Forest Ecology

Daniel M. Kashian, Professor of Biological Sciences, Department of Biological Sciences, Wayne State University, Detroit, USA.

Donald R. Zak, Alexander H. Smith Distinguished University Professor of Ecology; Arthur F. Thurnau Professor, School for Environment and Sustainability, University of Michigan, Ann Arbor, USA.

Burton V. Barnes (deceased) was an integral part of the authorship of Forest Ecology from its 2nd edition.

Stephen H. Spurr (deceased) authored the original Forest Ecology book and subsequent revisions.

Preface

PART 1 Forest Ecology and Landscape Ecosystems

CHAPTER 1 CONCEPTS OF FOREST ECOLOGY

Ecology

Landscape Ecosystems

Landscape Ecosystem and Community

Ecosystem Structure and Function

Examples of Landscape Ecosystems

An Approach to the Study of Forest Ecology

Applicability to Forest Management

Suggested Readings

CHAPTER 2 LANDSCAPE ECOSYSTEMS AT MULTIPLE SCALES

Overview of Spatial and Temporal Scales

Spatial Scales of Hierarchical Landscape Ecosystems

Climatic Classification

Physiography

Vegetation Types and Biomes

Distinguishing and Mapping Landscape Ecosystems at Multiple Spatial Scales

Regional Landscape Ecosystems

Regional Landscape Ecosystems of Michigan

Local Landscape Ecosystems

Local Landscape Ecosystems in Upper Michigan

Suggested Readings

PART 2 The Forest Tree

CHAPTER 3 FOREST TREE VARIATION

Components of Phenotypic Variation

Plasticity of the Phenotype

Sources of Variation

The Evolutionary Sequence

Sexual and Asexual Systems

Genetic Diversity of Woody Species

Genecology

Patterns of Genecological Differentiation

Genecological Categories

Factors Eliciting Genecological Differentiation

Growth Cessation

Growth Resumption

Examples of Genecological Differentiation

Eastern North American Species

Scots Pine

Wide-Ranging Western North American Conifers

Ponderosa Pine

Douglas-fir

Local Genecological Differentiation

Factors Affecting Differentiation: Gene Flow and Selection Pressure

Ecological Considerations at the Species Level

Niche

Hybridization

Polyploidy

The Fitness–Flexibility Compromise

Epigenetics

Suggested Readings

CHAPTER 4 REGENERATION ECOLOGY

Regeneration

Sexual Reproduction

Maturation and the Ability to Flower

Increasing Seed Production

Reproductive Cycles

Pollination

Periodicity of Seed Crops

Effects of Reproduction on Vegetative Growth

Dispersal

Seed Bank, Dormancy, and Germination

Establishment Following Sexual Reproduction

Post-Establishment Development

Vegetative Reproduction

Suggested Readings

CHAPTER 5 TREE STRUCTURE AND GROWTH

Tree Form

Architectural Models

Short and Long Shoots

Patterns of Intermittent Growth

Sylleptic and Proleptic Shoots

Roots

Kinds, Forms, and Occurrence

Fine Root Relations

Horizontal and Vertical Root Development

Periodicity of Primary Root Growth

Root Grafting

Specialized Roots and Buttresses

Stems

Xylem Cells and Growth Rings

Periodicity and Control of Secondary Growth

Control of Earlywood and Latewood Formation

Winter Freezing and Water Transport

Water Deficits and Tree Growth

Suggested Readings

PART 3 The Physical Environment

CHAPTER 6 LIGHT

Distribution of Light Reaching the Ecosphere

Plant Interception of Radiation

Canopy Structure and Leaf Area

Light Quality Beneath the Forest Canopy

               Sunflecks

Light and Growth of Trees

               Light and Seedling Survival and Growth

Light and Tree Morphology and Anatomy

               Light and Epicormic Sprouting

Photocontrol of Plant Response

Light and Ecosystem Change

Suggested Readings

CHAPTER 7 TEMPERATURE

Geographical Patterns of Temperature

Temperatures at the Soil Surface

Temperature within the Forest

