consulting original script): In petri
dishes, the floating gametophytes being isolated to flasks
and maintained at room temperature. Sub-culturing was
carried out from these gametophytes.
R.F. SCAGEL: Care must be taken when drawing con-
clusions from such techniques. All gametophytes look
alike (apart from the cell differences in male and female),
and the sporophyte must show secondary morphological
features before the species can be identified.
In reply to Dr. Papenfuss' enquiry, Dr. Scagel said that
his laboratory isolated sporangia to start their cultures.
228
VOLUME 4
BOTANY
CLEARING OLD TRAILS IN SYSTEMATIC PHYCOLOGYt
GEORGE F. PAPENFUSS
Department of Botany, University of California, Berkeley, California, U.S.A.
There is ample reason to believe that the current
interest in the algae as research material for many
kinds of problems in biology and biochemistry
and as possible sources of food and fuel will
continue. This interest has had and will continue
to have a stimulating effect on those aspects of
phycology that are primarily concerned with the
acquisition of basic knowledge about the algae
for its own sake. We may look forward, there-
fore, to expanded opportunities for fundamental
work on these plants and to an increase in the
number of phycologists in the world.
Because of its large size and the countless
variety of habitats that it provides, no other ocean
possesses such a rich and diversiiied flora as the
Pacific. Our knowledge of the algae of large parts
of the Pacific is still extremely meager, however.
Many of the genera and species have been col-
lected only once and are known only from the
brief and often inadequate original descriptions.
The flora of the Pacific thus provides us with
unlimited possibilities for the pursuit of basic
problems in all branches of phycology.
In anticipation of the development of oppor-
tunities for work on the algae of areas away from
the present centers of activity in the Pacific, I
have selected for discussion certain problems
relating to the taxonomy of green, brown, and
red algae occurring in some of the more remote
parts of this ocean. The problems chosen are
of interest because their solution not only will
advance our knowledge of a number of little-
known genera, some of which are also represented
in the floras of other parts of the world, but the
information gained will be useful in the inter-
pretation of natural relationships within the
groups concerned.
CHLOROPHYCEAE
Information is needed on the life history, the
morphology of the flagella of the motile cells,
the kinds of pigments present in the plastids, and
the nature of the food reserves of Pseudodicho-
tomosiphon in order to determine whether this
genus belongs to the chlorophycean family Dicho-
tomosiphonaceae, which was established by Feld-
mann (12) for the fresh-water genus Dichotomosi-
phon, or whether it is a member of the Vaucheria-
ceae, and hence representative of the class Xan-
thophyceae of the phylum Chrysophycophyta.
Pseudodichotomosiphon is a monotypic genus that
was established by Yamada in 1934 upon material
from the Ryukyu Islands. Tseng (33) has obtained
it also on the Island of Hainan. Luther (20, p.
36) is of the opinion that P. constricta (Yamada)
Yamada had best be retained in Vaucheria.
On the basis of material from Taiwan, Hey-
drich in 1894 erected the genus Rhipidiphyllon.
He regarded the type material as representative
of the species which Askenasy in 1888 described
under the name Anadyomene reticuhita upon
material from Western Australia. More recently
B^rgesen (4) and Taylor (31 ) have reported R.
reticulatum from Easter Island and the Marshall
Islands, respectively. Two problems relating to
this genus are: (1) Is the type material actually
representative of the plant from Western Austra-
lia? (2) Is Rhipidiphyllon an autonomous genus
or should it be merged in Microdictyon, with
which it appears to agree in all essential details?
Rhipidodesmis A. et E.S. Gepp (13), Boodleopsis
A. et E.S. Gepp (13) and Pseudochlorodesmis
B^rgesen (4) are three genera accredited to the
Codiaceae that perhaps are merely the filament-
ous, unconsolidated juvenile stages of other
genera of the Codiaceae, such as Udotea or
Avrainvillea, which are well represented in the
Pacific, or Penicillus, which has not yet been
reported from the Pacific. In this connection,
I might remark that I obtained in South Africa
filamentous, unconsolidated mats of Codium
that are easily mistaken for a filamentous genus
of the Codiaceae; in fact, they were so mistaken
by Dr. Egerod and me until Dr. Silva assured us
that they were only the juvenile stages of some
species of Codium.
Rhipidodesmis is based upon a species from
Ceylon, R. caespitosa (J. Agardh) A. et E.S.
Gepp, which has also been obtained on the
Island of Hainan by Tseng (33). Boodleopsis was
erected for a species from the Malayan Archi-
pelago, B. siphonacea A. et E.S. Gepp, and
Pseudochlorodesmis is based upon a plant from
the Canary Islands, P. furcellata (Zanardini)
t The preparation of this paper was aided by a Grant-in-aid from the National Science Foundation.
229
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
B^rgesen. This species was recently reported
from Vietnam and the Marshall Islands by
Dawson (7,9).
In the paper that I gave at the Seventh Pacific
Science Congress in New Zealand (23), I have
already remarked upon the uncertain status of
Rudicularia (? Codiaceae), which was described
by Heydrich (16) from material obtained in the
Ryukyu Islands, and of Bryobesia (? Derbesia-
ceae) which Weber-van Bosse (34) founded for
a plant from Java.
PHAEOPHYCEAE
Troll in 1931 (32) described under the generic
name Dictyotopsis a peculiar monostromatic
plant which grows by means of wedge-shaped
apical cells that cut off segments alternately on
two sides. The plant forms dense growths on
the aerial roots of mangroves in river mouths in
Sumatra and Amboina. Reproductive organs,
other than propagules, were not observed in any
of the material. In a general way, Dictyotopsis
bears some resemblance to the liverwort genus
Metzgeria, but it differs from this genus in a
number of important features. Dictyotopsis is
provisionally placed in the order Dictyotales,
but the type of apical cell in its thallus is quite
unlike that characteristic of the thalli of the
Dictyotales. Pigment analysis and the discovery
of reproductive organs will probably settle the
question of the systematic position of this inter-
esting genus.
RHODOPHYCEAE
At the generic level, the red algae of the Pacific
present more problems than any of the other large
groups of marine algae. The following are some
of the genera that would especially repay study.
Gracilariocolax Weber- van Bosse (38), which
is based on a species that occurs as a parasite on
Gracilaria in Java, is considered a genus of doubt-
ful systematic position by Kylin (17). It would be
instructive to make a comparative study of
Gracilariocolax, Gracilariophila Setchell et Wilson
(39), which is a parasite on Gracilaria along the
Pacific coast of North America, and Holmsella
Sturch (28), which is a parasite on Gracilaria in
England, to see whether there is a relationship
between these three genera. None of them has
been thoroughly studied, and it is conceivable
that they will be found to be closely related, if
not congeneric.
i De Toni fil. (10) has proposed the substitute name Drouetia for this genus.
230
The genus Weberella (Rhodymeniales: Rhody-
meniaceae) was described by Schmitz (25) upon
material from Flores Island in the Malayan
Archipelago. The type and only species, W.
micans, has also been reported from Timor and
Java by Weber-van Bosse (38) and from Taiwan
by Yamada (40). A study of the structure and
reproduction of this genus is highly desirable.
To judge from the original description by Schmitz
and the observations of Yamada, it appears likely
that Weberella and Herpophyllon Farlow 1 (11),
which is based upon a type from the Galapagos
Islands, are congeneric. Kylin (17) has placed
Herpophyllon in the Squamariaceae (Cryptone-
mialcs), but the structure of the thallus and es-
pecially the habit of these two genera (both have
a dorsiventral organization and produce hap-
teroid protuberances on the lower surface of the
thallus whereby the lobes of the thallus become
secondarily attached to one another or to the
substratum) suggest that they belong in the
Rhodymeniales. In connection with these two
genera, attention should also be drawn to Mali-
chrysis (J. Agardh) Schmitz (24), which is based
upon a type from Morocco, and Sciadophycus
Dawson (6), which was founded for a plant from
Cerros Island (off the west coast of Baja Califor-
nia), both of which appear to belong to the same
complex as Weberella and Herpophyllon.
On the basis of material from the Malayan
Archipelago, Weber-van Bosse (34) established
the genus Exophyllum. This plant has been re-
ferred, with doubt, to the Rhodymeniaceae.
One of its characteristic features is the production
of tetrasporangia in stichidia, a character which,
as Kylin (17, p. 558) has remarked, is entirely
foreign to the Rhodymeniaceae or for that matter
to the order Rhodymeniales. The determination
of the systematic position of this plant must
await a study of its reproductive organs. Recent-
ly the genus was discovered in southern Japan
by Tanaka (30).
Zellera and Opephyllwn are two of the little-
known genera of the Delesseriaceae. Zellera
was described by Martens in 1866 upon a plant
from the Island of Tawalli in the Moluccas and
was later reported from the same region by
Weber-van Bosse (36). The genus appears to be
related to the reticulate genera Claudea and
Vanvoorstia, but its true affinities will remain
uncertain until well-preserved material of the
type species, Z. tawallina, has been studied.
Martens stated that the blades tend to fuse with
others to form a reticulum; but this is not the
VOLUME 4
BOTANY
case, according to Sluiter (27) and Weber-van
Bosse (36). Knowledge of the structure and
reproduction of Z. tawallina may also assist in
the solution of a problem relating to the algae of
the West Indies. Sluiter (27) has referred a species
from that region to Zellera, but her figure of its
habit gives the impression that the plant is mis-
placed in Zellera.
Opephyllum was erected by Schmitz (25) for
a species collected by Martens at Zamboanga
on the Island of Mindanao. This plant has never
again been collected. The genus has not been
illustrated; but to judge from Schmitz's descrip-
tion, it seems likely that Opephyllum will be found
to be congeneric with Martensia Hering.
As is well known, Madame Weber-van Bosse
described a relatively large number of new genera
and species from the East Indies. Unfortunately,
she did not always furnish the detailed informa-
tion that is necessary for the correct placement
of her taxa. The following are genera of red
algae established by Weber-van Bosse, in addition
to some of those already referred to, whose
systematic position is uncertain: Dorella (1921),
Catenellocolax (1928), Microphyllum (1928),
Corallophila (1923), Mortensenia (1926), Chali-
costroma (1911), and Perinema (191 1).
To illustrate the benefits for systematic phy-
cology to be derived from a study of the old
taxa, I might review briefly a few of the recent
contributions that have materially advanced our
knowledge of certain genera.
In the paper that 1 gave at the Seventh Pacific
Science Congress (23), I remarked that the occur-
rence of the reproductive organs on filaments in
the New Zealand genus Microzonia is different
from the condition in all other Dictyotales, to
which order this genus had been assigned. A
study by Miss O'Donnell (22) of material of this
genus given me by Lindauer has shown not only
that the reproductive organs (unilocular spor-
angia) actually are produced on filaments but
that the thallus shows trichothallic growth. It is
evident, therefore, that Microzonia does not
belong to the Dictyotales but to the Cutleriales,
which order previously comprised only the
genera Cutleria and Zanardinia.
Levring (18,19, and in Svedelius, 29) studied
newly-collected material as well as the type of the
type species of Gloiophloea, G. scinaioides J.
Agardh (2), and found that our concept of this
genus is wrong. The previous concept was that
developed by Setchell (26) in his monograph on
the Scinaia complex and was based on a plant
from New Zealand which Setchell believed to be
representative of G. scinaioides, whose type (not
seen by Setchell) was obtained in Australia.
Levring established that the species from New
Zealand (and also the other species of Gloiophloea
described by Setchell) is representative of a new
genus for which he proposed the name Pseudo-
gloiophloea.
The genus Corallopsis was erected by Greville
in 1830 for Sphaerococcus salicornia C. Agardh
(1), whose type was collected by Chamisso during
the voyage of the Russian exploration ship
Rurik, allegedly in Unalaska. Several authors
have doubted that Unalaska actually was the
source of Chamisso's material. Dawson (8)
recently collected a plant in the Philippines that
agrees in all essential details with C. Agardh's
(I) illustration of the type of C. salicornia.
Examination of the log of the Rurik revealed,
furthermore, that the ship had stopped for six
weeks at Manila, which circumstance suggests
with a good deal of certainty that the type of
C. salicornia was obtained in the Philippines.
Dawson's study of the material that he collected
in the Philippines has shown that C. salicornia
is a species of Gracilaria. Thus Dawson in a
most satisfactory way cleared up the question
about the source of the type of C. salicornia and
at the same time showed that Corallopsis Greville
(14) is congeneric with Gracilaria Greville (14).
From the examples that have been discussed
of problems awaiting investigation and of prob-
lems solved, it is evident that the flora of the
Pacific offers challenging opportunities for fund-
amental work in systematic phycology.
I have confined my discussion to problems
relating to the type species of genera occurring
in more or less remote parts of the Pacific. In
addition, many taxa from such regions have been
described as new species of old genera and others
have been identified with species based on types
from other parts of the world. Detailed study of
these species will reveal that a large number of the
putative new species actually are not new and that
many of the others have been misidentified.
Therefore, on the basis of present knowledge of
the algae of the Pacific, little can be concluded
with any degree of certainty about the geographic
distribution of the majority of the species in this
large ocean.
I do not wish to convey the impression that in
my opinion nothing is to be gained from the
cataloguing of the floras of unexplored areas.
It is true, however, that in such works further
confusion will be created through the inevitable
description of new taxa, many of which will have
231
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
to be reduced when the old taxa have been care-
fully studied.
I should thus like to recommend to the phyco-
logist for whom an opportunity has developed to
work in remote parts of the Pacific, that he selects
as territory of operation one or more of the
regions where previous collecting has been done
rather than an area that has remained unexplored.
The unexplored parts can be taken care of much
more profitably after the forest of confusion has
been removed from the old trails.
REFERENCES
(1) Agardh,C. A., 1 820Jcones Algarum Ineditae,
fasc., 1 (4) pp., 10 pis., Lund.
(2) Agardh, J.G., 1872, Bidragtijl Florideernes
Systematik, Lunds Univ. Arsskr. Afd., 2,
8(6), 60 pp.
(3) Askenasy, E., 1888, Algen mit Unterstiit-
zung der Herren E. Bornet, A. Grunow,
P. Hariot, M. Moebius, O. Nordstedt
bearbeitet, in A. Engler (Ed.), For-
schungsreise S.M.S. "Gazelle", Theil 4,
Botanik, pp. 1-58, pis. 1-12, Berlin.
(4) B^rgesen, F., 1920, Marine Algae from
Easter Island, in C. Skottsberg, The
Natural History of Juan Fernandez
and Easter Island, 2:247-309, 50 figs.,
Uppsala.
(5) , 1925, Marine Algae from the
Canary Islands ... 1: Chlorophyceae.
K. Danske Vidensk. Selskab, Biol. Medd.
5(3), 123 pp., 49 figs.
(6) Dawson, E.Y., 1944, Some New and Unre-
ported Sublittoral Algae from Cerros
Island, Mexico, Bull. So. Calif. Acad.
Sci., 43:102-112, 20 figs.
1954a, Marine Plants in the
(7)
(8) -_- -
Vicinity of the Institut Oceanographique
de Nha Trang, Viet Nam, Pacific Sci.,
8:373-469, 63 figs., 1 map.
, 1954b, Notes on Tropical
Pacific Marine Algae, Bull. So. Calif.
Acad. Sci., 53:1-7, 4 figs.
, 1956, Some Marine Algae of
the Southern Marshall Islands, Pacific
Sci., 10:25-66, 66 figs.
(10) De Toni, J., 1938, A Note on Phycological
Nomenclature, Rhodora, 40:27.
(11) Farlow, W.G., 1902, Thallophytes and
Musci of the Galapagos Islands, Proc.
Amer. Acad. Arts Sci., 38:82-99, 102-104.
232
(12) Feldmann, J., 1946, Sur I'h6t6roplastie de
Certaines Siphonales et leur Classifica-
tion, C.R. Acad. Sci. (Paris), 222:752-
753.
(13) Gepp, A., and Gepp, Ethel S., 1911, The
Codiaceae of the Siboga Expedition . . .
Siboga-Exped. Monogr., 62, 150 pp., 22
pis., Leiden.
(14) Greville, R.K., 1830, Algae Britannicae, ...
88 + 218 pp., 19 pis., Edinburgh.
(15) Heydrich, F., 1894, Beitrage zur Kenntniss
der Algenflora von Ost-Asien Besonders
der Insel Formosa, Molukken-und Liu-
kiu-Inseln, Hedwigia, 33: 267-306, pis.
14, 15.
(16) , 1903, Rudicularia, ein Neues
Genus der Valoniaceen, Flora, 92:97-
101, 4 figs.
(17) Kylin, H., 1956, Die Gattungen der Rhodo-
phyceen, 15 + 673 pp., 458 figs., Lund.
(18) Levring, T., 1953, The Marine Algae of Aus-
tralia, I. Rhodophyta: Goniotrichales,
Bangiales and Nemalionales, Arkiv Bot.
ser. 2,2:457-530, 55 figs.
(19) , 1956, Contributions to the
Marine Algae of New Zealand, T. Rhodo-
phyta: Goniotrichales, Bangiales, Nema-
lionales and Bonnemaisoniales, Arkiv
Bot. ser. 2, 3:407-432, 15 figs.
(20) Luther, H., 1953, Vaucheria schleicheri de
Wild, neu ftir Nordeuropa, Mem. Soc.
Fauna Flora Fennica, 28:32-40, 5 figs.
(21) Martens, G., 1 866, Die Tange. Preuss. Exped.
Ost-Asien, Bot. Theil., (1) + 152pp., 8 pis.,
Berlin.
(22) O'Donnel, Elsa H.J., 1954, The Structure
and Taxonomic Position of the Brown
Alga Microzonia, Amer. Jour. Bot., 41:
380-384, 12 figs.
(23) Papenfuss, G.F., 1953, Outstanding Prob-
lems in the Morphology and Taxonomy
of the Marine Algae of the Aropical and
Southern Pacific, Proc. Seventh Pacific
Sci. Congress, 5, Botany, pp. 27-39.
(24) Schmitz, C.J.F., 1889, Systematische Cber-
sicht der bisher Bekannten Gattungen
der Florisdeen, Flora, 72:435-456, pi. 21.
(25) Schmitz, C.J.F., and Hauptfleisch, P.,
1896-1897, Rhodophyceae, in A. Engler
and K. Prantl, Die natiirlichen Pflanzen-
familien, Tell 1, Abt., 2, pp. 298-544, figs.
192-288, Leipzig.
(26) Setchell, W.A., 1914, The Scinaia Assem-
blage, Univ. Calif. Publ. Bot., 6:79-153,
incl. pis. 10-16.
(27) Sluitcr, Catharina P., 1908, List of the Algae
Collected by the Fishery-Inspection Cu-
racao, Rccueil Trav. Bot. Norland, 4:
231-241, 1 fig., pi. 8.
(28) Sturch, H.H., 1926, Choreocolax Polysipho-
niac Reinsch., Ann. Bot., 40:585-605,
15 figs.
(29) Svedelius, N., 1956, Are the Haplobiontic
Florideae to be Considered Reduced
Types? Sv. Bot. Tidskr., 50: 1-24, 14 figs.
(30) Tanaka, T., 1950, Studies on Some Marine
Algae from Southern Japan, 1. Jour.
Kagoshima Fish. College, 1:173-180,
4 figs., 1 pi.
(31) Taylor, W.R., 1950, Plants of Bikini and
Other Northern Marshall Islands, 15 +
227 pp., Frontispiece, pis. 1-79, Ann
Arbor.
(32) Troll, W., 1931, Botanische Mitteilungen
aus den Tropcn, IIT: Dictyotopsis pro-
pagulifera W. Troll, eine nene Brack-
wasseralge Ostindischer Mangrovege-
biete, Flora, 125:474-502, 18 figs., 1
table.
(33) Tseng, C.K., 1936, Studies of the Marine
Chlorophyceae from Hainan, Amov
Marine Blot. Bull., 1:129-200, 34 figs*.,
Ipl.
(34) Weber-van Bosse, Anna, 191 1, Notice sur
quelques Genres Nouveaux d'algues de
VOLUME 4 BOTANY
PArchipel Malaisien, Ann. Jar din Bot.
Buitenzorg, ser. 2, 9:25-33.
(35) ----- - -, 1921, Liste des
Algues du Siboga, H: Rhodophyceae,
partie 1, Protofiorideae, Nemalionales,
Cryptonemiales, Siboga-Exped. Monogr.
596, pp. 185-310, figs. 53-109, pis. 6-8,
Leiden.
(36) - -- , 1923, Liste des
Algues du Siboga, III: Rhodophyceae,
partie 2, Ceramiales. Siboga-Exped.
Monogr. 59c\ pp. 311-392, figs. 110-142,
pis. 9, 10, Leiden.
(37) , 1926, Algues de
['expedition danoise aux iles Kei. Viden-
skab. Medd. Dansk Naturhist. For.
Kfbenhavn, 81:57-155, 43 figs.
(38) - , 1928, Liste des
Algues du Siboga, IV: Rhodophyceae,
partie 3, Gigartinales et Rhodymeniales,
Siboga-Exped. Monogr. 59d, pp. 393-
533,71 figs., pis. 11-16, Leiden.
(39) Wilson, Harriet L., 1910, Gracilariophila,
a New Parasite on Gracllaria confer-
voides, Univ. Calif. Publ. Bot., 4:75-85,
incl. pis. 12, 13.
(40) Yamada, Y., 1932, Notes on Some Japanese
Algae IV, Jour. Fac. Sci. Hokkaido Univ.
ser. 5, 2:267-276, 3 figs., pis. 3-9.
(41) , 1934, The Marine Chlorophy-
ceae from Ryukyu, Especially from the
Vicinity of Nawa, Jour. Fac. Sci. Hokkai-
do Univ. ser. 5, 3:33-88, 55 figs.
DISCUSSION
M.S. DOTY: I suggest that the algal subcommittee
compile a list of problematic species and their localities.
G.F. PAPENFUSS: It is vital to clear up the confusion in
old species before erecting new ones.
j. FELDMANN: Few species are completely well-known.
More cytological studies should be carried out on genera
such as Vaucheria.
G.F. PAPFNFUSS: The problem becomes simpler if there
is a plentiful supply of material.
G.A. PROWSE: In the desmids, the frequent occurrence
of dichotypical cells has made the rc-asscssment of the
concept of species vitally essential.
M.S. DOTY: Have the pigments of Dichotomosiphan
been investigated?
j. FELDMANN: On the basis of the plastids, there is no
doubt that it is a green alga.
G.F. PAPENFUSS: Dr. Feldmann is fully justified in
erecting the family Dichotomosiphonaceae.
233
PROCLLD1NGS OF THE NINTH PACIFIC SCIENCE CONGRESS
ON THE STATE OF PHYCOLOGICAL KNOWLEDGE IN
THE PHILIPPINES!
GREGORIO T. VELASQUEzJ
Department of Botany, University of the Philippines, Quezon City, Philippines.
One finds the first general attempt to identify
the algae of the Philippines in Blanco's Flora
dc Filipinos, of which there were three editions,
in 1837, 1845, and in 1877 to 1883, the latter
edition posthumous. Blanco's identifications of
Philippine species with those of older authors
were of necessity unreliable as he worked in
botanical isolation and with very little available
literature. He proposed a few names for new
species himself in his first edition, of 1837, which
arc no more indefinite that some of those of his
contemporaries in other parts of the world.
These names were later changed (in one instant
by himself) or were considered by other botanists
as synonyms of older ones. In his second edition,
Blanco himself replaced the name Fucus Gidaman
Blanco by Fucus edulis, which was considered by
Georg von Martens to be Sphacrococcus gelatinus
Agardh. The latter name was based upon
Euchcuma gclatinae and Fucus gelatinus.i
Von Martens made a serious attempt to syno-
nymizc Blanco's algae and proposed the following
equivalents:
Fucus prolifcr = Halimeda discoidea Decaisne
Fucus denticulatus - Sargassum (spp.)
Fuc us Gulaman - Sphacrococcus gelatinus Ag.
An unnamed species similar to last - Sphaero-
carpus liclicnoides Ag.
Viva umbilicalis = Zonaria gymnospora Kiitzing
Viva compressa ^~ Enteromorpha complanata,
var. crinita Kilt zing
Viva intcstinalis - Enteromorpha intcstinalis
Link.
It must be noted that some of Blanco's names
are apparently older than the synonyms, and if
sufficiently definite in application to be synony-
mi/cd, should probably replace the later names.
This is particularly true as Blanco was rather
precise in indicating localities, where new collec-
tions at the places he indicated might enable his
species to be interpreted. This is a matter that
may rest for the present, but it is definitely indi-
cated that new and ample collections should be
made at all localities from which species were
described by the older authors, not only Blanco,
but also others. Interpretation of old species by
careful study of the florulas of type localities is
perfectly proper if specimens of the older authors
have not survived.
Not only did Georg von Martens look into the
identity of Blanco's species, he likewise compiled
an elaborate table showing the known geograph-
ical distribution in 1866 of all the Algae that had
been described or reported from tropical Asia
and the tropical Pacific. This table surely affords
a key to most of the literature bearing on Philip-
pine algae that appeared prior to 1866, but also ac-
counted for the collections of his son Dr. Eduard
von Martens who accompanied the Prussian
East Asia Expedition as zoologist. On account
of his father's special interest in the algae, Eduard
von Martens collected them whenever he had
the opportunity.
Only two Philippine fresh-water species were
described as new by Georg von Martens, namely
Cladophora diluta and Cladophora luzoniensis.
The seaweeds were all considered to have been
previously known, but there are among them
first records from various localities.
A few of the scanty early records of Philippine
algae were based upon collections of the
American Exploring Expedition under the com-
mand of Charles Wilkes. The botanical collec-
tions of the Wilkes Expedition were made by
Charles Pickering, general naturalist and ethnobo-
tanist, author of The Chronological History of
Plants: Man's Record of his own Existence
illustrated through their Names, Uses, and Com-
panionship, William D. Brackenridge, who
wrote a volume on the ferns, and William Rich,
who was primarily interested in collecting flower-
ing plants. Because there was no especialist in the
lower cryptogams among the naturalists of the
t Made possible through the Guggenheim fellowship and the University of the Philippines special detail fellowship during
1956-1957.
Presented by G.F. Papcnfuss.
t Secretary of the Graduate School and Associate Professor in Botany, University of the Philippines.
1 See J.G. Agardh, 1851, in Species Genera et Ordines Floridearum, 2: 268.
234
VOLUME 4
BOTANY
Expedition (and such were few at that early date),
the collections of algae were meagre. They were
determined by Bailey and Harvey 2 , in whose report
there are two lists, one of the large forms con-
taining numbers of Philippine reds and browns,
only one of them new, namely Dictyota dichotoma,
and a second longer list containing several new
species of diatoms, namely Amphitetras favosa,
Campy lodiscus Kutzingii, Lagena WilUamsonii,
and Triccratiwn orientate.
All of the diatoms were listed as having come
from the Sula Sea, without designation of place,
but are fairly considered as part of the Philippine
flora, as they were obtained as epiphytes on the
larger algae. There is to be found in the general
report of the Expedition, which was written by
Wilkes himself, a clear indication that shore
collecting was done at only one Philippine
locality. That was at Marongas Island, indicated
on Wilkes* map of Jolo (Sooloo) Island as lying
across a narrow strait of the northeast coast of
Jolo.
An English resident of Manila during the
Spanish regime was Hugh Cuming, who made
the most extensive collections down to the time
of the American occupation. His algal collections
were described by C. Montagne. They were
widely distributed from Kew, and it was possible
even as late as the early years of the present cen-
tury for Dr. E.D. Merrill to secure some of the
duplicates for the herbarium of the Bureau of
Science, which had been built up vigorously dur-
ing the period prior to World War II. The
Philippine algae of the famous British "Challenger
Expedition" were enumerated by Dickie in 1876
and 1877. The specimens were presumably in
English herbaria. A good many algae were
accumulated by the Bureau of Science collectors
and by botanists connected with the University
of the Philippines. This included Walter Shaw,
especially interested in the Volvocaceae, who
wrote on that group and published new Philip-
pine genera, namely: Campbellosphaeria, Janeto-
sphaeria, Merrillosphaeria, and Copelandosphaeria.
It was fortunate for the beginning of phycological
study that some of the algae of the Bureau of
Science and University herbaria had been sent out
as exchanges or on loan prior to World War II
and are still to be found in various herbaria a-
broad.
During the early part of the American period
the most extensive collecting of Philippine algae
was carried out by the Dutch Siboga Expedition.
There was a general report by Mme. Weber van
Bosse on all of the collections, and monographs
of special groups, the Codiaceae by A. and
E.S. Gepp, the genus Halimeda by E.S. Barton,
and the remarkably well represented corallines
by A. Weber van Bosse and M. Foslie. The
Siboga Expedition dredged very extensively at
charted stations throughout the Sulu Sea, and
the Foslie contributions to our knowledge of the
group are basic. The specimens are presumably
in Holland and elsewhere in Europe. They were
never represented in the Philippine herbaria.
The algae of the voyage of the Italian Vettor
Pisani were described by A. Picconc in 1886.
Later in 1889, three more species were added in
the record of Philippine algae. That ended the
period of visits of oceanographic expeditions, so
far as reports on algae were concerned.
During the incumbency of an exchange profes-
sorship at the University of the Philippines in 1935,
H.H. Bartlett began a systematic effort to collect
the algae. Many localities were visited by him,
his colleagues, and students. The collections have
so far been studied only partially, the Chlorophy-
ceae by Dr. W.J. Gilbert, the genus Galaxaura
by Dr. Ruth Chou, and various Myxophyceae
by the present writer. Fortunately, duplicates of
almost everything went to the University of
Michigan, and therefore remain as a basis for
further study. A collection of minute epiphytes
had been picked off of larger algae and mounted
on slides. These only were lost, but they can
doubtless be replaced from the larger herbarium
specimens. A series of plankton collections were
also lost. The localities represented have been
listed by Gilbert in his doctoral dissertation.
Most of the Myxophyceae were collected by the
present writer. He was induced by the urgency
of the nuisance caused in fish ponds by blue-
green algae to concentrate on that group. Hun-
dreds of collections were made, which were
largely studied taxonomically by Drouet, and
which appeared in numerous papers prepared by
the writer. But the more extensive is Drouet
and Daily's "Revision of the coccoid Myxo-
phyceae" (12) which cited all myxophycean
unicellular forms so far known from the Philip-
pines. Thanks to the circumstances of duplicates
having gone abroad, the vouchers for Philippine
collecting in this group down to 1941 have not
been lost.
2 Summary of the life of J.W. Bailey and W.H. Harvey, each a pioneer in his chosen botanical career at the time, may
be found respectively in Appleton's Cyclopaedia of American Biography, V. 1, 1888 and Dictionary of National Biography,
V. 25, 1891.
235
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Bartlett was again in the Philippines in 1940-41,
and although engaged in agronomic work, he
arranged for a pearl diver (Balhani, Moro of
Siasi) to make bulk collections at various places
in the Sulu Sea that could be reached by native
vinta from Zamboanga. The collecting was
supervised in large part by Professor Jose S.
Domantay of the Bureau of Fisheries. All of
these collections were preserved in formaldehyde
in 55 five-gallon cans. The huge task of making
them into herbarium specimens was undertaken
by the University of Michigan under the super-
vision of Dr. Wm. Randolph Taylor. They have
not yet been studied, but sets will be returned
eventually to the Philippines for the University
and the National Herbarium.
In spite of the discouraging loss of all of the
botanical collections and the libraries of the
Bureau of Science and the University of the
Philippines during the war for liberation of
Manila, an immediate effort was made in 1945
to make a new start. This was greatly aided by a
grant-in-aid to the present writer from the Ameri-
can Philosophical Society, which enabled him to
continue his own researches on the Myxophyceae.
Several of his students developed a sufficiently
strong interest so that they were able to publish
contributions or deposit unpublished master's
theses, notably J.D. Soriano, M. Cantoria,
V. Aligaen, E. Medina, M. Valero, and V.G.
Viola. Needless to say, the publications and
theses of these immediate post-war workers could
not have been prepared in the absence of all
reference materials without the friendly and
much appreciated collaboration of authorities
abroad, among whom should be especially
mentioned Dr. Francis Drouet at the Crypto-
gamic Herbarium of the Chicago Natural History
Museum and Dr. Wm. Randolph Taylor of the
University of Michigan.
In order to maintain impetus, the Algological
Society of the Philippines was organized in
June 1956, with sixteen initial members. Later
the names were added of several persons who
were interested in algae as sources of food, in
their uses in pharmacy and various fertilizer ana-
lyses in chemistry, or in other economic aspects
of the group such as their cultivation in fishponds.
Dean Patrocinio Valenzuela, Executive Secretary
of the National Research Council of the Philip-
pines, and Dr. Deogracias Villadolid, former
Director of the Bureau of Fisheries and presently
engaged in fish culture, were made honorary
members to furnish moral support and necessary
encouragement to the new organization. Among
236
the enthusiastic members are Professor Domantay
who collaborated with Bartlett in organizing and
supervising the Sula Sea collecting in 1940-41,
Mr. I. A. Ronquillo of the Bureau of Fisheries,
Mr. M. Palo of the Institute of Science and
Technology, Mr. Z.R. Torres of the University of
the East, and Dr. J.V. Santos of the Department
of Botany, University of the Philippines.
As the members of this Society could do little
at the start, except to collect and to make ecolog-
ical and economic observations, they have con-
ceived the idea of building up a "bank" of well
duplicated numbered specimens, only roughly
identified as to family or genus, from which
specialists abroad may receive material for use
in monographic or regional studies. It is hoped
that there will be requests from members of the
Subcommittee on Algae of the Committee on
Botany of Pacific Science Council for material to
be used in their various researches. As fast as
material from this "bank" is identified, remaining
specimens will be incorporated with the organized
herbaria of the Philippine National Museum,
the University of the Philippines, and any other
institutions that participate in the project. The
preparation of "Algae Philippinae Exsiccatae"
is soon to be started with the cooperation of
Dr. Mason Hale, Associate Curator of the Divi-
sion of Cryptogams, U.S. National Herbarium
of the Smithsonian Institution.
There is every reason why there should be
immediate cooperation with other countries of
Southeastern Asia and the Pacific. There is so
large a widely distributed element in the algal
floras of the tropical countries that are neighbors
to the Philippines that an understanding of geo-
graphical and ecological relationships requires
the most extended study possible. Furthermore,
there are various new political entities in the Old-
World tropics with newly established scientific
institutions which might like to cooperate closely
with the Philippines, or to organize regional
survey work independently in a similar manner.
In the rebuilding of Philippine references and
research collections an effort is being made to
secure for Manila institutions as much duplicate
prewar material as can be found abroad. The
fortunate circumstance that much still remains
for distribution from the University of Michigan
has already been mentioned. The greatest pos-
sible cooperation has been realized from Dr.
Drouet of the Cryptogamic Herbarium, Chicago
Natural History Museum, who has sent to Manila
whatever could be spared of duplicates sent him
prior to the destruction of the Philippine institu-
VOLUME 4
BOTANY
tions. Dr. Tiffany of Northwestern University
turned over to the writer all his accumulated
articles on the Myxophyceae for library refer-
ences. There could be no greater proof of the
value of collecting abundant material and distri-
buting it widely. What is destroyed in one place is
preserved in another. As a result of inter-institu-
tional cooperation between the Philippines and
the United States, for example, the voucher
specimens for 183 Philippine records of Myxo-
phyceae, including types, have been preserved.
BIBLIOGRAPHY
(1 ) Aligaen, V., 1954, Studies on the Algae of
the Brackish Water Fishponds of Iloilo
and Vicinity. Master of Science thesis
(in press), University of the Philippines.
(2) Bailey, J.W., and Harvey, W.H., 1851, Des-
cription of Algae from the Wilkes
United States Exploring Expedition in
Proceedings of the Meeting for Dec. 4,
1850, Proc. Boston Soc. Nat. Hist., 3:
369-378.
(3) - - , 1862, Uni-
ted States Exploring Expedition during
the Years 1838-1 842, under the Command
of Charles Wilkes, U.S.N., XVII, Phila-
delphia, pp. 153-182.
(4) Barton, E.S., 1901, The Genus Halimeda,
Siboga Expeditie Monographic, 60, 32
pp., 4 pis.
(5) Blanco, M., 1837, Flora de Filipinas, Segun
el Sistema Sexual de Linneo. LXXV11I-
887 pp., Manila. (There is also an
1845 edition.)
(6) Britton, M.E., 1948, New Species of Oedogo-
nium from Leyte, the Philippine Islands,
Amer. Jour. Bot., 30 (14): 715-719,
16 figs.
(7) Cantoria, M., Valenzuela, P., and Velasquez,
G.T., 1951, Pharmacopoeial Properties
of Agar from Three Philippine Seaweeds,
Jour. Phil. Pharm. Assoc., 38: 187-190,
3 figs.
(8) Chou, Ruth C, 1945, Pacific Species of
Galaxaura, I : Asexual Types. Papers of
the Mich. A cad. Sci. Arts and Letters,
30:35-55, 11 pis.
(9) -- , 1947, Pacific Species of
Galaxaura, II: Sexual Types, Ibid., 31
(1945): 3-24, 13 pis.
(10) Dickie, G., 1876, Contributions to the
Botany of the Expedition of H.M.S.
"Challenger" Algae Chiefly Polyne-
sian, Jour. Linn. Soc. Bot., 15: 235-246.
(11) __ ._. , 1877, Supplementary
Notes on Algae Collected by H.N.
Moseley, M.A. of H. M. S. "Challenger"
from Various Localities, Jour. Linn. Soc.
Bot., 15: 486-489.
