Interpreting Botanical Progress
July-September 1999
Editor's Note
Dennis Wm. Stevenson ......................................................187
Fossils and Ferns in the Resolution of Land Plant Phylogeny
Gar W. Rothwell...........................................................188
The Fossil Cycadales of Argentina
Analia E Artabe and Dennis Wm. Stevenson ..................................219
Implications of Fossil Conifers for the Phylogenetic
Relationships of Living Families
Charles N. Miller, Jr.....................................................239
Angiosperm Wood Evolution and the Potential Contribution
of Paleontological Data
Patrick S. Herendeen......................................................278
New Books Received............................................................300
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Editor's Note:
This issue of The Botanical Review contains papers given at a symposium called
"Practical and Theoretical Aspects of Incorporating Fossils in Analyses of Modern
Taxonomic Groups." The symposium was held in August 1995 at the Annual Meeting of the
Botanical Society of America in conjunction with The American Institute of Biological
Sciences (AIBS). This symposium, organized by William L. Crepet and Kevin C. Nixon,
of the Bailey Hortorium, Cornell University, was sponsored by the Paleobotanical
Section of the Botanical Society of America. This symposium originally had nine papers
presented. Only four manuscripts of these presentations were received and the delay
in their publication is in no way the fault of the authors of these authors.
However, suffice it to say that the four papers published here cover a great diversity
of the land flora, from ferns to flowering plants. Thus, Gar Rothwell discusses fossil
and extant ferns and the coding of salient characters for phylogenetic analyses.
Similarly, Artabe and Stevenson cover leaf and stem characters that are found in
fossil cycads from Argentina and compare them to extant cycads, none of which now
occur in Argentina. The utility and preservation of these characters is discussed in
terms of determining the relationships between fossil and extant cycad genera.
Charles Miller treats the fossil and extant conifers and demonstrates the utility of
including the fossil taxa in the same data matrix as extant taxa for elucidating
phylogenetic patterns of character distribution and generic relationships. In doing
so, he demonstrates the value of understanding fossil conifers in order to understand
the relationships among modern extant conifers. Herendeen, Wheeler, and Bass address
the topic of understanding the evolution of angiosperm wood by utilizing data from
fossil wood particularly when it can be correlated with fossil reproductive structures.
Moreover, they conclude that such correlations lead to not only a better understanding
of fossil woods but also the structural diversity within extant families.
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The Fossil Cycadales of Argentina
ANALIA E. ARTABE
Div. Paleobotánica
Fac. Cs. Nat. y Museo de La Plata
Paseo del Bosque s/n. 1900
La Plata
Argentina
DENNIS WM. STEVENSON
The New York Botanical Garden
Bronx, NY 10458, USA
Abstract
A survey of Cycadalean taxa of Argentina (including Antarctica section) is introduced.
The record of leaves represented by Nilssonia, Pseudoctenis, Ctenis, Mesodescolea,
Ticoa, Almargemia, Kurtziana and Zamia genera are summarized. Recent investigations
made of cuticles with transmission and Scanning Electron Microscopy are included. In
stems, a preliminary study of two forms found in the Upper Cretaceous of Rio Negro
Province is incorporated to the already described Michelilloa, Bororoa and Menucoa
genera. The fossil record shown some characters of leaf morphology, presence of
inverse xylem and medullary bundles, two kinds of leaf traces, and monoxylic and
polyxylic steles of systematic importance. Affinities of fossil taxa related to
extant families are suggested through the analysis of the above mentioned characters.
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Fossils and Ferns in the Resolution of Land Plant Phylogeny
GAR W. ROTHWELL
Department of Environmental and Plant Biology
Department of Biological Sciences
Ohio University
Athens, Ohio 45701
U.S.A.
