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Etextbook 978 1429219617 Raven Biology of Plants 8th Edition by Ray F Evert
Etextbook 978 1429219617 Raven Biology of Plants 8th Edition by Ray F Evert
The Result of Natural Selection: Adaptation 217 Reproduction and Gene Exchange 259
The Origin of Species 220 Endospores 261
How Does Speciation Occur? 221 Metabolic Diversity 261
ESSAY: Adaptive Radiation in Hawaiian Bacteria 263
Lobeliads 224 Archaea 269
The Origin of Major Groups of Organisms 229 Viruses 270
Viroids: Other Infectious Particles 275
14 Fungi 278
The Importance of Fungi 279
Characteristics of Fungi 281
ESSAY: Phototroprism in a Fungus 285
Microsporidians: Phylum Microsporidia 286
Chytrids: A Polyphyletic Group of Fungi
with Flagellated Cells 287
Zygomycetes: A Polyphyletic Group
of Filamentous Fungi 288
Glomeromycetes: Phylum
Glomeromycota 290
Ascomycetes: Phylum Ascomycota 291
Basidiomycetes: Phylum Basidiomycota 295
ESSAY: Predaceous Fungi 303
Symbiotic Relationships of Fungi 306
SECTION 4 DIVERSITY 233 ESSAY: From Pathogen to Symbiont:
Fungal Endophytes 307
12 Systematics: The Science of
Biological Diversity 234
15 Protists: Algae and Heterotrophic
Taxonomy: Nomenclature and
Protists 317
Classification 234
Ecology of the Algae 320
ESSAY: Convergent Evolution 239
ESSAY: Algae and Human Affairs 321
Cladistics 239
ESSAY: Red Tides/Toxic Blooms 323
Molecular Systematics 240
Euglenoids 324
ESSAY: Google Earth: A Tool for
Cryptomonads: Phylum Cryptophyta 324
Discovering and Protecting
Haptophytes: Phylum Haptophyta 326
Biodiversity 241
Dinoflagellates 327
The Major Groups of Organisms: Bacteria,
ESSAY: Coral Reefs and Global Warming 329
Archaea, and Eukarya 243
Photosynthetic Stramenopiles 330
Origin of the Eukaryotes 247
Red Algae: Phylum Rhodophyta 340
The Protists and Eukaryotic Kingdoms 248
Green Algae 345
Life Cycles and Diploidy 250
Heterotrophic Protists 358
18 Gymnosperms 430
Evolution of the Seed 430
Progymnosperms 432
Extinct Gymnosperms 433
Living Gymnosperms 435
Phylum Coniferophyta 437 SECTION 5 The Angiosperm Plant Body:
Other Living Gymnosperm Phyla: Structure and
Cycadophyta, Ginkgophyta, Development 525
and Gnetophyta 448 22 Early Development of the Plant Body 526
Formation of the Embryo 526
19 Introduction to the Angiosperms 457 The Mature Embryo 530
Diversity in the Phylum Anthophyta 457 Seed Maturation 532
The Flower 460 Requirements for Seed Germination 532
The Angiosperm Life Cycle 465 ESSAY: Wheat: Bread and Bran 533
ESSAY: Hay Fever 475 From Embryo to Adult Plant 534
20 Evolution of the Angiosperms 477 23 Cells and Tissues of the Plant Body 538
Angiosperm Ancestors 477 Apical Meristems and Their
Time of Origin and Diversification Derivatives 538
of the Angiosperms 478 Growth, Morphogenesis, and
Phylogenetic Relationships of the Differentiation 539
Angiosperms 478 Internal Organization of the Plant
Evolution of the Flower 482 Body 541
Evolution of Fruits 492 Ground Tissues 541
CONT ENTS xi
Phosphorus and the Phosphorus Cycle 700 Nutrient and Material Cycling
Human Impact on Nutrient Cycles Interactions between Organisms—
and Effects of Pollution 701 beyond Simple Trophic Relationships
Soils and Agriculture 702 Development and Change of Communities
ESSAY: Halophytes: A Future Resource? 703 and Ecosystems
ESSAY: Compost 704
Plant Nutrition Research 704 32 Global Ecology
ESSAY: The Water Cycle 705 Life on the Land
Rainforests
30 The Movement of Water and Solutes Deciduous Tropical Forests
in Plants 708 Savannas
Movement of Water and Inorganic
Deserts
Nutrients through the Plant Body 709
Grasslands
ESSAY: Green Roofs: A Cool Alternative 714
Temperate Deciduous Forests
Absorption of Water and Ions by Roots 717 Temperate Mixed and Coniferous Forests
Assimilate Transport: Movement of
Mediterranean Scrub
Substances through the Phloem 722
The Northernmost Forests—Taiga
and Boreal Forest
Arctic Tundra
A Final Word
Glossary G–1
Index I–1
SECTION 7 ECOLOGY
On the Web: www.whfreeman.com/raven8e
31 The Dynamics of Communities
and Ecosystems
Ecosystem Energetics—Trophic Structure
preface
• Chapter 10 (Recombinant DNA Technology, Plant The cedar-apple rust fungus alternates between two hosts, cedar
Biotechnology, and Genomics)—updates material on the and apple trees, causing damage to apple harvests (page 278).
