United States

Department of

Agriculture

Forest Service

Pacific Northwest

Research Station

General Technical

Report

PNW-GTR-457

May 1999

The Forest Ecosystem Study:

Background, Rationale,

Implementation, Baseline

Conditions, and Silvicultural

Assessment

Andrew B. Carey, David R. Thysell, and Angus W. Brodie
Authors ANDREW B. CAREY is a principal research wildlife biologist; DAVID R. THYSELL is a

botanist; and ANGUS W. BRODIE was a forester, Forestry Sciences Laboratory, 3625

93d Avenue, Olympia, WA 98512-9193.
Carey, Andrew B.; Thysell, David R.; Brodie, Angus W. 1999. The Forest Ecosystem

Study: background, rationale, implementation, baseline conditions, and silvicultural

assessment. Gen. Tech. Rep. PNW-GTR-457. Portland, OR: U.S. Department of

Agriculture, Forest Service, Pacific Northwest Research Station. 129 p.

The Forest Ecosystem Study (FES) came about as an early response to the need for

innovative silvicultural methods designed to stimulate development of late-successional attributes in managed forests—a need ensuing from the exceptional and longstanding controversies over old-growth forests and endangered species concerns in

the Pacific Northwest. In 1991, scientists with the FES applied experimental, variabledensity thinning to even-aged Douglas-fir (Pseudotsuga menziesii (Mirbel) Franco)

forests on the Fort Lewis Military Reservation in western Washington after having first

accumulated extensive baseline data on arboreal rodents, small mammals, trees and

other vascular plants, and fungi. Since thinning, further research elements have been

incorporated into the FES, including top rot fungal inoculation and soil food web

response to thinnings, in addition to the ongoing prey base, vegetation, fungal, and

silvicultural assessment investigations. We present study background, rationale,

baseline conditions, and selected preliminary responses, as well as a silvicultural

assessment of the variable-density thinning.

Keywords: Variable-density thinning, Pacific Northwest, Douglas-fir, biodiversity,

northern flying squirrel, truffle, Forest Ecosystem Study, experimental silviculture.

1 Introduction

3 Study Objectives

6 Study Area

6 Location

7 Vegetation

9 Experimental Design and Rationale

9 Overview

11 Variable-Density Thinning

18 Cavities

20 Underplanting

21 Top Rot Fungi Inoculation

21 Postthinning Stand Assessment

22 Implementation

22 Grid Establishment

22 Arboreal Rodent Trap Placement

23 Small Mammal Trapping Grid and Trap Placement

23 Cavity and Nestbox Placement

Abstract

Contents
23 Silvicultural Treatment

24 Underplanting

24 Top Rot Inoculation

25 Sampling Methods

25 Vegetation

29 Den Use

29 Fungi and Soil Food Webs

31 Coarse Woody Debris, Snags, and Potential Dens

32 Forest Floor and Arboreal Mammals

33 Predators and Birds

34 Baseline Conditions

35 Prethinning and Control Stand Conditions

43 Prethinning Coarse Woody Debris

43 Standing Dead Trees

43 Arboreal Rodents

43 Forest-Floor Small Mammals

46 Postthinning Conditions

46 Silvicultural Conditions

52 Postthinning Understory Vegetation

53 Postthinning Coarse Woody Debris

54 Den Site Availability and Snags

54 Supplemental Den Use

55 Fungi

57 Discussion

57 Retrospective Aspects

61 Prospective and Experimental Aspects

65 Variable-Density Thinning

66 Operational Aspects of Variable-Density Thinning

67 Summary

69 Acknowledgments

69 Metric and English Conversions
70 References

80 Appendix 1: Forest Ecosystem Study Species List

103 Appendix 2: Nestbox and Cavity Design

104 Appendix 3: Permanent Plot Locations

108 Appendix 4: Cell-by-Cell Stand Parameters

 (From Postthinning Assessment)

122 Appendix 5: Postthinning Stand Maps
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Document continues on next page.
1

Ancient forests have achieved preeminence in conservation issues in North America

because of their dwindling area, value as habitat for threatened species of plants and

animals, contributions as reservoirs of biodiversity, and aesthetic qualities. In the

Pacific Northwest, old-growth Douglas-fir forests have been at the center of exceptional public controversy because of their importance to spotted owls, marbled

murrelets, and other species.1 This controversy has resulted in numerous appeals of

proposed timber sales, production of several environmental impact statements followed by challenges in courts, and finally a succession of three major interagency

reports: the Interagency Scientific Committee (ISC) to address the conservation of the

northern spotted owl (Thomas and others 1990); the Scientific Analysis Team (SAT;

Thomas and others 1993) that addressed other wildlife associated with late seral

forests; and the Forest Ecosystem Management Assessment Team (FEMAT 1993) that

provided a comprehensive ecological, economic, and social assessment of forest

management in the Pacific Northwest.

The opportunity to maintain biodiversity in managed forests and the need to recreate

late-seral ecosystems was recognized more than a decade ago (Carey 1985,

Ruggiero and Carey 1984, Thomas and others 1990). Thomas and others (1990)

called for experimental approaches to forest management to determine if suitable

habitat for spotted owls could be created at the same time as commodity values are

extracted from managed forests. Substantial information on biodiversity in managed

and natural Douglas-fir forests had accumulated, including Carey (1989, 1995), Carey

and Johnson (1995), Carey and others (1990, 1992, 1996a), Forsman and others

(1984), Ruggiero and others (1991), and Spies (1991). By 1991, sufficient correlational data on biodiversity and elements of forest structure existed to design an

experiment in ecosystem manipulation, the Forest Ecosystem Study (FES).

The FES was initiated (Carey and Miller 1991) to address development of spotted owl

habitat through experimental manipulation of managed stands. Both silvicultural

treatments and direct interventions to enhance biodiversity were incorporated into the

experimental design. After a review of potential study locations, the Fort Lewis Military

Reservation was chosen because (1) the forests were predominantly Douglas-fir, thus

maximizing both potential for positive prey-base responses (Carey 1995) and applicability to the most common forest type in western Washington and Oregon; (2) the

forests were 50- to 70-year-old, even-aged, closed-canopy stands regenerated after

clearcutting and representative of commonly occurring second growth and thus

presented opportunities for ecosystem management; and (3) Fort Lewis had large,

forested areas that allowed selection of homogeneous blocks for replicates of treatments that would minimize confounding effects of heterogeneity and forest fragmentation on ecosystem processes.

