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PLANT SYSTEMATICS
Plant Systematics
Third Edition

Michael G. Simpson

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Notices

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Library of Congress Cataloging-in-Publication Data

Simpson, Michael G. (Michael George), 1953-
Plant systematics / Michael G. Simpson. – 3rd ed.
p. cm.
ISBN 978-0-12-812628-8
1. Plants–Classification. I. Title.
QK95.S566 2019
580.1'2–dc22
2010009204

British Library Cataloguing-in-Publication Data

A catalogue record for this book is available from the British Library.

For information on all Academic Press publications visit our

website at https://www.elsevier.com/books-and-journals

Printed in China
10 11 12 9 8 7 6 5 4 3 2
The first edition of this book was dedicated to three mentors I have been very fortunate to know: Albert Radford, who
taught critical thinking; P. Barry Tomlinson, who taught the fine art of careful observation; and Rolf Dahlgren, whose
magnetic personality was inspirational. I also wish to thank my many students who have provided useful suggestions
over the years, plus three writers who captured my interest in science and the wonder of it all: Isaac Asimov, Richard
Feynman, and Carl Sagan.

I wish to dedicate the second edition of this book to my wonderful family: Anna, Bonnie, Claire, Lee, and Lori.

The third edition is dedicated to Stella, Layla, and Quinn, who have brought so much joy into our lives.
 Contents

Preface……………………………………………………………………………………………………………………………………………………………ix

Acknowledgements…………………………………………………………………………………………………………….…………………........................xi

UNIT I SYSTEMATICS

Chapter 1 Plant Systematics: An Overview………………………………………………………………………………………...……………………………3

Chapter 2 Evolution and Diversity of Vascular Plants………………………………………………………………………………………………………….17

UNIT II EVOLUTION AND DIVERSITY OF PLANTS

Chapter 3 Evolution and Diversity of Green and Land Plants………………………………………………………………………………………...………..55

Chapter 4 Evolution and Diversity of Vascular Plants………………………………………………………………………………………...……………….75

Chapter 5 Evolution and Diversity of Woody and Seed plants………………………………………………………………………………………...……...131

Chapter 6 Evolution of Flowering Plants………………………………………………………………………..…………………………………………….167

Chapter 7 Diversity and Classification of Flowering Plants: Amborellales, Nymphaeales, Austrobaileyales, Magnoliids, Monocots, and Ceratophyllales..187

Chapter 8 Diversity and Classification of Flowering Plants: Eudicots………………………………………………………………………………………...285

UNIT III SYSTEMATIC EVIDENCE AND DESCRIPTIVE TERMINOLOGY

Chapter 9 Plant Morphology………………………………………………………………………………………...………………………………………...469

Chapter 10 Plant Anatomy and Physiology………………………………………………………………………………………...…………………………537

Chapter 11 Plant Embryology………………………………………………………………………………………...……………………………………….567

Chapter 12 Palynology…………………………………………………………………………...……………………………………………………………583

Chapter 13 Plant Reproductive Biology………………………………………………………………………………………...…………………………….595

Chapter 14 Plant Molecular Systematics………………………………………………………………………………………...……………………………607

vii
viii contents

UNIT IV RESOURCES IN PLANT SYSTEMATICS

Chapter 15 Plant Identification...…………………………………………………………………...…………………………………………………………625

Chapter 16 Plant Nomenclature………………………………………………………………………………………...……………………………………..631

Chapter 17 Plant Collecting and Documentation………………………………………………………………………………………...……………………647

Chapter 18 Herbaria and Data Information Systems………………………………………………………………………………………...………………...657

UNIT V SPECIES CONCEPTS AND CONSERVATION BIOLOGY

Chapter 19 Species and Conservation in Plant Systematics………………………………………………………………………………………...…………671

Appendix 1 Plant Description………………………………………………………………………………………...………………………………………..691

Appendix 2 Botanical Illustrations………………………………………………………………………………………...…………………………………...697

Appendix 3 Scientific Journals in Plant Systematics………………………………………………………………………………………...………………...701

Appendix 4 Statistics and Morphometrics in Plant Systematics………………………………………………………………………………………...…….703

