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Forest Ecology
Forest Ecology FIFTH EDITION
Daniel M. Kashian
Wayne State University,
Detroit, USA
Donald R. Zak
University of Michigan,
Ann Arbor, USA
Burton V. Barnes (deceased)

University of Michigan,
Ann Arbor, USA
Stephen H. Spurr (deceased)

University of Texas at Austin,
Austin, USA
This fifth edition first published 2023
© 2023 John Wiley & Sons Ltd
Edition History
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Library of Congress Cataloging-in-Publication Data

Names: Kashian, Daniel M., author. | Zak, Donald R., author. | Barnes,
Burton Verne, 1930-2014, author. | Spurr, Stephen H., 1918-1990, author.
Title: Forest ecology / Daniel M. Kashian, Wayne State University, Detroit,
USA, Donald R. Zak, University of Michigan, Ann Arbor, USA, Burton V.
Barnes (deceased), University of Michigan, Ann Arbor, USA, Stephen H.
Spurr (deceased), University of Texas at Austin, Austin, USA.
Description: Fifth edition. | Hoboken, NJ : Wiley, 2022. | Precedey by:
Forest ecology / Burton V. Barnes ... [et al.]. 4th ed. c1998. |
Includes bibliographical references and index.
Identifiers: LCCN 2022037810 (print) | LCCN 2022037811 (ebook) | ISBN
9781119476085 (paperback) | ISBN 9781119476054 (adobe pdf) | ISBN
9781119476146 (epub)
Subjects: LCSH: Forest ecology.
Classification: LCC QK938.F6 F635 2022 (print) | LCC QK938.F6 (ebook) |
DDC 577.3–dc23/eng/20220831
LC record available at https://lccn.loc.gov/2022037810
LC ebook record available at https://lccn.loc.gov/2022037811
Cover Design: Wiley

Cover Image: © U.S. Forest Service Photo
Set in 9.5/12pt STIXTwoText by Straive, Pondicherry, India

Dedication
We dedicate the 5th edition of Forest Ecology to the co-author of the 2nd and 3rd
editions and the lead author of the 4th edition, Burton V. Barnes (1930–2014). It would
be no easy task to find a more accomplished and humble leader in his field. He excelled
at his science, was a truly beloved teacher, and helped to shape the world view of
thousands of colleagues, friends, students, managers, and scientists alike, all with an
unmatched humor and a love of the natural world.
Burt Barnes was world-renowned as an expert in the ecology of North American aspens
and the ecological classification of forest ecosystems. His professional training was in
forest ecology, botany, and genetics, but he dabbled heavily in glacial geomorphology,
soil science, phytogeography, and woody plant physiology. Perhaps his greatest love,
however, was teaching, especially in the field, which drove his motivation for this
textbook. Generations of students have been touched by his love for the art and science
of teaching field ecology, which they will forever pass on to future generations. We, as
authors of this edition, have been personally and professionally shaped by him as a
mentor, colleague, and friend. His legacy is therefore unending. To him we say, in his
own words, “Thanks for everything you have done—and will do.”
Contents
Prefacexxiii
PA R T 1 Forest Ecology and Landscape Ecosystems
1 Concepts of Forest Ecology 3

Ecology, 4
Landscape Ecosystems, 4
Landscape Ecosystem and Community, 7
Ecosystem Structure and Function, 7
Examples of Landscape Ecosystems, 9
An Approach to The Study of Forest Ecology, 11
Applicability to Forest Management, 12
Suggested Readings, 13
2 Landscape Ecosystems at Multiple Scales 15

Overview of Spatial and Temporal Scales, 15
Spatial Scales of Hierarchical Landscape Ecosystems, 17
Climatic Classification, 18
Physiography, 20
Vegetation Types and Biomes, 20
Distinguishing and Mapping Landscape Ecosystems
at Multiple Spatial Scales, 25
Regional Landscape Ecosystems, 26
Regional Landscape Ecosystems of Michigan, 28
Local Landscape Ecosystems, 30
Local Landscape Ecosystems in Upper Michigan, 30
vii
viii C o n t e n ts
PA R T 2 The Forest Tree
3 Forest Tree Variation 35

Components of Phenotypic Variation, 35
Plasticity of the Phenotype, 36
Sources of Variation, 37
The Evolutionary Sequence, 38
Sexual and Asexual Systems, 39
Genetic Diversity of Woody Species, 39
Genecology, 40
Patterns of Genecological Differentiation, 41
Genecological Categories, 42
Factors Eliciting Genecological Differentiation, 42
Growth Cessation, 43
Growth Resumption, 46
Examples of Genecological Differentiation, 46
Eastern North American Species, 47
Scots Pine, 49
Wide-Ranging Western North American Conifers, 50
Ponderosa pine, 50
Douglas-fir, 52
Local Genecological Differentiation, 55
Factors Affecting Differentiation: Gene Flow and Selection Pressure, 56
Ecological Considerations at the Species Level, 57
Niche, 60
Hybridization, 60
Polyploidy, 63
The Fitness–Flexibility Compromise, 66
Epigenetics, 66
4 Regeneration Ecology 69
Regeneration, 69
Sexual Reproduction, 71
Maturation and the Ability to Flower, 72
Increasing Seed Production, 73
Reproductive Cycles, 74
Pollination, 75
Contents ix
Periodicity of Seed Crops, 76

Effects of Reproduction on Vegetative Growth, 78
Dispersal, 80
Seed Bank, Dormancy, and Germination, 82
Establishment Following Sexual Reproduction, 83
Post-Establishment Development, 90
Vegetative Reproduction, 90
5 Tree Structure and Growth 95

Tree Form, 95
Architectural Models, 97
Short and Long Shoots, 99
Patterns of Intermittent Growth, 100
Sylleptic and Proleptic Shoots, 102
Roots, 102
Kinds, Forms, and Occurrence, 103
Fine Root Relations, 105
Horizontal and Vertical Root Development, 107
Periodicity of Primary Root Growth, 108
Root Grafting, 108
Specialized Roots and Buttresses, 110
Stems, 110
Xylem Cells and Growth Rings, 111
Periodicity and Control of Secondary Growth, 112
Control of Earlywood and Latewood Formation, 114
Winter Freezing and Water Transport, 115
Water Deficits and Tree Growth, 116
PA R T 3 The Physical Environment

Forest Environment, 119
Site Factors, 120
Organization of Site Factors, 120
Interrelationships Among Site Factors, 121
Importance of Site in Forest Ecology, 122
0005430834.INDD 9 01-05-2023 17:43:22

x C o n t e n ts
6 Light 123
Distribution of Light Reaching the Ecosphere, 124
Plant Interception of Radiation, 125
Canopy Structure and Leaf Area, 125
Light Quality Beneath the Forest Canopy, 130
Sunflecks, 130
Light and Growth of Trees, 133
Light and Seedling Survival and Growth, 136
Light and Tree Morphology and Anatomy, 137
Light and Epicormic Sprouting, 139
Photocontrol of Plant Response, 139
Light and Ecosystem Change, 141
7 Temperature 143
Geographical Patterns of Temperature, 143
Temperatures at the Soil Surface, 145
Temperature within the Forest, 147
Temperature Variation with Local Topography, 148
Temperature and Plant Growth, 150
Cold Injury to Plants, 153
Dormancy, 155
Frost Hardiness and Cold Resistance, 156
Thermotropic Movements in Rhododendrons, 159
Winter Chilling and Growth Resumption, 161
Natural Plant Distributions and Cold Hardiness, 162
Deciduousness and Temperature, 164
8 Physiography 167
Concepts and Terms, 167
Characteristics of Physiography and their Significance, 168
Physiographic Setting, 169
Specific Landforms, 170
Elevation, 170
Form of Landforms, 170
Level Terrain, 170
Sloping Terrain, 171
Slope Characteristics, 171
Contents xi
Position on slope, 172

Aspect, 172
Slope inclination, 173
Parent Material in Relation to Landform, 173
Position of Landform in the Landscape, 174
Multiple Roles of Physiography, 174
Physiographic Diversity, Landscape Ecosystems, and Vegetation, 176
Mountainous Physiography, 176
Mountainous Terrain of California and the Pacific Northwest, 177
Physiography and Forests of the Central Appalachians, 180
Flatlands, 183
The Great Plains, 184
Pine Savannas of the Western Great Lakes Region, 185
Till Plains of the Midwest, 185
Southeastern and Southern Coastal Plain, 186
Floodplains, 186
Physiography and Firebreaks, 190
Microlandforms and Microtopography, 191
Tree Uprooting and Pit-and-Mound Microtopography, 191
Microtopography and Regeneration in Hardwood Swamps, 193
9 Soil 195
Parent Material, 195
Soil Formation, 197
Soil Profile Development, 197
Physical Properties of Soil, 200
Soil Texture, 200
Soil Structure, 201
Soil Color, 202
Soil Water, 202
Physical Properties of Water, 203
Soil Water Potential, 204
Chemical Properties of Soil, 206
Clay Mineralogy, 207
Cation Exchange and the Supply of Nutrients, 209
Soil Acidity, 210
Soil Organic Matter, 212
Soil Classification, 213
Landform, Soil, and Forest Vegetation: Landscape Relationships, 215
xii C o n t e n ts
10 Fire 219
Fire and the Forest Tree, 220
Causes, 220
Fire Regime, 220
Fire Types, Frequency, and Severity, 221
Fire Adaptations and Key Characteristics, 224
Strategies of Species Persistence, 227
Closed-Cone Pines, 228
Fire and the Forest Site, 230
Indirect Effects, 230
Direct Effects, 232
Organic Matter and Erosion, 234
Beneficial Effects of Fire, 236
11 Site Quality and Ecosystem Evaluation

and Classification237
Direct Measurement of Forest Productivity, 238
Tree Height as a Measure of Site, 239
Site-Index Curves, 240
Comparisons Between Species, 243
Advantages and Limitations, 243
Vegetation as an Indicator of Site Quality, 244
Species Groups of Ground Cover, 245
Indicator Plants of Coastal British Columbia, 246
Ecological Species Groups, 246
Plant Associations and Habitat Types in the Western United States, 247
Operational Site Classification Based on Vegetation, 247
Applications and Limitations of Vegetation, 250
Environmental Factors as a Measure of Site, 251
Climatic Factors, 251
Physiographic Land Classification, 252
Physiographic and Soil Factors: Soil-Site Studies, 252
Soil Surveys, 256
Multiple-Factor Methods of Site and Ecosystem Classification, 256
Ecosystem Classification and Mapping in Baden-Württemberg, 257
Applications of Multi-Factor Methods in the United States
and Canada, 258
Ecosystem Classification and Mapping in Michigan, 258
Ecosystem Classification in the Southeastern United States, 261
0005430834.INDD 12 11-23-2022 12:32:40

Contents xiii
Ecosystem Classification in the Southwestern United States, 262

National Classification Systems in the United States, 262
Ecological Land Classification in Canada, 263
Hills’ physiographic approach, 263
Other approaches used in Canada, 264
PA R T 4 Forest Communities
12 Animals in Forest Ecosystems 269

Plant Defense, 269
Investment in Plant Defense, 270
Plant Defense Against Insects, 272
Examples of Injury and Plant Defense, 273
Nutrition, 274
Plant Hybrid Zones as Reservoirs for Insect Diversity, 275
Plant Defense Against Mammals, 276
Roles of Animals in Plant Life History, 277
Pollination, 277
Seed Dispersal, 278
Fish and Reptiles, 278
Birds, 279
Mammals, 282
Germination and Establishment, 284
Decomposition, Mineral Cycling, and Soil Improvement, 285
Damage and Death, 286
Influence of Livestock on Forest Ecosystems, 288
13 Forest Communities 291

