North Central Research Station

Forest Service—U.S. Department of Agriculture

1992 Folwell Avenue

St. Paul, Minnesota 55108

www.ncrs.fs.fed.us
EIGHTH WORKSHOP ON 

SEEDLING PHYSIOLOGY AND GROWTH PROBLEMS IN OAK PLANTINGS

HIAWASSEE, GEORGIA, SEPTEMBER 9-12, 2001

Edited by

S. Sung; P. P. Kormanik, W. J. Ostrosina, and J. G. Isebrands

Sponsored by

USDA Forest Service, Southern Research Station

USDA Forest Service, Region 8

Georgia Forestry Commission

Held at

Lake Chatuga Lodge

Hiawassee, Georgia

Published by

North Central Research Station

USDA Forest Service

St. Paul, Minnesota

Mention of trade names does not constitute endorsement of products by the USDA Forest Service. Each author is responsible for the accuracy

and content of his/her abstract.
Dedication and Preface

Paul P. Kormanik

This General Technical Report (GTR) is dedicated to Dr. Paul P. Kormanik, a principal silviculturist, with the

Institute of Tree Root Biology, USDA Forest Service’s Southern Research Station in Athens, Georgia. We honor Paul for his

untiring research and development work promoting quality oak seedling production in forest nurseries and superior planting

methods in the Southern Region of the Forest Service. His contributions are also valued nationwide. Paul is pictured above

receiving an award from Fred Allen, director of the Georgia Forestry Commission, for his contributions to oak regeneration in

the South. The presentation was made at the opening of the 8th Workshop on Seedling Physiology and Growth Problems in

Oak Planting, held September 9-12, 2001, at the beautiful Lake Chatuga Lodge in the north Georgia Mountains near the

Chattahoochee National Forest. 

Organized by Paul and his colleagues from Athens, the workshop featured presentations on oak genetics and physiology, oak nursery practices, and oak silvicultural practices. Stephen Weaver of the Southern Region gave the keynote address

on the importance of oak in the Region. Clayton Black of the University of Georgia, the featured banquet speaker and Paul’s

long-time biochemist friend, talked about his travels to Mongolia. 

Little did the workshop participants know that during the field trip to nearby oak plantations on Tuesday,

September 11, that their trip would be interrupted by the horrific terrorist attacks that morning in New York City and

Washington, DC. The workshop went on, to the credit of the organizers, but none of us who attended will ever forget the 8th

Oak Workshop.

Abstracts of the papers and posters from the workshop are presented here in alphabetical order of senior authors.

J.G. Isebrands

Rhinelander, WI
ACKNOWLEDGMENTS

The authors wish to acknowledge the contributions of Linda Watson and Catharine Cook, who were in charge of

registration. Thank you to Jim Cunningham, Jeff Magniez, and Michael Thompson who helped arrange the field trips

and also to Jeff for his photography. We also thank the Southern Research Station and Region 8 of the USDA Forest

Service and the Georgia Forestry Commission for sponsoring the workshop.

REVIEW PROCEDURE

Each abstract in this general technical report was critically reviewed by at least two editors. Revised abstracts were

reviewed by each author and then submitted camera-ready to North Central Research Station, USDA Forest Service,

for publication. Individual authors are ultimately responsible for the accuracy of their papers.

DISCLAIMER

Mention of pesticides, fertilizer, equipment or trade names does not imply endorsement of these products or their

applications by the USDA Forest Service.
TABLE OF CONTENTS

Page

A Test of the Relative Importance of Canopy and Root System Gaps in a Bottomland Hardwood Forest 

Dominated by Laurel Oak (Quercus laurifolia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Marianne K. Burke and Charles A. Harrison

A Canker Disease Caused by Phytophthora ramorum Associated with Extensive 

Mortality of Lithocarpus and Quercus in California . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

M. Garbelotto and D. Rizzo

Establishing Bottomland Oak Plantations in the Lower Mississippi River 

Alluvial Valley . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

Emile S. Gardiner

Oak Regeneration in the Mid-Atlantic Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

Peter J. Gould

Guide to Microlepidoptera Feeding on Indigenous Oak of Central and Eastern 

North America North of Mexico . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

John R. Grehan, John A. Stein, Robert E. Acciavatti, Claudia A. Violette, 

Michael Sabourin, William E. Miller, and Denise Binion

Oak Regeneration in the Lake States: An Overview of 20 Years of Research . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

J.G. Isebrands and Ronald M. Teclaw

Grading Oak Seedlings for Assessing Their Competitive Ability Prior to Outplanting . . . . . . . . . . . . . . . . . . . . . . . . 7

Paul P. Kormanik, Shi-Jean S. Sung, and Stanley J. Zarnoch

Georgia Forestry Commission’s Nursery Protocol—Lessons to be Learned from Goldilocks . . . . . . . . . . . . . . . . . . . 9

Taryn Kormanik

Light Acclimation by Laurel Oak Seedlings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

Kenneth W. McLeod and Marianne K. Burke

Insects Threaten Oak Seedling Regeneration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .11

Daniel R. Miller

Minor Pathogens can have Major Impacts on Oaks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .12

William J. Otrosina, Shi-Jean S. Sung, and Paul P. Kormanik

Harvesting Intensity, Site Disturbance, and Understory Control Affect Survival

and Growth of Hardwoods 5 Years After Planting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .13

Felix Ponder, Jr.

Use of Oak in Riparian Forest Buffer Restoration in the Midwestern U.S. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14

Richard C. Schultz, Thomas M. Isenhart, William Simpkins, James Raich, 

Joe P. Colletti, Thomas L. Schultz, Jennifer Nelson, and Emily Stauffer
Growth and Development of First-Year, Nursery-Grown, Open Pollinated Half-Sib

Progeny of White and Southern Red Oak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16

Shi-Jean S. Sung, Paul P. Kormanik, and Stanley J. Zarnoch

Regenerating Northern Red Oak on a Rich Mesic Site in Northern Wisconsin . . . . . . . . . . . . . . . . . . . . . . . . . . . .17

Ronald M. Teclaw

Gentic Resource Management for Upland Oaks for the USDA Forest Service’s, Southern Region . . . . . . . . . . . . . . .18

Tom Tibbs

Response of Four Oak Species to Herbaceous Perennial Legumes in an Upland

Oak Savanna Restoration Project . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .19

J.W. VanSambeek, R.L. McGraw, N.E. Navarrete-Tindall, and P.R. Beuselinck

Bluejack, Blackjack, Southern Red, and Water Oaks After 25 Years of Growth

in Southern Mississippi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20

Charles H. Walkinshaw

The Upland Oak Resource and the Need for an Artificial Regeneration Option

in the USDA Forest Service’s Southern Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21

Stephen P. Weaver

Survival of Underplanted White Oak and Competitors Following a Prescribed Burn . . . . . . . . . . . . . . . . . . . . . . .22

Dale R. Weigel, Paul S. Johnson, and Daniel C. Dey

Abundance and Localization of Aluminum in the Fine Mycorrhizal Roots of Oak (Quercus petraea) . . . . . . . . . . . . .24

Christine Wilson and Joachim Block
A TEST OF THE RELATIVE IMPORTANCE OF CANOPY AND ROOT SYSTEM GAPS 

IN A BOTTOMLAND HARDWOOD FOREST DOMINATED BY 

LAUREL OAK (QUERCUS LAURIFOLIA) 

