Flowering Plant Origin, Evolution & Phylogeny

411 Pages·1996·9.69 MB·English

by  David Winship Taylor

Flowering Plant
,n~g~rO Evolution
& Phylogeny
Flowering Plant
0 rlgln, Evolution
& Phylogeny
EDITED YB
David Winship Taylor
Indiana University Southeast
& Leo .J Hickey
Yale University
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Library of Congress Cataloging-in-Publication Data
Flowering plant origin, evolution, and phylogeny / edited by David W. Taylor, Leo J. Hickey.
p. cm.
Papers from a symposium held at the American Institute of Biological Sciences
meeting at Ames, Iowa, on Aug. 4, 1993.
Includes bibliographical references and index.
ISBN 0-412-05341-1 (alk. paper)
.1 Flowering plant origin, Fossil--Congresses. 2. Angiosperms--Origin--Congresses.
3. Angiosperms--Evolution--Congresses. I. Taylor, David W., 1958-- . II. Hickey,
Leo J. III. American Institute of Biological Sciences, Meeting (I 993: Ames, Iowa).
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STNETNOC
Preface vii oo
Contributors ix
Chapter .1 Introduction: The Challenge of Flowering Plant History
David Winship Taylor and Leo J. Hickey
Chapter 2. A Survey of Reticulate Venation Among Fossil and
Living Land Plants
Mary Louise Trivett and Kathleen B. Pigg
Chapter 3. A New Gnetophyte from the Late Carnian (Late Triassic)
of Texas and its Bearing on the Origin of the Angiosperm
Carpel and Stamen 32
Bruce Cornet
Chapter 4. Wood Anatomy of Primitive Angiosperms: New
Perspectives and Syntheses 68
Sherwin Carlquist
Chapter 5. Evidence for the Earliest Stage of ~giosperm Pollen
Evolution: A Paleoequatodal Section from Israel 91
Gilbert J. Brenner
Chapter 6. The Origin and Evolution of the Angiosperm Carpel 116
David Winship Taylor and Gretchen Kirchner
Chapter .7 Floral Structure, Development, and Relationships of
Paleoherbs: Saruma, Cabomba, Lactoris, and Selected
Piperales 141
Shirley .C Tucker and Andrew .W Douglas
Chapter 8. Origin of the Angiosperm Flower 176
Leo J. Hickey and David Winship Taylor
Chapter .9 Evidence for and Implications of an Herbaceous Origin
for Angiosperms 232
David Winship Taylor and Leo Z Hickey
Chapter .01 Comparison of Alternative Hypotheses for the Origin of
the Angiosperms 267
Henry Loconte
vi Contents
Chapter .11 The Least Specialized Angiosperms 286
Robert F. Thorne
Chapter .21 Molecular Phylogenies and the Diversification of the
Angiosperms 314
Kenneth J. Sytsma and David A. Baum
References 341
Appendix. Stratigraphic Time Scale 387
Index
Preface
Questions about the origin, early evolution, and basal phylogeny of the flower-
ing plants have been areas of ferment in botanical research for nearly 25 years.
Rapid progress toward resolving these problems continues to be made, owing,
in part, to the unprecedented number of approaches now available, as well as
to the accumulation of carefully documented empirical data from both living
and fossil plants. During this time, a series of symposia and their published
volumes have periodically summarized the progress made and focused research
on many of the important issues that remain. The most influential of these
publications have been Origin and Early Evolution of Angiosperms, edited by
Charles B. Beck in 1976, Historical Perspectives of Angiosperm Evolution
edited by David L. Dilcher and William L. Crepet as Volume 71, (Issue 2) of
the Annals of the Missouri Botanical Garden in 1984, and ehT Origins of
Angiosperms and Their Biological Consequences edited by Else Marie Friis,
William G. Challoner, and Peter R. Crane in 1987. We feel that the time is
ripe for a new book that summarizes the recent research from fields of
paleobotany, comparative morphology, development, and structural and mol-
ecular phylogeny that bears on these problems.
The volume took shape at a symposium entitled ehT Origin, Early Evol-
ution, and Phylogeny of Angiosperms held at the American Institute of Bio-
logical Sciences meeting at Ames, Iowa on August 4, 1993. Our contributors
were invited because of their ability to provide the most current view of their
fields and to place this knowledge into a broad context. In an effort to bring a
greater degree of focus to the symposium and to the resulting book, we also
asked the authors to consider a set of questions relating to angiosperm evol-
ution. This book follows the same scheme as the symposium with the addition
of a geological time scale to aid those who are unfamiliar with stratigraphic
terminology, particularly at the level of the epoch and age divisions of the
geological periods. As with all efforts to summarize a rapidly developing field,
this publication is only a progress report, but one that we hope will serve to
further accelerate the process of resolving what seemed only a generation ago
to be intractable problems.
Finally, we wish to thank the following reviewers who gave generously of
their time to ensure that the contents of this volume met the standards of the
science: Pieter Bass, Robyn Burhnam, Peter R. Crane, David L. Dilcher, Jeff
H. Doyle, Peter K. Endress, Else Marie Friis, Dorian Q. Fuller, Richard C.
Keating, Sergius H. Mamay, Richard G. Olmstead, Robert Ornduff, Edward L.
oo
VII
rift Preface
Schneider, Taylor Steeves, Susan Swenson, Alfred Traverse, Thomas K. Wilson
and several anonymous reviewers. Very special thanks are due to Grady Web-
ster who acted as an associate editor for the two chapters co-authored by the
editors.
Contributors
David A. Baum Henry Loconte