Community Concept, 291
Grounding Communities, 292
Florida Keys, 292
Interior Alaska, 294
Southern Illinois, 294
View from the Past: Community Concepts, 296
Schools and Terminology, 296
Concepts of Clements and Gleason, 297
0005430834.INDD 13 11-23-2022 12:32:40

xiv C o n t e n ts
Phytosociology in Europe, 298

Continuum Concept, 298
Community as a Landscape Ecosystem Property, 299
Examples of Spatial Variation in Forest Communities, 300
Discrete Forest Communities, 300
Coastal California: Giant and Pygmy Forests, 301
Forest–Grassland Ecotone, 302
Alpine Tree Lines, 302
Merging Forest Communities, 303
Eastern Deciduous Forest—Southern Appalachians, 303
New England, 304
Competition and Niche Differentiation, 305
Interactions Among Organisms, 307
Mutualisms in Forest Ecosystems, 307
Symbiotic Mutualisms—Mycorrhizae, 307
Nonsymbiotic Mutualisms, 307
Competition, 308
Forest Community Structure and Composition, 308
Vertical Structure, 310
Stand Density, 312
Competition and Overstory Composition, 313
Competition in the Understory, 314
Understory Tolerance, 315
Characteristics of Understory-Tolerant and -Intolerant Species, 316
Tolerance Ratings of Tree Species, 317
Examples of Understory Tolerance in Forest Ecosystems, 319
Nature of Understory Tolerance, 319
Environmental Factors Relating to Understory Tolerance, 322
Physiological Processes Relating to Tolerance, 323
14 Diversity in Forests 327

Concepts of Biological and Ecosystem Diversity, 327
The Value of Species Diversity, 328
Value of Biodiversity, 329
Common Threats to Diversity, 331
Measuring Diversity, 331
Levels of Diversity, 331
Measurement, 333
Inventory Diversity: Alpha Diversity, 333
Differentiation or Beta Diversity, 335
0005430834.INDD 14 11-23-2022 12:32:40

Contents xv
Diversity of Landscape Ecosystems, 336

Examples of Diversity, 337
Ground-Cover Species Diversity in Northern Lower Michigan, 337
Ecosystem Groups, 337
Ecosystem Types, 340
Ecosystem Diversity, 342
Causes of Species Diversity, 343
Diversity at Continental and Subcontinental Scales, 343
Paleogeography and Continental Relationships, 343
Glaciation, 344
Latitude and Elevation, 344
Diversity at Local Scales, 346
Physiography and Soil, 346
Community Composition and Structure, 349
Disturbance and Succession, 349
Focal Species in Conserving Diversity, 351
Foundation Species, 352
Keystone Species, 352
Endemics and Rare and Endangered Species, 353
Diversity and the Functioning of Ecosystems, 354
Biodiversity–Productivity Relationship, 354
The Role of Biodiversity in Ecosystem Stability, 356
Forest Management and Diversity, 357
Effects of Traditional Forest Management on Diversity, 357
Preserving Diversity in Managed Forests, 358
Ecological Forestry: Incorporating Biodiversity
into Forest Management, 359
Variable-Retention Harvest System, 360
Designing a Variable-Retention Harvest System, 361
How Well Does Variable Retention Conserve
Biodiversity?, 361
Epilog: Conserving Ecosystem and Biological Diversity, 363
PA R T 5 Forest Ecosystem Dynamics
15 Long-Term Forest Ecosystem and Vegetation Change 367

Change Before the Pleistocene Age, 367
Pleistocene Glaciations, 368
Ecosystem and Vegetational Change Since the Last Glacial Maximum, 370
0005430834.INDD 15 11-23-2022 12:32:40

xvi C o n t e n ts
Eastern North America, 370

Overall Migration Sequence and Patterns, 371
Ecosystem Change in the Southern Appalachians, 373
Western North America, 373
Patterns of Tree Genera and Species Migrations, 375
Migration Irregularities and Disturbance, 377
Migration From Glacial Microrefugia, 377
Independent Migration and Similarity of Communities Through Time, 379
16 Disturbance 381
Concepts of Disturbance, 382
Defining a Disturbance, 382
Disturbance as an Ecosystem Process, 384
Sources of Disturbance, 386
Major Disturbances in Forest Ecosystems, 387
Fire, 387
Roles of Fire in Forest Ecosystems, 387
Pines in New England and the Lake States, 389
Western Pines and Trembling Aspen, 390
Southern Pines, 393
Douglas-Fir in the Pacific Northwest, 394
Giant Sequoia, 394
Fire History and Behavior, 395
Northern Lake States, 395
Boreal Forest and Taiga, 397
Northern Rocky Mountains, 398
Fire Suppression and Exclusion, 399
Wind, 400
Widespread and Local Effects, 401
Principles of Wind Damage, 401
Broadscale Disturbance by Hurricanes, 403
Gulf and Southern Atlantic Coasts, 403
New England—1938 Hurricane, 404
Wave-Regenerated Fir Species, 404
Floodwater and Ice Storms, 404
Insects and Disease, 406
Catastrophic and Local Land Movements, 407
Logging, 407
Land Clearing, 409
Disturbance Interactions, 409
0005430834.INDD 16 11-23-2022 12:32:41

Contents xvii
Biotic Composition Changes, 411

Elimination of Species, 411
Addition of Species, 412
17 Forest Succession 413

Basic Concepts of Succession, 414
Primary and Secondary Succession, 414
Biological Legacies, 414
Successional Pathways, Mechanisms, and Models, 414
Autogenic and Allogenic Succession, 416
How is Succession Determined?, 416
Evolution of the Concept of Forest Succession, 417
Formal Ecological Theory, 417
How Does Succession Work?, 418
Clementsian Succession, 419
Stages of Succession, 420
Primary Succession, 420
Secondary Succession, 423
Successional Causes, Mechanisms, and Models, 427
Key Characteristics and Regeneration Strategies, 427
Availability and Arrival Sequence of Species, 428
Facilitation, Tolerance, and Inhibition, 428
Change in Ecosystems, 431
End Point of Succession?, 431
Succession as an Ecosystem Process, 432
Examples of Forest Succession, 434
Primary Succession on Recently Deglaciated Terrain, 434
Succession Following the Eruption of Mount St. Helens, 437
Secondary Succession Following Fire in Ponderosa Pine Forests
of Western Montana, 439
Secondary Succession and Gap Dynamics, 439
Gap Specialists: American Beech and Sugar Maple, 444
Fire and Oak Dominance—Oaks at Risk, 446
18 Carbon Balance of Trees and Ecosystems 449

Carbon Balance of Trees, 450
Photosynthesis, Dark Respiration, and Leaf C Gain, 450
Light and Leaf C Gain, 452
0005430834.INDD 17 11-23-2022 12:32:41

xviii C o n t e n ts
Temperature and Leaf C Gain, 454

Water and Leaf C Gain, 455
Soil Nitrogen Availability and Leaf C Gain, 456
Construction and Maintenance Respiration, 456
Allocation to Structure, Storage, and Defense, 459
Light and C Allocation, 461
Soil Nitrogen Availability and C Allocation, 461
Carbon Balance of Ecosystems, 465
Biomass and Productivity of Forest Ecosystems, 466
Measurement of Biomass and Productivity, 469
Climate and Productivity, 473
Soil Properties, Forest Biomass, and ANPP, 475
Biomass Accumulation During Ecosystem Development, 477
Soil N Availability and Belowground Net Primary Productivity, 482
19 Nutrient Cycling 487

Nutrient Additions to Forest Ecosystems, 488
Mineral Weathering, 488
Atmospheric Deposition, 490
Biological Fixation of Nitrogen, 493
Nutrient Cycling within Forest Ecosystems, 497
Nutrient Transport to Roots, 497
Nutrient Uptake and Assimilation by Roots, 498
Root Architecture, Mycorrhizae, and Nutrient Acquisition, 501
Root Architecture, 501
Mycorrhizae, 502
Plant Litter and the Return of Nutrients to Forest Floor and Soil, 504
Leaf and Root Litter Production, 505
Nutrient Retranslocation, 507
Nutrients in the Forest Floor, 509
Organic Matter Decomposition and Nutrient Mineralization, 512
Biochemical Constituents of Plant Litter, 513
Dynamics of Decomposition, 515
Nitrogen Immobilization and Mineralization, 519
Nitrogen Availability in Forest Ecosystems, 521
Nitrification, 522
Nutrient Loss From Forest Ecosystems, 523
Nutrient Leaching from Forest Ecosystems, 524
0005430834.INDD 18 11-23-2022 12:32:41

Contents xix
Denitrification, 525
The Cycling and Storage of Nutrients in Forest Ecosystems, 527
Nutrient Storage in Boreal, Temperate, and Tropical Forests, 527
The Nitrogen and Calcium Cycle of a Temperate Forest Ecosystem, 528
Ecosystem C Balance and the Retention and Loss of Nutrients, 530
Forest Harvesting and Nutrient Loss, 532
PA R T 6 Forests of the Future
20 Climate Change and Forest Ecosystems 537

Climate Change Concepts, 537
Effects on Temperature, 540
Effects on Precipitation, 543
Climate Change Effects on the Forest Tree, 546
Tree Growth and Mortality, 546
Phenology, 548
Regeneration, 551
Climate Change Effects on Tree Species Distributions, 554
Observed Range Shifts, 554
Projected Changes in Tree Species Distributions, 556
Projected Changes in Forest Type Distributions, 559
Climate Change Effects on Forest Disturbances, 561
Fires, 563
Insects and Pathogens, 568
Wind, 570
Climate Change Effects on Forest Carbon, 571
Climate Change Effects on Carbon Gain: Primary Productivity, 572
Climate Change Effects on Carbon Loss: Heterotrophic Respiration, 573
Feedbacks Among Disturbance, Climate Change, and Carbon
in Forests, 575
Fire, Carbon, and Climate Change in Forests of Yellowstone
National Park, 577
Adapting to Climate Change Effects on Forests, 578
Assisted Migration, 579
Refugia, 581
Forest Carbon Management, 583
0005430834.INDD 19 11-23-2022 12:32:41

xx C o n t e n ts
21 Invasive Species in Forest Ecosystems 587

Concepts of Invasive Species, 587
Definition of Invasive Species, 587
Characteristic Traits of Invasive Plant Species, 589
Non-Plant Invasive Species in Forests, 594
Impacts of Invasive Species on Forests, 597
Impacts of Invasive Plants on Forests, 597
Competition, 598
Altered Fire Regimes, 598
Carbon and Nutrient Cycling, 599
Impacts of Invasive Insects and Pathogens on Forests, 600
Chestnut Blight, Dutch Elm Disease, and Forest Succession, 601
Impacts of Invasive Animals on Forests, 604
A Primer of Invasive Species Management in Forests, 605
Early Intervention Strategies, 607
Management Approaches for Established Invasive Species, 607
Novel Ecosystems and Invasive Species, 608
22 Forest Landscape Ecology 611

Concepts of Landscape Ecology, 612
Forest Fragmentation and Connectivity, 615
Patches in Forest Ecology, 616
Forest Fragmentation, 617
Ecological Effects, 617
Connectivity, 620
Disturbances on Landscapes, 622
Effects of Heterogeneity on Disturbances, 622
Hurricanes in New England, 622
Landscape Pattern Effects on Disturbance Spread, 625
Effects of Disturbances on Heterogeneity, 625
Stand-Replacing Wildfires in Yellowstone National Park, 627
Historical Range of Variability, 630
Interactions of Landscape Patterns and Ecological Processes, 632
Leaf Area and Productivity, 633
Forest Carbon Dynamics, 635
Nutrient Dynamics, 636
0005430834.INDD 20 11-23-2022 12:32:41