I. Abstract
II. Introduction
III. Nature of the Study
IV. Methods
V. Results
A. Tree Topology
B. Decay Analysis
C. Character Changes on Representative Tree
1. Stauropterid Ferns
2. Fern Clade #2
3. Fern Clade #3, with Living Representatives
4. Ophioglossalean Ferns
5. Marattialean Ferns
6. Leptosporangiate Ferns
D. Taxon Omission Experiments
1.Hydropteris
2.Extinct Taxa
VI.Comparison to Traditional Interpretations
and to Results of Analyses by Other Authors
A. Psilotum and Tmesipteris
B. Ophioglossales and Marattiales
C. Leptosporangiate Ferns
VII. Summary
VIII. Acknowledgements
IX. Literature Cited
X. Appendix 1 - Taxa Included in the Study
XI. Appendix 2 - Characters Used in the Analyses
I. Abstract
Fifty-two taxa of living and extinct vascular plants were evaluated in an unconstrained
numerical cladistic analysis using 101 morphological characters to simultaneously
resolve the phylogenetic relationships of ferns sensu lato. Included in the analysis
were ferns assignable to the Cladoxylales, Stauropteridales, Rhacophytales,
Zygopteridales, Ophioglossales, Marattiales, Filicales and Hydropteridales, as well
as a rhyniophyte, a trimerophyte, equisetophytes, lignophytes, and the psilotophytes
Psilotum and Tmesipteris. The results placed ferns and fern-like plants in three
distinct clades, indicating that ferns s.l. are a polyphyletic grade group. Fern
clades consist of (1) extinct stauropterids, (2) extinct cladoxylaleans, rhacophytaleans
and zygopteridaleans, and (3) eusporangiates and leptosporangiates with living and
extinct species. Psilotophytes occur near the base of the tree, rather than nesting
with the Filicales as hypothesized by some. These results place Ophioglossales as
the sister group to Marattiales plus the leptosporangiates, supporting the hypothesis
that Ophioglossales represent ferns rather than progymnosperms. These analyses are
a first attempt, that includes extinct plants, to develop cladistic hypotheses for
the overall topology of fern phylogeny, and to lay the groundwork for more detailed
analyses of relationships among the homosporous leptosporangiates.
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Angiosperm Wood Evolution and the Potential Contribution of Paleontological Data
PATRICK S. HERENDEEN
Department of Biological Sciences
The George Washington University
Lisner Hall 340
2023 G Street NW
Washington DC, 20052
USA
ELISABETH A. WHEEELER
Department of Wood and Paper Science
North Carolina State University
Raleigh, North Carolina 27695
USA
PIETER BAAS
Rijksherbarium/Hortus Botanicus
P.O. Box 9514
2300 RA Leiden
The Netherlands
I. Abstract
II. Introduction
III. Comparative Wood Anatomy and the Major Trends of Xylem Evolution
IV. Wood Anatomical Patterns in the Fossil Record
V. Major Trends vs. Details of Character Evolution
VI. Testing the Baileyan Trends: Wood Anatomical Characters in Cladistic
Analyses
A. Wood Structure in Magnoliidae and "Lower" Hamamelididae
B. Implications for Character Evolution
1. Vessel Presence/Absence
2. Perforation Plate Structure
3. Intervessel Pitting
4. Imperforate Element Pit Structure
VII. The Unrealized Potential Contribution of Fossil Woods in Angiosperm
Phylogeny
VIII. Acknowledgments
IX. Literature Cited
I. Abstract
Wood anatomy is often viewed as a source of independent data that may be used to
assess evolutionary relationships among angiosperms. Comparative anatomical studies
document suites of correlated characters that have been interpreted as general
evolutionary trends, of which several have been asserted to be irreversible.
Paleobotanical data summarized by Wheeler & Baas provide broad chronological
corroboration of some wood anatomical trends, such as evolution from scalariform to
simple perforation plates and long to short vessel elements. However, the focus on
general evolutionary trends rather than on analyzing character distribution patterns
in a cladistic phylogenetic context obscures a more detailed understanding of the
evolution of wood anatomical features. Patterns of character evolution, including
the assertions of irreversibility, need to be tested through cladistic analyses.
In this paper selected wood anatomical features from families of Magnoliidae and
"lower" Hamamelididae are summarized and mapped onto previously published cladograms
as a preliminary means of testing previous hypotheses of wood evolution. The results
show that many of the characters are homoplasious, and have evolved both in accord
with, and counter to, the hypothesized general trends in different groups of flowering
plants. In general, changes that confirm generalized trends are more common than
changes that are counter to those trends. Future studies should combine wood
anatomical characters with other features as part of a cladistic analysis. Fossil
woods have not yet contributed significantly to phylogenetic studies, but in the
very few cases where they have been linked to fossil reproductive structures, the
woods have provided a better understanding of wood anatomy in early members of some
families. Data from fossil wood expand the diversity of anatomical structure known
in some angiosperm taxa and thus provide additional evidence that might be used in
phylogenetic analyses. Fossil woods have the greatest potential to affect phylogenetic
analyses where they can be linked to other fossil organs. The best chance for
establishing such a linkage is through the study of fossil charcoalified woods that
co-occur with other dispersed mesofossils.
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Implications of Fossil Conifers for the Phylogenetic Relationships
of Living Families
CHARLES N. MILLER, JR.