Department of Organismic and Evolu- The New York Botanical Garden
tionary Biology Bronx, NY 10458-5126
Harvard University
Cambridge, MA 02138 Kathleen B. Pigg
Department of Botany
Gilbert Brenner Arizona State University
Department of Geological Sciences Tempe, AZ 85287-1601
State University of New York
College at New Paltz Kenneth J. Sytsma
New Paltz, New York 12562 Botany Department
University of Wisconsin
Sherwin Carlquist Madison, WI 53706
Santa Barbara Botanic Garden
1212 Mission Canyon Road David Winship Taylor
Santa Barbara, CA 92521 Department of Biology
Indiana University Southeast
Bruce Comet 4201 Grant Line Road
27 Tower Hill Avenue New Albany, IN 47150
Red Bank, NJ 07701
Robert .F Thome
Andrew Douglas Rancho Santa Ana Botanic Garden
Department of Botany 1500 North College Avenue
Louisiana State University Claremont, California 91711
Baton Rouge, LA 70803
Mary L. Trivett
Leo J. Hickey Department of Environmental and Plant
Department of Geology and Geophysics Biology
Yale University Ohio University
Box 6666 Athens, OH 45701-2979
New Haven, CT 06511
Shirley C. Tucker
Gretchen Kirclmer Department of Botany
Department of Biology Louisiana State University
Indiana University Southeast Baton Rouge, LA 70803
4201 Grant Line Road
New Albany, IN 47150
ix
Introduction: The
CHAPTER
1
Challenge of
Flowering Plant
History
David Winship Taylor and Leo .J Hickey
Angiosperms are well known for their incredible diversity in species number,
range of habitat, and morphology. Recent counts have shown that there are at
least 234,000 described species of flowering plants in some 437 families
(Thorne, 1992a). This number exceeds the total number of species of all other
photosynthetic land plants and algae combined (Spome, 1974; Prance, 1977).
Not only do angiosperms dominate in number of species, they are also found
in a far greater range of habitats than any other group of land plants. At their
extremes they range from the tropics to the tundra, and from freshwater and
marine environments to deserts. Not surprisingly, angiosperms also display an
unparalleled range in form, from tiny plants in the Lemnaceae to huge trees in
the Myrtaceae. Examples of their morphological variability can be found in
stems (e.g., Carlquist, Chapter 4), flowers (e.g., Tucker and Douglas, Chapter
7), carpels (e.g., Taylor and Kirchner, Chapter 6), and pollen (e.g., Brenner,
Chapter 5).
This great plasticity of the angiosperm body is the cause of contentious
debate over ancestral form (compare Taylor and Hickey, Chapter 9; Loconte,
2 .pahC 1 Introduction: The Challenge of Flowering Plant History
Chapter 10; Thorne, Chapter 11). No consensus on the ancestral states of many
morphological traits has been arrived at, especially when no single living taxon
seems to have all these states. Even determining the derived states shared by
all angiosperms has been difficult because of exceptions to nearly every char-
acter, even among angiosperms regarded as primitive. In addition, a growing
number of states that were thought to be uniquely angiospermous have now
been shown to exist in angiosperm outgroups and are, thus, symplesiomorphies
(see Taylor and Hickey, 1992, Chapter 9; Hickey and Taylor, Chapter 8).
Recognition of these shared ancestral states may help in reconstructing what the
ancestral angiosperm looked like but is of no assistance in determining whether
angiosperms are monophyletic or what the relationships are within the clade.
Nevertheless, a number of features appear to be nearly universal in angios-
perms and widespread in their putatively most basal group, the Magnoliidae.
These universal synapomorphies for angiosperms emerged both from conven-
tional and cladistical analysis of morphology (summarized in Taylor and Hick-
ey, Chapter 9). One of these synapomorphies is the existence of sieve-tube
members and companion cells. These cells are found in all angiosperms but in
no other living group. Carpels and stigmatic germination are also universal and
are another defining character. Other reproductive characters are the highly
reduced megagametophyte with three or fewer free cellular divisions and the
production of stored food for the embryo after fertilization. Several other
characters have been suggested as ancestral, yet either they are not universal in
the angiosperms or are possible symplesiomorphies because of their occurrence
in angiosperm outgroups. Some of these nonuniversal characters include vessels
with scalariform perforation plates, apocarpous carpels, wood fibers, and trip-
loid endosperm. Examples of angiospermous characters that may be
homologous with states from the outgroups include reticulately veined leaves,
ethereal oil cells, and vessels.
ANGIOSPERM ORIGIN
One aspect of phylogeny that has been important for understanding angiosperm
origin is their relationship to other seed plants. The consensus emerging from
recent phylogenetic analyses (Crane, 1985; Doyle and Donoghue, 1986, 1992;
Zimmer et al., 1989; Loconte and Stevenson, 1990; Hamby and Zimmer, 1992;
Chase et al., 1993; Rothwell and Serbet, 1994; Nixon et al., 1994) clearly shows
that the gnetopsids are the closest living relatives of the angiosperms, whereas
the closest extinct group is the bennettitaleans. Outgroup relationships provide
important clues for understanding the evolution of several distinctly angiosper-
mous organs such as leaves (Trivett and Pigg, Chapter 2), carpels (Taylor and
Kirchner, Chapter 6), and flowers (Hickey and Taylor, Chapter 8). Discovery