Contents xxi
23 Sustainability of Forest Ecosystems 639

Concepts of Sustainability, 639
The Prevalence of Human Values in Forest Ecology, 639
Historical Perspective of Sustainability in Forests, 641
Ecosystem Services, 642
Toward a Definition of Sustainability, 644
Where Do We Go From Here?, 646
Epilog: Earth as a Metaphor for Life, 648
References651
Scientific Names of Trees and Shrubs 731
Index739
0005430834.INDD 21 11-23-2022 12:32:41

Preface
F orest Ecology deals with forest ecosystems—spatial and volumetric segments of the Earth—and
their climate, landforms, soils, and biota. It is designed as a textbook for people interested in
forest ecosystems—either in the context of courses in forest ecology and environmental science or
as an ecological reference for those in professional practice. This book is meant to provide basic
ecological concepts and principles for field ecologists, foresters, naturalists, botanists, and others
interested in the conservation and restoration of forest ecosystems.
Ecology, in general, has undergone several sea changes since the appearance of the first
edition in 1964, with enormous increases in public interest and scientific development of theory
and research. Ecology and the issues associated with it have become part of our modern lexicon.
The great number of advances in our ecological knowledge, as well as increased public interest,
presents forest ecologists with both opportunities and challenges to sustainably manage ecosystems
using our best understanding of ecosystem properties and processes. This book will hopefully be
useful in that process by providing an understanding of the ecological relationships of individual
trees and forest ecosystems.
The book has six major subdivisions. “Forest Ecology and Landscape Ecosystems” introduces
forests as whole ecosystems rather than tree communities, and at multiple scales. “The Forest
Tree” considers the genetic variations among individual trees, the causes of diversity within and
between species, regeneration ecology, and selected aspects of tree structure and function. “The
Physical Environment” treats the physical factors of forest ecosystems that form the forest site—the
influences of light, temperature, physiography, soil, and fire on the individual forest plant and
on plant communities. The concluding chapter in this part considers methods of evaluation and
classification of the forest site and ecosystems. In Part 4, “Forest Communities,” we consider the
forest community of trees and associated plants and animals that form a key structural component
of forest ecosystems—one part of the whole. We also consider the importance and measurement of
diversity of species and ecosystems. In “Forest Ecosystem Dynamics,” we examine the functional
relationships of the physical environment and the biota. We first examine changes in communities
and ecosystems over tens of thousands of years. We then consider the extent to which disturbance,
an ecosystem process, initiates change (termed succession) over shorter time scales of centuries.
Chapters on carbon balance and nutrient cycling present a detailed consideration of the pattern
in which carbon (i.e., energy) and plant nutrients flow within forest ecosystems and how natural
and human-induced disturbances alter these patterns. Finally, “Forests of the Future” explores the
role of humans in the sustainability of forests. Here we emphasize two of the most pressing issues
in forest ecology today, climate change and invasive species, and present a review of landscape
ecology which has humans at its center. We end the book with a treatment of sustainability itself.
In this edition, we have made great attempts to maintain the core organization and readability
of previous editions while adding those areas most relevant to forest ecology that have developed
over the last quarter-century. We have retained the important focus on landscape ecosystems (rather
than organisms and communities) that was developed in the fourth edition and have added critical
new ecological concepts and research that have developed in genetics, diversity, climate change,
invasive species, and sustainability. The ecological literature has only become more voluminous
over the past 25 years, and as in previous editions our use of the literature was selective, rather
than exhaustive. New references were most often chosen based on their accessibility to students
xxiii
xxiv P r e fac e
and practitioners as understandable examples of important ecological concepts. At the same time,
we have retained many older references that still provide excellent examples of fundamental eco-
logical concepts, many of which would otherwise be lost in obscurity.
As before, we have integrated woody plant physiology into multiple chapters, rather than
developing it in a single chapter, and in this edition have done the same with climate because of its
overriding influence on so many ecosystem processes. We have also further limited our treatment
of forest ecology with examples from temperate and boreal forests, with special emphasis on North
America and Europe. With a primary focus on the ecological principles of forests that form the
basis for management, we have largely avoided specific treatments of forest management tech-
niques and strategies throughout the book, although by necessity we have provided some examples
of forest management in the chapters on diversity and invasive species.
The revised edition would not have come to pass without the patience and dedication of
many colleagues who helped in its preparation. We are especially indebted to the following for
their reviews of one or more chapters: Dennis Albert, Brian Buma, Mark Dixon, Tom Dowling,
Jennifer Fraterrigo, Stephen Handler, Donna Kashian, Doug Pearsall, David Rothstein, Madelyn
Tucker, and Chris Webster. Other important contributions, either through provision of material,
enlightening discussions, or enthusiastic encouragement, were made by Jonathon Adams, Virginia
Laetz, and Dan Binkley. Victoria Meller was extremely supportive of D. M. Kashian and his time
spent away from campus in the completion of this revision, as were the graduate students in the
Kashian lab. Perhaps without knowing it, the graduate and undergraduate students in the 2018
and 2020 Terrestrial Ecology classes at Wayne State had an immense role in the thought processes
and revisions made in the 5th edition. D. R. Zak was partially supported by the National Science
Foundation and the Department of Energy during the preparation of this text.
Daniel M. Kashian
Detroit, Michigan, USA
Donald R. Zak
Ann Arbor, Michigan, USA
Forest Ecology and PA R T 1
Landscape Ecosystems
F orest ecologists work to understand the dynamics, structure, and function of

forest ecosystems. The first step in doing so is to approach ecosystems as geo-
graphic units or landscape ecosystems—real, three-dimensional, defined locations at the
Earth’s surface. Landscape ecosystems include organisms, species, and communities, but
also the air above and the soil below these living things such that organisms are but
one important part of a larger landscape unit. Our perception of forest ecosystems in
this book is that ecosystems are not simply extensions of forest communities, whereby
ecosystems are conceived as organisms and their environment. Instead, units of whole
ecosystems that integrate factors of climate, landforms, soils, co-occurring biota, and all
the interactions between them are the most appropriate units of study rather than their
individual parts that too often claim our immediate attention.
In understanding forest ecosystems, particularly in the context of an ecology course
or in solving ecological problems, ecosystems need to be conceived at several spatial and
temporal scales. It is very daunting to attempt to gain such understanding, and certain
well-known ecologists have suggested that the environmental complex is unknowable
and inexpressible. However, conceiving landscapes as ecosystems and proceeding “from
above,” that is, from the biggest to the smallest units to understand their spatial relation-
ships, makes synthesis possible and manageable. Although local sites and stands with
their familiar species appear a convenient starting point, we know that these are, in turn,
affected by geological and climatic factors of higher levels within which they are embed-
ded. Landscape ecosystems are nested within one another in a hierarchy of spatial sizes
such that it is best to consider the big picture first—the comprehensive view from the
outside—rather than the minutia of details as seen from inside of the ecosystem. Our
perspective in understanding ecosystems involves establishing a framework for studying
the components—a framework of the spatial and temporal, hierarchical pattern of eco-
systems of all sizes making up the ecosphere.
Therefore, in the first part of this book, before immersing in the detail of organ-
isms, sites, and their interrelationships, we wish to first examine landscape ecosystems,
and then place them within a perspective of spatial levels and their processes at different
Forest Ecology, Fifth Edition. Daniel M. Kashian, Donald R. Zak, Burton V. Barnes, and Stephen H. Spurr.
© 2023 John Wiley & Sons Ltd. Published 2023 by John Wiley & Sons Ltd.
1
2 PA RT 1 Forest Ecology and Landscape Ecosystems
temporal scales. In Chapter 1, we present the basic concepts of forest ecology within a
framework of landscape ecosystems. In Chapter 2, we present the concept of scale and
consider the hierarchy of these landscape ecosystems. These fundamental concepts pro-
vide the basis for studying the variation, life history, and ecology of individual organisms
that follow in Part 2.
Concepts of Forest Ecology CHAPTER 1
A forest is an ecological system dominated by trees and other woody vegetation. More than
simply a stand of trees or a community of woody and herbaceous plants, a forest is a complex
ecological system, or ecosystem, characterized by a layered structure of functional parts. Ecology
is the study of ecological systems and their interacting abiotic and biotic components. Forest
ecology, therefore, addresses the structure, composition, and function of forests. In forest ecology,
we study forest organisms and their responses to physical factors of the environment across for-
ested landscapes. Forests are widespread on land surfaces in humid climates outside of the polar
regions. It is with forests in general, and with the temperate North American forest in particular,
that this book is concerned.
There are many ways to study forest ecosystems. Most simply, a forest may be considered in
terms of the trees that give the forest its characteristic aboveground appearance or physiognomy.
Thus, we think of a beech–sugar maple forest, a ponderosa pine forest, or of other forest types, for
which the naming of the predominant trees alone serves to characterize the forest ecosystem.
Forest types are often considered to be composed of forest stands, which are trees in a local
setting possessing sufficient uniformity of species composition, age, spatial arrangement, or
condition to be distinguishable from adjacent stands (Ford-Robertson 1983).
A broader concept of a forest may take into account the interrelationships that exist between
forest trees and other organisms. Certain herbs and shrubs are commonly found in beech–sugar
maple forests, and these may differ from those found in ponderosa pine or loblolly pine forests.
Similar interrelationships may be demonstrated, for example, for birds, mammals, arthropods,
mosses, fungi, and bacteria. Thus, part of the forest ecosystem is the assemblage of plants and ani-
mals living together in a biotic community. The forest community, then, is an aggregation of
plants and animals living together and occupying a common area. It is thus a more organismally
complex unit than the forest type.
A third approach is to focus on geographic or landscape ecosystems. This approach is
centered conceptually and in practice on whole ecosystems and not just their parts. When our pri-
mary focus is real live chunks of Earth space, that is, landscapes and waterscapes (oceans, lakes,
rivers; hereafter included as parts of a landscape), we can effectively study their parts (e.g., organ-
isms, soils, and landforms) while recognizing that each is but one part of a functioning whole. We
emphasize this focus on ecosystems rather than on the individual organisms and species that are
parts of them.
In the past, the forest stand or the species has been the focus in natural resource fields such
as forestry and wildlife. However, we are really managing whole forest ecosystems, despite their
incredible complexity, because the diverse biota is inseparable from the physical environment that
supports it. A consideration of the field of ecology from this viewpoint provides an overall
perspective.
3
4 Chapter 1 Concepts of Forest Ecology
ECOLOGY
Broader fields of scientific inquiry are difficult to limit and define, and ecology is one of the most
indistinct. In 1866, Ernst Haeckel proposed the term oecology, from the Greek oikos meaning
home or place to live, as the fourth field of biology dealing with environmental relationships of
organisms. Thus, ecology literally means “the knowledge of home,” or “home wisdom.” Since its
introduction, the term has been applied at one time or another to almost every aspect of scientific
investigation involving the relationship of organisms to one another or to the environment
(Rowe 1989). Haeckel’s organismal focus of ecology has since been redefined and expanded to
include the physical aspects of the environment that provide life for those organisms (Hagen 1992;
Golley 1993). Thus, Rowe (1989, p. 230) suggests:
Ecology is, or should be, the study of ecological systems that are home to organisms at the surface of the
earth. From this larger-than-life perspective, ecology’s concerns are with volumes of earth space, each con-
sisting of an atmospheric layer lying on an earth/water layer with organisms sandwiched at the
solar-energized interfaces. These three-dimensional air/organisms/earth systems are real ecosystems—the
true subjects of ecology.
This approach to ecology emphasizes whole ecosystems as well as organisms, both volumet-
ric and having structure and function.
LANDSCAPE ECOSYSTEMS
The British botanist–ecologist Arthur Tansley (1935) introduced the term ecosystem, writing with
an emphasis on “the whole ‘system,’ including not only the organism complex but also the whole
complex of physical factors.” He also noted that from the point of view of the ecologist, ecosystems
“are the basic units of nature on the face of the Earth.” Tansley was a biologist and vegetation ecol-
ogist, and so his idea of ecosystem was centered on organisms (species or communities) rather
than geographic or landscape entities. With this bioecosystem approach, “ecosystem” derives its
meaning from particular plant or animal organisms of interest, and an “abiotic” environment
defined by the organisms as relevant or not is considered with lesser emphasis. In this approach,
every organism defines its own ecosystem, nearly infinite in number and difficult to study and use
as a basis for management and conservation.
On the other hand, others (e.g., Rowe 1961a and Troll 1968, 1971) view ecosystems centered
on geographic or landscape units (i.e., geoecosystems) of which organisms are but one important
structural component (Rowe and Barnes 1994). We term these units landscape ecosystems in
part to differentiate them from geology-based units of study (e.g., Huggett 1995). Landscape eco-
systems are geographic objects, with a defined place on the Earth. Landscape ecosystems have
three dimensions (volume) just as organisms do, including landforms and biota at the Earth’s sur-
face as well as the air above them and the soils below them (Figure 1.1). Other terms have been
introduced to express the same idea, but are less commonly used, such as the ecotope
(Troll 1963a, 1968) and the ecoterresa (Jenny 1980). This geographic/volumetric concept has been
discussed and adapted by professional and academic ecological societies (Christensen et al. 1996),
and is useful to field ecologists, naturalists, foresters and other land managers, and natural resource
professionals. The concept is described in detail in Chapters 2 and 11.
In addition to being geographic and volumetric, landscape ecosystems are hierarchical,
extending downward from the largest ecosystem we know, the ecosphere (Cole 1958), through
multiple levels of ecological organization (Figure 1.2). These levels include macrolevel units of
continents and seas, each of which contains mesolevel units of regional ecosystems (major phys-
iographic units and their included organisms), which in turn contain local ecosystems
(Hills 1952), the smallest level of homogeneous environment with organisms enveloped in it. We
therefore conceive the ecosphere and its landscapes as ecosystems, large and small, nested
Ecosystem boundaries
Macroclimate
Topoclimate
Biota
Landform
Surface water
Soils
Ground water
Bedrock
F I G U R E 1 . 1 The three-dimensional, volumetric nature of a landscape ecosystem. Ecosystems comprise the