xiv PR EF A CE
The Chinese brake fern removes arsenic from contaminated soils • Chapter 27 (Regulating Growth and Development: The
(page 391). Plant Hormones)—expanded discussions of the role of auxin
PRE FACE xv
in vascular differentiation and on hormone receptors and online at www.whfreeman.com/raven8e. Fully illustrated,
signaling pathways for the plant hormones; new discussions these chapters have been thoroughly updated by Paul
of brassinosteroids as a major class of plant hormones and Zedler of the University of Wisconsin, Madison
of strigolactones, which interact with auxin in regulating
apical dominance
Media and Supplements
• Chapter 28 (External Factors and Plant Growth)— Companion Web Site
extensive revision of gravitropism, circadian rhythms, floral www.whfreeman.com/raven8e
stimulus, and thigmonastic movements; new discussion
of genes and vernalization, as well as hydrotropism, For students, this free interactive Web site includes:
phytochrome-interaction factors (PIFs), and the shade-
avoidance syndrome; new essay on “Doomsday Seed Vault:
Securing Crop Diversity”
• Interactive study aids to help foster understanding
of important concepts from the text, including
multiple-choice quizzes, flashcards, plus interactive
• Chapter 29 (Plant Nutrition and Soils)—discusses figures and tables
strategies involving nitrogen uptake by plants, along with
the new topics of beneficial elements, determinate and
indeterminate nodules, and strategies by plants for the
• Animations and live-action videos illustrate several
topics from the text for a deeper understanding of the
acquisition of phosphate; essay added on “The Water more difficult concepts
Cycle”
• Two ecology chapters in easily downloadable and
printable PDF format, fully illustrated and thoroughly
updated with major contributions by Paul Zedler,
University of Wisconsin, Madison.
Preparation Guide for Laboratory Topics in Botany George Johnson, Arkansas Tech University
Preparation Guide for Laboratory Topics in Botany offers Carolyn Howes Keiffer, Miami University
helpful information on how to cover each topic, the length Jeffrey M. Klopatek, Arizona State University
of time needed to complete the material, the sources of sup- Rebecca S. Lamb, Ohio State University
plies and ordering schedules, how to set up the laboratory, and Monica Macklin, Northeastern State University
ways to guide students in their laboratory work. Written by Carol C. Mapes, Kutztown University of Pennsylvania
Susan E. Eichhorn, University of Wisconsin, Madison; Joy B. Shawna Martinez, Sierra College
Perry, University of Wisconsin, Fox Valley; and Ray F. Evert, Austin R. Mast, Florida State University
University of Wisconsin, Madison. Wilf Nicholls, Memorial University of Newfoundland
Karen Renzaglia, Southern Illinois University
Frances M. Wren Rundlett, Georgia State University
Acknowledgments A. L. Samuels, University of British Columbia
S. E. Strelkov, University of Alberta
We are grateful for the enthusiastic response we have received Alexandru M. F. Tomescu, Humboldt State University
from readers who have used previous editions of Biology of M. Lucia Vazquez, University of Illinois at Springfield
Plants, either in English or in one of the six foreign languages in Justin K. Williams, Sam Houston State University
which it has been published. As always, we have appreciated the Michael J. Zanis, Purdue University
support and recommendations made by teachers who used the
last edition in their courses. We also wish to thank the following
people who provided valuable critiques of chapters or portions We are very grateful to our artist, Rhonda Nass, for the exquisite
of chapters for this edition: paintings that open each section and for her beautifully drawn
artwork. She has worked closely with us through many edi-
Richard Amasino, University of Wisconsin, Madison tions, and we value her ability to interpret our pencil sketches
Paul Berry, University of Michigan and render them into drawings that are instructive and accu-
James Birchler, University of Missouri rate, as well as attractive. We are grateful to Rick Nass, who
Wayne Becker, University of Wisconsin, Madison contributed a number of expertly produced graphs. We also
Clyde Calvin, Portland State University thank Sarah Friedrich and Kandis Elliot, Media Specialists,
Kenneth Cameron, University of Wisconsin, Madison Department of Botany, University of Wisconsin, Madison, for
Nancy Dengler, University of Toronto the preparation of digital images of photomicrographs and her-
John Doebley, University of Wisconsin, Madison barium specimens. Mark Allen Wetter, Collections Manager/
Eve Emshwiller, University of Wisconsin, Madison Senior Academic Curator, and Theodore S. Cochrane, Senior
Thomas German, University of Wisconsin, Madison Academic Curator, both of the Wisconsin State Herbarium,
Thomas Givnish, University of Wisconsin, Madison Department of Botany, University of Wisconsin, Madison, were
Linda Graham, University of Wisconsin, Madison very helpful in selecting and scanning herbarium specimens for
Christopher Haufler, University of Kansas our use.