Glossary of terms………………………………………………………………………………………...……………………………………………………..713

Index………………………………………………………………………………………...………………………………………………………………….749
 Acknowledgments
I sincerely thank Andy Bohonak, Bruce Baldwin, Lisa Campbell,
Travis Columbus, Gary Emberger, Lluvia Flores-Rentería,
Chrissen Gemmill, Matt Guilliams, Robert Hattaway, Bruce
Kirchoff, Eric Knox, Kristen Hasenstab-Lehman, Makenzie
Mabry, Lucinda McDade, Steve O’Kane, Kathleen Pryer (and
her lab group), Jon Rebman, Tanya Renner, P. van Rijckevorsel,
Paula Rudall, Dennis Stevenson, Livia Wanntorp, Annette
Winner, and several anonymous reviewers for their com-
ments on various chapters or appendices of the first, second,
or third editions of this book. I thank Anna C. Simpson and
Lee M. Simpson for technical help. I am grateful to Peter
Stevens for up-to-date information on higher level classifica-
tion of angiosperms from his excellent Angiosperm Phylogeny
Website. As always, none of these bear responsibility for any
mistakes, omissions, incongruities, misinterpretations, or
general stupidities.
Almost all of the illustrations and photographs are the
product of the author. I thank the following for additions to
these (in order of appearance in text):
The “tree” of Unit 1 opening page (at left) is from Augier, A.
1801. Essai d’une nouvelle classification des végétaux. Lyon,
Bruyset Ainé. (See Stevens, P. F. 1983. Taxon 32: 203–211.)
The cladogram (at right) was an unpublished tree from the
study by Simpson et al. 2017. Taxon 66: 1406–1420..
The Jepson Herbarium (University of California Press) gave
special permission to reproduce the key to the Oleaceae
(Thomas J. Rosatti, author) in Figure 1.7.
Madroño (California Botanical Society) gave permission
to reproduce Figure 4C of Simpson, et al. 2016. Madroño 63:
39-54, and Figure 2 of Dodero and Simpson. 2012. Madroño
59:223-229 in Figure 2.1 of Chapter 2.
Rick Bizzoco contributed the images of Chlamydomonas
reinhardtii in Figures 3.2C and 3.3A.
Linda Graham contributed the image of Coleochaete in
Figure 3.5B.
Figure 4.12A was reproduced from Kidston, R. and
W. H. Lang. 1921. Transactions of the Royal Society of
Edinburgh 52(4): 831–902.
Figure 4.20C was redrawn from Wakasugi, T., M. Sugita,
T. Tsudzuki, and M. Sugiura. 1998. Plant Molecular Biology
Reporter 16: 231–241, by permission.
xi
Figure 4.20D was reproduced from Banks et al. 1975.
Palaeontographica Americana 8: 77–126, with permission from
Paleontological Research Institution, Ithaca, New York.
John Braggins contributed the images of Figure 4.26H–J.
Figure 4.31 was redrawn from Smith, G. M. 1955.
Cryptogamic Botany, McGraw-Hill Book Company, Inc.,
New York.
Lawrence Jensen contributed the images of Figure
4.34A,B,E, 4.35D–E,H–J, 4.36C–D, and 4.37C–F.
Vera Svobodova contributed the images of 4.35A–C.
Gerald Carr contributed the image of 4.36F.
Figure 5.9 was reproduced and modified from Swamy,
B. G. L. 1948. American Journal of Botany 35: 77–88,
by permission.
Figure 5.13A,B was reproduced from: Beck, C. B. 1962.
American Journal of Botany 49: 373–382, by permission.
Figure 5.13C was reproduced from Stewart, W. N.,
and T. Delevoryas. 1956. Botanical Review 22: 45–80,
by per­mission.
Figure 5.19 was redrawn from Florin, R. 1951. Evolution in
Cordaites and Conifers. Acta Horti Bergiani 15: 285–388.
John Braggins contributed the images of Figure 5.22D and
5.24D–F,L.
Figure 5.26B was reproduced from Esau, K. 1965. Plant
Anatomy. J. Wiley and Sons, New York, by permission.
Mark Olson contributed the images of Welwitschia mirabilis
in Figure 5.27B–E.
Figure 6.5 was based upon Jack, T. 2001. Trends in Plant
Science 6: 310–316.
Figure 6.18A–C was redrawn from Thomas, H. H. 1925.
Philosophical Transactions of the Royal Society of London
213: 299–363.
Figure 6.18D–F was redrawn from Gould, R. E. and T.
Delevoryas. 1977. Alcheringa 1: 387–399, by permission.
Figure 6.19A was contributed by K. Simons and David
Dilcher (©); Figure 6.19B was contributed by David Dilcher (©)
and Ge Sun.
Stephen McCabe contributed the images of Amborella
in Figures 7.3A,C.
The Arboretum at the University of California-Santa Cruz
contributed the image of Amborella in Figure 7.3B.
xii Acknowledgments
Sandra Floyd provided the image of Amborella in
Figure 7.3D.
Jeffrey M. Osborn and Mackenzie L. Taylor contributed the
images of the Cabombaceae in Figure 7.4.
Jack Scheper contributed the image of Illicium floridanum
in Figure 7.6A.
Figure 7.18A was reproduced from Behnke, H.-D. 1972.
Botanical Review 38: 155–197, by permission.
Constance Gramlich contributed the image of Amorpho­
phallus in Figure 7.25C.
Wayne Armstrong contributed the image of a flowering
Wolffia in Figure 7.25G.
Benjamin Lowe contributed the image of Scoliopus bigelovii
(Liliaceae) in Figure 7.28H.
John Kress contributed the Zingiberales drawing in
Figure 7.54.
Will Cook contributed the images of Hamamelis virginiana
in Figure 8.13A–C and of Ulmus alata in 8.39G.
Gerald Carr contributed the images of Hillebrandia sand­
wicensis in Figure 8.42A,B and of Juglans hindsii in Figure
8.47D–H.
The Rampant Gardener contributed the image of Juglans
regia in Figure 8.47J.
Jerry Green contributed the image of Crossosoma bigelovii
in Figure 8.48A.
Reid Moran contributed the image of Crossosoma californi­
cum in Figure 8.48B–D, of Koeberlinia spinosa in Figure
8.57J–L, and of Eucnide urens in Figure 8.88B.
Mark Olson contributed the images of Moringa spp. in
Figure 8.57M–Q.
Figure 8.72B was reproduced from Behnke, H.-D. 1972.
Botanical Review 38: 155–197, by permission.
Gerald Carr contributed the images of Ardisia crenata in
Figure 8.93A,B.
Steven Swartz contributed the images of Heliamphora sp. in
Figure 8.97F,G.
Matt Guilliams contributed the image of Pholisma sonorae
in Figure 8.100F,G.
David G. Smith contributed the images of Phryma lepto­
stachya in Figure 8.116I,J.
Michael Silveira contributed the image of Linnaea borealis
in Figure 8.127C,D.
Michael Mayer contributed material of Scabiosa, photo-
graphed in Figure 8.128E–L.
On the Unit III opening page the brass microscope image is
courtesy of Allan Wissner (www.antique-microscopes.com),
and the image of an Illumina sequencer is a general use image
(https://www.illumina.com/company/news-center/multime­
dia-images.html)
Figure 9.6 was reproduced and condensed from Hallé, F. et
al. 1978. Tropical Trees and Forests: An Architectural Analysis.
Springer, Berlin., by permission.
Figure 9.13 was redrawn from Hickey, L. J. 1973. American
Journal of Botany 60: 17–33, by permission.
Darren Burton prepared several illustrations in Chapter 9.
Figure 13.4A was redrawn from Weberling. 1989. Morpho­logy
of Flowers and Inflorescences. Cambridge University Press,
Cambridge, New York, by permission.
Figure 13.4B was redrawn from Kohn et al. 1996. Evolution
50: 1454–1469, by permission.
Jon Rebman contributed the images in Figure 13.7D,E.
Figure 14.4 was redrawn from Wakasugi, T., M. Sugita,
T. Tsudzuki, and M. Sugiura. 1998. Plant Molecular Biology
Reporter 16: 231–241, by permission.
The Herbarium at the San Diego Natural History Museum
contributed the images in Figure 17.2.
Jon Rebman contributed the image of the herbarium sheet
in Figure 18.2.
Figure 19.6 was redrawn from Huang et al. 2005. Journal of
Plant Research 118: 1–11, by permission.
Figure 19.9 was redrawn from Baldwin. 2000. Madroño 47:
219–229, by permission.
Dinna Estrella contributed the stippled line drawing in
Appendix 2.
 Preface
Plant Systematics is an introduction to the morphology, evolu-
tion, and classification of land plants. My objective is to present a
foundation of the approach, methods, research goals, evidence,
and terminology of plant systematics and to summarize infor-
mation on the most recent knowledge of evolutionary relation-
ships of plants as well as practical information vital to the
field. I have tried to present the material in a condensed, clear
manner, such that the beginning student can better digest the
more important parts of the voluminous information in the
field and acquire more detailed information from the literature.
The book is meant to serve students at the college upper
undergraduate and graduate levels in plant systematics or
taxonomy courses, although portions of the book may be
used in flora courses and much of the book could be used in
courses in plant morphology, diversity, or general botany.
Each chapter has an expanded Table of Contents on the
first page, a feature my students recommended as very useful.
Numerous line drawings and color photographs are used
throughout. A key feature is that illustrated plant material is
often dissected and labeled to show important diagnostic fea-
tures. At the end of each chapter are (1) Review Questions,
which go over the chapter material; (2) Exercises, whereby a
student may apply the material; and (3) References for Further
Study, listing some of the basic and recent references.
Literature cited in the references is not exhaustive, so students
are encouraged to do literature searches on their own (see
Appendix 3). Websites are listed for some chapters.
The book is classified into units, which consist of one or
more chapters. Of course, a given instructor may choose to
vary the sequence of these units or the chapters within,
depending on personal preference and the availability of plant
material. There is a slight amount of repetition between chap-
ters of different units, but this was done so that chapters could
be used independently of one another.
Unit 1, Systematics, gives a general overview of the
concepts and methods of the field. Chapter 1 serves as an
introduction to the definition, relationships, classification,
and importance of plants and summarizes the basic concepts
and principles of systematics, taxonomy, evolution, and phy-
logeny. Chapter 2 introduces taxonomy as continuing, active
field of research and covers the details of phylogenetic sys-
tematics, and the theory and methodology for inferring phy-
logenetic trees or cladograms.
ix
Unit 2, Evolution and Diversity of Plants, describes in detail
the characteristics and classification of plants. The six chapters of
this unit are intended to give the beginning student a basic
understanding of the evolution of Green and Land Plants
(Chapter 3), Vascular Plants (Chapter 4), Woody and Seed Plants
(Chapter 5), and Flowering Plants (Chapters 6–8). Chapters 3–5
are formatted into two major sections. The first section presents
cladograms (phylogenetic trees), which portray the evolutionary
history of the group. Each of the major derived evolutionary fea-
tures (apomorphies) from that cladogram is described and illus-
trated, with emphasis on its possible adaptive significance. This
evolutionary approach to plant systematics makes learning the
major plant groups and their features conceptually easier than
simply memorizing a static list of characteristics. Treating these
features as the products of unique evolutionary events brings
them “to life,” especially when their possible functional signifi-
cance is pondered. The second section of Chapters 3–5 pres-
ents a survey of the diversity of the group in question.
Etymologies of the type genus for families are included. Rare
conditions and synonyms are enclosed by square brackets.
Exemplars within major groups are described and illustrated,
such that the student may learn to recognize and know the
basic features of the major lineages of plants.
Because they constitute the great majority of plants, the
flowering plants, or angiosperms, are covered in three chap-
ters. Chapter 6 deals with the evolution of flowering plants,
describing the apomorphies for that group and presenting a
synopsis of their origin. Chapters 7 and 8 describe specific
groups of flowering plants. In Chapter 7 the non-eudicot
groups are treated, including the earliest diverging angio-
sperm lineages and the monocotyledons. Chapter 8 covers
the eudicots, which make up the great majority of angio-
sperms. In these two chapters numerous flowering plant fam-
ilies (166 of the 472 recognized here) are described in detail,
accompanied by photographs and illustrations; these are
mostly families that are commonly encountered or for which
material is usually available to the beginning student.
Additional families are not described, but are illustrated with
one or more exemplars. I have tried to emphasize diagnostic
features a student might use to recognize a plant family, and
have included some economically important uses of family
members. Reference to Chapter 9 and occasionally to
Chapters 10–14 (or use of the comprehensive Glossary) may
x  Preface
be needed with regard to the technical terms. The Angiosperm
Phylogeny Group IV system of classification is used through-
out, with some exceptions. This system uses orders as the
major taxonomic rank in grouping families of close relation-
ship and has proven extremely useful in dealing with the tre-
mendous diversity of the flowering plants.
Unit 3, Systematic Evidence and Descriptive Terminology,
begins with a chapter on plant morphology (Chapter 9).
Explanatory text, numerous diagrammatic illustrations, and
photographs are used to train beginning students to precisely
and thoroughly describe a plant morphologically. Appendices
1 and 2 (see below) are designed to be used along with Chapter
9. The other chapters in this unit cover the basic descriptive
terminology of plant anatomy (Chapter 10), plant embryol-
ogy (Chapter 11), palynology (Chapter 12), plant reproduc-
tive biology (Chapter 13), and plant molecular systematics
(Chapter 14). The rationale for including these in a textbook
on plant systematics is that features from these various fields
are described in systematic research and are commonly uti-
lized in phylogenetic reconstruction and taxonomic delimita-
tion. In particular, in the last chapter on plant molecular
systematics, I have attempted to update techniques and meth-
odologies acquired in what has become in recent years the
most fruitful of endeavors in phylogenetic reconstruction.
Unit 4, Resources in Plant Systematics, discusses some
basics that are essential in everyday systematic research. Plant
identification (Chapter 15) contains a summary of both stan-
dard dichotomous keys and computerized polythetic keys
and reviews practical identification methods. The chapter on
nomenclature (Chapter 16) summarizes the basic rules of the
most recent International Code of Nomenclature for Algae,
Fungi, and Plants, including the steps needed in the valid
publication of a new species plus a review of botanical names.
A chapter on plant collecting and documentation (Chapter
17) emphasizes both correct techniques for collecting plants
and thorough data acquisition, the latter of which has become
increasingly important today in biodiversity studies and con-
servation biology. Finally, the chapter on herbaria and data
information systems (Chapter 18) reviews the basics of her-
barium management, emphasizing the role of computerized
database systems in plant collections for analyzing and syn-
thesizing morphological, ecological, and biogeographic data.
Unit 5, Species Concepts and Conservation Biology, con-
tains a chapter (Chapter 19) that reviews basic plant repro-
duction and the criteria and concepts of species and
infraspecies definitions. In addition, a section on conserva-
tion biology reviews the basic concepts of this field, how it
relates to taxonomy and systematics, and its importance to
biologists and society.
Lastly, four Appendices and a Glossary are included. I have
personally found each of these addenda to be of value in my own
plant systematics courses. Appendix 1 is a list of characters used
for detailed plant descriptions (available on the Plant Systematics
Resources website). This list is useful in training students to write
descriptions suitable for publication. Appendix 2 is a brief dis-
cussion of botanical illustration. I have found that drawing what
one sees helps to develop observational skills. Appendix 3 is a
listing of scientific journals in plant systematics, with literature
exercises. Appendix 4 gives a brief overview of statistical and
morphometric methods and how those may be applied in
addressing questions in taxonomy and phylogenetic systematics.
The Glossary defines all terms used in the book and indicates
synonyms, adjectival forms, plurals, abbreviations, and terms to
compare.
Three web sites will be available to be used in conjunction
with the textbook: (1) a Plant Systematics Resources site (http://
www.sci.sdsu.edu/plants/plantsystematics), with web links and
materials that are universally available; (2) a companion website
(http://www.elsevierdirect.com/companions/9780123743800)
that includes the chapter figures, appendix material from the
textbook, and links to the author’s website; and (3) an Instructor
Resources site (http://textbooks.elsevier.com/web/Login.aspx),
with material that is password protected. Please contact your
sales representative at textbooks@elsevier.com for access to the
Instructor Resources site.
Throughout the book, I have attempted to adhere to
W–H–Y, What–How–Why, in organizing and clarifying
chapter topics: (1) What is it? What is the topic, the basic def-
inition? (Many scientific arguments could have been resolved
at the start by a clear statement or definition of terms.)
(2) How is it done? What are the materials and methods, the
techniques of data acquisition, the types of data analysis?
(3) Why is it done? What is the purpose, objective, or goal?
What is the overriding paradigm involved? How does the
current study or topic relate to others? This simple W–H–Y
method, first presented to me by one of my mentors, A. E.
Radford, is useful to follow in any intellectual endeavor. It is a
good lesson to teach one’s students, and helps both in devel-
oping good writing skills and in critically evaluating a topic.
Finally, I would like to propose that each of us, instructors
and students, pause occasionally to evaluate why it is that
we do what we do. Over the years I have refined my ideas
and offer these suggestions as possible goals: (1) to realize and
explore the beauty, grandeur, and intricacy of nature;
(2) to engage in the excitement of scientific discovery; (3) to
experience and share the joy of learning. It is in this spirit that
I sincerely hope the book may be of use to others.
 I
SYSTEMATICS
 1
PLANT SYSTEMATICS:
AN OVERVIEW
PLANTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
What Is a Plant? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Phylogeny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Plants and the Evolution of Life . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Why Study Systematics? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Land Plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
REVIEW QUESTIONS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Why Study Plants? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
EXERCISES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
SYSTEMATICS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
What Is Systematics? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 REFERENCES FOR FURTHER STUDY . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Evolution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

This book is about a fascinating field of biology called plant cussion), defined by the common (but independently evolved)
systematics. The purpose of this chapter is to introduce the characteristic of photosynthesis. However, delimiting organismal
basics: what a plant is, what systematics is, and the reasons groups based on evolutionary history has gained almost universal
for studying plant systematics. acceptance. This latter type of classification directly reflects the
patterns of that evolutionary history and can be used to explicitly
test evolutionary hypotheses (discussed later; see Chapter 2).
PLANTS An understanding of what plants are requires an explanation
of the evolution of life in general.
WHAT IS A PLANT?
This question can be answered in either of two conceptual PLANTS AND THE EVOLUTION OF LIFE
ways. One way, the traditional way, is to define groups of Life is currently classified as three major groups (sometimes
organisms such as plants by the characteristics they possess. called domains) of organisms: Archaea (also called Archae-
Thus, historically, “plants” included those organisms that pos- bacteria), Bacteria (also called Eubacteria), and Eukarya
sess photosynthesis, cell walls, spores, and a more or less sed- or eukaryotes (also spelled eucaryotes). The evolutionary
entary behavior. This traditional grouping of plants contained relationships of these groups are summarized in the simplified
a variety of microscopic organisms, all of the “algae,” and the evolutionary tree or cladogram of Figure 1.1. The Archaea and
more familiar plants that live on land. A second way to answer Bacteria are small, mostly unicellular organisms that possess
the question “What is a plant?” is to evaluate the evolutionary circular DNA, replicate by fission, and lack membrane-bound
history of life and to use that history to delimit the groups of organelles. The two groups differ from one another in the
life. We now know from repeated research studies that some chemical structure of certain cellular components. Eukaryotes
of the photosynthetic organisms evolved independently of one are unicellular or multicellular organisms that possess linear
another and are not closely related. DNA (organized as histone-bound chromosomes), replicate by
Thus, the meaning or definition of the word plant can be mitotic and often meiotic division, and possess membrane-bound
ambiguous and can vary from person to person. Some still like organelles such as nuclei, cytoskeletal structures, and (in almost
to treat plants as a “polyphyletic” assemblage (see later dis- all) mitochondria (Figure 1.1).