atmosphere (macroclimate as well as the climate affected by surface relief), landforms and soils (underlain
by ground water and bedrock), and the biota that provide a physical connection between the air and the
Earth. Source: Bailey (2009) / Springer Nature.
Ecology
Organismal Biology
Cell
Organelle Cell Biology
olecu
Molecular Biology and
M
le
Biochemistry
F I G U R E 1 . 2 Objects of study from the most inclusive (ecosphere) to the least inclusive levels of organiza-
tion (cell and organelle and molecule below it). Note that each higher level envelops the lower ones as parts
of its whole. Some corresponding fields of study are also shown. Aggregates of organisms, such as popula-
tions and communities, are components of ecosystems at all scales. Like other components such as
atmosphere, landform, and soil, they do not appear in this diagram of first-order objects of study, but are
shown in Figures 1.3 and 1.4. Source: Modified from Rowe (1961a).
within one another in an ecological hierarchy, having processes at each level with their own
spatial and temporal scales (see Chapter 2).
Two landscape ecosystems in Figure 1.3 (Rowe 1984b) illustrate characteristic ecosystem
differences in hilly or mountainous terrain. The two ecosystems are distinguished by different geo-
morphologies (convex upper slope versus concave lower slope, and high versus low topographic
slope position) that mediate microclimate, soil water, and nutrient availability. The vertical dashed
line is placed at an ecologically significant boundary that spatially separates the two ecosystems.
Organisms in these ecosystems are sandwiched between the air and the Earth. Also shown in Figure 1.3
are the traditional fields of study in which individuals seek to understand each of the ecosystem com-
ponents, although forest ecologists aim to understand the integrated effects of all of these components.
In many parts of this book, we focus on organisms, species, and communities, but always
remembering that they are parts of volumetric, hierarchical ecosystems. For studies of organisms
in their immediate surroundings, a biological approach is often useful. Nevertheless, for
management of ecosystems, studies of forest productivity, and the conservation and restoration of
forest ecosystems, a landscape ecosystem approach is eminently practical and theoretically sound.
Forest ecologists not only study (i) organisms of these systems and their aggregates as communities
HILLSIDE SLOPE
Upper Lower
Fields of Study
Upper Air Meteorology

Layer Climatology
Lower Air Topoclimatology

Layer
Biota Ecology
Biology
Soil Pedology
Landform Geomorphology
& Surficial Geology
Mineral Strata Bedrock Geology
Water Table Hydrology
F I G U R E 1 . 3 Structural profile illustrating landscape ecosystems of upper and lower slopes. Air–earth
layers surround the organisms at the Earth’s energized surface. The vertical line is set at an ecologically
significant topographic break, dividing the upper and lower slope ecosystems. Source: Rowe (1984b) / United
States Department of Agriculture / Public Domain.
Landscape Ecosystems 7
and populations (see Chapters 3–5, 12, and 13), but also (ii) the functioning of local ecosystems
that involves complex interactions among organisms and their supporting environment
(Chapters 6–11, 13–14, and 16–19), and (iii) the spatial patterns of occurrence and interrelationships
of entire forest ecosystems (Chapters 2, 18, 19, and 22).
LANDSCAPE ECOSYSTEM AND COMMUNITY

The term ecosystem was introduced in 1935 by Tansley, but terms in various languages, such as taiga,
heath, bald, Auenwald (river floodplain forest), pampas, prairie, chaparral, maquis, hammock, mus-
keg, and bog, have long been used to depict interactions among air, organisms, and soil. Very often the
terms emphasize a distinctive plant community and may therefore imply that an ecosystem is simply
an extension of a community. Tansley’s definition of ecosystem as “organism + environment” leads to
this view, and general definitions such as “ecosystem = biotic community + environment” reinforce
this view. Although indispensable for forest ecologists and managers, vegetation may not always coin-
cide with climate–landform–soil-based ecosystems because of disturbance and/or unknown historical
factors. Thus, we emphasize the importance of geography and physiography within a regional climatic
setting as the basis of understanding not only vegetation but whole ecosystem structure and function.
This conceptual approach is not to say that communities are unimportant; they form the key
ecosystem component whose response is essential in affecting ecosystem change and indicating
the integrated effects of many site factors (Chapter 11). Communities and populations, however,
fundamentally differ from entities such as ecosystems, organisms, and cells because they are aggre-
gates of individuals but not functional systems. Entities such as ecosystem, organism, cell, and mol-
ecule (Figure 1.2) are “volumetric” levels of organization because they have structurally joined
parts that form a functioning unit (Rowe 1961a, 1992c). By contrast, communities and populations
are assemblages or aggregates of spatially separated trees and understory plants that have no
necessary, physical, structural connections.
ECOSYSTEM STRUCTURE AND FUNCTION

An ecosystem’s structure describes the spatial arrangement of its parts. In local forest ecosystems,
multiple strata of atmosphere and soil (Figure 1.3), as well as surface relief, influence species com-
position and its patterns of occurrence. Plants provide a physical connection between the soil strata
and the aboveground layers of air. The layered structure and related physical and chemical prop-
erties of soil are well studied and understood, but similar properties of the atmosphere are not
(Rowe 1961a; Woodward 1987), particularly as they affect forest organisms at the Earth’s surface.
Vegetation itself is also structured vertically, and the vertical structure and species composition of
communities as well as species interactions provide insights critical in understanding properties
and processes of ecosystems. In addition to species composition, vegetation structure is shaped by
tree form and stem density, canopy characteristics, shrub and herbaceous plant abundance, and the
amount and distribution of dead vegetation, among other characteristics. Thus, the physiognomy of
vegetation varies markedly in different regional and local ecosystem types (Chapters 5 and 6).
The horizontal spatial patterning of these structural components occurs at multiple spatial
scales, but is most often described across broad scales. Landscapes are structured horizontally into
mosaics of ecosystems that reflect differences in climate, geology, and physiography and their
relative effects on vegetation. The natural communities of these systems reflect limiting factors of
climate, soil water, nutrients, and disturbances, and in turn modify the physical factors. Different
ecosystem mosaics characterize mountains, plains, and river valleys due to fundamental differ-
ences in their physical factors and the vegetation adapted thereto (Chapter 8). The diversity of
landscape ecosystems can be assessed by understanding such mosaics (Lapin and Barnes 1995).
Such an understanding provides a spatial ecosystem framework for programs in biodiversity that
seek to conserve and manage the diversity of organisms and maintain and increase populations of
rare and endangered species (Chapters 14 and 22).
Local landscape ecosystems, besides having a structure of interconnected parts, are
functional units characterized by many processes that define their properties. Ecosystem-level
processes are part of the entire system and are not restricted only to physical or biotic parts. These
processes drive or mediate the flow of energy or matter and/or the cycling of materials in the sys-
tem. Organic matter decomposition; cycling of water, nutrients, and carbon; and biomass
accumulation are considered ecosystem-level processes. Other processes are often more associated
with plant species, populations, or communities, such as photosynthesis and respiration,
reproduction, regeneration, mortality, and succession. Despite their basis in organisms, these are
also ecosystem processes because they are mediated and regulated by characteristic ecosystem
factors of temperature, water, nutrients, and disturbances (such as fire, windstorm, and flooding).
Ecosystem function is often described with box-and-arrow diagrams, flow charts, and simulation
modeling as ways of disentangling very complex systems (Figure 1.4).
Landscape ecosystem structure and function are tightly coupled by the physical environ-
ment, the frequency and severity of disturbances that reset succession, and the life histories of the
plants and animals that comprise the biotic community. There is increasing evidence that many
aspects of an ecosystem’s function are linked to the diversity of its biota (Chapter 14). In turn, an
ABIOTIC ENVIROMMENT
SUBSTANCES OTHER RADIATION SPACE STRUCTURE
HUMANS
water, pressure, heat, media (soil, water,
as a super-
O2, CO2, fire, light air) and their motion,
organic
minerals pH, etc. area, substratum
factor
PRIMARY PRODUCERS
(other green plants
autotrophs*)
OTHER
ECO-
symbionts pollination,
SYSTEMS
distribution,
selection
by grazing
DECOMPOSERS DEAD
mineralizers organic
CONSUMERS OUTPUTS
substances
by respiration,
H scavengers herbivores H erosion, oozing
parasites away of water
bacterio- are omitted
predators H
phages soil
humus
‘limit’ of the ecosystem
dead living material or energy flow other influences
Humans as a partner *relations omitted

H
of the ecosystem
F I G U R E 1 . 4 A model of a landscape ecosystem detailing the flow of energy and matter among biota. The
model also describes the interactions between the physical environment and the biota. Source: Reprinted
from Ellenberg (1988) / Cambridge University Press.
Examples of Landscape Ecosystems 9
ecosystem’s biodiversity is strongly shaped by its physical factors such as climate, physiography,
and soil, which provide the context within which organisms survive, adapt, and evolve. In addition,
the functional aspects of an ecosystem are strongly affected by its geographical context and by its
spatial position relative to its surrounding neighbors on a landscape. Adjacent ecosystems, espe-
cially in mountainous or hilly terrain, affect one another by the lateral exchanges of materials and
energy. Water and snow, soil, organic matter, nutrients, and seeds are transported downhill; the
effects on other ecosystems depend on the size, shape, and composition of the systems. Therefore,
an understanding of the spatial pattern and configuration of landscape ecosystems can play an
important role in the management of ecosystems and their biota. An excellent overview of function
in terrestrial ecosystems is given by Chapin et al. (2012), its variation across landscapes considered
in Lovett et al. (2005), and the history of the ecosystem concept itself is reviewed by Golley (1993).
EXAMPLES OF LANDSCAPE ECOSYSTEMS