Marianne K. Burke and Charles A. Harrison

USDA Forest Service, Southern Research Station 

Charleston, SC 29414

When gaps are formed in the canopy of forests, gaps also are created in living root systems, thereby

enhancing the availability of soil moisture and nutrients in addition to light. The ability of advance

regeneration to respond to the more available resources will influence their probability of becoming

canopy trees. In the Lower Coastal Plain of South Carolina, laurel oak (Quercus laurifolia) is a dominant

species in many bottomland hardwood forests, and it is the dominant tree species in both the canopy

and advance regeneration at the Coosawhatchie Bottomland Ecosystem Study site along the

Coosawhatchie River in South Carolina. This is a long-term ecosystem study site managed by the USDA

Forest Service Southern Research Station on land owned by Westvaco Corporation. In 1997, 12 canopy

gaps were identified in 70-year-old bottomland hardwood forest stands. Half of the gaps were in an area

scheduled for regeneration the following year, and half were in an area not to be cut. In 1998, each gap

was enlarged to a 15-m diameter by girdling the large trees and removing the smaller individuals. In

each gap, four 1-m x 1-m plots were selected. Two of these plots were left intact and two were trenched

to a 40-cm depth to kill roots within the plots in order to reduce nutrient and moisture uptake by

canopy trees. A previous study had determined that almost no roots penetrated below 30-cm soil depths

due to the high bulk density of the lower soil. Aluminum flashing was placed in the trenches before they

were backfilled to eliminate regrowth of roots into trenched plots. Identical plots were also located in the

closed canopy forest 50 m from each gap. One and two years after girdling and trenching, laurel oak

seedlings in the gap plots were taller and had larger root crown diameter and greater root and stem biomass than saplings under the closed canopy. Biomass was predicted from allometric equations derived

from laurel oak seedlings collected on the site. Other understory vegetation groups also thrived in the

gap plots, which had greater cover and relative cover of herbaceous and especially grasslike (grass and

sedge) understory vegetation than the non-gap plots. Generally, trenching did not have an effect on

growth of the laurel oak seedlings, but it did increase the cover and relative cover of the other understory vegetation groups. Although laurel oak seedlings responded to the greater light availability, they

were not as responsive as understory vegetation to the enhanced edaphic resources. Based on these

observations, laurel oak seedlings in gaps should have an advantage in becoming canopy trees, but may

initially experience more competition with other understory plant species.
A CANKER DISEASE CAUSED BY PHYTOPHTHORA RAMORUM ASSOCIATED WITH EXTENSIVE

MORTALITY OF LITHOCARPUS AND QUERCUS IN CALIFORNIA

M. Garbelotto 

Department of Environmental Science, Policy, and Management, 

University of California 

Berkeley, CA 94720

and 

D. Rizzo 

Department of Plant Pathology 

University of California 

Davis, CA 95616

This paper describes a new canker disease of Lithocarpus densiflorus, Quercus agrifolia, Q. kellogii, and Q.

parvula var. shrevei caused by Phytophthora ramorum. This pathogenic oomycete has never been isolated

before in North America, and it is known to be present only in a small area of Europe, where it has just

been described. The earliest and most consistent symptom of the disease on larger trees is brown or black

discolored bark on the lower trunk followed by dark red sap seeping from the bark on the trunk.

Phytophthora ramorum was isolated from cankers on mature trees at the root crown to 20 m above the

ground. Although cankers are often initiated at the basal part of the tree, they did not enlarge below the

soil line and were not found extending into the roots. Phloem and xylem cankers were delimited by thin

black lines and ranged to over 2 m in length. The entire crown of affected trees often dies rapidly, and the

foliage turns from an apparently healthy green to brown within a few weeks. The disease is currently

known to be present from southern Oregon to the Big Sur region of California. Besides oaks and tanoaks,

the pathogen also infects leaves and branches of at least six more hosts in four different plant families. The

disease on these hosts appears as a foliar and twig die-back. Although Phytophthora ramorum is not as virulent on these further hosts, it appears that most inoculum infecting the oaks may come from them. The

high susceptibility of several indigenous plant species to this pathogen suggests either a significant and

new ecological predisposition in plants to infection by a native microbe, or the exotic nature of the

pathogen itself.

2
ESTABLISHING BOTTOMLAND OAK PLANTATIONS IN THE LOWER 

MISSISSIPPI RIVER ALLUVIAL VALLEY

Emile S. Gardiner

USDA Forest Service, Southern Research Station

Center For Bottomland Hardwoods Research

Stoneville, MS 38776

In the 1960s, researchers at the USDA Forest Service, Southern Hardwoods Laboratory in Stoneville, MS,

initiated a sustained research program to address artificial regeneration of bottomland oaks (Quercus spp.)

in the Lower Mississippi River Alluvial Valley. Through this research effort, basic techniques for direct seeding and establishing seedlings of bottomland oaks were developed. These techniques were applied to establish the earliest oak plantations on public property for restoration of forest cover and production of hard

mast for wildlife habitat. Concurrent to these early afforestation activities on public holdings, drainage programs and agricultural commodity prices provided incentive for private landowners to emphasize agricultural land uses. High commodity prices, especially for soybeans (Glycine max [L.] Merrill), supported additional deforestation in the region where the original bottomland forest was reduced from an estimated 21

million acres at the time of European settlement to about 5 million acres. Signing of the “Swampbuster”

legislation in 1985 slowed bottomland hardwood deforestation and current governmental incentives are

encouraging land use shifts back to forest cover. More than three decades after the first oak plantations

were established, there is a strong demand for reliable techniques to establish bottomland hardwoods on

economically marginal agricultural land. Afforestation efforts over the last decade resulted in the establishment of more than 300,000 acres of hardwood plantations on public and private land in the Lower

Mississippi River Alluvial Valley. Bottomland oaks are the most common species established in these plantations, making up over 75% of the planting stock. A summary of techniques and associated problems of

establishing bottomland oaks on economically marginal agricultural land in the Lower Mississippi River

Alluvial Valley will be presented at the workshop.

3
OAK REGENERATION IN THE MID-ATLANTIC REGION

Peter J. Gould

Pennsylvania State University

School of Forest Resources

University Park, PA 16802

The objectives of this presentation are to (1) describe recent trends in growing-stock composition in the

region; (2) examine the species composition, abundance, height, and diameter of advanced regeneration;

and (3) describe how oak and non-oak advanced regeneration develop after overstory manipulation.

Permanent plot data collected under a number of treatments will be used to illustrate the development of

advanced regeneration before and after overstory manipulation.

4
GUIDE TO MICROLEPIDOPTERA FEEDING ON INDIGENOUS OAK OF 

CENTRAL AND EASTERN NORTH AMERICA NORTH OF MEXICO

John R. Grehan and Claudia A. Violette

Frost Entomological Museum, Department of Entomology

University of Pennsylvania

University Park, PA 16802

John A. Stein and Denise Binion

USDA Forest Service

Forest Health Technology Enterprise Team

Morgantown, WV 26505

Robert E. Acciavatti

USDA Forest Service

Forest Health Protection

Morgantown, WV 26505

Michael Sabourin

Grantsburg, WI 54840

and

William E. Miller

Department of Entomology

University of Minnesota

St. Paul, MN 55108

A guide to oak-feeding Microlepidoptera is currently being developed to focus attention on the smaller

moth species that belong to the community of oak-feeding insects. These moths represent a collection of

various families comprising many species that have small bodies and often cryptic larval stages that either

burrow within the host or live within shelter constructed by the larva. Microlepidopterans are often less

visible than the more well known Macrolepidoptera that have relatively large external leaf-feeding larvae.

The guide will present an overview of the biodiversity and taxonomic composition of the microlepidopteran fauna known to inhabit the indigenous oak species of eastern North America north of Mexico.