PLANT SYSTEMATICS
https://doi.org/10.1016/B978-0-12-812628-8.50001-8, Copyright © 2019 Elsevier Inc. All rights reserved. 3
4  CHAPTER 1 PLANT SYSTEMATICS: AN OVERVIEW

Eukarya (Eukaryotes)

Viridiplantae (Green plants)

Stramenopiles

Rhodophyta (Red algae)

Excavata

Phaeophyta (Browns)
Alveolates

(incl. slime molds)

Ampicomplexans

Dinoflagellates

Chlorobionta/
Opisthokonts

Kinetoplastids

(water molds)
Bacteria

Archaea

Diplomonads

Euglenozoa
Parabasalids
Amoebozoa

Oomycota
Animalia

Rhizaria
Ciliates
Fungi

*chloroplast “red” chloroplast

*chloroplast “green” chloroplast
*chloroplast

chloroplast origin
Primary
Endosymbiosis

mitochondria (by endosymbiosis)

cytoskeletal/contractile elements (actin, myosin, tubulin )
other membrane-bound or ganelles (endoplasm. retic., golgi, lysosomes)
mitosis (+ meiosis in sexually reproducing organisms)
nucleus (membrane bound), enclosing chromosomes
DNA linear, bound to histones

= endosymbiotic origin of mitochondrion

and chloroplast from ancestral Bacterium
* = secondary chloroplast endosymbioses

Figure 1.1 Simplified cladogram (evolutionary tree) of life (modified from Parfrey et al. 2010; see also Williams et al. 2018), illustrating
eukaryotic apomorphies (the relative order of which is unknown) and the hypothesis of a single origin of mitochondria and chloroplasts via
endosymbiosis (arrows). Note modification of chloroplast structure in the red and green plants, and subsequent secondary endosymbiosis in
numerous other lineages (indicated by *). Eukaryotic groups with photosynthetic members in bold.

Some of the unicellular bacteria (e.g., the Cyanobacteria, or How did chloroplasts evolve? It is now largely accepted that
blue-greens) carry on photosynthesis, a biochemical system chloroplasts of eukaryotes originated by the engulfment of an
in which light energy is used to synthesize high-energy com- ancestral photosynthetic bacterium (probably a cyanobacteri-
pounds from simpler starting compounds, carbon dioxide and um) by an ancestral eukaryotic cell such that the photosynthetic
water. These photosynthetic bacteria have a system of internal bacterium continued to live and ultimately multiply inside the
membranes called thylakoids, within which are embedded pho- eukaryotic cell (Figures 1.1, 1.2). (Mitochondria also evolved
tosynthetic pigments, compounds that convert light energy to by this process, from an ancestral, non-photosynthetic bacte-
chemical energy. Of the several groups of eukaryotes that are rium; see Figure 1.1.) The evidence for this is the fact that chlo-
photosynthetic, all have specialized photosynthetic organelles roplasts, like bacteria today, (a) have their own single-stranded,
called chloroplasts, which resemble photosynthetic bacteria circular DNA; (b) have a smaller sized, 70S ribosome; and (c)
in having pigment-containing thylakoid membranes. replicate by fission. These engulfed photosynthetic bacteria

Ancestral . .. .
.
. . . . . . . ..
photosynthetic
bacterium
. . . .
. .
. .
. .
. . .
. .. .
. . . . . . .. . . .
.. . .. . . . . . . . .
. .
. . . . . . .. . . . . . . . .. . .
. . .. . . .
. .
Eukaryotic cell
Figure 1.2 Diagrammatic illustration of the origin of chloroplasts by endosymbiosis of ancestral photosynthetic bacterium within ancestral
eukaryotic cell.
provided high-energy products to the eukaryotic cell; the
“host” eukaryotic cell provided a beneficial environment for
the photosynthetic bacteria. The condition of two species living
together in close contact is termed symbiosis, and the process
in which symbiosis results by the engulfment and retention of
one cell by another is termed endosymbiosis. Over time, these
endosymbiotic, photosynthetic bacteria became transformed
structurally and functionally, retaining their own DNA and the
ability to replicate, but losing the ability to live independently
of the host cell. In fact, over time there has been a transfer of
some genes from the DNA of the chloroplast to the nuclear
DNA of the eukaryotic host cell, making the two biochemi-
cally interdependent.
Although knowledge of eukaryotic relationships is still in flux,
the most recent data from molecular systematic studies indicates
that this so-called “primary” endosymbiosis of the chloroplast
probably occurred one time, a shared evolutionary novelty of
the red algae (Rhodophyta) and green plants (Viridiplantae
or Chlorobionta; Figure 1.1). This early chloroplast became
modified with regard to photosynthetic pigments, thylakoid
structure, and storage products into forms characteristic of the
red algae and green plants (see Figure 1.1). In addition, several
lineages of photosynthetic organisms – including the euglenoids
(Euglenozoa), dinoflagellates, and brown algae (Phaeophyta),
and a few other lineages – may have acquired chloroplasts via
“secondary” endosymbiosis, which occurred by the engulfment
of an ancestral chloroplast-containing eukaryote by another
eukaryotic cell (Figure 1.1).
LAND PLANTS
Of the major groups of photosynthetic eukaryotes, the green
plants (Viridiplantae or Chlorobionta) are united primarily by
distinctive characteristics of the green plant chloroplast with
respect to photosynthetic pigments, thylakoid structure, and
storage compounds (see Chapter 3 for details). Green plants
include both the predominately aquatic “green algae” and a
UNIT I SYSTEMATICS    5
Self-replicating
chloroplasts
.
.. .
. . .
. . . . . . .
. . ..
. . . .
. . . . .
. .. .
.. . . . . .. . . . . .. .
.. . .. . . . . . . .
. .
. . . . .
. . .
Photosynthetic
eukaryotic cell
group known as embryophytes (formally, the Embryophyta),
usually referred to as the land plants (Figure 1.3). The land
plants are united by several evolutionary novelties that were
adaptations to the transition from an aquatic environment to
living on land. These include (1) an outer cuticle, which aids in
protecting tissues from desiccation; (2) specialized gametangia
(egg and sperm producing organs) that have an outer, protec-
tive layer of sterile cells; and (3) an intercalated diploid phase
(sporophyte) in the life cycle, the early, immature component
of which is termed the embryo (hence, “embryophytes”; see
Chapter 3 for details).
Just as the green plants include the land plants, the land
plants are inclusive of the vascular plants (Figure 1.3), the latter
being united by the evolution of an independent sporophyte
and xylem and phloem vascular conductive tissue (see Chapter
4). The vascular plants are inclusive of the seed plants (Figure
1.3), which are united by the evolution of wood and seeds
(see Chapter 5). Finally, seed plants include the angiosperms
(Figure 1.3), united by the evolution of the flower, including
carpels and stamens, and by a number of other specialized
features (see Chapters 6–8).
For the remainder of this book, the term plant is treated as
equivalent to the embryophytes, the land plants. The rationale for
this is partly that land plants make up a so-called monophyletic
group, whereas the photosynthetic eukaryotes as a whole are not
monophyletic, and do not accurately reflect evolutionary history
(see later discussion, Chapter 2). And, practically, it is land plants
that most people are talking about when they refer to “plants,”
including those in the field of plant systematics. However, as
noted before, the word plant can be used by some to refer to
other groupings; when in doubt, get a precise clarification.
WHY STUDY PLANTS?
The tremendous importance of plants cannot be overstated.
Without them, we and most other species of animals (and of
many other groups of organisms) would not be here. Photosyn-
6  CHAPTER 1 PLANT SYSTEMATICS: AN OVERVIEW

Chlorobionta (Viridiplantae) – green plants

Embryophytes – land plants*
Tracheophytes – vascular plants
Monilophytes Spermatophytes – seed plants
Gymnosperms Angiosperms

Ophioglossales
Polypodiales
Marattiales
Lycophytes

(incl. Gnetales)
Equisetales
Liverworts

Hornworts
"Green Algae"

Monocots
Psilotales

Eudicots
Conifers
Ginkgo
Cycads
Mosses
Flower, carpels,
stamens (+ sev.
other features)
Seeds
Wood

Xylem & phloem vascular tissue

Independent sporophyte

Cuticle, gametangia, embryo (sporophyte)

Green plant chloroplast

Figure 1.3 Simplified cladogram (evolutionary tree) of the green plants, illustrating major extant groups and evolutionary events
(or “apomorphies,” hash marks). *Embryophytes are treated as “plants” in this book.

thesis in plants and the other photosynthetic organisms changed
the earth in two major ways. First, the fixation of carbon di-
oxide and the release of molecular oxygen in photosynthesis
directly altered the earth’s atmosphere over billions of years.
What used to be an atmosphere deficient in oxygen underwent
a gradual change. As a critical mass of oxygen accumulated
in the atmosphere, selection for oxygen-dependent respiration
occurred (via oxidative phosphorylation in mitochondria),
which may have been a necessary precursor to the evolution
of many multicellular organisms, including all animals. In ad-
dition, an oxygen-rich atmosphere permitted the establishment
of an upper atmosphere ozone layer, which shielded life from
excess UV radiation. This allowed organisms to inhabit more
exposed niches that were previously inaccessible.
Second, the compounds that photosynthetic species produce
are utilized, directly or indirectly, by nonphotosynthetic (heterotro-
phic) organisms. For virtually all land creatures and many aquatic
ones as well, land plants make up the so-called primary producers
in the food chain, the source of high-energy compounds such as
carbohydrates, structural compounds such as certain amino acids,
and other compounds essential to metabolism in some hetero-
trophs. Thus, most species on land today, including the millions
of species of animals, are absolutely dependent on plants for their
survival. As primary producers, plants are the major components,
and definers, of many communities and ecosystems. The survival
of plants is essential to maintaining the health of those ecosystems,
the severe disruption of which could bring about rampant species
extirpation or extinction and disastrous changes in erosion, water

flow, and ultimately climate.
To humans, plants are also monumentally important in
numerous, direct ways (Figures 1.4, 1.5). Agricultural plants,
most of which are flowering plants, are our major source of
food. We utilize all plant parts as food products: roots (e.g., sweet
potatoes and carrots; Figure 1.4A,B); stems (e.g., yams, cassava/
manioc, potatoes; Figure 1.4C); leaves (e.g., cabbage, celery,
lettuce; Figure 1.4D); flowers (e.g., cauliflower and broccoli;
Figure 1.4E); and fruits and seeds, including grains such as
rice (Figure 1.4F), wheat (Figure 1.4G), corn (Figure 1.4H),
rye, barley, and oats, legumes such as beans and peas (Figure
1.4I), and a plethora of fruits such as bananas (Figure 1.4J),
tomatoes, peppers, pineapples (Figure 1.4K), apples (Figure
1.4L), cherries, peaches, melons, kiwis, citrus, olives (Figure
1.4M), and others too numerous to mention. Other plants
are used as flavoring agents, such as herbs (Figure 1.5A–D)
and spices (Figure 1.5E), as stimulating beverages, such as
chocolate, coffee, tea, and cola (Figure 1.5F), or as alcoholic
drinks, such as beer, wine, distilled liquors, and sweet liqueurs.
Woody trees of both conifers and flowering plants are used
structurally for lumber and for pulp products such as paper
(Figure 1.5G). Non-woody plants, such as bamboos, palms,
and a variety of other species, serve as construction materials
for a great variety of purposes. Plant fibers are used to make
thread for cordage (such as sisal), for sacks (such as jute for
burlap), and for textiles (most notably cotton, Figure 1.5H, but
also linen and hemp, Figure 1.5I). Extracts from plants, which
include essential oils, latex (for rubber or balata), vegetable
oils, pectins, starches, and waxes, have a plethora of uses in
industry, food, perfume, and cosmetics. In many cultures,
plants or plant products are used as euphorics or hallucinogen-
ics (whether legally or illegally), such as marijuana (Figure
1.5I), opium, cocaine, and a great variety of other species that
have been used by indigenous peoples for centuries. Plants
are important for their aesthetic beauty, and the cultivation of
plants as ornamentals is an important industry. Finally, plants
have great medicinal significance, to treat a variety of illnesses
or to maintain good health. Plant products are very important
in the pharmaceutical industry; their compounds are extracted,
semisynthesized, or used as templates to synthesize new drugs.
Many “modern” drugs, from aspirin (originally derived from
the bark of willow trees) to vincristine and vinblastine (ob-
tained from the Madagascar periwinkle, used to treat childhood
leukemia; Figure 1.5J), are ultimately derived from plants. In
addition, various plant parts of a great number of species are
used whole or are processed as so-called herbal supplements,
which have become tremendously popular of late.
The people, methods, and rationale concerned with the
plant sciences (defined here as the study of land plants) are
as diverse as are the uses and importance of plants. Some of
UNIT I SYSTEMATICS    7
the fields in the plant sciences are very practically oriented.
Agriculture and horticulture deal with improving the yield or
disease resistance of food crops or cultivated ornamental plants,
e.g., through breeding studies and identifying new cultivars.
Forestry is concerned with the cultivation and harvesting of
trees used for lumber and pulp. Pharmacognosy deals with
crude natural drugs, often of plant origin. In contrast to these
more practical fields of the plant sciences, the “pure” sciences
have as their goal the advancement of scientific knowledge
(understanding how nature works) through research, regard-
less of the practical implications. But many aspects of the
pure sciences also have important practical applications, either
directly by applicable discovery or indirectly by providing the
foundation of knowledge used in the more practical sciences.
Among these are plant anatomy, dealing with cell and tissue
structure and development; plant chemistry and physiology,
dealing with biochemical and biophysical processes and
products; plant molecular biology, dealing with the structure
and function of genetic material; plant ecology, dealing with
interactions of plants with their environment; and, of course,
plant systematics.
Note that a distinction should be made between “botany” and
“plant sciences.” Plant sciences is the study of plants, treated
as equivalent to land plants here. Botany is the study of most
organisms traditionally treated as plants, including virtually
all eukaryotic photosynthetic organisms (land plants and the
several groups of “algae”) plus other eukaryotic organisms
with cell walls and spores (true fungi and groups formerly
treated as fungi, such as the water molds, Oomycota, and the
slime molds, included within the Amoebozoa; Figure 1.1).
Thus, in this sense, botany is inclusive of but broader than the
plant sciences. Recognition of both botany and plant sciences
as fields of study can be useful, although how these fields are
defined can vary and may require clarification.
SYSTEMATICS
WHAT IS SYSTEMATICS?
Systematics is defined in this book as a science that includes
and encompasses traditional taxonomy, the description, iden-
tification, nomenclature, and classification of organisms, and
that has as its primary goal the reconstruction of phylogeny, or
evolutionary history, of life. This definition of systematics is
not novel, but neither is it universal. Others in the field would
treat taxonomy and systematics as separate but overlapping
areas; still others argue that historical usage necessitates what
is in essence a reversal of the definitions used here. But words,
like organisms, evolve. The use of systematics to describe an
all-encompassing field of endeavor is both most useful and
8  CHAPTER 1 PLANT SYSTEMATICS: AN OVERVIEW