Temperature Variation with Local Topography

Temperature and Plant Growth

Cold Injury to Plants

Dormancy

Frost Hardiness and Cold Resistance

Thermotropic Movements in Rhododendrons

Winter Chilling and Growth Resumption

Natural Plant Distributions and Cold Hardiness

Deciduousness and Temperature

Suggested Readings

CHAPTER 8 PHYSIOGRAPHY

Concepts and Terms

Characteristics of Physiography and Their Significance

Physiographic Setting

Specific Landforms

Elevation

Form of Landforms

Level Terrain

Sloping Terrain

Slope Characteristics

Position on Slope

Aspect

Slope Inclination

Parent Material in Relation to Landform

Position of Landform in the Landscape

Multiple Roles of Physiography

Physiographic Diversity, Landscape Ecosystems, and Vegetation

Mountainous Physiography

Mountainous Terrain of California and the Pacific Northwest

Physiography and Forests of the Central Appalachians

Flatlands

The Great Plains

Pine Savannas of the Western Great Lakes Region

Till Plains of the Midwest

Southeastern and Southern Coastal Plain

Floodplains

Physiography and Firebreaks

Microlandforms and Microtopography

Tree Uprooting and Pit-and-Mound Microtopography

Microtopography and Regeneration in Hardwood Swamps

Suggested Readings

CHAPTER 9 SOIL

Parent Material

Soil Formation

Soil Profile Development

Physical Properties of Soil

Soil Texture

Soil Structure

Soil Color

Soil Water

Physical Properties of Water

Soil Water Potential

Chemical Properties of Soil

Clay Mineralogy

Cation Exchange and the Supply of Nutrients

Soil Acidity

Soil Organic Matter

Soil Classification

Landform, Soil, and Forest Vegetation: Landscape Relationships

Suggested Readings

CHAPTER 10 FIRE

Fire and the Forest Tree

Causes

Fire Regime

Fire Types, Frequency, and Severity

Fire Adaptations and Key Characteristics

Strategies of Species Persistence

Closed-Cone Pines

Fire and the Forest Site

Indirect Effects

Direct Effects

Organic Matter and Erosion

Beneficial Effects of Fire

Suggested Readings

 

CHAPTER 11 SITE QUALITY AND ECOSYSTEM EVALUATION AND CLASSIFICATION

Direct Measurement of Forest Productivity

Tree Height as a Measure of Site

Site-Index Curves

Comparisons between Species

Advantages and Limitations

Vegetation as an Indicator of Site Quality

Species Groups of Ground Cover

Indicator Plants of Coastal British Columbia

Ecological Species Groups

Plant Associations and Habitat Types in the Western United States

Operational Site Classification Based on Vegetation

Applications and Limitations of Vegetation

Environmental Factors as a Measure of Site

Climatic Factors

Physiographic Land Classification

Physiographic and Soil Factors: Soil-Site Studies

Soil Surveys

Multiple-Factor Methods of Site and Ecosystem Classification

Ecosystem Classification and Mapping in Baden-Württemberg

Applications of Multifactor Methods in the United States and Canada

Ecosystem Classification and Mapping in Michigan

Ecosystem Classification in the Southeastern United States

Ecosystem Classification in the Southwestern United States

National Classification Systems in the United States

Ecological Land Classification in Canada

Hills’ Physiographic Approach

Other Approaches Used in Canada

Suggested Readings

PART 4 Forest Communities

CHAPTER 12 ANIMALS IN FOREST ECOSYSTEMS

Plant Defense

Investment in Plant Defense

Plant Defense Against Insects

Examples of Injury and Plant Defense

Nutrition

Plant Hybrid Zones as Reservoirs for Insect Diversity

Plant Defense Against Mammals

Roles of Animals in Plant Life History

Pollination

Seed Dispersal

Fish and Reptiles

Birds

Mammals

Germination and Establishment

Decomposition, Mineral Cycling, and Soil Improvement

Damage and Death

 