The species are primarily drawn from the literature, and some additional records are drawn from museum collections. The guide will provide foresters and forest entomologists with an outline of current information on the diversity, identification, and biology of the oak-feeding Microlepidoptera community. We

anticipate future studies will continue to develop a more detailed and precise characterization of the oak

fauna with respect to host-plant preferences, species abundance and persistence, and the challenge of

accurate identification. 

The guide will include two main sections. 

1) The microlepidopteran species with notes on the morphology of the adult and larva (the latter

where known), the known hosts, larval biology, and species distribution. Adults and larvae will be

illustrated in color plates where possible, and the distribution range will be mapped. A general summary of host-plant relationships will be presented. 

2) An introductory guide to the identification of 50 indigenous oak species in eastern North America

with illustrations and notes on habitat, growth form, bark, twigs, buds, leaves, and acorns. Also

included with species description is the most current distribution information mapped to the county

level. An electronic version of the oak key with species description will be published on the Forest

Health Technology Enterprise Team Web site within the next year.

5
OAK REGENERATION IN THE LAKE STATES: AN OVERVIEW

OF 20 YEARS OF RESEARCH 

J.G. Isebrands and Ronald M. Teclaw

USDA Forest Service, North Central Research Station

Rhinelander, WI 54501

In the early 1980s we established an interdisciplinary research program to address the lack of oak regeneration in Lake States forests. The program included fundamental and applied studies conducted in the laboratory, growth chamber, greenhouse, nursery, and field. Ten field studies were established on county,

State, and Federal lands in cooperation with forest managers. These studies included overstory manipulation studies across a gradient of habitat types, underplanting studies in hardwood and pine stands, and

exclosure studies. The funding for the program ended in 1991, but the field studies remain as a living

legacy of the program. In addition, more than 82 publications resulted from the program. In this contribution we give an overview of the most significant applied results and lessons learned from the 20 years

of research.

Our most significant findings were in eight categories. 

1) Herbivory—herbivore browsing is the most important problem affecting oak regeneration in the Lake

States. Tree shelters, fencing, use of large nursery stock, and repellents provide some protection. 

2) Frost—frost in the establishment phase and during the early growing season seriously affects oak

regeneration. Silvicultural treatments that provide canopy cover ameliorate the frost problem. 

3) Competition Control—competing vegetation control with mechanical or chemical methods significantly

improves oak artificial regeneration success. 

4) Nursery Stock Quality—improvements in nursery stock quality are needed in Lake States. Improved

seed sources, seed handling, and nursery cultural practices result in larger caliper, higher quality oak

nursery stock. Containerized stock also shows promise. 

5) Oak Hybridization—northern red oak, black oak, and northern pin oak routinely hybridize naturally

throughout the Lake States. Hybrid oak seeds do not perform as well in the nursery and often

exhibit poorer form after establishment. 

6) Light Intensity—once established, oak seedlings perform better at higher light intensities provided by

silvicultural treatment or natural disturbance gaps.

7) Tree Shelters—large tree shelters under a 50% crown cover show promise for providing herbivore protection and for improving early growth during establishment. However, results are variable and

material and maintenance costs are high. 

8) Oak Under Pine—we have had much success in establishing oak seedlings naturally and artificially

under pine by emulating the natural ecological life history relationship of oak/pine in the Lake

States. Pine overstories provide favorable overstory and understory conditions for natural as well as

artificial oak regeneration.

Our findings suggest that to successfully regenerate oak in the Lake States more intensive management is

needed than is currently employed—including time, effort, and money.

6
GRADING OAK SEEDLINGS FOR ASSESSING THEIR COMPETITIVE 

ABILITY PRIOR TO OUTPLANTING 

Paul P. Kormanik and Sung, Shi-Jean S.Sung 

Institute of Tree Root Biology, USDA Forest Service, 

Southern Research Station

Athens, GA 30602

and

Zarnoch, Stanley J.

USDA Forest Service, Southern Research Station

Asheville, NC 28802

It is well known that artificial regeneration of many oak species has been less than satisfactory and there

are no standard criteria by which seedlings are either grown or evaluated prior to outplanting. In the

past, most nursery produced oak seedlings were of inferior planting stock as 1-0 material and various

methods of using 1-1, 2-0, and even 2-1 were evaluated to improve planting stock quality. These

attempts of extending the nursery production period were expensive and produced seedlings deemed too

large for outplanting. Various combinations of root and top prunings were undertaken to reduce shipping and planting costs as well as to attempt to improve survival and growth after outplanting.

Characteristically, while these clipping treatments were sometimes effective in improving survival, growth

was not improved. Thus, it took 3-5 years for the seedlings to obtain the heights that existed prior to

clipping. During this period, the seedlings became overtopped and were no longer competitive. Few

successful plantings have been reported. It was generally conceded that large diameter seedlings were

most competitive and that seedlings would become more competitive in the field as they became a meter

or more in height. Unfortunately, most did not.

At the USDA Forest Service Institute of Tree Root Biology (ITRB) in Athens, Georgia, we developed a

nursery protocol (described elsewhere in these proceedings) that has proven to be effective in producing

high quality planting stock of most of the oaks native to the Eastern and Southeastern United States. We

developed a seedling evaluation procedure that is very effective in identifying the future competitive

seedlings based upon their nursery development as 1-0 stock. This protocol has the additional benefit of

being a first step in screening that can identify poor families as well as poor performers within a family, as

illustrated in table 1 for northern red oak (Quercus rubra) and table 2 for white oak (Q. alba). In addition, the protocol has produced comparable data in tests on five other southern oak species. Table 1 was

taken from a particular year’s experiment where 24 or more open pollinated half-sib progenies were evaluated before outplanting. 

What our data clearly indicate is that if a seedling has more first-order-lateral-roots (FOLR) than the mean

number for that family, when produced by our nursery management protocol, its root collar diameter

(RCD) and height (HGT) will both be greater than their respective family means. These individuals

would then meet our standards for outplanting stock. We use minimum FOLR number, RCD, and HGT

of 4, 8.0 mm and 90 cm, respectively, as our criteria for northern red oak planting stock. In this example, mother tree 2-23-850 would probably be excluded from further testing since in reality only about

20% of its progeny exhibited desirable nursery competitiveness (table 1).

7
Table 1.—Morphological and growth characteristics of 1-0 Quercus rubra seedlings

% of

FOLR# Seedlings with RCD, mm HGT, cm

Family x Range FOLR < x x Range x Range

2-23-850 2.7 0-25 72 6.2 1.0-16.9 73 8-207

4-2-902 3.6 0-15 62 8.4 3.6-15.0 124 46-223

4-27-100 4.2 0-19 56 9.3 2.9-17.9 141 33-246

4-14-2459 5.9 0-20 52 10.5 3.4-18.9 125 22-253

Table 2 presents white oak data from the 2000 growing season. The mother trees have been followed for over

25 years at the Georgia Forestry Commission Arrowhead Seed Orchard. Records have been maintained and

individuals that have not consistently produced acorns have been gradually eliminated. Progenies from the

remaining individuals are consistently the most competitive seedlings in the nursery and have been developing

very well in our outplanting trials. The performance of the progeny from the Arrowhead Seed Orchard, when

compared to that of the wild collection, indicates significant potential for improving seedling quality through

judicial selections in seed orchards. For example, most half-sib progeny groups (35 in this particular trial)

had at least 45% plantable seedlings (table 2). In our normal wild collections, we characteristically obtain 25-

30%.