Following site selection in 1991, a memorandum of understanding (MOU) between the

Pacific Northwest Research Station and Fort Lewis was developed that protected the

study sites for a minimum of 20 years. In 1992, Fort Lewis became a Designated

Conservation Area (DCA) for the northern spotted owl with publication of the final draft

Introduction

1 Scientific names, authorities, and sources of all species not

provided in the body of the report are given in appendix 1. Where

common names have not been recognized or assigned, the

scientific name is used throughout.
2

Table 1—Schedule of Forest Ecosystem Study activities

Year and quartera

1991 1992 1993 1994 1995 1996

FES activity 123412341234123412341234

Treatments and assessments:

 Site selection/MOU •

 Grid establishment •

 Arboreal rodent trap placement •

 Small mammal trap placement • •

 Cavity and nestbox installation • •

 Silvicultural treatment • •

 Underplanting •

 Top rot inoculation •

 Silvicultural assessment • •

Sampling of response variables:

 Prethinning vegetation • •

 Postthinning vegetation • •

 Cavity tree age and growth • • • •

 Den-site use by flying squirrels ••••••••

 Epigeous fungi • •••••

 Hypogeous fungi (truffles) ••••••••••• •

 Soil food webs ••••

 Prethinning coarse woody debris •

 Postthinning coarse woody debris • •

 Den-site availability •

 Arboreal rodent trapping •••••••••••

 Small mammal trapping • • • •

 Owl survey •

 Winter bird survey •

 a Calendar quarters: 1 = January-March; 2 = April-June; 3 = July-September; 4 = October-December.

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3

of the northern spotted owl recovery plan, because it was “an important location to

assist in reestablishing demographic interchange between owls in the Cascade

Mountains and owls on the Olympic Peninsula” (Lujan and others 1992).2 Subsequently, a consultation (under Section 7 of the Endangered Species Act) was undertaken with the Olympia, Washington, office of the U.S. Fish and Wildlife Service, as to

the suitability of the proposed habitat manipulation. The experiment was deemed

compatible with management of Fort Lewis as a DCA, and thinning operations were

performed during January-April 1993. Variable-density thinning (VDT), a cornerstone

of our experiment, was a new silvicultural technique (Carey 1994b, Carey and Curtis

1996, Carey and Miller 1991) devised to create spatial heterogeneity in composition

and structure that would mimic conditions found in old-growth forests. Supplemental

funding that allowed expansion of research was obtained from the U.S. Army and the

U.S. Department of Agriculture, National Research Initiative.

In this report, we provide the background, historical context, and objectives of the FES;

an outline of the experimental design, implementation schedule (table 1), and sampling methods; pretreatment and baseline conditions; lists of species identified during

the project (appendix 1); an assessment of the posttreatment silvicultural conditions;

and opportunities for potential research collaboration. We hope this information will be

useful to future research collaborators, managers seeking to design or implement

adaptive management programs, and readers of related publications who want further

detail on some aspect of the study.

Statistics and statistical tests are presented for illustrative purposes only. Because

presented data are preliminary, we primarily use the nonparametric Mann-Whitney

U-test and Wilcoxon matched-pairs signed-rank test (Norus ̆is 1993) to contrast,

respectively, two independent samples or two paired samples. We also use the twosample t-test where necessary assumptions were met. We recognize many of our

data are amenable to the use of analysis of variance (ANOVA), but because this

publication is intended as an establishment report rather than a report of research

results, such analyses are not presented herein. More refined and detailed analyses

and discussions of specific study components will be published in the future.

The FES is an application of experimental management techniques to second-growth

forests. The techniques are intended to hasten development of late-seral “composition, structure, function, ecologic products, and economic products” (Carey 1994a).

The study was designed to test whether late-seral forest attributes3 can be developed

from stem-exclusion-phase (Oliver and Larson 1990) forests through silviculture, and

whether providing supplemental dens can increase the density of northern flying

squirrels (the primary prey of the northern spotted owl) in second-growth forests. The

experiment was formulated to address specific applied and basic objectives relative to

forest structure, composition, and ecological function.

Study Objectives

2 Currently, northern spotted owls are not known to occur within

Fort Lewis.

3 Late-seral attributes include large-diameter trees, well-developed

understories, high vegetation site type diversity, and associated

characteristics of high fungal diversity and abundant small mammal

populations.
4

The applied research objectives of the study are to determine if:

1. Woody plant species diversity, spatial heterogeneity in vegetation, and vertical

diversity in vegetation can be manipulated through variable-density thinning and

underplanting that do not require replacing the existing stand.

2. Enhancing the growth and diversity of woody plants also will be accompanied by

increased abundance and diversity of ectomycorrhizal fungi.4

3. Increasing den availability through direct intervention (creating cavities in live trees

and adding nest boxes) will increase flying squirrel populations.

4. Inoculation of live trees with stem decay fungi will hasten their use by primary

excavators (woodpeckers) and thus hasten the development of cavities useful to

other secondary cavity dwellers (including the northern flying squirrel).

5. Increasing flying squirrel numbers and woody plant diversity will then have a

synergistic effect in promulgating an abundance and diversity of ectomycorrhizal

fungi and concurrently increase forest productivity.

6. Wood production is compatible with advancing late-seral forest conditions.

7. Variable-density thinning compares favorably with traditional commercial thinning in

promoting tree growth and yield.

The basic research objectives include:

1. Elucidating the community ecology of ectomycorrhizal fungi in relation to seasonal

and annual climatic variability, composition and abundance of the rodent community,

composition and abundance of the understory vascular plant community, coarse

woody debris, and silvicultural treatments.

2. Evaluating the relative influence of den availability, abundance and diversity of food,

abundance of protective cover (understory vegetation), and abundance and diversity of predators on populations of northern flying squirrels, Douglas’ squirrels, and

Townsend’s chipmunks to determine factors limiting sciurid populations and regulating arboreal rodent communities.

3. Elucidating factors controlling composition and abundance of the forest-floor

mammal communities, especially the influence of understory vegetation and coarse

woody debris.

4. Testing hypotheses about the utility of vertebrate communities as ecosystem

indicators. The indices, developed and explained by Carey and others (1996a),

include:

a. biomass of arboreal rodents as an indicator of ecosystem productivity; and

b. integrity of the forest-floor mammal community as an index of forest-floor

function.

4 The hypogeous sporocarps of ectomycorrhizal fungi (truffles) are

the primary food of the northern flying squirrel. The flying squirrel

disseminates the spores of the fungi and serves as the primary

prey of the northern spotted owl. The sporocarps also are

important food for several other animals (e.g., voles, chipmunks,

other squirrels).
5

The following tabulation indicates which methods will be used to meet specific objectives and the pages on which the methods are discussed:

Objective Method to meet specific objectives (with page nos.)

Applied

1 Variable-density thinning (11, 26); underplanting (20, 24); vegetation

sampling (25, 27).

2 Vegetation sampling (25, 27); fungal sampling (29-31); fecal pellet

analysis (33).

3 Supplemental den establishment and use (18, 23, 32); arboreal rodent

trapping (22, 32).

4 Top-rot inoculation, monitoring (21, 24).

5 Arboreal rodent trapping (22, 32); vegetation sampling (25, 27); fungal

sampling (29-31); fecal pellet analysis (33).

6 Vegetation sampling (25, 27); fungal sampling (29-31); trapping (22, 23,

32, 33).

7 Stand assessment (21, 26); vegetation sampling (25, 27).

Basic

1 Vegetation sampling (25, 27); fungal sampling (29-31); trapping (22, 23,

32, 33); coarse woody debris sampling (31); fecal pellet analysis (33).

2 Supplemental den establishment and use (18, 23, 32); vegetation

sampling (25, 27); fungal sampling (29-31); arboreal rodent trapping (22,

32).

3 Vegetation sampling (25, 27); small mammal trapping (23, 33); coarse

woody debris sampling (31).