Influence of Livestock on Forest Ecosystems

Suggested Readings

CHAPTER 13 FOREST COMMUNITIES

Community Concept

Grounding Communities

Florida Keys

Interior Alaska

Southern Illinois

View from the Past: Community Concepts

Schools and Terminology

Concepts of Clements and Gleason

Phytosociology in Europe

Continuum Concept

Community as a Landscape Property

Examples of Spatial Variation in Forest Communities

Discrete Forest Communities

Coastal California: Giant and Pygmy Forests        

Forest–Grassland Ecotone

Alpine Tree Lines

Merging Forest Communities

Eastern Deciduous Forest—Southern Appalachians

New England

Competition and Niche Differentiation

Interactions Among Organisms

Mutualisms in Forest Ecosystems

Symbiotic Mutualisms—Mycorrhizae

Nonsymbiotic Mutualisms

Competition

               Forest Community Structure and Composition

                              Vertical Structure

                              Stand Density

Competition and Overstory Composition

Competition in the Understory

Understory Tolerance

Characteristics of Understory-Tolerant and -Intolerant Species

Tolerance Ratings of Tree Species

Examples of Understory Tolerance in Forest Ecosystems

Nature of Understory Tolerance

Environmental Factors Relating to Understory Tolerance

Physiological Processes Relating to Tolerance

Suggested Readings

CHAPTER 14 DIVERSITY IN FORESTS

Concepts of Biological and Ecosystem Diversity

The Value of Species Diversity

Value of Biodiversity

Common Threats to Diversity

Measuring Diversity

Levels of Diversity

Measurement

Inventory Diversity: Alpha Diversity

Differentiation or Beta Diversity

Diversity of Landscape Ecosystems

Examples of Diversity

Ground-Cover Species Diversity in Northern Lower Michigan

Ecosystem Groups

Ecosystem Types

Ecosystem Diversity

Causes of Species Diversity

Diversity at Continental and Subcontinental Scales

Paleogeography and Continental Relationships

Glaciation

Latitude and Elevation

Diversity at Local Scales

Physiography and Soil

Community Composition and Structure

Disturbance and Succession

Species of Concern in Conserving Diversity

               Foundation Species

               Keystone Species

               Endemics and Rare and Endangered Species

Diversity and the Functioning of Ecosystems

               Biodiversity–Productivity Relationship

               The Role of Biodiversity in Ecosystem Stability

Forest Management and Diversity

               Effects of Traditional Forest Management on Diversity

               Preserving Diversity in Managed Forests

               Ecological Forestry: Incorporating Biodiversity into Forest Management

                              Variable-Retention Harvest System

                                             Designing a Variable-Retention Harvest System

                                             How Well Does Variable Retention Conserve Biodiversity?

Prolog: Conserving Ecosystem and Biological Diversity

Suggested Readings

PART 5 Forest Ecosystem Dynamics

CHAPTER 15 LONG-TERM FOREST ECOSYSTEM AND VEGETATION CHANGE

Change Before the Pleistocene Age

Pleistocene Glaciations

Ecosystem and Vegetational Change Since the Last Glacial Maximum

Eastern North America

Overall Migration Sequence and Patterns

Ecosystem Change in the Southern Appalachians

Western North America

Patterns of Tree Genera and Species Migrations

Migration Irregularities and Disturbance

Migration from Glacial Microrefugia

Independent Migration and Similarity of Communities Through Time

Suggested Readings

CHAPTER 16 DISTURBANCE

Concepts of Disturbance

               Defining a Disturbance

Disturbance as an Ecosystem Process

Source of Disturbance

Major Disturbances in Forest Ecosystems

Fire

Role of Fire in Forest Ecosystems

Pines in New England and the Lake States

Western Pines and Trembling Aspen

Southern Pines

Douglas-fir in the Pacific Northwest

Giant Sequoia

Fire History and Behavior

Northern Lake States

Boreal Forest and Taiga

Northern Rocky Mountains

Fire Suppression and Exclusion

Wind

Widespread and Local Effects

Principles of Wind Damage         

Broadscale Disturbance by Hurricanes

Gulf and Southern Atlantic Coast

New England—1938 Hurricane

Wave-Regenerated Fir Species

Floodwater and Ice Storms

Insects and Disease

Catastrophic and Local Land Movements

Logging

Land Clearing

Disturbance Interactions

Biotic Composition Changes

Elimination of Species

Addition of Species

Suggested Readings

CHAPTER 17 FOREST SUCCESSION

Basic Concepts of Succession

Primary and Secondary Succession

Biological Legacies

Successional Pathways, Mechanisms, and Models

Autogenic and Allogenic Succession

How Is Succession Determined?

Evolution of the Concept of Forest Succession

Formal Ecological Theory

How Does Succession Work?

Clementsian Succession

Stages of Succession

Primary Succession

Secondary Succession

 Successional Causes, Mechanisms, and Models

Key Characteristics and Regeneration Strategies

Availability and Arrival Sequence of Species

Facilitation, Tolerance, and Inhibition

Change in Ecosystems

End Point of Succession?