Table 2.—Morphological and growth characteristics of 1-0 Quercus alba seedlings

% of

FOLR# Seedlings with RCD, mm HGT, cm

Family x Range FOLR < x x Range x Range

ASO-5 6.7 0-28 53 9.3 2.7-18.6 86 14-198

ASO-16 5.8 0-27 56 10.3 2.0-19.0 89 12-180

ASO-18 4.8 0-28 56 9.8 2.3-19.8 98 22-183

ASO-42 5.2 0-42 55 9.2 2.1-22.4 90 8-188

8
GEORGIA FORESTRY COMMISSION’S NURSERY PROTOCOL – 

LESSONS TO BE LEARNED FROM GOLDILOCKS

Taryn Kormanik

Department of Agricultural and Environmental Science

University of Georgia

Athens, GA 30602

Nursery management raises many questions that scientists, both basic and applied, have yet to answer.

High quality seedling production, with minimal or no deleterious impacts on the local environment, would

be much easier if nursery managers knew everything about the soil and plant processes that constitute a

crop of seedlings. The Institute of Tree Root Biology (ITRB) has only scratched the surface of these questions. However, even in the absence of complete knowledge, the Institute has developed nursery management guidelines that consistently yield high quality seedlings for reforestation. The guidelines wield a

multi-pronged approach, covering everything from procurement of suitable seed stock to outplanting

issues. One major emphasis is the manipulation of soil fertility levels. Through several years of testing and

fine-tuning, the following levels of extractable micro and macro soil nutrients have proven optimal for

seedling production in the Flint and Walker GFC nurseries:

Phosphorous 100 ppm

Potassium 130 ppm

Calcium 400 – 600 ppm

Magnesium 75+ ppm

Copper 0.3 – 3 ppm

Zinc 5 – 15 ppm

Boron 0.4 – 1.2 ppm

Various aspects of these guidelines will be discussed.

9
LIGHT ACCLIMATION BY LAUREL OAK SEEDLINGS

Kenneth W. McLeod

Savannah River Ecology Laboratory

University of Georgia

Aiken, SC 29802

and 

Marianne K. Burke

USDA Forest Service, Southern Research Station

Center for Forested Wetland Research, 

Charleston, SC 29414

Forest gap dynamics predicts that species that can most fully use the new light environment of a gap will

become the replacement individuals. When a gap is created, as when a tree falls, all seedlings in the existing regeneration layer theoretically have an equal opportunity to become the next canopy individual in the

new gap environment. But the loss of a single canopy individual frequently begins a process whereby the

gap slowly enlarges as the opening affects the canopy trees bordering the gap. When this happens, the

light and rhizosphere environment slowly changes. If a species can acclimate to the new environment,

then it will have a competitive advantage. 

Along the Coosawhatchie River in South Carolina, 12 canopy gaps were identified in each of two sections

of a bottomland hardwood forest, dominated in the canopy and advanced regeneration layer by laurel oak

(Quercus laurifolia). Each gap was enlarged to a 15-m diameter by girdling large trees and removing the

smaller ones. In each gap, four 1-m x 1-m plots were established. Two plots were trenched to kill roots

within the plots to reduce nutrient and water uptake by canopy trees, and two were left intact. Aluminum

flashing was placed in the trench to inhibit root regrowth before the trench was backfilled. Another set of

four plots was located in the adjacent closed canopy forest near the enlarged canopy gap and treated like

these four above. To determine the response of laurel oak seedlings, photosynthetic light response curves

(from 75 to 1,200 μmole/m2/s) were determined during the first two growing seasons after gap enlargement. Photosynthetic rates were significantly greater in the gaps than under closed canopy for all light levels greater than or equal to 150 μmole/m2/s. Seedlings exposed previously to the higher light levels in the

gaps were better able to use higher light levels. However, maximum photosynthetic rates of these

seedlings were still only 50% of those observed in seedlings growing in full sunlight. Since the photosynthetic rate of gap and non-gap seedlings differed even when stomatal conductance was similar, seedlings in

gaps or under closed canopy must differ in leaf structure. Trenching did not affect seedling photosynthesis. Based on these observations, laurel oak seedlings previously exposed to higher light levels in gaps

should have an initial advantage of greater potential photosynthetic rates over seedlings that existed under

full canopy.

10
INSECTS THREATEN OAK SEEDLING REGENERATION

Daniel R. Miller

USDA Forest Service, Southern Research Station

Athens, GA 30602-2044

Paul Kormanik asked me to speak to you today and leave you with one message: forest managers dealing

with oak regeneration need to “fear the weevils” because weevils feed on acorns and have no controls.

Acorn insects, my focus in this paper, are one of four groups of insects that impact mast production and

regeneration. Defoliating insects are by far the most recognized and diverse of these groups, causing highly visible damage at various times of the year. Major control efforts are geared to defoliators such as gypsy

moths throughout the range of oaks. Impacts of defoliating insects are generally associated with several

years of severe defoliation and include growth losses, branch dieback, and reduction in mast production.

Under stress, extensive tree mortality can also occur, often due to activities of a third group of insects:

woodborers. Woodborers are a diverse group and generally attack dead or dying trees, leading to direct

loss of mast production. Regeneration pests are the last group of insects that impact seedlings directly by

causing growth loss or mortality.

Acorn insects can be divided into primary or secondary insects. The primary insects, such as Curculio

weevils, filbertworms, and gall wasps, damage the acorn directly to introduce a larva. Secondary insects,

such as acorn moths, midges, and Conotrachelus weevils, take advantage of existing holes or damage.

There are 23 species of Curculio weevils that feed on oak in North America and 11 of these species are in

the East. Any forest stand or seed orchard generally has three to four species at the same time. Insect

damage to acorns can result in average losses of 50% of mast crops with the Curculio weevil accounting

for most of the insect damage. Adult weevils generally emerge in August/September, mate, and climb trees

to lay eggs inside acorns. Larvae feed inside the acorn, often with numerous other larvae inside the same

acorn, until the acorn falls and the larvae leave the acorns and burrow into the ground to depths

approaching 12 inches. Larvae remain underground for about 2 years and then pupate and reemerge as

adults.

An effective integrated control program is required to address three major concerns: (1) the lack of acorns

of specific families; (2) cost of handling acorns to remove damaged ones; and (3) germination failures in

nursery beds. Effective physical control tactics for weevils will likely involve sanitation procedures. The

removal of all acorns in the fall will reduce population levels and impacts in 2 years. Removal of unwanted trees in seed orchards will also reduce population levels. Harvesting directed to trees with large crops

late in the season may yield fewer damaged acorns in harvests. Prescribed fire is not likely to be effective

against acorn weevils since the larvae are protected underground. Biological control tactics are not promising at this time but in the future may include host resistance, parasites, predators, diseases, and

pheromones. No chemical controls are registered for oak acorn weevils. Various pyrethroids, organophosphates, and carbamates are registered for pecan and chestnut weevils, cone and seed pests in pine seed

orchards, defoliators, and ornamental pests. Work is required to expand labels to include oak acorn

insects as well as to determine effective spray programs. Timing and frequency of sprays, monitoring tools

and models, spray coverage, and pre-harvest restrictions need to be addressed. Use of systemic insecticides is possible, but further research is needed on insecticides and doses, method and timing of application, entry paths, and effects on germination and tree health.