4 Arboreal rodent and small mammal trapping (22, 23, 32, 33).

The increases in productivity of plants, foliage, fruits, and seeds predicted to result

from VDTs should increase abundance and diversity of invertebrates, insectivores,

amphibians, birds, and mammals. Many small mammals, especially Townsend’s

chipmunks, northern flying squirrels, and Douglas’ squirrels, are important prey for

predators such as weasels, marten, owls (e.g., spotted and great-horned), and raptors

(e.g., northern goshawk). Spotted owl abundance, requirements for late-seral forests,

and use of second-growth forests differ with prey diversity and total prey biomass

(Carey and others 1992, Carey and Peeler 1995). Thus, experimental treatments

should affect a hierarchy of ecosystem processes from symbioses between fungi and

plant roots to predation at the highest trophic level (the spotted owl, marten, fisher,

great-horned owl, and northern goshawk). Response variables representative of this

hierarchy of ecosystem biodiversity that will be evaluated during the study include:
6

1. Composition and diversity of selected soil microbial communities (including fungi,

bacteria, and nematodes)

2. Abundance and diversity of hypogeous and epigeous sporocarps of ectomycorrhizal

fungi in or on the forest floor

3. Diversity of fungi in the diets of the primarily mycophagous northern flying squirrel and

the facultatively mycophagous Townsend’s chipmunk and southern red-backed vole

4. Abundance and diversity of understory vegetation

5. Abundance of Townsend’s chipmunks and northern flying squirrels

6. Space and den use by northern flying squirrels

7. Diversity and abundance of the forest-floor community of insectivorous, frugivorous,

granivorous, herbivorous, and mycophagous small mammals (shrews, shrewmoles, voles, and deer-mice)

8. Frequency of encounter of predators of small mammals (e.g., weasels, owls, hawks)

9. Tree diameter at breast height (d.b.h.), height to live crown, total height, crown

width, and growth rates

10. Stand basal area and volume of wood

These response variables comprise the basis for an ecosystem study based on trophic

pathways supporting spotted owls and other predators and will provide a template for

evaluating the degree to which our applied and basic research objectives are met.

Meeting many FES objectives will be contingent on a thorough understanding of actual

postthinning forest conditions, including stand density, relative density, and basal area.

The FES is primarily a stand-level experiment, and eventually all response variables

will be assessed in relation to actual posttreatment conditions. Some response

variables (understory vegetation, hypogeous and epigeous fungi) will further be

evaluated in relation to postthinning conditions at a specific plot or at plots of similar

thinning intensity. Thus, we incorporated a complete, posttreatment assessment of

stand conditions.

Finally, the study site, experimental design, and sampling framework meet our operational objective of creating a foundation and stimulus for interdisciplinary studies of

ecosystem processes.

The FES is in the Rainier Training Area, Fort Lewis Military Reservation, south of the

Nisqually River, approximately 30 km5 east of Olympia, Washington, in the southern

Puget Trough Physiographic Province (Franklin and Dyrness 1973). The present-day

landforms and substrate were deposited when the Puget Lobe of the Vashon Stade of

the continental glacier retreated 13,000-15,000 years ago (Kruckeberg 1991). Topographic relief is moderate to rolling, with occasional steeper areas surrounding small

kettle depressions. The experimental sites contain no year-round creeks, ponds, or

wetlands. The study blocks are on glacial uplands, terminal moraines, and glacial till

Study Area

Location

5 Metric units are used except where English units were the

operational units. A conversion table is given on p. 69.
7

plains; root penetration into the soil is moderately shallow, and drainage is classified

as somewhat excessive (Pringle 1990). The maximum elevation at Fort Lewis is 168 m

above sea level; the study blocks range from 120 to 165 m in elevation. Annual

precipitation for Fort Lewis is 800-900 mm with only 10 to 15 percent of the annual

precipitation falling during the peak growing months of June-September (Pringle

1990). Soils are coarse-textured gravelly and gravelly sandy loams (of glacial till and

glacial outwash origin) of Tenino and Everett soil series (Pringle 1990). The excessively drained soils combine with low summer rainfall to create droughty conditions

where Douglas-fir typically dominates and also regenerates in the understory (Franklin

and Dyrness 1973, Pringle 1990).

Fort Lewis is an area of relatively isolated forests, prairies, and wetlands bounded on

the northwest by Puget Sound, and on the north, east, and south by agricultural and

urban areas. The forests of Fort Lewis provide the only forested connection between

the Black Hills and Olympic Mountains to the west, and the Cascade Range to the

east; its unique position is the primary reason for its being a Designated Conservation

Area for the northern spotted owl (Lujan and others 1992).

Large portions of Fort Lewis forests were clearcut in the early 20th century and regenerated from natural seeding; the second growth is now actively managed. Some areas

contain residual Douglas-fir as legacies from the previous stand. Where present,

these large, dominant Douglas-fir project above a second, dense layer of codominant

Douglas-fir. Many stands at Fort Lewis are now in the stem exclusion phase of forest

development (Oliver and Larson 1990) or competitive exclusion stage of forest ecosystem development (Carey and others 1996a). In this stage of development there are

abundant dying suppressed trees, small-diameter fallen trees, and standing dead trees

throughout the stands. In contrast, intensively managed stands at Fort Lewis are

uniform in tree diameter and canopy height, contain few old-growth legacy trees,

contain few small diameter, suppressed, or dead trees, and are in the understory

reinitiation stage (Oliver and Larson 1990, Carey and others 1996a).

The FES is divided between two forested tracts that have experienced widely disparate management histories. The two portions of the study are about 3 km apart and are

referred to as the Star/Stellar (SS) forest (fig. 1) and the Farley/Hill (FH) forest (fig. 2).

The FH forest, comprised of the Farley and Hill blocks, was clearcut in 1925 and,

following natural regeneration, was lightly thinned in 1972 and again between 1979

and 1989. During these commercial thinnings, live, recently dead but salvageable,

and fallen trees were routinely removed; 3.0-5.8 thousand board feet (MBF) per acre

were removed per entry. In 1991 the FH blocks were dominated by 55- to 65-cm

d.b.h. Douglas-fir (39-43 m tall) with small amounts of black cottonwood, red alder, and

bigleaf maple (3 to 8 percent total stems). Trees in FH appeared to have been grown

under moderate stocking levels since stand initiation, with many trees having ample

crowns and lateral branches to near ground level. The low understory was well

developed and was dominated by salal, California hazel, swordfern, and bracken fern.

Other common Douglas-fir series associates (Henderson and others 1989), such as

serviceberry, western fescue, baldhip rose, and creeping snowberry, were common.

Vegetation
8

Figure 1—Blocks 1 (Star 101-104) and 2 (Stellar

201-204) show the spatial arrangement of the four

main treatments. Each of the eight stands is a

280- by 280-m square, with 80 m between stands.