(12) Drouet, F., and Daily, W.A., 1956, Revision
of the Coccoid Myxophyceae, Butler
Univ. Bot. Stud., 12: 1-218.
(13) Gepp, A., and Gepp, E.S., 1911, The
Codiaceae of the Siboga Expedition,
Siboga Expeditie Monographic, 62, 150
pp., 22 pis.
(14) Gilbert, W.J., 1942, Studies on the Marine
Chlorophyceae of the Philippines. Doc-
tor of Philosophy thesis (unpublished),
University of Michigan.
(15) Martens, G. von, 1866, Die Proussische
Expedition nach Ost-Asien , Bot.
Thcil. Die Tange 4 plus 152 pp., 3 pis.
K. Geheime, Berlin.
(16) Medina, E., 1955, Studies on the Siphonales
of Puerto Galera, Oriental Mindoro.
Master of Science thesis (in press),
University of the Philippines.
(17) Merrill, E.D., 1918, Species Blancoanae, a
Critical Revision of the Philippine
Species of Plants Described by Blanco
and Llanos, Dept. Agr. and Nat. Res.,
Bu. Sci., Manila, 12, 423 pp., 1 map.
(18) Piccone, A., 1886, Alghe del Viaggio di
Circumnavigazione della Vettor Pisani,
Genoa, 97 pp., 1 map.
(19) , 1889, Nuove Alghe del
Viaggio di Circumnavigazione della Vet-
tor Pisani, Roma, R. Ace. Lined Mem.
Series 4, 6: 10-63.
(20) Rabanal, H.H., 1949, The Culture of Lab-
lab, the Natural Food of the Milkfish Fry
and Fingerlings under Cultivation, The
Technical Bulletin, 18, Bu. of Printing,
Manila.
(21) --, and Montalban, H.R., 1953,
The Growing of Algae or "Lumut" in
Bangos Fishponds, Philippine Fisheries,
pp. 142-152, 3 figs.
(22) Shaw, W.R., 1919-1922, Papers on Volvo-
caceae of the Philippines, four new
genera in separate articles, Phil. Jour.
Sci., respectively: Cawpbellosphaeria,
15 (6): 493-520, 2 pis., 1 fig.; Janetos-
phaeria, 20 (5): 477-508, 5 pis., 5 figs.;
237
PROChhDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Merrillosphaeria, 21 (1): 87-129, 8 pis.,
1 fig.; and Copelandosphaeria, 21 (2):
207-232, 4 pis., 2 figs.
(23) Soriano, J.D., 1953, Myxophyceae of Panay
and Ncgros Islands, Nat. and A pp. Set.
Bull., 13(1 and 2): 3-57, 3 pis.
(24) , and Velasquez, G.T., 1952,
Studies on the Myxophyceae of Manila
and Vicinity, Ibid., 12 (1): 1-93, 4 pis.
(25) Tiffany, L.H., 1951, Two New Oedogonia
from the Philippines, Chicago Acad.
Sci., 82: 81-83, 1 pi.
(26 ) Valero, M., 1956, Preliminary Studies on
the Algae of U.P. Site, Quezon City.
Master of Science thesis (unpublished).
University of the Philippines.
(27) Velasquez, G.T., 1940-41, Papers on Fila-
mentous Myxophyceae of the Philip-
pines 1, II, and III, Nat. and App. Sci.
Bull., respectively: 7 (3): 269-271 : 8 (2):
189-200: and 8 (3): 203-210.
(28)
(29)
(30)
(31)
, 1948, Survey of the Algae
and Economic Algal Resources of the
Philippines, The Amcr. Philosophical
Society Year Book, pp. 154-155.
, 1950, Studies on the Myx-
ophyceae of Puerto Galera and Vicinity,
Nat. und App. Sci. Bull., 10(4): 309-328.
, 1952, Algal Pollutions
from the Ponds of Puerto Galera, Orien-
tal Mindoro, Ibid., 12 (3): 239-251.
, 1952,
of the Philippines,
Seaweed Resources
Proc. of the First
International Seaweed Symposium, In-
stitute of Seaweed Research, Scotland,
pp. 100-101.
(32) , 1953, Studies on the
Marine Algae of the Philippines. Ab-
stract, Proceedings of the Eighth Pac.
Sci. Cong., pp. 205-206.
(33) Villadolid, D.V., and Villaluz, O.K., 1953,
A Preliminary Study on Bangos Cultiva-
tion and Its Relation to Algal Culture in
the Philippines, Bu. of Printing, Manila,
Pop. Bull., 30, pp. 3-16.
(34) Viola, V.G., 1956, The Chlorophyceae of
Puerto Galera and Vicinity, Their Dis-
tribution and Reproduction. Master of
Science thesis, University of the Philip-
pines. (With permission to be read in
the Ninth Pacific Science Congress,
1957.)
(35) Weber van Bosse, Ann., 1913-28, Listc des
algues du Siboga, I: Myxophyceae,
Chlorophyceae, Phaeophyceae , pp.
1-1 86, pis. 1-5(1913). II:Rhodophyceae,
premiere partie, Protoflorideae, Nema-
leonales, Cryptonemiales, pp. 187-310,
pis. 6-8 (1921); secondc partie, Cerami-
ales, pp. 311-392, pis. 9-10 (1923); troisi-
cmc partie, Gigartinales et Rhodymeni-
ales , pp. 393-533, pis. 11-16 (1928),
Siboga Expeditie Monog., 59, E.J. Brill,
Leiden.
(36) , and Foslie, M.,
1900, The Corallinaceae of the Siboga
Expedition, Sihoga Expeditie Mono-
graphie, 61, 110 pp., 16 pis., 34 text figs.
DISCUSSION
G.F. PAPENFUSS: Perhaps Dr. Santos can tell us what
has happened to Blanco's material.
j.v. SANTOS:
the war.
It was destroyed by fire in Manila during
I should explain that the algal society mentioned by
Dr. Velasquez has been formed from members from
zoology, medical, and fisheries departments. The algal
collection is increasing from time to time, and Dr.
Velasquez is organizing the identifications.
238
VOLUME 4
PHYLOGENETIC RELATIONSHIPS OF
CERTAIN DORSIVENTRAL RHODOMELACEAE
BOTANY
ROBERT F. SCAGEL
University of British Columbia, Vancouver, Canada.
Although Schmitz (14) and Schmitz and Fal-
kenberg (15), as well as earlier authors, had sub-
divided the family Rhodomelaceae into a number
of subfamilies, Falkenberg's (2) monograph on
the Rhodomelaceae was the first noteworthy
attempt at a broad phylogenetic study of this
largest family of Rhodophycophyta in which
there are some 1,200 species now recognized.
Oltmanns (8, 9), Rosenberg (11), Kylin ?tf, 7),
Fritsch (3), and Scagel (12) have attempted to
bring the group in line with more modern con-
cepts of classification in the Florideophycidae,
but each, with some modification, has essentially
followed Falkenberg's division of the family into
subfamilies. Rosenberg (II) removed the Dasy-
aceae from the Rhodomelaceae and established
this group as an autonomous family, the Dasy-
aceae, and Scagel (12) has attempted, on the basis
of a study of certain of the dorsiventral Rhodo-
melaceae, to further assess phylogenetic relation-
ships in the family. Kylin (7 ) has departed to the
greatest extent from the system of classification
proposed by Falkenberg (2). It has been possible
by further study of some of the genera in this
large group to place a number of entities formerly
of uncertain position. However, there still exists
in this family a broad field for research on a
generic level before a satisfactory approach can
be made to the solution of the many specific
problems and before the merits of the phylogene-
tic relationships can be adequately weighed. As
Oltmanns (8, 9) has stressed, one must have a
detailed knowledge of the family as a whole before
a reassessment of such a large group can be made.
Many of the genera, even as they have now been
rearranged or subdivided by Kylin (7), still in-
clude obscure and little-known plants which
have never been adequately studied. There seems
little doubt that further rearrangements and sub-
divisions of this large family will be warranted,
but until more of the entities have been studied
in greater detail and in all stages of development
there seems little justification in proceeding fur-
ther at this time in an attempt to regroup genera
into subfamilies.
The position of the Rhodomelaceae in the
classification system, as for the majority of the
groups in the Florideophycidae, is based essen-
tially on embryological characters proposed by
Schmit/ (13). More recent studies, particularly
on the female reproductive apparatus, have
further emphasized the importance of using
these structures for the interpretation of phy-
logenetic relationships on both ordinal and family
levels. The studies of Kylin (4,5 ) and Pa pen fuss
(10) have particularly stressed the importance
of the use of the female reproductive system in
the delimitation and interrelationships of the
families of algae now included in the Order
Ceramialcs. It is apparent from these studies
that the Ceramiales is a highly evolved group.
In the members of this order the carpogonial
branch, auxiliary cell, supporting cell, and often
the central cell, the sterile cells, and in some
instances even the inner cells of the pcricarpic
layer, all contribute in varying degree to the
nutrition of the carposporophyte. Of the four
families in this order, however, this phylogenetic
trend in the nourishment and protection of the
developing carposporophyte is most clearly
evident in the Rhodomelaceae. In this family,
the direct fertilization of an auxiliary cell, which
is formed only after fertilization of the carpog-
onium, and the early formation and high degree
of development of the pericarp are well-
established features. With respect to both re-
productive and vegetative structures, not only
the Order Ceramiales, but also the Family Rhodo-
melaceae is the most clearly defined and most
highly developed of all groups of Florideophy-
cidae.
Chief among the distinguishing characteristics
of the life-histories of the Rhodomelaceae, as well
as for the majority of the Florideophycidae, is
the succession of somatic phases, different from
that of most other algal groups. This is due to
the development of the carposporophyte, fol-
lowing the fertilization of the carpogonium.
The gametophyte, which bears the sex organs,
is haploid, and the fertilized carpogonium gives
rise indirectly to a carposporophyte with diploid
sporogenous tissue. No reduction division ac-
companies the production of the carpospores,
which germinate to give rise to the diploid tetra-
sporophyte. This tetrasporophyte bears the
tetrasporangia in which reduction division occurs
239
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
and the haploid tetraspores give rise to male and
female gametophytes. Thus there are four soma-
tic phases, two haploid and two diploid. The
carposporophyte, which is almost like a parasitic
generation, is much reduced in vegetative develop-
ment and remains attached to the female game-
tophyte. Although the tetrasporophyte and the
male and female gametophytes are quite separate
phases in the life history, they cannot generally
be identified morphologically as such, or even
anatomically until the reproductive cells are
formed. For this reason, there is an even greater
opportunity in such a group for studying vege-
tative characteristics both in the haploid and
diploid phases.
In spite of the large size of the family Rhodo-
melaceae, there is probably no group of algae
which shows such a remarkable degree of
uniformity in the female reproductive organs
and in the post-fertilization development. Al-
though the uniformity of the female reproductive
system is an outstanding feature of the Rhodo-
melaceae, on the other hand the family shows a
diversity in vegetative organization not met
elsewhere in a family of multicellular algae. The
strictly apical development of the thallus can be
traced in the young plant by means of the primary
pit-connections between cell lineages, although
in older stages secondary pit-connections may
be formed between adjacent pericentral cell
derivatives and result in a pseudoparenchymatous
thallus whose development and organization may
soon be obscured. Although the development of
the mature thallus may be completely evident
from a study of its apices, there are instances, as
in Placophora, in which the evanescent juvenile
stages have been found to exhibit an organization
quite different from that of the mature thallus.
For this reason it is instructive to study not only
the mature plants but also the juvenile plants and
their development before completing a phylo-
genetic arrangement of genera in this group.
Although the range in construction of the thalli
varies from comparatively simple, radially sym-
metrical plants to elaborate and complex types
with bilateral symmetry or dorsiventral organiza-
tion, through a study of the apex of the mature
plants the polysiphonous theme on which these
variations are superimposed is readily ascertained
even in the most highly modified types.
The family can be subdivided into a number
of subfamilies which can be arranged in several
lines of evolution based primarily on symmetry.
Accumulated evidence supports the theory that
the subfamily Polysiphonieae is the basic group
from which all other subfamilies have been
240
derived. The ancestral type was probably a sim-
ple, radially symmetrical, Polysiphonia-like alga
with five pericentral cells in the polysiphonous
segments. On this basis the evolutionary lines
are : the Potysiphonia-series, the basic group from
which all Rhodomclaccae are probably derived,
the Lophothalia-, Chondria-, Pterosiphonia-, Her-
posiphonia-, and Amansia-series. The Polysipho-
nia-, Lophothalia-, and Chondria-series include
chiefly radial forms, whereas the Pterosiphonia-
series includes primarily bilateral forms. The
Herposiphonia- and Amans la-series include pri-
marily dorsiventral forms. It is a part of the
Herposiphonia-series which I shall use particularly
to illustrate the emphasis which may be placed
on vegetative characteristics in the group.
According to Falkenberg (2) this series com-
prises two subfamilies the Herposiphonieae and
the Polyzonieae. However, Kyi in (7 ' ) has further
split up the Herposiphonieae into three groups
which may be referred to as the Herposiphonieae,
Placophoreae, and the Streblocladieae. The
genera included in this series are chiefly dorsi-
ventrally organized. In some, as in Metamorphe
(Herposiphonieae), the axes are simple, more or
less erect, and Polysiphonia-Yike with dorsiven-
trality very slightly developed. In more advanced
genera, as Placophora and Amplisiphonia (Placo-
phoreae), dorsiventrality is very pronounced and
the mature thalli, which are almost entirely
prostrate, show extensive congenital coalescence
of the axes. The Polyzonieae includes genera
which show strongly pronounced dorsiventrality.
Many of the latter have acquired the form of
jungermanniaceous liverworts as the result of
various types of wing-development (Leveillea),
the coalescence of determinate laterals (species
of the "cuneifoliate" group of Dasyclonium
(=Euzoniella) or a combination of both.
The initial stages of the prostrate phase of many
of these dorsiventral forms arise from the basal
part of an erect, radially or bilaterally symmetri-
cal, determinate and evanescent juvenile or pri-
mary axis. But even where there is pronounced
dorsiventrality, as in Placophora, when repro-
ductive branches are produced, the latter may
revert to an erect radial habit. If we interpret
these juvenile stages as evidence of ontogeny
repeating phylogeny, then we have additional
features supporting the theory that the erect,
radial, Polysiphonia-type represents the prototype
of the group. One might postulate that in time
the erect form lost its ability to produce exogenous
indeterminate laterals. The production of a pros-
trate indeterminate lateral from the lower part of
the juvenile axis consequently has survival value,
VOLUME 4
BOTANY
and the plant is able to produce an extensive
thallus despite the fact that it may have lost the
capacity of producing exogenous indeterminate
laterals at the apex of an erect primary axis. As
a result of the development of holdfasts and a
prostrate habit, this successful, basal, indeter-
minate branch has gained a still better chance
of survival.
To illustrate how one can carry the analysis of
the vegetative characteristics to the specific level
I should like to consider in somewhat greater
detail the genus Dasyclonium (= Euzoniella).
As far as is known, the dorsiventral phase of the
thallus of the species of this genus is initiated by
a primary lateral which arises from the basal
part of a diminutive, comparatively evanescent,
radially or bilaterally symmetrical primary axis.
The two groups within this genus, which have
been designated (12) as the "incisate" and "cunei-
foliate" groups, are characterized primarily on
the basis of the structure of the determinate
branches. The "incisate" group includes those
species in which the secondary and subsequent
orders of branches formed by the determinate
axes are more or less free from one another at
maturity. The "cuneifoliate" group includes
those species in which the secondary and higher
orders of branches formed on the determinate
axes are congenitally fused to varying degrees
at maturity. Kylin (7) has chosen to split up the
group of species formerly included in the genus
Euzoniella Falk. 1901 (in the broad sense as the
name was proposed for conservation as against
Dasyclonium J.Ag.,7; by Silva,/6,/7) into two
groups (Dasyclonium J.Ag., sensu Kylin, 7 and
Euzonia Kylin, 7). These two groups are those
which have already been designated as the
"incisate" and "cuneifoliate" groups. I do not
believe that the separation of this group of species
into two genera is justified. Further confusion
results in properly assigning certain entities, one
of which Kylin apparently overlooked, [Dasyclo-
nium palmatifidum (Grun.) n. comb. (= Euzoniella
palmatifida (Grun.) Cuoghi-Cost.), D. harveya-
num (Decne ex Harv.) Kylin (= Euzoniella har-
veyana (Decne ex Harv.) Falk.) and D. ocellatum
(Yendo) n. comb. (= Euzonia ocellata (Yendo)
Kylin) ] to Dasyclonium sensu Kylin and his new
genus Euzonia. Although a good case may be
made for conserving Euzoniella as proposed by
Silva (16), as this proposal was subsequently
rejected in favour of Dasyclonium, it would now
seem best to reject Kylin's division of the group
and refer all species to the one genus Dasyclonium
J.Ag. (= Euzoniella Falk.). On the basis of the
structure and coalescence of the determinate
branches, the species of this genus can be arranged
in an evolutionary series whose equal for com-
pleteness is probably not encountered elsewhere
in the family ( 12). Any attempt to separate these
two groups serves only to obscure their natural
relationships. It should be noted that in the
"incisate" group, which Kylin (7) refers to Dasy-
clonium, he has included D. harveyanum (Decne
ex Harv.) Kylin. However, the lowermost
lateral of the determinate branch of this species
is further branched, a feature which is character-
istic of the species of the "cuneifoliate" group
and, although Dasyclonium harveyanum might
be regarded as a transitional species, it should
more properly have been placed by Kylin in the
genus Euzonia. On the other hand he has placed
Dasyclonium ocellatum (Yendo) n. comb. (= Euzo-
niella ocellata Yendo ; = Euzonia ocellata (Yendo)
Kylin), in which there is no coalescence of mono-
siphonous laterals, with the "cuneifoliate" group,
whereas this species is most closely related to
D. incisum (J.Ag.) Kylin (= Euzoniella incisa
(J.Ag.) Falk.). Furthermore, Kylin has omitted
any mention of D. palmatifidum (Grun.) //. comb.
which also forms a natural link between the
"incisate" and "cuneifoliate" species. Dasyclo-
nium palmatifidum has only incipient stages of
coalescence in the laterals on the determinate
branch, but it has the branching habit of the
lowermost lateral of the determinate branch
typical of the "cuneifoliate" group. Thus it can
be shown that there are species of the genus
Dasyclonium which illustrate all stages in transi-
tion from the monosiphonous, uncoalesced con-
dition of secondary laterals through to a poly-
siphonous (or partially polysiphonous) and
completely coalesced condition. The develop-
ment of the spermatangia axes in Dasyclonium
incisum, D. palmatifidum, and D. cuneifolium
(Mont.) n. comb. (= Euzonia cuneifolia (Mont.)
Kylin; = Euzoniella cuneifolia (Mont.) Falk.) are
also identical, as are the number and arrangement
of cover cells formed during the development
of the tetrasporangium. All of these features
further support the argument that these entities
are all species of the same genus.
Dasyclonium incisum apparently is one of the
most primitive species in the genus, and may
have served as the prototype from which D.
flaccidum (Harv.) Kylin and D. ocellatum have
been derived by reduction of the polysiphonous
laterals on the determinate branches to the mono-
siphonous unbranched condition. On the other
hand, it seems likely that a monosiphonous con-
dition throughout may represent the prototype
for the genus as a whole. Dasyclonium bipartitum
241
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
(Hook. f. et Harv.) Kylin [= Euzoniella bi-
partita (Hook. f. et Harv.) Falk.] may also be
considered as having been derived through re-
duction from D. incisum. In this instance it is
the number of secondary laterals that has been
reduced rather than the polysiphonous character
of the laterals that has been lost. In D. biparti-
tum only the two lowermost secondary laterals
are initiated.
The next probable step in the evolution of the
determinate branch is indicated in Dasyclonium
harveyanum. In this species several orders of
branches, which remain free from one another at
maturity, are formed from the lowermost lateral
of the determinate appendage. This branching
characteristic is also found throughout the
"cuneifoliate" group and suggests that D.
harveyanum forms a connecting link, as has
already been suggested, between the "incisate"
and "cuneifoliate" groups. In the "cuneifoliate"
group, various degrees of congenital coalescence
of the secondary laterals have occurred, resulting
in the evolution of a monostromatic (for the
most part) determinate appendage. The incipient
stages of this coalescence and monostromatic
character are indicated in D. palmatifidum. This
congenital coalescence of the laterals is more
evident in D.flabellifera (J.Ag.) n. comb. [-Euzo-
niaflabellifera (J.Ag.) Kylin; = Euzoniella flabel-
lifcra (J.Ag.) Laing] and is even more pronounced
in D. ovalifolium (Hook. f. et Harv.) n. comb.
[=Euzonia ovalifolia (Hook. f. et Harv.) Kylin;
^Euzoniella ovalifolia (Hook. f. et Harv.) Falk.],
D. cuneifolium and D. adiantiformis (Decne) n.
comb. [-Euzonia adiantiformis (Decne) Kylin;
^Euzoniella adiantiformis (Decne) Falk.].
In summary one can say that the female re-
productive structures, which are strikingly uniform
throughout the family Rhodomelaceae, appear
to be of least phylogenetic significance at the
subfamily, generic, and specific levels. The
diversity in vegetative construction that is asso-
ciated with this uniformity in the structure of the
reproductive organs in the Rhodomelaceae, as
well as in the Ceramiales as an order, indicate
that the group is a long-established one, whose
representatives have undergone pronounced
evolutionary development in the vegetative system
in comparatively recent times. Of the families
constituting this order, the Rhodomelaceae is
the most highly evolved.
Symmetry relationship is the most fundamental
feature on which a separation of the subfamilies
into the various probable lines of evolution can
be made. Bilaterality and dorsiventrality appear
242
to have been achieved more than once within the
family along entirely different lines of evolution,
but in general there appears to have been an
advance from simple forms with a radial organ-
ization as in the Polysiphonieae to bilateral types
as in the Pterosiphonieae, and finally to elaborate
forms with dorsiventral organization as in the
Polyzonieae.
The method of formation, type, pattern of
distribution, and the degree of congenital fusion
or subdivision of the laterals are of secondary
importance, but are significant at the generic and
specific levels in distinguishing taxa.
The genus Euzonia Kylin is rejected because
the species included in it form only a part of an
evolutionary series of closely related entities
hitherto referred to the genus Euzoniella Falk.
Furthermore, because of the additional confusion
that has resulted by referring only a part of this
same series to Dasyclonium J.Ag. (sensu Kylin),
and because an earlier proposal to conserve Euzo-
niella has been rejected, it would seem best to
refer all entities to the one genus Dasyclonium
J.Ag.
REFERENCES
(1) Agardh, J.G., 1894, Analecta Algologica
Contin. IL Lunch Univ. Arsskr. 30 (Afd.
2, Nr 7), 98 + 1 pp., 1 pi.
(2) Falkenberg, P., 1901, Die Rhodomelaceen
des Golfes von Neapel und Angrenzen-
den Meeresabschnitte, Fauna und Flora
des Golfes von Neapel, 26: i-xvi, 1-754,
10 figs., 24 pis.
(3) Fritsch,F.E., 1945, The Structure and Repro-
duction of the Algae, Vol. 2, Cambridge,
xvi -f 939 pp., 336 figs.
(4) Kylin, H., 1928, Entwicklungsgeschichtliche
Florideen Studien., Lunds Univ. Arsskr.
N.F. 24 (Avd. 2, Nr 4), 127 pp., 64 figs.
(5) _ , 1930, Uber die Entwicklungsges-
chichte der Florideen, Lunds Univ.
Arsskr. N.F. 26 (Avd. 2, Nr 6), 104 pp.,
56 figs.
(6) _., 1937, Anatomic der Rhodophy-
ceae. In Linsbauer, K., Handbuch der
Pflanzenanatomie, II. Abt. Band VI,
2 Teilband, Berlin, viii -f 347 pp., 252
figs.
(7) , 1956, Die Gattungen der Rhodo-
phyceen, xv + 673 pp., 458 figs, (in-
cluding Nachtrag by E. Kylin), Gleerup.
Lund.
VOLUME 4
BOTANY
(8) Oltmanns, F., 1904, Morphologic und Bio-
logic der Algcn, Ed. 1, Spezieller Teil.
Jena, vi + 733 pp., 476 figs.
(9) , 1922, Morphologic und Bio-
logic der Algen, Bd. 2, 2nd ed. Jena,
iv + 439 pp., 325 figs.
(10) Papenfuss, G.F., 1944, Structure and Taxo-
nomy of Taenioma\ including a discus-
sion on the Phylogeny of the Ceramiales.
Madrona, 7: 193-214, inch pis. 23 and
24, 1 fig.
(11) Rosenberg, T., 1933, Studien iiber Rhodo-
melaceen und Dasyacecn, Lund (Thesis),
87 pp., 25 figs.
(12) Scagel, R.F., 1953, A Morphological Study
of Some Dorsiventral Rhodomelaceae,
Univ. Calif. Publ Bot., 27 (1): 1-108,
20 figs.
(13) Schmitz, F., 1883, Untersuchungen iiber die
Befruchtung der Florideen, Sitzungsber.
K. Preuss. Akad. Wiss. Berlin, 1883:
215-258, pi. 5.
(14) , 1889, Systematische Ubersicht
der Bisher Bekannten Gattungen der
Florideen, Flora, 72: 433-456, pi. 21.
(15) , and Falkenberg, P., 1897, Rho-
domelaceae. In Engler, A., and Prantl,
K., Die Natiirlichen Pflanzenfamilien....
Teil l,Abt.2: 421-480, figs. 240-266.
(16) Silva, P.C, 1950, Generic Names of Algae
Proposed for Conservation, Hydrobio-
logia, 2:252-280.
(17) , 1953, A Review of Nomenclatu-
ral Conservation in the Algae from the
Point of View of the Type Method, Univ.
Calif. Publ. Bot., 25 (4): 241-324.
243
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Symposium: Ethnobotany of Thailand and Contiguous Countries
OPENING REMARKS ON ETHNOBOTANY AND ECOLOGY
THEODORE P. BANK II
Institute for Regional Exploration, Ann Arbor, Michigan, U.S.A.
The subject matter of the current symposium
is ethnobotanical. It seems, however, that this is
also the subject matter of at least several other
sessions of the Congress. I have counted two
dozen papers given elsewhere which are directly
concerned with ethnobotany, including such
topics as the origin of paddy field culture in
ancient Japan, the study of variation of taros and
kumaras and its possible ethnobotanical signi-
ficance, a symposium on vernacular names (Bota-
ny Division), several papers on the geographical
differentiation and origin of barley, banana trees,
and other cultivated plants (Agriculture) as well
as papers forming a part of the symposium on
the humid tropics, sessions on coconut problems
and on shifting agriculture.
The dispersion of ethnobotanical reports
throughout the Congress is unavoidable, of course,
for it is probably the only way of including
ethnobotanical data in specialized symposia. In
other respects it is unfortunate. It means that
many papers will not reach certain persons who
are interested in them until after they are pub-
lished, which may be a matter of years. Further-
more I have noticed in attending sessions
that many discussions suffer because they lack
authoritative opinions which probably would be
present if there were a concentration of ethno-
botanical material in a single, jointly-sponsored
session or sessions.
Today, I would like to begin our symposium
on the ethnobotany of Thailand and contiguous
countries by offering a definition of our subject.
It is evident, I think, that there is a difference of
opinion concerning the nature and scope of
ethnobotany as witnessed, for example, by the
diversity of subjects covered in our symposium
this morning. For purposes of discussion, I
propose a definition of ethnobotany that is largely
accepted in America, that is: ethnobotany is a
study of the interrelations of primitive man and
plants.
There is, I believe, a distinction between eth-
nobotany and economic botany. Ethnobotany
should deal with verbal traditions only, except
for those traditions which have recently found
244
their way into the literature, whereas economic
botany is more concerned with modern concepts
of the science of plants and their uses. The
word "primitive" often carries with it unfortunate
social implications; it has different meanings for
different people. Yet it is nevertheless our best
key to the real content of ethnobotanical study.
Ethnobotany is primarily concerned with the
interrelations of plants and primitive man in the
sense that "primitive" denotes only a lack of any
written language and therefore the preservation
of traditions by verbal means alone. This is,
I think, the accepted anthropologicall view.
Chemical analysis of useful plants is not usually
considered a part of ethnobotanical work, where-
as the following are: the study of plant debris in
archaeological deposits, studies of plant patterns
in the vicinity of old village sites, pollen analysis,
and radiocarbon dating.
The earliest use of the term "ethnobotany" was
by J. W. Harshberger in 1895. It was first used
narrowly in reference to the use of plants by
aborigines. More recently, most authors agree
that ethnobotany should deal "not only with
plant uses, but with the entire range of relations
between primitive man and plants." (V. H. Jones,
1941, p. 219).
It is evident, therefore, that our study must be
a broad one, for man and plants are co-existent
over a broad spectrum. There is necessarily
ecological interaction between them, as Jones
points out. Perhaps of even greater significance
is the fact that there are cultural forces that must
be reckoned with, and these make our study
uniquely different from animal ecology. It is
necessary that we make use not only of bio-ecolo-
gical concepts in dealing with plants and man in
any ethnobotanical work, but also we must bring
into our study the various concepts which an-
thropologists have regarding cultural dynamics.
It goes without saying that ethnobotany is an
interdisciplinary study that overlaps a number of
established scientific fields. Perhaps, as some
have suggested, the real function of ethnobotanic-
al work is to iilucidate the interaction of primi-
tive man and plants on a cultural level, not on a
VOLUME 4
BOTANY
biological level, and we should leave the biologi-
cal interpretations to the plant and animal scien-
tists. To my mind, however, such a restriction
on the scope of ethnobotany is a step in the
wrong direction. Ethnobotanical study will be
of far greater significance if it seeks to overlap,
if it brings together various approaches of various
disciplines for the better understanding of both
the cultural and the biological factors involved
in the interrelations of plants and primitive man.
The main point I would like to leave from this
perhaps overly-long dwelling on already pub-
lished ideas is that ethnobotany is really an inte-
gral part of the newly developing science which
various authors have called "human ecology"
and "cultural ecology." The content, philoso-
phies, and methods of this new science are
yet to be satisfactorily defined. Various authors
have their own views of just what "human
ecology" is. But from the strides which bio-
ecology has made toward illucidating for us the
forces of Nature, and their effects upon biota
and environment, I feel certain that the study we
are beginning to call human ecology will be able
to do the same for a better understanding of the
forces of culture. It is significant that the term
"ecology" implies that specialists will be involved
in the study not only from anthropology but from
the biological sciences as well. Ethnobotanists
will undoubtedly be called upon to make major
contributions to the combined effort.
245
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
THE NATURE AND STATUS OF ETHNOBOTANYt
VOLNEY H. JONES
University of Michigan, Ann Arbor, Michigan, U.S.A.
(Abstract)
"The Nature and Status of Ethnobotany" was
published by Professor Jones in 1941, when it
appeared in Chronica Botanica, Volume VI,
Number 10, pp. 219-221. Because it contains
some pertinent thoughts on the content and status
of ethnobotany which arc worth repeating for
the benefit of this symposium on "The Ethno-
botany of Thailand and Contiguous Countries,"
I have taken the liberty, with Professor Jones'
permission, of extracting portions for this paper.
Jones points out in his original article that a
large amount of data associated in some manner
with the relationships of primitive man and plants
has accumulated in the various literatures of such
diverse disciplines as botany, anthropology,
linguistics, agriculture, horticultural science, and
geography. These data are concerned with econo-
mic botany, plant lore, properties and value of
economic plants, the origins of cultivated plants,
plant remains in archaeological sites, and plant
names and plant knowledge of primitive peoples.
In America these various aspects of the study of
the relationship of aboriginal peoples and plants
have come to be known under the term "eth-
nobotany."
Among the students of ethnobotany, the plant
scientist is primarily interested in knowing what
plants were and are used by primitive man, how
they are gathered and utilized, and what effect
man has had upon the dispersal of plants. The
former distributions of plants as indicated by
archaeological deposits should offer the botanist
valuable data to supplement his own research in
plant geography. The lists of plant names and
ethnological and archaeological occurrences of
plants have been shown to be of value in the study
of the origin and dispersal of cultivated plants.
The plant ecologist is interested in the influence
of primitive man on the plant environment,
especially the effect of man's activities in dis-
turbing the otherwise normal processes of plant
succession. The anthropologist, on the other
hand, is more interested in the manner in which
primitive man adapts himself to his plant en-
vironment, what plants he uses, and how his
economy, activities, and thoughts are influenced
t Presented in abstract by T.P. Bank II.
246
by the plant world. The cultural implications,
in other words, are paramount over the botanical.
The ethnobotanist, speaking of the professional
worker who specializes in ethnobotany, is most
useful perhaps in correlating the data on these
and similar problems and in presenting his results
in a form that is useful to either the plant scientist,
the anthropologist, or both.
Few individuals, either in the United States or
elsewhere, have given their entire attention to
ethnobotany alone. Ethnobotany for its own
sake is practiced by perhaps a handful of workers.
On the other hand, anthropologists have gone
into ethnobotany to solve certain problems, and
botanists have become part-time anthropologists
for the same reason. Recently there has developed
in America a healthy and productive cooperation
among botanists, anthropologists, and linguists,
and a more concerted attack has been made
upon ethnobotanical problems of mutual interest.
Although many ethnobotanical observations
were made much earlier, it is only since about
1850 that any great amount of substantial pro-
gress has been made in the correlation of ethno-
botanical information. In Europe the most
notable early work was that of such men as
Alphonse de Candolle, Ungcr, Targioni-Tozzetti,
Bretschneider, and Wittmack in applying ethno-
botanical data to the solution of problems of the
origins and distributions of cultivated plants.
More recently, emphasis in Europe has been on
the development and application of techniques
for the removal and study of archaeological plant
materials. Some of the early work in America
was done by Europeans; but since about 1875,
American ethnobotany has progressed more or
less independently. A rather extensive and valu-
able literature bearing on the relations of the
Indians of North America and plants has accu-
mulated. American archaeological plant material
has yet to receive the attention which it deserves.
Although little has been done on the ethnobotany
of South America except in Peru, there would
seem to be remarkable opportunities in that
continent.
VOLUME 4 BOTANY
The results of the intensive surveys of the world. In the collecting of such products and
expeditions of the Soviet Union under the particularly in the obtaining and recording of
direction of Vavilov seem to indicate that many the knowledge of the natives concerning uses,
kinds and varieties of cultivated plants distinct properties, cultural treatment, and other such
in their genetic constitution, adaptations, and information, ethnobotanical experience and the
virtues from any now in the agriculture of civilized ethnobotanical approach should be valuable,
man may yet be obtained in various parts of the
247
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
MATERIALS USED FOR THATCHING IN THAILAND
TEM SMITINAND
Chief, Section of Botany and Zoology, Forest Products Research Division, Royal Forest Department, Bangkok, Thailand.
In rural parts of Thailand, thatches are still
in use, as they are cheap and easy to procure.
The materials used vary in different parts, but
fall roughly into three categories:
TREE LEAVES
In northern Thailand, people collect mature
leaves of young trees of Mai Pluang (Dipterocar-
pus tuber culatus ) , abundant in the dry deciduous
forests. In thatching, these leaves are arranged
in an overlapping row on a thin piece of bamboo,
about one metre long, fastened together with
bamboo or rattan. This kind of thatch, when
properly seasoned, is not susceptible to insect
attack and is not inflammable. The thatch lasts
for two to three years.
PALM LEAVES
At least four varieties of these leaves are used
in thatching:
Kaw (Livistona sp.) is a tall, erect palm of 15 to
20 metres in height, common in evergreen forest.
The leaves are fan-shaped and are much used
by hill tribes who fold them in half into a
roughly triangular shape which are then ar-
ranged to overlap each other, being secured to
the building framework with rattan. The
thatch does not last long, but as these hill tribes
move every three or four years, this does not
matter.
Wai (Daemonorops sp.) is a kind of rattan,
known in certain places as Wai Chak. There
are two or three species of this rattan whose
leaves are feather-like in shape with thin spines
growing on the pinnae and larger spines grow-
ing on the petioles. The leaves are folded in half
and placed in pairs to three long rattan canes
to which they are secured by split rattan bind-
ing at intervals of 5 cm as illustrated below.
When a required length has been made, the
whole sheet is rolled up, ready for laying upon
the building framework to which it is secured
by split rattan binding. These thatches last
from two to three years.
Chak (Nipa fruticans), a feather-leaved palm
with underground stem, grows gregariously in
tidal forests and provides material for roofing
for people living in the central plain and coastal
area. The pinnae of matured leaves are cut
from the axis, one-third of their length is then
248
VOLUME 4
BOTANY
folded and placed overlapping each other on a
bamboo split and bound with Wai nam (Flagel-
laria indica), a riperian species of scandent
shrub.
Sakhu (Metroxylon sp.), known commercially
as sago palm, is another form of thatching pre-
pared in a like manner to Chak. This is com-
monly used in the Malay peninsular.
GRASS LEAVES
Two kinds are used, Kha (Imperata cylindri-
cata), commonly known as Alang-Alang, and
Faek (Vetiveria sp.).
Kha is extensively used by local people in every
part of the country, whereas the use of Faek is
limited to central and northeastern parts. The
method of preparation is similar. After the
grasses are reaped and dried in the sun, they are
made up into bundles. Six to eight leaves are
folded along one-third of their length on a piece
of bamboo about one meter long and are secured
by a string of Paw (Sterculia sp.). In certain
parts, branches of Khon Tha (Harissonia per-
forata) are used instead of bamboo. The thatch
lasts for two to three years, and it is interesting
to note that after the grasses are reaped the whole
area is burned down, as it is thought that this
will produce a good crop for the next season.