Division of Biological Sciences
University of Montana
Missoula, Montana 59812
USA
I. Abstract
II. Introduction
III. Methodology
IV. Results
A. The Basal Conifers: Emporiaceae, Utrechtiaceae, Majonicaceae
B. Relationship of Other Conifers with the Basal Groups
1. Basic Patterns
2. Group 1 Conifers: Cupressaceae, Sciadopityaceae, Taxodiaceae
3. Group 2 Conifers: Araucariaceae, Cephalotaxaceae, Pinaceae, Podocarpaceae
4. Higher Fossil Conifers
V. Discussion
A. Use of Ovulate Cones as a Basis for Phylogenetic Inference
B. Characters
C. Outgroup selection
D. Phylogenetic Relationships of the Basal Conifers
E. Phylogenetic Relationships of Extant Conifer Families
F. Phylogenetic Relationships of the Cheirolepidiaceae, Ullmanniaceae,
and Voltziaceae
VI. Summary and Conclusions
VII. Acknowledgments
VIII. Literature Cited
I. Abstract
Fossils have played a central role in our understanding of the evolution of conifers.
Interpretation of the seed cone as a compound strobilus and the homologies of the
ovuliferous scales of modern conifers with the axillary dwarf shoot of Pennsylvanian
forms are based on fossils. Similarly, early evolutionary trends involving the
reduction, fusion and planation of the fertile and sterile elements of the axillary
dwarf shoot, leading to structures characteristic of modern families, are documented
in Late Permian and Triassic conifers. However, a phylogeny elucidating the derivation
of modern families from fossil forms based on shared derived features has been elusive.
The present cladistic treatment using eleven characters of ovulate cones and one of
pollen grains suggests three phylogenetic groups of Late Paleozoic conifers represented
loosely by the Emporicaceae, Utrechtiaceae, and Majonicaceae of Mapes and Rothwell.
The Taxaceae appears to have diverged from ancestors within the Utrechtiaceae while
the other modern families owe their origins to the Majoniaceae. The origin of the
Taxodiaceae appears to have been biphyletic. Taxodium, Cupressus and Sciadopitys
are strongly linked to Dolmitia of the Majonicaceae, but Cryptomeria, Cunninghamia
and Araucaria are grouped together and diverge basal to the former taxa. Pinus
branches from a position basal to the known general of the Majonicaceae and all
modern families except the Taxaceae. Podocarpus also diverges basal to Majonica
but may share an anscestor with this genus, while Cepahalotaxus diverges basal to
the Dolmitia-Pseudovoltzia subclade but distal to Majonica. Similarly, the
Cheirolepidiaceae originated from basal members of the Majonicaceae and shows no
close phylogenetic relationship with any modern family. Excepting a strong linkage
between Cycadocarpidium and the Araucaria-Cunninghamia subclade, genera of the
Voltziaceae appear to have branched more or less independently from within the
Majonicaceae and show no strong affinity with modern conifers. Thus, differences
between modern conifer families are mainly due to their divergence from different
Paleozoic ancestors.
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Allen, A. 1977. Steps toward better scientific illustrations. Ed. 2. Allen Press, Lawrence,
Kansas.
Alston, R. E. 1968. The genetics of phenolic compounds. Pages 171-204 in J. B. Harborne (ed.),
Biochemistry of phenolic compounds. Academic Press, New York.
CBE Style Manual Committee. 1983. CBE style manual: A guide for authors, editors, and
publishers in the biological sciences. Ed. 5. Council of Biology Editors, Bethesda,
Maryland.
Dahlgren, R. M. T., H. T. Clifford & F. F. Yeo. 1985. The families of monocotyledons:
structure, evolution, and taxonomy. Springer-Verlag, New York.
Funk, V. A. 1982. Systematics of Montanoa Cerv. (Compositae). Mem. New York Bot. Gard. 36:1-133.
_____& D. R. Brooks. 198 1. Advances in cladistics. The New York Botanical Garden, Bronx,
New York.
Gifford, E. M. & A. S. Foster. 1988. Morphology and evolution of vascular plants. Ed. 3.
W. H. Freeman, New York.
Takhtajan, A. 1980. Outline of the classification of flowering plants (Magnoliophyta). Bot.
Rev. 46:225-359.
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The Botanical Review thanks the publishers who have provided books for this listing.
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Stuessy, Tod F., M. Ono. 1998. Evolution and Speciation of Island Plants.
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Taylor, Ronald J. 1998. Desert Wildflowers of North America. (ISBN: 0-87842-376-1 PB).
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Roth, Ingrid & A. Giménez de Bolzón. 1997. Argentine Chaco Forest, Dendrology, Tree
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