Figure 2—Blocks 3 (Farley 301-304) and

4 (Hill 401-404) show the spatial

arrangement of the four main

treatments. Each of the eight stands is

a 280- by 280-m square, with 80 m

between stands.

  yyy yyy  

yyy yyy yyy y



A

1

104

2345678

BCDEFGH

1

A

102

2345678

BCDEFGH

1

A

101

2345678

BCDEFGH

A

1

103

2345678

BCDEFGH

A

1

202

2345678

BCDEFGH

A

1

204

2345678

A BCDEFGH

1

203

2345678

BCDEFGH

A

1

201

2345678

BCDEFGH

STAR

(101-104)

STELLAR N

(201-204)

280 m

Control (no treatment) Cavities only (no thinning)

Thinning only Thinning and cavities 280 m y



yyyy    yy

(301-304)

FARLEY yy

HILL

(401-404)

N

280 m

Control (no treatment) Cavities only (no thinning)

Thinning only Thinning and cavities

280 m

402

403

404

401

301

302

303

ABCDEFGH

12345678

ABCDEFGH

12345678ABCDEFGH

12345678

ABCDEFGH

12345678 ABCDEFGH 12345678

ABCDEFGH 12345678

304 ABCDEFGH

12345678

ABCDEFGH 12345678
9

Though the likelihood of laminated root rot (Phellinus weirii 6) in these blocks was high,

actual occurrence was difficult to judge because diseased or infected trees were

probably removed during the thinnings. Because of past thinning entries, and because

very few trees were left standing during the 1925 harvest, the FH forest had few

residual Douglas-fir trees and snags and minimal cover of coarse woody debris (Carey

and others 1996b).

The SS forest, consisting of the Star and Stellar blocks, was harvested in 1937 and

had not been further manipulated. In 1991, SS was a closed-canopy forest dominated

by 30- to 45-cm d.b.h. Douglas-fir (30-35 m tall), with a few western hemlocks, western

redcedars, and Pacific yews. Trees killed by suppression and trees with small crowns

were abundant, suggesting a very high stocking level since stand initiation. Understory vascular vegetation was sparse, with only small amounts of salal and Cascade

Oregongrape. Douglas-fir series associates (Henderson and others 1989), such as

serviceberry, western fescue, baldhip rose, and creeping snowberry, also were found

in this forest. The forest floor typically was dominated by mosses, particularly

Kindbergia oregana (Sull.) Ochyra and Hylocomium splendens (Hedw.) B.S.G.7

Exceptions to the high canopy closure and sparse understory were found in areas of

laminated root rot infection. In these root rot centers, understory shrubs, forbs, and

ferns were well developed. In SS, some Douglas-fir trees and snags were left during

the 1937 entry, as was a substantial amount of large coarse woody debris (Carey and

others 1996b).

We used a complete randomized block design of four blocks with four treatments per

block, to which were applied the three principle manipulative components of the

experiment: (1) a VDT treatment (with three different thinning density subtreatments);

(2) den augmentation with cavities and nestboxes; and (3) underplanting to increase

species and structural diversity. The 13-ha stands, each containing a 280- by 280-m,

8 by 8 grid (with 40- by 40-m, 0.16-ha grid cells), were the operational units for application of treatments. The four main treatments, each randomly assigned to one of the

four stands per block (figs. 1 and 2) in all blocks, were:

1. no treatment (control = CT);

2. VDT with 3 thinning intensities and underplanting;

3. den-augmentation-only (cavities in live trees and nest boxes);8 and

4. den augmentation plus VDT and underplanting.

Experimental

Design and

Rationale

Overview

6 Larson and others (1994) have proposed the name, Inonotus

sulphurascens (Pilat) M. Larsen and others, for Douglas-fir-infecting

laminated root rot. We recognize the distinction between the

Douglas-fir form and the western redcedar form of laminated root

rot (Thies and Sturrock 1995); however, to avoid confusion, we use

the name Phellinus weirii for the Douglas-fir form.

7 Moss names and authorities are from Schofield (1992).

8 Because only baseline and preliminary responses of selected

variables are presented, we do not herein present or evaluate the

effect of den-augmentation-only. As such, unless specifically noted,

control will refer to both no treatment (1) and den-augmentationonly (3).
10

Figure 3—Actual and potential applications of variable-density thinning. Star 101 (A) shows random

assignment of subtreatments with actual root rot pockets. Farley 302 (B) shows random assignment of

subtreatments with simulated root rot pockets. Systematic assignment (C. & D.) of light and heavy thins (as

possible operational applications) in a 2 to 1 ratio of LT to HT.



















A

1

2

3

4

5

6

7

8

A. Star 101: random with actual

root rot

BCD E F GH

A

1

2

3

4

5

6

7

8

C. Systematic: 2 LT to 1 HT

BCD E F GH

A

1

2

3

4

5

6

7

8

B. Farley 302: random with

simulated root rot

BCD E F GH

A

1

2

3

4

5

6

7

8

D. Systematic: 2 LT to 1 HT

Root rot thin

Heavy thin

Phellinus pocket

Light thin

BCD E F GH
11

Subtreatments were randomly assigned to the forty-nine 40- by 40-m grid cells in each

of the two treated stands (thinning and thinning plus dens) per block (see fig. 3 for

example applications). Thus, the entire study consisted of 16 stands, 13 ha each, for a

total study area of 208 ha. Grids were templates for treatments (figs. 1, 2 and 3), for

cavity and nestbox installation (fig. 4), for sampling arboreal rodents and forest-floor

mammals (fig. 5), fungi, soil food webs, and vegetation (fig. 6), and for evaluating

prethinning and postthinning stand conditions.

Background—Development of the silvicultural prescription took place in two stages.

First, a desired future condition was defined that would provide a suitable environment

for foraging and roosting by spotted owls. Second, the study areas were surveyed to

determine the steps necessary to achieve the desired future condition in the shortest

time possible. The desired future condition was based on information summarized in

Carey and others (1991b, 1992), Gutiérrez and Carey (1985), Ruggiero and others

(1991), USDA Forest Service (1988), and Thomas and others (1990), and research on

characteristics of the habitat of spotted owls and their prey (Carey 1995, Carey and

Johnson 1995, Carey and Peeler 1995).

The desired future condition was an environment suitable for spotted owl foraging.

Suitability for spotted owl foraging encompasses both an abundance of prey sufficient

to provide owls with a reasonable return per unit effort of foraging and vegetation

structure amenable to the sit-and-wait foraging strategy used by spotted owls. The

structure also should be amenable to spotted owl roosting. Silvicultural manipulation

has the potential to produce the structure desired and to benefit prey by providing a

diversity of woody plants that support ectomycorrhizal fungi of the types consumed by

flying squirrels and other small mammals. Both herbaceous and woody understory

vegetation also contribute other food types (foliage, seeds, fruits, nuts, associated

fungi) that are used by ancillary prey of owls (hares, rabbits, mice, and voles).

The desired future structure consists of the spatial arrangement of vegetation; spatial

scale is also important. Areas of high vertical diversity of vegetation should alternate

with areas of sparse understory. Sparse understory allows owls to spot, track, and

attack prey; a column of vegetation provides a vertical array of perches from which

owls can pounce or fly short distances to prey. Data collected from a variety of plot

radii ranging from 5.6 to 25 m, and grids and transects with distances between stations

ranging from 20 to 100 m in old forests (Carey 1995; Carey and others 1991b, 1992;

Carey and Johnson 1995), suggested patterning in the overstory should be on a scale

of 80 m with areas of sparse and variable understory twice as abundant as areas of

dense understory.

