Plant Biosystematics

676 Pages·1984·15.263 MB·English

by  William F. Grant (Eds.)

Academic Press Rapid Manuscript Reproduction
Based on the Symposium of the
International Organization of Plant Biosystematists,
McGill University, Montreal, Canada,
July 17 - 21, 1983.
Plant
Biosystematics
Editedby
William E Grant
Department of Plant Science
Macdonald College of McGill University
Ste. Anne de Belle vue, Quebec, Canada
1984
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Library of Congress Cataloging in Publication Data Canadian Cataloguing in Publication Data
Main entry under title: Main entry under title:
Plant biosystematics. Plant biosystematics
Proceedings of a symposium sponsored by the Inter- Based on a symposium sponsored by the International
national Organization of Plant Biosystematists and held Organization of Plant Biosystematists, held at McGill
at McGill University in July 1983. University, Montreal, Quebec, July 17-21, 1983.
Includes index. Includes index.
1. Botany—Classification—Congresses. I. Grant, ISBN 0-12-295680-X
William F. II. International Organization of Plant
Biosystematists. I. Botany—Classification—Congresses. I. Grant,
QK95.P545 1984 580'.12 84-6490 W.F., 1924- II. International Organization of
ISBN 0-12-295680-X (alk. paper) Plant Biosystematists.
QK95.P42 1984 580 C84-098685-8
PRINTED IN CANADA
84 85 86 87 9 8 7 6 5 4 3 2 1
Contributors
Numbers in parentheses indicate the pages on which the authors' contributions begin.
P. S. Ashton (497), The Arnold Arboretum, Harvard University, Cambridge, Mas-
sachusetts 02138, U.S.A.
Sven Asker (237), Institute of Genetics, University of Lund, S-223 62 Lund, Sweden
B. A. Barlow (117), Division of Plant Industry, C.S.I.R.O., G.P.O. Box 1600,
Canberra City, ACT 2601, Australia
R. N. Beachy (321), Department of Biology, Washington University, St. Louis,
Missouri 63130, U.S.A.
Michael D. Bennett (41), Plant Breeding Institute, Maris Lane, Trumpington,
Cambridge CB2 2LQ, England
Liv Borgen (477), Botanical Garden and Museum, University of Oslo, Trondheims-
veien 23B, Oslo 5, Norway
Céline Boutin1 (293), Département de biologie, Faculté des sciences, Université
Laval Québec G1K 7P4, Canada
y
David Bramwell (633), Jardin Botanico "Viera y Clavijo," Aparto de Correos 14,
Tafira Alta, Las Palmas de Gran Canada, Canary Islands, Spain
Willem A. Brandenburg (617), Department of Taxonomy of Cultivated Plants and
Weeds, The Agricultural University, Haagsteg 3, Wageningen NL-6708 PM,
The Netherlands
D. M. Britton (543), Department of Botany, University of Guelph, Guelph, Ontario
N1G2W1, Canada
J. Contandriopoulos (141), Laboratoire de Cytotaxinomie végétale, Université de
Provence, Centre de Saint-Charles, 3 Place Victor-Hugo, 13331 Marseille,
Cedex 3, France
K. E. Denford (359), Department of Botany, University of Alberta, Edmonton,
Alberta T6G 2E9, Canada
Present address: School of Plant Biology, University College of North Wales, Bangor,
Gwynned LL 57 2UW, U.K.
ix
x CONTRIBUTORS
John F. Doebley (587), Department of Genetics, North Carolina State University,
Raleigh, North Carolina 27650, U.S.A.
J. J. Doyle (321), Department of Biology, Washington University, St. Louis, Mis-
souri 63130, U.S.A.
C. Favarger (453), Institut de Botanique, Université de Neuchâtel, 2000 Neuchâtel,
Switzerland
I. K. Ferguson (377), The Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3DS,
England
Ichiro Fukuda (97), Division of Biology, Tokyo Woman's Christian University,
Zempukuji, Suginami, Tokyo 167, Japan
Fred R. Ganders (179), Department of Botany, University of British Columbia,
2075 Wesbrook Mall, Vancouver, British Columbia V6T 1W5, Canada
L. D. Gottlieb (343), Department of Genetics, University of California, Davis,
California 95616, U.S.A.
Hugh H. Iltis (587), Herbarium and Department of Botany, University of Wisconsin,
Madison, Wisconsin 53706, U.S.A.
R. C. Jackson (67), Department of Biology, Texas Technical University, Lubbock,
Texas 79409, U.S.A.