There are many examples of landscape ecosystems, some of which are easily discerned in the field,
whereas others must be carved out of geographic continua. For example, bogs in the glaciated ter-
rain of northern and boreal regions are easily discernible ecosystem types. They exhibit distinctive
physiography, soil, and vegetation, and they recur in the landscape. The distinct terraces of river
floodplains distinguish local ecosystem types that differ in microclimate, soil, drainage, vegetation,
and their dynamics. Mountains are characterized by gradients of elevation, aspect, and slope steep-
ness along which strikingly different ecosystems can be purposefully delimited and mapped,
although their boundaries may not be sharply demarcated. On old lake-bed terrain, dry, sandy
beach ridges are easily distinguished from adjacent depressions of swampy land. Differences in
water drainage and oxygen availability in these adjacent systems result in major differences in
their native tree-and ground-flora vegetation. Unseen but critical are the very different processes
that also distinguish these ecosystems.
Ecosystem components will change over time, giving rise to a sequence of different landscape
or waterscape ecosystems on a given place on the Earth’s surface. Striking changes in the fossil
record illustrate long-term changes in physiography, soil, and biota (Chapter 15), but short-term
changes also occur. Shorter-term changes are most obvious when natural or human-caused distur-
bances affect vegetation. A site-specific location may exhibit a range of different forest commu-
nities over 100 to 300 years, young and old, as disturbances or lack thereof affect the biota. Ecosys-
tem structure and function at this site change over time. The landforms and associated pattern of
parent material, soil, and climate also change over time, but more slowly than the suite of species
available to recolonize the disturbed site. Thus, what we term the ecosystem site type or simply
ecosystem type (land area supporting potentially equivalent ecosystems) can be distinguished
and mapped regardless of the forest community currently present.
One such example is illustrated in Figure 1.5 where deciduous forests occur on glaciated ter-
rain in southern Michigan. In this setting, three local ecosystem types are distinguished by differ-
ences in physiography (outwash and moraine landforms); soil; drainage; and overstory, under-
story, and ground-cover vegetation. The relatively fine-textured, silty soil on the outwash plain
(type 1) supports forest dominated by white oak, whereas the drier, coarse-textured, sandy out-
wash soil (type 2) supports a black oak community. The fine-textured, moist, clayey soil on
the rolling moraine landform (type 3) supports a community dominated by northern red oak. The
moraine formerly supported a beech–sugar maple forest. However, recurrent fires through the drier
white and black oak ecosystems killed the fire-sensitive mesic species of the adjacent moraine
ecosystem and led to dominance of northern red oak. An occasional beech is still found, and red
maple has invaded the shaded understory.
The western portion of ecosystem type 1 was clear-cut about 90 years ago, and a cover type
markedly different from the adjacent old-growth white oak forest has formed (Figure 1.5, type
(a) Ecosystem Type

1 2 3
American T'
T 1a-cut 1b-uncut Black Oak Northern Red Oak beech
White Oak
Fine Outwash
– Silt
and
sh – S
Outwa
Coarse
End Moraine
(Till)
(b) Forest Cover Type

1a 1b 2 3
cut uncut Black Oak Northern Red Oak
T T'
1 2 3
Ecosystem Type
Cutting Type Transect

Boundary Boundary T-T'
F I G U R E 1 . 5 Three local landscape ecosystem types in glacial terrain that are differentiated by landform,
parent material, soil, and vegetation: (a) Lateral transect from T to T′ showing vegetation and underlying
geological parent material. (b) Top view showing distribution of forest trees, cover types, and ecosystem
types. Following clear-cutting of part of type 1, two forest cover types (1a—early successional oak forest;
1b—old-growth white-oak forest) are distinguished. The diagram illustrates that different forest cover types
(1a and 1b) are not necessarily different ecosystem types. They represent two of many possible ecosystem
derivatives (disturbed in 1a versus relatively undisturbed in 1b) of a given ecosystem type. See text for
discussion.
An Approach to the Study of Forest Ecology 11
la). The overstory tree layer of the disturbed area is now dominated by white, black, and red oaks,
white ash, black maple, American elm, black cherry, and sassafras. These species either sprouted
from the base of cut trees, were already present in the white oak forest understory, or seeded in
from adjacent communities following cutting. Today, we can recognize two (types 1a and 1b) of
the many compositionally different forest communities that might occur at the site of ecosystem
type 1; these easily could be mapped as two different forest cover types. Because the cut-over area
(type 1a) has physiography and soil like type 1b, its vegetation may gradually become similar in
structure to those of ecosystem type 1b, providing that no major changes in climate, soil, or
ecosystem processes (including changed browsing pressure by herbivores) were caused by clear-
cutting. Thus, a given local ecosystem type, defined by relatively stable features of physiography
and soil, may have a suite of disturbance-induced cover types (Simpson et al. 1990). Therefore,
recognizing forest cover types alone is not necessarily likely to provide a useful estimate
of site potential for management or conservation. However, cover types are extremely use-
ful in management planning for wildlife, timber, water, and recreational use of existing forest
communities.
It is often the conspicuous forest cover type that receives our immediate attention. However,
the enormous complexity of geographic space and changes in its component ecosystems through
time require that major attention be directed to atmospheric, geologic, physiographic, and soil
properties of forest landscapes. In summary, for every landscape, a combination of factors should
be used to distinguish the pattern of local and regional landscape and waterscape ecosystems that
have similar ecological potential in the long run. Understanding ecological units at multiple spatial
scales (Chapters 2, 11, and 22) is needed to provide the basis for monitoring ecosystem change over
time and for ecosystem management.
AN APPROACH TO THE STUDY OF FOREST ECOLOGY

This book presents the scope of forest ecology as having at least two major parts. The first is
analysis: studying the details of ecosystem components (atmosphere, organisms, and Earth). The
second part is synthesis: understanding the broad ecological framework within which ecosystem
components are integrated. We therefore emphasize a “big picture” approach even as we discuss
much smaller details of the organic and inorganic parts of ecosystems, their interactions, and the
structure and function of entire ecosystems. The explicit separation and analysis of the biotic and
physical parts of ecosystems are not realistic or desirable from an ecological perspective, but may
serve as reasonable subdivisions to provide both analysis and synthesis. The sequence of presen-
tation is (i) understanding forest ecosystems at multiple spatial scales, (ii) the variation, life
history, and structure of the forest tree, (iii) the physical environment of forests, (iv) forest com-
munities, (v) forest ecosystem dynamics, and (vi) the outlook for and consideration of future forests.
In Part 1 of this book, landscape ecosystems are the primary objects of interest, but we
observe the importance of understanding spatial scale in examining them. Spatial scale influ-
ences the observations we make about ecosystems because ecological processes vary as spatial
scale changes. Advances in hierarchy theory in ecology have helped us to understand ecosys-
tems as volumetric segments of the ecosphere within which smaller volumetric units
are nested.
In Part 2, we consider that the forest tree largely owes its appearance, rate of growth, and size
to the environment in which it has grown throughout its life. In other words, the phenotype, the
individual as it appears in the forest, is the product of the environment acting on its genotype, its
individual genetic constitution. An understanding of trees as parts of ecosystems depends, in part,
upon the genetics of the trees themselves and the way their genetic heritage affects their response
to the environment they live in. In addition, life history features of regeneration as well as anatomical
and physiological aspects of forest trees in relation to environment are considered.
We examine the physical factors of forest ecosystems in Part 3. The site is the sum total of
environmental factors surrounding and available to the plant at a specific geographic place. These
factors are primarily the atmospheric, physiographic, and soil components of the physical environ-
ment, but also include important influences of growing vegetation which itself affects the micro-
climate and soil. Climate includes various atmospheric factors that vary across hours, days,
months, and years, such as solar radiation, air temperature, precipitation, humidity, wind, and
carbon dioxide content. We provide special consideration to sunlight (solar radiation) and tem-
perature, and we examine the relations and dynamics of water throughout the text in appropriate
contexts. Physiological processes related to photosynthesis, respiration, and growth are included in
Part 3 in chapters on light and temperature, but also in later chapters on carbon balance and nutri-
ent cycling.
Physiography—comprising landforms and soil parent material——plays a key role in
affecting climate as well as the amount and rates at which radiation and moisture are received and
distributed in forest ecosystems. Below the ground surface, the supply of soil moisture and nutri-
ents, the physical structure of the soil, microbial communities in the soil, and the nature and
decomposition pattern of organic matter, that is, factors pertaining to edaphic characteristics of
soil, all affect the growth and development of plants. We also treat fire as a site factor. The study of
the environmental factors and their effects on individual plants constitutes the field of autecol-
ogy, and we summarize those aspects of forest autecology most pertinent to an understanding of
forest ecosystems. Finally, we examine site quality and its evaluation, focusing on the degree to
which individual site factors and combinations of them are used to estimate the productivity of
forest ecosystems and determine management prescriptions for them.
In Part 4, we consider forest communities and their plants and animals as integral parts of
ecosystems. First, we examine the important roles of animals in affecting all phases of plant
development, as well as their effects on forest communities and ecosystem processes. Forest com-
munities are then treated, emphasizing their composition, occurrence, and the interactions (with
emphasis on competition and mutualism) of their constituent individuals. Finally, we examine
concepts of biological and ecological diversity, including their importance, measurement, and
conservation.
Forest ecosystems are ever changing, and we examine their dynamics in Part 5. We first con-
sider change in ecosystems over hundreds to millions of years. Primary bases for this change come
from geologic and physiographic studies of the land itself and determination of plant species and
their communities as deduced from the paleoecological record. Disturbance and succession as eco-
system processes are treated next, followed by chapters on whole ecosystem functioning, including
the carbon balance of trees and ecosystems and the dynamics of nutrients.
Part 6 examines future forest ecosystems in considering their sustainability. We first examine
the effects of climate change on forest ecosystems, including effects on individuals, populations,
communities, and whole ecosystems. We also explore the potential for mitigation of climate change
using forest management. We next consider the importance of invasive plants and animals for
forest ecosystem structure and function. Finally, we place humans in an appropriate context of
forest ecosystems in considering forest landscape ecology and the emerging concept of forest
sustainability.
APPLICABILITY TO FOREST MANAGEMENT