The spatial heterogeneity observed in natural old forests could have resulted from a

combination of three different processes: suppression of subordinate trees in densely

stocked stands; gap formation resulting from the breakage of trees with top rot infestation or death of trees from senescence, windthrow, lightning strikes, disease, insects,

or other causes; and gaps resulting from locally intense, laminated root rot infestations

or small-scale catastrophic disturbances (fire, windthrow). Variable-density thinning

was designed to simulate these processes. Light thinning simulates suppression

Variable-Density

Thinning
12

Figure 4—Nestbox, cavity,

and top rot inoculation

locations in Star 102, 103;

Stellar 201, 203; Farley 303,

304; and Hill 403,404.

Squares (nestboxes) and

circles (cavities) are locations

of supplemental den sites.

Open boxes and circles are

cavities and nestboxes from

fall 1991. Solid boxes are the

additional eight nestboxes

added in winter 1995. Filled

circles mark locations of top

rot inoculations in addition to

cavities. All grid corners also

served as stations for trapping

arboreal mammals.

Figure 5—Design of live

trapping grids (LTG) used

for trapping arboreal rodents

(A-H, 1-8), and small mammal

grids (SMG, in bold) used for

trapping forest floor small

mammals (Q-Z, 1-10). Both

grid types are located on all

16 stands.

A BCDE FGH

1

2

3

4

5

6

7

8

280 m

280 m

A B CD E F GH

1

2

3

4

5

6

7

8

280 m

280 m

Q R STU V WX Y Z

1

2

3

4

5

6

7

8

9

10

20 m

40 m

40 m

20 m

SMG

LTG

200 m
13

mortality; heavy thinning simulates gap formation as a result of individual tree breakage or death; and the root rot treatment, contributing to, but not essential to our goals

of creating spatial heterogeneity, mimics small-scale catastrophic disturbance (e.g.,

root rot, fire, windthrow).

Natural or management-induced canopy openings

can produce a vertical array of vegetation—a

column of vegetation from the forest floor to the

canopy. The vegetation can consist of low shrubs

and ferns (e.g., salal, Oregongrape, swordfern),

tall shrubs (e.g., huckleberry, oceanspray, hazel,

vine maple), conifer seedlings and midstory trees

(e.g., western hemlock, western redcedar),

deciduous trees (e.g., red alder, bigleaf maple,

Scouler’s willow), and lower branches of overstory

conifers with deep crowns. Low understory

shrubs provide food and cover for spotted owl

prey; tall shrubs, saplings, understory trees,

midstory trees, and overstory trees provide the

array of foraging perches owls use to exploit the

prey base. The column of vegetation and perches

provides protected roost sites for owls. Undesirable structure includes large vertical gaps between the lower crown and understory vegetation,

Figure 6—Permanent

vegetation plot configuration.

Plots were marked with a 1-mlong PVC pipe inserted 66 cm

in the ground at plot center.

The inner 5.6-m-radius plot

was used for understory

composition, cover, and

structure and for epigeous

fungal composition and

biomass. The outer, 12.6-mradius plot was used for

overstory tree composition

and tally. Truffle plots were

located systematically, 10 m

apart after a random start.

Plots were identified by stand

and cell number, in this case

D/E 2/3.



 









A

1

2

3

4

5

6

7

8

BCDE FGH

40 m

5.6 m

12.6 m

40 m

Permanent plots

Truffle plots

(4 m ) 2

D/E 2/3
14

Table 2—Block descriptions of live trees in the Star, Stellar, Farley, and Hill blocks based

on prethinning prism plots

Height

to live

Block Year of Basal crown

name origin Harvest activities area Douglas-fir ± S.E. Other tree speciesa

m2/ha(ft2/acre) Percent Meters

Star 1937 Unthinned since 42.2 98 18.8 Western redcedar,

1937 harvest (184) ± 0.3 western hemlock,

Pacific yew

Stellar 1937 Unthinned since 50 99 18.7 Western hemlock,

1937 harvest (218) ± 0.5 big-leaf maple

Farley 1925 Clearcut 1925. 45 97 20.9 Red alder,

Thinned twice: (196) ± 0.6 black cottonwood,

1972, 1979-89 big-leaf maple,

western hemlock

Hill 1925 Clearcut 1925. 43.6 92 21.4 Red alder,

Thinned twice: (190) ± 0.6 black cottonwood

1972, 1979-89

a Scientific names are provided in appendix 1.

an absence of understory vegetation, and a continuous, dense cover of low shrubs.

The most difficult aspect of producing the desired future structure is stimulating

vegetation to fill intermediate heights (2 m to base of live crown). Height to live crown

for dominant Douglas-fir trees averaged 16 to 21 m in Star/Stellar stands and 18 to 24

m in Farley/Hill stands (tables 2 and 3), suggesting shade-tolerant conifers or understory deciduous trees would be required to fill this gap where subordinant and codominant trees were removed by thinning. Our goals for the desired future structure in

areas of high vertical vegetative diversity were:

1. Vertical diversity of vegetation using the Berger-Parker Index9 > 2 (old-growth

averages 1.7; spotted owl roosts, 2.3)

2. Shrub cover (≤ 2 m tall) > 40 percent

3. Understory cover (subcanopy, tall shrubs > 2 m tall) > 24 percent

9 A measure of foliage-height diversity. See Magurran (1988) and

Carey and others (1991b). In this case, four strata are considered:

forbs, ferns, and trailing vines; low shrubs; subcanopy trees and tall

shrubs; and canopy trees. During forest development (after the

stem exclusion phase), canopy cover becomes less dominant in

relation to the cover of the other strata, and thus the index becomes

larger. We use the reciprocal form of the B.P. Index so that it

increases with increasing evenness among the strata.
15

Table 3—Mean height, mean d.b.h., and mean height to live crown for Douglas-fir

site trees from pretreatment prism plotsa

Stand Height to live

Block name number Height ± S.E. D.b.h. ± S.E. crown ± S.E.

Meters Centimeters Meters

Star 101 31.5 ± 0.6 38.6 ± 2.0 16.5 ± 0.9

102 34.1 ± 0.7 43.4 ± 1.5 20.7 ± 0.7

103 32.3 ± 0.7 46.5 ± 1.8 18.5 ± 0.5

104 32.9 ± 0.7 45.0 ± 2.0 18.4 ± 0.7

Stellar 201 31.9 ± 0.6 42.0 ± 1.7 16.4 ± 0.7

202 34.5 ± 0.8 46.6 ± 1.3 17.9 ± 0.8

203 32.3 ± 0.8 37.3 ± 1.8 19.7 ± 0.9

204 33.1 ± 0.2 41.5 ± 1.6 20.8 ± 1.0

Farley 301 43.1 ± 0.8 61.6 ± 1.9 19.1 ± 0.7

302 42.0 ± 0.5 59.7 ± 1.5 23.6 ± 1.1

303 41.8 ± 0.8 63.8 ± 1.8 22.0 ± 1.1

304 41.4 ± 0.6 62.3 ± 1.3 17.9 ± 1.1

Hill 401 39.1 ± 1.6 56.2 ± 3.4 21.7 ± 1.3

402 40.9 ± 1.4 62.6 ± 2.3 19.9 ± 1.3

402 43.2 ± 0.8 65.7 ± 1.9 22.7 ± 1.1

404 41.6 ± 1.3 60.2 ± 2.1 21.2 ± 0.9

a 30 site trees per stand were sampled.
16

Relative density—We adopted Curtis’ (1982) relative density (RD) as a broadly

applicable index of stand density that is also useful in designing silvicultural

prescriptions. Relative density varies along a continuum from zero (no trees) to a