249
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
SOME FOOD PLANTS IN THE FORESTS OF THAILANDt
K. SAMAPUDDHI
Royal Forest Department, Bangkok, Thailand.
Ancient Thai literature often described the
forests as abounding in food plants and fruit
trees. In the well-known story of Vesandorn
Chadok, for instance, the part of the Himapharn
forest into which Prince Vesandorn was exiled by
his Royal Father, King Son Chai of Siplee, was
described as full of fruit trees like wild banana,
langsat (Lansium domcsticun i ) , Mafai (Baccau-
rea sapida), Krathon (Sandoricwn koetjape),
Lamyai (Euphoria longana) etc., on which
the Prince could manage to survive without much
trouble. It is felt that there is some truth in this,
for, in the northeastern region of Thailand,
especially, villagers more or less depend on the
forest for their daily meals. Greens, bamboo
shoots, mushrooms, etc., are gathered from the
nearby woodland. They are cooked, roasted, or
fried with either fish or meat and eaten with
rice, the staple food of the teeming millions of
the East.
During his tours through the forests of Thai-
land, the writer has always been interested in
food plants, and, in order to give travellers and
adventurers an idea of how one could survive
when lost in the forest, he has compiled a list of
plants from which food material can be obtained.
The list is by no means exhaustive, and the com-
mon herbs or fruit trees which are already well
known and not actually growing wild have been
omitted. The edible parts of the plants are either
the flowers, fruits, shoots, leaves, stem, roots,
bulbs, or tubers.
For practical purposes, the forests of Thailand
may be classified into four main types, namely,
the evergreen forest, the mixed deciduous forest
(including teak forest), the deciduous dipterocarps
forest, and the mangrove forest. For the sake
of convenience, the plant names listed here have
been grouped under the various types of forests
in which they are found.
Table I.
Food plant of the evergreen forest.
No.
Local name
Bot. name
Edible part
How prepared
Taste
Remarks
1.
Kum Nam
Crataeva nurvala
Young leaves
Pickled in vinegar
Almost taste-
A medium-sized
/ ' * \
Ham.
less
tree
2.
Kl'uey-pa
Musa spp.
"Stem" as a
Taken fresh
Wild banana is ra-
^nmfii M^
source of water,
ther full of seeds
also fruit
3.
Chamuang
Garcinia cowa Roxb.
Young shoots and
By boiling with
Slightly sour
A medium-sized
f \
leaves
meat or pork to
tree in Southern
VTJ/JJ 13)
form an appetiz-
Thailand
ing broth
4.
Chaluerd
Premna integrifolia
Young shoots
To be boiled or
Crispy and al-
A medium-sized
y 4
Linn.
baked before be-
most tasteless
tree
^ '
ing taken with
chilli paste
5.
Tao
Arenga pinnata Merr.
Young shoots
May be taken
Sweetish
A palm with simi-
(T3)
fresh or boiled
lar appearance to
nipa palm
6.
Book
Amorphophallus spp.
Young stems
May either be
Sweetish
A herb; must not
( \ i f*^
fried or used as
be washed with
an ingredient in
coldwater. The
the preparation
contact with cold
of Thai curry
water creates a reac-
tion that causes a
irritable itch in the
throat when eaten.
Only boiled water
must be used.
t Presented by T.P. Bank II.
250
VOLUME 4
BOTANY
Bot. name
Edible part
How prepared , Taste
Remarks
Mo.
Local name
7.
Bua
On)
Nymphaea lotus L.
Petioles
May be taken ! Almost taste-
fresh or fried or less
boiled '
Aquatic; found in
ponds or swamps
8.
Phak Koot
(wnnw)
Athyrium esculent um
Copel.
Young leaves
To be boiled and
prepared as an or-
dinary vegetable
Almost taste-
less
A fern
9.
Mafai-pa
(Uvlr^lll)
Baccaurea sapida Mu-
ell-Arg.
Fruits
Taken fresh
Sweet or acid
A forest fruit tree
10.
Madua-nam
F/CWJ scandens Roxb.
Fruits
Taken fresh
Sweetish
Sort of a fig tree
11.
(jJ^lflOUl)
Madua-din
(USLttOftlj)
Aganosma marginata
G. Don
Young shoots
and leaves
Taken fresh with
chilli paste or
pla-ra (fish paste)
Slightly astrin-
gent
A scandcnt climber
12.
13.
Wild Raspberry
Canes
Omo)
Rub us spp.
Calamus spp.
Fruit
Young shoots
Taken fresh
To be boiled and
taken with chilli
paste or Pla-ra
(sort of sal ted fish)
Acidic
Slightly bitter
A shrub usually
found in mountain
evergreen forest
14.
Soke
flHin)
Saraca indica Linn.
Young leaves and
inflorescence
To be used as an
ingredient for pie-
paring aThai curry
Slightly sour
Small tree growing
along stream banks
15.
Hucha-niang or
Niang
OlTfEllim)
Pithecolobium jiringa
Prain.
Seeds both young
and germinated
Slightly astrin-
gent
Taken raw with
chilli paste and
curry
Tree 15-20m tall
16.
Kariang
(nsimrn)
Parkia javanica Merr.
Germinated seeds
Slightly astrin-
gent
Taken raw with
chilli paste and
curry
Tree 30-40 m tall
buttressed
17.
Sataw
(ttswo)
Parkia speciosa Hassk.
Seeds
Slightly astrin-
gent
Taken raw or
pickled, with
chilli paste and
curry
18.
Kra
(ms)
Elateriospermum tapos
Bl.
Mature seeds
Slightly sweet
Pickled and tak-
en with curry
or used in salad
Tree 20-30 m tall,
lacticiferous
Table 2.
Food plants of the mixed deciduous forest.
Taste
Remarks
No.
Local name
Bot. name
Careya arborea Roxb.
Edible part
How prepared
1.
Kradone or pui
Young leaves
Taken fresh
Slightly sour
A medium-sized
tree
2.
3.
Krathon
(msnbw)
Kae-foy
(Uflftati)
Sandoricum koetjape
Merr.
Dolichandrone crispa
Seem.
Fruit
Flowers
Taken fresh
To be boiled or
roasted before
being taken with
sauce or chilli paste
Sour and slight-
ly astringent
Almost taste-
less
A big forest tree
4.
Chakan
(vrfrw)
Pi/w sp.
Young shoots
To be boiled be-
fore being taken
with sauce or
chilli paste
Aromatic
A vine
251
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Local name
Bot. name
Edible part
How prepared
Taste
Remarks
Matat
(USfllfl)
Dillenia indica Linn.
Young unripe
fruit
Used as an ingre-
dient in prepar-
ing a Thai curry
Sour
A large tree
Teiw-klicng
%
Cratoxylon polyan-
thum Korth.
Young leaves
May be taken
fresh
Slightly sour
A medium- sized
tree
Bamboo shoots
Bamboo Seed
Dendrocalamus spp.
Bambusa spp. & Oxy-
tcnanthera spp.
Shoots
Seeds in the form
of grains like rice
grains
Usually boiled be-
fore being eaten
Boiled like rice
Almost taste-
less
Like rice; may
be substituted
for rice in time
of famine
Found scattered
about on the forest
floor in bamboo
flowering areas
Kloy
(miou)
Dioscorca hispida
Dennst.
Tuber
Boiled with salt
water
Starchy like
potato
May cause nasty
itch in the throat
if not properly
prepared
Peka or Marid-
Mai
(wmmfl
Oroxylum indicum
Vent.
Seed in the young
pods
By boiling
Almost taste-
less
A shrub
wsiflljj)
Makok
(wsnon)
Tamarind
(usinu)
Spondias pinnata
Kurz.
Tamarindus indica
Linn.
Young leaves and
fruits
Young leaves and
fruits
May be eaten
fresh
May be eaten
fresh
Slightly sour
and astringent
Acidic
A big tall tree
Found growing
wild in old village
sites; a big tree
Lcb yciw
(ifl 111)10 in)
Zizyphus oenoplia
Mill.
Fruits
May be eaten
fresh
Acidic
A thorny shrub
Maroom
(wsijj)
Moringa oleifera
Lamk.
Young pods and
flowers
To be boiled be-
fore eating; may
be eaten fresh
Almost taste-
less
Found growing in
old village sites
with chilli paste
Wan Poh
i
Kaempferia galanga
Linn.
Tuber
May be eaten
fresh with chilli
Crispy and al-
most tasteless
A herb
paste
Wah
(VI VI)
Sadao-pa
Syzygium cumini
Skeels.
Antelaea azadirachta
Adelb.
Fruits
Young leaves and
inflorescence
Eaten fresh
By boiling
Sweet
Rather bitter
A big tree
A medium sized
tree
Samaw-Thai
(mje)1nij)
Terminalia chebula
Retz.
Fruit
Taken fresh
Slightly acid
and astringent
A big tree
In-ta-nin
(Surma)
Lagerstroemia macro-
capa
Young shoots
To be boiled be-
fore eating with
chilli paste or pla-
ra (fish paste)
Almost taste-
less
Siew
(isttn)
Piliostignaa malaha-
sica Benth.
Young leaves
May be boiled
with meat or pork
to make a tasty
broth
Acid
A large shrub
Cha-om
(tftou)
Acacia insuavis Lace
Young shoots
and inflorescence
May be boiled or
baked or taken
fresh with chilli
Slightly bitter
Large wood and
thorny, scandent
shrub
paste or sauce
Kracheo
(msrotn)
Curcuma sp.
Inflorescence
Taken fresh or
steamed with chil-
li paste
Slightly sweet
and aromatic
Herb with bright
red bracts
252
VOLUME 4
BOTANY
Table 3.
Food plants of the deciduous dipterocarps forest.
No.
Local name
Bot. name
Edible part
How prepared
Taste
Remarks
1.
Kabok or Mameun
Irvingia ma lay ana
Endorsperm of
By roasting until
Like melon
A big tree
(ntunvnouswu)
Roxb.
the seed
well cooked
seed
2.
Pak-wan
Meliantha suavis
Young leaves
By boiling into
Sweetish, de-
A shrub. At times,
(whvmu)
Pierre
a broth or used
licious
especially in the
in a curry
rainy season, it
may turn deadly
poisonous a
mystery to both
villagers and bo-
tanists
3.
Phan-ngu
Amorphophallus spp.
Stems By boiling or
Like lotus
A herb with flow-
(wu)
frying with meat
or pork (if avail-
stem, almost
tasteless
ers in the form
of spadises
i
able)
4.
Uang-mai-na
Costus speciosus
Young shoots
By boiling until
Like asparagus
A monocot herb
(ioo<Miw"iuvn)
Smith
well cooked
i
5.
Phayom
Shorea talura Roxb.
Inflorescence
Used in a curry
Slightly astrin-
A big tall tree
(WSUOIJ)
i
gent
Table 4.
Food plants of the mangrove forest.
No.
Local name
Bot. name
Edible part
How prepared
Taste
Remarks
1.
Kong-kang or
Panga
(Irurmmomrn)
Phizophora mucronat
Lamk.
Young shoots
By boiling and
taking with sauce
or chilli paste
Just crispy and
almost tasteless
2.
Kilek-pa or
Samaesarn
,* d ft *
(umfininmo
Cassia garrettiana
Craib.
Young leaves
and flowers
By boiling
Slightly bitter
A small tree
3.
ummn)
Chik
(in)
Barringtonia asiatica
Kurz.
Young shoots
May be eaten
fresh
Slightly astring-
ent
Found in swamps
4.
Samet Kao
(ffljjflim)
Melaleuca leucaden-
dron Linn.
Young shoots
By boiling be-
fore taking with
chilli paste
DISCUSSION
T.P. BANK ii : Would botanists and ethnobotanists
study the ritual and folklore attached to these plants, and
not leave it at a list of plant names?
253
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
THE USE OF TALIPOT PALM LEAVES AS
WRITING MATERIAL IN THAILAND
KASIN SUVATABANDHU
Department of Agriculture, Ministry of Agriculture, Bangkok, Thailand.
Leaves of the palm known in Sanskrit as tala
or tal were used in India as writing material for
a very long time. The name tala or tal was con-
fined to Borassus flahellifer, Corypha umbraculi-
fera and Phoenix syhestris. When the Indians
brought the custom of writing on the tala leaves
to the Malesian area where Borassus flabellifer
was plentiful, they also introduced that word
into the local language. Tala became ron-tal
and later, by metathesis, lontar; that is how the
common name lontar, or lontar palm for Boras-
sus flabellifer was derived.
Whereas in Ceylon the leaves of Corypha wn-
braculifera, the talipat (tali or tal, and pat =
leaves), were used, because the Indian who
brought the custom of writing on the tala leaves
used the Corypha leaves, Corypha unibraculifera
being plentiful in Ceylon. The word talipat
later became talipot and the tree is called by the
Anglo-Indian the talipot palm.
The use of the talipot palm leaves as writing
material was very likely adopted in Ceylon before
B.E. 900, because it was known at that time that
the manuscripts of all the teachings of the Lord
Buddha had been written on talipot palm leaves.
Thailand had for a long time been connected
with Ceylon through religious channel. She
accepted Buddhism from Ceylon as national
religion. Some Thai monks were sent for training,
and on return they were supposed to bring back
authentic Buddhism to Thailand. It was recorded
that religious manuscripts written on talipot palm
leaves were brought into the Thai Kingdom just
a little before B.E. 1800. Copies were then made
using locally prepared talipot palm leaves vernacu-
larly known as "Bi-larn". The Thai have used
"Bi-larn" for writing religious manuscripts from
that period up to the present.
Talipot palm (Corypha umbraculifera) is found
indigenously in Thailand. It forms great groves
usually in the mixed, more or less dry and ever-
green forests. It is a big tree of erect cylindrical
trunk, 2-3 ft in diameter and 30-80 ft in height,
clothed throughout with petiole bases. Leaves
with stout petioles 5-10 ft long, lamina large 8-16
ft in diameter palmately pinnatifid, plicate, cleft
to about the middle into 80-100 linear lanceolate
254
acute or bi-fid lobes. The lobe is web-liked blade
in between two small ribs (veins). Flowering and
fruiting occur after the age of 30 years. The plant
dies after fruiting.
Talipot palm is classified as one of the forest
products which is a national treasure. Those
who want to exploit this particular palm must
apply to the Royal Forest Department for
licenses. Under the rules and regulations of the
Department, a license is valid for 3 years. After
a permission is granted the exploiter may enter
the Corypha grove and starts cutting juvenile
plicate leaves with a curved and long handled
knife. There arc 2 or 3 usable leaves in a tree,
but 1 or 2 leaves only can be cut near the base of
the lamina, leaving the youngest for future growth.
All cut leaves are then recut at both ends. Only
the middle part of the leaves which are about
27 inches in length are gathered and dried in open
air for 3 days, and then split along the ribs into
bi-lobed slips (2 lobes with 1 rib in between).
Bundles of one thousand bi-lobed slips tied up
near both ends are prepared and shipped to
Bangkok for sale.
The Bi-larn makers purchase these raw bi-
lobed slips, and grade them according to the
width. There are four grades varying from 2.25
inches to 1.75 inches wide. The undersized slips
are not used as writing material but are sold as
plaiting and weaving fibre for hats and other
articles.
After grading, the thin ribs of the bi-lobed
slips are removed, so that one raw bi-lobed slip
produces 2 sheets. These sheets are again selected
in order to eliminate poor specimens.
Each sheet is cut to standard size and two
holes, seven inches apart, used for binding purpose,
are made along the centre line of the sheet. (The
normal standard sizes of the sheets in inches are
1.75 x 20.5, 2.0 x 20.5, 2.0 x 21.0, and 2.25 x 22.5.)
Five hundred sheets are bound up into one packet
with talipot ribs piercing through the holes for
guiding. The packet is then placed in a wooden
frame and tightly pressed in order to straighten
and flatten the sheets. At the same time the edge
are planed for smoothness.
After that, the whole frames (with pressed
packets of palm sheets) are placed in a smoking
oven to be dried and coloured for about 48 hours.
Rice husk or the palm leave waste may be used
as fuel. The dried smoked sheets are then ready
for writing.
The so prepared talipot palm leaves are called
"Bi-larn." In writing, this Bi-larn is placed on a
metal table and characters are engraved with an
iron stylus. Usually there are only 4 to 5 written
lines on a single slip. To render the characters
more legible, cow-dung or miller charcoal or soot
mixed with coconut oil is smeared on the engraved
leaf so that the written lines are blackened.
The written Bi-larn are cleaned with sand and
cloth and then bound into a packet by running
through the holes two small cords, one for each
hole, which are formed into two loops. The
number of leaves in a packet depends on the size
of the article or story. After binding into a book
form, the edges are painted with a red dye-stuff
known as "Chard", varnished with lacquer and
then plated with gold leaves. These are very
VOLUME 4 * ' BOTANY
useful as a protection against fungi and insects
which may occasionally attack Bi-larn.
Nowadays, most of the religious articles or
manuscripts are not transcribed, but printed.
The use of iron stylus for writing on the palm
leaf will be lost from the sight of younger gener-
ation in the very near future.
In printing, the prepared Bi-larn must first be
roasted over a fire to remove excess oil in the
leaf before it can be used by a printer. The
characters are printed on both sides of the leaf,
using generally 4 to 6 lines, and are arranged in
three columns.
The printed Bi-larn are brought together article
by article and then bound into books. The edges
are again dressed by planing, painting with red
dye-stuff, and gold plated.
The preparation of Bi-larn becomes a flour-
ishing home industry in the Thai Kingdom;
the palm leaves are used not only for religious
and language writings, but also for making into
cords, ropes, and plaiting or weaving material.
Fig. 1. Talipot palm leaf.
255
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Fig, 2*' --Grading of the
256
Fig. 3. Removing of the ribs from the bi-lobed slips.
VOLUME 4
BOTANY
4 Cutting to
Fig. 5. Trimming for uniformity.
257
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
;;fr~ ,
V''.- j 2. ^ >j"'
6, Binding in ihc
258
Fig. 7. Planing for smoothness.
S{ VOLUME 4
BOTANY
In hot air
Fig. 9. Removing oil over a fire before printing.
259
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Fig.
260
Fig. 1 1 .Sorting the printed leaves for further binding.
VOLUME 4
DISCUSSION
BOTANY
F.R. FOSBERO: I would like to ask Mr. Kasin about
the geographic range of Talipot palm in Thailand.
K. SUVATABANDHU: So far only in the North and
Northeast, but in the South another species occurs, and
all species are used for writing.
C.G.G.J. VAN STEENIS: I am glad to have heard the
detail on the talipot palm, that its use for writing has been
imported to Thailand from Ceylon. I only know it from
Bali where, however, the leaves used are derived from the
lontar palm ( Borassus flahellifer ) which has been used for
it since Hindu times and has been introduced by Hindu
culture and wandered together with the invaders. The Bali
lontars are mainly on religion, medicinal prescripts, history,
art, etc. and thus the old ones contain much written
information. They have been a great subject for study by
ethnologists."
F.R. FOSBLRG: I would ask Prof. Van Steenis if the
famous Sumatran manuscripts, such as Prof. Bartlett has
collected, are written on this sort of material.
C.G.G.J. VAN STEENIS: As to papers in North Sumatra,
I do not know particulars. The palms (Corypha, Borassus)
are not growing in the mountains where the Batah live,
but they occur on the sub-seasonal northern coastal
plains of Atjeh (northernmost Sumatra).
M.L. STEINER: In the Philippines palm leaves and
bamboo strips are used.
261
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
NOTES MADE FROM LOCAL KNOWLEDGE OF
THE USE OF POISONOUS PLANTS BY THE THAI PEOPLE
TEM SMITINAND
Chief, Section of Botany and Zoology, Forest Products Research Division, Royal Forest Department, Bangkok, Thailand.
The Thai people are, by nature, migratory, as
can be seen from their history. During their
migrations, they learned much from nature
around them. At one time, however, they did
settle down, but hostile and aggressive neigh-
bours forced them to move. Their migration
over a period of thousands of years stretch from
the Altai Mountains in Mongolia to the present
Thailand.
In their history, the Thai people made much use
of poison in both war and peace time. Most of
the poisons used came from plants, and in spite
of the impact of Western civilization upon the
Thais, they are still used today.
The poisons now in use are for poisoning arrow
tips, insecticides, and fish-poisoning, as follows:
Parts usec
1
Local name
Botanical name
Habit
Habitat
arrow
insect
fish
Khamin kru's
Anamirta cocculus
Liane
Fruits
Fruits
Evergreen
& Seeds
& Seeds
forest
Nong
A ntiaris toxicaria
Tree
Latex
.
Evergreen
forest
Sakae dong
Cocculus laurifolius
Liane
Bark
Bark
Evergreen and
mixed forests
Saba
Entada phaseoloides
Liane
,
Bark
Evergreen
forests
Prik pa
Ervatamia corymbosa
Shrub
Bark&
,
. .
Evergreen
Roots
forests
Bua khru'ng sik
Lobelia chinensis
Herb
Leaves
Evergreen for-
est & cultivated
Makham di khwai
Sapindus rarak
Tree
Aril
Evergreen
forest
Tatum bok
Sapium insigne
Tree
Latex
- .
Evergreen and
mixed forests
Rak pa
Seme car pus curtisii
Tree
Latex
Evergreen
forest
Nawn tai yak
Stemona tuherosa
under-
Roots
Evergreen and
shrub
mixed forests
Khika daeng
Trichosanthes brae teat a
Climber
Fruits
Leaves
Mixed forests
Nong khru's
Strophanthus scandens
Liane
Latex
Evergreen
forest
Salawt
Croton tiglium
Shrub
Bark
Seeds
Mixed forests
or small
tree
Rak
Melanorrhoea usitata
Tree
Later
, .
Mixed forests
Lian
Melia azedarach
Tree
Fruits
Bark &
Mixed forests
& Seeds
Roots
Salaeng chai
Strychnos nuxivomica
Tree
Seeds
,
Mixed forests
Wan nam
Acorns calamus
Herb
Rhizome
Riparian
Thawn
Albizzia procera
Tree
.
Bark
Mixed forests
Lai nam
Derris elliptica
Climber
Stem
Stem
Stem
Riparian
Khao san
Phyllanthus columnaris
Tree
Bark
Riparian
262
VOLUME 4
BOTANY
Local name
Tatum thale
Man kaew
Botanical name
1 Excoecaria agallocha
Pachvrhizus erosm
Habit
Parts used
arrow j insect [ fish
Tree j Latex i Latex
Habitat
Tuber-
ous
climber
Mangrove
swamps
Leaves Cultivated
i & seeds
263
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
TEA IN THAILAND
PRASIDHI PHUMXUSRT
Mae Malai Farm, Mae Rim, Chiang Mai, Thailand.
The genus Camellia was originally thought to
be indigenous to China, but later discovered that
the location of Nature's original tea-garden is
in the monsoon districts of Southeast Asia. The
monsoon, one of the most vital producing factors
to this part of the world, is called "Morasoom
Tawan Tok Xieng Tai" or Southwest monsoon
in Thailand, it blows across the Indian Ocean
towards eastern Tibet, world's top-most plateau,
carries moisture-laden clouds over the northern
mountain ranges of Thailand, precipitates along
its route as rains to an average of well over sixty
inches per annum.
The climate in Chiang Mai, Chiang Rai,
Lampang, Mae Hongson, Nan and Phrac may be
divided into three distinct seasonal categories;
the cool and dry lasting from November to
February, the hot and dry from March to early
May, and the hot and humid from late May to
October. The season in these vicinities is fa-
vorably comparable to many tea producing
countries.
The topography of northern Thailand may be
divided into three regions: (1) The low-land or
paddy field region, 'Tung Na." (2) The dipter-
ocarpus region, "Pa Daeng." (3) The evergreen
region, "Pa Dong Dib," with an altitude ranging
from 200 to 300 metres MSL, 300 to 800 metres
MSL, and 800 to 1,500 metres MSL respectively.
The one that is concerned most in this report is
the evergreen mountain region, the soil with a
pH value of 4.5 to 5.5 of which has its origin
from Gneisses rock and Quartzitic sandstone.
Among these high hill ranges, there are tracks
after tracks of Tea-trees growing. Villages after
villages sprang up with the sites, people engaged
in preparing the tea-leaf into "Miang," this name
also implies to Shan, Laos, "Leppetso" to
Burmese, and "Pickled-tea" to the English speak-
ing world.
"Miang" is consumed by Thai, Burmese, Lao,
Shan, and Singpho (a tribe inhabiting the upper-
reach of the Chindwin river in Northwest Bur-
ma).
The miang trees have not at all been cultivated,
they were allowed to grow in their natural states
and some acquired a height of from 3 to 6 metres,
when they are found too high for Plucking, the
264
top branches may be broken off or given a light
skipping.
Tn preparing "miang" the leaf is plucked, in
halves without stalk after the buds are all opened
and not as a bud and two leaves, four times a
year. The first plucking is locally called "Miang
Hoa Pee," lasting from April to June, the second
plucking "Miang Klang" from July to August,
the third plucking "Miang Soi" from September
to October and the fourth plucking "Miang Moey"
(Dew Tea) lasting from November to December.
After plucking and bundled, the leaves are
put into a round wooden container "Hai Nueng"
with plaited bamboo base and placed over a pan
of boiling water for steaming which last from 10
to 15 minutes. When the steaming process is
through the content is thrown on to bamboo
mat, people, sitting around kneading and retight-
ening the bundles and let cool for 5 to 10 minutes;
then the bundles or the day's plucking are placed
and stamped down in bamboo silo 5 to 6 feet
in diameter and 8 to 10 feet high.
The silo is made of plaited bamboo lined
heavily with wild plantain leaves, and kept adding
to the content with steamed-leaf throughout the
season ; at each filling the contents must be pressed
down by tramping and planked with heavy stones.
Under pressure, no doubt, the juice is oozed out,
it is collected, boiled to a very thick consistency
called "Nam miang" which is used as one of
"Miang" delicacies. The leaf may be left in the
silo for a period varying from 15 days to several
months or until the commencement of the next
season's plucking.
Raw "miang" is very pungent and people pre-
fers longer pickled one that has a sour taste.
Miang has the smell of wet humus decomposed
in a limited air supply (ensilage). It also has a
slight sweet smell of fermented with yeast steamed
glutinous rice (Khao mark) or alcoholic fermen-
tation (zymurgy). The colour is bright yellowish
green when taken out from the silo and rapidly
becomes red when exposed for any length of
time, the quality deteriorated since then. For
preservation and handyness in packed-bull tran-
sport from the producing hills down to the nearest
motorized road or bullock-cart track, miang is
repacked into bamboo baskets shaping like a
VOLUME 4
BOTANY
mortar in lots of 60 to 80 kilograms. To assure
of having proper insulation the basket is both sides
smeared with viscid cow-dung, and heavily
lined with plantain leaves. Once the basket is
packed it never can be inspected in any transac-
tion until retailing to consumers. Quality is
guaranteed through honesty, the number of
bundles in the basket is recorded by a numerical
system engraved on a piece of bamboo stripe in
the shape of a multiplication sign, a plus sign,
a minus sign and dot sign ; each symbol represent-
ing the amount is well known to all engaged in
the trade.
The marts of the pickled tea are in Mae Rim,
Doi Saket, Ban Mae Ai in Chiang Mai district,
Cheh Horn, Lampang district and Wieng Papao
in Chiang Rai district. The storage for miang
in these marts is a very interesting sight, a huge
ditch swimming-pool-like is dug within the com-
pound and kept filled with water, all tea baskets
are kept submerged under water to keep them
from contacting with the open atmosphere.
Miang has a small export through Raheng-
Mae Sot land route to South Burma, the rest of
which is consumed locally. The common way of
consuming "miang" is to wrap the leaf with
salt and just hold in the mouth like lozenges, in a
deluxe fashion "miang" is wrapped with hog's
fat, pickled garlic, cocoa-nut-chips, ground-nuts
and salt. The fashion of consuming miang is
slowly dying out.
Thailand is undoubtedly a country where tea
could be grown in plantations commercially and
economically. Hill-tribes labour is plentiful and
who when employed to permanent agriculture will
be a great asset in minimizing the destruction
of forest wealth through "clearing culture"
(Kaingining) or "Rai luen loi."
In 1938 a small plantation was started the
progress of which is rather slow. At Huey
Tart a factory was built postwar time with modern
equipment to make black tea with leaf purchased
from smallholders. The production in 1952 was
nearly 1 50,000 Ibs. The product is sold in Thailand
only.
Developments and researches can be carried
on and on for further advancement.
265
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
THE MANUFACTURE OF SUGAR FROM THE SUGAR PALM IN
UPPER MANDAILING, SUMATRA!
ELISE TUGBY
Australian National University, Canberra, A.C.T., Australia.
INTRODUCTION
The sugar palm (Bargot), Arenga pinnata
(Wurmb) Merr., is found throughout Upper
Mandating, Southern Tapanuli, Sumatra 1 . The
area is one of complex mountain ranges bisected
by a rift valley. Height varies from 400 to 2,000
metres. The more temperate, wetter mountains
contrast with the hotter, drier valley. The Medan-
Padang road follows the latter, linking the main
market centres and providing the import and
export channel for the region. As the major
subsistence crop, wet-rice in the major and tri-
butary valleys, contrasts with dry-rice on the hill
slopes. Rubber and coffee are the important
cash crops.
Table 1 shows the altitude and general soil
type of the fifteen villages producing sugar for
the extra-village market, throughout the year.
The total number of villages in Upper Mandailing
is eighty-seven. From the table it can be seen
that economically significant palms grow on a
clay soil usually of volcanic origin. The first
nine villages listed are sited on lava flows ori-
ginating from the volcano Sorik Merapi. It may
be noted that most villages below the 500 metre
level occur in valleys on soils derived from sandy
sediments.
The highest village in the area, Pagargunung,
1,140 metres, is enclosed by hills rising to 1,310
metres. Here the palm was previously utilised,
but although the sap was sweeter than elsewhere,
the flow is now considered too little to warrant
collection. Here, too, coconuts were grown but
did not bear. This situation is concomitant
with Burkill's remarks that "the palm can be
Table 1.
Approximate height and general soil type of villages producing for the extra village market.
Name of Village
Si Banggor Djai
Si Banggor Djulu
Huta Tinggi
Huta na Male
Pasar Maga
Angin Barat
Si Antona
Pangkat
Si Ladang Djulu
Simangambat
Torrumbi
Si Bio Bio
Batahan Djulu
Simpang Duhu Dolok
Simpang Banjak Djulu
Approximate height
above S.L. in metres
760
960
800
950
580
620
600
480
430
480
950
800
800
930
1,060
Note: Term used throughout the paper are those current in upper Mandailing.
General Soil Type
Heavy clay derived from lava
Clay derived from limestone
and schist
from volcanic
breccia
from volcanic
breccia
,, from limestone
,, from slate
t Presented by C.S. Christian.
1 Upper Mandailing includes the Ketjamatan Kotanopan and the three villages forming the Si Ladang complex at the foot
of the mountain, Tor Sihite.
266
VOLUME 4
BOTANY
sjown at greater elevations than the coconut..." 2
During the month of fasting (Ramadan) in at
east eight other villages there is a little produc-
ion of sugar to supply the extra demand in that
jeriod, and output is also increased in villages
producing throughout the year.
Formerly, nearly all villages in the area pro-
iuced some sugar, but, with the introduction of
'ubber, production has become concentrated in
i few. The industry was stimulated during the
Japanese occupation, when a ready market
existed for sugar, but not for rubber.
In recent years the planting of palms has become
are. Distribution depends on wild animals,
iuch as the civet cat, which eat the fruit and void
he seeds. It is reported in many villages that
he number of palms has increased in the past
en years. The highest concentration of palms
s found in the Si Ladang villages, the scattered
elements of which lie within what is virtually one
arge grove. Elsewhere in Upper Mandailing
mly small groves occur and these outside the
dllage proper.
TAPPING THE PALM
A palm comes into use when about ten years
)ld. Before tapping commences, the common
)eduncle of the male inflorescence (Sajatan) is
>eaten with a wooden mallet ten times in a month,
causing the flowers (Apili) to increase in size
ind darken. This is taken as a sign that the
ap (Aek Ni Bargot) is flowing into the inflores-
cence and that it is ready for cutting.
On cutting, one flower-spike (Arirang) is left.
The heel of the knife blade is struck repeatedly
vith a mallet to force it through the peduncle.
Three days elapse between the first cutting and
mnging a bamboo container (Taguk) below to
collect the sap. Each morning a slice of the
>eduncle about two cm thick is cut off. The
low is then considerable, and a large container
s set to catch the sap at night when the quantity
exuded is large, while during the day a smaller
container is used.
The sap is collected every twelve hours. Each
norning the peduncle is given two taps with the
cnife handle before a slice is cut off, the size of
vhich may be progressively increased to about
hree cm. A leaf is tied round the cut to channel
he sap into the container below.
Access to a high peduncle is gained by climb-
ng the palm or by means of a notched bamboo
>ole.
Substances may be placed in the container
before it is fixed below the peduncle to retard
fermentation, e.g., the fruit of the mangosteen
(Manggis) or a pulverised root (probably ginger)
from the forest. A further preventative measure
is the washing out and smoking of all the bamboo
containers before reuse. Clean containers are
stored adjacent to the hearth.
The length of time one inflorescence will yield
is usually three months; but, where use is irregu-
lar, it may extend to nine. The yield is measured
in terms of the number of kilogrammes of
sugar produced per day. Inflorescences in use
(1956) were each yielding from one and a half to
three kilogrammes of sugar per day. A very high
price was being asked for an inflorescence yield-
ing five kilogrammes. The female inflorescence
(Alto) is distinguished, but never cut as the flow
of sap from it is considered too insignificant.
COOKING THE SAP
The fresh sap is boiled in large open pans in
a special out-of-doors cooking place. After
about one and a half hours, a thick liquid
(Tangguli) is obtained which is returned to a
bamboo container. This part of the preparation
reduces the volume of liquid (one large pan of
sap yields only one kilogramme of sugar) and
prevents the sap from souring due to fermentation.
When there is sufficient Tangguli on hand, a
second boiling for one to one and a half hours,
depending on the strength of the fire, produces
a red liquid "free of water." To the boiling
liquid a little coconut oil or crushed kernels of
the candle nut (Aleurites moluccana Willd) or
kernels from a bush (Djarat possibly Jatropha
curcas L.) is added to prevent the liquid boiling
over. A spoon with a perforated bowl may be
used to remove froth in the early stages of cooking
and any foreign materials. As the liquid thickens,
a little is tested in the air on the stirring spoon
to ascertain its setting properties and when ready
it is ladled into moulds.
The plank of wood on which the moulds are
placed is washed down to prevent contamination
from bacteria resulting from its prior use; this is
seen in Mandailing eyes as a sickness of the sugar.
The board may also be rubbed over with crushed
candle nut kernels. The moulds (Bila) are hoops
of bamboo made in three sizes, approximately
7, 14, and 21 cm in diameter and 1, 2.5, and 3.5
cm in height, respectively.
I.H. Burkill: A Dictionary of the Economic Products of the Malay Peninsula, 1935, Vol. 1, p. 231.
267
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
When set, the cakes (Paske) are pressed from
the moulds and allowed to cool and harden.
Cakes of like size are wrapped together in the
dried fibre of the banana trunk and tied in parcels
of a marketable weight. On account of the hy-
groscopic nature of the sugar the parcels are
stored above the hearth until marketed. Type and
quantity of paske produced are adjusted to the
market aimed at; the first, to the small scale
intra-village spending of child and household;
the second, to the market town and the largest to
the requirements of bulk export. In 1956, the
producers 1 selling price of a parcel of twenty
medium sized cakes, was about Rp. 10, at about
Rp. 2.50 a kilogramme.
Variations in the process of cooking are mainly
the result of differences in aptitude of the
producers, in particular, in their ability to judge
when the sugar has been sufficiently boiled. In
the local markets the quality of the sugar varies
considerably and, as a consequence, so does its
price. Sugar may be marketed black in colour
due to overcooking or smoke, moist and crum-
bling from undcrcooking, or adulterated by vari-
ous foreign bodies such as bees and palm fibre.
Or, again, though carefully prepared, it may be
held back until the market price rises, and then
finally be sold in a deteriorated state.
The site of the cooking place affects the quality
of the final product. Cooking takes place in
the precincts of the village, at the edge of the
rice fields, or in the arboreal gardens. In the
former case, unless sheltered by a roof and walls,
dust may be carried by wind from the bare
village streets into the cooking pans. In the
garden and field sites it is usual to build a shelter
to house the hearth and implements, but greater
care has to be taken to prevent bees and other
insects from being trapped in the liquid. But the
construction of a cooking place in gardens is
considered justified only where a number of palms
owned or hired by the tapper are clustered
together.
LABOUR FORCE
The care of the palm and the preparation of
the sugar are male occupations. Women assist
indirectly by providing some of the wood for the
cooking and directly in guarding the simmering
liquid and tending the fire. Although women
never participate in collecting the juice, in the
Si Ladang villages women do climb the palms to
collect the fibre.
268
OWNERSHIP OF PALMS
Ownership of the palms is established by
inheritance, by planting, and by finding palms
on land jointly owned by the village. Where a
person desires usufruct of another's palm, several
alternative arrangements exist. They are as
follows:
1. The hiring of a tree and repayment with
approximately a third of the final product.
2. The buying of the usufruct of an inflores-
cence at a fixed price after it has been estab-
lished that the palm is yielding.