The management of forests is, as it should be, a continuously evolving effort as new science, tech-
nology, and consideration of the role of humans in forest ecosystems develop. Management of
forests may include decisions made at broad or local scales, but consideration is increasingly given
to whole systems, their complexity and heterogeneity, as well as their long-term sustainability and
integrity (Franklin et al. 2018) rather than single-focus use of their parts for recreation, wildlife, or
Applicability to Forest Management 13
timber. To ecologists, understanding ecosystems for their own sake has significant value in itself,
but the principles of forest ecology are essential for their guidance in conservation and management
practices. Spurr (1945) defined silviculture, or applied forest ecology, as the theory and practice
of controlling forest establishment, composition, and growth. Modern texts suggest that current
silviculture is most concerned with managing forests for future products and services in a manner
that has social stability (Ashton and Kelty 2018). Nevertheless, we explicitly emphasize that silvi-
culture is the theory and practice of controlling forest ecosystem composition, structure, function,
and heterogeneity, often at multiple spatial and temporal scales. Even local management opera-
tions may have significant and often long-term effects on surrounding landscapes far removed
from the specific site of human activity.
The management, conservation, and restoration of forested landscape ecosystems rely
heavily on understanding the structure and function of forest ecosystems and the autecology of
their organisms (i.e., forest ecology). More than ever, understanding how ecological systems
work is critical for making sound decisions regarding human intervention in forests. Under-
standing the structure and function of whole ecosystems will also equip us to best deal with the
ecological consequences of past human activities, such as deforestation and its wide range of
impacts, habitat loss, and fragmentation. Human activities associated with industrialization,
such as air pollution or the introduction of destructive new species, have resulted in local tree
mortality and, in certain ecosystems, widespread death and decline of forest trees and other
organisms. Increasing the carbon dioxide concentration in the atmosphere due to the burning
of fossil fuels and deforestation are creating climatic changes and concomitant changes in
forest ecosystems that are quickly increasing in their severity. Invasive plants and animals
introduced into forests are well known for disrupting ecosystem structure and function on
relatively short time scales. In all cases, one must understand the system in order to understand
the extent to which it is disrupted.
At the time of the last edition of this textbook over 20 years ago, a new paradigm in for-
estry had occurred that shifted the view of the forest from one of single commodities (timber,
wildlife, water, and recreation) to forestry understood as sustaining ecosystems and their
structure, diversity, heterogeneity, and function (Rowe 1994; Kohm and Franklin 1997). That
paradigm emphasized maintaining the integrity of ecosystems across landscapes by sustaining
their natural patterns and processes even when heavily manipulated by humans. This paradigm
has been further refined over the past two decades to include an emphasis on considering or
even incorporating natural disturbances, increasing ecosystem resistance and resilience,
embracing structural complexity, and maintaining a range of ecosystem conditions across
broad scales (Franklin et al. 2018). Whether intentionally or otherwise, this paradigm has the
landscape ecosystem approach at its core because it focuses on the maintenance of landscapes
as complete ecosystems. The only way to assure the sustained yields of forests, wildlife, and
water, now and in the future, is to maintain the integrity of their processes and keep their
biota in a healthy state. Such a practice demands an intimate familiarity with forest ecology as
we have described it.
SUGGESTED
R E A D I N G S
Bailey, R.G. (2009). Ecosystem Geography, 2e. (Chapters 1 Golley, F.B. (1993). A History of the Ecosystem Concept
and 11). New York: Springer-Verlag 251 pp. + 1 map. in Ecology. New Haven, CT: Yale Univ. Press 254 pp.
Chapin, F.S. III, Matson, P.A., Vitousek, P., and Chap- Franklin, J.F., Johnson, K.N., and Johnson, D.L. (2018).
in, M.C. (2012). Principles of Terrestrial Ecosystem Ecological Forest Management. (Chapter 1). Long
Ecology, 2e. New York: Springer 520 pp. Grove, IL: Waveland Press 646 pp.
Kohm, K.A. and Franklin, J.F. (ed.) (1997). Creating a Rowe, J.S. (1994). A new paradigm for forestry. For.
Forestry for the 21st Century. Washington, D.C: Island Chron. 70: 565–568.
Press 475 pp. Tansley, A.G. (1935). The use and abuse of vegetational
Rowe, J.S. (1961). The level-of-integration concept and concepts and terms. Ecology 16: 284–307.
ecology. Ecology 42: 420–427.
Rowe, J.S. (1992). The ecosystem approach to forestland
management. For. Chron. 68: 222–224.
Landscape Ecosystems at
CHAPTER 2
Multiple Scales
F orest ecologists often by default are also landscape ecologists—people whose interests and
practices necessarily encompass ecosystems of many different spatial scales. A landscape
is a heterogeneous land area composed of a group of intermeshed ecosystems, each with inter-
acting atmosphere, earth, and biota. Landscape ecology is the study of the spatial patterns of
ecosystems, the causes of the pattern, and how the pattern affects ecological processes. Many
of the core concepts of landscape ecosystems overlap with those of landscape ecology, but the
domain of interest of the two fields is not identical. In this chapter, we examine the spatial occur-
rence and complexity of landscape ecosystems, whereas the discipline of landscape ecology is
considered in Chapter 22.
OVERVIEW OF SPATIAL AND TEMPORAL SCALES

Forest ecologists ask questions, conduct field experiments, and resolve problems at different scales
of space and time, and these scales directly influence the problems posed, the questions asked, and
the techniques employed. Scale describes the dimensions of an object or process in space or time
(Turner and Gardner 2015). Scale is often confused with biological or ecological levels of organiza-
tion, such as cells, tissues, organs, organisms, and ecosystems (see Figure 1.2, Chapter 1), but the
concepts are quite distinct. For example, organisms are obviously an important level of organiza-
tion, but they may occur over very different spatial scales (e.g., 1–10 μm for bacteria to 84 m tall
giant sequoia) with very different life spans (e.g., 24 hours for mayflies to nearly 5000 years for
bristlecone pine). A given scale is defined by its grain (its finest spatial or temporal resolution) and
its extent (the size of a study area or the length of time considered), and there is some trade-off
between the two characteristics. For example, forest ecologists might increase grain size (e.g.,
sample plot size) in a larger study area (e.g., the size of a county) because of their interest in
capturing variation in species composition at a larger spatial scale. However, they might reduce
grain size (using smaller plots or quadrats) if they are interested in describing species variation
across a 1 ha woodlot. As a result, no single set of spatial and temporal scales is applicable to all the
questions addressed by forest ecologists (Turner and Gardner 2015).
Processes that occur at a given scale do so within a hierarchy. The process in question is
provided context (influenced and constrained) by the processes at the hierarchical level above it,
while the level below it provides its mechanisms. For example, it is vital in understanding the
growth of trees in forested wetlands to examine those processes that occur at broader and finer
spatial scales than those of an individual tree. Wetland tree growth is constrained by broader-scale
processes such as landscape position and soils, but the mechanisms that influence tree growth,
such as tree physiology, occur at finer scales. A classical hierarchical model shown in Figures 2.1
and 2.2 suggests that many forest managers, ecologists, and naturalists would have an interest at a
microscale (up to 100 ha and 500 years), which would encompass the size and life span of a typical
forest in North America (Delcourt and Delcourt 1988). Disturbance events such as clear-cuts, some
wildfires, and wind events, as well as biotic responses such as gap-phase replacement, competition,
15
16 Chapter 2 Landscape Ecosystems at Multiple Scales
SCALE PARADIGM F I G U R E 2.1 General spatial–temporal scales for

the hierarchical characterization of disturbance
109 regimes, ecosystem and biotic responses, and
megascale
vegetation units. Quaternary studies at the macro-
scale refer to investigations of species migrations and
extinctions in the time frame of 10 000–1 000 000 years
in the Quaternary Period (see Chapter 15) on a
TEMPORAL SCALE (yr)
continental scale. Source: After Delcourt and

106 Delcourt (1988) / Springer Nature.
macroscale
(QUATERNARY STUDIES)
mesoscale
103 (LANDSCAPE ECOLOGY)
microscale
100
100 104 108 1012
SPATIAL SCALE (m2)
and productivity might occur at this spatiotemporal scale (Figure 2.2). Many landscape ecologists
are concerned with the mesoscale (100–10 000 ha and 500–100 000 years), which encompasses
longer-term processes such as fire regimes, insect or pathogen outbreaks, and soil development, as
well as biotic processes such as secondary succession. Seasonal patterns of precipitation and
temperature and longer-term (decades to centuries) weather trends and climatic fluctuations affect
the distribution, growth, and interactions of plants and animals at microscale and mesoscale.
Macroscale and megascale (Figures 2.1 and 2.2) extend in space and time to include not only
events of ecosystem change and plant migration occurring at the level of physiographic regions
(macroscale), but also plate tectonics and evolution of biota on the spatial scale of continents and
the ecosphere (megascale).
The disturbance regimes, ecological responses, and vegetation units that are of concern to
forest ecologists may occur over a range of spatiotemporal scales rather than cleanly within one
shown in the hierarchical framework in Figure 2.2. Disturbance events such as wildfire or insect
and pathogen outbreaks may occur at a local site or over thousands of square kilometers. Wind
may uproot a single tree in an old-growth forest or may affect huge land areas for much longer
intervals, such as the 150 000 ha (370 000 acres) of forest blown down in the Boundary Waters
Canoe Area in northern Minnesota in 1999 (Mattson and Shriner 2001). Likewise, geomorphic
processes such as soil creep, debris avalanches, stream transport, and deposition of sediments
affect vegetation at levels from individual plants to large forest stands and occur on areas extending
from the size of sample plots (m2) and stream watersheds (ha) to mountain ranges (km2). Vegeta-
tion change or succession is another good example of an ecosystem process that may be examined
at different spatial and temporal scales. For example, vegetation change may occur in just a few
years in small tree-fall gaps, as widespread secondary succession over decades or centuries, or as
evolution over millennia on a continental spatial scale. Even the same disturbance type may occur
at different scales among ecosystems. For example, subalpine lodgepole pine forests of Yellowstone
National Park were historically burned by large fires (>5000 ha) every 100–300 years (Romme and
Despain 1989), which spans both the microscale and mesoscale in Figure 2.1. By contrast,
hemlock-northern hardwood forests of the Lake States were likely burned by very small fires
Spatial Scales of Hierarchical Landscape Ecosystems 17
ENVIRONMENTAL ECOSYSTEM AND

DISTURBANCE REGIMES BIOTIC RESPONSES VEGETATIONAL PATTERNS
100 104 108 1012 100 104 108 1012 100 104 108 1012
108 Global Terrestrial 108

Vegetation
Plate Evolution of
tectonics the Biota
Temporal Scale (years)
FM.
Zone
106 108
Glacial– Ecosystem Change
Interglacial Forma-
Biota Landforms tion
Climatic Cycles
Soil Development Speciation Soils
Macro-and-
Migration Microclimate
Climatic Fluctuations
103 Stand
Type 108
Human Activities Secondary Subtype
Succession
Tree
Gap-phase
Fire Regime Replacement
Pathogen Outbreak Competition Shrub
*Disturbance Events Productivity Herb
100 108
100 104 108 1012 100 104 108 1012 100 104 108 1012
*Examples: Wildfire, Spatial Scale (m2)
Wind Damage,
clear-cut, Flood,
Earthquake
F I G U R E 2 . 2 Disturbance regimes, ecosystem and biotic responses, and vegetational units viewed in the
context of four spatial–temporal scales shown in Figure 2.1. Source: Modified from Delcourt and Delcourt, 1988 /
with permission of Elsevier.
(<5 ha) over very long intervals (approximately 1000 years; Frelich and Lorimer 1991), which also
spans both the microscale and mesoscale.
SPATIAL SCALES OF HIERARCHICAL LANDSCAPE ECOSYSTEMS