A B C D E

F G H I

J K L M
Figure 1.4 Examples of economically important plants. A–E. Vegetables. A. Ipomoea batatas, sweet potato (root). B. Daucus carota, carrot
(root). C. Solanum tuberosum, potato (stem). D. Lactuca sativa, lettuce (leaves). E. Brassica oleracea, broccoli (flower buds).
F–H. Fruits, dry (grains). F. Oryza sativa, rice. G. Triticum aestivum, bread wheat. H. Zea mays, corn. I. Seeds (pulse legumes), from top,
clockwise to center: Glycine max, soybean; Lens culinaris, lentil; Phaseolus aureus, mung bean; Phaseolus vulgaris, pinto bean; Phaseolus
vulgaris, black bean; Cicer areitinum, chick-pea/garbanzo bean; Vigna unguiculata, black-eyed pea; Phaseolus lunatus, lima bean. J–M. Fruits,
fleshy. J. Musa paradisiaca, banana. K. Ananas comosus, pineapple. L. Malus pumila, apple. M. Olea europaea, olive.
 UNIT I SYSTEMATICS    9

A B C D

E F

G H I J
Figure 1.5 Further examples of economically important plants. A–D. Herbs. A. Petroselinum crispum, parsley. B. Salvia officinalis, sage.
C. Salvia rosmarinus, rosemary. D. Thymus vulgaris, thyme. E. Spices and herbs, from upper left: Cinnamomum cassia/zeylanicum, cinnamon
(bark);Vanilla planifolia; vanilla (fruit); Laurus nobilis, laurel (leaf); Syzygium aromaticum, cloves (flower buds); Myristica fragrans, nutmeg
(seed); Carum carvi, caraway (fruit); Anethum graveolens, dill (fruit); Pimenta dioica, allspice (seed); Piper nigrum, pepper (seed).
F. Flavoring plants, from upper left, clockwise. Theobroma cacao, chocolate (seeds); Coffea arabica, coffee (seeds); Camellia sinensis, tea
(leaves). G. Wood products: lumber (Sequoia sempervirens, redwood), and paper derived from wood pulp. H. Fiber plant. Gossypium sp.,
cotton (seed trichomes), one of the most important natural fibers. I. Euphoric, medicinal, and fiber plant. Cannabis sativa, marijuana, hemp;
stem fibers used in twine, rope, and cloth; resins contain the euphoric and medicinal compound tetrahydrocannabinol. J. Medicinal plant.
Catharanthus roseus, Madagascar periwinkle, from which is derived vincristine and vinblastine, used to treat childhood leukemia.
10  CHAPTER 1 PLANT SYSTEMATICS: AN OVERVIEW
represents the consensus of how most specialists in the field use
the term, an example being the journal Systematic Botany, which
contains articles both in traditional taxonomy and phylogenetic
reconstruction. Plant systematics is studied by acquiring, ana-
lyzing, and synthesizing information about plants and plant
parts, the content and methodology of which is the topic for
the remainder of this book.
Systematics is founded in the principles of evolution, its
major premise being that there is one phylogeny of life. The
goal of systematists is, in part, to discover that phylogeny.
EVOLUTION
Evolution, in the broadest sense, means “change” and can be
viewed as the cumulative changes occurring since the origin of
the universe some 14 billion years ago. Biological evolution,
the evolution of life, may be defined (as it was by Charles Dar-
win) as “descent with modification.” Descent is the transfer of
genetic material (enclosed within a cell, the unit of life) from
parent(s) to offspring over time. This is a simple concept, but one
that is important to grasp and ponder thoroughly. Since the time
that life first originated some 3.8 billion years ago, all life has
been derived from preexisting life. Organisms come to exist
by the transfer of genetic material, within a surrounding cell,
from one or more parents. Descent may occur by simple clonal
reproduction, such as a single bacterial cell “parent” dividing
by fission to form two “offspring” cells or a land plant giving
rise to a vegetative propagule. It may also occur by complex
sexual reproduction (Figure 1.6A), in which each of two parents
produces specialized gametes (e.g., sperm and egg cells), which
contain half the complement of genetic material, the result of
meiosis. Two of the gametes fuse together to form a new cell,
the zygote, which may develop into a new individual (as oc-
curs in plants; see Chapter 3) or may itself divide by meiosis
to form gametes. Descent through time results in the formation
of a lineage (Figure 1.6B), a set of organisms interconnected
through time and space by the transfer of genetic material from
parents to offspring. So, in a very literal sense, we and all other
forms of life on earth are connected by descent, the transfer of
DNA (actually the pattern of DNA) from parent to offspring
(ancestor to descendant), generation after generation.
The modification component of evolution refers to a change
in the genetic material that is transferred from parent(s) to
offspring, such that the genetic material of the offspring is dif-
ferent from that of the parent(s). This modification may occur
either by mutation, which is a direct alteration of DNA, or by
genetic recombination, whereby existing genes are reshuffled
in different combinations (during meiosis in eukaryotes, by the
genetic processes of crossing over and independent assortment).
Systematics is concerned with the identification of the unique
modifications of evolution.
It should also be asked, what evolves? Although genetic
modification may occur in offspring relative to their parents,
individual organisms do not generally evolve. This is because
a new individual begins when it receives its complement of
DNA from the parent(s); that individual’s DNA does not
change during its/his/her lifetime (with the exception of rela-
tively rare, nonreproductive “somatic” mutations). The general
units of evolution are populations and species. A population
is a group of individuals of the same species that is usually
geographically delimited and that typically have a significant
amount of gene exchange. Species are groups of populations
that are related to one another by various criteria and that have
evolutionarily diverged from other such groups. There are a
number of different species concepts of definitions, dependent
on the biological system and on the criteria used to recognize
them (see Chapter 19). With changes in the genetic makeup of
offspring (relative to parents), the genetic makeup of popula-
tions and species changes over time.
In summary, evolution is descent with modification occurring
by a change in the genetic makeup (DNA) of populations or spe-
cies over time. How does evolution occur? Evolutionary change
may come about by two major mechanisms: (1) genetic drift, in
which genetic modification is random; or (2) natural selection,
in which genetic change is directed and nonrandom. Natural
selection is the differential contribution of genetic material
from one generation to the next, differential in the sense that
genetic components of the population or species are contrib-
uted in different amounts to the next generation; those genetic
combinations resulting in increased survival or reproduction
are contributed to a greater degree. (A quantitative measure
of this differential contribution is known as fitness.) Natural
selection results in an adaptation, a structure or feature that
performs a particular function and which itself brings about
increased survival or reproduction. In a consideration of the
evolution of any feature in systematics, the possible adaptive
significance of that feature should be explored.
Finally, an ultimate result of evolution is speciation, the
formation of new species from preexisting species. Speciation
can follow lineage divergence, the splitting of one lineage into
two, separate lineages (Figure 1.6D). Lineage divergence is
itself a means of increasing evolutionary diversity. If two,
divergent lineages remain relatively distinct, they may change
independently of one another, into what may be designated as
separate species (see Chapter 19).
TAXONOMY
Taxonomy is a major part of systematics that includes four
components: Description, Identification, Nomenclature, and
Classification. (Remember the mnemonic device: DINC.) The
general subjects of study are taxa (singular, taxon), which are
 UNIT I SYSTEMATICS    11

egg sperm egg sperm

fusion of fusion of
gametes gametes

LINEAGE

Sporophyte
Gametophytes(n) Gametophytes(n) (2n)
Female Male Female Male

spores spore
meiosis meiosis
Sporophytes
(2n)

A B TIME
SPOROPHYTE PERENNIAL SPOROPHYTE ANNUAL
HETEROSPOROUS HETEROSPOROUS

SPECIES 1 SPECIES 2

DIVERGENCE

LINEAGE LINEAGE

LINEAGE

C
TIME
Figure 1.6 A. Simplified diagram of descent in sexually reproducing land plants, in which diploid sporophytes give rise to haploid spores
(through meiosis), which develop into haploid gametophytes; the latter produce egg and sperm, fusing to form a diploid zygote, which devel-
ops into a diploid sporophyte. B. A lineage, the result of transfer of genetic material over time and space. C. Divergence of one lineage into
two, which may result in speciation (illustrated here).
12  CHAPTER 1 PLANT SYSTEMATICS: AN OVERVIEW

defined or delimited groups of organisms. Ideally, taxa should
have a property known as monophyly (discussed later; Chapter
2) and are traditionally treated at a particular rank (see later
discussion). It should be pointed out that the four components
of taxonomy are not limited to formal systematic studies but
are the foundation of virtually all intellectual endeavors of all
fields, in which conceptual entities are described, identified,
named, and classified. In fact, the ability to describe, identify,
name, and classify things undoubtedly has been selected for
in humans and, in part, in other organisms as well.
Description is the assignment of features or attributes to a
taxon. The features are called characters. Two or more forms
of a character are character states. One example of a character
is “petal color,” for which two character states are “yellow”
and “blue.” Another character is “leaf shape,” for which pos-
sible character states are “elliptic,” “lanceolate,” and “ovate.”
Numerous character and character state terms are used in plant
systematics, both for general plant morphology (see Chapter 9)
and for specialized types of data (Chapters 10–14). The purpose
of these descriptive character and character state terms is to
use them as tools of communication, for concisely categorizing
and for delimiting the attributes of a taxon, an organism, or
some part of the organism. An accurate and complete listing
of these features is one of the major objectives and contribu-
tions of taxonomy.
Identification is the process of associating an unknown
taxon with a known one, or recognizing that the unknown is
new to science and warrants formal description and naming.
One generally identifies an unknown by first noting its charac-
teristics, that is, by describing it. Then, these features are com-
pared with those of other taxa to see if they conform. Plant taxa
can be identified in many ways (see Chapter 15). A taxonomic
key is perhaps the most utilized of identification devices. Of the
different types of taxonomic keys, the most common, used in
virtually all floras, is a dichotomous key. A dichotomous key
consists of a series of two contrasting statements. Each state-
ment is a lead; the pair of leads constitutes a couplet (Figure
1.7). That lead which best fits the specimen to be identified is
selected; then all couplets hierarchically beneath that lead (by
indentation and/or numbering) are sequentially checked for
agreement until an identification is reached (Figure 1.7).
Nomenclature is the formal naming of taxa according to
some standardized system. For algae, fungi, and plants, the
rules and regulations for the naming of taxa are provided by
the International Code of Nomenclature for algae, fungi, and
plants. These formal names are known as scientific names,
which by convention are translated into the Latin language.
The fundamental principle of nomenclature is that all taxa
may bear only one “correct” scientific name (Chapter 16).
Although they may seem difficult to learn at first, scientific
names are much preferable to common (vernacular) names.
(Note that another system of nomenclature, the Phylocode, is
also available and has been used especially for names of higher
rank; see Chapter 16.)
The scientific name of a species traditionally consists of two
parts (typically underlined or italicized): the genus name, which
is always capitalized, e.g., Quercus, plus the specific epithet,
which by general consensus is not capitalized, e.g., agrifolia.
Thus, the species name for what is commonly called Califor-
nia live oak is Quercus agrifolia. Species names are known as
binomials (literally meaning “two names”) and this type of
nomenclature is called binomial nomenclature, first formalized
in the mid-18th century by Carolus Linnaeus.
Classification is the arrangement of entities (in this case,
taxa) into some type of order. The purpose of classification is
to provide a system for cataloguing and expressing relation-
ships between these entities. Taxonomists have traditionally
agreed upon a method for classifying organisms that utilizes
categories called ranks. These taxonomic ranks are hierar-
chical, meaning that each rank is inclusive of all other ranks
beneath it (Figure 1.8).
As defined earlier, a taxon is a group of organisms typically
treated at a given rank. Thus, in the example of Figure 1.8,
Magnoliophyta is a taxon placed at the rank of phylum; Liliopsida
is a taxon placed at the rank of class; Arecaceae is a taxon placed
at the rank of family; etc. Note that taxa of a particular rank