3. The buying of the right to tap an untried
palm. As there are many unproductive
palms in the area, this represents a specula-
tion. However, the slightly lower price is
attractive, and it is said that some men have
acquired the skill of being able to estimate
the likelihood of a palm yielding.
ECONOMIC STATUS OF PALM SUGAR
Palm sugar plays an insignificant part in the
subsistence economy of the area. As a cash crop,
its actual and potential use is severely limited by
its storage properties which enforce a delicate
adjustment between production and market
demands. It has to compete with rubber and
coffee for which a market is assured and which
can be stored in anticipation of a favourable price.
Hence the industry is usually combined with the
production of other cash crops. In most of
Upper Mandailing it is the preferred occupation
of only a few and is irregularly practised. In the
Si Ladang villages, on the other hand, tapping
is regularly engaged in, and there is a greater
standardisation in the quality of the product.
In most areas the investment of labour and
capital in the cultivation of the palm is slight as
compared with that involved in coffee or rubber.
Thus the palm is utilised when the demand is
high, e.g., during the fasting month, and other-
wise neglected. There is no loss due to idle
capital.
Since the introduction of cheap white sugar
with its better storage qualities, the use of this
commodity has spread with a concomitant decline
in the utilisation of the palm for sugar.
OTHER USES OF THE PALM
The multipurpose functions of the palm are
appreciated in Mandailing, all parts other than
VOLUME 4 BOTANY
the roots being utilised, e.g., the fibre for roofing, food, the trunk for conduits, the fronds for
cord, and brooms, the pith of trunk and seeds for shelters, and the leaves for decorative purposes.
Note: Terms used throughout the paper are those current in Upper Mandailing.
DISCUSSION
T. R. MCHALH: Is there any knowledge there concerning it was probably extracted from Arenga pinnata, not cane,
the use of earthcrnware vessels for boiling sugar juices? If Did sugar techniques evolve from India or from Southeast
sugar as a crystalline product was known before 600 A.D., Asia before Persian techniques were invented ?
269
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
POSSIBLE SEPARATE ORIGIN AND EVOLUTION OF THE LADANG
AND SAWAH TYPES OF TROPICAL AGRICULTURE!
HARLEY HARRIS BARTLETT
University of Michigan, Ann Arbor , Michigan^ U.S.A.
In reviewing a vast amount of literature on
primitive tropical agriculture, the writer has been
impressed by the fact that there are a few chief
types which must have been adapted to the early
pre-agricultural environments of those tribes who
became agriculturists. One finds himself spe-
culating about where agriculture could have
originated and what environments would have
favored its development in succession to a purely
food-gathering economy. It is a certain conclu-
sion that primitive man in the humid tropics was
not pastoral. The safest habitat for him would
have been the shores of salt water, where he
lived largely on molluscs and other marine pro-
ducts, supplemented by what fruits and vegetables
he gathered in the forest, as well as by hunting
and by fishing. The first horticulture of the
seaside dweller might have been the planting of
such food-producing trees as coconut and bread-
fruit; his first vegetable gardening the actual
planting of edible plants that first presented the
idea to him (or to her!) by springing up as weeds
at the home site on rubbish heaps and other
accidentally fertilized areas. Permanent or long-
continued occupation of a site would result in
constant accession of fertilizer derived from the
debris of living, and so we might think of house-
hold gardening and seaside horticulture as
growing out of the sort of a subsistence that
prevailed among people who built up shell
mounds, inhabiting one place through many
generations. It is easy to conceive of their
migration along streams and into the forest as
soon as they invented or came into possession
of even simple tools and weapons for fishing,
hunting, and defence.
The fertility of burned-over .places would
suggest the clearing of land, and moving the
living site into the clearing would start an entirely
new cultural evolution and population mobility,
in which various forms of agriculture would
coexist with the more ancient house-site gardening
and horticulture. Early shifting dry-land (ladang)
agriculture must have developed in areas of the
most tropics where no irrigation was necessary,
in forest clearings prepared by deadening trees
without felling them and by the burning of slash.
At first the crops must have been the same species
that were gathered in the wild, such things as
Tacca (in Polynesia) and yams (throughout the
eastern tropics), and the tubers of edible aroids.
A century ago heavy, edged, blade-like clubs not
unlike canoe paddles were reported to be used in
Fiji for slashing down the forest undergrowth in
making agricultural clearings; and similar im-
plements, still surviving elsewhere, indicate some-
thing of the technique of clearing prior to the
introduction of metal tools. Trees were deadened
by ringing them with stone axes or adzes, or by
piling and burning slash about them. The ashes
of slash and the superficial organic debris of the
forest floor fertilized the soil. Cultivation was
abandoned when weeds and sprout growth
became too rampant. There was no regular
rotation of cropping with forest fallow.
Ladang agriculture extended rapidly when the
introduction of metal blades permitted the more
complete clearing which is general in shifting
agriculture today and which is characteristic of
the Malay ladang. The greater area that could be
cleared with the Malay metal-edged adze (beliung)
or its equivalent enabled the chief men to have
larger than ordinary clearings with more sub-
stantial houses, built by the joint labor of the
people. In Indonesia, the primitive clearing in
which there would have been no tillage may have
been called utna, and this term was retained in
the original sense or as a name for the entire
homestead, including clearing and house.
The more gregarious people would retain one
permanent house site, and around it the original
ladang or uma would become stocked with fruit
trees and little patches of disorderly mixed kitchen
garden. Round about the old original inhabited
site the repeated clearing of land for shifting
cultivation might result in a regular orderly
sequence of cropping and forest fallowing
permanent land use of a sort. Over-population
might result in the flocking away of a colony,
to repeat the same process. In Sumatra such a
colony would have been called a dusun.
t Presented by T. P. Bank II.
270
VOLUME 4
BOTANY
On the contrary, a less socially-minded, less
gregarious population with plenty of land to
spread over might never develop village life or
orderly land use, but remain in a semi-nomadic
state, utilizing previously unused forest as long as
it lasted, and migrating as the old forest retreated.
They would not have practiced tillage, but would
merely have sown in the soft ash-covered soil.
Chitemene and Rab agriculture grew out of
typical shifting agriculture. In remotely separate
places there has arisen the technique of increasing
productivity of a clearing by burning upon it
more slashings and other vegetable debris than it
has produced during a period of forest fallow.
Thus, in eastern tropical Africa a clearing is made,
and in it are strewn branches cut from trees in
the surrounding forest. With the debris thus
supplemented, the ashes, sometimes dug into the
soil with spades or hoes, are sufficient to produce
good crops on sadly depleted soils. The technique
is called chitemene. In India, an essentially similar
form of supplementary fertilization by the ash of
extra slashings from outside the cultivated area
characterizes rah agriculture.
The clearest transitions from ladang to per-
manent dry-land agriculture by the introduction
of tillage have been observed over the course of
years in Java. Here the population has grown so
enormously that the typical ladang has all but
disappeared. Its place has been taken by gogo (or
gaga) agriculture which is characterized by the
introduction of tillage. Its counterpart in other
parts of the world is generally designated as hoe
agriculture or, in Africa, Bantu agriculture.
It is unusual to find good descriptions of the
primitive tillage of pre-plow agricultural peoples.
Too often one runs across the expression "hoe
agriculture" as indicating a stage of agriculture
higher than the more primitive types of shifting
cultivation, in which there is no tillage, but with
no description whatever. The hoe agriculture of
tropical Africa seems to have developed typically
in forest lands marginal to savanna and more
readily transformed by fire into grassland than
likely to become regenerated forest if abandoned.
So hoe agriculture has to have tillage and be able
to combat grass as well as brush. It may have
various developments before it develops into
plow agriculture, one of which has developed
both in India and Africa, namely sod-burning for
securing ash from grass rhizomes and stubble,
where wood is no longer available. The sods,
dug by hoe or digging stick, are dried, then piled
up and burned. There are enough rhizomes and
roots so that the mass burns through, leaving a
mixture of ashes and burned-out soil which is
spread over the ground and dug in.
So far all of the types of primitive agriculture
considered may have evolved from a single point
of origin. Two main groupings are on the one
hand homestead or village horticulture with
vegetable gardening which is more or less con-
tinuous, and shifting agriculture, which may
likewise give way to a type of permanent land use
if population increases to the point where the use
of land falls into a regular progressive rotation
with alternate cropping and bush or forest fallow.
Still left for consideration is wet-land rice
growing, or sawah agriculture. What evolu-
tionary sequence could have led to this?
A chief objective of this communication is to
point out the possibility that there may not have
been any linear or connected sequence in the
evolution of dry-land and wet-land types of
agriculture which we may call the ladang and
sawah types, respectively, using the convenient
and rather familiar Malay terms, but rather,
that wet-land rice agriculture may have had a
direct origin from a food-gathering phase of
human culture. Such a possibility is suggested
by the existence in various countries of seasonally
inundated grass lands upon which the vegetation
is chiefly grass after the water subsides. It is
frequently thought that all tropical grasslands,
with the most minor exceptions, result from
repeated clearing followed by fire, and this is
undoubtedly true in general, for many have
observed degradation to grassland of originally
forested lands. There are, however, extensive
tropical areas, seasonally flooded, which show no
trace of ever having been forested. Such occur
in central Africa, where certain of the lakes are
surrounded by great areas of marsh which,
depending upon fluctuation in regional rainfall,
have the soil surface submerged to a greater or
less depth during several months of the year.
Such areas are resorted to for the harvesting of
grain from wild grasses, including various kinds
of wild rice. Extending from the Sudan south-
ward and westward through tropical Africa, there
are various instances of such harvesting of a wild
grain, and of easy transition to agriculture.
In a state of nature many grass inflorescences
have a highly developed mechanism for disarti-
culating below the spikelets or at other nodes as
fast as the grain ripens. They are therefore self-
sown so promptly that the birds (and man) are
not likely to secure most of the seed. There are
occasional mutations, however, to hereditary
sorts of which the inflorescence does not shatter.
271
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
In order to be harvested economically, a cultivated
rice or grass would have to be of this sort. It is
impossible that primitive man would not have
noticed such mutations and have taken advantage
of them. Such mutational types in nature would
have been at a definite disadvantage in the struggle
for existence, but with man's intervention might
have been artificially sown in appropriate places
and guarded during the ripening period, just as
the cultivated crops are guarded from the rice
birds in the tropics.
It must be noted that the habitat of wild rice
and other edible subaquatic grass grains would
have been open, not requiring the clearing of
forest, and that a type of agriculture might readily
have evolved on seasonally submerged lands that
would have required a minimum of tools and
implements, as it would have consisted chiefly in
the selection and sowning of especially valuable
grass mutations in habitats that already existed.
Some of these habitats in Africa were so defi-
cient in natural vegetation when the seasonal
flood water subsided that they were described as
expanses of mud that quickly became covered
with grass and other herbage, sometimes
becoming so dry as to split into hexagons during
the rainless months.
Other African habitats had a continually
emersed marsh vegetation. Why they did not
become forested is perhaps because they were
markedly saline during the dry season, but only
slightly brackish when the salt was in more dilute
solution during high water. At any rate there is
no sufficient reason to believe that marsh grass-
lands with highly varying water level were always
man-made, through the agency of fire. The
present writer has collected several instances,
from the literature on Africa, of types of primitive
agriculture on seasonally flooded land that could
well have developed directly by tribes at a food-
gathering level. Such types of cultivation, hardly
removed from the artificial increase of natural
products by selection and sowing in unprepared
habitats, must have preceded the modification of
habitats by diking, elimination of useless vege-
tation, and soil preparation.
Turning to Asia, we can see how sawah culti-
vation, the growing of subaquatic rice in diked
land, seasonally submerged, might have come
about in such areas as the Sunderbans at the
mouth of the Ganges. From here up into Assam
there are reports of wild rices, often harvested,
and similar to those generally planted in shallow
water, after the partial subsidence of flood^ water.
Needless to say, the management of artificially
modified habitats would in time have resulted in
the clearing of forest to extend the area of what
in Sumatra (East Coast) would be called bandjir
land into seasonally wet swamp forest, in which
flood water could be retained by diking.
Needless to say, after wet-land cultivation had
once come about, the techniques of irrigation
and diking would be developed to transform dry
habitats into wet. It is reasonable to suppose
that the terracing of irrigable mountain sides was
a relatively late development, carrying sawah
agriculture upward from the flood plains, where
the subaquatic rices would have naturally oc-
curred and would have been first grown, into
areas that had to be profoundly modified in order
to be suitable for sawah rice. All kinds of con-
fusing intermediates have been developed between
dry-land (ladang) and wet-land (sawah) agricul-
ture, such, even, as diking land to retain rainwater,
and planting a mixture of sawah and ladang rices
in order to secure a crop regardless of how much
or how little rain there might be.
The most extraordinary type of rice cultivation
that exists is that of the floating rice which is
found throughout flood-land areas from India
to Indochina, has attracted especial attention in
Thailand and Cambodia, and has not been un-
known in Java. (For the literature, see "floating
rice" and "wild rice" in the indexes to the writer's
Annotated Bibliography.) This cultivation takes
advantage not of subsiding or of shallow flood
water, but of rising flood water, which is event-
ually many feet deep. The growth of the floating
rice keeps pace with the rise of the flood water,
and the stems may be several meters long when
the grain ripens. It is harvested from boats.
This type of grain harvesting closely resembles
the grain gathering of the Indians of the Great
Lakes region of the United States, who gather
the excellent grain of Zizania aquatica directly
into their canoes from its shallow-lake habitat.
These Indians may have extended the range of
the useful Zizania by artificial seeding of appro-
priate places where it did not grow. Far from
the tropical Orient, we have a suggestion of how
a type of aquatic cultivation might well have
become important and have had no relation
whatever to dry-land maize cultivation. So the
ladang and sawah types of agriculture might have
had entirely different beginnings, depending upon
the habitats occupied by primitive agriculturists.
272
VOLUME 4
DISCUSSION
BOTANY
C.G.G.J. VAN STEENIS: As far as I have understood
from the reading of the paper, Prof. Bartlett assumes a
continuum of ladang (huma) and irrigated rice-field.
I doubt this generality. The irrigated rice culture is agri-
culturally a complicated thing requiring great skill and a
suitable soil. As Prof. Beyer showed in the Philippines,
the rice terraces are laid from the Gulf of Lingeyen into
the mountains, a process requiring very many centuries;
it came from an invading people from Formosa or Indo-
china, and the culture travelled with the invaders. The
same thing holds for Java which was originally called the
''Millet Island." I think the original shifting cultivators
(Proto-Malays) planted their huma with millets (Sctaria
italica, Coix, Panicnm, etc.) as cereal; later the rice was
added or replaced the millet in bulk. There was no con-
tinuum between the shifting cultivators and the wet-rice
settlers. There was also certainly not an independent
(polytopic) origin of the wet-rice culture, but the custom
travelled with human migration waves from the Asiatic
mainland.
Furthermore, I want to state that huma (ladang) is also
employed in the seasonal areas of the tropics, for example,
in the Lesser Sunda Islands, where the grassland is worked
by overthrowing the sod by means of pointed sticks by the
combined village population. This is obviously also done
in Africa where the huma is found mostly in the seasonal
or sub-seasonal marginal savanna part of the rain forest.
Besides, ladangs are more easily made under sub-seasonal
conditions which facilitate clearing the land by fire. La-
dang is not confined to the ever-wet rainforest area.
T.R. MCHAIT: Taro was cultivated long before rice in
the Philippines and elsewhere in Southeast Asia; rice is a
more recent introduction than most people assume. The
Philippine terraces may be old and made for wet-taro
cultivation.
F.R. FOSBERG: The Mariannas mark the eastern limit
of Pacific nee cultivation. There seems to have been little
contact with Asia, yet rice may tentatively be estimated
to have grown there for 3,000 years.
T.R. MCHALE: Is this wet- or dry-rice cultivation?
F.R. FOSBERG: I do not know.
T.R. MCHALE: Many of the wet-rices can be grown on
dry-land, and vice-versa.
273
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
SOME WORDS USED IN CONNECTION WITH
PRIMITIVE AGRICULTURE IN SOUTHEAST ASIAt
HARLEY HARRIS BARTLETT
University of Michigan, Ann Arbor, Michigan, U.S.A.
This paper deals with certain words that recur
in various languages and dialects and that pertain
to shifting agriculture. Sometimes when cognate
words are found far apart they show considerable
phonetic change. Likewise unrelated words of
different origin may converge into deceptive
similarity.
The writer has traced the geographical distri-
bution of Common Malayo-Polynesian uma,
which would appear to have had the basic mean-
ing "forest agricultural clearing." This meaning
has persisted in some languages without change,
but has changed somewhat in others to mean
"clearing and house" (i.e., "homestead"),
"home," "household," "persons descended from
one household," and "ceremonial center of a
local population of related persons." It is a word
widely distributed from Madagascar to Samoa
and represented in many languages, which must
indicate the extent of some ancient cultural drift,
and one suspects that it is related to rwnah,
house, another widely dispersed world.
Although uma is more widespread than other
words for "forest clearing" it is not so generally
well known as Malay lacking, which would seem
to have spread with the Malay language, and to
be, in its present wide distribution, comparatively
modern in comparison with uma.
In Assam, several languages have the word
jhum, which the writer has vainly sought to equate
with uma because of the occurrence in the Batak
languages of Sumatra of the word Juma (djuma;
djocma). The latter word, however, is considered
by students of the Sumatran dialects (such as
Joustra and Warneck) as being a condensation of
di uma, meaning "at the uma." Therefore, unless
the derived word juma spread to the continent
from the islands rather than in the other direction,
it must be concluded that the similarity to jhum
probably does not indicate a common origin.
The probable correctness of the explanation of
Batak Juma finds support from the occurrence
of a parallel derivative of uma in Toba Batak,
namely, hauma, presumably ha uma ("at the
uma"), for Toba has h instead of the k of Com-
mon Malayan ha, meaning "at," "towards," or
t Presented by T.P. Bank II.
274
"associated with." Nevertheless, the similarity of
juma -and jhum should be kept in mind by philolo-
gists, who will find a disturbing recurrence of
jum for shifting agriculture as far away as Africa.
Another word associated with shifting agricul-
ture is Kaingin of the Philippine languages and
dialects. Its origin and distribution should be
carefully looked into, especially in view of the rare
occurrence of startlingly similar words in Indo-
china, which, if the words of the two areas were
found to be cognate, would constitute a bit of
new evidence on possible early cultural drift
directly from Indochina to the Philippines,
rather than round about by way of more southern
islands.
Another word of key significance from the
standpoint of locating centers of cultural drift
is the Indochinese ray. Its history and distribu-
tion in local languages should be given attention
by persons favorably situated to do so, and a point
of especial importance would be to find out what
languages and dialects have some other and
possibly more ancient word for ray (rai). The
latter, like Malay ladang and Philippine kaingin,
may be suspected of having had a relatively
recent extension of occurrence because of selec-
tive use by administratively or politically domi-
nant groups.
The term taungya (toungyah; yah) originally
meant, in Burma, nothing more than Malay
ladang, but it has had a very curious dispersal and
change of meaning in the course of a century.
In Burma, as elsewhere, during the occupation of
a forest clearing (taungya), a good many fruit
trees might be planted, and this might lead to the
transformation of the clearing into a permanently
occupied site. The transformation of an old
clearing, occupied by a chief or important man
(who would have a more than ordinarily valuable
house and unusually extensive house-site plant-
ings) into a disorderly orchard interspersed with
garden spots, may be observed in many places
in Southeast Asia. Such a process was observed
by Kurz in Burma where (exceptionally, not
generally) the interplanting of a clearing with
fruit trees was done early, intentionally, and
VOLUME 4
BOTANY
somewhat systematically. It gave him and others
the idea that this procedure, with the planting of
forest trees instead of fruit trees, might be adapted
to the needs of foresters as a regular silvicultural
system. The trials met with considerable success,
and the word taungya, from the tongues of
foresters, quite lost its proper general meaning as
a temporary clearing for shifting agriculture
(only rarely transformed into a grove of fruit
trees) and came to designate a systematic silvi-
culture procedure. In the latter sense, the word
has spread far and has been especially attractive
to British foresters in tropical Africa. The
reader who wishes to trace the change of meaning
should consult the indexes of Vols. 1 and 11 of
the writer's Annotated Bibliography, and sub-
sequent volumes when they appear.
Although only one term of several which apply
to agricultural implements, and to which attention
might be called, tliere is one word that may prove
to be of special importance from the standpoint
of cultural drift. That is tuwai or tuai (Malay),
the name for a curious little reaping implement
consisting of a blade mounted lengthwise in the
edge of a flat, thin vertically perforated piece of
wood. The hole is for the vertical insertion of a
stick of wood or bamboo which enables the
implement to be held and operated in one hand,
for harvesting the rice straw by straw, cutting
shortly below the inflorescence. Its use instead of
the sickle often has a ceremonial significance, for
it is concealed in the hand and so "does not
alarm the soul of the rice." The cult of the
rice spirit is widely diffused in the Southeast
Asian area and probably was associated with the
exceedingly important cultural migration that
originally carried rice cultivation with it. So
vestigial beliefs and primitive techniques connected
with rice cultivation are especially important to
the investigator of prehistory, and it is merely
significant that the curious little tuwai, so familiar
in Indonesia and Malaya, extends up into Indo-
china. It should receive attention from ethnolo-
gists and linguists. It is doubtless younger by
millenia than the simple digging stick (which
should also have more attention); but neverthe-
less archeologists should be on the lookout for
symmetrical shell, stone, and metal blades with
a median perforation, which might have preceded
the blade mounted in wood. It is, of course,
possible that bamboo blades preceded the present
tuwai.
DISCUSSION
T. HARRISSON: I doubt the value of drawing conclu-
sions from word comparisons in different languages; I
believe that "i/malf is derived from the Malay "Rumah"
as also may be the words used by the Kalabits, the Kayans,
and the Kenyans, and the related words used in other
languages. The word means not a ladang, but a group
of people and the place they move to in the three languages
mentioned. These people came from the Southeast, and
the word is used by a people therefore accustomed to
moving through the forest as a community.
275
PROCFFD1NGS OF THE NINTH PACIFIC SCIENCE CONGRESS
ETHNO-ECOLOGICAL RELATIONSHIPS BETWEEN
THE MAIZE PLANT AND MAN IN WESTERN ANCIENT AMERICA
PAUL WEATHERWAX
Indiana University, Bloornington, Indiana, U.S.A.
It is generally conceded that the growing of
maize was the principal key to civilization in
ancient America. Those Indian races which
developed the three great cultures centered,
respectively, in Mexico, Central America, and
Peru, all had well developed systems of agricul-
ture. Their arts, folklore, and religions consist-
ently reflected their dependence on maize.
We ordinarily attribute this close relationship
between maize and man's cultural accomplish-
ments to the fact that the use of this superlatively
productive plant left to the Indian enough time for
the leisurely thought and exercise of the imagina-
tion necessary for cultural development. No
one would seriously question this point, but this
simple statement of cause and effect falls far short
of telling the whole story. For many centuries
before the discovery of America by Columbus,
there had been a complex interplay between man
and his principal agricultural plant. Man did
something to maize, and it, in turn, did some-
thing to him, and this exchange of stimuli had a
profound influence on both. At the dawn of
modern history, man and maize had traveled the
same road so long, and had exchanged influences
in so many ways, that, to understand either, we
must know a great deal about the other.
As this drama of cultural and biological
evolution was enacted mostly near the western
shores of the Americas, on the eastern borders
of the Pacific, the development of this theme
would seem to be an appropriate topic for con-
sideration in this Congress. It is further justified
by the fact that we hear from time to time rumors
of ancient exchange routes across the Pacific,
between the Old World and the New, and even
the occasional suggestion that the whole process
had its origin in Asia and not in America.
In many ways, maize is unique among the
great food plants of the world. It is so prolific
that, even under conditions of primitive agri-
culture, it sometimes has a ratio of increase of
almost a thousand fold. Per unit of land occupied
and per hour of labor spent on it, it gives enor-
mous yields of food.
More significantly than this, maize is one of
the few annual crops in which each plant stands
276
out with sufficient distinctness to attract indi-
vidual attention. In a field of beans, squashes,
or quinua or of the small grains, in which basal
branching makes it difficult to determine just
how much constitutes one plant, the individual
is submerged in the mass. A single maize plant,
however, has a discrete individuality. It may
produce enough grain to feed a man for a day.
It is worth individual care and attention. More-
over, it completes its life cycle in a short time,
and anything that is clone to it produces quick
results. We can imagine the slow steps by which
man learned to give it a better environment and
even the slower process of giving it a better
heredity. If it were longlived like a tree, the lag
between cause and effect would obscure their
relationships.
The collective accomplishment of the native
American races in shaping the maize plant into
what European found it to be in 1492 has never
been fully appreciated. It will ultimately come
to be regarded as one of the half-dozen or so
greatest accomplishments of applied genetics of
all time. Knowing nothing of genes or chromo-
somes or of the fertilization process, and usually
very little about even the rudiments of reproduc-
tion, native Americans employed methods of
trial and error which changed the simple wild
species into a most complex cereal completely
dependent on man for its existence.
At one end of this long process was probably
a perennial plant having something of the appear-
ance of modern tripsacum or teosintc, but a plant
so handicapped by certain maizoid characteristics
that, when it was first taken into custody by the
Indian, it was well on the way toward extinction.
At the other end we find a highly efficient cul-
tivated species displaying a morphological and
genetic complexity probably unequalled in any
other crop plant. It grew on a great variety of
soils, at altitudes ranging from sea level to 13,000
feet, and in climates from the humid tropics to
deserts, from perpetual summer to latitudes and
altitudes where the frost-free season was scarcely
more than two months in length.
The stature of the plant might be 18 inches, or
it might be more than 20 feet. Some ears were
less than an inch in length, while others are said
to have measured no less than 30 inches. The
largest grains weighed at least 60 times as
much as the smallest. The endosperm ranged
from flinty hardness to a texture so soft that the
mature, dry grain could be crushed between the
fingers, and its chemical structure showed cor-
responding variations in protein and starch con-
tent, with a few extra features, such as the high
sucrose content of the sweet varieties, the side-
chain starches of the glutinous forms, or the
ability to pop, all thrown in for good measure.
There was a relatively high correlation between
these characteristics of the plant and the ways in
which it was grown or used. The adjustment of
varieties to soil and climate was imperative. The
character of the grain dictated methods of popping,
parching, boiling, milling, or brewing. The soft,
floury grain, for example, was preferred for dry
grinding, and the flinty kinds were used mainly
in the processes involving treatment with quick-
lime or wood ashes. Almost any variety might
be parched, but there were special advantages for
this method with the sweet corns and pop corns.
The complex pigmentation of the grain merits
more consideration than it has ordinarily received.
Indeed, some modern investigators speak dis-
paragingly about any interest in this characteristic
except insofar as it presents good subject matter
for color photography. For the Indian, however,
color of the grain was not only a thing of beauty
but also often a guide to his practice of genetics,
even though he did not know what he was doing.
The Hopi maintained one of their favorite vari-
eties through careful attention to its blue color,
and the Navajo used an aleurone pattern in
maintaining a "sacred" variety. The sara misa
ears, regarded with much awe by some of the
Andean races, displayed a baffling chimera of
pericarp colors. The husking games, popular in
the South American highlands and many other
places, in which prizes went to those who found
certain color combinations, undoubtedly had the
effect of maintaining hybrid vigor in highly
heterozygous seed stocks.
In shaping the environment of the plant, the
Indian showed a skill and experienced an evolu-
tion similar to those incident to his work on
heredity. It was a long road from the first ob-
servation that a grain placed in the soil would
produce a new plant, or that the plant grew better
if the weeds were removed from around it, to the
elaborate practices of drainage, irrigation, ter-
racing and fertilizing which were in use at the end
of the fifteenth century.
_VOLUME_4 BOTANY
An especially significant progressive trend was
initiated when it was discovered that there were
good times and bad times for planting corn.
This led to observations of the movements of the
sun and stars, with the ultimate formulation of
permanent systems of counting time.
It is well that at this point we repeat the primary
thesis of this discussion: the maize plant had a
unique civilizing effect which far surpassed its
importance as a mere source of food. As the
Indian observed the plant and made decisions
about seed selection, planting, and cultivation,
he experienced many successes and many failures.
As he was dealing with a species in which the
individual was taller than a man and economically
very important, it was easy for him to test his
decisions. In other words, intelligent judgment
paid prompt and impressive dividends, and this
was conducive to further straight thinking. The
cultural value of this was immeasurable. A few
illustrations may help to clarify this point.
Maize agriculture was the original germ out
of which grew the elaborate mathematical system
of the Mayas. In order to know when to plant
corn, the Indian observed the movements of the
sun and discovered the solstices and the equinox.
Numerous sundials scattered among the ruins
from Arizona to Peru testify to the accuracy with
which these determinations were made. The
next step was the discovery that the length of the
day and the length of the year were not integrally
related. Their leapyear correction for this was
unusually accurate for its time. Out of this came
the division of the year into months, the recog-
nition of the decimal and duodecimal systems,
and the discovery of the 52-year cycle. All this
called for a system of notation, which ultimately
involved place values and the use of zero. By
this time the fascination of abstract mathematics
had left the humble agricultural origin of the
system far behind.
As the Indian worked in his fields, he came to
have a friendly regard and a feeling of awe about
this important plant. It could be nothing less
than the work of a deity or even a deity itself.
From this, came an emotional expression in
pageantry, story, and song, in ornamentation of
sculpture, architecture, and ceramics, and in an
endless array of religious rites and ceremonies.
Even the human sacrifices which shocked the
early explorers could often be traced directly to
agricultural origins.
To tie this theme more closely to the purposes of
this Congress, a word must now be said about
the time and place of the origin of the maize
277
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
plant and maize agriculture. This involves some
differences of opinion which arose as the New
World first became known to Europe. Because
the writer of this article is not primarily an an-
thropologist, he refuses to be drawn into the
controversy between the independent inventionists
and the diffusionists, but will limit himself to the
problem as presented by maize.
Soon after the discovery of America by Colum-
bus, attempts were made to identify maize with
plants mentioned by ancient writers. There were
good reasons for this error. The new-born
science of western Europe still stood in awe of
the authority of the classics and expected to find
there the answers to most of their questions. The
world was believed to be much smaller than it
later proved to be, and the newly discovered
lands were regarded as eastern outposts of Asia.
And methods of collecting and evaluating data
were not yet well standardized.
The idea that maize was known in Asia in
ancient times, or even that it originated there, still
breaks out in our literature from time to time,
but it is our opinion that there is no objective
evidence to support it. The varieties of maize
recently described from upper Burma and ad-
jacent areas could have reached there by Portu-
guese trade routes in the early part of the six-
teenth century, and the testimony of an illiterate
people that they have always had maize is not to
be taken seriously when other evidences are
against it. The earliest definite mention of maize
in western China is late enough to allow plenty
of time for it to have come in from eastern India,
and it is significantly characterized in these early
reports as something new. The report of an
early African potsherd ornamented with maize is
beset with much doubt both as to its age and as
to the identity of its ornamentation. The linguis-
tic arguments of similar import have largely
backfired against those who have advanced them.
Probably the weakest of all the evidences is the
one which points out the sweet potato and
possibly one or two other crop plants as having
been present in both hemispheres and then asks
why not also maize. As far as we know at present,
Asia does not hold a single fragment of the maize
plant, a single statement about it, or a single
artifact connected with it which is definitely dated
earlier than almost a century after 1492.
No very near relative of maize was native of
the Eastern Hemisphere. In spite of various
theories which have been held in the past, it now
seems that there is no close relationship between
maize and the sorghums or others of that section
of the Andropogoneae\ and, although Coix and
a few other oriental genera have been placed in
the tribe Maydeae, this classification is to be
regarded as only an artificial one.
In addition to all these things, we note the
rapid migration of maize after it reached Asia
in the sixteenth century. It filled a great need in
countries suffering from a shortage of food, and,
if it had been there before that time, why had not
its value been recognized ?
We are safe in assuming that maize has been
cultivated in America for at least the past 4,000
years, and fossil pollens indicate that it and Trip-
sacum and Euchlaena, the only plants closely
enough related to be able to hybridize with it,
have been associated there for many thousands
of years. The ancient human cultures based on
maize agriculture testify to its having been under
domestication for many centuries before the dawn
of history. In view of this array of evidences all
pointing in one direction, it seems futile to
speculate further about the origin of maize in
Asia or about its having been there before the
early years of the sixteenth century.
278
__ VOLUME 4
ETHNOBOTANY OF NORTHERNlPEOPLES AND
THE PROBLEM OF CULTURAL DRIFT
THEODORE P. BANK II
Museum of Anthropology, University of Michigan, Ann Arbor, Michigan, U.S.A.
BOTANY
( Abstract)
A few writers, notably de Laguna (1934),
Collins (1937), Birket Smith and de Laguna
(1938), and Heizer (1943), have suggested that
prehistoric Aleut contact with Kamchatka and
the Kuriles is indicated by archaeological and
ethnographic evidence, including the presence of
aconite poison whaling among the Ainu, Kam-
chadal, Aleut and Koniag. Japanese archaeolo-
gists declineate an Okhotsk Sea cultural sphere,
more recently a Moyoro phase (Kodama, 1948;
Oba, 1955), which shows strong Aleut-Eskimo
characteristics, including use of the labret, bone
arrow heads, toggle heads, stone lamp, compound
fish hook, and needle cases. Further, Kurile and
Kamchatkan sites yield artifacts which some
(Baba, 1934, 1936; Nakayama, 1933, 1934)
believe indicate a definite prehistoric contact with
the Aleutian area.
Historical references to Aleut-Koniag voyages
are on firmer ground. In 1621 the Japanese Duke
of Matsumae described non-Ainu Kurilcans who
apparently had skin boats and other Eskimoid
traits. Following Bering's voyage, Russian fur
hunters transported Aleuts to Kamchatka, and
the first definite record of Aleuts and Koniags in
the Kuriles appears in 1800 (Kondo). They were
probably brought by Russians in 1789 (Baba,
1943). In 1826, the Russian- American Fur
Company settled about a hundred Kodiak is-
landers at two villages in the middle Kuriles, and
in 1869 another group was taken to Sakhalien
(Stejonger). After the Russo-Japanese exhange
in 1867, all Aleuts and Koniags were presumably
resettled in Kamchatka or the Kommandorskies.
It is well known that the Aleuts undertook long
voyages on the American side of the Pacific. Aleut
sea mammal hunters, forced by the Russians,
undertook voyages in historical times to the
Pribilofs, Kodiak, Sitka, and as far south as
California. Even in pre-Russian times, they
probably traveled at least as far as Kodiak for
trading purposes.
Aleut-Koniag voyages, both in America and
in Asia, during the Eighteenth Century are of
more than mere historical interest; they may
account for a few of the cultural similarities
found among the Ainu, Kamchadal, Aleut,
Koniag, and Northwest Coast American Indian,
including the presence of aconite poison whaling
in the Aleutian-Kodiak area.
A recent study by the writer shows that there
are similarities in method of preparation, actual
use, or beliefs surrounding use of plants among
Aleuts and Ainu in approximately 55 per cent of
the instances where the same plant genera are
found in both areas. Some similarities of use are
striking, but the body of lore and beliefs surround-
ing these plant uses is much more highly devel-
oped among the Ainu. These correspondences
can largely be explained in terms of independent
invention and historical contact, and there is no
indication of prehistoric cultural drift via the
Aleutians.
DISCUSSION
C.G.G.J. VAN STtENis: Dr. Bank defines ethnobotany
as the study of the relationship between plants and primi-
tive man. In Holland there are such relationships that
have arisen only within the last few hundred years.
T.P. BANK: But this would come within the realm of
economic botany.
C.G.G.J. VAN STEENIS: I disagree. In the instance men-
tioned, the people of East Holland make a soup with
herbs introduced only 300 years ago. This is an ethnobo- ,
tanical problem, but "primitive" has no meaning here.
T.P. BANK: I agree. The economic botanist may be
an anthropologist as well. There is no real distinction
between the two terms.
F.R. FOSBFRG: In this definition of ethnobotany, I
wonder where such studies as of the plant relationships of
the Kentucky mountaineers or of the Maldive Islanders,
who do have written languages, come in. These studies
would scarcely be of interest to economic botany in its
modern sense, but are important botanically and ethno-
logically. Perhaps the way to adapt the definition would
be to omit the word "primitive."
C.G.G.J. VAN STEENIS: The definition should be the
relations between plants and man, other than where econo-
mics comes in.
279
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
F.R. FOSBERO: What about the word "primitive"? our present culture? Our distant descendants may perhaps
p.w. RICHARDS: Would it not be correct to regard regard us as "primitive'* people.
economic botany as a specialized branch of ethnobotany T>P> BANK: All would agree but the ethnobotanists.
that which deals with the relations of man and plants in
280
VOLUME 4 ___
MEDICINAL PLANT LORE OF THE ALEUT
BOTANY
THEODORE P. BANK II
Museum of Anthropology, University of Michigan, Ann Arbor, Michigan, U.S.A.
In the early Russian accounts there are frequent
references to "concoctions of bitter plants" and
"astringent roots" used by the Aleuts in the
treatment of various ailments, but plant names are
rarely given, even in later anthropological
reports. 1 Knowledge of medicinal uses of local
plants and animals was probably well developed
among the native doctors, but, as with the use
of local plants for food, these uses are now
mostly forgotten by the Aleuts. A few of the
older villagers still remember and occasionally
resort to traditional remedies, however, especially
to some of the plant infusions, which are known
to be effective tonics and astringents. They still
use the aromatic yarrow (Achillca borealis), for
example, for treatment of chest and muscular
pains. Yarrow is also said to give some aid, when
crushed and placed over a bleeding wound, in
bringing about clotting. Fox oil, derived from
the rendered fatty portions of newly killed foxes,
is a favorite cure all for skin lesions and itches.