The ecosphere is the largest and most all-inclusive ecosystem that includes within it volumet-
ric segments (i.e., landscape ecosystems). Three general levels of landscape ecosystems are
typically identified (Rowe and Sheard 1981; Rowe 1992b; Bailey 2009). Macroscale landscape
ecosystems (scale in this context refers to spatial scale) are seas and continents and their major
units, essentially encompassing the macro- and mega- scales shown in Figure 2.2. Continents
contain mesoscale landscape ecosystems, usually major physiographic features such as broad
plains, rolling hills, and mountain ranges. These major physiographic features contain microscale
landscape ecosystems that are local landforms supporting upland forests, swamps, and lakes.
This hierarchical framework proceeds from the more complex and heterogeneous ecosystem
units at broad spatial scales to less complex and relatively homogeneous at fine spatial scales.
Note the three levels of the hierarchy cited in the earlier text are arbitrary, as the complexity of a
landscape and the ecological questions at hand will determine the number of levels necessarily
distinguished.
Ecosystems at each level, whether at the broad regional scale (macroscale and mesoscale
landscape ecosystems) or at a local scale (microscale), all include climate, physiography, soil, and
organisms. It would be ideal to delineate and map landscape ecosystems by integrating all these
components at all levels, but this formidable task has been accomplished only at local scales (e.g.,
Barnes et al. 1982; Pregitzer and Barnes 1984; Spies and Barnes 1985a; Simpson et al. 1990). Despite
failing to integrate multiple factors, single-component classifications of ecosystems by climate,
vegetation, or physiography are useful to illustrate the spatial pattern of the focal component. In
the sections provided in the following text, we provide examples of these approaches as an avenue
to emphasize the complexity of landscape ecosystems at multiple spatial scales.
18 Chapter 2 Landscape Ecosystems at Multiple Scales
CLIMATIC CLASSIFICATION
Climate is the source of energy and water for all ecosystems, and thus it is the most important eco-
logical factor at the macroscale. Together with physiography, climate strongly controls genetic
differentiation of organisms and hence their distribution in space and time. Climate may be classi-
fied at the broadest spatial scale, such as the “ecoclimatic zones” of Bailey (1988; Figure 2.3), to
illustrate the pattern of climate across the ecosphere. In this way climate may be used to distin-
guish macroscale and mesoscale ecosystem units to which other ecosystem components can be
linked. Each of the 13 ecoclimatic zones in Figure 2.3, adapted from the climatic classification of
Köppen (1931) (modified by Trewartha 1968), is characterized by a particular climatic regime (the
characteristic seasons and daily ranges in temperature and moisture). Each climatic zone essen-
tially corresponds to a general zonal soil type (usually at the level of soil order) and a broad, late-
successional vegetation type (Table 2.1). These climatic zones span the continents and in many
cases include high mountains. Mountains provide vertical stacks of ecoclimatic zones or sequences
of altitudinal belts that are related to elevation, slope aspect, and associated climatic and soil char-
acteristics (Figure 2.4) within the context of the horizontal ecoclimatic zones. The sequence of
altitudinal belts may differ according to the geographic position of the zone in which it is located.
In this example, climate is the primary factor for classification of mountainous landscapes
into horizontal and vertical zones at the broadest spatial scales. At the scale of continents, gross
physiographic features and vegetative cover also play a major role in affecting climate both hori-
zontally and vertically. At the mesoscale, Bailey (2009, 1988) subdivides climatic zones using
physiography (specific landforms and their surficial form and parent material) as the key factor.
Physiographic features affect the distribution and composition of vegetation by influencing both
macroclimate and microclimate (Chapter 8).
0°
Ar Tropical Wet Cf Subtropical Humid

Aw Tropical Wet-Dry Do Temperate Oceanic
BSh Tropical/Subtropical Semi-arid Dc Temperate Continental
BWh Tropical/Subtropical Arid E Boreal
BSk Temperate Semi-arid Ft Polar Tundra
BWk Temperate Arid Fi Polar Ice Cap
Ca Subtropical Dry Summer H Highlands
FIGURE 2.3 Ecoclimatic zones of the world. Source: Bailey (2009) / Springer Nature.
Spatial Scales of Hierarchical Landscape Ecosystems 19
Table 2.1 Zonal relationships between climate, soil orders, and vegetation.a
Ecoclimatic zoneb Soil orderc Vegetation

Ar Oxisols (latosols) Evergreen tropical rainforest (selva)
Aw Oxisols (latosols) Tropical deciduous forest or savanna
BS Mollisols, aridisols (chestnut, brown soils, and Shortgrass
sierozems)
BW Aridisols (desert) Shrubs or sparse grasses
Cs Entisols, inceptisols Sclerophyllous woodlands
(Mediterranean brown earths)

Cf Ultisols Coniferous and mixed coniferous–
deciduous forest
(red and yellow podzolic)
Do Alfisols (brown forest and gray-brown podzolic) Coniferous forest
Dc Alfisols Deciduous and mixed coniferous–
deciduous forest
(gray-brown podzolic)
E Spodosols and associated histosols (podzolic) Boreal coniferous forest (taiga)
Ft Entisols, inceptisols, and associated histosols Tundra vegetation (treeless)
(tundra humus soils with solifluction)
a
After Walter (1984, p. 3).
b
Abbreviations of the zones are shown in Figure 2.3.
c
Names in parentheses are soil orders (USDA Natural Resources Soil Conservation Service, 1999).
Source: Bailey (1988) / United States Department of Agriculture / Public Domain.
N
6000 m
4000
2000
0
A B C D
70° 65° 55° 45° 35°

Edmonton Denver
1 2 3 4 5 6 7 8
F I G U R E 2 . 4 Vertical zonation (cover types 1–8) within boreal (B) and temperate semi-arid (D) ecoclimatic
zones (Figure 2.2) from 70° to 35° N latitude along the eastern slopes of the Rocky Mountains from approxi-
mately 102° to 115° W longitude. Climatic zones: A, transition; B, boreal; C, transition; D, temperate
semi-arid. Vertical zones: 1, ice region; 2, mountain vegetation above tree line; 3, boreal and subpolar open
coniferous woodland; 4, boreal evergreen coniferous forest; 5, boreal evergreen mountain coniferous forest;
6, coniferous dry forest; 7, shortgrass dry steppe; 8, boreal evergreen coniferous forest with cold-deciduous
broad-leaved trees. Source: Rowe (1988) / United States Department of Agriculture / Public Domain.
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The Project Gutenberg eBook of Joseph
Hergesheimer, an essay in interpretation
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Title: Joseph Hergesheimer, an essay in interpretation
Author: James Branch Cabell
Release date: December 2, 2023 [eBook #72292]
Language: English
Original publication: Chicago: The Bookfellows, 1921
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*** START OF THE PROJECT GUTENBERG EBOOK JOSEPH

HERGESHEIMER, AN ESSAY IN INTERPRETATION ***
Joseph Hergesheimer
An Essay in Interpretation
Joseph Hergesheimer
An Essay in Interpretation
By
James Branch Cabell
“And we dreamed a dream in one night, I and he: we dreamed

each man according to the interpretation of his dream.”
CHICAGO
THE BOOKFELLOWS
1921
One thousand small paper and ninety-nine tall paper copies of this monograph
have been printed for The Bookfellows in August, 1921. The edition is the first;
Mr. Cabell the author is Bookfellow No. 513 and Mr. Brewer the printer is
Bookfellow No. 14.
Copyright 1921 by
James Branch Cabell
To
JOSEPH HERGESHEIMER
with friendship and large admiration

as goes the past, and with
cordial faith in what
is to come.
ONE
O say they, speak they, and tell they the tale, in “literary
gossip,” that Joseph Hergesheimer “wrote” for a long while
before an iota of his typing was transmuted into “author’s proof.” And
the tale tells how for fourteen years he could find nowhere any
magazine editor to whose present needs a Hergesheimer story was
quite suited.
It is my belief that in approaching Mr. Hergesheimer’s work one
should bear constantly in mind those fourteen years, for to me they
appear, not uncuriously, to have shaped and colored every book he
has thus far published.
The actual merit of the writing done during that period of
“unavailability” is—here, at least—irrelevant. It is not the point of the
fable that he high-heartedly wrote a story to which, when completed,
his unbiased judgment could not quite honestly deny such deference
as is due to a literary masterpiece; and which, through some odd
error, was rejected by a magazine that every month was publishing
vastly inferior stories; and which was later declined by another
magazine, and by a host of magazines, with a dispiriting bland
unanimity not unsuggestive of editorial conspiracy. Meanwhile—of
course—he had written another tale, which was much better than the
first, and which proved to be an equally faithful chaperon of return
postage. So story followed story, each dreeing the same weird....
And he used to wait for the postman, no doubt, and to note from
afar that it was a large envelope; and would open the damned thing
with a faint hope that perhaps they just wanted some slight changes
made; and would find only the neat, impersonal, and civilly
patronizing death-warrant of hope. So Joseph Hergesheimer kept on
with his foolishness, without any gleam of success, or even (they
report) any word of encouragement. And doubtless his relatives said
the customary things....
Yet none of these circumstances, either, is the point of the
apologue, because in all save one detail the comedy has been
abraded into pointlessness by over-constant repetition; and is, of
course, being futilely performed at this moment in one prefers not to
reflect how many thousand homes. The leading rôle, though, is too
unprofitable and irksome for any quite sane person to persist in
enacting it for fourteen years. This Joseph Hergesheimer did: and
that is the fable’s significant point.
TWO
ES, it is the boy’s illogical pertinacity that is the fable’s point,

because it so plausibly explains why nearly all the men in Mr.
Hergesheimer’s books are hag-ridden by one or another sole desire
which spurs them toward a definite goal at every instant of their
mimic lives. These men but variously reflect, I take it, that younger
Hergesheimer’s “will to write,” that unconquerable will. To Mr.
Hergesheimer, even to-day, it probably seems natural that a man’s
whole living should be devoted to the attaining of one desire quite
clearly perceived, because his own life has been thus dedicated. The
more shrewd mass of practical persons that go about in flesh are
otherwise; and comfortably fritter through the day, with no larger
objective at any time in mind than the catching of a car, the rounding
off of a business transaction, the keeping of an engagement for
luncheon, and the vesperal attendance to some unmental form of
recreation, with one small interest displacing another in endless
succession, until bedtime arrives and the undertaker tucks them in.
It explains to me the Hergesheimer women, too, those troublingly
ornamental odalisques. They are fine costly toys, tricked out in
curious tissues: and, waiting for the strong male’s leisure, they smile
cryptically. They will divert him by and by, when the day’s work is
dispatched, maintaining their own thoughts inviolate, even in the
instant of comminglement wherein the strongest man abates
reserve: but their moment is not daylit, for the Hergesheimer women
are all-incongruous with what is done during office hours, nor are
they to be valued then. Sometimes they are embodied ideals, to be
sure, remotely prized as symbols or else grasped as trophies to
commemorate the nearing of the goal: but for the most part they rank
candidly as avocational interests. I find nowhere in Joseph
Hergesheimer’s stories any record of intimacy and confidence
between a man and a woman.... And this too, I think, reflects that all-
important formative fourteen years wherein, whatever may have
been Mr. Hergesheimer’s conduct of his relatively unimportant
physical life, his fundamental concernments were pursued in a
realm, of necessity, uninhabited by women. Indeed, no woman can
with real content permit the man whom she proprietorially cherishes
to traffic in this queer lonely realm, and she cannot but secretly
regard his visits thereto as a personal slight. So the creative literary
artist is (when luckiest) at silent feud with his women, because the
two are perpetually irritated by the failure of their joint effort to ignore
the fact that she ranks necessarily as an avocational interest.
THREE
HAT, though, was the precise goal of the fourteen years of

visually unproductive “writing”? Those earlier stories have
never been printed, so that one perforce advances on a bridge of
guesswork. But certainly, in all that is to-day accessible of Mr.
Hergesheimer’s creative feats—with one exception duly noted
hereinafter—there is a patent negligence, and indeed an
ostentatious avoidance, of any aiming toward popularity. That during
the fourteen years young Hergesheimer labored toward the applause
and cheques of a “best seller,” is to the considerate inconceivable.
Nor could that well have been a motive strong enough to sustain him
thus long, since the maker of reading-matter, like any other
tradesman, has need of quick returns where the artist battens on
immediate rejections.
No, Mr. Hergesheimer’s monomania, one estimates, was then, just
as it seems to be to-day, to write for his own delectation—in large
part because he could not help it, and in part with the hope of,
somehow and some day, obtaining an audience with the same or, at
any rate, a kindred sense of beauty.... This, to be sure, is always a
vain aspiration. That which, in effusions such as this, we loosely talk
about as “beauty” probably does not exist as a vital thing save here
and there in the thoughts of not too many and not to be too seriously
taken persons. In life, rather frequently, one appears to catch a
glimpse of something of the sort just around the corner or over the
way, but it is rarely, and perhaps never, actually at hand. Sometimes,
of course, one seems about to incorporate the elusive thing into
one’s daily living; and, striving, finds the attempt a grasping at an
opalescent bubble, with the same small shock, the same disrupting
disillusionment.
“Beauty,” thus, is by the judicious conceded to be an
unembodiable thought, not even quite to be grasped by the mind;
and certainly never nicely nor with any self-content to be
communicated via the pages of a book, wherein are preserved, at
best, the faded petals and the flattened crumbling stalks of what
seemed lovely once to somebody who is as dead as are these
desiccated relics of his ardor and of his disputable taste.
In brief, it may be granted—and by Mr. Hergesheimer most
cheerfully of all persons—that during these fourteen years Mr.
Hergesheimer was attempting the preposterously impossible.
FOUR
OW, to my thinking, there is something curiously similar to

