The Botanical Review 65(3)The Botanical Review 65(3)
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. Click Here to Go to Back to Top 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. Click Here to Go to Back to Top 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. Click Here to Go to Back to Top 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. Click Here to Go to Back to Top INSTRUCTIONS TO CONTRIBUTORS These guidelines are intended to increase the uniformity of manuscripts and will speed the editorial and review process. Authors are encouraged to contact the Editor (THE BOTANICAL REVIEW, the NYBG Press, The New York Botanical Garden, Bronx, NY 10458-5126, USA) whenever a contribution has some special requirement that is not discussed in the following guidelines. 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The Botanical Review thanks the publishers who have provided books for this listing.
Publishers wishing to have their books considered for inclusion in this list should 
write to The Botanical Review/New Books Received/NYBG Press, The New York Botanical 
Garden, Bronx, NY 10458-5126.
Flora of Australia-Volume 16, Elaeagnaceae, Protecaceae 1.  1995.  (ISBN: 0-643-05693-9
  Paperback].   522 pp. (CSIRO Publishing, 150 Oxford St, P.O. Box 1139, Collingwood, 
  Victoria, 3066, Australia).   Price: $64.95 Paperback.

Bernhardt, Peter. 1999.  A Natural History of Flowers.  (ISBN: 1-55963-564-9 Paperback).  
  267 pp. (Island Press, 1718 connecticut Avenue NW, Ste 300, Washington, DC  20009).   
  Price: $24.95 Paperback

Brown, James & M.V. Lomolino. 1998.  Biogeography, 2nd Edition.  (ISBN: 0-87893-073-6 
  Paperback).  560 pp. (Sinauer Associates, Inc. 23 Plumtree Road, Sunderland, MA  
  01375).   Price: $67.95 Paperback

Bryant, J.A., M.M. Burell & N.J. Kruger. 1999.  Plant Carbohydrate Biochemistry.  
  [ISBN: 1-85996-112-6 HB).  340 pp. (BIOS Scientific Publishers Ltd., 9, Newtec Place, 
  Magdalen Road, Oxford OX4 1RE, UK).   Price: £67.50 HB

Evans, L.T.  1998.  Feeding the Ten Billion Plants and Population Growth. 
  (ISBN: 0-521-64081-4 HB/ 0-521-64685-5) 247 pp. (Cambridge University Press, 40 
  W. 20th St., New York, NY 10011-4211). Price: $54.95 HB/19.99 PB. 

Hall, Franklin R. & J. J. Menn. 1998.  Biopesticides. (ISBN: 0-896-03515-8 HB) 626 
  pp. (Huymana Press, 999 Riverview Drive, Suite 208, Totowa, NJ  07512). Price: $119.50 HB. 

Hopkins, William G.  1998.  Introduction to Plant Physiology Second Edition.  
  (ISBN: 0-471-19281-3 HB)  512 pp. (John Wiley & Sons, Inc. 605 Third Avenue, 
  New York, NY 10158-0018). Price: $97.95 HB

Janick, Jules.  1999.  Horticultural Reviews. Vol. 23. (ISBN: 0-471-25445-2 HB)  
  366 pp. (John Wiley & Sons, Inc. 605 Third Avenue, New York, NY 10158-0018). 
  Price: $145.00 HB

Judd, Walter, C. S. Campbell, E. A. Kellog, & P. F. Stevens. 1999.  Plant Systematics: 
  A Phylogenetic Approach.  (ISBN: 0-87893-404-9 HB).  410 pp. (Sinauer Associates, Inc. 
  23 Plumtree Road, Sunderland, MA  01375).   Price: $67.95 HB

Lowry Judith L. 1998.  Gardening with a Wild Heart. (ISBN: 0-520-21516-8HB/ 0-520-21517-6PB)
  252 pp. (University Of California Press, 2120 Berkeley Way, Berkeley, CA  94720). 
  Price: $35.00 HB/17.95 PB. 

Slatkin, Montgomery.  1998.  Exploring Evolutionary Biology.  (ISBN: 0-87893-764-1 PB).  
  280 pp.  (Sinauer Associates, Inc. 23 Plumtree Road, Sunderland, MA  01375). Price: $24.95

Stuessy, Tod F., M. Ono.  1998.  Evolution and Speciation of Island Plants. 
  (ISBN: 0-521-49653-5 HB) 358 pp. (Cambridge University Press, 40 W. 20th St., 
  New York, NY 10011-4211). Price: $80.00 HB. 

Taylor, Ronald J.  1998.  Desert Wildflowers of North America.  (ISBN: 0-87842-376-1 PB).  
  (Mountain Press Publishing Company, P.O. Box 2399, Missoula, MT  59806). 

Tollrian, R. & C. D. Harvell.  1999.  The Ecology and Evolution of Inducible Defenses.  
  (ISBN: 0-691-01221-0 HB/0-691-00494-3 PB). (Princeton University Press, 41 William 
  Street, Princeton, NJ  08540). 232 pp.  Price: $79.50 HB/ $29.95 PB.

Roth, Ingrid & A. Giménez de Bolzón.  1997.  Argentine Chaco Forest, Dendrology, Tree 
  Structure, and Economic Use.  ENCYCLOPEDIA OF PLANT ANATOMY. (ISBN:  3-443-14024-6)  
  184pp. (E. Schweizerbart'sche Verlagsbuchhandlung (Nagele u. Obermiller) 
  Johannesstr. 3A, D-70176 Stuttgart).

Singh, Hardev.  1978.  Embryology of Gymnosperms. ENCYCLOPEDIA OF PLANT ANATOMY. 
  (ISBN:  3-443-14011-4)  302pp. (E. Schweizerbart'sche Verlagsbuchhandlung (Nagele u. 
  Obermiller) Johannesstr. 3A, D-70176 Stuttgart).  Price: $85.55 HB

Martens, P.   1971.  Les Gnétophytes.  ENCYCLOPEDIA OF PLANT ANATOMY. (ISBN:  3-443-
  14005-X)  295pp. (E. Schweizerbart'sche Verlagsbuchhandlung (Nagele u. Obermiller) 
  Johannesstr. 3A, D-70176 Stuttgart).  Price: $83.69 HB