David Hibbett, Clark University We would especially like to thank Sally Anderson, our
Robin Kurtz, University of Wisconsin, Madison talented developmental editor, who has worked with us for five
Ben Pierce, Southwestern University editions now. We are grateful for her many contributions at
Scott Russell, University of Oklahoma, Norman every stage of the process, from the early stages of planning
Dennis Stevenson, New York Botanical Garden the new edition through the manuscript and proof stages to the
Joseph Williams, University of Tennessee, Knoxville finished book. We have worked well together over the years,
Paul Zedler, University of Wisconsin, Madison and we thank her for her dedication to all aspects of making
this edition the most accurate and accessible of all.
We would also like to thank Richard Robinson, who has
The following people gave us helpful feedback as we planned written the engaging ecology-oriented essays that have been
this edition: added to this edition. The essays are highlighted in the text by
a green leaf, and they cover such topics as green roofs, invasive
Richard Carter, Valdosta State University plants, coral bleaching, the development of biofuels, and the use
Sara Cohen Christopherson, University of Wisconsin, Madison of Google Earth to map and study biodiversity.
Les C. Cwynar, University of New Brunswick The preparation of the eighth edition has involved the
Brian Eisenback, Bryan College collaborative efforts of a large number of talented people at
Karl H. Hasenstein, University of Louisiana at Lafayette W. H. Freeman and Company. Particular thanks go to Peter
Bernard A. Hauser, University of Florida Marshall, Publisher of Life Sciences, whose vision and support
Jodie S. Holt, University of California Riverside have made this new edition possible; to Vivien Weiss, who has
PREF ACE xvii
skillfully managed the production process; to Elyse Rieder and coordinated the complicated illustration program. Our gratitude
Bianca Moscatelli, who enthusiastically tracked down pho- also goes to Debbie Clare, Associate Director of Marketing,
tographs; and to Blake Logan, who applied her design talents who has been tireless in managing the sales and market-
to give this edition a new look. We especially want to thank ing efforts for this edition, and to Susan Wein, Production
Sheridan Sellers for the remarkable work she has done in mak- Coordinator, for her many contributions during the complex
ing up the pages of this book—with her aesthetic and peda- stages of production.
gogical talents, she has worked a miracle in placing the many A great number of people, only some of whom are men-
large and complicated illustrations into a coherent layout. And tioned here, have contributed in many essential ways to this
we are grateful to Linda Strange, our long-time copyeditor revision, and we extend to them our deepest appreciation and
who, with good humor and a steady hand, manages to hold us gratitude.
to a high standard in consistency and accuracy. We also thank
Marni Rolfes, associate editor, who competently handled the Ray F. Evert
day-to-day matters and kept us on track, and Bill Page, who Susan E. Eichhorn
INTRODUCTION
of food. They provide us with fiber for clothing; wood for fur-
chapter outline niture, shelter, and fuel; paper for books (such as the page you
Evolution of Plants are reading at this moment); spices for flavor; drugs for medi-
Evolution of Communities cines; and the oxygen we breathe. We are utterly dependent on
Appearance of Human Beings plants. Plants also have enormous sensory appeal, and our lives
are enhanced by the gardens, parks, and wilderness areas avail-
able to us. The study of plants has provided us with great in-
Evolution of Plants
Life Originated Early in Earth’s Geologic History
Like all other living organisms, plants have had a long history
during which they evolved, or changed, over time. The planet
Earth itself—an accretion of dust and gases swirling in orbit
around the star that is our sun—is some 4.6 billion years old
(Figure 1–1). It is believed that Earth sustained a lethal meteor
bombardment that ended about 3.8 to 3.9 billion years ago.
Vast chunks of rubble slammed into the planet, helping to keep
it hot. As the molten Earth began to cool, violent storms raged,
accompanied by lightning and the release of electrical energy,
and widespread volcanism spewed molten rock and boiling
water from beneath the Earth’s surface.