An infusion made from Matricaria matricarioides
is frequently used by Aleut elders for gas pains
and general stomach disorders. The root of
Geum calthifolium, the leaves of Conioselinum
Gmelini and Angelica lucida, and the roots of
Plantago macrocarpa are all employed occasional-
ly in tonics. Leaves of the cow parsnip (Hera-
cleum lanatum) and of Artemisia unalaskensis
are sometimes heated and placed over sore
muscles and minor cuts. The leaves of Geranium
erianthum are boiled in water, which is then used
both as a tea and as a gargle. A number of other
plants find occasional medicinal use in the Aleut
household when the local teacher is not at hand.
For the most part these plants grow near the vil-
lages and do not necessitate any major trek
inland when the need arises for employing them.
A list of the chief ingredients in native medi-
cines formerly or still used by the Aleuts includes:
Yarrow, Achillea borealis (Atkan SAMIKA-
YAX). 2 An infusion was made from
the leaves and taken as a cure for stomach
and throat pains, also for colds. In post-
Russian times it was thought to be espe-
cially good for consumptives. Even today,
the leaves are plucked, rolled between the
palms, and placed over open cuts as a
coagulant. Leaves are also crushed and
stuffed into the nostrils, for nose-bleed.
Matricaria matricarioides [all Aleut dialects,
RAMASKAN (Russian)]. This plant
was greatly used by Aleuts and Russians
alike as a cure-all. It was thought that an
infusion of the leaves was good for stom-
ach pains, especially for gas on the stom-
ach. The infusion was also quite beneficial
1 Waldemar Jochclson, History, Ethnology and Anthropology of the Aleuts (Washington, D.C. : Carnegie Institution of
Washington, 1933).
2 The letters in the alphabet adopted for Aleut words (essentially that of R.H. Geoghegan, The Aleut Language, Washington:
U.S. Dept. Interior, 1944) are pronounced as follows:
A generally like the a in father
C like the ch in charm and chew
D like the hard d in day; sometimes like a softly voiced th
G usually like the hard g in garden
I generally like the i in machine; sometimes like the i in kid
K as in kite and kiss
L softly, like the first 1 in bull; sometimes with a suggestion of a y before the vowels a, i, u
M as in man and moon
N as in noon and naughty
NG as a single sound uttered through the nose, nga; somewhat like the ng in sing and ring, with nasal exaggeration
(very difficult for non-Aleuts)
Q like a deep clearing of the throat; often like a ch spoken far back in the throat; at the end of words, hard, like
the German ch in ach
S as in seen and soon; at times like z between vowels
T as in tune and tea
U generally like the oo in moon and moo
W like the w in we and wash
X like the German ch in noch and auch, but sometimes especially at the end of words, uttered in a more gutteral
fashion
Y like the y in you and young.
281
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
as a laxative. Today, the Nikolskians rely
upon this plant as a tonic.
Artemisia unalaskensis aleutica (Atkan SISI-
KAX). This plant was heated over hot
stones and used as a switch to beat the
part of the body where pain was felt. A
few older Aleuts, particularly rheumatic
individuals, still use the plant as a switch
during steam baths. It was also heated
and used as a poultice, and sometimes the
leaves were boiled in water for a tonic
which was thought especially beneficial for
dying persons.
Pondweed, Menvanthes trifoliata [Nikolskian
TRILISKAN (Russian)]. -Modern
Nikolskians remember that the roots of
this plant were a powerful ingredient in a
tonic. They say that it was especially
valuable for gas pains, constipation, and
rheumatism.
Pond scum (fresh-water algae and oils given
off by plants bordering ponds) (Atkan
UQUXLUX). The scum is believed
by modern Aleuts to be especially good in
the treatment of eye troubles. It is
gathered and used for bathing the sore or
inflamed parts. The oils are more often
used than the actual algae.
Avens, Geum calthifolium (Atkan AMIDUXIX
or AM I DUG IX). The roots are boiled,
and the infusion is drunk as a tonic for
colds and sore throat. The Aleuts once
used the leaves on sores that refused to
heal naturally, placing the wet or boiled
leaves over the wounds and binding them
in place. It was thought that the plant
helped to dry out the sore and aided scab
formation.
Umbelliferae. Much used in the past in
tonics for colds and in soothing drinks
for sore throats were: cow parsnip,
Heracleum lanalum (PUTSCH KE of the
Russians); wild parsnip, Angelica lucida
(SAKUDAX of all Aleut dialects); and
hemlock parsley, Conioselinum Gmelini
(Atkan CIKJGALUX). Wild parsnip
leaves were also used as a poultice, and
the older natives would slice the roots into
two parts, heat the halves, and place them
over the area of the body that hurt. If the
pain was deep within the body, merely
placing the heated roots over the skin in
the general region was supposed to bring
relief.
Ragwort, Senecio Pseudo - Arnica (Atkan
UKCUDAX). The leaves are gathered
when the plant is in flower. They are
placed directly over cuts and boils to aid
in drainage.
Clubmoss, Lycopodium clavatum (informan?
has forgotten local name). After deli-
very of a bady, the mother, if she is in
pain, is given an infusion of this plant to
drink.
Rumex acetosella (Atkan TANGAX UQUX).
The leaves are steamed and placed over
a wart, or over bruised skin.
Leptarrhena pyrolifolia (Atkan ALIXSISIX).
The leaves used to be brewed, and the
infusion was taken internally in the treat-
ment of sicknesses such as influenza.
Iris, Iris setosa (Unalaskan NASANCAGA-
DAX). The root is boiled in water, and
the infusion taken as a laxative.
Geranium erianthum (Atkan CUNUSIX).
The leaves are used in a gargle for sore
throat.
Anemone, Anemone narcissiflora var. villosis-
sima (Unalaskan CIXUDANG1X). The
root is boiled until all of the juice is
extracted, and the juice is then given to
patients suffering from hemorrhage.
Plantago macrocarpa (name not recalled).
The root is boiled in water, and the
infusion is used as a tonic.
Reindeer moss, Gladonia spp. (Atkan KINA-
DAM AIYUKAX). One informant
said that this plant is used as a tea for
internal chest pains. It is also said that
hunters who are climbing hills eat it in
order to maintain their wind.
Raw fish. One informant at Atka said that
her people occasionally eat raw salmon
or halibut in the belief that it stops heart-
burn.
Soap. Brown laundry soap is shaved,
mixed with bread and milk, and made
into a paste [Atkan PIPARKIX (Rus-
sian) ], which is placed on boils to draw
out the pus.
MAGICAL AND POISONOUS PLANTS
According to one informant, if the juice of the
buttercup Ranunculus occidentalis (Atkan KAN-
GLAGIM AMAGI) could be slipped into the
282
VOLUME 4
BOTANY
food of a well man, he would soon take sick and
would "dry up" until he had wasted away into
nothing. But the deed was dangerous to the
perpetrator, for it was thought that constant
plucking of the buttercup would bring on bad
rain storms, which would drown the wrongdoer
when he was at sea. Another informant men-
tioned feeding a soup made with mare's-tail,
Equisetum (Atkan LAGIS KALGADAX), to a
guest who was hated. It is also said that fox
trappers used to carry a "sparkling sand" and
pieces of mare's-tail in their pockets as a special
charm to help them catch more animals. One of
the red-capped mushrooms (Russula) is thought
to spring up where human blood has been spilled.
These are the only bits of magical plant lore we
gathered concerning poisonous or possibly
poisonous plants, except for what relates to monk-
shood, Aconitum maximum, the one actually
poisonous plant which we discovered. Aconitum
has been found in a number of cultures as the
primary ingredient of poison for harpoon points,
and it was apparently employed at one time by
Aleut hunters as a whale poison. 3 We made
every effort with living Aleuts to track down this
tradition, but, except for one individual, none
recalled the use of Aconitum as a poison, or
otherwise.
Not only do the Aleuts know of no use for the
plant, but most of them deny that the monk-
shood is in any way poisonous. To prove their
contention, several of them chewed and swallowed
small portions of the plant. As far as we are
aware, they suffered no ill effects. One of the
Nikolski natives declared that the old harpoon
poisons were made from decayed human fat
mixed with crushed bodies of small, poisonous
worms found in a fresh-water lake near the vil-
lage. He was, however, not able either to des-
cribe or to collect the worm for us. One of the
Aleuts at Unalaska vaguely recalled hearing that
the monkshood had once been employed as a
poison in conjunction with decayed human fat,
but he was unable to tell us anything more.
It seems obvious that the use was not widespread
among the Aleuts and that it was discontinued
soon after the arrival of the Russians. Probably,
as has been suggested by Heizer, 4 the use was
restricted to a few whalers, who kept the secret
of the poison to themselves. Interrogation of the
oldest living Aleuts near the ancient whaling
center of Akutan might yield fuller information
regarding this matter, which is of considerable
interest in comparative ethnology. Collections
of the Aleutian Aconitum from various localities
are to be made for an alkaloid assay. It is possible
that not all races of Aconitum have a sufficiently
high alkaloid content, or even the right alkaloid,
to be particularly poisonous.
The author recently had an opportunity to
study Ainu plant uses at first hand during one
year of research in northernmost Japan. An
extremely comprehensive monograph on the
subject of Ainu plant lore has already been
written 5 but it is available only in Japanese.
Of the more than one hundred fifty plants used
by the Ainu for food, medicine, poison or in
handicraft, approximately half of the genera
occur in the Aleutian Islands, and of these, a
majority found use among the Aleuts in similar
fashion to Ainu plant uses. In a few instances,
especially among plants used for medicines and
poisons, the similarities are striking.
It is impossible, however, to draw from the
ethnobotanical evidence at hand any concrete
suggestion, certainly no proof, that there were
prchistorical cultural contacts between Ainu and
Aleut via the Kuriles and the Aleutian Islands.
Aleut plant lore can be viewed as having its
origin 1) in environmentally determined needs
and independent invention among the Aleuts;
2) in cultural exchanges with American aborigi-
nals, primarily with Kodiak Islanders and
Southeast Alaskan Indians; and 3) through his-
torical contacts and cultural exchanges after 1741
with Kurile Island Ainu and Kamchadals. Simi-
larities with northern Alaskan Eskimo plant
lore are few in number, largely due to cultural
differences that are environmentally determined.
The Aleuts had access to and an abundance of
many kinds of useful plants, whereas the northern
Eskimos were much more restricted to a few plant
uses which were not highly developed among
them because of severe limitations of climate,
terrain and a much more nomadic form of life.
BIBLIOGRAPHY
Baha, Osamu, 1934, Archaeological Investigations
in the Shimushu Islands, Jour. Anthro. Soc.
Tokyo, 49: 39-62, (in Japanese).
3 R.F. Heizer, "Aconite Poison Whaling in Asia and America: An Aleutian Transfer to the New World, "Bull. Bureau
American Ethnology, (1943) No. 133, 415-68.
4 Ibid.
5 M. Chiri, Systematic Ainu Dictionary, Pt. I. Plants (Tokyo: Nippon Jomin Bunka Kcnkyu-sho, 1953), 394 pp.
283
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
, 1936, A Second Archaeological
Investigation at Shimshir, an Island of the
Kuriles, Jour. Anihro. Soc. Tokyo, 51: 91-115,
(in Japanese).
, 1943, The Aleuts in the Kuriles, I,
Jap. Jour. EthnoL, 1: 773-794, (in Japanese).
Birket-Smith, K., and de Laguna, F., 1938, The
Eyak Indians of the Copper River Delta,
Alaska, Hist.-FiloI. Mcdd. Udgivne Kgl.
Dansk Vid. Seh. Copenhagen.
Collins, H.B., 1937, Archaeology of St. Lawrence
Island, Alaska, Smithsonian Miscel. Coll.,
39: 375-384.
Heizer, R.F., 1943, Aconite Poison Whaling in
Asia and America: an Aleutian Transfer to
the New World, Bur. Amer. Ethnol. Bui.,
133:415-468.
Kodama, S., 1948, Moyoro Shell Heap, Monogr.
Res. Assoc. Pre-hist. Cul. of Hokkaido,
Sapporo, Japan, (in Japanese).
Kondo, Morishige, 1804, Henyo Bunkai Zuko
(Illustration of Japan Frontier), Tokyo, (in
Japanese; references in Baha, 1943, pp.
776-777).
de Laguna, F., 1934, The Archaeology of Cook
Inlet, Alaska. Philadelphia. ^
Nakayama, E., 1933, Excavation of the Pit Dwell-
ing Sites of Ust-Kamchatka on the Eastern
Coast of the Kamchatka Peninsula, Jour.
Anihro. Soc. Tokyo, 48: 63-72, (in Japanese).
, 1934, Neolithic Remains from the
Western Coast of Kamchatka Peninsula,
Jour. Anthro. Soc. Tokyo, 49: 375-388, (in
Japanese).
Oba, T., 1955, On the Bone Implements of the
Moyoro Shell Mound at Abashiri, Studies of
Res. Inst. Northern Cultures, 10: 173-249.
Stejneger, Leonhard, The Asiatic Fur Seal Islands
and Fur Seal Industry, (quoted in Baba,
1943; Kodama, 1948; and other Japanese
Sources; Reference to Original Publication
by Stejneger is obscure).
284
VOLUME 4 BOTANY
^HlTPROBLEM OF VERNACULAR NAMES~OF PLANTS
IN THE PACIFIC AND ITS SOLUTION
MONA LISA STEINER
Pasay City, Philippines.
Scientists are mainly concerned with scientific
nomenclature and pay little attention to verna-
cular names. For teachers, linguists, anthro-
pologists, ethnobotanists, foresters and numerous
other researchers vernacular names, on the other
hand, are significant. Foodplants, lumber trees,
weeds and forage plants, ornamentals and medi-
cinal plants, they all concern a tremendous num-
ber of people, and with growing contacts and
economic international research those names are
often essential to collectors and scientists as well.
In Europe and the United States the problem
of common names has been solved to some extent,
most European countries, for instance, have
produced floras or lists of plants with their cor-
responding common names. Dictionaries of
plant names exist, and particularly economic
plants have been well covered. In the Pacific,
however, numerous languages exist and these
plant names can only be found in regional lists or
floras. These lists of plant names are so much
in demand, as a rule, that they are frequently
out of print and therefore not available to the
general public. In many instances collectors or
people without proper botanical background
are responsible for such enumerations and the
scientific equivalents may be outdated or in-
correct.
The problem of vernacular names in the Pacific
is too complex to be treated by one or a few in-
dividuals; results can only be obtained by co-
operation, coordinated cooperation, carried on
over an extended period. Three phases of this
work are to be considered: (a) collecting of un-
recorded regional names together with their
Latin equivalents and herbarium specimens,
(b) inducing individual countries to publish lists
of plant names if no other botanical up-to-date
references with vernacular names are in print in
this region, (c) the coordination of the recorded
material, published and unpublished, in the form
of a dictionary of economic plant names in the
Pacific.
The first attempt of such a vast undertaking
was initiated by the Pacific Science Association
in the form of a standing committee on Common
Names of Economic Plants in the Pacific, which
should work towards the compilation of lists of
common names of plants in active use in the
Pacific Basin together with their botanical equi-
valents. Since such a vast task cannot be ac-
complished uno actu the project was divided into
several parts such as: foodplants, medicinal
plants, forest trees, weeds and forage plants, and
ornamentals. For the Ninth Pacific Science
Congress only foodplants were worked on.
A preliminary list of foodplants was prepared
by the chairman. This list of scientific names of
foodplants, together with their synonyms, was
sent to representatives of botanical institutions
of the various countries in the Pacific. Numerous
responses have been received from all parts of the
Pacific, and all the available literature in Manila
was scanned for vernacular names of foodplants.
Through the facilities of the National Research
Council in the Philippines, courtesy of Dr. Pa-
troncinio Valenzuela, 250 copies were mimeo-
graphed. One page was reserved for every plant.
Next to the known Latin synonyms the vernacular
name in the major languages were given: Chinese,
Dutch, English, French, German, Japanese, and
Spanish. Then the Pacific Basin was divided into
various regions such as : Malay Peninsula, Thai-
land, Indonesia, North Borneo, Philippines, For-
mosa, East coast of China, Okinawa, Japan, New
Guinea, Micronesia, Melanesia, Polynesia, Central
America, and North America. The respective
vernacular names with their location in parenthesis
were then given.
Since this mimeographed compilation was
done mainly in 1956, empty spaces indicated that
many regions did not send in sufficient material
before 1956. After this preliminary compilation
was sent to various institutions in the Pacific
much new material was collected by the chairman.
During her leave in Europe the chairman was
also able to gather numerous vernacular names
particularly in the major languages. All these
materials are going to be incorporated into the
Dictionary of Vernacular Names of Foodplants
in the Pacific. Supplied with a cross index it will
become a valuable reference work.
Before the final publication all plant names
have to be checked and rechecked by residents
of that certain region. Only a person familiar
with the Thai language, for example, can correct
285
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
the common names of Thailand, leave out extinct
names or those of hybrids. An overall dictionary
can naturally not include varieties and hybrids,
plants as the banana, rice, etc., would alone fill a
book. Regional collaborators are also to select
the preferred names of a region, and mark them.
Although standardization is not planned, the
selection of a preferred name will help the readers
to pick out the most commonly used name. In
the Philippines, for instance, about 80 languages
are reported, yet one of them, namely Tagalog is
the National Language, and this one name is
generally understood in most provinces. The
problem of the multiplicity of vernacular names
in many countries is thus solved to some extent,
the various common names of a plant will be
recorded in the dictionary, but the preferred name
will be set ofT by a different type of print.
Although the project was started with phase
(c), as previously mentioned, it initiated indirectly
also phase (a), namely the listing of hitherto
unrecorded common names of economic plants
in the Pacific. Various members of the committee,
such as Mr. Jacques Barreau, Mr. Jean Huon de
Navrancourt, Kamulai, Papua, have sent in
valuable lists of sofar unpublished material which
have been included. From several communi-
cations it became apparent, that the work of the
committee on vernacular names has been a
stimulus for further recording of lists of common
names.
Phase (b) was not yet attempted, namely in-
ducing individual countries to publish lists of
plant names, but this work is part of the future
program.
Future program: In case this committee should
be continued by the Pacific Science Association
the following procedure is suggested:
The publication of the Dictionary of Common
Names of Foodplants of the Pacific. Besides the
cross-index of the vernacular names, a bibliog-
raphy will be included to furnish detailed in-
formation too voluminous to be incorporated.
The publication must be well printed and pro-
perly distributed to be of real value. Much of its
usefulness depends on its wide distribution and
availability.
Foodplants are, however, only a part of eco-
nomic plants, and forest trees, medicinal plants,
and ornamentals have also to be worked on.
The chairman intends to cover foodplants and
ornamentals, and has appointed specialists to
handle forest trees and medicinal plants. After
the individual publications on forest trees, medi-
cinal plants, etc., have been issued, all these
286
information should be fused into one compre-
hensive dictionary.
This is a very far reaching project to be ac-
complished only by several persons, one scientisi
alone can not reach its aim, it requires adequate
funds and most of all international support. This
is not a work that can be "accomplished" within
a few years, but working, recording, listing and
improving has to be continued over a long period.
Working members have to be supplying the
committee in every country, and the expenses of
mail alone are sizeable.
One of the main purposes of this committee
should be to act as a clearing center for common
names. They should be recorded and evaluated
to some degree. Overall standardization is neither
possible nor intended. One common name can
not be used by different language groups, and the
general usage can naturally not be altered by any
group of people-language lives, grows, and
changes.
It is quite obvious that vernacular names can
never be perfectly under control such as Latin
names, but a serious effort nevertheless should
be made to bring some order in this most con-
fusing matter of vernacular names, first by
recording, and secondly by selecting preferred
names for a certain region, if feasible. Standard-
ized names of lumber trees appear on many lists
in the Pacific, but they are not circularized suffi-
ciently, maybe the committee could eventually
combine these various lists, and lumbermen will
be able to survey the various common names and
know their scientific equivalents, and can check
with the help of the dictionary its validity.
Landscape architects dealing with tropical
plant material are frequently faced with lack of
common names, eventually each one coins a new
one. Standardized Plants Names, prepared by
the American Joint Committee on Horticultural
Nomenclature has proven very effective, but
although numerous tropical plants are listed with
their common names, many new ones are not
included, and it is felt that tropical material should
be treated only after all common names have
been recorded, and one can select the common
usage for Hawaii, India, etc. One name can
eventually be preferred in the English language,
and once a dictionary has become available the
preferred names will be used in references, in
literature and employed by teachers and laymen
as well.
From the enthusiastic response of the project
on vernacular names from various institutions,
and also from the well attended symposium during
VOLUME 4
the 9th Pacific Science Congress and its dis-
cussion, it became apparent that this undertaking
is of substantial interest to numerous branches of
applied and pure science. Continuation and
further advances are highly recommended.
SUMMARY
The complex problem of vernacular names is
particularly complicated in the Pacific, where
numerous language groups meet and tropical
plantlife is extremely rich. A Standing Committee
on Vernacular Names of Pacific Plants has been
established after the 8th Pacific Science Congress
and produced a "Preliminary Compilation of
Vernacular Names of Foodplants in the Pacific"
which was sent to various institutions and eco-
nomic botanists all over the Pacific to be checked
and filled in. Published and unpublished refer-
ences and lists respectively have been used and
incorporated. It is to be hoped that a Dictionary
of Vernacular Names of Foodplants in the Pacific
will be published soon, including a cross-index
and substantial bibliography.
It is hoped that through the initiative of the
committee members and through international
cooperation lists of hitherto unpublished common
names will be recorded and published. Individual
countries should also be induced to publish their
own lists of common names.
Besides food plants, forest trees, medicinal
plants, weeds and forage plants, and ornamentals
should be worked on. Specialists in each field
should take over each field and publish an in-
dividual dictionary. Eventually all special dic-
tionaries can be combined into one complete
dictionary of vernacular names of Pacific Plants.
List of members of the Committee on Verna-
cular Names of Pacific Plants and other who
have sent in lists of vernacular names or
references :
Mr. H. ANDO
Hiroshima Universi-
ty, Sendamachi, Hi-
roshima, Japan
(member, Japan)
Mr. JACQUES BARRAU
Laboratoire d'Agrono-
mie Tropicale, Facult6
de Sciences, Marseille,
France
(member, Melanesia)
checked and cor-
rected Preliminary
Compilation
sent several published
and unpublished lists
New Caledonia, Trust
Territories
Miss N. BURBIDGE
Canberra, Division of
Plant Industry
(member, North Aus-
tralia)
Mr. R.E. DWYER
Director of Agriculture
Papua, Port Moresby
(member, New Guinea)
Dr. SHIU-YING Hu
Arnold Arboretum, Ja-
maica Plain, Mass.
U.S.A.
Mr. M. C. LAKSNAKARA
KASIIEMSANTA
Department of Agri-
culture, Bangkok, Thai-
land
(member, Thailand)
Mr. FAUSTINO MIRANDA
Institute de Bilogia
Casa del Lago, Chapul-
tepec, Mexico
(member, Mexico)
Mr. H.W. Li
Director Academia Si-
nica, Taiwan, China
(member, Formosa)
Miss M. NEAL
Bishop Museum, Ho-
nolulu
(member, Polynesia)
Mr. JEAN HUON DE NAV-
RANCOURT
Kamulai, Papua
Mr. B.E.V. PARHAM
Department of Agri-
culture, Suva, Fiji
(member, Fiji)
Mr. HAROLD ST. JOHN
University of Hawaii,
Honolulu
Dr. MONA LISA STEINER
2833 Park Avenue, Pas-
ay, Philippines
(chairman)
Mr. TEM SMITINAND
Royal Forest Depart-
ment, Thailand
BOTANY
sent lists of New Gui-
nea plants, and refer-
ence on Australia
list of Solomon Is-
land, Comparative
list of Micronesia,
Polynesia, Melanesia
sent mi meographed
copies of a specially
prepared extensive
enumeration of the
food plants of China
sent reference on
Thai names
sent list of Mexican
plant names
sent list of Formosan
names
sent list of Hawaiian
names
sent unpublished list
of names of plants of
Papua
sent Fijian plant
names, published
sent several publica-
tions with vernacu-
lar names on Micro-
nesia
checked Philippine
names
checked and filled in
Preliminary Compi-
lation.
287
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
DISCUSSION
F.R. FOSBERG: The fact that lists of vernacular names
are generally out of print seems a significant fact in indi-
cating a need for such lists. This situation may be very
useful when the dictionary is ready for publication in per-
suading a publisher to accept it.
Attention should be called to the existence of a card file
said to contain one and one half million vernacular names.
This was compiled by a man named Williams and is de-
posited at the Los Angeles Museum Herbarium. It should
prove to be a valuable source of information.
E.H. WALKHR: In connection with compilation of
vernacular names, there should be compiled a bibliography
of sources of these names. These entries should be well
annotated with special stress on comments and interpre-
tations of the works.
Preliminary treatments such as Dr. Steiner's list are
valuable by their unavoidable omissions and errors in
stimulating others to cooperate in correcting them.
In compiling lists of vernacular names, careful consider-
ation must be made to document their origins. The source
of each name must be given. It is important to start this
now on an understanding basis because the project will
undoubtedly snowball to unexpected proportions.
T. TUYAMA: Concerning Dr. Steiner's "A Preliminary
Compilation of Vernacular Names of Food Plants in the
Pacific" I wish to indicate that Japan has a large number
of vernacular names that are still unrecorded. J also
suggest that Okinawan names be included in Japanese when
they are so used.
p.s. ASHTON: In my opinion, to get this work
(1) representative of the whole area under study,
(2) to get it in some form of completion in the near
future, all those at present concerned with the project
should use all the opportunities at their disposal to:
a. Contact and appoint willing people in all areas under
study, who are more or less permanently living in these
areas, to take on the duties of regional investigation.
Short-term investigators rarely get valuable material for
this type of study as, through ignorance of the language
and the people, they do not get their full cooperation.
b. To find people in Europe and America who can find
time to spend in the herbaria and libraries in order to:
(i) gather information concerning early European
names of Pacific plants and
(ii) gather information of Pacific vernacular names
already recorded both to help this present work and also
to compare with the field work being done by the regional
representative that recorded mistakes may be eliminated.
c. We need one person or one more or less cenj
tralized committee (as we have at present) to goad on the
regional and herbarium library representatives.
F.R. FOSBERG: Obsolete or archaic names would be
of much more use to me than modern names. It is probable
in special instances that we will not get any one botanist to
handle this, perhaps not a botanist at all. It may be desir-
able to bring in people with historical interests, linguists,
etc. and to help them to correlate the vernacular names
with Latin binomials.
J.H. HURLIMANN: Recording of Portuguese vernacular
names seems important, especially for etymologic under-
standing of alterations in indigenous languages, as Portu-
guese were amongst the first Europeans to come to the
Western Pacific area.
T.P. BANK: Dr. Fosbcrg's remarks are pertinent to
what I'd like to say. Why don't we use this excellent
work of Mrs. Stciner and her group to enlarge the scope
to include not only vernacular names of current usage,
and names of obsolete usage (very important to linguists and
ethnobotanists), but also an ethnobotanical dictionary!
This is something which we have been planning for some
time, and now we have in this compilation of vernacular
names a focus to give us a start.
We might do this by forming a sub-subcommittee a
group of interested botanists, linguists and anthropolog-
ists to obtain local traditional uses for these plants.
This would make the compilation much more useful to
those working at a cultural level. Much of the necessary
data is already available but hidden away in numerous
field reports and publications by anthropologists and
others. The result of their combined effort would be an
ethnobotanical dictionary for the Pacific area.
j.v. SANTOS: The introductory remarks of Dr. Steiner
presents a problem on the vernacular names of many
plants introduced into the Philippines. There are instances
when the vernacular names of plants in the country from
which they were believed to have been introduced, were
also "imported" and adopted in the Philippines. We have,
for example, the following plants which are known in the
Philippines in their Spanish-Mexican names: Antrogonon
legtopus (Cadena de amor), Delonix regia (Caballero).
At the same time we have also a large number of intro-
duced plants without vernacular names in the Philippines.
If these plants should be given vernacular names, then
the problem will be how to go about it.
288
VOLUME 4
BOTANY
AN ENUMERATION OF THE FOOD PLANTS OF CHINA WITH
VERNACULAR NAMES
SHIU-YING HU
Arnold Arboretum, Harvard University, Cambridge, Massachusetts, U.S.A.
This enumeration is prepared in response to a
request of Dr. Mona Lisa Steiner, Chairman of
the Committee for the Vernacular Names of
Pacific Plants, who has asked me to join the
committee and to participate in its activities by
supplying Chinese plant names for the forth-
coming dictionary of Pacific plants of economic
importance. It contains over five hundred species
of edible plants. Each entry covers the scientific
name and the vernacular name or names in
Chinese and their transliterations both in sound
and in meaning.
The distribution of the enumerated species in
the plant kingdom is as follows: 15 algae, 11
fungi, 7 ferns and fern-allies, 8 gymnosperms,
86 monocota and the rest are dicots. The arrange-
ment of the species follows the Engler and Prantl
System of classification. It seems that the rela-
tively small number of the n on- vascular plants
does not call for subdivisions down to the families
to insure clarity. Only higher hierarchical cate-
gories are adopted for the algae and the fungi.
For example, all the red algae are arranged
alphabetically under the Rhodophyceae, and all
the mushrooms and their allies are alphabetically
arranged under the Basidiomycetes.
The Chinese plant names are generally com-
posed of two or three, and rarely more monosyl-
labic words with one or two representing a noun
and the others being modifiers. As China has
many dialects, these names can be pronounced
differently by different people. In this enumera-
tion the sound of each Chinese character is tran-
sliterated in accordance to the pronunciation
given in the Revised American Edition of Mat-
hew's Chinese-English Dictionary. The sounds of
a name with several characters are connected by
hyphens.
China is vast both in space and time. A plant
of some economic importance is often widely
distributed and generally has several vernacular
names which are used simultaneously in various
parts of the country. For example, "maize or
com" is called yu-shu-shu in East China, pao-ku
in West China, and pang-tzu in North China.
Moreover, a name used once in historical time
may have been changed and the current name for
the same species are different from the book
names. For example, la-la-yang and chu-chu-
fung are both current vernacular names for
Humulus japonica. In books lu-ts'ao is always
given for this species. In this enumeration the
names which to my knowledge are the most widely
used in this generation have priority while the
other follow. The names which appear in print
only are marked "book-name."
In regard to the food plants of China, there are
several words which appear repeatedly in this
enumeration. They are often seen in other
references, and are worthy of remembering. These
are kua for melon or squash, kuo for fruit, // for
pear, mei for wheat, mi for grains especially for
rice, Cao for peach, ts'ai for vegetable, ts'ao for
herb, and tzit for seed. Wild form of edible
plants are often designated as yeh by the northern
people and as shan (which means mountain) by
the southern people. For example, yeh-fen-t*
wan-erh is the northern name of Aster brinervius
while shan-po-chu is the southern term for the
same. Both refer to the white flower. Introduced
plants are designated as hu if they entered China
overland by the ancient silk route, as yang if
they have come to China by the sea route and
reached the Yangtze area, and as fan, if they
have come to China through Hong Kong,
Macao, and Canton. For example, hu-fao for
walnut, hu-tou for broad bean, and hu-kua for
cucumber, are all ancient introductions. Yang"
szu-tzu undfan-ch'ieh are synonymous for tomato
and Yang-yu and fan-shu are synonymous for
Irish potato. These are more recent introductions
which entered China through the seaports like
Shanghai, Foochow, or Canton.
The Chinese language is an old and compli-
cated but living language. Various combinations
can be made to meet the need of the people. It
has been proved to be quite sufficient to convey
the idea of the Chinese people from the ancient
historical time to the present atomic age. Never-
theless, there are certain limitations. The words
are monosyllabic. As the number of sounds
are limited and the number of combination of
parts to make new words unlimited, every sound
covers many characters. Take Mathew's Chinese-
English Dictionary as an example, this book
gives 366 variations of sounds and 7,773 words.
289
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Thus each would average more or less 20 charac-
ters. When the first persons recorded the plant
names in writing, they were usually people without
much education. They may have given the correct
character or they may have put in any character
that sounded like what the illiterate people
pronounced. This often happened and conse-
quently the written names of certain plants are
far from expressing the meaning of the people
who used them. For a long time I could not
understand why the pokeweed was called "shang-
lu. jo] . " Finally on a collecting trip with a
medicine digger, he casually called the plant the
mountain radish in Chinese, and I began to
realize that shang-lu was derived as a corruption
ofshan-lo-pu. In this enumeration there are several
such names. In these cases no translation for the
meaning of the name is attempted.
Many of the species in this enumeration appear
for the first time in the English language as edible
plants. To these species a short note concerning
the parts of the plant used and also as to whether
they are eaten raw or cooked is given. Wild
edible plants are specially marked with E,
meaning emergency food. The more expensive
food plants are marked "delicacy" for this may
avoid embarrassment or shock when the bills
are received. It is worthy of note that the most
expensive item in this enumeration is Tremella
It is sold by dollars
fuciformis, a Basidiomyctes.
per ounce.
There are quite a number of plants which have
recently been introduced in the tropical regions
of China such as Taiwan and Canton. The*
distribution is limited to experimental stations
and their names are usually straight translations
of the scientific or the common English names.
These species are marked by NI, meaning newly
introduced, and now translation of the meaning
of the names are necessary.
yu-shu-shu
la-la-yang
chu-chu-t'ung
lu-ts'ao $ Q
kua i\\
kuo ^
met ^
mi fa
t'ao fa
ts'ai
ts'ao i^.
tzu ^~
yeh ft
shan J4
yeh-fen-f wan-erh
shan-po-chu
hu Jfl
yang >^
fan ife
hu-t'ao
hu-tou
hu-kua ifl ;
yang-szu-tzu
fan-cKieh -
yang-yu
fan-shu
shang-lu
shan-lo-pu
tremella fuciformis
^f*>(i
290
VOLUME 4
BOTANY
THE PROBLEM ON THE VERNACULAR PLANT NAMES IN
JAPANESE
TAKASI TUYAMA
Ochanomizu University, Otsuka, Tokyo, Japan.
The first appearance of plant names in the
Japanese literature can be traced back to about
100 A.D. These names are essentially on plants
of economic importance. At that time, the
Japanese alphabets had not been invented. These
names were written in Chinese characters. Among
them there were many names pertaining to plants
introduced from abroad, principally from or
through China. For example, Prunus mune from
China, Cannabis saliva from Central Asia through
China were all mentioned.
About 700 A.D., the first Japanese poetical
collection as well as an encyclopedia was pub-
lished in Japanese alphabets. One section of it
was devoted to plants. For the first time the
Japanese pronunciation of plants were recorded.
Of these plant names, some remain unaltered
while others have been more or less changed in
pronunciation and spelling. So that the plant
names in Japan, as in the other countries, reflex
the culture and history of the country. For
example, "Cotton" or "Wata" in older Japanese
literature means the kapok-like product, while
in later usage (about 1600 A.D.) "cotton" refers
to the newly imported Gossypium. Similarly,
"Potato" or "Imo" in the older records refers to
Dioscorea nipponica which is wild in Japan. But
in later years, after the introduction of Colocasia
antiquorum from the south, the name of the
wild Dioscorea nipponica was distinguished by
the addition of "Yamatsu," meaning mountain.
Thus we have "mountain Potato" or "Yamatsu
Imo" for D. nipponica and "House Potato" or
"letsu Imo" for C. antiquorum. Still later, after
the introduction of Ipomoea batatas and Solanum
tuberosum into Japan in the fifteenth century,
there are four kinds of plants bearing the name
"Imo." They are as the follows:
Dioscorea nipponica with the addition of new
yams introduced thereafter. . . "Yama-nolmo
Colocasia antiquorum "Sato Imo"
Ipomoea batatas "Satsuma Imo"
Solanum tuberosum "Jaga Imo".
At the present Ipomoea batatas is the most
commonly cultivated species. It is often simply
called as "Imo".
This is a brief history of the changes in "Imo."
However the actual courses of development are
more complicated.
Of course, there are local differences in the
vernacular names at the present as well as in the
past. Some old names are remaining occasion-
ally in certain places and are used with the recently
and sporadically developed names. Some com-
pilation of such vernacular names has been under-
taken by herbalists. Today we have many out-
standing works compiled by botanists and eth-
nologists. The present day Japanese vernacular
names of plants can roughly be classified as the
follows:
1 . Standard Japanese names : These names are
used for educational and scientific purposes.
There are occasionally several standard
names used in accordance to the opinions
of different botanists.
2. Commercial names: These names are
used principally by dealers of horticural
plants such as floriculturists, nursery men,
etc.
3. Local names and names used only by
children: These are names used by people
in different localities and those by children
in their plays.
In fact, there are collective names of different
species or genera used in popular periodicals or
in general conversation. For example, "Sakura"
means the flowering cherry of any kind when it is
not accompanied by any modifiers.
There are also special problems concerning
the name represented by Chinese characters.
Even after the establishment of Japanese alpha-
bets, the Chinese characters have been used in
mixture with the Japanese. Chinese characters
were used exclusively and extensively in Pharma-
cognostics because this subject has been developed
in China very greatly since the ancient days. It
has been a difficult task for the Japanese herba-
lists to find the identities of the Chinese plant
names and the actual plants wild or cultivated in
Japan or imported from China.
Owing to the extensive development of general
education in Japan, standard names of plants
291
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
are gradually taking the place of the local ones.