S. H. James (169), Department of Botany, University of Western Australia, Ned-
lands, WA 6009, Australia
Keith Jones (25), Jodrell Laboratory, The Royal Botanic Gardens, Kew, Richmond,
Surrey TW9 3DS, England
Bengt Jonsell (159), Bergius Botanic Garden, Stockholm University, Box 50017, S-
104 05 Stockholm, Sweden
Peter G. Kevan (271), Department of Environmental Biology, University of Guelph,
Guelph, Ontario NIG 2W1, Canada
Walter H. Lewis (321, 561), Department of Biology, Washington University,
St. Louis, Missouri 63130, U.S.A.
Nicole J. Martin (117), Cytogenetics Unit, Royal Brisbane Hospital, Brisbane,
Queensland 4029, Australia
J. McNeill (395), Department of Biology, University of Ottawa, Ottawa, Ontario
KIN 6N5, Canada
Pierre Morisset (293), Département de biologie, Faculté des sciences, Université
Laval, Québec, Québec G1K 7P4, Canada
David L. Mulcahy (229), Department of Botany, University of Massachusetts,
Amherst, Massachusetts 01002, U.S.A.
Kenneth M. Nagata (179), Harold L. Lyon Arboretum, Honolulu, Hawaii, U.S.A.
J. B. Phipps (417), Department of Plant Science, University of Western Ontario,
London, Ontario N6A 3K7, Canada
H. Rees (87), Department of Agricultural Botany, University College of Wales,
Penglais, Aberystwyth, Cardiganshire, Wales SY23 3DD, U.K.
CONTRIBUTORS xi
Barbara A. Schaal (439), Department of Biology, Washington University, St. Louis,
Missouri 63130, U.S.A.
John C. Semple (655), Department of Biology, University of Waterloo, Waterloo,
Ontario N2L 3G1, Canada
Ernest Small (195), Biosystematics Research Institute, Saunders Building, Central
Experimental Farm, Agriculture Canada, Ottawa, Ontario K1A 0C6, Canada
Ann Stoneburner (519), Department of Botany, University of Georgia, Athens,
Georgia 30602, U.S.A.
Krystyna M. Urbanska (211), Geobotanisches Institut ETH, Stiftung Rubel,
38 Zürichbergstrasse, C H-8044 Zürich, Switzerland
R. Verlaque (307), Laboratoire de Cytotaxinomie végétale, Centre de Saint-Charles,
3 Place Victor-Hugo, 13331 Marseille, Cedex 3, France
Robert K. Vickery, Jr. (1), Department of Biological Sciences, University of Utah,
Salt Lake City, Utah 84112, U.S.A.
Warren H. Wagner, Jr. (643), Department of Botany, University of Michigan, Ann
Arbor, Michigan 48109, U.S.A.
C. J. Webb (249), Botany Division, D.S.I.R., Private Bag, Christchurch, New
Zealand
Robert Wyatt (519), Department of Botany, University of Georgia, Athens, Georgia
30602, U.S.A.
Daniel Zohary (579), Department of Botany, The Hebrew University, Jerusalem,
Israel
Preface
Plant Biosystematics is the proceedings of a four-day symposium held at McGill
University, Montreal, in July, 1983. The symposium was sponsored by the Inter-
national Organization of Plant Biosystematists in their continuing role to foster
international cooperation in plant biosystematics. Delegates from 16 countries par-
ticipated. The theme of the symposium was "Plant Biosystematics: Forty Years
Later," in recognition of the introduction of the term "biosystematy" by Camp
and Gilly in 1943. Camp and Gilly considered "biosystematy" as a descriptive
term to cover newer approaches to "(1) delimit the natural biotic units and (2)
apply to these units a system of nomenclature adequate to the task of conveying
precise information regarding their defined limits, relationships, variability, and
dynamic structure." The types of studies they advocated were to provide a system
of classification which was to form a basis for the organization of information of
all types concerning living organisms, and ultimately to provide solutions to ques-
tions concerning biological diversity within the framework of evolution.
Until the 1960s biosystematics was viewed as the ' 'alpha and omega" of biology
and as an "unending synthesis." It was considered that the "processes of micro-
evolution and the origin of higher plant diversity" would be readily resolved and
that these would be able to be expressed "satisfactorily in our systematic arrange-
ments."
In the 1960s chemistry and statistics became increasingly important and often
very powerful analytical tools. But by 1970 there was a growing awareness among
biosystematists that they were not in a position to offer unequivocal solutions to
the quandaries faced by classical taxonomy, and that it was not possible to set up
a system of classification which reflected all types of data with equal fidelity.
Plant systematics in the 1970s matured to the point where it is generally under-
stood that taxonomy provides a general-purpose classification, incorporating and
summing up characters of many kinds, which can serve as a framework for many
types of investigation, including the biology of plant populations. Biosystematics
xiii
Xiv PREFACE
has placed more emphasis on an understanding of processes at the level of the
population, and of evolutionary forces, especially those of microevolution, and on
all aspects of the biology of extant populations and the individual organisms of
which they consist.