The earliest known fossils are found in rocks of Western 1–2 The earliest known fossils Obtained from ancient rocks
Australia about 3.5 billion years old (Figure 1–2). These micro- in northwestern Western Australia, these fossilized prokaryotes
fossils consist of several kinds of small, relatively simple fila- are dated at 3.5 billion years of age. They are about a billion years
mentous microorganisms resembling bacteria. About the same younger than the Earth itself, but there are few suitable older
age as these microfossils are ancient stromatolites—fossilized rocks in which to look for earlier evidence of life. More complex
microbial mats consisting of layers of filamentous and other organisms—those with eukaryotic cellular organization—did not
microorganisms and trapped sediment. Stromatolites continue evolve until about 2.1 billion years ago. For about 1.5 billion years,
to be formed today in a few places, such as in the warm, shal- therefore, prokaryotes were the only forms of life on Earth. These
low oceans off the shores of Australia and the Bahamas (see so-called microfossils have been magnified 1000 times.
Chapter 13). By comparing the ancient stromatolites with mod-
ern ones, which are formed by cyanobacteria (filamentous pho-
tosynthetic bacteria), scientists have concluded that the ancient on Mars, for example, whose early history paralleled that of
stromatolites were formed by similar filamentous bacteria. Earth. Strong evidence, first discovered by the Opportunity
Whether life originated on Earth or reached Earth through rover in 2004, indicated that water once flowed across the
space in the form of spores (resistant reproductive cells) or planet, raising the possibility that, at one time, Mars could
by some other means is problematic. Life may have formed have supported life (Figure 1–3). In 2008, the Phoenix Mars
1–1 Life on Earth Of the nine planets in our solar system, 1–3 Life on Mars? This color-enhanced image shows a portion
only one, as far as we know, has life on it. This planet, Earth, is of the Jezero Crater, a 25-mile-wide impact crater on northern
visibly different from the others. From a distance, it appears blue Mars that once held a lake. Claylike minerals (indicated in green)
and green, and it shines a little. The blue is water, the green is were carried by ancient rivers into the lake, forming a delta.
chlorophyll, and the shine is sunlight reflected off the layer of Because clays are able to trap and preserve organic matter,
gases surrounding the planet’s surface. Life, at least as we know it, deltas and lakebeds are promising areas in which to search for
depends on these visible features of Earth. signs of ancient life on Mars.
4 C h a p te r 1 Botany: An Introduction
Lander found water ice in abundance near the surface. More- that all living things share a common ancestor: a DNA-based
over, its instruments monitored a diurnal water cycle: water microbe that lived more that 3.5 billion years ago. Near the end
vapor, originating from the shallow subsurface water ice and of On the Origin of Species, Charles Darwin wrote: “Probably
from water clinging to soil grains, is released into the Martian all the organic beings which have ever lived on this earth have
atmosphere during the morning hours, and at night it condenses descended from some one primordial form, into which life was
and falls out by gravity. Most of the ice crystals evaporate as first breathed.”
they fall through the atmospheric boundary layer, but snowfall
on Mars has been observed. Autotrophic Organisms Make Their Own Food, but
No organic molecules or traces of previous or present bio-
logical activity were detected at the Phoenix landing site. How-
Heterotrophic Organisms Must Obtain Their Food from
ever, one would expect organic molecules to be present in the External Sources
Martian soil, given the steady influx of certain types of mete- Cells that satisfy their energy requirements by consuming the
orites that contain considerable quantities of organic material. organic compounds produced by external sources are known
Meteorites that fall to Earth contain amino acids and organic as heterotrophs (Gk. heteros, “other,” and trophos, “feeder”).
carbon molecules such as formaldehyde. We will continue to A heterotrophic organism is dependent on an outside source of
assume, however, that life on Earth originated on Earth. organic molecules for its energy. Animals, fungi (Figure 1–4),
In 2011, NASA’s Mars Reconnaissance Orbiter satellite and many of the one-celled organisms, such as certain bacteria
found evidence of liquid water flowing down slopes and crater and protists, are heterotrophs.
walls during the warm month on Mars. The liquid is thought to As the primitive heterotrophs increased in number, they
be highly salty and to occur just below the surface, where it is began to use up the complex molecules on which their existence
protected from freezing in the frigid temperatures found on Mars depended—and which had taken millions of years to accumu-
and from evaporating in the planet’s low air pressures. These late. Organic molecules in free solution (that is, not inside a cell)
findings further raise the possibilities of finding life on Mars.
became more and more scarce, and competition began. Under the oceans and lakes reacted with dissolved iron and precipitated
pressure of this competition, cells that could make efficient use of as iron oxides (Figure 1–6). From about 2.7 to 2.2 billion years
the limited energy sources now available were more likely to sur- ago, oxygen began gradually to accumulate in the atmosphere.
vive than cells that could not. In the course of time, by the long, By about 700 million years ago, atmospheric levels of oxygen
slow process of elimination of the most poorly adapted, cells increased markedly, and began to approach modern levels dur-
evolved that were able to make their own energy-rich molecules ing the Cambrian period (570–510 million years ago).
out of simple inorganic materials. Such organisms are called This increase in oxygen level had two important conse-
autotrophs, “self-feeders.” Without the evolution of these early quences. First, some of the oxygen molecules in the outer layer
autotrophs, life on Earth would soon have come to an end. of the atmosphere were converted to ozone (O3) molecules.