species-specific biological maximum in extremely heavily stocked stands (Curtis

1982). Though empirically derived data relating understory vegetative development (and other forest conditions) to stand relative density are not extant, the

range of relative density for a given age stand encompasses recognizable stages

of forest development and of responses to natural forest disturbance (Carey and

others 1996a, Curtis 1997, Oliver and Larson 1990). For 50- to 60-year-old, evenaged Douglas-fir stands, we suggest the following RD ranges as being indicative

of particular forest conditions.10

1. RD < 3.25 (metric) represents larger gaps in the forest canopy, which, under

natural conditions, could be formed by multiple tree blowdown, small fires,

localized insect outbreaks, or root rot pockets. In such areas, there is little

intertree competition, and abundant insolation is available to understory vegetation. Depending on the size of such areas, and on age and species composition of the canopy trees, these gaps may fill by lateral branch growth, by growth

of shade-tolerant trees and shrubs, or by shade-intolerant trees such as Douglas-fir.

2. RD 3.25-4.75. Within this range, intertree competition increases rapidly.

Subcanopy insolation is reduced, thereby resulting in less understory development. Natural canopy gaps within this range of RD may be the result of smallscale endogenous disturbances such as the death, defoliation, tree-fall, or

lightning strike of one or a few trees. Stand-level tree growth within this range

is somewhat sacrificed to understory development, though forest floor vegetation is less well developed than when RD is < 3.25. The major source of light

below the canopy is from short-duration sunflecks through a moderately open

canopy.

3. RD 4.75-6.75. Within this range, the canopy is increasingly closed, with little

direct light reaching the forest floor. Intertree canopy competition increases,

resulting in near-suppression mortality conditions at the upper end of this range.

Understory vegetation is typically sparse and poorly developed. Many evenaged stands under commercial management for timber production are managed

within this range; stand tree growth is maximized and tree form and development are both quite uniform.

4. RD > 6.75. Above RD 6.75-7.00, forest stands enter a state of suppression

(Curtis 1982), where severe crowding results in reduction of tree growth and

significant tree mortality. Individual trees are under extreme intertree competition, crown development is restricted, and understory vegetation is typically

quite sparse.

10 Continued sampling and analysis on the Forest Ecosystem Study

will evaluate and further refine these suggested relative density

categories.
17

Subtreatment descriptions—Two thinning entries were scheduled as part of the

FES. The first entry was made during January-April 1993; the second is scheduled for

2003 but is provisional. Prescription targets for the first entry were formulated for

forest stands under “normal”11 conditions for 50- to 60-year-old, even-aged, unmanaged

Douglas-fir forests. An average stand-level residual tree density target was specified

as 220 trees/ha, representing a relative density of between RD 4 and RD 5 (metric) for

such stand conditions. To achieve this stand density and the desired spatial diversity

of densities within stands, the prescription for the first entry had two primary components (subtreatments):

1. A thinning of subordinant and codominant trees > 20 cm d.b.h. to reduce the

density of overstory to RD 6 (metric) with a thinning ratio d/D12 of 0.8-1.0. For a

50- to 60-year-old forest under such normal conditions, this represents a tree

density of approximately 310 trees/ha with an average spacing13 of 5.8 m

between trees (= light thin, LT).

2. A thinning of subordinant and codominant trees > 20 cm d.b.h. to reduce the

density of overstory trees to RD 4 (metric), or about 185 trees/ha with an average

spacing of 7.3 m between trees (= heavy thin, HT).

The prescription specified the removal of suppressed and subdominant trees of

the current cohort (e.g., thinning from below); this equates to a thinning ratio (d/D)

of 0.8-1.0. Marking guidelines specified retention of all large standing dead trees

and all deciduous trees (trees retained included black cottonwood, bigleaf maple,

red alder, Pacific madrone, and Scouler’s willow). Similarly, deciduous shrubs

were favored for retention; these included red huckleberry, oceanspray, and

California hazel. Existing roads and skid trails were used wherever possible to

reduce impacts on stands; where skid trails were lacking, they were preferentially

located in the more heavily thinned areas.

11 A “normal” forest is described as a stand where every tree is

using all its available space, and where competition, in terms of

individual tree growth, represents an “average maximum” (Curtis

1970). The choice of the term “normal” may mislead some,

because it does not refer to modal or average conditions in

naturally regenerated forests of the Pacific Northwest, and does not

refer to the normal or Gaussian distribution. Actual thinning

densities were applied by using foresters’ professional judgment to

avoid severely reduced wind resistance in dense stands and to

create variable-density in the more open stands.

12 d/D is the ratio of the quadratic mean d.b.h. of the cut trees (d) to

the quadratic mean d.b.h. of the original stand of trees (D). The

ratio, d/D, indexes the type of thinning by comparing the original

stand diameter distribution with that of the cut trees. When d/D =

1.0, this indicates a systematic thinning; when < 1.0, a thinning

from below; and when > 1.0, a crown thinning (Bennett and

Maguire 1995).

13 Our tree spacing targets do not imply that we sought to achieve

even or regular spacing conditions. Even at the 40- by 40-m scale,

we attempted to capitalize on naturally occurring clumps and

openings. As such, spacing values are averages only.
18

Prethinning forest cruise surveys of the study area revealed that pockets of laminated

root rot occupied 7 to 15 percent of the area of the younger, unmanaged stands (Star

and Stellar); root rot may have been present in the older, managed stands (Farley and

Hill) but infection pockets were not apparent. Because root rot can lead to canopy

openings, we incorporated a root rot treatment in the prescription. This became a third

component (subtreatment) of the thinning prescription:

3. Removal of low-vigor trees from root rot infection centers to 10 m beyond the

perimeter of the pocket of dead and dying trees; apparently healthy, dominant trees

were retained. Approximate residual tree density for the root rot areas was 40

trees/ha > 20 cm d.b.h., or an RD 1.5 (= root rot thin, RT).14

Because this is an experimental study, we applied the root rot treatment equally

(15 percent of stand area) to all stands selected for thinning. Where root rot pockets

were apparent, they were mapped and treated; where not apparent, the treatment was

randomly assigned. Thus, the same overall treatment was applied to all treated

stands.

The spatial arrangements of the treatments consisted of first treating existing root rot

pockets in the most heavily infected stand—15 percent of the area. Natural root rot

pocket size averaged 0.07 ha. Where root rot pockets comprised < 15 percent of the

remaining stands, root rot treatments were randomly applied to the 0.16-ha blocks to

achieve consistency in the experimental treatment. Next, the light and heavy thinnings

were applied to the 0.16-ha blocks (40 by 40 m) in a 2:1 ratio over the remaining area

by random assignment.15 Additional entries (at about 10-year intervals) will be necessary to achieve the desired future condition: eventually the 2:1 ratio will be reversed to

heavy:light.