Lead: 1. Leaf simple of generally pinnately compound, leaflets 0 or (1)3–9; fruit winged achene ............. Fraxinus
Couplet:
Lead: 1' Leaf simple; fruit capsule or drupe
2. Leaves generally alternate; fruit a deeply 2-lobed capsule ............................................................ Menodora
2' Leaves opposite or clustered; fruit a drupe
3. Flowers unisexual; corolla 0; stamens 0 or 4–5 ............................................................................ Forestiera
3' Flowers bisexual; corolla funnel-shaped, salverform, or rotate; stamens 2
4. Corolla funnel-shaped or salverform; fruit 1–4-seeded ........................................................ Ligustrum
4' Corolla rotate; fruit 1-seeded ................................................................................................ Olea

Figure 1.7 Dichotomous key to the genera of the Oleaceae of California, by Thomas J. Rosatti, Oleaceae, in Jepson Flora Project (eds.)
Jepson eFlora, http://ucjeps.berkeley.edu/cgi-bin/get_IJM.pl?key=207, accessed on May 31, 2019, reprinted by special permission.

Major Taxonomic Ranks
Kingdom
Phylum (“Division” also acceptable)
Class
Order
Family
Genus (plural: genera)
Species (plural: species)
Figure 1.8 The primary taxonomic ranks accepted by the International Code of Nomenclature for algae, fungi, and plants, with some
examples of taxa.
generally end in a particular suffix (Chapter 16). There is a trend
among systematic biologists to eliminate the rank system of
classification (see Chapter 16). In this book, ranks are generally
used for naming groups but not emphasized as ranks.
There are two major means of arriving at a classification of
life: phenetic and phylogenetic. Phenetic classification is that
based on overall similarities. Most of our everyday classifica-
tions are phenetic. For efficiency of organization (e.g., storing
and retrieving objects, like nuts and bolts in a hardware store) we
group similar objects together and dissimilar objects apart. Many
traditional classifications in plant systematics are phenetic, based
on noted similarities between and among taxa. Phylogenetic
classification is that which is based on evolutionary history, or
pattern of descent, which may or may not correspond to overall
similarity (see later discussion, Chapter 2).
PHYLOGENY
Phylogeny, the primary goal of systematics, refers to the
evolutionary history of a group of organisms. Phylogeny is
commonly represented in the form of a cladogram (or phylo-
genetic tree), a branching diagram that conceptually represents
the evolutionary pattern of descent (see Figure 1.9). The lines
of a cladogram represent lineages, which (as discussed earlier)
denote descent, the sequence of ancestral-descendant popula-
tions through time (Figure 1.9A). Thus, cladograms have an
implied (relative) time scale. Any branching of the cladogram
represents lineage divergence, the diversification of lineages
from one common ancestor.
Changes in the genetic makeup of populations, i.e., evolu-
tion, may occur in lineages over time. Evolution is recognized
as a change from a preexisting, or ancestral, character state to
a new, derived character state. The derived character state is an
evolutionary novelty, also called an apomorphy (Figure 1.9A).
Phylogenetic systematics, or cladistics, is a methodology for
inferring the pattern of evolutionary history of a group of organ-
isms, utilizing these apomorphies (Chapter 2).
As cited earlier, cladograms serve as the basis for phyloge-
netic classification. A key component in this classification sys-
tem is the recognition of what are termed monophyletic groups
of taxa. A monophyletic group, or clade, is a group consisting
UNIT I SYSTEMATICS    13
Taxa
Plantae
Magnoliophyta
Liliopsida (Monocots)
Arecales
Arecaceae
Cocos
Cocos nucifera
of a common ancestor plus all (and only all) descendants of
that common ancestor. For example, the monophyletic groups
of the cladogram in Figure 1.9B are circled. A phylogenetic
classification recognizes only monophyletic groups. Note that
some monophyletic groups are included within others (e.g., in
Figure 1.9B the group containing only taxa E and F is included
within the group containing only taxa D, E, and F, which is
included within the group containing only taxa B, C, D, E,
and F, etc.). The sequential listing of clades can serve as a
phylogenetic classification scheme (see Chapter 2).
In contrast to a monophyletic group, a paraphyletic group
is one consisting of a common ancestor but not all descendants
of that common ancestor; a polyphyletic group is one in which
there are two or more separate groups, each with a separate
common ancestor. Paraphyletic and polyphyletic groups distort
the accurate portrayal of evolutionary history and should be
abandoned (see Chapter 2).
Knowing the phylogeny of a group, in the form of a cladogram,
can be viewed as an important end in itself. As discussed earlier,
the cladogram may be used to devise a system of classification,
one of the primary goals of taxonomy. The cladogram also can be
used as a tool for addressing several interesting biological ques-
tions, including biogeographic or ecological history, processes
of speciation, and adaptive character evolution. A thorough
discussion of the principles and methodology of phylogenetic
systematics is discussed in Chapter 2.
WHY STUDY SYSTEMATICS?
The rationale and motives for engaging the field of systemat-
ics are worth examining. For one, systematics is important in
providing a foundation of information about the tremendous
diversity of life. Virtually all fields of biology are dependent on
the correct taxonomic determination of a given study organism,
which relies on formal description, identification, naming, and
classification. Systematic research is the basis for acquiring,
cataloguing, and retrieving information about life’s diversity.
Essential to this research is documentation, through collection
(Chapter 17) and storage of reference specimens, e.g., for plants
in an accredited herbarium (Chapter 18). Computerized data
entry of this collection information is now vital to cataloguing
14  CHAPTER 1 PLANT SYSTEMATICS: AN OVERVIEW

Taxon D
Taxon B

Taxon C

Taxon E

Taxon F
Taxon A
(Present)

Lineage Apomorphy
(for taxon D)

Apomorphies
TIME (for taxa B & C) Lineage
Lineage

Lineage divergence,
followed by speciation
Apomorphy:
Common ancestor Represents evolutionary change:
(of taxon A & taxa BñF) Ancestral state Derived state

(Past)
A
Taxon D
Taxon B

Taxon C

Taxon E

Taxon F
Taxon A

(Present)

Common ancestor
(of taxa B & C)

Common ancestor
(of taxa E & F )
TIME
Common ancestor
(of taxa D–F)

Common ancestor
(of taxa B–F )

Common ancestor
(of taxa A–F )

(Past)
B
Figure 1.9 Example of a cladogram or phylogenetic tree for taxa A–F. A. Cladogram showing lineages and apomorphies, the latter indicated
by thick hash marks. B. Cladogram with common ancestors shown and monophyletic groups (clades) circled.
 UNIT I SYSTEMATICS    15

and retrieving the vast amount of information dealing with
biodiversity (Chapter 18).
Systematics is also an integrative and unifying science. One
of the “fun” aspects of systematics is that it may utilize data
from all fields of biology: morphology, anatomy, embryology/
development, ultrastructure, paleontology, ecology, geography,
chemistry, physiology, genetics, karyology, and cell/molecular
biology. The systematist has an opportunity to understand all
aspects of his/her group of interest in an overall synthesis of
what is known from all biological specialties, with the goal
being to understand the evolutionary history and relationships
of the group.
Knowing the phylogeny of life can give insight into other
fields and have significant practical value. For example, when
a species of Dioscorea, wild yam, was discovered to possess
steroid compounds (used first in birth control pills), exami-
nation of other closely related species revealed species that
contained even greater quantities and diversity of these com-
pounds. Other examples corroborate the practical importance
of knowing phylogenetic relationships among plant species.
The methodology of phylogenetics is now an important part
of comparative biology, used by, for example, evolutionary
ecologists, functional biologists, and parasitologists, all of
whom need to take history into account in formulating and
testing hypotheses.
The study of systematics provides the scientific basis for
defining or delimiting species and infraspecific taxa (subspe-
cies or varieties) and for establishing that these are distinct
from other, closely related and similar taxa. Such studies are
especially important today in conservation biology (Chapter
19). In order to determine whether a species or infraspecific
taxon of plant is rare or endangered and warrants protection,
one must first know the limits of that species or infraspecific
taxon. In addition, understanding the history of evolution and
geography may aid in conservation and management decisions,
where priorities must be set as to which regions to preserve.
Finally, perhaps the primary motivation for many, if not
most, in the field of systematics has been the joy of exploring
the intricate complexity and incredible diversity of life. This
sense of wonder and amazement about the natural world is
worth cultivating (or occasionally rekindling). Systematics
also can be a challenging intellectual activity, generally requir-
ing acute and patient skills of observation. Reconstruction of
phylogenetic relationships and ascertaining the significance of
those relationships can be especially challenging and rewarding.
But today we also face a moral issue: the tragic and irrevocable
loss of species, particularly accelerated by rampant destruction
of habitat, such as deforestation in the tropics. We can all try
to help, both on a personal and a professional level. Systemat-
ics, which has been called simply “the study of biodiversity,”
is the major tool for documenting that biodiversity and can be
a major tool for helping to save it. Perhaps we can all consider
reassessing our own personal priorities in order to help conserve
the life that we study.

REVIEW QUESTIONS

PLANTS
1. What is a “plant”? In what two conceptual ways can the answer to this question be approached?
2. What are the three major groups of life currently accepted?
3. Name and define the mechanism for the evolution of chloroplasts.
4. Name some chlorophyllous organismal groups that have traditionally been called “plants” but that evolved or acquired
chloroplasts independently.
5. Draw a simplified cladogram showing the relative relationships among the green plants (Chlorobionta/Viridiplantae), land plants (em-
bryophytes), vascular plants (tracheophytes), seed plants (spermatophytes), gymnosperms, and angiosperms (flowering plants).
6. Why are land plants treated as equivalent to “plants” in this book?
7. List the many ways that plants are important, both in evolution of life on earth and in terms of direct benefits to humans.

SYSTEMATICS
8. What is systematics and what is its primary emphasis?
9. Define biological evolution, describing what is meant both by descent and by modification.
10. What is a lineage?
11. Name and define the units that undergo evolutionary change.
12. What are the two major mechanisms for evolutionary change?
13. What is a functional feature that results in increased survival or reproduction called?
14. Name and define the four components of taxonomy.
15. Define character and character state.
16  CHAPTER 1 PLANT SYSTEMATICS: AN OVERVIEW

16. Give one example of a character and character state from morphology or from some type of specialized data.
17. What is a dichotomous key? A couplet? A lead?
18. What is a scientific name?
19. Define binomial and indicate what each part of the binomial is called.
20. What is the difference between rank and taxon?
21. What is the plural of taxon?
22. Name the two main ways to classify organisms and describe how they differ.
23. Define phylogeny and give the name of the branching diagram that represents phylogeny.
24. What does a split, from one lineage to two, represent?
25. Name the term for both a preexisting feature and a new feature.
26. What is phylogenetic systematics (cladistics)?
27. What is a monophyletic group or clade? A paraphyletic group? A polyphyletic group?
28. For what can phylogenetic methods be used?
29. How is systematics the foundation of the biological sciences?
30. How can systematics be viewed as unifying the biological sciences?
31. How is systematics of value in conservation biology?
32. Of what benefit is plant systematics to you?