In some cases the vernacular names are accepted
as standard ones. For example, in the islands
of Ryukyu and Izu archipelago, there are local
names used generally by the natives. Many of
these have been authorized by the modern
botanists as standard names.
In the Bonin islands the vernacular names of
plants are strongly influenced by the introduction
of Polynesian language, especially that of the
Hawaiian.
I must add here that there are some researche^
on the Micronesian vernacular plant names oy
Japanese botanists, some of which are not yet
published.
D/SCUSSION
F.R. FOSBERG: I would like to call attention to the fact
that vernacular names are a part of everyday language.
Creation of pseudo-vernacular names has no significance
and is worse than useless. True vernacular names belong
to folk language and are useful in studying languages and
in getting clues to other things, such as presence of species
in a flora, tracing relationships of peoples, languages,
etc. Botanical precision is served by Latin binomials,
not by vernacular names. The study of vernacular names
is a part of linguistics more than of botany.
T.P. BANK: T would like to suggest that workers with
vernacular names in Japan attempt to determine whether
or not the common names for plants arc derived in any
way from Ainu words. Place names, for example, Fuji
yama mountain of fire god throughout most of Hon-
shu are often Ainu words. I believe we would find this to
be true, also, for many plant names.
292
VOLUME 4
BOTANY
COMMON VERNACULAR NAMES OF PLANTS OF
THE PACIFIC BASIN
J.W. PURSEGLOVE
Professor of Botany, Imperial College of Tropical Agriculture, Trinidad.^
The compilation of vernacular names with
their scientific equivalents for the whole of the
Pacific Basin would be a formidable undertaking,
and if done with any completeness would run
into many hundreds of thousands of names.
With an unlimited staff available, this would be
an interesting and instructive exercise and would
undoubtedly be of use to those interested in the
plants of the area, more especially if it could be
printed under the various vernacular languages,
rather than all the languages printed together in
an alphabetical list.
To be authentic, however, it would be essential
that the work of collecting and compiling the
names should be done by competent botanists
with access to herbarium material and who are
also competent linguists in the language in which
the names are recorded. Quite frankly, unless
this is done critically, the mere compiling and
printing of vernacular names, as recorded in the
literature and on herbarium specimens would
be very time consuming and of comparatively
little use. It might well lead to increased con-
fusion. With so few capable taxonomists working
in the area, I doubt very much whether they can
spare the time to undertake the task; they would
in fact be more profitably employed on other
work, unless the lists were limited to a few eco-
nomic plants (names of which are likely to be
given in the dictionaries any way), a few orna-
mental or other restricted groups of plants.
This does not mean that I believe vernacular
names to be of no use. On the contrary I believe
that they can be of great value, provided they
are critically recorded. They may even help the
taxonomist in his work and often provide in-
formation as to the local uses of the plants. They
may sometimes provide the key, if correctly
interpreted, to the movement and spread of
cultivated and other introduced plants in an area.
Undoubtedly, they can be of great value to the
forester, as he can inflict upon the small number
of his local indigenous staff vernacular names,
which his department has standardised for the
commoner timber trees and timbers of an area,
more especially when such staff would have much
, Director oTfiotanic Gardens, Singapore. " ~
greater difficulty with the scientific names of the
trees concerned. It does not follow, however,
that the standardised names provided by a Forest
Department will be in general use by the mass of
the population of an area.
What I think is essential is that every effort
should be made to collect reliable vernacular
names for as many as possible of the plants of an
area, as it is likely that with the spread of educa-
tion and sophistication that many of these names
will be lost in the future. I would emphasize,
however, that these vernacular names should be
very carefully checked by those who are recording
them, and it is desirable that they should have
some knowledge of the language in which the
names are given.
Nevertheless, there are innumerable snags in
this work, as I have found from past experience,
when I recorded several thousands of vernacular
plant names in various Bantu and Nilotic
languages in Uganda during my 16 years there.
I can only presume that some of the difficulties
encountered there must also arise in the Pacific
Basin and from the perusal of the literature it
would appear that they do. It is worthwhile,
I think, to list some of them.
/. Unreliable Information
It frequently happens in collecting vernacular
names, and in approaching the people who live
in the area about them, that they will not admit
that they do not know and will either make up
a name for the satisfaction of the person recording
them or will give a wrong name. Unless the names
are carefully checked and rechecked, a large
percentage of them may be completely useless and
in fact misleading. Usually the older members
of the community have a better and more reliable
knowledge of such names and will usually give
reliable information when they know well the
person who is recording them, more especially
when this person can point out information which
has been wrongly given. An unsuspecting person
with no knowledge of the local language will
often record such names as "I don't know,"
293
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
"It is a plant," etc., and these appear on her-
barium sheets.
2. Spelling
The person recording the names usually writes
them down phonetically and it is not surprising
to find that quite reliable recorders will spell the
same name in different ways. It seems to me that
the person recording them should have a reason-
able knowledge of the vernacular language before
he attempts to record this type of information.
Often the names have been misheard by the
European and great care must be taken not to
confuse names that sound alike to Western ears,
but which are quite distinct to the local person
who is giving the names. It is also likely that
there would be divergent spelling according to
the nationality of the person recording the names.
Thus it often happens that different spelling is
given by the British, Dutch and French authori-
ties, etc., even when the name recorded is in the
same vernacular language. In some cases,
certainly in Africa, the orthography of the
language has not been fixed. Committees are
formed from time to time to try to reach a finalised
orthography, which invariably results in changes
of spelling of the language and, of course, of
plant names, so that one has constantly to be
revising the method of writing these names, thus
leading in some cases to confusion.
3. Prefixes, etc.
In Bantu languages, and I understand in Malay,
names are frequently changed by changing the
prefixes, which if recorded incorrectly is often
a cause of confusion. Care must be taken to
record the singular of the vernacular names.
4. Names of Various Parts of the Plant
It sometimes happens that parts of a plant are
given quite distinctive names, and one man will
record the name of the fruit, another of the leaves
and another of the complete plant, etc. It is
obvious for any list of vernacular names that the
name of the complete plant should be recorded.
5. Multiplicity of Names for the Same Plant
Very often the same plant has several quite
distinct names even within one area and certainly
the names change as one moves to other parts of
the country, although the inhabitants may be of
the same race. This can add greatly to the con-
fusion and mitigates against the value of verna-
cular names. If it is proposed to standardise them
and have only one vernacular language all over
294
the country for the one plant, who is to do the
standardisation? Even if this is done by a com-
mittee such as yours, it is unlikely to be accepted
by the inhabitants themselves, who will continue
to use the names to which they are already 3
customed. I have come across cases in Africa
in which the witch-doctor will use his own peculiar
names for the commonest plants, in order that
the mass of the population should not know to
which plant he is referring.
6. Generic of Family Names
Indigenous peoples often recognise closely
allied species and will use a general generic name
for a number of similar species. Sometimes the
grouping is even bigger and the name will be
given to a number of genera or even to a family.
This becomes very confusing when very different
species belonging to different families are given
the same vernacular name. Corner (1951) quotes
the name of Tangisong Burong "It is given to
any tree, shrub, climber or epiphyte the fruit of
which is yellow or red and has large, often pulpy
seeds so that it is attracted to birds, yet because it
is poisonous or unpalatable, the birds weep with
disappointment."
7. Degeneration of Foreign Words
When a new plant or crop is introduced into
an area, the English or scientific name of the
plants may be changed considerably to form the
basis of the new vernacular names for these plants.
Thus, in one area in Uganda, Black Wattle
became in the vernacular Bull Kofi, while Euca-
lyptus became Kalitunsi. The variety of sweet
potato was known in one area as Disi, which is
how the people of those parts describe the D.C.
or to give him his full title District Commis-
sioner. This variety gets its name because the
D.C.'s on tour tried to persuade the people to
plant this particular variety, because of its superi-
ority to those which they were already growing.
I well remember introducing a certain grass for
soil conservation into a district and which spread
widely over the area; this grass became known
as Pulesigulufu, which was how the people of that
area attempted to pronounce my name. Thus
unless the historic aspect is considered it is often
difficult to see how some vernacular names have
been derived and the final result bears little
resemblance to the original name.
8. Vulgar Names
Quite a number of plants in any area are given
vulgar names, which are often a source of amuse-
ment to a certain section of the community but
are a source of embarassment to others. As one
dictionary put it when asterisking such words
"they are not used in polite conversation."
9. English Plant Names
Attempts have been made from time to time in
popular works on the plants of a tropical area
to give each plant a common English name, in
the hope that this may prove to be of use to the
non-botanist. This was done by Henderson in
"Malayan Wild Flowers" and by Corner in
"Wayside Trees of Malaya." Sometimes the
name given is a direct translation of the scientific
name of the plant. Thus Lettsomia rnaingayi
is called Maingay's Lettsomia. Surely this is a
little unnecessary. Sometimes common English
plant names are taken and given to a tropical
plant which bear no relationship whatsoever to
the plant so named in Britain. Thus Passiflom
foetida is called "Love-in-a-mist." Both these
examples were taken at random from Henderson.
To me this merely adds to confusion. Similarly,
common cultivated trees and shrubs are given
completely different English names in various
parts of the world, which was a great cause to
confusion to me, coming as 1 do from Africa to
Malaya. Thus Delonix regia is known as Flame
of the Forest in Malaya. In Africa it was known
as Flamboyante, while in Africa Spathodea cam-
panulata (syn. S. niloticd) is known as Flame of
the Forest, but in Singapore as Tulip tree. Even
if your Committee decided which plant should be
called the Tulip tree, I cannot see that this will be
accepted by other countries, where a completely
different plant is currently known by that name,
e.g. Liriodendron Thespesia, etc.
JO. Different Language Groups
A further difficulty is that in a cosmopolitan
region such as Singapore with a total area of just
over 200 sq. miles, it would be necessary to list
vernacular names in several languages Chinese
(several dialects), Malay (often several names for
the same plant), Tamil, Javanese, English and
VOLUME 4 BOTANY
other European languages, etc. Furthermore,
the flora of Singapore includes a great number
of introduced species, which have now become
naturalised, in addition to many introduced crops
and variety of crops. A list for this small area
would be formidable and difficult to finalise
and to use.
//. Varietal Names
The number of local vernacular names for the
many varieties of cultivated groups is legion and
the work of collecting them would be prodigious.
It is very easy to record the name of an obscure
variety instead of the general name for the species
as a whole.
12. Sources of Information
A very great number of vernacular names have
been recorded and published for Malayan Plants.
The mere physical effort of extracting these names
into an alphabetical list with the scientific equi-
valents would take many months.
The principal sources are as follows :
Burkill, I.H., 1935, Dictionary of the Economic
Products of the Malay Peninsula, 2 vols.,
London.
Corner, E.J.H., 1952, Wayside Trees of Malaya,
2 vols., Singapore.
Henderson, M.R., 1949-54, Malayan Wild
Flowers, 4 parts, Kuala Lumpur.
Holttum, R.E., 1953, Gardening in the Low-
lands of Malaya.
Ridley, H.N., 1922-24, Flora of the Malay
Peninsula, 4 vols., London.
Watson, J.G., 1928, Malayan Plant Names,
Mai. For. Rec. 5.
A list of Botanical and Vernacular Equivalents.
Govt. Gazette, F.M.S. XXIX, 5884, 1937.
Numerous articles in Gardens' Bulletins, Agri-
cultural Bulletins. F.M.S. Malayan Forester, etc.
295
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
NOTES ON THE SIGNIFICANCE OF SOME VERNACULAR NAMES
OF FOOD PLANTS IN THE SOUTH PACIFIC ISLANDS
JACQUES BARRAU
South Pacific Commission, Noumea, New Caledonia.
Our knowledge of the vernacular names of
food plants in the South Pacific Islands is far
from complete. The Polynesian and Micro-
nesian names are rather well known, but a lot
remains to be done where Melanesia is con-
cerned. There the multiplicity of language and
dialects makes it sometimes difficult to record
plant names.
Even in this preliminary stage, it may be
useful, in these notes, to study very briefly the
meanings of those names already known.
Besides, a comparison between lists of South
Pacific vernacular names shows apparent
similarities between names in different Mela-
nesian, Micronesian and Polynesian territories.
A study of these resemblances could lead to better
understanding of the significance of vernacular
names.
It may be of interest to consider first the verna-
cular names of food plants introduced into the
South Pacific Islands since their discovery by
Europeans. Their meaning could help under-
stand how the plants used by the islanders in old
times were named. It seems that the vernacular
names of the more recently introduced food
plants can be classified as follows:
/. Plants completely unknown by the islanders in
the p re- European era:
a. The name adopted is the vernacular name
used in the territory from which the plant
was first introduced.
Examples: The sweet potato, Ipomoea batatas
(L.) Lam. was most probably introduced
into Guam Island during the 17th century
from Mexico. The vernacular name in this
island is CAMOTE (or KAMOTE) a name
derived from CAMOTLI (or CAMOTL)
as the Mexican Nahuatl Indians called the
sweet potato. CAMOTE is also used in
the Philippines where /. batatas was probably
introduced from the same area. The first
European discoverers who visited Guam
(cf. PIGAFETTA) mentioned sweet potato
tubers among the local food. I think that
there was a confusion with the tubers of
Dioscorea eseulenta (Lour.) Burk. one of
296
the main subsistence crops in ancient Guam.
Le Gobien's book, "Histoire des ilcs
Mariannes nouvellement converties a la
religion chretienne" published in Paris in
1700 does not mention the sweet potato
among the Guamanian subsistence plants
but says that the bases of the vegetable food
pattern were tubers of NIKA, Dioscorea
eseulenta, and SUN1, Colocasia eseulenta
(L.) Schott.
Another example: Xanthosoma brasilicnse
(Desf.) Engl., an aroid grown for its edible
leaves was introduced into Tahiti from the
West Indies probably at the beginning of the
present century. Today, its vernacular name
in French Oceania is CALALUO, which in
the Caribbean area is the name of a popular
dish, a kind of soup, of which the leaves of
X. brasiliense are the basic ingredient.
b. The South Pacific Islanders found that the
new plant had some resemblance with one
of their traditional food plants, and used
the latter's vernacular name for the new
comer.
Examples: Xanthosoma sagittifolium (L.) Schott
is called TARUA in the Cook Islands and
French Polynesia. This name originally
designated a local pre-European variety of
the banana Musa sapientum L. The cooked
lateral tubers of X. sagittifolium, according
to the islanders, tasted and looked like the
cooked bananas of the variety MEIKA
TARUA. The aroid still bears this last
varietal name.
Another example: The sweet potato, /. bata-
tas, is called KAV by the Marind Papuans of
the Southern coast of Netherlands New
Guinea. KAV as a matter of fact is the local
name of Dioscorea eseulenta the tubers of
which, according to the Marind, look like
those of the sweet potato. Similarly in some
dialects in Indonesia, the sweet potato is
called GUMBILI or KUMBILI, which is the
name of D. eseulenta. This emphasizes the
frequent confusion between the tubers of
the two plants already indicated in the case
of the Guam name of/, batatas.
VOLUME 4
BOTANY
2. Plants belonging to a species (or a variety) of
a genus (or species) already present in the area
before European times.
a. The vernacular name includes the name used
for the genus (or species) plus one or several
words indicating that the new comer was
introduced by Europeans.
Example : The bananas introduced in Samoa by
Europeans, particularly Musa nana Lour.,
are called FA'I-PAPALAGI (VUDI-NI-
PAPALAGI in Fiji), meaning banana (FA'I)
of the European (PAPALAGI). The same
is the case in the Loyalty Islands where
introduced bananas are called WA-WIW1
meaning bananas (WA) of the WIW1 (OUI-
OUI), a nickname given to Frenchmen
because the Loyalty Islanders were appa-
rently much amused by the exaggerated use
of the affirmative GUI in French speech.
b. The vernacular name of the new plant
includes the name used for the species (or
genus), plus the name, more or less trans-
formed, of the territory where the plant was
thought to have come from.
Example: Xanthosoma sagittifoliwn is called
in Ponape SAWA-N'AWAI, literally, the
Colocasia taro (SAWA) of Hawaii
(N'AWAl). The same aroid is called
BISECH-LARUK in Palau, meaning appa-
rently the Colocasia (BTSECH) from Truk
(LARUK), whence the Palau Islanders said
it was originally introduced.
When the islanders had to name a cultivar of
an already known food plant, the vernacular
name was, of course, more complex. In such a
case, in addition to the name for the species, one
or more words indicate :
Either the place where the cultivar was
originally found or grown, or whence it was
originally introduced. Thus one very common
variety of Dioscorea alata L. in New Caledonia
is called KU (or KUK, WUKUK, WU, etc.
according to the name of D. alata in the dialect)
PAPUA. A New Caledonian teacher of the
London Missionary Society brought it back from
New Guinea during the last century.
Or the name of the islander who first used,
grew or introduced the cultivar. Thus a taro,
Colocasia esculenta, grown in Ateu, a highland
village of New Caledonia is called WA (Colocasia)
ENEILOKAPU. The meaning of this last word
is that a New Caledonian woman named KAPU
used to eat only the tubers of this cultivar.
Or, more frequently, a particular character-
istic (shape, size, color) of a part of the plant.
As an example, a variety of the yam Dioscorea
alata is called THAMBWA in Wunjo, a coastal
village of New Caledonia. Among the THAMB-
WA yams, the islanders distinguish:
THAMBWA TANA BWALl: red-fleshed
(TANA), long (BWALI) tubers.
THAMBWA TANA XUMAT: red-fleshed,
short (XUMAT) tubers.
THAMBWA FWAT BWALI: white-fleshed
(FWAT), long tubers.
THAMBWA FWAT XUMAT: white-
fleshed, short tubers.
Or, a word reminiscent of a particular cir-
cumstance associated with the discovery or intro-
duction of the cultivar: Thus a cultivar of
Colocasia esculenta grown in the vicinity of
Canala in New Caledonia is called ENLA-
POCHE. This seems to be an alteration of the
French "DANS LA POCHE," literally, IN THE
POCKET. The islanders said that the first pro-
pagating material of this cultivar was stolen in
another village by a man who brought it to the
Canala area concealed in his pocket.
Sometimes the vernacular name of a cultivar
is even more complex and combines several of
the ways of naming a plant listed above.
Several of the names used in the South Pacific
Islands for subsistence plants grown for their
tubers or as a source of edible starch show some
likeness. Many of them seem to be related to the
BIAH or PIAH of Malaya. Thus Alocasia ma-
crorrhiza Schott is called in Malaya BIAH,
B1RAH or BRAK. Apparently BIAH is also
used for the same plant by the Semang negritos
of Malaya. In Indonesia, we found among other
names, BIRA and WIRE. In Melanesia, this
aroid is called :
In some dialects of New Guinea: ABIR and
WERIAK.
In one dialect of the New Hebrides: BARU.
In New Caledonia: AWERE, PERA, PI A,
PIRA, VIA, VIE.
In Fiji: VIA.
In the latter island group, VIA is also used for
Cyrtosperma chamissonis (Schott) Merr., VIA
KANA. The same aroid is called BRAK in the
Palau Is., this name being used for Alocasia
macrorrhiza in Malaya. In Micronesia, we found
as names for Cyrtosperma PURA, PUNA, and
PULA (Truk), PWOLOK (Ulithi), LOK (Yap).
The PURA series may well be related to the
297
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
BIRA, BRAK, and BIAH series used in Malaya
for Alocasia macrorrhiza. Finally the names of
Cyrtosperma chamissonis in Polynesia, BROKKA
(Ellice Is.), PULA'A, PULAKA or PURAKA
(from Samoa to the Cook Islands) apparently
have also the same origin.
East of the Cook Islands, the names for Cyrtos-
perma chamissonis are different: thus we find in
the Leeward Islands the name APE-VEO, literally
Alocasia (APE) yellow (VEO) and a similar case
in the Marquesas Islands where the name for
Cyrtosperma is TAO-KAPE, literally Colocasia
(TAG) Alocasia (KAPE).
Returning to the vernacular names apparently
related to the Malayan P1AH, BIAH, and BIRAH,
there is, according to H.B. Guppy (1897, the
Polynesians and Their Plant Names, Jour. Trans.
Viet. Inst., 29: 135-170, London) some resem-
blance between some of the Indonesian names of
the sago palm, Metroxylon spp. (LAPIA, P1HIR,
RAMBIA), the Melanesian names of the same
palms (ABI, BIA, IABIA, RABIA) and some of
those used for Tacca leontopetaloides (L.) OK
in Melanesia and Polynesia (PIA, RABIA,
YABIA). Both Metroxylon and Tacca are the
source of edible starch. In the case of Tacca,
it may be further significant that its Polynesian
name PIA has been given to the introduced
Cassava which has replaced Tacca as a source of
starch in many Polynesian islands.
The vernacular names of Tacca are different
in Micronesia where we find the following series
of names from the Marianas to the Gilberts:
GAOGAO, GABGAB, SOBOSOB, MOGMOG,
MOKMOK, MOKIMOK, MUGAMUK, TE-
MAKAMAKA. In the Philippines, Tacca"&
known, among other names, as GAUGAU and
YABYABAN. This last name may possibly
be related to the YABIA of the Fiji Is. In the
British Solomon Islands, there is one name for
Tacca related to this series: MAMAGO.
PIA is not the only name of Tacca in Polynesia
where, in the Samoa-Tonga-Rotuma area names
such as MASOA, MAAENA, and MARA are
found, MASOA being used also in Tikopia.
However, it seems that PIA was also used in
Samoa in ancient times.
Not being a linguist, I am unable to draw
any conclusion from all these names. However,
it seems that the apparent common origin of
vernacular names used for some different basic
sustenance plants of ancient Oceania could
profitably be the subject of further investiga-
tion. Listing the vernacular names of plants
is useful, but a better knowledge of their signi-
ficance seems necessary. In this respect, a
more careful study of the ways used at present
in the islands to name recently introduced
plants could help us understand the meaning
and origin of some of the ancient names.
DISCUSSION
C.G.G.J. VAN STEENIS: It does not follow from the
existence of a field vernacular name that the plant is native
or of ancient introduction. I observed with a plant which
I introduced myself. Use is now made of it as a mild
laxative, and has acquired a fixed vernacular name (En-
phorbia geniculata).
J.H. HURLIMANN: In the Tougan Islands, the name of
the country is sometimes added to a plant name, in order to
characterize a species or variety as the best available, not
for indicating the true origin. One must, therefore, be
careful with conclusions from this type of names.
298
VOLUME 4
BOTANY
ETHNOBOTANICAL PROBLEMS IN
THE COMPARATIVE STUDY OF FOLK TAXONOMY
HAROLD C. CONKLIN
Department of Anthropology, Columbia University, New York, New York, U.S.A.
With specific reference to Malaysian ethno-
botany I would like to discuss briefly six general
problems faced by the ethnographer when
analyzing and comparing cultural systems of
categorizing diverse forms of natural phenomena.
/. The nature of folk taxa (cf. scientific taxa).
Several important differences distinguish folk
taxa from scientific taxonomic categories. The
former are concerned only with locally observable
phenomena. They are based on criteria which
may differ greatly from culture to culture. In
folk taxonomy there are no formal rules for the
recognition or rejection of taxa, though new
categories can be added with considerable ease.
Tn respect to any particular flora, there is no
reason to expect the local folk categories to equal
those of systematic botany either in number or
in size. The Hanunoo of Mindoro, whose system
of folk botany I have studied recently in the field,
classify their local plant world into more than
1,600 categories while botanists catalogue the same
flora species-wise into less than 1,200 groups.
2. Translation and semantic structure. At least
at the level of obligatory categories, native names
for folk taxa always comprise a segment of the
everyday vocabulary of a particular language.
The associated semantic structure of such a set
of lexical items is something to be discovered for
each language, not to be prescribed through
familiarity, in another system, with the concrete
designata of the set. In ethnobotanical terms,
this means that a local system of plant classifica-
tion cannot be described accurately by merely
trying to obtain vernacular "equivalents" for rec-
ognized botanical species. Translation labels
are frequently necessary, but in a strict sense
they can serve neither as definitions nor as
exact equivalents. This well-established, and
perhaps almost obvious, semantic principle is
sometimes forgotten where the assumed absolute
nature of "scientific" names is involved.
3. Hierarchies of generalization. Folk systems
vary in the number and position of their levels
of contrastive usage, and in their methods of
increasing or decreasing specification. Without
the conventions of taxonomic nomenclature, the
investigator of such systems must discover these
levels and the rules determining their use by
internal means, i.e., by intracultural analysis of
varying contexts. The kind of data involved can
be illustrated easily. A Hanunoo, for instance,
may answer correctly a question about the
identity of a particular yam vine with any one of
at least seven answers depending on the context
in which the question is asked, and upon the
degree of specificity required :
(1) kayu a plant, i.e., not a rock, etc.
(2) halaman a domesticated plant
(3) wakat a plant having a vinelikc
stem
(4) tanwn a plant propagated horticul-
turally
(5) 'tihi a yam
(6) 'ubi dungun a dungun yam
(7) dungun a dungun (meaning dark-
colored, of certain root
crops).
In this case, two crosscutting systems of plant
categorization are exemplified, one based on
treatment (2 levels: 2,4), and the other based on
morphological type (3 levels: 1, 3, 5-6).
4. Partial vs. unit concepts. In folk taxonomies,
growth stages or parts of individual biotic units
are often classified in much the same way as unit
concepts of distinct animal or plant types. Con-
fusion can result when these categories differ
widely from the investigator's own folk or scien-
tific concepts, yet it is in these very differences that
clues to the significant features of folk classifica-
tion may be found. A few examples may help
to illustrate this point. One form of Diospyros
discolor is known to the Hanunoo as kamagung
mab'iru. When this tree matures fully its heart
wood is much sought after for spear handles,
and canes because it becomes exceedingly tough
and hard. Conceptually, the plant then becomes
a new entity, baluntlnaw. When the low cattle
sedge pddang (Cyperus kyllinga) sends up tall
slender stems to fruit, it becomes paray-mdya
'sparrow rice.' These and numerous other in-
stances reflect the primary categorization of
plants by the Hanunoo according to contrastive
habits of stem growth.
5. Linguistic structure. The shape and combi-
natorial structure of the linguistic forms which
299
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
designate folk taxa are irrelevant, in a strict sense,
to the analysis of the system of classification
itself, i.e., to the semantic structure. Labels and
categories can change independently and, there-
fore, must be analyzed separately. On the other
hand, a knowledge of the linguistic structure
involved is essential for understanding folk
nomenclature, and in working out this structure,
clues for isolating folk taxa and eliciting informa-
tion about them may be found. In the compara-
tive use of native names, however, particular
care should be exercised to avoid confusing forms
with meanings. In many Philippine and Indone-
sian languages, for example, bdgo or some simi-
lar form refers to the bast-providing vine
Gnetum indicum. The expected Hanunoo cognate
bdgu occurs, but not as the name for Gnetum
indicum which the Hanunoo call sandari. In-
stead bdgu refers specifically only to the inner
bark of another species Gnetum gnemon, known
as lungud, and is the probable root of the plant
name malihdgu which designates a bast-providing
shrub ( Wikstroemia polyantha ) of another family
(Thymaceac not Gnetaceae). Numerous similar
examples can be expected in comparing the
plant names and associated vocabularies of
such related languages. Malay pinang and
Hanunoo pinang, for example, are cognate forms,
but the first is a unit name for the arcca palm,
the second a name for only one type of fallen
areca fruit. In Tagalog kawdyan is a general term
for bamboo, while in Hanunoo kawdyan refers
specifically to only one type of bamboo (Bamhusa
spinosa), etc. In short, plant names are subject
to the same processes of historical change as
other linguistic forms.
6. Synonomy. In a narrow sense, there are
no true synonyms, but terms with overlapping
areas of associated meaning are common in all
languages. At lower levels of generalization,
vernacular names for folk taxa often include
such "partial" synonyms, which must be distin-
guished from terms for separate ontogenetic
stages on one hand and from those indicating
greater or lesser specification on the other. While
such alternative labels may not modify the basic
system of classification involved, they may suggest
historical connections or associated behavior
patterns of interest to the comparativist or to the
ethnographer in other contexts. Over 200
specific plant name synonyms have been recorded
in Hanun6o. In part, they indicate religious
taboos, ritual and literary usage, metaphoric
extensions, and the degree of contact with neigh-
boring speech communities.
300
Some of the implications of the preceding
remarks for the use of vernacular plant names in
reference to, or as a substitute for, scientific
botanical nomenclature (with particular reference
to the Indo-Malaysian area), are as follows:
(a) Without other evidence, one cannot assume
that similar vernacular names, in different local
languages, refer to the same scientific taxa. The
chances for error in this type of guess work are
very great especially where names are short,
traditional orthographic transcriptions are highly
simplified (ignoring many phonemic distinctions),
and many closely related languages are involved.
(b) Most vernacular terminologies include
numerous loans, the shapes of which have been
modified to adjust to the phonological require-
ments of borrowing languages. In quoting or
referring to these forms as vernacular names,
they should be treated like the other forms in
the languages concerned. No advantage is
achieved in trying to represent such labels in the
form of the supposed etymons.
(c) When a local name is used as an equivalent
of a scientific binomial, it ceases to be a vernacular
lable as such, in that it no longer designates a
segment of a folk botanical classification. In other
words, when a local term is selected as an "official
name," it becomes a synonym in the language of
systematic botany and thus it cannot be con-
sidered a native plant name, except historically.
Scientific vocabulary can be prescribed, stan-
dardized, and at least relatively stabilized;
vernacular usage can only be described. Folk
botanical terminology is subject to the same
kinds of modification that affect other parts of the
lexicon of any spoken language.
For the culture historian or phytogeographer
the vernacular names for plants and plant pro-
ducts in the Pacilic-Indo-Malaysian area comprise
a very rich, and as yet almost untapped, source
of indirect evidence of the pre-European spread
of cultigens, weeds, and agricultural practices.
Watt, Heyne, Merrill, Burkill, and other workers
have furnished excellent compilations of local
plant names in restricted parts of this vast and
luxuriant botanical province, but many areas
remain completely undocumented, and very
little use has yet been made of the available
materials. Those of us who are particularly
concerned with the relation of plants and man in
the Pacific hope that recent interest in vernacular
terminology will stimulate further descriptive
and comparative research in this field.
VOLUME 4 BOTANY
DISCUSS/ON
A. CAPELL: The term lona is applied in some Polynesian Newcomers often name local plants after one of their
languages to imported plants, lona being a compass point homeland to which it bears physical resemblance, though
as well as of a specific island. This usage is noticeable, there may be little or no botanical relationship. This fact
especially about Futuna-Aniwa in New Hebrides, and must be taken into consideration in any subsequent study,
is borrowed in some neighboring Mclanesian languages.
301
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
A LINGUISTIC APPROACH TO
THE ETHNOBOTANY OF SOUTH-SAHALIN
ROBERT AUSTERLITZ
Columbia University, New York, New York, U.S.A.
(Abstract)
The area discussed is the southern half of the
island of Sahalin (Japanese: Karahuto), specifical-
ly, the area along the Gulf of Taraika. The ethnic
groups involved are the Gilyaks (G), the Oroks
(O), and the Ainus (A). The G language is not
known to be genetically related to any other lan-
guage. O forms part of the large Manchu-Tungus
stock which occupies many areas from the Enisei
eastwards to Sahalin. No genetic affiliations have
as yet been found for A. Population data: G:
total 4,000 (mostly around the mouth of the
Amur), of which 100 in South-Sahalin. O: total
420 (probably all on Sahalin), of which 250 in
South-Sahalin. A : 1 5,000 in Hokkaido and 1 ,000
in Sahalin. (Figures are approximate and based
on pre-war Japanese and Soviet sources; geo-
graphical distribution is pre-war.)
The material adduced consists of (1) about 170
plant names in (7, O and A, accompanied by
(a) a discussion of the linguistic aspects of each
plant-name and (b) ethnological data concerning
each plant; (2) a list of about 60 terms concerning
plant anatomy, folk-botanical notions, and plant-
artefacts; (3) a list of about 30 topographic terms,
and (4) nine names of plant-commodities (to-
bacco, rice, etc.). The investigation has a Gilyak
bias: information concerning O and A cultures
and languages is considered ancillary although
the optimum, namely, complete information con-
cerning all three cultures, has been striven for.
The information has been mainly collected from
one (7, one 0, and one A informant, but other
informants, when available, have also been con-
sulted. All work was carried out in Hokkaido,
Japan. Reference to earlier sources has been made.
The information was elicited by letting the in-
formants associate freely (singly and in groups),
by asking them to procure specimens from nature,
by reference to older sources (G and A), and by
showing them illustrations in Japanese and West-
ern books on northern flora. The entire collection
of plant names was finally identified (from descrip-
tions and specimens) and provided with scientific
binomials by Prof. M. Tatewaki (Sapporo).
The following points are especially noteworthy
from the viewpoint of ethnobiology: (1) Plant
302
uses: nutritional, industrial, medicinal, ritualistic;
(2) The folk-botanical distinctions made in the
cultures in questions; (3) The etymologies of plant
names and flora terminology in (7, <9, and A.
Accordingly, the preliminary conclusions of
the investigation reveal the following: (1) G, O,
and A plant uses in South-Sahalin overlap to a
great extent. Since we assume a different geo-
graphical and cultural origin for each of the three
groups we must here reckon with cultural borrow-
ing. In the case of drugs, especially, we assume
that the lending party has been the A. In the case
of ritualistic uses, G and O show similarities,
whereas A stands apart. Industrial uses are highly
developed among the A (weaving, dyeing), much
less developed among the O, and primitive among
the G. (2) G culture shows a predilection for so-
phisticated terminology for mosses, lychens and
conifers; this points to a tundra-origin. The same
distinctions are primitive among the A. The O,
who form a part of the large reindeer-culture com-
plex (Tungus) have the terminology to be expected
under these circumstances. Hunting and trapping
are equally represented everywhere, but it seems
that the G terminology concerning topology is the
most developed. (This conclusion may result from
the above-mentioned G bias.) (3) The etymolog-
ical investigation of plant names shows that much
borrowing in all directions has taken place, per-
haps mostly from A to G/O. 1
The overall picture gained confirms and
strengthens previous hypotheses reached by other
means: (1) The G and O moved to Sahalin from
the Amur area. (2) The (9, being reindeer-breeders
came from an originally more temperate zone
than the G (whose reindeer terminology is Tungus/
but not 0\/) (3) The A moved from Hokkaido to
Sahalin and are the most plant-conscious group.
(4) The O seem to have borrowed many cultural
features from the G; this implies a relatively
remote separation from the main Tungus stock,
(5) G plant names are very often analytic, i.e.,
names can be analyzed into components which
are not basically botanical: this would indicate a
relatively recent "awakening of a folk botanical
consciousness." Reindeer-culture terminology
VOLUME 4 BOTANY
and sea-mammal distinctions should be scrutinized broader area of Northeast Asia's flora remains to
together with plant-terminology. On a larger be carried out.
scale, the evaluation of the data in terms of the
DISCUSSION
T.P. BANK: When last I saw Austerlitz's manuscript waki of Hokkaido University (professor of botany) to
(in November, 1957) it had already grown to enormous obtain authoritative plant identifications. When finished
proportions. He is a linguist who has developed a recent (this winter, 1 think) his volume will form a most useful
interest in ethnobotany and he is doing an excellent job of and valuable contribution to the botany and anthropology
his project. Not only is he working with Gilyak, Ovok of the North Pacific region and Northeast Asia. It definitely
and Ainu informants, but he is collecting the plants should be published and 1 hope Dr. Austerlitz can be
wherever possible, recording his data on the plant collection encouraged to continue his interest in ethnobotanical
sheets, and working closely with Professor Misao Tate- linguistics.
Ut is very difficult to reach definite conclusions about the direction in which a loan travelled without disposing over a
greater body of data than we have at present. For the time being, we employ the following methodology which is based
on four foci:
Dialects:
cG sG cG -continental (Amur) Gilyak
sG -Sahalin Gilyak
hA -Hokkaido Ainu
hA sA sA -Sahalin Ainu
If a term in sG and sA contains features which lead us to believe that we are dealing with a loanword and if this term also
exists in cG we conclude that the loan travelled from G to A . If the term exists in sA and sG as well as in HA, we assume that,
it travelled from A to G. If the term exists in all four (sG, c<7, sA 9 hA) we are faced with a larger problem. This method can,
of course also be carried out in respect to A and (employing Tungus cognates for terms). There are, of course, also
other indices, by which loans can be determined. The most useful of these is the evaluation of the structure of the word
within the framework of the structure of the respective language as a whole. Thus, anyone familiar with the structure of G
will conclude that a term like orarif "rosa rugosa Thunb." is not originally G in structure and must be considered a loan.
Another index is the analytic structure of certain plant names: if a term can be decomposed into such meaningful terms as
have a relevance to the plant in question and if the same term reappears in another language, we may assume that the lan-
guage in which the analytic term appears is the lending party, e.g., A otarup "rosa rugosa Thunb.'*, decomposable into ota
"sand" and ru-p "round (thing)". In such cases, however, there is always an imminent danger of folk-etymology on the
part of the borrowing party. Still another index is the number of names per plant : if a language has three or four names for
the same plant (in one dialect), at least one of these is certainly a loan. Etc.
The large collection of A plant names by M. Tiri is now available. For G, however, we only have the century-old col-
lection of Grube (Schrenck and Glehn). A source on cG plant names would be a most welcome aid.
303
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
VERNACULAR OR COMMON NAMES OF PLANTS
R.E.P. DWYER
Department of Agriculture, Stock and Fisheries, Port Moresby, Papua.