Succession as an Ecosystem Process

Examples of Forest Succession

Primary Succession on Recently Deglaciated Terrain

Succession Following the Eruption of Mount St. Helens

Secondary Succession Following Fire in Ponderosa Pine Forests of Western Montana

Secondary Succession and Gap Dynamics

Gap Specialists: American Beech and Sugar Maple

Fire and Oak Dominance—Oaks at Risk

Suggested Readings

CHAPTER 18 CARBON BALANCE OF TREES AND ECOSYSTEMS 

Carbon Balance of Trees

Photosynthesis, Dark Respiration, and Leaf C Gain

Light and Leaf C Gain

Temperature and Leaf C Gain

Water and Leaf C Gain

Soil Nitrogen Availability and Leaf C Gain

Construction and Maintenance Respiration

Allocation to Structure, Storage, and Defense

Light and C Allocation

Soil Nitrogen Availability and C Allocation

Carbon Balance of Ecosystems

Biomass and Productivity of Forest Ecosystems

Measurement of Biomass and Productivity

Climate and Productivity

Soil Properties, Forest Biomass, and ANPP

Biomass Accumulation During Ecosystem Development

Soil N Availability and Belowground Net Primary Productivity

Suggested Readings

CHAPTER 19 NUTRIENT CYCLING

Nutrient Additions to Forest Ecosystems

Mineral Weathering

Atmospheric Deposition

Biological Fixation of Nitrogen

Nutrient Cycling within Forest Ecosystems

Nutrient Transport to Roots

Nutrient Uptake and Assimilation by Roots

Root Architecture, Mycorrhizae, and Nutrient Acquisition

Root Architecture

Mycorrhizae

Plant Litter and the Return of Nutrients to Forest Floor and Soil

Leaf and Root Litter Production

Nutrient Retranslocation

Nutrients in the Forest Floor

Organic Matter Decomposition and Nutrient Mineralization

Biochemical Constituents of Plant Litter

Dynamics of Decomposition

Nitrogen Immobilization and Mineralization

Nitrogen Availability in Forest Ecosystems

Nitrification

Nutrient Loss from Forest Ecosystems

Nutrient Leaching from Forest Ecosystems

Denitrification

The Cycling and Storage of Nutrients in Forest Ecosystems

Nutrient Storage in Boreal, Temperate, and Tropical Forests

The Nitrogen and Calcium Cycle of a Temperate Forest Ecosystem

Ecosystem C Balance and the Retention and Loss of Nutrients

Forest Harvesting and Nutrient Loss

Suggested Readings

PART 6 Forests of the Future

CHAPTER 20 CLIMATE CHANGE AND FOREST ECOSYSTEMS

Climate Change Concepts

               Effects on Temperature

               Effects on Precipitation

Climate Change Effects on the Forest Tree

               Tree Growth and Mortality

               Phenology

               Regeneration

Climate Change Effects on Tree Species Distributions

               Observed Range Shifts

               Projected Changes in Tree Species Distributions

               Projected Changes in Forest Type Distributions

Climate Change Effects on Forest Disturbances

               Fires

               Insects and Pathogens

               Wind

Climate Change Effects on Forest Carbon

               Climate Change Effects on Carbon Gain: Primary Productivity

               Climate Change Effects on Carbon Loss: Heterotrophic Respiration

               Feedbacks Among Disturbance, Climate Change, and Carbon in Forests

                              Fire, Carbon, and Climate Change in Forests of Yellowstone National Park

Adapting to Climate Change Effects on Forests

               Assisted Migration

               Refugia

               Forest Carbon Management

Suggested Readings

CHAPTER 21 INVASIVE SPECIES IN FOREST ECOSYSTEMS

Concepts of Invasive Species

               Definition of Invasive Species

               Characteristic Traits of Invasive Plant Species

               Non-Plant Invasive Species in Forests

Impacts of Invasive Species on Forests

               Impacts of Invasive Plants on Forests

                              Competition

                              Altered Fire Regimes

                              Carbon and Nutrient Cycling

               Impacts of Invasive Insects and Pathogens on Forests

                              Chestnut Blight, Dutch Elm Disease, and Forest Succession

               Impacts of Invasive Animals on Forests

A Primer of Invasive Species Management in Forests

               Early Intervention Strategies

               Management Approaches for Established Invasive Species

               Novel Ecosystems and Invasive Species

Suggested Readings

CHAPTER 22 FOREST LANDSCAPE ECOLOGY

Concepts of Landscape Ecology

Forest Fragmentation and Connectivity

               Patches in Forest Ecology

               Forest Fragmentation

               Ecological Effects

               Connectivity

Disturbances on Landscapes

               Effects of Heterogeneity on Disturbances

                              Hurricanes in New England

                              Landscape Pattern Effects on Disturbance Spread

               Effects of Disturbances on Heterogeneity

                              Stand-Replacing Wildfires in Yellowstone National Park

               Historical Range of Variability

Interactions of Landscape Patterns and Ecological Processes

               Leaf Area and Productivity

               Forest Carbon Dynamics

               Nutrient Dynamics

Suggested Readings

CHAPTER 23 SUSTAINABILITY OF FOREST ECOSYSTEMS

Concepts of Sustainability

               The Prevalence of Human Values in Forest Ecology

               Historical Perspective of Sustainability in Forests

               Ecosystem Services

               Toward a Definition of Sustainability

Where Do We Go from Here?

               Prologue: Earth as a Metaphor for Life

Suggested Readings

LITERATURE CITED

SCIENTIFIC NAMES OF TREES AND SHRUBS

INDEX