Alternating the heavy and light thinnings should produce highly variable light regimes

on the forest floor that result in a mosaic of vegetation site types, some with a high

degree of vertical layering, and some with simple structure. Variable-density thinning

has the additional advantage of maintaining stand wind resistance while increasing the

amount of light available to the understory and maintaining the stand composition

necessary for wood production. The multiple entries that are a part of VDT are necessary to achieve the final desired future condition (Curtis and Carey 1996).

The amount of old forest used by spotted owls in the Pacific Northwest reflects regional variation in prey abundance; selection of stands for foraging also reflects prey

abundance (Carey and others 1992). The primary prey of the spotted owl in the

Douglas-fir and western hemlock forests of western Washington is the northern flying

squirrel (Carey 1995; Forsman and others 1984, 1991), which uses cavities in trees

14 Our choice of root rot treatment was guided by Thies and

Sturrock (1995). Due to site-specific windthrow considerations and

the 40- by 40-m size of our operational thinning units, we chose

10-m buffers rather than their suggested 15 m.

15 Figure 3 illustrates examples of VDT application with actual RT

pockets (3a), VDT with simulated RT (3b), and systematic HT and

LT applications, such as could be used in an operational, rather

than an experimental, setting (3, c and d).

Cavities
19

(both live and dead), stick nests, and moss-leaf-twig nests as dens (Carey 1991,

Carey and others 1997). Previous research shows that both large snags and trees

with natural cavities are much less abundant in young, managed stands than in old

growth (Carey 1995, Carey and others 1991a). Numerous studies of cavity-using

squirrels outside the Pacific Northwest have shown that addition of supplemental dens

(usually nest boxes) to managed stands can increase squirrel populations (see Carey

and Sanderson 1981 for a review). Thus, adding supplemental dens to managed

stands allows testing of the hypothesis that dens are limiting to flying squirrels. If

providing dens can raise flying squirrel populations, the potential exists for attracting

owls to managed stands, thus increasing available owl foraging habitat.

Supplemental dens can be provided in two forms: nestboxes and cavities created in

trees (Carey and Gill 1983). Nestboxes increase the number of flying squirrels in other

areas (see Carey and Sanderson 1981 for a review) but have the disadvantages of

being relatively susceptible to decay and damage and, to some observers, appearing

unnatural. Nestboxes generally benefit only a few species besides target species.

Cavities created in live trees have the advantage of potentially being available for

many years and of introducing top rot that will eventually provide a condition suitable

for excavation by woodpeckers and subsequently be used by a great variety of cavityusing birds and mammals. Because nestboxes increase flying squirrel abundance

whereas excavated cavities, as designed for this study, have not, both forms were

used in this study. High flying squirrel densities are those > 3/ha (Carey 1995, Carey

and others 1992), with flying squirrels using multiple dens and dens being used by

several flying squirrels simultaneously (Carey and others 1997). Thus, we initially

added sufficient cavities and nestboxes so that with natural dens we would ensure that

dens were not a limiting factor for squirrel populations.

Cavities and nestboxes were systematically and alternately placed over the treatment

areas at a density of 3/ha. They were placed in half of the experimentally thinned

stands and half of the unthinned stands (fig. 4). Nestbox and cavity designs are

shown in Carey and Gill (1983), in a video (Carey 1993), and in appendix 2. Sizes of

boxes and cavities were chosen to be the minimum that could contain three or more

flying squirrels.16 Interior dimensions (below the shelf) of Flyger nestboxes (Carey and

Gill 1983) were 6 by 7 by 7 inches (height, width, depth) and 7 by 7 by 6 inches for the

cavities. Cavity depth was a minimum of 6 inches but often extended to 10 inches if

the tree was sufficiently large. The nestboxes are more predator-proof and more

thermally beneficial (because of the shelf) than the created cavities. If nestboxes are

used more than created cavities, cavities may be modified by adding a shelf below the

entrance as was done for nestboxes. Because addition of a shelf to a cavity reduces

interior volume, the cavity might need to be deepened or lengthened. We placed

boxes and cavities 6 to 7 m aboveground for ease of inspection. Application in a

management scenario would call for placing cavities as high as practical with maximum

height being limited by tree diameter; trees should be at least 30 cm in diameter at

the cavity site. Increasing the height of cavities aboveground is desirable not only

because of increased attractiveness to flying squirrels and reduced access to predators

16 Up to five flying squirrels have used the boxes simultaneously.
20

but also because of timber values of cavity trees destined for harvest. Cavities will

induce top rot, but trees should compartmentalize the rot and one or two 16-foot logs

could be kept clear of rot if the cavity is placed high enough.

Reconnaissance showed insufficient tree regeneration and insufficient seed sources to

provide a midstory of shade-tolerant trees to fill gaps between lower crowns of dominant and codominant trees and upper layers of the crowns of tall shrubs. Accordingly,

we incorporated underplanting for species and architectural diversity into our experimental design. Because part of the underplanting of seedlings was to go into

Phellinus weirii infection centers, we included species immune to, highly resistant to,

or tolerant of root rot (red alder, western redcedar, and white pine, respectively) and

grand fir because of its shade tolerance, seed production, and retention of a dense

crown (to provide architectural diversity) (Burns and Honkala 1990a, 1990b; Thies and

Sturrock 1995).17 We limited our choice to species that commonly occur or were

17 Western hemlock regeneration was found infrequently in both

forests. Though western hemlock occurs in other areas with similar

glacial soils, the combination of excessively permeable soils (and

concurrent low water storage capacity) and low summer rainfall

found at Fort Lewis creates substantially greater annual soil

moisture deficit than in other areas where western hemlock

dominates (Packee 1976). Fort Lewis is also an area of historically

short fire-return intervals, and the thin-barked western hemlock is

very susceptible to fire (Burns and Honkala 1990a). Because

western hemlock also is frequently infected with Phellinus weirii,

especially where Douglas-fir is or was a major component (Burns

and Honkala 1990a), we chose not to underplant western hemlock.

Grand fir was chosen as an ecological (shade-tolerant) equivalent

to western hemlock; grand fir is common in the Puget Trough and it

was thought that it would grow better in such a dry environment.

Underplanting
21

historically common in west-side forests. We chose to plant only root rot thin and

heavy thin cells to retain the desired open understory conditions in the remaining light

thin cells.

These four tree species will enhance the tree diversity of the study site and in the long

term will begin to fill the intermediate canopy spaces between the tall shrubs and the

base of the overstory canopy. Natural regeneration of Douglas-fir also will occur, but

its growth in the understory probably will be limited (Oliver 1995). The production of

abundant Douglas-fir cone crops is highly variable, and thus the mix of underplanted

species will in the long term provide a more diverse and more reliable source of seed

for small mammals and birds (Burns and Honkala 1990a, 1990b; USDA Forest Service

1974).