It is considered a great honour to be asked to
present a paper even so limited as this one to the
members of this Ninth Pacific Science Conference,
especially to that Sub-Committee or section,
dealing with vernacular names of Pacific plants.
At the request of Dr. Mona Lisa Steiner of the
Philippines, Convenor of the Symposium on
Vernacular Names of Pacific plants, lists of
common names of, mainly, Papua and New
Guinea plants have been compiled. These have
been done from a number of sources, by Depart-
mental ofilcers of the Department of Forests,
particularly the Forest Botanist, and the Depart-
ment of Agriculture, Stock and Fisheries, and the
author (see acknowledgments). The actual
compilation and collation of the plant lists for
this submission is definitely the main purpose
envisaged.
The attached nomenclature lists 1 are presented
in three phases:
(1) A list of native, or vernacular names of
plants, from Papua and New Guinea,
including the out-lying islands.
(2) A shorter, comparable list from the neigh-
bouring British Solomon Islands and other
South Sea Islands showing some common
names which were collated while compiling
the above list. This list is supplementary
and most incomplete.
(3) A list of some comparative names, chiefly
of food plants, from Micronesia, Polynesia,
Melanesia, Indonesia and Malaya is also
included. See Barrau (1 and 2) Merrill
(7) Ochse (9). This is a limited attempt at
comparative nomenclature to try and find
out whether any affinity in the common
names used exists for specific plants for the
various territories and districts represented.
Refer to the more comprehensive list by
Dr. Steiner, Philippines, 1957 (11).
The lists of plants submitted herewith are
generally presented under the following headings:
(a) Scientific name.
(b) Native name.
(c) Dialect and locality.
(d) District.
l These lists are not published in the Proceedings but will form part of the dictionary of vernacular names being compiled
by the sub-committee on Vernacular Names of Pacific Plants, Standing Committee on Botany, Pacific Science Association.
304
It will be seen that widely-used, ordinary,
common names such as bananas, coconuts,
breadfruits etc. have been omitted as these are
readily available.
The present author was formerly working as
an economic botanist, up to 1942, in the old,
New Guinea Department of Agriculture, so that
the assembled scientists will realise that the com-
parable data deals with food and important eco-
nomic plants, but not purely in timber, or forest
trees, as such, or with plants of mainly botanical
interest.
The lists of plant common names, although
comprising an extensive collection, can only be
regarded as being most incomplete. From the
experience presently gained in these compilations,
it would appear that it would take months, or
longer to collect and present the data in such a
way as to be of maximum use to the working
botanist. The immensity of the task involved
in collecting vernacular names of Papua and
New Guinea plants is highlighted by the following
considerations:
(1) The very diverse physiography present over
the Territory of Papua and New Guinea as
briefly there are dry savannah areas, such as
in East Papua and very wet areas, up to
nearly 300" per annum, in some districts.
The average rainfall for the whole Territory
would probably exceed 80" per annum.
The Territory can therefore be generally
regarded as wet, except in the south east
coast of Papua, in the dry belt.
(2) Variations in altitude ranging from sea
level to mountains of 15,400 ft in height,
with immense, highland valleys of around
6,000 ft. It is estimated that more than one-
third of the Territory is above medium to
high-altitude terrain.
(3) The hundreds of islands present, including
coral atolls, archipelagoes, etc. Thus
diverse genera and species of plants are
found in some islands in this Territory
which are not present in others, due to
different ages of geological separation,
difficulties and types of dispersal etc.
VOLUMF 4
BOTANY
(4) The great diversity of peoples and languages
represented in the Territory's population,
presents a great difficulty in compiling
reliable data on plant names. This is
perhaps the most difficult obstacle to get-
ting anything like a complete survey of all
the local names used by the indigenous
inhabitants. It is necessary to point out
that the, approximately, 1 J million inha-
bitants here speak several hundred major
group languages, many of which include a
number of different dialects as well as
"lingua franca" languages such as Motu,
Tolai, Kote, Yabim, Pidgin English etc.
The peoples here arc roughly classed as Me-
lanesians by anthropologists, yet there are true
Microncsians in the Western islands, Polynesians
in the outer islands (in the far Eastern part of the
Bougainville District), also Negrito people in the
hinterland, and also mixtures of these types in
Papua and New Guinea. The present data are
presented with some trepidation, therefore, es-
pecially as lack of time has prevented the listing
of the plants under their botanical orders, or
generally the names of describing authorities.
There has been no presentation of brief descrip-
tions as has been so well done by Record dealing
with Watcrhouses Collection (10) 1932-36. Lane
Poole (4), Barrau (1, 2) and many other authors
see (3.4,5,6,7,8,9).
The native peoples of the Territory are parti-
cularly adept at recognising and naming plants,
in the field, as they are so important for their
subsistence. They can even recognise and name
various fungi, both those which can be used for
food and those which are poisonous. Even
luminous fungi are recognised and used in games.
It would be valuable to know how far they go in
recognising sea-weeds, marine plants etc. It is
known that some coastal natives do use sea-weed
as food and also in their totems.
Professional botanists are well aware of the
various main Flora, far too extensive to be quoted
in the present bibliography, which describes the
Flora of the South Pacific and Indo-Malaysian
region, e.g., Flora Malesiana; Flora Papuasian,
Flora Kaiser Wilhelmsland, etc. Few of these
flora or the numerous, separate, botanical mono-
graphs in this region present vernacular names.
However, a limited bibliography of books and
articles, where common names are quoted, is
presented.
There is also a wide field not touched on here,
where widely-used common names are given in
the main languages to represent plant life, or
groups of flora, as a whole, so just a few examples
are given:
"KUMLTor^KUMUL"
A Pidgin term for all green
vegetables
"P1CUS" Pidgin for all species of Ficus
"KAPIAK" Pidgin for
Artocarpus Spp.
44 BIN" Pidgin for Legumes
"KUNDA" Pidgin for all rattans, Calamus
Spp.
"RET' Motu for all grasses, Grami-
neae
"M ULIS" Pidgin for all limes and lemons
Citrus Spp. generally
"SEPOR A" Motu corresponds with above-
all Citrus Spp.
cf. "DJERUK" for all Citrus in Indonesia.
Where important food and economic plants are
widely used for a number of purposes, as with
the coconut, Cocos nucifera all parts of the plants
receive vernacular names which fact, in itself,
can be very confusing, as can be the practice that
food products made from the fruit, or plant,
itself, also receive separate names, see Massal
and Barrau (6). It is the author's considered
opinion that a properly-conducted survey of the
range and usage of common names and their
implication and comparisons would yield much
valuable data as to plant origins and plant migra-
tions in the South Pacific area. The indications
are very strong, though not definitely proved,
that where plants have been introduced and are
not widely used, only a few common names,
often similar in sound and derivation, are used
and even where the variations occur, they are
readily traceable. However, even with intro-
duced plants such as sweet potato, when they are
spread over a wide area range of Territories,
many common names may be applied, especially
as numerous varieties of sweet potato Ipomoea
batatus are cultivated. Even in such cases,
according to Barrau (loc. cit.) and other authors,
it is possible to trace a similarity in common
names between those used in Micronesia, Melane-
sia and Polynesia. It is an interesting fact that
the sweet potato has no known representative in
the wild plant sector. It would appear that, his-
torically, it is not many centuries ago this plant
was introduced to the South Pacific region.
There are numerous introduced plants in this
Territory which arc so ubiquitous that they are
practically accepted as indigenous. Examples
305
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
are well known, such as the pawpaw, Carica
Papaya, the betel nut, Areca catechu, pineapple,
Ananus cosmosus, tobacco, Nicoticma tabacum,
which was probably brought here not long after
it was introduced to Malaya and the Philippines,
about 1610, and there are many other examples
too numerous to mention. In the reverse direction,
the position exists where various types of taros
have been distributed from this region to other
parts of the world, particularly the South Pacific
Region, bread fruits Artocarpus spp., sugar canes
Saccharum spp., okari nuts Terwinalia spp. which
have similar common names even as far afield
as Malaya and Indonesia. Again, these only
present a suggestion of the movements which
have taken place and would need more investiga-
tion as in fact has been tried by Merill (7) and
Barrau (loc. cit.) and others.
There is some evidence from the study of com-
mon names of coconuts and from local folk-lore
that they have been introduced into the Pacific
Islands and particularly to New Guinea within
the last two or three centuries and enquiries from
the inland people here seem to denote this. The
common name "Niu" (New) is very commonly
used throughout the area. This may, or may
not, be very significant. The original introduction
of the ordinary, tall coconut should not be con-
fused with the spread of the Malayan and dwarf
types from the East which can be traced to within
a relatively few years ago.
The study of the common names of plants used
in the Territory indicates that often where the
names seem to vary very much, they have rela-
tively the same meaning though expressed in
different languages. Examples are: in Bougain-
ville a number of different names which all mean
"possum's ear" as applied to the peculiar shape of
the small flower of Aporosa papuana and there
is no douht about the similarity which has been
noticed. Tn some cases, the names have magic,
or taboo meanings, e.g. Codiaeum variegatum has
the Siwai name, "tatovo" which refers to the use
of the live plants to define prohibited areas;
Entada phaseohides, the Siwai name, "aija"
means crooked, also for Boehmeria platyphylla,
"Omoro" equals tail of a fish, and numerous
examples could be used to show the derivation
of the common names which have reference to
their appearance, their usage, their ceremonial
importance, economic importance, and so on. Tn
some cases, the inhabitants use common names to
separate large divisions in the one genus; e.g. the
use of the derivations "moi" and "pii" to separate
the large divisions of the Genus Canariwn.
306
One interesting feature is that a large number
of villages and plantations and, in a few instances,
districts and subdistricts have been named after
the common names used for wellknown plants.
It is possible that the plants are named after the
place names, or naming occurs in both directions.
This subject would warrant further study though,
of course, there are examples of where this has
been done deliberately, as in one suburb of Port
Moresby, where common names of trees are
used to denote the streets and avenues, e.g.
Okari Avenue, Hohabu Avenue etc.
ACKNOWLEDGMENTS
The Author is indebted to the Director of
Forests, Mr. J.B. MacAdam; Mr. J.A. Womer-
sley, Forest Botanist, Lae; of the Territory
Department of Forests for their very helpful
assistance.
In addition, Mr. W.L. Conroy, Chief of the
Division of Agricultural Extension and Mr. F.J.
Broekhals, A/Plant Introduction Officer, both of
the Department of Agriculture, Stock and Fish-
erics, Papua and New Guinea; have greatly
assisted.
The idea of these compilations, however,
originated from Dr. Mona Lisa Steiner as stated
in the text and, last of all my thanks to Dr. J.J.
Szent-Ivany, who is to present the brief, introduc-
tory article.
BIBLIOGRAPHY
7. Barrau, J., 1955, Subsistence Agriculture in
Melanesia, South Pacific Commission,
Noumea, New Caledonia.
2. , 1956, Polynesian and Micronesian
Subsistence Agriculture; South Pacific
Commission, Noumea, New Caledonia.
3. Cranwell, L.M, Green, J.E., Powell, H.W.B.,
1944, Food is Where You Find It,
Auckland Institute and Museum; Fourth
Impression.
4. Lane-Poole, C. E., 1925, The Forest Re-
sources of The Territories of Papua and
New Guinea, Report, Govt. Printer,
Melb.
5. Malcolm, S., and Barrau, J., 1956, Yams,
South Pacific Commission Quarterly Bul-
letin, Noumea.
6. Massal, E., and Barrau, J., 1954-1956, Bread-
fruit; Taros; Sweet Potatoes; Casava;
The Banana; The Coconut; Various
articles; October, 1954 - April, 1956,
South Pacific Quarterly Bulletin,
Noumea.
7. Merrill, E.D., 1943, Emergency Food Plants
and Poisonous Plants of the Islands
of the Pacific, War Dept. Tech. Manual
T.M., 10:420.
8. Ochse, J.J., 1927, Indische Vruchten, Depart-
ment Landbouw, Buitenzorg, Uitg.-
Volkslectuur.
9.
_, 1931, Indische Groenten, Depart-
J/OLUMF 4 BOTANY
ment Landbouw, Buitenzorg, 1931, Uitg.-
Volkslectuur.
10. Record, M., 1945, A Collection of Woody
Plants from Melanesia, (Collection
J.H.L. Waterhouse) New Guinea, 1932-
-36: Tropical Woods, No.81, Yale
University.
11. Steiner, M.L., 1957, A Preliminary Compila-
tion of Vernacular Names of Food
Plants in the Pacific, Pac. Sci. Assocn.
Sub-Committee, on Vernacular Plants,
Manila, Philippines.
DISCUSSION
F.R. FOSBPRG: The author has pointed out the matter of
collective names for groups of plants with some general
resemblance, some point of utility in common, or some
other feature. This idea should be emphasi/ed. It seems
to be a general feature of all languages, at least those with
which I am familiar to have both general and specific
names. This situation may often mislead the botanist or
collector, who may put on his label a name which only
indicates a grass-like plant, an edible fruit, a potherb, or
some similar term, not knowing that it is not a spccitic
vernacular name for the plant in hand.
p. WEATHER WAX: I have found this a most interesting
and stimulating symposium, and it is to be hoped that it
may provide an incentive and a pattern for further work.
One interesting study might consider what we are to do
about vernacular names which are obviously incorrect in
application. Example: the yam sweet potato potato
complex.
Despite the fact that studies of this sort can never be
completed and that they often arouse feelings of disgust
or at least impatience in workers in experimental botany,
they certainly fill a great need and merit consideration.
A warning against "forcing" vernacular names. Many
species, many grasses for example, are not clearly recog-
nized as separate entities, and it is foolish to try to find
common names for them. It is even worse to create com-
mon names for such plants when none exist.
Another warning about the informant who, wishing to
please you or feeling that it is necessary to save face,
"manufactures" information for you.
307
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
ORIGIN OF THE SUSTENANCE PLANTS OF POLYNESIA, AND
LINGUISTIC EVIDENCE FOR THE MIGRATION ROUTE OF
THE POLYNESIANS INTO THE PACIFIC
HAROLD ST. JOHN
Senior Professor of Botany, University of Hawaii, Honolulu, Hawaii.
(Abstract)
One of the riddles of the Pacific is the origin of
the Polynesians and how they crossed the Pacific
to discover and colonize the distant isles of the
tropic sea. It happened more than a thousand
years ago. They were a people of a stone age
culture and without a written language, so there is
no record in written history of their maritime
explorations. Yet, in the central, and especially
in the eastern Pacific they found and settled all
fertile islands of the tropics and even some down
into the south temperate zone. In the many
earlier attempts to unravel the history of the
Polynesian migrations, much evidence has been
studied, most of it of an ethnological nature.
The writer has approached the problem from a
botanical angle, attempting to solve the riddle by
a study of the crops that the Polynesians culti-
vated and ate.
The Polynesians had a highly developed agri-
culture based upon the growing of 27 species of
crop plants. One, Piper methysticum, was a bever-
age plant, the others were food plants. By the
known source and by phylogeny, the home land
of these crops can be determined. One, Ipomoea
batatas, is demonstrably of American origin, but
in aboriginal times was carried by native people as
far west as New Guinea. One was domesticated
in Polynesia; three in Polynesia and adjacent
Melanesia, two in Melanesia, and one in Micro-
nesia. These central Pacific ones make 25 per cent.
The great majority of them came from farther
west, 7 from Malaysia, 6 from Malaysia and
southeast Asia, 3 from the shores of the southwest
Pacific and the Indian Oceans, and 3 from India
or Ceylon. These Oriental ones total 70.4 per cent.
Hence, the geographic origin of their crops im-
plies that the people brought them from South-
east Asia.
Evidence of origin can also be found in the
vernacular names used by the many tribes of
aborigines. It was pointed out by S.H. Ray
(1912) that the coconut, Cocas nucifera, was
known by the name "niu," in that same or in a
cognate form of the word, all the way from
Madagascar to Hawaii. The same wide usage is
true of the name "taro" for Colocasia esculenta.
Others of the 27 crops have names with a wide
use, but over area of less size than the whole
tropical Indo-Pacific.
Since all but one of these crops are of Asiatic
or Pacific origin, it would be of interest to find the
route over which they were imported, that is,
either through Micronesia direct to Polynesia,
or through Indonesia and Melanesia to Polynesia.
The aboriginal occurrence and use of the crop
plants gives the best evidence. In aboriginal
times, in all Micronesia 4 of the crops were miss-
ing, while in the Marshall, Gilbert, and Ellice
Islands, all of which are atolls or cora islands,
1 1 were missing. On the other hand, all 27 were
in use by the natives in all or in several parts of
the East Indies. This supports the theory that
Polynesians, emigrating from Southeast Asia,
followed the chain of islands of the East Indies or
skirted and touched them while migrating
through Malaysia and Melanesia to Polynesia.
308
VOLUME 4
BOTANY
EDIBLE YAMS OF THE SOUTH SEA ISLANDS,
SPECIES PRESENT, VERNACULAR NAMES AND DISTRIBUTION
JACQUES BARRAU
Museum National d'Histoire Naturelle, Labor at oirc d* Agronomic Tropicale, Pat is ; France.
Yams are important food-producing plants to
be found in all the high Pacific islands. It may
be of interest to give here a list of the species
together with the vernacular names known and
the distribution of the plants. A more complete
study has been given elsewhere (Journal of Tro-
pical Agriculture and Applied Botany, III, 7-8,
1956). The enumeration which follows is based,
from the botanical point of view, on BurkilPs
monography, "Dioscoreaccae" (in Flora Male-
siana, I, 4, 3- 1951).
I. Section Combilium
Dioscorea csculenta (Lour.) Burk. [D. sativa
Linne, D. aculeata Linne, D. fasciculata Roxb.,
D. spinosa (Roxb. ex) Hooker, D. papuana Warb.]
Two botanical varieties of these are present
in the South Sea Islands:
spinosa var. (Roxb.) Prain and Burkill,
corresponding to the very prickly yams of the
species abundantly found in the wild state in the
island of Guam in the Marianas archipelago.
fasciculata var. (Roxb.) Prain and Burkill,
which groups together all the less prickly horti-
cultural varieties common from New Guinea to
the Samoa islands inclusive.
Vernacular names :
1. Melanesia:
New Guinea: Diba, Kav, Kava, Nak\ Ncu\
Nemu, Taitu. Taitukava
Trobriand islands: Taitu
New Caledonia: Diwada, Kudik, Kudim, Ku
din, Si, Ubin, Wale, Walei,
Ware, Warei
2. Micronesia:
Yap: Dai
Marianas islands: Nika
3. Polynesia: Of a lei, Ufi lei, Uhlei.
The species is only found plentifully as far as
the islands of Samoa included. Further eastwards
it is rare and its introduction perhaps recent.
II. Section Paramenocarpa
According to Sasuke, "Breadfruit, yams and
taros of Ponape island," Proceedings of the
Seventh Pacific Science Congress, VI, 1953, one
of the yams of Ponape is D. flabellifolia Prain
and Burkill, the presence of this yam in the Palau
islands having already been pointed out by
Burkill.
HI. Section Opsophvton
Disc area hulhifera [D. sativa (non Linne sp.
pi.) Thunb., D. sercideal mentioned by Langley
in the "Report of the 1947 New Guinea nutrition
survey expedition," 1951].
Four varieties are present in the South Sea
Islands:
(a) Wild tuberculous and bulbous plants which
are bitter and poisonous,
hulhifera Burk. var. with cordate leaves,
hcterophylla Burk. var. with more elon-
gated leaves.
These two varieties are present in almost all the
Upper Pacific Islands.
(b) tuberculous and bulbous plants with a
less bitter taste and only slightly or non-
poisonous, probably the outcome of empiric
selections in ancient times,
suavior Burk. var. bulbils with greyish and
uneven epidermis, present in almost all the Upper
South Sea Islands,
-sativa Burk. var. bulbils with smooth and
light coloured epidermis, to be found in New
Guinea.
Vernacular names :
I. Melanesia:
Trobriand islands: Kasiena
New Caledonia: Ae, Ave, Dema, Dimoa,
Dimwe, Eja, Niwa, Nua,
Numwe, Soa, Sua, Samoa,
Waeki, Wasuma
Fiji islands: kaile
II. Micronesia:
Palau: Belhi
Yap: Rok, Yoi
Truk: Apuereka
Ponape: Palai
309
PROCEEDINGS OI THE NINTH PACIFIC SCIENCE CONGRESS
III. Polynesia: Hoi, Soi, Oi.
IV. Section Lasiophyton
1) Dioscorea pentaphylla Linne
Two varieties are present in the South Sea
Islands:
papuana Burk. var. for hardy plants with
prickly stems and tubers carrying numerous
roots,
-palmata Burk. var. for the plants with short,
pyriform tubers which are sometimes flattened
and lobcd and which have a smooth epidermis.
The "cultivar Piia" described by St. John, "The
Hawaiian variety of D. pentaphylla, an edible
yam," (Journal of the Polynesian Society, 63, 1,
1954), seems to correspond to this latter variety.
The former variety is found principally in New
Guinea, the latter is common as far as Eastern
Polynesia.
Vernacular names :
T. Melanesia:
New Caledonia: Tue, Wahneor, Xota
Fiji islands: Bulou, Kaile, Tokatolu
II. Micronesia:
Yap: Duol
III. Polynesia:
Samoa island to the Cook islands at Raro-
tonga: Pint a
Cook islands to the East of Rarotonga:
Pakatiro
Society islands: Parauara
Marquesas islands: Utau.
2) Dioscorea hispida Dennst. (D. triphylla L.,
D. hirsuta Dennst.)
This variety is to be found only in New Guinea
and at Ponape, Caroline islands. I have found it
in the cultivated state in the latter island.
V. Section Enantiophyllum
1 ) Dioscorea alata L.
The most important species with multiple
clones in the South Sea Islands.
Vernacular names:
I. Melanesia:
New Guinea: Ambi, Dari, Kav, Krenup, Nar
Trobriand islands : Knvi
New Hebrides : Dem, Ngaka, Pae, Taufi
310
New Caledonia: Ko, Koko, Ku, Kuik, Kuk,
Kuvik, Nuvuk, U, Ufi, Uvi,
Uvuk, Wakoko, Wu 9 Wu-
kuk, Wuvuk
II. Micronesia:
Palau : Telngot
Yap : Duok
Marianas islands: Dago
Caroline islands a) Truk: Ep
b) Ponape to Pingelap:
Kap, Kep
111. Polynesia: Pttauhi, Ufi, U/ri.
D. alata is present and cultivated in all the
high South Sea Islands. This yam is a basic food
plant in certain territories of Melanesia only
(Trobriand islands and New Caledonia, for
example) and at Ponape in Micronesia. Else-
where, in eastern Polynesia in particular, it is
a subsidiary food plant.
2) D. Nummularia Lamk. (D. pirita Nadeaud,
D. glabra (non Roxb.) Koorders, D. seernanii
Prain and Burkill, D. hebridensis Knuth, D. palau-
ensis Knuth, D. angulata Knuth, D. lufensis
Knuth, etc.)
This yam was identified by Brown, "Flora
of southeastern Polynesia, 1931," who gathered
it in the Marquesas as D. cayenensis Lamk. His
conclusion was that its presence indicated ancient
contacts with America, an argument which was
taken up by Heyerdahl "American Indians in the
Pacific, 1952."
Vernacular names :
I. Melanesia:
New Guinea: Baku, Sinngo
Salomon islands: Karausu, Pifuitiko
New Hebrides: Loopitiko, Telewo
Fiji islands: Rauva, Rauvadna, Tivoli, Tikau
II. Micronesia:
Yap: Sep
Ponape: Kapnair
III. Polynesia:
from Niue to Rarotonga : UhiPalai, UfiParai,
Ufi Palai
from Atiu to, and including, the Society
islands: Pirita
Marquesas islands: Puauhi-Peahi.
D. nummularia has, so far, never been found
in New Caledonia. Although this species is
present in almost all the high South Sea Islands,
it has never been extensively cultivated. Just
a few plants are to be found growing often
_VOLUME_4 BOTANY
in the villages, sometimes in gardens, more by usage of which almost completely disappeared
tradition it seems than by necessary. It may on the introduction of food crops easier to pro-
be a case, as suggested by Guppy, "Observations duce and prepare. Guppy classed D. nummu-
of a naturalist in the Pacific 1906," of a food loria among what he called "pre-polynesian"
plant of more importance at one time but the food plants.
311
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
Symposium: Vernacular Name of Plants
GENERAL DISCUSSION
F.R. FOSBERG: Dr. Carter of Johns Hopkins University,
on seeing the preliminary compilation of names of food
plants, brought up the desirability of extending this work
to include meanings and derivations of these names. This
would provide additional clues to the origins of the plants,
migration of peoples, development of languages, etc. This
extension might also secure for us the cooperation of able
people in these fields who might not otherwise be interested.
J.H. HURLIMANN: A joint botanic, linguistic, and
anthropologic study of the plants of Tonga seems to be
feasible, because materials have been gathered in the three
fields of approach in 1951-1952.
A. CAPI:LL: Linguists arc in the process of organizing
symposia for the next Congress. I suggest that arrange-
ments may be made for a joint session between Botany and
Linguistics on ethnobotany. There is still time to arrange
this through the relevant sub-committees.
T. TAKESI: I wish to call attention to the problem of
Pandanus. There are many edible species of rather limited
distribution. As an entry in Dr. Steiner's project, I suggest
that the generic name is enough, after the vernacular names
of different localities, specific names may be better recorded
in parenthesis.
F.R. FOSBFRG: The matter of vernacular names of
Pandanus will not be straightened out to everyone's satis-
faction. The Pandanus tectonics group includes a myriad
of cultivated and wild forms comparable to those of the
domestic apple.
These have vernacular names just as do apple varieties.
No one agrees on which arc to be regarded as species.
Edibility is no criterion. When seeds of some of the large
edible varieties are planted, the resulting plant may be the
small fruited with plant. There are reasons for the diversity
of names, in that the fruits are of different quality. It is
desirable to record these things, but we should not delude
ourselves about getting anywhere with it.
T.P. BANK: I don't wish to take sides in this argument
except to point out again that from an anthropological
point of view some of these periphereal names, these syno-
nyms for the same species, are most significant. For
example, among the Ainu there are many names for the
same tree, depending upon whether or not it is scared
(and the spirits have escaped). The Ainu have a whole
complex of ritual associated with tree spirits, and the
accompanying names arc most important in gaining an
insight for anthropologists.
j.v. SANTOS: With regards to the accuracy of vernacu-
lar names of plants, I have the impression that there may
be an occasion when there is danger of considering seriously
some names furnished by the informers. Based on my
little experience on collecting trips in the Philippines,
1 gathered the information that there were times when a
guide did not know the name of a plant presented to him
and in his desire to satisfy the mquisitivcness of the
collector, he (the guide) supplied any vernacular name
which came to his fancy.
312
VO1.UMI: 4
ACETO-CARMINE STAIN IN THE CYTOGENETICAL
INVESTIGATION OF SOME MARINE ALGAE OF
THE PACIFIC COASTt
BOTANY
KATHLEEN COLE
Department oj Biology and Botany, The University of British Columbia, Vancouver, Canada.
There exist many problems concerning the
speciation of marine algae on the Pacific Coast
of North America. A cytogenetical study is fun-
damental to an understanding of speciation
and, in conjunction with culturing, helps to
correlate the presence or absence of morpholog-
ical alternation of generations with cytological
phenomena.
Information pertaining to the life histories of
the green, brown and red marine algae has been
obtained by improved culturing techniques and
has assisted in the interpretation of phylogenetic
relationships within these groups. Investigations
within the Chlorophyta have proved that, in the
order Ul vales, the genus Monostroma lacks an
alternation of generations, thus differing from
the genera Ulva and Enteromorpha (Kunieda,
1934; Moewus, 1938; Yamada and Saito, 1938).
Kornmann (1938) claimed that Halicystis and
Derbesia, genera of two distinct families, Hali-
cystidaceae and Derbesiaceae, of the order
Siphonales, are phases in the life history of one
and the same alga. Others, including Hollenberg
(1935), have shown that they are distinct genera.
It is evident that Kornmann's work is in need
of repetition since the conclusions regarding the
life histories differ so greatly. Cultural studies
have also made it apparent that the algae olfer
possibilities for genetical studies such as the
lieritability of sex determination already demon-
strated in Dictyota and Laminaria saccharina (L.)
Lamour. (Schreibcr, 1930, 1935), two members
of the Phaephyta.
There is a notable lack of algal cytological
literature dealing with problems such as the
nature of cell division and the nature and function
of the pyrenoids, ccntrioles and nucleolus. This
is understandable when it is realized that technical
difficulties arise in dealing with many of the algae
and especially those having gelatinous wall layers,
large vacuoles and very small nuclei. This small
size of the nuclei mitigates against critical obser-
vation of certain important phases of cell division.
In spite of this, more or less typical mitotic
divisions similar to those of higher plants have
t Presented by R.F. Scagel.
been reported for representatives of all the algal
groups.
A cylogenetical investigation of some marine
algae of the Pacific Coast was begun recently
at the University of British Columbia. This
study includes the culturing of several forms in
an attempt to obtain complete life cycles. The
iron-aceto-carmine squash technique (Belling,
1926) which has been used successfully with
many dilferent animals and plants, including
certain fresh water algae (Cave and Pocock, 1951),
was selected for the cytological observation of
all species. Squash preparations have the advan-
tage over sections in that they permit rapid fix-
ation and handling of materials and the exami-
nation of single layers of cells in totality. The
combined stain-fixative is the simplest of all
chromosomal treatments since cells are directly
mounted and studied in the fixative. The mounts
may be readily prepared in the field and the stain
applied to living material or to material which
has been previously fixed. Prefixation of the
material reduces the staining of the cytoplasm
and allows more rapid absorption of the stain
by the chromatin.
A general survey of the Chlorophyta, Phaeo-
phyta and Rhodophyta, with emphasis on the
benthonic marine algae, was conducted first and
it was soon evident that the iron-aceto-carmine
effectively stained the nuclei in all examples
studied. Certain modifications of the technique
were required in some instances. There follows
an account of the techniques found most suitable
for genera of the three major algal divisions.
I. Chlorophyta. Species of the following
genera were studied: Chaetomorpha, Cladophora,
Codium, Derbesia, Dunaliella, Haematococcus,
Halicystis, Rhizoclonium, Spongomorpha and
Urospora. Prefixation in l-and-3 acetic alcohol
for a period of 12 to 24 hours facilitated nuclear
staining. A small portion of the material was
placed on a slide with several drops of iron-
aceto-carmine stain and a cover slip applied.
The slide was heated gently over a spirit lamp
313
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
for several minutes to flatten the cells, stick them
to the slide and speed up penetration of the stain.
Slight pressure was exerted on the cover slip
with the thumb or the end of a pencil. Temporary
slides were made permanent rapidly and easily
by means of the quick-freeze technique (Conger
and Fairchild, 1953). Following this treatment
most cytological features such as flagellae, cell
walls, cytoplasmic inclusions, pyrenoids, nuclear
membranes, nucleoli and chromosomes were
readily observed.
II. Phaeophyta. Species of the following
genera were studied: Nereocystis, Pleurophycus,
Soranthcra and Sphacelaria. Slides were pre-
pared as for the Chlorophyta with the exception
that when the iron-aceto-carmine was applied
to living material it was necessary to add one
drop of iron alum solution (23 ml of 3 per cent
aqueous iron alum in 77 ml of 90 per cent alcohol)
to the slide in order to speed up absorption of
the stain. Nuclei and mitotic figures were observ-
able when this procedure was followed.
III. Rhodophyta. Species of the following
genera were studied : Antithamnion, Bangia, Ccilli-
thamnion, Ceramium, Corallina, Cryptopleura,
Dasyopsis, Euthora, Fryecla, Gonimophyllum,
Herposiphonia, Nitophyllum, Platythamnion, Poly-
coryne, Polyneura, Polysiphonia, Porphyra, Ptero-
siphonia, Sarcodiotheca, and Wccksia. Certain
of the red algae studied required more modifi-
cation of the staining technique than the greens
and browns. Slides of Antithamnion, Callitham-
nion, Herposiphonia^ Plalythamnion, Polysiphonia,
Ptcrosiphonia and Dasyopsis were prepared as
for the Phaeophyta. Species of the other genera
listed above were fixed in either alcohol-formalin
or Karpcchenko's chrom-acetic-formalin, since
decolorization of the tissues was necessary.
Before staining, treatment with 50 per cent HC1
softened the calcified thallus of the corallines and
facilitated separation of the cells of others whose
tissue had hardened during fixation. Porphyra
and Bang ia required pre-mordanting in the iron
alum solution. A small portion of the material
was placed on a slide together with a drop of
iron-aceto-carmine stain, a drop of iron alum
solution and a drop of liquid detergent. Portions
of the thalli of the genera studied required
vigorous heating before pressure was applied
to the cover slip. The use of this technique
allowed the clear observation of nuclei and mitotic
figures.
In conclusion, it has been demonstrated that
the iron-aceto-carmine squash technique, with
modifications, may be used as a nuclear stain
on marine forms of the Pacific Northwest. It is
particularly suitable for a cytogenetical survey
because it is simple, rapid, stains the nucli*-
effectively and may be used in the field if neces-
sary. The survey is being continued and the study
intensified in order to obtain chromosome counts
and a more detailed knowledge of the cytological
aspects of the life cycles of many of the marine
algae of this region.
BIBLIOGRAPHY
Belling, J., 1926, The Iron-Aceto-Carmine Method
of Fixing and Staining Chromosomes, BioL
Bull., 50: 160-162.
Cave, M.S., and Pocock, M.A., 1951, The Aceto-
Carmine Technic Applied to the Colonial
Volvocales, Stain Techn., 26: 173-174.
Conger, A.D., and Fairchild, L.M., 1953, A
Quick-Freeze Method for Making Smear and
Squash Slides Permanent, Stain Techn.,28:
281-283.
Hollenberg, G.J., 1935, A Study of Halicystis
ovalis, I: Morphology and Reproduction,
Amer. Jour. Bot., 22: 782-812.
Kornmann, P., 1938, Zur Enthwicklungsgeschi-
chte von Derbesia und Halicystis, Planta,
28: 464-470.
Kunieda, H., 1934, On the Life-History of Mono-
stroma, Proc. Imp. Acad., Tokyo, 10: 103-106.
Moewus, F., 1938, Die Sexualitat und der Gener-
ationswechscl der Ulvaccen und Unter-
suchungen iiber die Parthenogenese der
Gameten, Arch.f. Protistenk, 91: 357-441.
Schreiber, E., 1930, Untersuchungen iiber Par-
thenogenesis, Geschlechtsbestimmung und
Bastardierungsvermogen bei Laminarien,
Planta, 12: 331-353.
_, 1935, l)ber Kultur und Geschle-
chtbestimmung von Dictyota dichotoma,
Planta, 24: 266-275.
Yamada, Y., and Saito, E., 1938, On some culture
experiments with the swarmers of certain
species belonging to the Ulvaceae, Sci. Pap.
Inst. Alg. Research Hokkaido Imp. Univ.^
2(1): 35-51.
DISCUSSION
j.v. SANTOS: Has the method been used to examine the
chromosome figures in the algae?
314
R.F. SCAGEL: There are numerous problems in the stages
of division, since some cells are multinucleate and the
VOLUMH 4 BOTANY
nuclei small. Some nuclear divisions have been seen with G.K PAPLNFLISS: Columbia is lucky in having two algo-
chromosome figures, other without. In addition there logists and a cytologist interested in algae. Valuable work
was a lack of well-developed nuclear membrane. should result from this team.
315
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
THE 1956 TYPHOON SEASON ON OKINAWAt
JACK C. ELLIOTT
Natural Science Consultant, Michigan State University's Mission to the University of the Ryukyus, Naha, Okinawa.
(Abstract)
Between 31 July and 25 September, 1956, five
typhoons visited the Ryukyu Islands. Four of
these (Table 1) hit Okinawa, the largest island of
the archipelago.
Although damage to man-made objects was
considerable, time will not permit an accounting,
excepting as damage had effect on vegetation, i.e.,
destruction of sea-walls permitting flooded paddy
lands.
Crops were considerably retarded m their
maturing (sugar cane, tea, banana) or the normal
growing season was a loss (sweet potato and other
vine crops, papaya).
Windbreaks of Australian Pine (Casucirina
cquisctifolia), Pandanus, Hibiscus (Hibiscus rosa-
sincnsis), Garcinia (Garcinia spicata) withstood
the typhoons and salt-spray with a varying degree
of success. Most resistant was Garcinia. Pan-
danus withstood the ussualt well, however, by the
last typhoon much of its effectiveness had been
lost because of wind and salt spray damage. Both
Australian Pine and Hibiscus suffered intensely,
being severely burned and broken with the first,
Wanda. Australian Pine, an introduced species,
was much used. However, because the majority
of them, which were approximately six years old,
were broken-off in the top half or third, some
other materials will have to be used. Garcinia
is best, but slow growing. Perhaps a combination
of Pandanus and Garcinia may be workable.
Sprout forests in northern Okinawa were
affected by both wind damage and salt burn. The
co-dominant trees of the area, Castanopsis cus-
pidata and Schima liukiuensis did not produce a
seed crop because of salt burning. Ryukyuan
Pine, (Pinus luchuemis) was badly burned. Life
cycles, especially those in woody vegetation,
were completely upset. Many were the species
which started anew after each typhoon, never to
realize their destiny.
Following a typhoon season such as that of
1956 all Okinawa was brown, bleak and in some
cases barren. It was not until seven months later,
April 1957, that again the countryside took on
the healthy green look which characterizes the
island when no typhoons have rent it so.