The most successful of the autotrophs were those in which When there is a sufficient quantity of ozone in the atmosphere,
a system evolved for making direct use of the sun’s energy— it absorbs the ultraviolet rays—rays highly destructive to living
that is, the process of photosynthesis (Figure 1–5). The earli- organisms—from the sunlight that reaches the Earth. By about
est photosynthetic organisms, although simple in comparison 450 million years ago, the ozone layer apparently protected or-
with plants, were much more complex than the primitive het- ganisms sufficiently so they could survive in the surface layers
erotrophs. Use of the sun’s energy required a complex pigment of water and on the shores, and life emerged on land for the
system to capture the light energy and, linked to this system, a first time.
way to store the energy in an organic molecule.
Evidence of the activities of photosynthetic organisms
has been found in rocks 3.4 billion years old, about 100 mil-
lion years after the first fossil evidence of life on Earth. We can
be almost certain, however, that both life and photosynthetic
organisms evolved considerably earlier than the evidence sug-
gests. In addition, there seems to be no doubt that heterotrophs
evolved before autotrophs. With the arrival of autotrophs, the
flow of energy in the biosphere (that is, the living world and its
environment) came to assume its modern form: radiant energy
from the sun channeled through the photosynthetic autotrophs
to all other forms of life.
20 μm
1–8 Stomata Open stomata on the surface of a Apical meristem
of shoot
tobacco (Nicotiana tabacum) leaf. Each stoma in the aerial
parts of the plant is regulated by two guard cells.
Leaflets
(a) (b)
(e) (f)
the study of cell structure, function, and life histories; genet- mycorrhizal fungi) form important, mutually beneficial sym-
ics, the study of heredity and variation; genomics, the study of biotic relationships with their plant hosts. Virology, bacteriol-
the content, organization, and function of genetic information ogy, phycology (the study of algae), and mycology (the study
in whole genomes; molecular biology, the study of the struc- of fungi) are well-established fields in their own right, but they
ture and function of biological molecules; economic botany, the still fall loosely under the umbrella of botany.
study of past, present, and future uses of plants by people; eth-
nobotany, the study of the uses of plants for medicinal and other
purposes by indigenous peoples; ecology, the study of the rela-
A Knowledge of Botany Is Important for Dealing
tionships between organisms and their environment; and paleo- with Today’s—and Tomorrow’s—Problems
botany, the study of the biology and evolution of fossil plants. In this chapter, we have ranged from the beginnings of life on
Included in this book are all organisms that have tradi- this planet to the evolution of plants and ecosystems to the de-
tionally been studied by botanists: not only plants but also velopment of agriculture and civilization. These broad topics
prokaryotes, viruses, fungi, and autotrophic protists (algae). are of interest to many people other than botanists, or plant bi-
Nonphotosynthetic eukaryotes and protists have tradition- ologists. The urgent efforts of botanists and agricultural scien-
ally been the province of zoologists. Although we do not re- tists will be needed to feed the world’s rapidly growing human
gard algae, fungi, prokaryotes, or viruses as plants, and shall population (Figure 1–12), as discussed in Chapter 21. Modern
not refer to them as plants in this book, they are included here plants, algae, and bacteria offer the best hope of providing a re-
because of tradition and because they are normally considered newable source of energy for human activities, just as extinct
part of the botanical portion of the curriculum, just as botany plants, algae, and bacteria have been responsible for the mas-
itself used to be considered a part of medicine. Moreover, both sive accumulations of gas, oil, and coal on which our modern
prokaryotes (e.g., nitrogen-fixing bacteria) and fungi (e.g., industrial civilization depends. In an even more fundamental
1–12 Growth of the human population Over the last 10,000 years, the human population has grown from
several million to approximately 6.5 billion. A significant increase in the rate of population growth occurred as
a result of the cultivation of plants as crops, and an even more dramatic increase began with the advent of the
Industrial Revolution, which started in the middle of the eighteenth century and continues to the present.
The consequences of the rapid growth of the human population are many and varied. In the United States
and other parts of the developed world, they include not only the sheer numbers of people but also heavy
consumption of nonrenewable fossil fuels and the resulting pollution—both as the fuels are burned and as a
result of accidents such as oil spills at drilling sites and during transport. In less developed parts of the world,
the consequences include malnutrition and, all too often, starvation, coupled with a continuing vulnerability to
infectious diseases. The consequences for other organisms include not only the direct effects of pollution but
also—and most important—the loss of habitat.