11
MINOR PATHOGENS CAN HAVE MAJOR IMPACTS ON OAKS

William J. Otrosina, Shi-Jean S. Sung, and Paul P. Kormanik

USDA Forest Service, Southern Research Station

Institute of Tree Root Biology

Athens, GA 30602

Numerous fungal pathogens have been described that attack various oak species. The majority of these

fungi are regarded by pathologists as minor curiosities that are either rare or have very little impact on

oak survival and growth. Two examples of such diseases are Botryosphaeria canker and powdery mildew.

Again, these diseases are not regarded as serious threats to oak regeneration; however, we had a catastrophic loss due to Botryosphaeria canker in a northern red oak experimental field study. The study

contained over 12 genetically unrelated families totaling over 1,400 trees. By age three, occasional

symptoms such as branch flagging and limited dieback began to appear. Within 3 years after the initial

appearance of symptoms, all the trees in these plantings had died. Why would an essentially secondary

pathogen, normally limited to attacking weakened trees, attack vigorously growing trees planted on a

high quality site? We are uncertain of the mechanisms involved in this disease outbreak, but we theorize

a large amount of available inoculum, coupled with high tree growth rates, may have played a role. The

rapid growth rates may produce large amounts of more susceptible young tissue that can facilitate disease spread.

Other pathogens, such as powdery mildew of white oaks, do not kill their host but affect photosynthetic

rates. They affect foliage by invading and impairing cells that are involved in photosynthesis, causing

affected foliage to be distorted and smaller in size. We conducted a study in which we measured photosynthetic activity of white oak foliage in relation to amount of powdery mildew infection. Our data indicate photosynthetic activity is reduced significantly as powdery mildew infection increases (r2 = 0.89).

While not directly causing mortality, powdery mildew infection in young trees outplanted in the field

can adversely impact growth and, thus, competitiveness. Diseases such as these underscore the need for

awareness of potential disease threats affecting oak regeneration. Research that addresses the impact of

this and other diseases in the nursery and field will provide critical information necessary to ensure successful artificial regeneration of oaks.

12
HARVESTING INTENSITY, SITE DISTURBANCE, AND UNDERSTORY 

CONTROL AFFECT SURVIVAL AND GROWTH OF 

HARDWOODS 5 YEARS AFTER PLANTING

Felix Ponder, Jr.

USDA Forest Service, North Central Research Station 

Jefferson City, MO 65102.

Five years after planting, survival and growth of northern red oak (Quercus rubra L.) and white oak (Q.

alba L.) were examined for differences between treatment combinations that included three levels each of

harvesting intensity (organic matter removal) and site disturbance (soil compaction), and two levels of

understory vegetation control (weed control). This research is part of the USDA Forest Service initiative

to study the long-term soil productivity (LTSP) of forested lands in the United States. The study consists

of three levels of organic matter removal and soil compaction as main effect treatments in a factorial

design and weed control as a secondary treatment applied in a split arrangement. The levels of organic

matter removal included bole only (BO), whole tree (WT), and whole tree plus forest floor (WT+FF),

and the levels of compaction included none, moderate, and severe. The understory was present (U+) or

absent (U-). Organic matter removal treatments had little effect on survival for either tree species. But

both soil compaction and understory control did. Percent survival was better for the moderate and

severe treatments than for the uncompacted treatment for both species, but differences were significant

(α=0.05) only for northern red oak. Survival was significantly higher for the U- treatment than for the

U+ treatment for both tree species. Organic matter removal significantly affected 5-year height and diameter of both oak species. Both height and diameter were greater for BO and WT than for WT+FF. Soil

compaction treatments affected the height of northern red oak and the height and diameter of white oak.

Also, both northern red oak and white oak were significantly taller and larger in diameter in the U- treatment than in the U+ treatment. Some two-way interactions and the three-way interaction were significant

for some measurements for both species. These results indicate that the effects of soil compaction, organic matter removal, and understory control on survival and growth were apparent at age 5. But perhaps

because of weeds’ depressing effect on soil moisture and nutrient availability, understory vegetation survival and growth were affected more by differences in levels of understory control than by organic matter

removal or soil compaction on this site.

13
USE OF OAK IN RIPARIAN FOREST BUFFER RESTORATION 

IN THE MIDWESTERN U.S.

Richard C. Schultz, Thomas M. Isenhart, William Simpkins, James Raich, Joe P. Colletti, 

Thomas L. Schultz, Jennifer Nelson and Emily Stauffer

Agroecology Issue Team, Leopold Center for Sustainable Agriculture 

Iowa State University

Ames, IA 50011

Intensive row crop agriculture and grazing have modified the hydrology of much of the Midwestern U.S.

Corn Belt. Most of the natural prairie, wetland, and forest ecosystems have been replaced by annual row

crops and introduced cool-season grass pastures. Soil organic matter has been reduced and surface soil

has been compacted, reducing infiltration and increasing surface runoff. As a result, streams have had to

carry higher storm flows and lower base flows. Many meandering stream channels have been straightened resulting in faster flows and more incision. The incision has produced vertical stream banks in

danger of collapse. Water tables have been lowered, producing aerated rooting zones that are deeper

than they were 150 years ago. Because the riparian zone (flood plain) often contains some of the most

fertile soil in the landscape, farmers also have removed most of this native plant community and replaced

it with annual crop plants or cool-season grasses. As a result of these changes, sediment, fertilizers, and

other agricultural chemicals have easy access to stream channels, producing sediment and chemical laden

streams that are responsible for major siltation of streams and reservoirs and the hypoxia problems of the

Gulf of Mexico.

The USDA Conservation Reserve Program under the 1996 Farm Bill created an opportunity for farmers

to improve stream water quality and to provide wildlife and aquatic habitat and potential alternative

products by restoring perennial riparian plant communities. The program has two major options: 1)

grass filters and 2) riparian forest buffers. The latter consist of one or more woody plant zones adjacent

to the stream and a native or cool-season grass filter strip between the woody zones and the crop field.

The woody zone(s) must make up at least one-third of the width of the buffer where restoration takes

place on previous crop ground and all but 7 m of the width on restored pastures. 

Members of the Agroecology Issue Team of the Leopold Center for Sustainable Agriculture have played a

key role in developing riparian buffer designs for intensive agricultural landscapes. Their basic model

was one of several used to develop the Riparian Forest Buffer Conservation Standard of the USDA

Natural Resources Conservation Service. Their model consists of a wide variety of trees and shrubs

planted parallel to the channel. They also use a mixed native grass and forb community as the grass filter between the tree/shrub zone and the crop field. For the most part, fast growing bottomland tree

species are planted right along the banks of the channel. However, because of the modified hydrology of

many of the streams in the region, the rooting zone often is deep enough to support high quality upland

species such as oaks (Quercus spp.) and black walnut (Juglans nigra). Red, white, bur, and swamp white

oak are used extensively in these plantings. These trees can provide high quality timber as well as hard

mast for species such as turkey that use the restored corridors to move from one island habitat to the

next.

Oaks are planted in mixtures with other species, but they are planted on micro-sites that are suitable for

them. Since the goal of buffers is to provide a plant community with a rough surface throughout, coolseason grasses such as timothy and rye are planted with the trees on restored crop field sites. If the site

was a pasture, glyphosate kill-strips are used for planting sites. While these understory plants provide

some competition, they are necessary to provide the frictional surface necessary to slow surface runoff

and trap sediment and associated chemicals. As a result, growth rates of the oaks and other trees may

not be as rapid as expected in timber plantations. 