Table 1.
1 yphoon
Name
Wanda
Babs
Diana
Emma
Harriet
Period Typhoon in Area of circle
Approx. 240 miles radius with center at Naha
Entered
Date
July 31
Aug. 14
Sept. 1
Sept. 7
Sept. 25
Time
0200
1000
1100
1200
2400
Departed
Time
Aug. ;
Aug. 16 |
Sept. 9 I
Sept. 26 i
1900
1400
1100
1800
Wind Velocity
Average
: 73
64
35
97
i 87
dedat
Approx. Total
i ha Precipitation
at
Naha dur-
Maxi-
ing Typhoon
period
mum
(Gusts)
111
"
8.0
94
7.4
52
0.32
165
15.2
129 |
3.96
t Presented by R.F. Carlson.
316
VOLUME 4
DISCUSSION
BOTANY
Egbert W. Walker and P.R. Wycherley referred to
typhoons as one of the limiting factors in the Ryukyus
vegetation, due in part to the damaging wind, but perhaps
more to the injury from salt spray, especially in dry ty-
phoons. More extensive investigations are needed to
determine the distribution of the salt spray in respect to
altitude and windbreaks. There is room for more inves-
tigation of windbreaks, especially as to their materials,
construction and penetrability as well as the exact nature
of damage in their lee due to special swirling action from
non-penetrable windbreaks. The reported highly effective
use of Casuarina eqniseti folia in windbreaks seems as yet
not to have been fully reali/ed and needs further study.
F.R. Fosberg referred to observations elsewhere on
breakage in this species from wind. It is noticeable that
trees in areas not affected by typhoons have a very dif-
ferent growth form. There is need of further observations
on plants whose blooming seems dependent on or stimu-
lated by typhoons.
317
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
THE EVOLUTION OF THE GERMINATIVE POWER OF RICE~SEEDS~
TRAN-NGOC-TIENG
Department of Botany, Faculty of Sciences, University of Saigon, Saigon, Vietnam.
Although rice seeds appear to be mature at time
of harvest, when left to germinate, even in best
conditions, only a few of them can grow imme-
diately, while others fail to do. The former are
physiologically mature, the latter have to undergo
a post-maturation after which they become
mature. Even so, they do not remain always
viable, their longevity varies with the ways of
conservation, age, humidity, temperature.
Jn South Vietnam, rice seeds kept without any
precaution, lose all their germinative power in
a year.
In this paper, we study the Evolution of the
germinative power as related with time, so as the
influence of ways of conservation, humidity,
temperature, amylase activity of seeds on it.
We used two local, purified varieties of Rice:
Nang-Phet-Muon and Nang-Quat respectively
harvested at the beginning of the dry and rainy
seasons.
Experiments were conducted at once after the
harvest as follow:
Rice seeds were divided into four series;
The first series was left in the open-air but
sheltered from sun-light.
The second was exposed to the sun 3 hours
daily. Its mass was mixed after every exposure
and its temperature was taken. It varied between
28 and 30C.
The third scries was composed of lots of
10 g of seeds put in test-tubes sealed with
paraffin. They were conserved at different
temperatures : 28, 1 5, at alternate temperatures
of 28 - 40 U C. Each tube was used once.
The fourth series was made with a mass of
250 g of seeds kept in a dessicator with sulfuric
acid.
The germinative power, the rate of humidity,
the amylase activity of seeds were determined
every week on these lots.
Fifty seeds apparently identical were put to
soak during 24 hours, to germinate during 48
hours on Petri dishes fitted with a ring of cotton
covered by two rings of filter-paper and mois-
tened with 15 ml of water from a tap. The value
of germinative power is given by the number of
germinated seeds reported to 100.
318
The rate of humidity in seeds was calculated
on 5 g of every lot which had been dried until
constant weight (about 16 hours) in a drying-
stove of 110 -115 C. The difference between
fresh and dry weights reported to 100 expresses the
rate of humidity of each lot.
As rice seeds are rich in starch, we thought to
compare their germinative power to their amylase
activity. We used Wogelhmuth's method which
consists in determining the minimum quantity of
malt extract necessary to provoke a complete
decoloration of 5 ml of a 2% starch solution
after 30 minutes in the presence of Iodine 1/1200.
From the results we can see that the evolution
of the germinative power of rice seeds is divided
into two periods : an acquisition of maturity which
lasted 7 weeks for Nang-Phet-Muon and 10 weeks
for Nang-Quat, a second period the duration of
which was variable.
The germinative power of Nang-Phet-Muon
harvested in wet season became null after 20
weeks of conservation in the open-air in the
shadow, that of the same variety dried in the sun
was 3 weeks longer; time did not affect seeds kept
in scaled tubes or in the dessicator.
Nang-Quat harvested in dry season, left in I lie
shadow, maintained its power much longer than
Nang-Phet-Muon kept in the same way.
With regard to the rate of humidity we notice
that when it decreased from 14.50% to 11.40%
germinative power increased from 12% to 100%
for seeds in the open-air.
So, when seeds are in contact with air, the
atmosphere humidity, the rate of humidity of
seeds have an important influence on their ger-
minative power. Humidity favors the coagula-
tion of proteins, the diffusion and hydrolysis of
albumen substances. It makes the seed coat
more permeable and accelerates respiration
rhythm.
But, seeds kept in sealed tubes and in dessicator
could maintain their germinative power to a
maximum even when the rate of humidity
reached 14%. In confined atmosphere, gazeous
exchanges are quickly limited by the accumula-
tion of carbon dioxide and the rarefaction of
oxygen. In the dessicator, we must add the
VOLUME 4
BOTANY
effect of dessication which makes the seed coat
more impermeable and stop respiration.
Experiments on the influence of temperatures
were begun on Nang-Phet-Muon at once after
harvest when its germinative power was to a
minimum, and on Nang-Quat 10 weeks after
harvest when its germinative power was to a
maximum.
Results show that alternatives temperatures had
a retarding effect on Nang-Phct-Muon and a
destructive one on Nang-Quat. The responsible
temperature is the higher because seeds kept at
28 C had a normal evolution. It is supposed to
strengthen the respiration rhythm and enzyme
destruction.
Low temperatures occasion a cooling on the
acquisition of seeds maturity as well as on
enzyme activity. But the raising of Nang-Quat
power to 100% and the slow evolution of Nang-
Phet-Muon power to a maximum show that
low temperature have the same effect then
vernalisation, bringing insoluble glucides and
nitrogen compounds to soluble and useful ones.
Seeds possessed two amylases : the a amylase
or amylase of activity appearing when germina-
tion occurs and the /? amylase or amylase of
dormancy present in seeds at rest.
Results show that amylases are destroyed by
age and by cold. Seeds of Nang-Phet-Muon
harvested in 1955 lost all their amylases activities
in 1957, amylases activity of seeds kept at 15C
was lessened too. We notice that amylases activ-
ities were closely related to germinative power,
a amylase was more abundant than /? amylase.
Seeds which has such an activity that 0.2-0.4 cc
of their malt extract could bring a complete
decoloration of starch has the highest germinative
power.
Besides those experiments we also studied the
influence of the date of harvest: None of the seeds
collected 8 days before the normal date could
germinate. Seeds harvested 15 days later had a
higher energy. Insolation, ventilation would
provide seeds on plants with favorable condition.
The effect of a long washing was determined
too. Seeds were submitted as soon as harvested
to a washing of 6 to 96 hours. They had a higher
power than those of the same age but not washed ;
the longer washing lasted the higher their energy
was.
We suppose that fresh seeds would keep sub-
stances which delay germination, substances
which would act as inhibitors or as activators
according to their concentration.
So, one variety of Rice seeds can see its ger-
minative power vary with the way of conserva-
tion, for instance Nang-Phet-Muon left in the
open-air, cannot germinate after three months,
but kept in sealed tubes, they are still alive after
sixteen months.
Once inhibitors taken away, factors like age,
humidity, light, temperature, oxygen rarefaction
and carbon dioxide accumulation, act through the
medium of biological phenomena, in disturbing
the respiration rhythm, intensifying oxydations,
degradations, olden in one word the living matter,
and thus lessen germinative power and energy.
319
PROCTEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
STIMULATION OF YIELD IN H EVE A BRASILIENSISt
P. DE JONGE
Botanical Division, Rubber Research Institute of Malaya, Kuala Lumpur, Federation of Malaya.
It is not the intention of this communication
to discuss the different theories which have been
brought forward in an attempt to explain the high
increase in yields obtained after application of
a yield stimulant. Much work has been done
and still more will have to be done to explain this
phenomenon. Here we limit ourselves to dis-
cussing results obtained in field experiments
carried out at the Experiment Station of the
Rubber Research Institute of Malaya, and on
cooperating estates.
LONG TERM EXPERIMENTS
I. (a) Single cut 100% intensity tapping systems
Our experiments on trees initially 18 years
old have shown that periodical applications of
a yield stimulant can be continued over a period
of five years without inducing any signs of de-
terioration of the trees. However, young trees
should not be treated with a yield stimulant.
In an experiment carried out on clonal seedlings
six years old, girthing was 30% less after appli-
cation of a stimulant containing 2,4,5-1. In-
creased yield was obtained at the expense of
growth. After about the 15th year of life, by
which time the growth curve has flattened out,
carbohydrate in excess of that required for growth
can be mobilised for rubber production. On
such trees we have not been able to detect any
retardation in growth following application of
a yield stimulant.
In an experiment carried out on clonal seedlings
which were 18 years old at the time of the first
application, the stimulant was applied at intervals
of six months to a three inch strip of scraped
bark below the cut. The mean yields shown in
table 1 were obtained in nine half-year periods,
the tapping system being alternate-daily half
spiral (S/2.d/2.100%) throughout, the figures
adjusted for pre-treatment differences. It is clear
that the response to successive applications on
a given tapping panel falls off progressively while
continuous alternate-daily tapping is done. The
falling oiT in response is not observed on budded
trees.
t Presented by P.R. Wycherley.
320
Table 1 .
The effect of stimulant on the yield of clonal
seedling trees.
Control
Treated with 2,4,5-T
g/trcc/
g/trce/
as % of
tapping
tapping
control
1st 6 months
27.4
48.3
176%
2nd
25.7
44.6
174%
3rd
30.6
42.3
138%
4th
31.4
42.5
135%
5th
34.1
41.7
122%
6th
35.9
41.5
116%
7th
42.5
46.2 109%
8th
30.5
42.0 1 138'> *
9th
33.7
41.4 123%
* New panel in hark of second renewal.
In an experiment carried out on buddings of
nine clones, planted in 1929, highly satisfactory
yield increases were still obtained after four years
of continuous half-yearly treatments (table 2).
Throughout this experiment tapping was once
again alternate-daily on a half-spiral cut on bark
of first renewal.
A further experiment has been set up to inves-
tigate the optimum frequency of application. This
experiment is being conducted on clonal seedling
trees planted in 1930 and tapped on renewed
bark, to determine at what intervals stimulant
should be applied on trees tapped alternate-daily
on a half-spiral cut. The results covering the first
experimental year of tapping are summarised in
table 3, in which the figures have been adjusted
for pre-treatment differences. We have already
completed another 10 months of tapping, but the
data can only be validly compared over yearly
periods. The figures show that, during the first
year, treatment 4 has produced the highest yield;
but the extra yield obtained is not large when
compared with treatment 6. Treatment 5 would
have yielded less than treatment 6 if no accidental
scraping had been done.
This incomplete experiment suggests that half-
yearly applications may prove more profitable
in the long run than more frequent applications.
VOLUME 4
BOTANY
Table 2.
The effect of stimulant on the yield of budded
trees.
Control
Treated Trees
Yield in
g/tree/
tapping
g/tree/
tapping
as % of
control
32.6
35.2
108%
32.2
46.7
148%
31.4
43.6
139%
26.8
35.4
132%
27.3
40.3
148%
30.5
43.1
141%
33.7
49.1
146%
30.7 40.6
132%
Pretreatment
1st 6 months
2nd
3rd
4th
5th
6th
7th
I. (b) Double cut 200% intensity tapping systems
Our experiments on the tapping of high cuts
(ladder tapping) of unselected seedling trees and
budded trees, combined with the use of yield
stimulants and later intensive tapping, have
been very successful. The recommendations
based on these experiments are now applied on
a large scale all over Malaya. Indeed, the high
yields obtained from old unselected seedling
trees tapped at 200% intensity (often exceeding
1 ,000 Ib/acre) has led to the replanting programme
being postponed on several estates.
The results of an experiment carried out on
unselected seedling trees planted in 1929 are
summarised in table 4 as adjusted mean yields
over a period of 20 months. The experiment is
a tree plot design, in which each tree is considered
as a treatment plot. There are 50 replications,
hence fifty trees are tapped on each of the systems
listed in table 4. The treatments are randomised
over the field. A result of this is that the control
trees yield more than usual, it is believed because
small amounts of the stimulant are transferred
on the tapping knife to trees not treated. The
effect of a yield stimulant is therefore, if anything,
better than is shown by an experiment of this
design.
The trees are tapped alternate-daily on a high
V/2 cut on virgin bark and a low S/2 cut on bark
of second renewal 2C/2.d/2.200% both on the
same day. Yields of the high and low cuts have
been recorded separately by coagulation of the
latex in the cup on each tapping day. The fol-
lowing conclusions can be drawn from table 4.
1 . Trees tapped S/2.d/2. 1 00 % in bark of second
renewal show a significant but small response to
the yield stimulant. If it is applied below the
tapping cut, a high increase is noted during the
first month but the yield sometimes drops below
control level during later months.
2. Ladder tapping V/2.d/2.100% results in
slightly lower yield than low cut tapping. This
is partly caused by the shorter length of the high
cut on seedling trees, partly by the high cut
moving towards the junction of virgin bark and
bark of second renewal where yields are always
low. The yield on high cuts shows a continuous
downwards trend. Applying a stimulant below
a high cut gives a clearly better response than
below a low cut. We therefore consider that old
unselected seedling trees with poor renewed bark
should be tapped on high cuts and that a stimu-
lant should be applied at half-yearly intervals
as an ordinary estate practice.
3. The yield of the high and low cuts of un-
stimulated trees tapped at 200% intensity (treat-
ment 7) is practically identical with the yield
Table 3.
The effects of frequency of application of stim-
ulant treatment.
Treatment
1. Control (S/2.d/2. 100%)
2. as 1. plus monthly sti-
mulation above\
the cut
3. as 1. plus bi-monthly sti-
mulation above
the cut
4. as 1. plus stimulation be-
low the cut at
3 months in-
tervals
5. as 1. plus stimulation be-
low the cut at
4 months in-
tervals
6. as 1 . plus stimulation be-
low the cut at
6 months in-
tervals
7. as 1 . plus stimulation be-
low the cut at
12 months in-
tervals _ 86.9* I_J26%
* All trees belonging to these treatments were accidentally scraped
when treatments 3 and 6 were due for application. This has resulted
in a temporary highly increased yield level.
321
Mean
yield in
Ib/acre/
month
as %of
control
69^2
~ioo%
96.2
139%
94.2
136%
117.7
170%
105.1*
152%
105.7
153%
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
of trees tapped on a high cut only (treatment 4)
and of trees tapped on a low cut only (treatment
1). There is no interference between cuts on
these low yielding trees if they are kept 4 feet or
more apart. A further considerable boost in
yield can be obtained when tapping at 200%
intensity by applying a stimulant and, as might
be expected, the highest yields are recorded when
both cuts are treated. Even under these treat-
ments, no cases of brown bast have yet been
recorded. A stimulant should not be applied
above the low cut on trees tapped on two cuts
(treatment 11) for this greatly reduces the yield
of the high cut.
The 200% double cut tapping system requires
double the number of tappers employed on nor-
mal tapping. If labour or tapping costs are the
limiting factors, periodic double cut systems
with stimulant application should be considered,
with tapping to be done during the periods of
flush yields after stimulant application.
Another experiment on intensive tapping is
being conducted on the site of an old clone trial.
Nine clones are being compared, in four replica-
tions. Before the experiment started the trees,
which had been planted in 1929, were tapped on
bark of second renewal. The figures given in table
5 show that high level tapping of budded trees
results in a greatly increased yield which, how-
ever, falls off as the cut moves towards renewed
bark. There are large clonal differences. Those
clones which were considered high yielding before
the war have generally responded outstandingly
to high level tapping, whereas the moderate and
low yielding clones yielded only slightly more
on high cuts, as is shown in table 6. The pro-*
gressive fall in yield on high cut tapping is pro-
nounced for the high yielders, but in both classes
the high cuts yielded less than the low cuts after
two years of tapping.
An inter-action between cuts on trees tapped
at 200% intensity, depressing the combined yield,
is suggested more strongly for the high than for
the low yielding class, as might have been
expected, but in neither group has there been an
increase in brown bast incidence. Though
it might reduce the inter-action to tap the cuts
on different days, this is difficult in practice
because tasks tapped on high cuts are only about
70% as large as those tapped on low cuts.
On this budded material, the worst of which
still yields appreciably more than the unselected
seedling trees discussed earlier, it might be desired
to separate the tapping cuts by more than four
feet, but this is not feasible.
The experiment has not been sub-divided, in
part the tapping treatments being combined with
application of a stimulant, but the results are
not complete.
Table 4.
The eflect of stimulant treatment and tapping systems on old unselected seedlings.
Tapping system
1. S/2.d/2.100% in bark of second renewal
2. as 1. plus half yearly application of stimulant below the cut
3. as 1. plus monthly application of stimulant above the cut
4. V/2.d/2.100% in virgin bark opened at 90 inches from the
ground
5. as 4. plus half yearly application of stimulant below the cut
6. as 4. plus monthly application of stimulant above the cut
7. 2C/2.d/2.200% one high V/2. cut in virgin bark
one low S/2. cut in renewed bark
8. as 7. plus half yearly application of stimulant below high cut
9. as 7. plus monthly application of stimulant above high cut
10. as 7. plus half yearly application of stimulant below low cut
11. as 7. plus monthly application of stimulant above low cut
12. as 7. plus half yearly application of stimulant below both cuts
13. as 7. plus monthly application of stimulant above both cuts
14. as 7. plus application of a stimulant to alternating high and
low cuts at three months intervals.
Yield in Ib/acre/month
%of
control
46
100%
55
117%
55
117%
40
87%
63
137%
51
111%
Low cut
High cut
Total
44
41
85
185%
44
54
98
213%
46
55
101
219%
57
40
97
211%
47
32
79
172%
54
61
115
250%
49
49
98
213%
54
54
108
235%
VOLUME 4
BOTANY
Table 5.
The effects of tapping systems on older clones.
Tapping
system
Mean yield in Ib/month/acre
1st 6
months
2nd 6
months
3rd 6
months
4th 6
months
82
85
150
5th 6
months
70
85
131
Trees tapped
on high cut
(V/2.d/2.
100%)
Trees tapped
on low cut
(S/2.d/2.
100%)
105
62
'
95
70
90
80
Trees tapped
(2C/2.d/2.
200%)
155
150
147
Table 6.
The effect of tapping of high and low yielding
clones.
"Low and Moderate" Yielders
Clones AVROS 152, AVROS 49,
AVROS 50, PB 23, and BD 5.
Tapping
system
Mean yie
1st 6 2nd 6
months months
d in Ib/a
3rd 6
months
cre/mon
4th 6
months
th
5th 6
months
Trees tapped
1
on high cut
(V/2.d/2.
100%)
60 71
76
70
57
Trees tapped
on low cut
(S/2.d/2.
100%) , 58 63
72
78
77
Trees tapped i
2C/2.d/2. !
200%) 117 129 129 134 121
"High" Yielders
Clones PB 186, Tjir 1, Pil A 44, and Pil B 84
Trees tapped
on high cuts
(V/2.d/2.
100%)^
Trees tapped
on low cuts
(S/2.d/2.
100%)
Trees tapped
(2C/2.d/2.
200%)
153
68
203
125
79
173
109
89
169
96
94
170
86
95
143
The major conclusion from these experiments
with double cut tapping systems is that fully
mature rubber trees can stand up to more in-
tensive tapping than has been practised for a long
period in the past. A more intensive tapping
policy is now practised on a large scale in Malaya
in areas due for replanting in the foreseeable
future, and the results obtained are generally
highly satisfactory.
SHORT TERM EXPERIMENTS
II. (a) Optimum concentration of 2,4,5-T in a
stimulant
An experiment has been conducted on low
yielding unselected seedling trees tapped on
bark of second renewal to find the optimum
concentration of 2,4, 5-T (normal butyl ester)
to be used in a palm oil/petrolatum carrier.
Knowing from previous experience that a 1 %
concentration was about right, we tried the fol-
lowing concentrations: 0.5%, 1%, 1.5%, and 2%
acid equivalent of 2,4, 5-T. The results of this
experiment, covering a period of 16 months,
are given in table 7. The figures confirm the
soundness of the present composition of the
commercial 2,4,5-T yield stimulant, which is
made up according to our prescription and con-
tains 1 % acid equivalent of 2,4,5-T. Not only
did the 2% concentration result in severe burring
of the bark, but it did not induce as high a peak
yield as the 1 and 1.5% concentrations.
II. (b) Upward versus downward tapping on
seedling trees
Upward tapping has been carried out in our
experiments during the final year before replant-
ing and has proved successful when combined
with application of a yield stimulant. Its major
disadvantage is that it leads to high bark con-
sumption (a) because thicker shavings are taken
and (b) because it is necessary to leave a one-
inch strip of bark untapped at intervals of half
a year, to guide the latex into the channel.
Upward tapping has been compared with
downward tapping on low yielding unselected
seedling trees planted in 1930. For downward
Table 7.
The effects of different concentrations of 2,4,5-T.
Mean
Yield
in grams/
tree/
tapping
as % of
control
Con- | 0.5% 1 1% i 1.5%
trol ! 2,4,5-T : 2,4,5-T , 2,4,5-T
2%
2,4,5-T
33
44
53
56
100% 1 133% | 180% ( 170%
50
151%
323
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
tapping a high V/2 cut was opened in the virgin
bark at 90 inches from ground level. For upward
tapping a V/2 cut was opened in the virgin bark
with the lowest point one inch above the renewed
bark. Tapping was done alternate-daily. The
results summarised in table 8 show a slight
initial advantage for upward tapping without
stimulant, when compared with downward tap-
ping especially during the first two months after
opening. The difference may prove to be more
pronounced on higher yielding material. The
advantage of upward tapping becomes clearly
apparent when it is combined with application
of a stimulant. However, we do not recommend
this method for practical application except
during the final year of high intensity tapping,
because of the high bark consumption, which
was 39 inches for upward tapping against 27
inches for downward tapping during a period
of 23 months.
Table 8.
The effects of stimulant treatment and upward
tapping.
Mean yield in Ib/acre/month
Tapping system
1. V/2.d/2.100%-
downward
2. as 1 . plus half year-
ly stimulant appli-
cation
3. V/2.d/2.100% up-
ward
4. as 3. plus half year-
ly stimulant appli-
cation
1st
6 months
2nd
6 months
3rd
6 months
37
36
37
50
53
50
40
39
33
67
68
55
324
VOLUME 4 BOTANY
FLOWERING TREES THAT FAIL TO FLOWER IN ALIEN LANDS
EDWIN A. MENNINGER
Stewart, Florida, U.S.A.
Plantsmen wanting their plants to bloom is
scarcely a new aspiration; trying to force them to
bloom is a process almost as old. It is significant
that to the present day, almost the only success
attendant upon this struggle has been achieved
by physical means; chemical procedures have
scored practically a complete failure.
The author has been introducing to South
Florida (where temperatures remain above the
freezing point) all the showy flowered trees of the
tropics that have been found available, either
seeds or plants, and over a period of 25 years he
has established there more than 1,000 species,
constituting probably the largest collection of
"flowering" trees in the world. With plenty
of examples to draw from, he is confronted with
the fact that a good many of the most beautiful
flowering trees in the world do not want to flower
when they get moved to this new environment.
Growers have observed that many plants will
not flower at all, or will flower only after consi-
derable delay, unless certain environmental con-
ditions are met which simulate conditions that
prevailed where the plants previously or normally
grew. The two Temperate Zone conditions to
which most study have been given are the period
of low temperature storage required between
growth periods (peony roots, carrots, gladiolus
bulbs, etc.) and the photoperiod of length of day
(chrysanthemums). These conditions have been
studied chiefly in connection with non-woody
plants. Similar environmental conditions are
the basis of extensive research done by fruit trees
physiologists to discover nitrogen-carbohydrate
relationships preceding flowering and fruiting.
They have learned much. But aside from physical
control measures that range from barking to
root pruning, they have not come up with any
formula to force bloom that is sure fire even for
a single species. Nobody, in the history of
research in this field, has ever succeeded in making
a plant come into bloom by chemical means,
i.e., spraying it with a "sex-life-awakener" or
adding this to the soil, except in one isolated
instance. The pineapple is the only plant ever
to be brought into flower as the result of chemical
treatment.
Today the plant world is being invaded by new
chemical compounds that may change this pic-
ture. Chief of these at the moment are gibberel-
lic acid and some of its derivatives, and triiodo-
benzoic acid (called TIBA in the trade). The uses
of these, as well as the several physical methods
of inducing bloom will be discussed briefly, but
first the author cites some outstanding showy-
flowered tropical trees that are being grown suc-
cessfully in Florida but which so far are worth-
less as ornamentals because they wilt not or do
not flower normally. No research work has been
done in the United States to determine the reasons
(environmental or others) that these and other
tropical transplants do not bloom, or whether
these conditions could be corrected. To remove
ancestry and geography as influencing factors,
six trees are cited belonging to different natural
orders and coming from different countries.
Jacaranda mimosacfolia (Bignoniaceae). This
beautiful species is widely planted as a street
tree in southern California, blooms for nursery-
men there when still 3 feet high in a can, and in
the belt across central Florida from Tampa to
Daytona Beach it blooms at an early age, often
2 years. However, south of this belt, the trees
ordinarily will not bloom at all until they are 25
years old or more. I have a 40-foot tree in my
yard that is 27 years old and bloomed last year
for the first time. This hesitancy to bloom is
being overcome in south Florida by grafting 40-
year-old wood onto seedlings and these bloom
the second or third year. But this is an expedient
rather than a solution. What factor is it that
awakens the sex instinct of the Jacaranda tree
and makes it want to bloom? There are excep-
tions. One of my friends has a tree that bloomed
beautifully when it was 4 years old but has failed
to throw a single blossom the succeeding two
years. In my nursery I have a hundred trees and
they do not bloom. I am familiar with the
theory that, like peaches, the Jacaranda requires
a certain number of cold nights to ripen the
flowering buds (in contradistinction to the leaf
buds) and that it gets sufficient cold for this
purpose in Central Florida but does not in South
Florida. 1 doubt the validity of this theory; at
least I have seen no evidence of any connection
between unseasonably cold winters in south
Florida and the vigor of the Jacaranda trees they
still do not bloom. Jacaranda sagreana, a Cuban
325
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
form, bloomed when 5 feet high; now, 6 years
later, it is 18 feet high and has not bloomed again.
Contrariwise, Jacaranda chelonia from Argentina,
started blooming when 6 years old and has
flowered vigorously every year.
Phyllocarpus septentrional is (Caesalpiniaceae),
a Guatemalan was introduced by Dr. Wilson
Popenoe because in bloom it is such a spectacular
affair that he hoped to brighten Florida's land-
scape with it. That was 50 years ago almost.
Trees were planted all over Florida, but although
I am familiar with the location of at least 50 of
them, only 3 or 4 have ever bloomed. Last year
in St. Petersburg I was called on to identify
a huge tree spreading over a filling station on a
busy corner. It was covered with red flowers,
it had a spread of 50 feet or more, and undoubted-
ly was one of the trees Popenoe sent in so long ago.
"Nobody knows what it is," 1 was told by the
park superintendent. Not much wonder, for
it was the first time the tree had bloomed, and none
of the present generation had ever seen anything
like it. Two years ago I was giving an illustrated
lecture on flowering trees to perhaps a hundred
persons at a home in Coconut Grove, and I made
the mistake of showing a kodachrome of the
fiery flowers of Phyllocarpus. The slightly alco-
holic but intensely interested host demanded
that I stop the lecture; he turned on the flood
lights in the garden and insisted that the entire
audience troop out on the grass and look at his
50-foot Phyllocarpus that had never bloomed.
Now by contrast, consider another leguminous
tree that can hardly wait to emerge from the
pot before it starts to bloom Chamacfistula an-
tillanwn. Dr. Britton's book on Puerto Rico
says this is a vine, but in Florida it is a handsome
15-foot tree that blooms wildly from December
into April. It is cauliflorous all over the trunk
and main branches as well as producing axillary
and terminal clusters among the leaves. It
just insists on blooming, and it starts when 6
inches high! I call attention to the fact that it
blooms all through our coldest season.
Metrosideros tomentosa (Myrtaceae). The New
Zealand Christmas tree does exceedingly well in
California around Los Angeles, is highly resistant
to salt spray and cold wet winds, and there are
fine specimens to 35 feet high. In Florida the
tree is sparingly grown from Homestead as far
north as Winter Haven in the center of the
state, but to the best of my knowledge it has never
bloomed in Florida. I doubt that temperature
has anything to do with flowering inclinations,
as the mercury drops to 25 in Winter Haven
326
and Homestead on occasion. I doubt if high
humidity is the answer, because we have 5 times
the rainfall they have in Los Angeles and it is
reasonably well distributed through the year.
Occasionally we have prolonged dry spells but we
also have month-long wet spells that would
awaken the sexual activity if only water were
needed. The plants grow well enough in Florida,
although mortality without obvious reason is
above the average. In contrast, there are a good
many Myrtaceous plants that grow and bloom
enthusiastically in South Florida, like the Callis-
temon and Melaleuca, a score of Eugenia, etc.
Triplaris americana (Polygonaceae). This is
one of 4 species in cultivation in South Florida,
and the female tree is one of the tropics' breath-
taking sights when in bloom. Unfortunately
Florida is not sharing the enjoyment of this
spectacle because, although there are thousands
of these trees on the Florida landscape, I have
yet to see one of the female trees in bloom, and
I have been looking a good many years. Contrast
with this the common seagrape (Coccoloba
uvifcra) in the same group, which is native on
our shores, blooms and fruits plentifully, and is
highly ornamental even if the bloom is not showy.
Millet tia (Papilionaceac). There are 8 species
of this tree in my garden, one from India (M.
ovalifolia) which blooms beautifully every March,
and 7 from Africa, not one of which has ever
bloomed though several of them are old enough
and large enough. They grow well, their foliage
is handsome, but there are no flowers.
Stenocarpus sinuatus (Proteaceae). Grows
splendidly through Florida but does not bloom.
In its native Australia blooming is usual at 8 years
but is sometimes delayed until 1 5 years old, which
is bad enough. Here in Florida, there are a good
many specimens, but the only one in the State
that blooms is at Lake Placid. Contrast this
with Grevillea banksi foster! , of the same Natural
Order, which blooms and fruits most precocious-
ly all over Florida and I have had it in flower in
a gallon can when 8 inches high from seed.
The dividing line between juvenility in trees
and the period in which they have achieved matu-
rity and are ready for flowering and fruiting, is
ordinarily not well marked. But in the unique
flora of Australia and New Zealand, many trees
have this change period or phase in their growth
exceedingly well indicated, especially in their
leaf structure.
Many Eucalyptus trees have leaves in the
juvenile stage so entirely different in size, shape
VOLUME 4
BOTANY
and arrangement from the mature tree of the
same species, that many observers have fallen
into the trap of believing they are different trees.
Most New Zealand trees are so markedly different
in the juvenile and mature stages as to seem like
different plants. Tt is conceivable that this divid-
ing line between youth and middle age is equally
as important in other trees, though not so apparent,
and inability to cross that line easily in a new
environment might be an explanation for absence
of flowering. Is something more than age
required to push the tree across the line of
maturity ?
There are other avenues that should be ex-
plored. Several experimenters with the tran-
splanting of the Proteas an exceedingly difficult
operation in most instances have found a meas-
ure of success by transferring soil cultures from
the old location to the new, along with the plant.
The presumption is that certain microorganisms
complement the plant, perhaps are required to
make available the food supply in a form the
plant can use, perhaps have other functions
peculiar to the species. The same reasoning
might be applied to the non-flowering of certain
trees in an alien environment.
It is conceivable that a change in the degree
of acidity from that enjoyed in the original
location, might accelerate the growth of leaf
buds at the expense of flowering tissue and create
an imbalance that might even develop into an
antipathy for flowering. The peculiarities of the
problems involved might even pose the question
whether sex is a physical or a metaphysical de-
monstration.
Dr. William J. Robbins, director of the New
York Botanical Garden, has written me in this
connection.
"The question of inducing trees to flower and
fruit is one which has received a great deal of
attention, and only rule of thumb methods have
resulted without too much real basic knowledge.
In general, conditions which favor vegetative
growth interfere with flowering. Too much
nitrogen, too much water, too much of anything
which favours the general vegetative growth of
a plant may prevent flowering. There seems to
be an antagonism between vegetative growth and
flowering, but just why is still a mystery.
On this basis various procedures have been
followed to induce flowering. Root pruning is
one. In some instances actually digging up the
tree and exposing its roots for period has been
used. Ringing a twig or a branch has been
found to be effective sometimes. Limiting
applications of nitrogenous fertilizers also works
on occasion. Your comment on Jacaranda
blooming in California in cans suggests that the
slower growth which probably occurs in the dry
air of California and in the root bound condition
in a can, explains the difference. 1 note also,
how well the trees grow in Florida. I wonder
whether they may not grow too well and if you
could find some practical method of limiting
growth you might make them flower.
For some plants day length is the controlling
factor in determining blooming. Though some
are indifferent to day length others bloom only
when the days are long and some only when
the days are short. I don't believe this is of any
importance for your problem. Strong sunlight
seems to be important in some instances in shade
those plants fail to bloom and in strong light
they do. Temperature is important also. In
general, the temperature limits for blooming are
narrower than for vegetative growth. At one time
a good deal of attention was given to the impor-
tance of the ratio between carbon and nitrogen
in the plant. It was thought that if the relation
between these two elements did not fall within
certain limits, the plant would not flower. This
idea is less popular now than it once was.
There is some evidence for a flowering 'hor-
mone' which plants must produce in sufficient
quantity before they flower, but it is still uniden-
tified. It would not surprise me if in the next
few years substances were found which, sprayed
on a plant, would induce it to flower. As yet,
however, we do not have such substances and can
only use methods of the sort 1 mentioned in the
beginning of this letter.
Is there any marked difference in soil between
the areas where the trees in which you are inter-
ested bloom and those where they do not? The
amount of minor elements like zinc, manga-
nese, boron, etc., your plants are able to absorb
might be important, especially if your soil is
calcareous. It might be worth while to see what
would happen if you sprayed your plants with
one of the leaf sprays now on the market which
furnish these minor elements."
Besides the physical measures suggested by
Dr. Robbins to induce flowering, several others
are noteworthy. Sometimes thinning a fruit
tree helps to make it bloom by letting more light
into the crown, thus increasing the production
of carbohydrates. Bending the branches to a
horizontal position often helps. Both these
procedures have the same effect as girdling as
suggested by Dr. Robbins. Many persons have
327
PROCEEDINGS OF THE NINTH PACIFIC SCIENCE CONGRESS
tried spraying trees with growth-promoting
compounds, but there is no evidence that this
has induced flowering.
Perhaps the most interesting physical procedure
is that suggested by Dr. Karl Sax, professor of
botany at Harvard University. In a letter to me,
Dr. Sax briefly explained his procedure to induce
flowering, thus:
"We do it by inverting a ring of bark on the
trunk or branch of a tree. We do this in mid to
late June and always get flowers the next year
with apple varieties which have been grafted, but
not with seedling trees.
The technique is tricky, the bark must fit
closely when inverted, it must be bound tightly
with a wide rubber band, and all cut seams must
be covered to exclude air. Try it on some sur-
plus trees first."
In considering chemical compounds that are or
might be useful in inducing flowering, we are
confronted immediately with the fact that almost
no experimental work has been done to date on
woody plants, and the two chief chemicals being
tried are still too new to permit any conclusions.
Chief of these are the gibberellins, a group
of these closely related compounds derived from
a fungus disease of rice. Almost all plants treated
with any of the gibberellins, respond with
accelerated growth, and there are other effects.
The gibberellins have overcome to some extent
the two Temperate Zone environmental obstacles
to flowering heretofore cited cold storage delay
and photo-periodism. Gibberellic acid has
greatly increased certain seed yields. An experi-
mental work done at the University of California
has prompted Anton Lang of the Department of
Botany to predict: "The flower-promoting effect
of gibberellin may prove of practical value . . .
raising certain ornamental plants in such climatic
regions where they ordinarily do not give reliable
results."
The other chemical compound being tried
experimentally is TIBA (2, 3, 5-triiodobenzoic
acid) in extremely dilute concentrations. Dr.
A.C. Leopold and Dr. Frances S. Guernsey of
Purdue University, Agricultural Experiment
Station, Lafayette, Indiana, are pioneers in this
work. One of their papers says: "In view of the
fact that treatment of seeds or seedlings with
various chemical materials plus low temperatures
appears to be promotive of flowering in plants
generally, it may be justifiable to designate the
treatment with a simple name. We tentatively
propose the term chemical vernalization''' How-
ever, the authors have done almost all their work
with herbaceous species, and this is true of others
who have been experimenting with TIBA.
The foregoing discussion leads back to the
original observation that the phenomena of
flowering trees that do not flower remains an
enigma.
328
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