12 C h a p te r 1 Botany: An Introduction
sense, the role of plants, along with that of algae and photo- has been seriously depleted by the use of chlorofluorocarbons
synthetic bacteria, commands our attention. As the producers (CFCs), and damaging ultraviolet rays penetrating the depleted
of energy-containing compounds in the global ecosystem, these layer have increased the incidence of skin cancer in people
photosynthetic organisms are the route by which all other liv- all over the world. Moreover, it has been estimated that, by
ing things, including ourselves, obtain energy, oxygen, and the the middle of this century, the average temperature will have
many other materials necessary for their continued existence. increased between 1.5° and 4.5°C due to the greenhouse ef-
As a student of botany, you will be in a better position to assess fect. This global-warming phenomenon—the trapping of heat
the important ecological and environmental issues of the day radiating from the Earth’s surface out into space—is intensi-
and, by understanding, help to build a healthier world. fied through the increased amounts of carbon dioxide, nitrogen
In this second decade of the twenty-first century, it is clear oxides, CFCs, and methane in the atmosphere resulting from
that human beings, with a population of 6.5 billion in 2010 and human activities. And most seriously, a large portion of the
a projected population of 9 billion by 2050, are managing the total number of species of plants, animals, fungi, and micro-
Earth with an intensity that would have been unimaginable a organisms is disappearing during our lifetime—the victims of
few decades ago. Every hour, manufactured chemicals fall on human exploitation of the Earth—resulting in a loss of biodi-
every square centimeter of the planet’s surface. The protec- versity. All of these trends are alarming, and they demand our
tive stratospheric ozone layer formed 450 million years ago utmost attention.
(a) (b)
(a) (b)
1–15 Greening of abandoned industrial sites (a) The High Line in New York City is built on an
abandoned railroad track elevated above a newly resurrected neighborhood of restaurants, galleries,
and shops. Remnants of the original track can be seen among the shrubs, perennials, grasses, and trees
planted along the popular promenade. (b) A recently created wetlands situated on a former airbase
at Magnuson Park in Seattle, Washington, offers a rich habitat of native plants and various species of
wildlife, including dragonflies, frogs, ducks, owls, hawks, shorebirds, and warblers.
of all kinds of plants—as individual species and as components Heterotrophic Organisms Evolved before Autotrophic
of the life-support systems into which we all have evolved— Organisms, Prokaryotes before Eukaryotes, and
depend on our knowledge and ability to critically assess the in- Unicellular Organisms before Multicellular Organisms
formation we are given. Thus, this book is dedicated not only
Heterotrophs, organisms that feed on organic molecules or on
to the botanists of the future, whether teachers or researchers,
other organisms, were the first life forms to appear on Earth.
but also to the informed citizens, scientists, and laypeople alike,
Autotrophic organisms, those that could produce their own food
in whose hands such decisions lie.
by photosynthesis, evolved no less than 3.4 billion years ago.
Until about 2.1 billion years ago, the prokaryotes—archaea and
bacteria—were the only organisms that existed. Eukaryotes, with
larger, much more complex cells, evolved at that time. Multicel-
Summary lular eukaryotes began to evolve at least 650 million years ago,
Photosynthesis Is the Process by Which the Sun’s Energy and they began to invade the land about 450 million years ago.
Is Captured to Form Organic Molecules With the advent of oxygen-producing photosynthesis, in
Only a few kinds of organisms—plants, algae, and some bacte- which water molecules are split and oxygen is released, oxygen
ria—have the capacity to capture energy from the sun and use began to accumulate in the atmosphere. The presence of this
it to form organic molecules by the process of photosynthesis. free oxygen enabled organisms to break down the energy-rich
Almost all life on Earth depends, directly or indirectly, on the products of photosynthesis by aerobic respiration.
products of this process.
Colonization of the Land Was Associated with the
The Chemical Building Blocks of Life Accumulated in the Evolution of Structures to Obtain Water and Minimize
Early Oceans Water Loss
The planet Earth is some 4.6 billion years old. The oldest Plants, which are basically a terrestrial group, have achieved
known fossils date back 3.5 billion years and resemble today’s a number of specialized characteristics that suit them for life
filamentous bacteria. Although the process by which living or- on land. These characteristics are best developed among the
ganisms arose is a matter of speculation, there is general agree- members of the dominant group known as the vascular plants.