14
The oak and other upland species help diversify the riparian buffer plantings in a landscape that is often

devoid of diversity. These narrow restored riparian ecosystems can trap over 90% of the surface sediment

and 80% of the chemicals associated with that sediment before they get to the stream. Infiltration rates in

these undisturbed soils are five times greater than in adjacent crop fields. They also can remove as much

as 90% of the nitrate in the shallow groundwater. These buffer plantings shade the channel, cooling the

water and elevating oxygen levels. In addition, they provide habitat for eight times more bird species

than cool-season grass filters as well as deer, beaver, and other large mammals. After seeing the potential

of these buffers, most farmers become ardent supporters even though they have lost some productive crop

ground. Riparian forest buffers with oak and other species work. For more information, visit our riparian

buffer homepage at: http://www.buffer.forestry.iastate.edu/

15
GROWTH AND DEVELOPMENT OF FIRST-YEAR, NURSERY-GROWN, 

OPEN POLLINATED HALF-SIB PROGENY OF WHITE AND 

SOUTHERN RED OAK

Shi-Jean S. Sung and Paul P. Kormanik

USDA Forest Service, Southern Research Station

Institute of Tree Root Biology

Athens, GA 30602 

and

Stanley J. Zarnoch

USDA Forest Service, Southern Research Station

Asheville, NC 28802

White oak (Quercus alba L., WO) acorns from individual mother trees from the Arrowhead Seed Orchard

(ASO, Milledgeville, GA), Beech Creek Seed Orchard (BSO, Murphy, NC), and Savannah River Station

(SRS, New Ellenton, SC) were sown in December 1999 at the Whitehall Experiment Forest Nursery

(Athens, GA). All six mother trees from BSO were grafted. By early April, germination exceeded 80% for

all but six families. Five of these six families were from BSO. Seedlings that emerged after mid-April generally were much smaller than those that emerged earlier. Ten percent of the progeny of one SRS family

were albino, i.e., their leaves contained very low levels of chlorophyll. Furthermore, the rest of the surviving seedlings from this family were much smaller than seedlings from other families. Buds for the first

flushes started swelling near the end of April for most WO seedlings. Time from current flush bud

swelling to next flush bud swelling in most seedlings was approximately 33 days for all flushes.

Regardless of seed sources, elongation of the third, fourth, and fifth flushes occurred mainly between 4

and 12 days post bud break (dpbb), and the most active elongation occurred approximately 10 dpbb.

About 89, 55, and 9% of ASO and SRS seedlings had three, four, and five flushes, respectively. Only 60,

15, and 2% of BSO seedlings had three, four, and five flushes, respectively. More than 60% of WO

seedlings from each seed source group had fewer than the mean number of first-order lateral roots

(FOLR). White oak seedlings with first flush length shorter than 5 cm generally had lower values in

growth parameters including height, root collar diameter, flush number, and number. Based on germination time and percent and first flush length, it may be possible to assess progeny quality of given WO

mother trees as early as mid-May. Except for root collar diameter, progeny from grafted WO mother trees

performed poorly in several growth parameters in nursery as compared to progeny from other groups. 

In mid-March 2001, southern red oak (Q. falcata Michaux, SRO) acorns from 12 mother trees from SRS

were sown at the Whitehall Experiment Forest Nursery. By mid-May, germination for 10 SRO families

ranged from 62 to 90%. The other two families had 52 and 14% germination. Similar to late germinated

WO, the SRO seedlings that emerged after mid-May remained small. First flush bud of SRO seedlings

began swelling 3 weeks later than WO first flush buds. However, similar to WO, time from current flush

bud swelling to next flush bud swelling in most SRO seedlings was approximately 31 days. Based on data

collected so far, elongation patterns of the second, third, and fourth flushes in SRO seedlings were similar

to each other and to WO flushes. About 98, 85, and 15% of randomly selected SRO seedlings had three,

four, and five flushes, respectively. Final assessment of progeny quality for individual SRO mother trees

will be made at lifting. Feasibility of using germination percent and time and first flush length as early

(e.g., early June) assessment criteria for SRO progeny quality will be determined at lifting when other

growth parameter data are available.

16
REGENERATING NORTHERN RED OAK ON A RICH MESIC SITE IN NORTHERN WISCONSIN

Ronald M. Teclaw

USDA Forest Service, North Central Research Station

Rhinelander, WI 54501

A regeneration study was established in 1989 to evaluate the effects of overstory reduction and understory

vegetation control on the establishment and growth of northern red oak. The study is located on the Park

Falls District of the Chequamegon-Nicolet National Forest in northern Wisconsin, (45oN, 90oW) and is

referred to as the Willow Springs Ecosystem Processes Study Site. The site is located within a northern

hardwood stand with a northern red oak site index of 65 feet (base 50 years). The habitat classification is

Acer/Viloa-Osmorhiza (AViO), a nutrient rich mesic site. Bareroot (1-0) and containerized northern red

oak seedlings were planted in a clearcut, in two shelterwood stands (one thinned to 50% crown cover and

one to 75% crown cover), and under a fully closed canopy that served as a control. Understory vegetation was controlled in the clearcut and two shelterwood stands according to a factorial design of spraying

RoundupTM herbicide (glyphosate) and disking in 180-foot-wide strips. The bareroot seedlings were rootgraded by the number of first-order lateral roots greater than 1 mm in diameter at the point of attachment

to the tap root. Four-foot-tall tree shelters of two diameters (4 inch and 8 inch) were installed on a subset

of both the bareroot and containerized seedlings.

Survival and growth trends started to develop as early as 2 years after planting; the best survival and

growth were in the 50% crown cover shelterwood stand followed by the 75% crown cover shelterwood

and the clearcut. The greatest mortality and poorest growth were under the closed canopy stand (100%

crown cover). These trends continued to where in 2000 there were significant differences in seedling

height relative to overstory cover; the largest seedlings were under the 50% canopy (avg. ht.=2.5 m), followed by seedlings under the 75% canopy (avg. ht.=1.8 m). Clearcut data have yet be to summarized.

Furthermore, there was significant difference in seedling height relative to understory vegetation control

under the 50% shelterwood canopy. Seedlings were largest in the spray and disk plots (avg. ht.=3.9 m)

and smallest in the control plots (avg. ht.=1.4 m). Seedling height under the 75% shelterwood canopy

was relatively unaffected by understory vegetation control. During the study, the seedlings planted under

the fully closed canopy steadily declined until they were essentially nonexistent at the time of the last survey.

A complete analysis of seedling growth relative to overstory density, understory vegetation control, root

grade, and tree shelter size will be presented at the workshop meeting.

17
GENETIC RESOURCE MANAGEMENT FOR UPLAND OAKS

FOR THE USDA FOREST SERVICE’S, SOUTHERN REGION

Tom Tibbs 

USDA Forest Service, Southern Region

Atlanta, GA 30367

The USDA Forest Service has implemented a management program for two species of upland oaks, northern red oak and white oak. The objectives of this program are genetic conservation and development of

sources of acorns for regeneration, gene conservation, and research. The program is based on genetic

selection of families and seedlings in the nursery and on the establishment of seedling seed orchards and

demonstration areas. 

18
RESPONSE OF FOUR OAK SPECIES TO HERBACEOUS PERENNIAL LEGUMES IN AN UPLAND

OAK SAVANNA RESTORATION PROJECT

J. W. Van Sambeek

USDA Forest Service, North Central Research Station

Columbia, MO 65211-7260

R. L. McGraw and N. E. Navarrete-Tindall

Department of Agronomy

University of Missouri

Columbia, MO 65211-6130

and 

P. R. Beuselinck

USDA Agriculture Research Service

Columbia, MO 65211-6130

One of the new practices included in the Conservation Reserve Program is oak savanna restoration with

widely spaced oak trees established with a ground cover of perennial legumes and warm-season grasses.

