Table Of ContentEcological Restoration and
Management of
Longleaf Pine Forests
Ecological Restoration and
Management of
Longleaf Pine Forests
Edited by
L. Katherine Kirkman
Steven B. Jack
CRC Press
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Library of Congress Cataloging-in-Publication Data
Names: Kirkman, L. Katherine, editor. | Jack, Steven B., 1960- editor.
Title: Ecological restoration and management of longleaf pine forests/ editors, L. Katherine
Kirkman and Steven B. Jack.
Description: Boca Raton : CRC Press, 2017. | Includes bibliographical
references and index.
Identifiers: LCCN 2017010051 | ISBN 9781498748186 (hardback : alk. paper)
Subjects: LCSH: Longleaf pine--Southern States. | Longleaf
pine--Conservation--Southern States. | Pine--Southern States. |
Pine--Conservation--Southern States. | Forest ecology--Southern States. |
Forest restoration--Southern States.
Classification: LCC SD397.P59 E36 2017 | DDC 634.9/7510975--dc23
LC record available at https://lccn.loc.gov/2017010051
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Dedication
Photograph courtesy of Richard T. Bryant.
This book is dedicated to Dr. Robert J. “Bob” Mitchell (1955–2013). Bob was
among the initial cohort of scientists recruited to the Joseph W. Jones Ecological
Research Center at Ichauway and was instrumental in helping establish its
multifaceted program focused on the ecology, management, and restoration of the
longleaf pine ecosystem. As a long-time colleague and collaborator to those of us
at Ichauway, Bob’s intellectual “thumbprint” and scientific legacy is pervasive
throughout this volume. He was a creative senior scientist, dedicating his
research program to furthering a scientific understanding of the abiotic and biotic
processes that structure longleaf pine ecosystems and applying that knowledge
to the development of sustainable management and restoration strategies.
vi
Bob thought broadly and deeply about longleaf pine—studying the ecophysiology,
population dynamics, fire ecology, biogeochemistry, roots, mycorrhizal
partnerships, and competitive interactions of the ecosystem—and was a
prominent figure in the development of ecological forestry approaches for natural
resource management. Through his careful reasoning, rigorous investigation,
critical evaluation of ideas, and numerous collaborations with other eminent
scientists, Bob challenged long-held conventional paradigms about the
ecology of longleaf pine forests, and in the process, fundamentally changed the
understanding and management of the ecosystem. Bob’s numerous contributions
to longleaf pine ecology, conservation, and management are widely recognized
by scientific colleagues and by many natural resource practitioners throughout
the Southeast. Those of us who were fortunate to have worked closely with
him recognize the marked influence he had on our investigations, careers, and
the research program at the Joseph W. Jones Ecological Research Center.
Kay Kirkman and Steve Jack
–Editors
Contents
Foreword ...........................................................................................................................................ix
Preface...............................................................................................................................................xi
Acknowledgments ............................................................................................................................xv
Editors ............................................................................................................................................xvii
Contributors ....................................................................................................................................xix
Reviewers ........................................................................................................................................xxi
PART I The Background for Ecological Restoration
Chapter 1 The Fire Forest of the Past and Present ........................................................................3
L. Katherine Kirkman, Steven B. Jack, and R. Kevin McIntyre
Chapter 2 Biogeography: An Interweave of Climate, Fire, and Humans ...................................17
Michael C. Stambaugh, J. Morgan Varner, and Stephen T. Jackson
Chapter 3 The Social and Economic Drivers of the Southeastern Forest Landscape ................39
R. Kevin McIntyre, Barrett B. McCall, and David N. Wear
PART II The Ecological Basis for Restoration
Chapter 4 Regeneration Dynamics, Competition, and Seedling Response ................................71
Steven B. Jack and Stephen D. Pecot
Chapter 5 Mechanistic Controls of Community Assembly and Biodiversity .............................89
L. Katherine Kirkman and Jonathan A. Myers
Chapter 6 The Role of Fuels for Understanding Fire Behavior and Fire Effects .....................107
E. Louise Loudermilk, J. Kevin Hiers, and Joseph J. O’Brien
Chapter 7 Ecosystem Processes and Restoration ......................................................................123
Lindsay R. Boring, Joseph J. Hendricks, R. Scott Taylor, and Daniel Markewitz
Chapter 8 Considering Herbivory and Predation in Forest Management .................................141
L. Mike Conner and Michael J. Cherry
Chapter 9 Geographically Isolated Wetlands: Embedded Habitats ..............................................153
Lora L. Smith, Amanda L. Subalusky, Carla L. Atkinson, and
L. Katherine Kirkman
vii
viii Contents
PART III Ecosystem Restoration: Linking Ecological
Understanding and Management
Chapter 10 Restoring and Managing the Overstory: An Ecological Forestry Approach ...........175
Steven B. Jack and R. Kevin McIntyre
Chapter 11 Restoring and Managing a Diverse Ground Cover ............................................207
L. Katherine Kirkman and Lisa M. Giencke
Chapter 12 Management and Restoration for Wildlife ...............................................................233