ment that life as we know it probably emerged on Earth only Among these features are a waxy cuticle, penetrated by special-
once—that is, all living things share a common ancestor. ized openings known as stomata through which gas exchange
Questions 15
I see increasing reason to believe that the view formed some time
back as to the origin of the Makonde bush is the correct one. I have
no doubt that it is not a natural product, but the result of human
occupation. Those parts of the high country where man—as a very
slight amount of practice enables the eye to perceive at once—has not
yet penetrated with axe and hoe, are still occupied by a splendid
timber forest quite able to sustain a comparison with our mixed
forests in Germany. But wherever man has once built his hut or tilled
his field, this horrible bush springs up. Every phase of this process
may be seen in the course of a couple of hours’ walk along the main
road. From the bush to right or left, one hears the sound of the axe—
not from one spot only, but from several directions at once. A few
steps further on, we can see what is taking place. The brush has been
cut down and piled up in heaps to the height of a yard or more,
between which the trunks of the large trees stand up like the last
pillars of a magnificent ruined building. These, too, present a
melancholy spectacle: the destructive Makonde have ringed them—
cut a broad strip of bark all round to ensure their dying off—and also
piled up pyramids of brush round them. Father and son, mother and
son-in-law, are chopping away perseveringly in the background—too
busy, almost, to look round at the white stranger, who usually excites
so much interest. If you pass by the same place a week later, the piles
of brushwood have disappeared and a thick layer of ashes has taken
the place of the green forest. The large trees stretch their
smouldering trunks and branches in dumb accusation to heaven—if
they have not already fallen and been more or less reduced to ashes,
perhaps only showing as a white stripe on the dark ground.
This work of destruction is carried out by the Makonde alike on the
virgin forest and on the bush which has sprung up on sites already
cultivated and deserted. In the second case they are saved the trouble
of burning the large trees, these being entirely absent in the
secondary bush.
After burning this piece of forest ground and loosening it with the
hoe, the native sows his corn and plants his vegetables. All over the
country, he goes in for bed-culture, which requires, and, in fact,
receives, the most careful attention. Weeds are nowhere tolerated in
the south of German East Africa. The crops may fail on the plains,
where droughts are frequent, but never on the plateau with its
abundant rains and heavy dews. Its fortunate inhabitants even have
the satisfaction of seeing the proud Wayao and Wamakua working
for them as labourers, driven by hunger to serve where they were
accustomed to rule.
But the light, sandy soil is soon exhausted, and would yield no
harvest the second year if cultivated twice running. This fact has
been familiar to the native for ages; consequently he provides in
time, and, while his crop is growing, prepares the next plot with axe
and firebrand. Next year he plants this with his various crops and
lets the first piece lie fallow. For a short time it remains waste and
desolate; then nature steps in to repair the destruction wrought by
man; a thousand new growths spring out of the exhausted soil, and
even the old stumps put forth fresh shoots. Next year the new growth
is up to one’s knees, and in a few years more it is that terrible,
impenetrable bush, which maintains its position till the black
occupier of the land has made the round of all the available sites and
come back to his starting point.
The Makonde are, body and soul, so to speak, one with this bush.
According to my Yao informants, indeed, their name means nothing
else but “bush people.” Their own tradition says that they have been
settled up here for a very long time, but to my surprise they laid great
stress on an original immigration. Their old homes were in the
south-east, near Mikindani and the mouth of the Rovuma, whence
their peaceful forefathers were driven by the continual raids of the
Sakalavas from Madagascar and the warlike Shirazis[47] of the coast,
to take refuge on the almost inaccessible plateau. I have studied
African ethnology for twenty years, but the fact that changes of
population in this apparently quiet and peaceable corner of the earth
could have been occasioned by outside enterprises taking place on
the high seas, was completely new to me. It is, no doubt, however,
correct.
The charming tribal legend of the Makonde—besides informing us
of other interesting matters—explains why they have to live in the
thickest of the bush and a long way from the edge of the plateau,
instead of making their permanent homes beside the purling brooks
and springs of the low country.
“The place where the tribe originated is Mahuta, on the southern
side of the plateau towards the Rovuma, where of old time there was
nothing but thick bush. Out of this bush came a man who never
washed himself or shaved his head, and who ate and drank but little.
He went out and made a human figure from the wood of a tree
growing in the open country, which he took home to his abode in the
bush and there set it upright. In the night this image came to life and
was a woman. The man and woman went down together to the
Rovuma to wash themselves. Here the woman gave birth to a still-
born child. They left that place and passed over the high land into the
valley of the Mbemkuru, where the woman had another child, which
was also born dead. Then they returned to the high bush country of
Mahuta, where the third child was born, which lived and grew up. In
course of time, the couple had many more children, and called
themselves Wamatanda. These were the ancestral stock of the
Makonde, also called Wamakonde,[48] i.e., aborigines. Their
forefather, the man from the bush, gave his children the command to
bury their dead upright, in memory of the mother of their race who
was cut out of wood and awoke to life when standing upright. He also
warned them against settling in the valleys and near large streams,
for sickness and death dwelt there. They were to make it a rule to
have their huts at least an hour’s walk from the nearest watering-
place; then their children would thrive and escape illness.”