that unreasonable endeavor to be found in all the
Hergesheimer novels. Here always I find portrayed, with an
insistency and a reiteration to which I seem to detect a queer
analogue in the writings of Christopher Marlowe, men laboring
toward the unattainable, and a high questing foiled. No one of the
five novels varies from this formula.
Anthony Ball, of The Lay Anthony, strives toward the beauty of
chastity—not morally concerned one way or the other, but resolute to
preserve his physical purity for the sake of a girl whose body, he
finds at last, has long ago been ravished by worms. Again there is in
Mountain Blood no hint of moral-mongering—for Mr. Hergesheimer
is no more concerned with moral values than is the Decalogue—
when Gordon Makimmon toils toward the beauty of atonement, to
die in all a broken man, with his high goal yet gleaming on the
horizon untouched. The three black Pennys flounder toward the
beauty of a defiant carnal passion, which through the generations
scorches and defiles, and burns out futilely by and by, leaving only
slag where the aspiring lovely fire was. And through the formal
garden ways of Java Head pass feverishly at least five persons who
struggle (and fretfully know their failure to be foredoomed) toward
the capturing of one or another evincement of beauty, with the
resultant bodily demolishment of three of them and the spiritual
maiming of the others.
That which one, for whatever reason, finds most beautiful must be
sought; it is a goal which one seeks futilely, and with discomfort and
peril, but which one seeks inevitably: such is the “plot” of these four
novels. Such is also, as I need hardly say, the “plot” of the
aforementioned fourteen years wherein not anything tangible was
achieved except the consuming of youth and postage....
Nor does the dénouement differ, either, in any of these novels: the
postman comes with the plethoric envelope which signals from afar
that the result of much high-hearted striving is not quite suited to the
present needs of this world’s editor; and sometimes the postman is
Age, but more often he is Death.
FIVE
OW the fifth, and incomparably the finest and loveliest, of the

Hergesheimer novels is Linda Condon, which renders self-
confessedly a story of “the old service of beauty, of the old gesture
toward the stars”—“here never to be won, never to be realized”—of
the service which “only beauty knows and possesses”.... For Linda
Condon is to be valued less as the life-history of a woman than as
the depiction—curt, incisive, and yet pitying—of a shrine that,
however transiently, was hallowed.
At the exacting workaday pursuit of being a human being this
Linda fails, fails chilled and wistful. She has, like more of us than
dare proclaim the defect, no talent whatever for heart-felt living, so
that most persons seem but to pass grayly upon the horizon of her
consciousness, like unintelligible wraiths gesticulating,—and always
remaining somehow disjunct and not gravely important,—the while
that all the needs and obligations of one’s corporal life must be
discharged with an ever-present sense of their queer triviality.
Toward nobody, neither toward Linda Condon’s mother nor lover, nor
husband, nor children, may she, the real Linda, quite entertain any
sense of actual attachment, far less of intimacy....
Meanwhile she has her loveliness, not of character or mind, but a
loan of surpassing physical beauty. And to Linda Condon her own
bright moving carcass becomes a thing to be tended and preserved
religiously, because beauty is divine, and she herself is estimable, if
at all, as the fane which beauty briefly inhabits.... And by and by,
under time’s handling, her comeliness is shriveled, and her lovers
are turned to valueless dust: but first, has Linda’s lost young beauty
been the buried sculptor’s inspiration, and it has been perpetuated in
everlasting bronze. The perfection of Linda Condon’s youth is never
to perish, and is not ever to be dulled by old age or corrupted in
death. She comprehends this as she passes out of the story, a
faded, desolate and insignificant bit of rubbish, contented to know
that the one thing which really meant much to her is, as if by a
miracle, preserved inviolate. The statue remains, the immutable child
of Linda’s comeliness and Pleydon’s genius, the deathless offspring
of transitory things.
Beauty is divine; a power superior and even elfinly inimical to all
human moralities and rules of thumb, and a divinity which must
unflinchingly be served: that, in this book as always, is Mr.
Hergesheimer’s text. For this is the divinity which he, too, serves
unflinchingly, with strangely cadenced evocations, in striving to write
perfectly of beautiful happenings.
It is an ideal here approached even more nobly than in the
preceding Hergesheimer books. Nowhere has Joseph Hergesheimer
found an arena more nicely suited to the exercise of his most
exquisite powers than in this modern tale of domnei,—of the worship
of woman’s beauty as, upon the whole, Heaven’s finest sample of
artistic self-expression, and as, in consequence, the most adequate
revelation of God; and as such a symbol, therefore, a thing to be
revered above all else that visibly exists, even by its temporary
possessor. That last is Mr. Hergesheimer’s especial refinement upon
a tenet sufficiently venerable to have been nodded over by Troy’s
gray-bearded councillors when Helen’s skirts were rustling by,—and
a refinement, too, which would have been repudiated by Helen
herself, who, if one may trust to Euripides’ report of her sentiments,
was inclined less elevatedly to regard her own personal appearance,
as a disaster-provoking nuisance.
Well, and to Linda, also, was beauty a nuisance—“a bitter and
luxurious god,” that implacably required to be honored with sacrifices
of common joys and ties and ruddy interests, but was none the less
divine. Sustained by this sole knowledge, Linda Hallet passes out of
the story, when youth is over, regarding not very seriously that which
is human and ephemeral, even as embodied in her lovers and her
children, nor in herself, but rather always turning grave blue eyes
toward that which is divine; passes, at once the abandoned
sanctuary, the priestess, the postulant, and the martyr, of that beauty
to which fools had once referred as “hers”; passes not as the
wreckage of a toy but as an outworn instrument which has helped to
further the proud labor of a god; and passes, as all must pass,
without any sure comprehension of achievement, but with content.
That, really, is The Happy End....
SIX
HICH reminds me that for the most part I am rattling very old
bones. Those seemingly unfruitful fourteen years are to-day at
one with those other fourteen years which brought an elder Joseph
into Egyptian publicity. Mr. Hergesheimer has “arrived”: his books
have found their proper and appreciative audience; whereas his
short stories are purchased, and probably read, along with the
encomiums of ready-made clothing and safety razors, by the I forget
how many million buyers of the world’s most popular magazine....
Now, here, I think, one finds stark provocations of uneasiness. I
speak with diffidence, and am not entirely swayed, I believe, by the
natural inclination of every writer to backbite his fellow craftsman. In
any event, dismissing Gold and Iron (after some reflection) with
unqualified applause, I take up The Happy End; and of the seven
stories contained therein six seem to me to display a cornerstone of
eminently “popular” psychology, ranging from the as yet sacrosanct
belief that all Germans are perfectly horrid people, to the axiom that
the quiet and unrespected youngest brother is invariably the one to
exterminate the family enemies, and duly including the sentiment
that noble hearts very often beat under ragged shirts. And I am
made uneasy to see these uplifting faiths—these literary baking-
powders more properly adapted to the Horrible Trites and the
Gluepot Stews among reading-matter confectioners—thus utilized by
a Joseph Hergesheimer.
I am made uneasy because I reason in this way: when Mr.
Hergesheimer consciously is writing a short story to be printed next
to advertising matter in some justly popular periodical, Mr.
Hergesheimer, being rational and human, cannot but think of the
subscribers to that popular periodical. I forget, I repeat, how many
millions of them have been duly attested upon affidavit to exist, but
certainly not many thousands of our fellow citizens can regard Mr.
Hergesheimer at his best and purest with anything save bewildered
abhorrence. So he must compromise,—subconsciously, I believe,—

Forest Ecology 5Th Edition Daniel M Kashian Full Chapter

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Forest Ecology 5th Edition Daniel M.

Burton V. Barnes (deceased)

Stephen H. Spurr (deceased)

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Library of Congress Cataloging-­in-­Publication Data

Cover Design: Wiley

Set in 9.5/12pt STIXTwoText by Straive, Pondicherry, India

PA R T 1 Forest Ecology and Landscape Ecosystems

1 Concepts of Forest Ecology 3

2 Landscape Ecosystems at Multiple Scales 15

PA R T 2 The Forest Tree

3 Forest Tree Variation 35

Periodicity of Seed Crops, 76

5 Tree Structure and Growth 95

PA R T 3 The Physical Environment

0005430834.INDD 9 01-05-2023 17:43:22

Position on slope, 172

11 Site Quality and Ecosystem Evaluation

0005430834.INDD 12 11-23-2022 12:32:40

Ecosystem Classification in the Southwestern United States, 262

12 Animals in Forest Ecosystems 269

13 Forest Communities 291

0005430834.INDD 13 11-23-2022 12:32:40

Phytosociology in Europe, 298

14 Diversity in Forests 327

0005430834.INDD 14 11-23-2022 12:32:40

Diversity of Landscape Ecosystems, 336

PA R T 5 Forest Ecosystem Dynamics

15 Long-­Term Forest Ecosystem and Vegetation Change 367

0005430834.INDD 15 11-23-2022 12:32:40

Eastern North America, 370

0005430834.INDD 16 11-23-2022 12:32:41

Biotic Composition Changes, 411

17 Forest Succession 413

18 Carbon Balance of Trees and Ecosystems 449

0005430834.INDD 17 11-23-2022 12:32:41

Temperature and Leaf C Gain, 454

19 Nutrient Cycling 487

0005430834.INDD 18 11-23-2022 12:32:41

PA R T 6 Forests of the Future

20 Climate Change and Forest Ecosystems 537

0005430834.INDD 19 11-23-2022 12:32:41

21 Invasive Species in Forest Ecosystems 587

22 Forest Landscape Ecology 611

0005430834.INDD 20 11-23-2022 12:32:41

23 Sustainability of Forest Ecosystems 639

Scientific Names of Trees and Shrubs 731

0005430834.INDD 21 11-23-2022 12:32:41

F orest ecologists work to understand the dynamics, structure, and function of

F I G U R E 1 . 1 The three-­dimensional, volumetric nature of a landscape ecosystem. Ecosystems comprise the

Upper Air Meteorology

Lower Air Topoclimatology

Mineral Strata Bedrock Geology

Water Table Hydrology

LANDSCAPE ECOSYSTEM AND COMMUNITY

ECOSYSTEM STRUCTURE AND FUNCTION

dead living material or energy flow other influences

Humans as a partner *relations omitted

EXAMPLES OF LANDSCAPE ECOSYSTEMS

(a) Ecosystem Type

(b) Forest Cover Type

Cutting Type Transect

AN APPROACH TO THE STUDY OF FOREST ECOLOGY

Library of Congress Cataloging-in-Publication Data

1 Concepts of Forest Ecology 3

2 Landscape Ecosystems at Multiple Scales 15

3 Forest Tree Variation 35

5 Tree Structure and Growth 95

12 Animals in Forest Ecosystems 269

13 Forest Communities 291

14 Diversity in Forests 327

15 Long-Term Forest Ecosystem and Vegetation Change 367

17 Forest Succession 413

18 Carbon Balance of Trees and Ecosystems 449

19 Nutrient Cycling 487

20 Climate Change and Forest Ecosystems 537

21 Invasive Species in Forest Ecosystems 587

22 Forest Landscape Ecology 611

23 Sustainability of Forest Ecosystems 639

Scientific Names of Trees and Shrubs 731

F I G U R E 1 . 1 The three-dimensional, volumetric nature of a landscape ecosystem. Ecosystems comprise the