Lora L. Smith, James A. Cox, L. Mike Conner, Robert A. McCleery, and
Elizabeth M. Schlimm
PART IV Restoration and Practical Issues
Chapter 13 Air Quality and Human Health Challenges to Prescribed Fire ...............................255
Mark A. Melvin and R. Kevin McIntyre
Chapter 14 Ecosystem Monitoring and Adaptive Management .................................................267
Robert D. Sutter and Brandon T. Rutledge
PART V Longleaf Pine Ecosystem Restoration Perspectives
Chapter 15 Planning for an Uncertain Future: Restoration to Mitigate Water Scarcity and
Sustain Carbon Sequestration ..................................................................................291
Steven T. Brantley, James M. Vose, David N. Wear, and Larry Band
Chapter 16 Longleaf Pine Restoration in Context: Comparisons with
Frequent-Fire Forests ...............................................................................................311
Seth W. Bigelow, Michael C. Stambaugh, Joseph J. O’Brien,
Andrew J. Larson, and Michael A. Battaglia
Chapter 17 Longleaf Pine Ecosystems: The Path Forward ........................................................339
Steven B. Jack and L. Katherine Kirkman
Literature Cited ...........................................................................................................................349
Index ..............................................................................................................................................421
Foreword
The longleaf pine ecosystem is one of the most extraordinary of the major forest ecosystems in
North America, notable for its incredibly rich biological diversity and the extreme role that fire
plays in its maintenance. It is without question the most biologically diverse of any temperate forest
ecosystem in North America. The richness of the biological diversity is evident in both its plant and
animal life. The ground layer of longleaf pine forests typically includes hundreds of vascular plant
species, providing an incredible plant identification challenge to the amateur botanist and even the
professional plant ecologist. Vertebrate life in the longleaf ecosystem is rich with reptiles, amphib-
ians, birds, and mammals, including many species of special interest, such as the gopher tortoise
and red-cockaded woodpecker.
The longleaf pine ecosystem depends fundamentally upon the frequent occurrence of fire for its
sustainability. These frequent fires maintain open, savanna-like forests dominated by longleaf pine
and herbaceous ground cover; as a longleaf forester friend of mine once commented, “Jerry, you
just need to understand that this ecosystem is a tallgrass prairie with some pine trees stuck in it!”
This statement is true to a degree because the longleaf pine ecosystem requires frequent fire, and
eliminating fire or reducing its frequency causes the forest to transition to another state with greatly
reduced biodiversity, similar to the effects of removing fire from prairie grass systems. In actuality,
however, the basic ecological processes and their linkages with structural characteristics are vastly
different in longleaf pine forests and tallgrass prairies. Arguably, the longleaf pine ecosystem repre-
sents an extreme example of a forest’s sensitivity to its disturbance regime; it also exemplifies what
happens when frequent fire interacts with a productive forest ecosystem. Hence, my characteriza-
tion of longleaf pine as the “bookend,” or most extreme example of a frequent-fire forest ecosystem.
Some of the characteristics of longleaf pine and of the ecosystem that it inhabits have been
known for some time, such as the unique grass stage of the young longleaf pine seedlings and the
importance of frequent fire. However, beyond some of the species’ silvics and the richness of plants
and animals, the longleaf pine ecosystem has received relatively little scientific attention, despite
its historic extent in the Southeast. After the virgin stands were logged, longleaf pine was largely
rejected as a commercial tree species by production foresters because the trees grew too slowly,
making it poorly adapted to short-rotation plantation forestry. Instead, southern forestry focused on
the faster-growing and shorter-lived loblolly, slash, and shortleaf pines. With few exceptions, the
only advocates for longleaf pine for many decades were a community of hunters and a few foresters
and wildlife managers who managed some private estates as hunting grounds, particularly for quail.
Interest in the longleaf pine ecosystem exploded in recent decades, however, particularly among
those interested in the maintenance of regional biodiversity, with the red-cockaded woodpecker
as a flagship organism. However, there was very little science to inform efforts at restoration and
management of the ecosystem as a whole; much of the effort has been focused on individual spe-
cies (e.g., quail, or more recently, woodpeckers) or managing gamebird habitat with prescribed fire.
Our general perspectives on forests in North America have been dramatically altered by
scientists since the 1960s, from thinking of them as a collection of trees to understanding them as
rich and complex ecosystems. Much of this research occurred in the hardwood forests of the east-
ern United States, such as at the Coweeta Experimental Forest in North Carolina, Hubbard Brook
Experimental Forest in New Hampshire, and Harvard Forest in Massachusetts. In my home region
of the Pacific Northwest, ecosystem-level science largely began with studies in old-growth Douglas-
fir and western hemlock forests with support from the International Biological Program. Much of
the ecosystem science has been funded by the National Science Foundation. However, during this
period, there was no center of research focused on learning about longleaf pine as an ecosystem and
using it to develop approaches to restoring and managing such ecosystems.
ix