Growth of hardwood seedlings is reduced by competing vegetation. Forage legumes have been shown to

improve growth of black walnut seedlings and saplings by reducing grass competition and adding nitrogen to the soil. Less is known about the response of oaks to legumes or the potential benefits of using

native legumes as a ground cover in hardwood plantings.

In spring 1999, an upland 1.5-ha area was plowed, cultivated, and seeded using a randomized block

design with 3 replications of 12 treatments with and without warm-season grasses in combination with 6

legume treatments. Legume treatments included seeding with Illinois bundle flower (Desmanthus illinoensis (Michaux) MacMillian), panicled tick trefoil (Desmodium paniculatum (L.) DC), slender bush clover

(Lespedeza virginica (L.) Britton), roundhead bush clover (Lespedeza capitata Michaux), a mixture of all

four native Missouri legumes, or natural succession of resident vegetation. In spring 2000, six rows of

trees were planted approximately 13 m apart across each ground cover plot to create six four-tree plots

each planted with one seedling each of bur oak (Quercus macrocarpa Michx.), white oak (Q. alba L.),

northern red oak (Q. rubra L.), and Shumard oak (Q. shumardii Buckl.) or black walnut (Juglans nigra L.).

Extensive rodent and deer damage precluded meaningful measurements of stem height or basal diameter;

however, we were able to evaluate survival and seedling vigor.

Overall first-year seedling survival was 85% for bur oak, 74% for black walnut, 70% for white oak, 55%

for northern red oak, and less than 10% for severely dehydrated planting stock of Shumard oak. Bur and

northern red oak had higher percentages of seedlings girdled by mice than white oak and black walnut

(27% vs. 18%). Although no differences were found among species or legume treatments, deer browse

was higher in plots without warm-season grasses than with warm-season grasses (68% vs. 58%). On a

scale of 0 (dead) to 5 (multiple flushes with large healthy leaves), bur oak had the most vigorous

seedlings (3.3) followed by black walnut (2.9), white oak (2.2), and northern red oak (1.9). Legume

treatments with panicled tick trefoil reduced tree seedling vigor compared to a ground cover of resident

vegetation. After two growing seasons, only panicled tick trefoil had produced dense stands that excluded most other vegetation.

19
BLUEJACK, BLACKJACK, SOUTHERN RED, AND WATER OAKS AFTER 25 YEARS 

OF GROWTH IN SOUTHERN MISSISSIPPI

Charles H. Walkinshaw

USDA Forest Service, Southern Region

Tree Root Biology Team

Athens, GA 30602

In the early 1970s the need for ground-truth measurements using remote sensing tools was defined by

W.G. Cibula and others at the National Space Technology Laboratories. I responded to this by planting six

oak species on a site approximately 20 miles north of Gulfport, MS. These trees are now 25 years old and

satisfactory for aerial photography, video and satellite scanning. This report explores the genetic variation

encountered in the water oak population at the site.

Acorns were collected in 1974 at the Biloxi and Red Creek wildlife management areas in southern

Mississippi. Six species were included: blackjack oak (Quercus marilandica), bluejack oak (Q. incana), post

oak (Q. stellata), southern red oak (Q. falcata), turkey oak (Q. laevis), and water oak (Q. nigra). Caps were

removed and the acorns were heated to 45-50oC for 45 minutes, dried, and stored at 3-5oC for 6 months.

The acorns were germinated in sand:vermiculite (1:1 v/v), planted in quart milk cartons of sandy soil, and

transplanted in the field (cleared tree site). A thinning was made after 5 and 10 growing seasons.

More than 90% of the turkey oak and post oak seedlings died 5 years after planting. Bluejack oak had

high mortality after 10 years, but the remaining living trees averaged 4.2 inches in d.b.h. Blackjack, southern red, and water oaks had d.b.h. values from 1.0 to 7.2 inches after 25 years. Mean diameters were 3.6

± 0.24, 3.6 ± 0.29, and 4.1± 0.31 inches, respectively. The largest trees for the three species had diameters

of 5.7, 7.2, and 6.8 inches. When all d.b.h. values for the three oak species were plotted as a histogram,

they followed a normal distribution pattern. Bluejack oak appeared superior after planting, and it produced many acorns. By 25 years, it showed a mortality of approximately 60%. The diameters of bluejack

oak, ranging from 1.4 to 7.6 inches, also followed a normal distribution curve. Differences among the

diameters of bluejack, southern red, water, and blackjack oaks were not significant.

A parallel group of water oak seedlings were inoculated with spores from single fusiform rust galls in six

experiments. As a result, 28.3 to 69.6% of the seedlings had dense telia on their lower surface. Many oak

families failed to form telia with a number of field isolates of the fusiform fungus. However, susceptibility

of oak families did not parallel that of slash pine families.

Results of this study indicate that growing blackjack, southern red, and water oaks in southern Mississippi

is easy. The dense vegetation in the plots did not deter these oaks. I was surprised that some of the water

oaks inoculated with fusiform rust were totally resistant. Thus the role of oaks in natural infections may be

significant in creating new races of the fungus.

20
THE UPLAND OAK RESOURCE AND THE NEED FOR AN 

ARTIFICIAL REGENERATION OPTION IN THE 

USDA FOREST SERVICE’S SOUTHERN REGION

Stephen P. Weaver

USDA Forest Service, Southern Region

Forest Management Unit

Atlanta, GA 30309

This paper examines the upland oak resource managed by the USDA Forest Service in the Southeastern

U.S. Current management plans identify an increased need for additional acres in oak type at the same

time that the oak type is decreasing and being converted to other forest types due to oak regeneration failures and major catastrophic events. The factors influencing the change of oak types are discussed.

Seedling quality standards and acorn availability are identified as being important in the development of a

successful oak-planting program. 

21
SURVIVAL OF UNDERPLANTED WHITE OAK AND COMPETITORS 

FOLLOWING A PRESCRIBED BURN

Dale R. Weigel 

USDA Forest Service, North Central Research Station

Bedford, IN 47421

and 

Paul S. Johnson and Daniel C. Dey

USDA Forest Service, North Central Research Station

Columbia, MO 65211

Based on our 15 years or more of research on underplanting oaks beneath shelterwoods in the Central

Hardwood Region, we have developed prescriptions for regenerating northern red oak (Quercus rubra L.)

in Missouri and Indiana and for regenerating white oak (Quercus alba L.) in Missouri. Our research suggests that there is little need for controlling competing vegetation after final removal of the shelterwood

overstory and release of the underplanted oaks in upland forests in Missouri. In contrast, survival of

northern red oak underplanted in an Indiana shelterwood can be as low as 13 to 26% after 13 years if

competing vegetation is not controlled. Oak has low regeneration potential after overstory removal when

competing with aggressive and fast growing species such as yellow-poplar (Liriodendron tulipifera L.) in

Indiana. Therefore, we recommend that competing vegetation be controlled sometime during the first 6

years following removal of the shelterwood overstory to increase the competitiveness of oak reproduction.

We are interested in how fire can be used to control oak’s competitors and to increase the developmental

potential of oak reproduction. In Indiana, we are testing the use of prescribed burning to control competing vegetation after complete removal of the overstory by adding fire as a treatment to an ongoing research

study that was designed to regenerate white oak by underplanting in a shelterwood. The study was established on the Martin State Forest (MSF) and the Paoli Experimental Forest (PEF) in southern Indiana in

1991. The prescription used in the original study included: 1) a reduction in overstory stocking to 60%,

2) treatment of competing woody vegetation with an herbicide at the time of harvesting, 3) underplanting

white oak bareroot seedlings after the initial shelterwood harvest, and 4) removal of the shelterwood overstory after 3 years.