EXERCISES

1. Obtain definitions of the word plant by asking various people (lay persons or biologists) or looking in reference sources, such as
dictionaries or textbooks. Tabulate the various definitions into classes. What are the advantages and disadvantages of each?
2. Take a day to note and list the uses and importance of plants in your everyday life.
3. Pick a subject, such as history or astronomy, and cite how the principles of taxonomy are used in its study.
4. Do a Web search for a particular plant species (try common and scientific name) and note what aspect of plant biology each
site covers.
5. Peruse five articles in a systematics journal and tabulate the different types of research questions that are addressed.

REFERENCES FOR FURTHER STUDY

GENERAL
Daly, M., P. S. Herendeen, R. P. Guralnick, M. W. Westneat, and L. McDade. 2012. Systematics agenda 2020: The mission evolves. Systematic
Biology 61:549-552.
Parfrey, L. W., J. Grant, Y. I. Tekle, E. Lasek-Nesselquist, H. G. Morrison, M. L. Sogin, D. J. Patterson, and L. A. Katz. 2010. Broadly sampled
multigene analyses yield a well-resolved eukaryotic tree of life. Systematic Biology 59:518–533.
Reaka-Kudla, M. L., D. E. Wilson, and E. O. Wilson (eds.). 1997. Biodiversity II: Understanding and Protecting Our Biological Resources.
Joseph Henry Press, Washington, DC.
Sauquet, H. and S. W. Graham. 2016. Planning the future of plant systematics: Report on a special colloquium at the Royal Netherlands
Academy of Arts and Sciences. American Journal of Botany 103:2022-2027.
Simpson, B. B., and M. C. Ogorzaly. 2001. Economic Botany: Plants in Our World. 3rd ed. McGraw-Hill, New York.
Stevens, P. F. 1994. The Development of Biological Systematics: Antoine-Laurent de Jussieu, Nature, and the Natural System. Columbia
University Press, New York.
Systematics Agenda 2000: Charting the Biosphere. 1994. Produced by Systematics Agenda 2000. Wilson, E. O. (ed.), and F. M. Peter (assoc.
ed.). 1988. Biodiversity. National Academy Press, Washington, DC.
Williams, T. A., P. G. Foster, C. J. Cox, and T. M. Embley. 2018. An archaeal origin of eukaryotes supports only two primary domains of life.
Nature 504:231-236.

OTHER PLANT SYSTEMATICS TEXTBOOKS

Judd, W. S., C. S. Campbell, E. A. Kellogg, P. F. Stevens, and M. J. Donoghue. 2015. Plant Systematics: A Phylogenetic Approach, 4th edi-
tion. Sinauer Associates, Sunderland, Mass.
Singh, G. 2004. Plant Systematics: An integrated approach. Science Publishers, Enfield, N.H.
Woodland, D. W. 2009. Contemporary Plant Systematics, 4th edition. Andrews University Press, Berrien Springs, Mich.
Walters, D. R., and D. J. Keil. 1996. Vascular Plant Taxonomy, 4th edition. Kendall/Hunt Publishing Co., Dubuque, Iowa.
 2
TAXONOMY AND
PHYLOGENETIC SYSTEMATICS
TAXONOMY. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
PHYLOGENETIC SYSTEMATICS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
TAXA AND CHARACTERS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Taxon Selection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20
Description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20
Character Selection and Definition . . . . . . . . . . . . . . . . . . . . . . . . . . .20
Character State Discreteness . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21
Character Correlation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21
Homology Assessment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21
Transformation Series and Polarity . . . . . . . . . . . . . . . . . . . . . . . . . . .22
Character Weighting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .23
Character Step Matrix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .23
Character × Taxon Matrix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24
FEATURES OF CLADOGRAMS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Apomorphies as Evolutionary Novelties. . . . . . . . . . . . . . . . . . . . . . .24
Recency of Common Ancestry. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24
Monophyly. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .27
Unrooted Trees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .28
Polytomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .28
Reticulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .28
Taxon Selection and Polymorphic Characters. . . . . . . . . . . . . . . . . . .30
TAXONOMY
As introduced in Chapter 1, taxonomy is defined as the field
of science dealing with description, identification, nomencla-
ture, and classification. By itself, plant taxonomy is not con-
cerned with the evolutionary relationships of a group of
species, which is in the realm of systematics. Yet, taxonomic
research is still a valid and vibrant field of endeavor. Almost
all of our named species have been and continue to be defined
based on taxonomic studies, generally with no direct knowl-
edge of evolutionary relationships, although closeness of
relationship may certainly be inferred from similarities noted
from those studies.
Taxonomic research generally entails determining if a par-
ticular set of populations, generally studied from specimens,
is discrete enough from others to warrant its own taxonomic
PLANT SYSTEMATICS
https://doi.org/10.1016/B978-0-12-812628-8.50002-X, Copyright © 2019 Elsevier Inc. All rights reserved.
CLADOGRAM CONSTRUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
Maximum Parsimony. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .30
Outgroup Comparison . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .32
Ancestral versus Derived Characters . . . . . . . . . . . . . . . . . . . . . . . . .33
Consensus Trees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .34
Long Branch Attraction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .34
Maximum Likelihood. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .34
Bayesian Inference. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .38
Coalescent Methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .38
Measures of Homoplasy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .39
Cladogram Robustness. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .39
CLADOGRAM ANALYSIS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Phylogenetic Classification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .40
Character Evolution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .43
Biogeography, Ecology, Dating Methods, and Speciation Rates. . . . .43
Ontogeny and Heterochrony . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .47
REVIEW QUESTIONS. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
EXERCISES. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
REFERENCES FOR FURTHER STUDY. . . . . . . . . . . . . . . . . . . . . . . . . . . 51
PHYLOGENY COMPUTER PROGRAMS . . . . . . . . . . . . . . . . . . . . . . . . 52
status, e.g., as a separate species, infraspecies (subspecies,
variety, or form), or higher ranked taxon. A taxonomic study
may include a clarification of already named taxa, or may
likely entail the formal naming of a new taxon. Comparisons
are made between and among the taxa of study with respect
to overall similarity, generally using morphological features
(see Chapter 9), but other types of features may be assessed
as well (Chapters 10–13). These comparisons may be qualita-
tive and/or quantitative; the latter often include measure-
ments that are analyzed by various statistical methods (e.g.,
Figure 2.1A; see Appendix 4). Illustrations or photographs
are often used to document the distinctiveness of a new taxon
(Figure 2.1B). Any new taxon name must be validly pub-
lished according to specified rules (see Chapter 16,
Nomenclature and Botanical Names). Proposing a new taxo-
nomic entity includes all of the components of the field:
17
 18  CHAPTER4
  2 TAXONOMY AND PHYLOGENETIC SYSTEMATICS
4
 
lepida
 
lepida
 
3
  m
 var.
 grandi6lora
 
3
  m
 var.
 grandi6lora
 
m
 var.
 micrantha
 
m
 var.
 micrantha
 
4
  2
 
E. lepida
2
 
lepida
 
E. m. var. micrantha 3
 
3
  m
 1
 
var.
 grandi6lora
  lepida
 
1
  var. pseudolepida
E. m. 2.5
 
m
 var.
 micrantha
  m
 var.
 grandi5lora
 

2   0   2   m  var.  micrantha  
-­‐4   -­‐3   -­‐2   0   -­‐1   0   1   2   3  
3   -­‐2   -­‐1   0   1   2   3  
1.5  
1   -­‐1  
-­‐1  
1  

0   -­‐2  
0.5  
-­‐2  
-­‐1   0   1   2   3  
0  
-­‐1   -­‐5   -­‐4   -­‐3   -­‐3   -­‐2   -­‐1   0   1   2   3   4  
-­‐3  
-­‐0.5  

-­‐2   -­‐4  
-­‐4   -­‐1  

A B
-­‐5   -­‐1.5  
-­‐3  
-­‐5    
-­‐2    
 
-­‐4  
Figure 2.1 Examples of results of taxonomic studies. A. Principal components analysis, a statistical procedure demonstrating the
-­‐5  
distinctiveness of a new plant variety (from Simpson et al. 2016. Madroño 63: 39–54). B. Line drawings of vegetative and floral parts,
illustrating distinctiveness from closely related   taxa (from Dodero and Simpson 2012. Madroño 59:223-229). Courtesy of Madroño.

describing the plant, suggesting a means of identification often referred to simply as “branches.” Lineages represent the
(usually providing a taxonomic key), formally naming it, and sequence of ancestral-descendant populations through time,
designating, as part of that name, its classification within ultimately denoting descent. (The term “lineage” is treated
higher ranked taxa. here as a single branch; “clade” is defined as a given common
Taxonomic research will continue to provide the founda- ancestor plus all descendants, including two or more lineages,
tion of our knowledge of the biodiversity of life. It can stand essentially equivalent to a monophyletic group.)
alone, or it may be viewed as a starting point for future phy- Evolution may occur within lineages over time and is rec-
logenetic studies that aim to assess more precise species ognized as a change from a preexisting ancestral (also called
delimitations or to elucidate the evolutionary relationships of plesiomorphic) condition to a new, derived (also called apo-
the taxa of study. morphic) condition. The derived condition, or apomorphy,
represents an evolutionary novelty. As seen in Figure 2.2A,
an apomorphy that unites two or more lineages is known as a
PHYLOGENETIC SYSTEMATICS synapomorphy (syn, together); one that occurs within a
single lineage for a single terminal taxon is called an autapo-
Also as introduced in the previous chapter, systematics is the morphy (aut, self). Either may be referred to simply as an
field that is inclusive of taxonomy but having the primary apomorphy, a convention used throughout this book. (Note
objective of elucidating phylogeny, the evolutionary history that many people use synapomorphy in this general way.)
or pattern of descent of a group of organisms. Phylogenetic Any branching of the cladogram represents lineage diver-
systematics, or cladistics, is that specific branch of systemat- gence or diversification, the formation of two separate lin-
ics concerned with inferring phylogeny. Ever since Darwin eages from one common ancestor. The two lineages could
laid down the fundamental principles of evolutionary theory, diverge into what would be designated separate species, the
a major goal of the biological sciences has been the elucida- process of forming two species from one termed speciation.
tion of the pattern of life’s history of descent. This phylogeny The point of divergence of one clade into two (where the
of life, visualized as a branching pattern, can be inferred by most recent common ancestor of the two divergent clades is
an analysis of characters from living or fossil organisms, uti- located) is termed a node; the region between two nodes is an
lizing phylogenetic principles and methodology. internode (Figure 2.2A). Cladograms may be represented in
Recall that a phylogeny is commonly represented in the different ways. Figure 2.2B shows the same cladogram as in
form of a cladogram, or phylogenetic tree, a branching dia- Figure 2.2A, but shifted 90° clockwise and with the lineages
gram that conceptually represents an estimate of phylogeny drawn perpendicular to one another and of a length reflective
(Figure 2.2). The lines of a cladogram are known as lineages, of the number of apomorphic changes. These two representa-
 UNIT I SYSTEMATICS    19

A B C D E F

Lineage Apomorphy
(autapomorphy
for taxon D) Node

Apomorphies Lineage
(synapomorphies
for taxa B & C)
Internode
TIME Apomorphy
(synapomorphy for taxa D, E, F)

Evolutionary divergence,
followed by speciation
Apomorphy: represents evolutionary change:
Common ancestor ancestral state derived state
(of taxon A & taxa B–F)

A
A

B D

Figure 2.2 A. Example of a cladogram or phylogenetic tree for taxa A–F, with apomorphies indicated by thick hash marks. See text for
explanation of terms. B. Same cladogram topology but drawn horizontally, with branch lengths scaled to number of apomorphic changes.

tions of a cladogram have the same topology, which is the
structure of the branching diagram, i.e., how lineages, includ-
ing those terminating in taxa, are connected together.
Cladograms have an implied, but relative, time scale. For
example, in Figure 2.2A, the common ancestor giving rise to
taxa E and F occurred later in time than that giving rise to
taxa D, E, and F, but we do not know when the lineage split
at these nodes occurred or how long the lineages are in terms
of real time. The term phylogram is often used for a clado-
gram that has an estimated absolute time scale, such that
nodes and branch lengths are calibrated and correspond more
closely to real elapsed time. (See later discussion.)
Why study phylogeny? Elucidating the pattern of descent,
in the form of a cladogram, can be viewed as an important
end in itself. The branching pattern derived from a phyloge-
netic analysis may be used to infer the collective evolutionary
changes that have occurred in ancestral/descendant popula-
tions through time. Thus, a knowledge of phylogenetic rela-
tionships may be invaluable in understanding structural or
DNA sequence evolution as well as in gaining insight into the
possible functional, adaptive significance of hypothesized
evolutionary changes. The cladogram can also be used to
classify life in a way that directly reflects evolutionary his-
tory. Cladistic analysis may also serve as a tool for inferring
biogeographic and ecological history, assessing evolutionary
processes such as species diversification rates, and making
decisions in the conservation of threatened or endangered
species (see Chapter 19).
The principles, methodology, and applications of phyloge-
netic analyses are described in the remainder of this chapter.
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 CHAPTER XIII
TORONTO—THE CITY OF PUBLIC
OWNERSHIP