The explanation of the name Makonde given by my informants is
somewhat different from that contained in the above legend, which I
extract from a little book (small, but packed with information), by
Pater Adams, entitled Lindi und sein Hinterland. Otherwise, my
results agree exactly with the statements of the legend. Washing?
Hapana—there is no such thing. Why should they do so? As it is, the
supply of water scarcely suffices for cooking and drinking; other
people do not wash, so why should the Makonde distinguish himself
by such needless eccentricity? As for shaving the head, the short,
woolly crop scarcely needs it,[49] so the second ancestral precept is
likewise easy enough to follow. Beyond this, however, there is
nothing ridiculous in the ancestor’s advice. I have obtained from
various local artists a fairly large number of figures carved in wood,
ranging from fifteen to twenty-three inches in height, and
representing women belonging to the great group of the Mavia,
Makonde, and Matambwe tribes. The carving is remarkably well
done and renders the female type with great accuracy, especially the
keloid ornamentation, to be described later on. As to the object and
meaning of their works the sculptors either could or (more probably)
would tell me nothing, and I was forced to content myself with the
scanty information vouchsafed by one man, who said that the figures
were merely intended to represent the nembo—the artificial
deformations of pelele, ear-discs, and keloids. The legend recorded
by Pater Adams places these figures in a new light. They must surely
be more than mere dolls; and we may even venture to assume that
they are—though the majority of present-day Makonde are probably
unaware of the fact—representations of the tribal ancestress.
The references in the legend to the descent from Mahuta to the
Rovuma, and to a journey across the highlands into the Mbekuru
valley, undoubtedly indicate the previous history of the tribe, the
travels of the ancestral pair typifying the migrations of their
descendants. The descent to the neighbouring Rovuma valley, with
its extraordinary fertility and great abundance of game, is intelligible
at a glance—but the crossing of the Lukuledi depression, the ascent
to the Rondo Plateau and the descent to the Mbemkuru, also lie
within the bounds of probability, for all these districts have exactly
the same character as the extreme south. Now, however, comes a
point of especial interest for our bacteriological age. The primitive
Makonde did not enjoy their lives in the marshy river-valleys.
Disease raged among them, and many died. It was only after they
had returned to their original home near Mahuta, that the health
conditions of these people improved. We are very apt to think of the
African as a stupid person whose ignorance of nature is only equalled
by his fear of it, and who looks on all mishaps as caused by evil
spirits and malignant natural powers. It is much more correct to
assume in this case that the people very early learnt to distinguish
districts infested with malaria from those where it is absent.
This knowledge is crystallized in the
ancestral warning against settling in the
valleys and near the great waters, the
dwelling-places of disease and death. At the
same time, for security against the hostile
Mavia south of the Rovuma, it was enacted
that every settlement must be not less than a
certain distance from the southern edge of the
plateau. Such in fact is their mode of life at the
present day. It is not such a bad one, and
certainly they are both safer and more
comfortable than the Makua, the recent
intruders from the south, who have made USUAL METHOD OF
good their footing on the western edge of the CLOSING HUT-DOOR
plateau, extending over a fairly wide belt of
country. Neither Makua nor Makonde show in their dwellings
anything of the size and comeliness of the Yao houses in the plain,
especially at Masasi, Chingulungulu and Zuza’s. Jumbe Chauro, a
Makonde hamlet not far from Newala, on the road to Mahuta, is the
most important settlement of the tribe I have yet seen, and has fairly
spacious huts. But how slovenly is their construction compared with
the palatial residences of the elephant-hunters living in the plain.
The roofs are still more untidy than in the general run of huts during
the dry season, the walls show here and there the scanty beginnings
or the lamentable remains of the mud plastering, and the interior is a
veritable dog-kennel; dirt, dust and disorder everywhere. A few huts
only show any attempt at division into rooms, and this consists
merely of very roughly-made bamboo partitions. In one point alone
have I noticed any indication of progress—in the method of fastening
the door. Houses all over the south are secured in a simple but
ingenious manner. The door consists of a set of stout pieces of wood
or bamboo, tied with bark-string to two cross-pieces, and moving in
two grooves round one of the door-posts, so as to open inwards. If
the owner wishes to leave home, he takes two logs as thick as a man’s
upper arm and about a yard long. One of these is placed obliquely
against the middle of the door from the inside, so as to form an angle
of from 60° to 75° with the ground. He then places the second piece
horizontally across the first, pressing it downward with all his might.
It is kept in place by two strong posts planted in the ground a few
inches inside the door. This fastening is absolutely safe, but of course
cannot be applied to both doors at once, otherwise how could the
owner leave or enter his house? I have not yet succeeded in finding
out how the back door is fastened.