At both of the study sites, we conducted a spring surface prescribed burn to control competing vegetation

after final overstory removal. Due to weather factors the sites were not burned the same year. The MSF

site was burned 5 years after shelterwood removal, and the PEF site was burned 6 years after shelterwood

removal. Although we did not make detailed observations on fire behavior, the fire at the MSF site burned

hotter and more completely than the fire at the PEF site did.

To quantify the response of the planted white oak seedlings and their competitors to fire, we measured

basal diameter and height of the planted oak reproduction and the tallest woody competitor within 1 m of

the oak seedling. The reproduction was inventoried in the winter before burning and after the first growing season following the fire. Before burning, white oak reproduction, on average, was taller (286 cm vs.

217 cm) and had larger basal diameters (28 mm vs. 27 mm) at the PEF site than at the MSF site.

Similarly, competitors, on average, were taller (326 cm vs. 293 cm) and had larger basal diameters (38 mm

vs. 29 mm) at the PEF site than at the MSF site. Slightly more than 20% of the white oak seedlings at PEF

suffered shoot dieback compared to more than half (58%) of the oaks at MSF. Complete mortality of

white oak seedlings was similar at both sites where 7% (PEF) and 8.8% (MSF) of the seedlings died the

first year after burning. The basal diameter and height of the planted white oak reproduction were significantly related to the probability that a stem experienced either shoot dieback or complete mortality.

22
Smaller diameter and shorter white oaks tended to be more susceptible to topkill by the fire than the larger diameter and taller stems. A similar number of competitors were present at PEF (18) and MSF (19).

However, yellow-poplar, white ash (Fraxinus americana L.), and sassafras (Sassafras albidum (Nutt.) Nees)

comprised 67% of the competitors at PEF, while only American beech (Fagus grandifolia Ehrh.) (13%)

comprised more than 10% of the competition at MSF. Initial size of competing reproduction was less

strongly related to post-fire shoot dieback or mortality than the initial size of white oak reproduction was

smaller diameter and shorter trees to be more susceptible to fire damage or mortality. Shoot dieback in

the oak competitors averaged 16.1% at PEF and 67.1% at MSF. Mortality of competitors was significantly

lower at PEF (9.3%) than at MSF (30.4%).

23
ABUNDANCE AND LOCALIZATION OF ALUMINUM IN THE FINE MYCORRHIZAL 

ROOTS OF OAK (QUERCUS PETRAEA)

Christine Wilson

Institut für Allgemeine Botanik

Johannes Gutenberg-Universität 

Mainz, Germany 55099

and

Dr. Joachim Block

Abeitlung C, Waldschutz

Forstliche Versuchsanstalt Rheinland-Pfalz 

Trippstadt, Germany 67705

This study focused on the seasonal abundance and diversity of ectosymbiotic mycorrhizae, associated with

Quercus petraea (Matt) LIEBL, at different depths in the soil of limed and unlimed forest plots at Merzalben

research station in the high elevation Palatinate Forest of Rheinland-Pfalz, Germany. The ultimate goal

was to determine the localization and abundance of potentially toxic aluminum within fine roots and their

associated mycorrhizae.

In Rheinland-Pfalz, oak trees are under increasingly severe stress. In 1984, only 8% of the oaks were in

damage classes 2 to 4, but by 1999, 45% of the oaks were in damage class 2 and 5% were in classes 3 and

4 (Kronauer 1999). Many trees are living only to 125-150 years, or less than half of their expected lifespan (Block, pers. comm.). The leading cause of forest decline in the region is generally thought to be

atmospheric pollutants and acid rain. As the soil pH drops below 4.5, continued deposition of atmospheric acids from the mountain mists into the ground water may cause argilaceous minerals such as aluminum to be solubilized into potentially toxic forms (Al 3+) (Cunningham and Saigo 1995). Aluminum

toxicity symptoms may include damage to root caps and meristems and similar restricted growth in plant

shoots with mottling and necrosis of leaves (Bollard 1993). Aluminum may bind to meristem DNA, thus

interfering with cellular division (Horst et al. 1983), and it may also precipitate essential phosphate, preventing translocation and leading to phosphate deficiency (Bollard 1993). 

Using MorinTM, a fluorescent chemical marker, according to fluorescent microscopic techniques described

by Vogelei and Rothe (1993), the extracellular and intracellular locations of aluminum can be determined.

Total cytoplasmic and bound aluminum concentrations in roots can be determined by standardized low

and high pressure extraction procedures. 

Freed acidic aluminum may enter the ectomycorrhizal tips, be deposited outside the walls or incorporated

into the mycorrhizal sheath, or may go directly into the rootlet cells where damage can occur. Certain

mycorrhizal species seem to act as a defensive barrier to the translocation of aluminum into the absorptive

zone, while other species seem to allow free passage. 

Cunningham, W.; Saigo, B.W. 1995. Environmental science: a global concern. 3d ed. Boston, MA: Wm C.

Brown Publ.: 385-387.

Horst, W.J.; Wagner, A.; Marschner, H. 1983. Effect of aluminum on root growth and mineral element

contents of roots of Vigna unguiculata genotypes. Zeitschrift fuer Pflanzenphysiologie. 104: 95-103. 

Kronauer, H. 6 Dez. 1999. Waldschadenserhebung 1999. AFZ 25: B 1089 D,: 1370-1373.

Volgelei, A.; Rothe, G.M. 1993. Anreicherung und Lokalisation von Aluminum in Wurzelspitzen von

Fichten (Picea abies [L.]. Karst.) Mitteilungen für Wissenschaft und Technik Bd X: 6: 197-204.

24
Mission Statement

We believe the good life has its roots in clean air, sparkling water, rich soil, healthy

economies and a diverse living landscape. Maintaining the good life for generations

to come begins with everyday choices about natural resources. The North Central

Research Station provides the knowledge and the tools to help people make informed

choices. That’s how the science we do enhances the quality of people’s lives.

For further information contact:

North Central Research Station

USDA Forest Service

1992 Folwell Ave.

St. Paul, MN 55108

Or visit our web site:

www.ncrs.fs.fed.us

The U.S. Department of Agriculture (USDA) prohibits discrimination in all its programs and activities on the basis of race, color, national origin, gender, religion,

age, disability, political beliefs, sexual orientation, and marital or family status. (Not all prohibited bases apply to all programs.) Persons with disabilities who

require alternative means for communication of program information (Braille, large print, audiotape, etc.) should contact USDA’s TARGET Center at (202) 720-

2600 (voice and TDD).

To file a complaint of discrimination, write USDA, Director, Office of Civil Rights, Room 326-W, Whitten Building, 14th and Independence Avenue, SW,

Washington, DC 20250-9410, or call (202) 720-5964 (voice or TDD). USDA is an equal opportunity provider and employer.
U.S. Department of Agriculture, Forest Service.

2002. Sung, S.; Kormanik, P.P.; Ostrosina, W.J.; Isebrands, J.G., eds. Eighth workshop on seedling physiology and growth problems in oak plantings

(abstracts); 2001 September 9-12; Hiawassee, GA. Gen. Tech. Rep. NC-224. St.

Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Research

Station. 24 p.

Research results and ongoing research activities in field performance of planted

trees, seedling propagation, physiology, genetics, acorn germination, and natural

regeneration for oaks are described in 21 abstract.

KEY WORDS: Plantations, propagation, regeneration.

Printed on recyclable paper.