In the brief history of this discipline, biosystematists have used a wide variety
of techniques, in addition to those traditional to classical taxonomy, in an attempt
to resolve these issues. Two of these, cytology and hybridization, have been sig-
nificant contributors to biosystematic studies. Karyological studies have from the
beginning had a central role in biosystematics. The importance of chromosomes
has been twofold: they have been used as a diagnostic character in classification
and to provide insights into genetic phenomena and evolutionary processes leading
to speciation. Three characteristics of chromosomes have been found to have es-
pecially high information content, namely, chromosome number, chromosome mor-
phology, and chromosome behavior at meiosis. Likewise, hybridization, observed
either in wild populations, or carried out artificially, has played a central role in
biosystematic studies. It has been used both to test for reproductive barriers and to
explain the transfer of genetic information from one taxon to another. Since species
differ from each other genetically, the consequences of hybridization have also
been much linked to the biological species concept.
The program was planned to consider biosystematics in its broadest sense in
which not only traditional themes of speciation, karyology, polyploidy, hybridi-
zation, and specialized systems (apomixis, incompatibility, pollination) would be
considered, but also some of the current methodologies and future trends, as well
as some of the practical aspects (agriculture, conservation, horticulture, and medicine).
The volume has provided a review of the current field of biosystematics from
which botanists, geneticists, agriculturalists, evolutionists, and biologists in general,
who are interested in the evolution of natural biota, may obtain some perception
of the present status of plant biosystematics and how this discipline contributes to
human well-being. Of the 38 papers presented at the meeting, 37 are reproduced
herein. Those by J. Contandriopoulos, C. Favarger, P. Morisset, and R. Verlaque
were presented in French but were translated into English for publication with the
authors' consent. Although discussions were not recorded, J. Semple has sum-
marized in a final chapter some of the flavor of the meeting.
Acknowledgments
Although the symposium was organized by the Executive and Council of the
International Organization of Plant Biosystematists, many other individuals were
responsible for suggestions and help with arrangements. The active and enthusiastic
participation during the initial organizational phase of two of my former students,
C. W. Crompton and A. E. Stahevitch of the Biosystematics Research Institute,
Ottawa, was most fruitful. Walter Lewis not only contributed to the program but
delivered the post-banquet talk on * 'Recent Contributions of Plants to Medicine and
Dentistry." Hugh Iltis supplied the Camp and Gilly cartoon which decorated the
cover of the program. Luc Brouillet made all the arrangements for the tour of the
Montreal Botanical Gardens, and the reception by the City of Montreal. Pierre
Morisset translated the abstracts for the program into French. Paulette Lachance
served most efficiently as Executive Secretary and headed the registration desk.
Others who helped during the symposium included Lawrence Goldstein, Pierre St.
Marseille, John Raelson, Kate Merlin and Nancy MacLean. Finally, financial con-
tributions, which made the meeting possible, are gratefully acknowledged from
McGill University, the Genetics Society of Canada, l'Association Canadienne-
Française pour l'Avancement des Sciences, and the Natural Sciences and Engi-
neering Research Council of Canada.
William F. Grant
XV
Biosystematics 1983
Robert K. Vickery, Jr.
Department of Biological Sciences
University of Utah
Salt Lake City, Utah, U.S.A.
INTRODUCTION
1983 finds biosystematics alive, well and thriving! However, as
it has developed and grown over the last several decades, it has
faced several challenges and new opportunities· But first, let us
see what the roots of biosystematics are and how it has developed
before we assess its status, challenges and opportunities today.
Biosystematics or biosystematy as it was originally defined by
Camp and Gilly in 1943 "seeks (1) to delimit the natural biotic
units and (2) to apply to these units a system of nomenclature
adequate to the task of conveying precise information regarding
their defined limits, relationships, variability and dynamic
structure." Purists may argue that Camp and Gilly should have
used the term biotaxonomy inasmuch as their stated emphasis
falls heavily on the philosophic consideration of "what is a
species?" or, more accurately, "what are species?". In practice,
their approach consisted of using the methods of cytology and
genetics to experimentally ascertain species relationships, that
is, it was, in fact, a systematic approach, but to a taxonomic
problem. What they were doing was attempting to define and
formalize the growing emphasis on the use of experimental studies
as the foundation for a better, more biological classification.
HISTORICAL BACKGROUND
Historically, classification had been highly artificial from the
days of the "Father of Botany," Theophrastus (3rd and 4th Century
B.C.), and the celebrated physician, Dioscorides (1st Century
A.D.) down to the 16th century. Theophrastus grouped plants by
form: herbs, undershrubs, shrubs and trees (Lawrence 1951).
Trees were plants with a woody main stem, e.g., the olive, fig and
vine. Dioscorides grouped plants by use: pot herbs, sharp herbs,
PLANT BIOSYSTEMATICS Copyright « 1984 by Academic Press Canada
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ISBN 0-12-295680-X