Said an American whom I met in Toronto the other day:

“I don’t care for this place; it’s too much like home. When I travel
I want to see something different.”
I don’t know just what this man hoped to find here in the second
largest city in Canada. I fear that he expected to find Toronto so
inferior that he would be able to indulge in some boasting at the
expense of the Canadians. If so, he came to the wrong place, for,
judged by American standards, Toronto is thoroughly alive, first
class, and up-to-date.
Located on the north shore of Lake Ontario, and the political and
commercial capital of Ontario province, Toronto is the “Chicago of
Canada.” It is larger than Buffalo or San Francisco, and nearly as big
as Los Angeles. It is the greatest live-stock market of all Canada,
and the chief butcher shop of the Dominion. Like Chicago, it is on the
route of the transcontinental railroad lines. It is the centre of tourist
travel to Niagara Falls, the Thousand Islands, and the vacation lands
of the North. It supplies the mines, the mills, and the farms of a
region rich in natural resources, and fast becoming as highly
industrialized as New England. Ontario does more than half of the
manufacturing of Canada, and one third of the factories of the
province are located in Toronto. Seven of the great chartered banks
of the Dominion have their home offices here, and the city is second
only to Montreal in its financial strength.
In Toronto, I find myself again in a city of twenty-story
skyscrapers, big department stores, and American “hustle.” It is, I
suppose, because it does not seem “foreign” that visitors from the
States find this city disappointing. The people are mostly of British
extraction, and, unlike Montreal, there are but few French, and
comparatively few Catholics.
The city was founded by Tories from New York just after our
Revolutionary War, and it soon became the capital of Upper Canada.
Our soldiers burned it once and captured it twice during the War of
1812. Its name Toronto, an Indian word meaning “place of meeting,”
was chosen about a century ago. Since then the city has doubled in
population and wealth every fifteen years.
In the residential districts, I saw scores of magnificent homes
that compare favourably with those of any of our large cities. The
town is built entirely of brick, and sixty-seven per cent. of the homes
are occupied by their owners. The residents, all of whom seem to
belong to a boosters’ club, tell me that they have the lowest death
rate but one of any city of five hundred thousand population in North
America, and that they have fewer deaths from tuberculosis than
anywhere else on the hemisphere.
I have been out to Queen’s Park to see the provincial
government buildings. Here also is Toronto University, the largest in
the British Empire, with several thousand students of both sexes.
The park is approached by University Avenue, a broad street with
rows of elm and chestnut trees on each side. There are many other
schools and colleges, making Toronto the educational centre of
Ontario.
It was at the University of Toronto that Dr. F. G. Banting
discovered insulin, the new treatment for diabetes obtained from the
pancreas of cattle. Doctor Banting and his associates have since
received many honours. The Dominion government gave him
seventy-five hundred dollars a year for life, so that he might continue
his investigations, while the provincial government has established
him in a chair of medical research at Toronto University paying ten
thousand dollars a year. Instead of commercializing his discovery,
the doctor had it patented in the name of the university, and the
royalties are devoted to research.
 Toronto is about equidistant from New York and Chicago, and
nearly midway between Winnipeg and Halifax. It is only three
hundred and thirty-four miles from Montreal, but between the two
cities are the rapids of the upper St. Lawrence, which so far have
prevented the lake port from becoming accessible to large ocean-
going vessels. The present canals along the St. Lawrence can
accommodate ships up to twenty-five hundred tons, but Toronto has
a plan for bringing ten-thousand-ton steamers to her front door. She
proposes to overcome the rapids and shallows with lakes and
canals, and at the same time utilize the fall of water, which exceeds
two hundred feet, to generate electricity.
The locks of the new and larger Welland Canal around Niagara
Falls have been built thirty feet deep and eight hundred feet long.
When this work is completed, the improvement of the St. Lawrence
will be the only thing needed to make possible the passage of deep-
water ships from the Atlantic to Lake Superior. The St. Lawrence
project has the enthusiastic support of the people of middle Canada,
who see their grain of the future going direct to Liverpool in steamers
loaded at the lake ports. This will cut down the freight charges on
every bushel and add millions to the farmers’ profits.
Our own middle western states also want this Lakes-to-the-
Atlantic waterway, but New York and Buffalo, which have grown fat
on handling freight from the Great Lakes, oppose it. So does
Montreal, for fear that her port might suffer, just as Quebec did when
the St. Lawrence was dredged out from that city to Montreal.
Since the St. Lawrence, for part of its course, borders the state
of New York, the project requires the coöperation of the United
States. The International Joint Commission, representing both
Canada and the United States, after investigation, unanimously
approved it. It recommended the construction of nine locks, thirty-
three miles of canals, forty miles of lake channel, and one hundred
miles of river channel improvements. It also recommended the
construction of a hydro-electric power plant near Ogdensburg, New
York, which, it is estimated, would produce sixteen hundred and forty
thousand horse-power, to be divided between the United States and
Canada. To do all this is comparable to the building of the Panama
Canal. It is estimated that the job will take about eight years and will
cost more than a quarter of a billion dollars.
Meanwhile, Toronto is so sure that the project will be carried out
that she has already spent more than twenty million dollars in getting
her harbour ready for the business she expects in the future. Her
port to-day is like a newly built palace, awaiting the birth of an heir to
the throne, with the king still a bachelor.
An island lying about a mile offshore from the city gives Toronto
a natural harbour. The Harbour Commission has built breakwaters,
channels, and anchorages, and erected piers and berthing spaces to
accommodate fleets of large tonnage vessels. So far, however, these
improvements are used mostly by passenger steamers handling the
summer tourist travel to points on the lakes and along the St.
Lawrence. In part the work of the Harbour Commission has already
paid for itself. It has reclaimed a large tract of marshland along the
eastern shore of the harbour and converted it into industrial sites,
equipped with docks, railroad tracks, and other facilities. There are
now more than eight million dollars’ worth of buildings and machinery
in operation on this area.
The Harbour Commission has developed the lakeside not only
for commercial purposes, but also for the use of the people. West of
the city it has built Sunnyside beach, a half mile long, with
accommodations of all kinds for seventy-five hundred bathers.
Across the harbour is Island Park, another great playground.
Toronto was the first city in the world to establish a municipal
athletic commission to promote sports and outdoor games. Though
baseball is not native to Canada, six thousand Toronto boys played
in regularly organized leagues last summer, and eight thousand
soccer or association football players were listed with the
commission. The city maintains two public golf courses, and there
are country clubs, canoe clubs, and yacht clubs.
Another publicly owned institution in Toronto is an abattoir, built
and operated by the city. Here any cattle dealer or local marketman
may have his animals killed under the most sanitary conditions. The
city owns also its waterworks and has a Hydro-Electric Commission
which furnishes power to its factories and homes at low rates. It has
invested more than two million dollars in grounds and buildings for
the Canadian National Exhibition, held here every September with
an attendance of over a million.
Its street railway system is Toronto’s latest and largest venture in
public ownership. Both the cars and the service are by far the best I
have seen anywhere in Canada, and few of our cities can show
better. The city paid forty-five million dollars for the property, and
within two years it had doubled the single fare area, increased the
mileage twenty-five per cent., built extensions out to the suburbs,
replaced antiquated cars with the newest and best, and speeded up
service. On the main lines, the cars are very large and during rush
hours they are run in twos, coupled together. In the newer cars the
conductor sits perched in a cage in the middle. Passengers enter by
the front door, and if they pass down the aisle to sit in the rear they
pay the conductor as they go by. If they take seats in the front half,
they do not pay their fares until they get up to leave by the door in
the middle. It is interesting to know that the first electric street car in
America was operated in Toronto.
Conservative Montreal looks upon Toronto’s plunges into public
works as the height of folly, and sometimes gives her sister city a
lecture. Replying to such criticism, a local paper said the other day it
supposed Montreal would have every Torontoan go to bed at night
saying these verses:

Oh, let us love our occupations,

Bless the squire and his relations,
Live upon our daily rations,
And always know our proper stations.
 Unlike Montreal and Quebec, Toronto is a city of
sky-scrapers, and the Yonge Street canyon makes the
American visitor feel much at home. Toronto has
hustle, enterprise, and the courage to do whatever it
pleases.
 Flax raising has become important in
southwestern Ontario. The crop competes with the
best Russian product. The Canadians use labour-
saving devices to keep costs down to European
levels.
 But Toronto comes honestly by its independent spirit and bold
experiments for the public welfare. The entire province of Ontario is
imbued with the same tendency. With an area eight times that of
New York, it is, next to Quebec, the largest province of Canada, and
with three million people, mostly of British extraction, excels them all
in population. It is richer in mineral wealth, agricultural resources,
and industrial development than any other province. The people
believe in their future and they show the courage of their convictions
when it comes to going in debt to back public enterprises.
The province owns a railroad that taps the Cobalt silver mining
district and the northern agricultural lands. The main line of the road
extends from the Canadian Pacific and Canadian National lines at
North Bay two hundred and fifty-three miles northward to Cochrane,
where it meets the northernmost of the three transcontinental routes.
A few years ago Ontario increased its expenditure for good
roads from two million dollars a year to nine millions. It created the
Ontario Hydro-Electric Commission, which generates and distributes
more electric power at lower rates than any other similar body in the
world. The province pensions needy mothers, and its public health
service furnishes serums and toxins free to the public.
The Ontario parliament has no upper house, but only a single
chamber to which members are elected by the votes of both men
and women. Not long ago the farmers’ organizations captured
enough seats to give them control of the government.
In Toronto I have seen so many familiar names on the factory
buildings that I have had to ask myself whether I was in a British
Dominion or back in the United States. These are the “branch plants”
of American firms, established here to be inside Canada’s tariff wall
and to get the benefit of the preferential tariffs conceded by Great
Britain and her dominions to Canadian products. From automobiles
to silverware, and from bridge steel to fountain pens, many of our
best known American goods are not only used but made in Canada.
Some of the branch plants bear the same names as at home, but
many adopt for Canadian use designations that give no trace of their
American origin. For example, world-famous corporations that use
“United States” or “American” as part of their names, are the
“Dominion” this or the “Imperial” that in Canada. This policy caters to
the growing movement among the people to buy only goods “made
in Canada.” The American branch-plant system accounts in part for
the resemblance of Toronto to American cities. On every hand I see
electric signs, window displays, and bill-boards bearing the same
appeals to buy the goods that are so extensively advertised at home.
No one knows just how many American branch factories Canada
has, but their number is well over one thousand. There are more
than two hundred in Toronto alone, and as many more elsewhere in
southern Ontario. Montreal has many American branch plants and
American owned enterprises. Its largest hotel belongs to an
American syndicate, and so does my hotel in Toronto.
Americans control nine tenths of the automobile accessory
business of Canada, and in their branch plants they make three fifths
of the Dominion’s automobiles. Practically all of our well known firms
devoted to low and moderate priced cars have big factories in
Canada, and they do practically all their exporting to Australia, New
Zealand, Great Britain, and South Africa through their Canadian
branch plants. This export business amounts to more than twenty-
five million dollars a year, while the cars made here for the Canadian
market represent a value three times as great.
In other lines American capital is conspicuous. Half of the
Canadian rubber factories are owned by Americans, and nearly half
of the meat packing, paint, brass, condensed milk, car construction,
and electrical apparatus industries represent American money.
American controlled concerns do more than half of all the oil refining,
while two hundred and fifty million dollars of our money is invested in
the pulp and paper industry.
Altogether, it is estimated that American investments in
government loans, corporation bonds, land mortgages, and industrial
enterprises amount to two thousand five hundred million dollars. Our
stake in Canada has been increasing rapidly ever since 1914, and
now it nearly equals that of the British. Within a few years it will
probably be much greater. Nearly one sixth of all the money we have
invested in foreign countries is in Canada, and in return for the
capital Canada is now buying from us more than three fifths of her
annual thousand million dollar purchases abroad. In fact, her people
are our best customers; their purchases of us amount to eighty-three
dollars per capita a year as compared with five dollars for all Europe
and fifty cents for China.
 CHAPTER XIV
WATERFALLS THAT WORK FOR THE PEOPLE

How much do you pay for the electric current that lights your
home and runs your heater, vacuum cleaner, and washing machine?
In Washington I am charged ten cents a kilowatt hour, and unless
you are especially favourably located I venture your bills are figured
at about the same rate. To be sure, the monthly expense is not great,
but wouldn’t you feel free to use more electricity if your bills were cut
down one half or two thirds?
That is just what has happened to hundreds of thousands of
people in southern Ontario, and I have been devoting the last couple
of days to finding out how it was done. I have made a special trip
from Toronto to Niagara Falls, and am now writing almost in sight of
those mighty waters. I have visited the world’s biggest power station
and talked here also with the engineers of the Ontario Hydro-Electric
Power Commission, which sells the current to the public at cost.
Again I am impressed with the enterprise and the courage of the
people of this province, and the way they use their government to
get what they want.
Canada’s water-power is, as you know, among her chief assets,
and the Dominion is one of the greatest water-power countries on
earth. Although the United States has more hydraulic power
available for development, yet in proportion to her population
Canada actually uses three times as much as we do. She has in her
rivers and streams a total of about eighteen million low-water
available horse-power, of which about one sixth is now being used.
Quebec and Ontario possess between them nearly two thirds of the
total water-power resources, of which Ontario has the larger amount
developed, and Quebec the more in reserve. These two provinces,
like the eastern United States, contain also the greater part of
Canada’s industries, seventy per cent. of which are now run by
water-power. This cheap power is one of the principal reasons why
Ontario and Quebec do most of Canada’s manufacturing and have
so many American branch plants.
Ontario’s biggest single water-power is in the waters of Niagara
Falls. As you know, the Niagara River connects Lake Erie and Lake
Ontario. It forms also a part of the boundary between the United
States and Canada. For many years commercial power companies
have operated on both sides of the Falls. By treaty, Canada and the
United States have limited the commercial use of the Falls, and have
fixed the amount of water that can be diverted from them at twenty
thousand cubic feet a second on the American side, and thirty-six
thousand feet on the Canadian side. This apportionment was made
because the major part of the Falls is on Canada’s side of the
boundary, and much power is imported by the United States from
Canada. The engineers say that if the governments will let them they
can divert much more of the water in such a way that the beauty of
the Falls will not be impaired.
The Ontario Hydro-Electric Power Commission has
revolutionized the situation on the Canadian side. The Falls have
been put to work directly for the people, private corporations and
profits have been eliminated, and, through the building of a giant
new power station, the mighty forces of the falling waters have been
made more productive than ever before. The Commission is
distributing Niagara power to points two hundred and fifty miles
away, and in addition is operating more than twenty additional power
producing stations. It has built up a super-power system that covers
southern Ontario. It also supplies power at the head of the Great
Lakes. Through it more than three hundred cities, towns, and smaller
municipalities are supplying themselves with power “at cost.”
The Commission is the world’s largest publicly owned power
enterprise, having assets worth two hundred and fifty million dollars.
It is claimed that nowhere else on earth do so many people, spread
over so large a territory, enjoy such low-cost electricity as is the case
under Ontario “Hydro.” It distributes six hundred and fifty thousand
horse-power in electrical energy, and, when all its present projects
are in full operation, the daily output will be more than a million
horse-power. Canada’s coal bill would be three hundred million
dollars a year more if “Hydro” power were produced by fuel from
mines.
I can make you see in a flash just what all this means to the
people. Below the mists of Niagara Falls the river is crossed by the
Railway Arch Bridge. Half this bridge is lighted by an American
company, and the other half by the current from the Canadian side.
The lighting load for the lamps is the same in both sections, yet the
cost last year of lighting the Canadian portion was one hundred and
ten dollars, while the American company charges at its regular rates
totalled two hundred and forty. Now you know why citizens in Ontario
go to the polls and vote their municipalities into partnership with
“Hydro,” as the Commission is popularly called.
The “Hydro” was created because the towns of southern Ontario
felt that they were not getting the full benefit of Nature’s great power
station at their doors. They then secured a charter from the
provincial legislature at Toronto to form a partnership for the purpose
of buying power and selling it to themselves. The Ontario Hydro-
Electric Commission was created to handle the business. In 1910 it
began operations by distributing one thousand horse-power,
purchased under contract from a commercial company at Canadian
Niagara. Four years later it was selling seventy-seven thousand
horse-power, and in 1917 it purchased outright the company from
which it had been buying current. Meanwhile, more and more cities
and towns were joining the partnership, additional power stations
were being bought and built to supply the growing demand, and as
consumption rose rates went down. The provincial authorities are
now talking about a shortage of power in the near future unless the
scheme for building dams and canals along the St. Lawrence is
started at once.
I am told the operations of “Hydro” have not cost the taxpayers
of Ontario a cent in interest charges or capital investment. The whole
scheme actually pays for itself as it goes. This is how it is done: The
provincial government acts as banker for the Commission, loaning it
money with which to build power stations and transmission lines.
These loans are covered by the pledges, in the form of bond issues,
of the cities, through the Commission, to meet the interest and make
repayment of the capital investment. Each city issues twenty-year
bonds for its local central station and distributing system. Payments
on both interest and principal are met out of each year’s receipts, so
that eventually the entire power system will be free of debt. Nearly
fifty cities, in fact, are already in the clear on their investment, and
each year brings others to the same situation.
The basic principle of “Hydro” is a partnership of municipalities to
obtain power at cost. The Commission makes all the expenditures
necessary to develop power and deliver it to the cities. It determines
the rates at which the cities may re-sell the power to local
consumers. Each year the probable cost of power is estimated for
each city, with allowances for interest, depreciation, reserves, and
contingencies. This fixes the rates for the next twelve months. At the
end of the year, the actual cost is calculated. If the expense has
been more than the rates previously fixed, the cities are called upon
to make up the difference; if the cost has been less, the partners get
rebates on the year’s bills.
When I looked for the directing mind behind all this amazing
development, I quickly found it. It is in the person of Sir Adam Beck,
formerly a box manufacturer of London, Ontario, and the only
chairman the Commission has ever had. He is one of the most
popular figures in all Ontario, and it is largely to the organizing
genius and driving power of his personality that the success of
“Hydro” is due. He sits in Parliament, where he keeps on the alert for
the welfare of the Commission’s work, and manages in person its
relations with the provincial government.
 The Niagara Peninsula, along the north shore of
Lake Erie, is one of the finest fruit-growing districts in
the world. Among other advantages the farmers here
have cheap power from the Ontario “Hydro”.
 The big ditch that feeds the giant power station
was cut through solid rock for miles and carries a
stream of water thirty-nine feet deep and fifty feet
wide, or more than the flow of a river.
 A bucket of water dropped down this cliff to the
Niagara River would strike with the force of thirty
horse-power. Imagine 20,000 buckets dropped every
second and you have the capacity of this 600,000
horse-power station.
The cold facts about “Hydro” make it easy for Sir Adam to
demonstrate its achievements. In his home town of London, for
example, where formerly the monthly consumption of electricity
averaged less than twenty kilowatt hours for each household,
seventy-five kilowatt hours are now used. To an extent formerly
undreamed of, power has been put into the homes of the people,
and they are using electric appliances in far greater number than
persons of similar circumstances in the United States. Electric
cooking stoves are being installed in southern Ontario at the rate of
one thousand a month. Workingmen’s wives have toasters, electric
washers, electric fans, electric heaters, curling irons, and everything
else the appliance companies can devise.
The Commission is especially proud of the way it has taken
electricity out to the farms. Private companies usually find rural
extensions too expensive, on account of the long distances between
installations, and the relatively small volume of current used. The
Commission has now enough rural lines to reach from New York to
Atlanta, and it serves as many country homes as there are people
living on farms in Rhode Island. Although the rates are higher than in
the cities, I find that a farmer can light his house and barns, run an
electric stove and household appliances, and a three-horse-power
motor besides, for from six to eight dollars a month. The farmers of
Ontario can afford to use electricity to run their pumps, separators,
churns, milking machines, saw-mills, choppers, and threshers—not
because they are richer than other farmers, but because of low-cost
power.
I have before me a schedule of the rates charged in the twelve
largest cities of the province. For domestic service the average net
cost ranges from 1.3 cents per kilowatt hour to 2.8 cents per kilowatt
hour. The rates vary chiefly with distance. Toronto, ninety miles from
Niagara, pays 2.1 cents per kilowatt hour, while Windsor, two
hundred and fifty miles to the west, opposite Detroit, Michigan, is
charged 2.6 cents. Commercial users, such as stores and office
buildings, pay slightly higher rates, while factories are charged from
$11.75 to $28.66 per horse-power per year. As with commercial
companies, the “Hydro” rates decrease with the amount of power
used. For households the secondary or larger-user rate is nowhere
more than 1.8 cents, and in most places it is less. In Toronto, the
average household electric light bill is less than a dollar and a
quarter a month. At Windsor it is less than a dollar and three
quarters. In the city of London, Ontario, the household consumption
of electricity has under “Hydro” increased more than four hundred
per cent. and the average cost has been reduced to less than one
fourth of the former charges.
 CHAPTER XV
NIAGARA’S GIANT POWER STATION

“Hydro’s” biggest feat in physical construction is the great

development, known as the Queenston Chippewa plant, on the
Niagara River. This station is designed to produce six hundred
thousand horse-power, or about one sixth as much as all the
electrical energy now generated in Canada. Suppose we visit it with
one of the engineers. Stepping into an automobile, we drive first
toward the falls, now partly obscured in the clouds of mist from the
tumbling, roaring, boiling waters. Our way lies through the park the
Canadians have made so that the people may enjoy for all time the
approaches to this monarch among the wonders of Nature.
We stop at Chippewa, at the mouth of the Welland River. This
stream used to empty into the Niagara River above the falls, but to-
day its channel carries the water diverted from the Niagara for the
supply of the power station. The river was deepened and widened
for a distance of four and a half miles, and then a canal was dug
through the remaining eight and a half miles to the site of the plant.
Now we turn back, and as our car passes over one of the numerous
bridges across the big ditch, we look down upon a miniature Panama
Canal, fifty feet wide at water level, and thirty-nine feet deep. In
many places it was cut through hills of rock to a depth of more than
one hundred feet. When the station is operated at full capacity, the
flow of water through the canal is more than twice that of the
Connecticut River.
Seven miles below the falls we come to the power plant. If your
nerves are steady, walk out to the cliff and see where we are. Look
first across the great gorge. Those hills over there are in the state of
New York. See their steep, rocky sides, with van-coloured strata
exposed by the wearing action of the Niagara River through millions
of years. To our left, and hardly a mile away, the high plateau
suddenly drops off; below it are the glistening waters of Lake
Ontario. We are near the end of the escarpment over which Niagara
Falls once plunged into the lake.
Now look straight down; the Niagara River is three hundred feet
below us. From this height it looks like an innocent stream. It is really
a raging torrent in the final throes of its mad struggle to get into Lake
Ontario. The workmen clinging to the sheer, rocky face of the cliff
under our feet, and boring into it with their drills, seem like huge
insects using their stingers. That big block of concrete, resting on a
shelf carved out of the rock and washed by the current, is the power
station. It has the dimensions of an eighteen-story office building,
and part of it is lower than the surface of the river.
The huge steel penstocks, or pipes, through which the water
rushes to the waterwheels below, are set in grooves cut in the rock.
Each of them is twice the diameter of a big dining-room table. The
water enters the penstocks from a great pool fed by the canal and,
dropping, creates this mighty stream of electrical energy, which
surpasses any that can be produced right at the Falls. The reason
for this is that while the total drop in the Niagara River is three
hundred and twenty-seven feet, the height of the Falls is only one
hundred and sixty-seven feet. That is the maximum “head” of water
available to the power stations located right at the Falls. But from
where we stand the drop to the waterwheels is exactly three hundred
and five feet, or nearly twice as great as that at the falls. This station,
in other words, utilizes all except about twenty feet of the difference
in level between Lake Erie and Lake Ontario. It does this by taking in
the water of the Niagara River at Chippewa and carrying it for
thirteen miles nearly to the end of the Niagara escarpment. There is
a fall of only twelve feet in the whole length of the canal.
Every cubic foot of water per second that flows through the
penstocks generates thirty horse-power, as compared with about
sixteen horse-power per foot per second developed by the stations
nearer the falls. The Canadians are thus making the water of