Flying squirrels, other arboreal mammals, and birds make substantial use of woodpecker holes for nest sites. Our work has shown natural nest sites (cavities and

others) to be of limited availability in many managed Douglas-fir forests, including

those of the study site (Carey 1995, Carey and others 1997). Artificial inoculation of

decay fungi in managed stands has been successful in inducing decay (Conner and

others 1983, Parks and others 1995); fungal decay is a prerequisite for excavation by

many primary cavity nesters such as woodpeckers (Neitro and others 1985). To

hasten development of trees and snags with acceptable decay conditions for woodpecker use, we incorporated tree inoculation with stem-decaying fungi into our experimental design.18 We isolated two commonly occurring wind-dispersed fungi, Fomitopsis cajanderi and Phellinus pini, from the study site; Fomitopsis is generally found

at greater heights in the stem than is Phellinus. Both are widespread, naturally

occurring, and weakly pathogenic fungi. Isolates were obtained from decaying Douglasfir trees, grown in pure culture, and inoculated into sterilized softwood dowels to

facilitate later placement in trees.

Because response variables will be related to specific forest conditions as they exist

after experimental thinning, an evaluation was performed to describe actual, postthinning variability at the stand and grid-cell level. Specific goals of the assessment

were:

1. To estimate prethinning and current stand stocking densities19 and basal area by

sampling each thinning subtreatment in each stand

2. To estimate logging damage and stand health

3. To estimate incidence of root grafting

4. To estimate stocking density of underplanted seedlings and natural regeneration

Top Rot Fungi

Inoculation

Postthinning Stand

Assessment

18 The FES inoculations are part of a much larger and more

widespread series of field trials throughout different forest

conditions in the Western United States. Background, methods,

and locations of the larger study are available from: Catherine

Parks, Pacific Northwest Research Station, 1401 Gekeler Lane,

La Grande, OR 97850.

19 The terms “stocking density,” “stocking,” and “tree density” are

used interchangeably and refer to number of trees per hectare

(N/ha).
22

5. To provide a database of estimates of forest parameters (e.g., retained basal area,

retained relative density, current density) for each grid cell within thinned stands

6. To reclassify thinning subtreatments based on retained relative density and to

provide estimates of areas occupied by each category in each thinned stand

7. To provide forest parameter maps for each of the thinned stands

8. To provide the basis for designing the 2003 thinning

A schedule of the major activities, treatments, and sampling efforts from the inception

of the Forest Ecosystem Study in 1991 until the end of 1996 is shown in table 1.

After on-the-ground reconnaissance identified sufficiently large areas of homogeneous

vegetation to contain the study, the four stands per block were located with a minimum

of 80 m between the stands. Blocks and stands were given numerical designations:

the Star stands were designated 101-104; Stellar, 201-204; Farley, 301-304; and Hill,

401-404. Maps of the four blocks (Farley, Hill, Star, and Stellar) showing the arrangement of treatments and the grid coordinate designations are shown in figures 1 and 2.

Grids were surveyed and established in early fall 1991; each grid consisted of 64

stations aligned in an 8 by 8 matrix with 40-m spacing between stations. Each station

was identified by a letter (A through H, west to east) and a number (1 through 8, north

to south). For example, the northwestmost station of every grid was always “A1.” The

grids were 280- by 280-m squares, with each station point (= grid cell corner) marked

with a 1-m length of white plastic PVC pipe (1-3/4 in OD, Schedule 40, inserted 66 cm

in the ground). The aboveground portion of the stake was painted fluorescent orange.

Each station also was marked with two strips of orange fluorescent flagging, and the grid

station was visibly marked at the base of both strips with permanent ink. In addition to the

64 station points, the north-south 20-m points between stations were identified by a pin

flag inserted into the ground and with a single strip of fluorescent orange flagging. The

resulting forty-nine 40- by 40-m grid cells in each surveyed stand were the operational

units to which the thinning subtreatments were randomly applied (fig. 3).

The 8 by 8 matrices were used as a template for live trapping grids (LTG), which

determined the placement of arboreal rodent traps (fig. 5). In 1991, two Tomahawk20

201 live traps were placed at each of 64 grid stations in each of the 16 stands (2,048

traps total)—one trap was placed on the ground and one was attached to a tree as

outlined in Carey and others (1991a). A 4-in-square aluminum tag with the grid and

station number etched on it was nailed to each trap tree. Traps were placed in the

grids in early fall 1991; trap locations were identified by stand (i.e., 201, 303, 404) and

grid station (i.e., A1, B3, G7). Trapping of all grids has taken place twice yearly (spring

and fall) since 1991.

Implementation

Grid Establishment

Arboreal Rodent Trap

Placement

20 The use of trade or firm names in this publication is for reader

information only and does not imply endorsement by the U.S.

Department of Agriculture of any product or service.
23

In 1992, 10 by 10 small mammal grids (SMG) with 20-m spacing between stations

were established within the center of each of the eight stands designated as not-to-bethinned (e.g., control and den-augmentation only). Each SMG was located within the

LTG, with the northwestmost SMG station being identical with the LTG point B-2, and

the SMG extending grid-east and -south for 200 m from that point. Because of the

20-m spacing, every second SMG station coincided with an arboreal rodent trapping

station. To avoid confusion with the arboreal rodent trapping station designations, a

system of identification was used for the SMG stations whereby each station was

identified by a letter (Q through Z, west to east) and a number (1 through 10, north to

south). Because many of the grid points were both an arboreal mammal trap station

and an SMG station, the station number used for recording data was based on the

type of trapping being performed at the time (fig. 5).

Two Sherman live traps were positioned on the ground within 2 m of each SMG

station; one large 3- by 3.5- by 9-in trap, and a smaller 2- by 2.5- by 6.5-in trap. Trap

placement was prioritized by proximity to large, fallen, decayed logs and other coarse

woody debris.

Trapping was performed at the eight unthinned SMGs in 1992, 1993, and 1994. In

summer 1996, SMGs were placed in the eight experimentally thinned grids as well

(fig. 5). Trapping of all 16 SMGs began in July 1996.

In April 1992, 16 nestboxes and 16 cavities (illustrated in appendix 2) were systematically distributed at 80-m intervals throughout eight grids. A cavity or nestbox was

placed on the largest tree (> 30 cm d.b.h.) within 5 m of the selected gridpoint, or on

the nearest tree > 30 cm d.b.h. if no large tree could be found within 5 m. Beginning

with the A1 station, alternating cavities and nestboxes were placed at every other grid

station on the north-south lines (fig. 4). Construction of nestbox and cavity faceplates,

installation of cavities, and placement of nestboxes were performed by contractors.

Nestboxes were secured with aluminum nails at a height of 6 m. Cavities were

created at this same height with the use of a chainsaw equipped with a special tip.

The faceplate was then nailed over the opening of each cavity.

Because most supplemental nestboxes showed signs of use by spring 1995, an

additional eight nestboxes per den-augmented stand (64 boxes total) were added in

December 1995. The additional eight nestboxes were placed 12 m above the ground

rather than at the original 6-m height. They were placed at the C4, C6, D3, D5, E4,

E6, F3, and F5 grid stations on den-augmented stands (fig. 4). At that time, modifications were made to the cavity size and the faceplate door placement (appendix 2).

Experimentally thinned stands (thin, thin plus den-augmentation) were cruised during

tree marking. We employed cut-tree marking; only trees > 20 cm d.b.h. were marked

for harvest. Thinning operations took place between January and April 1993 with

tractor skidders. Net harvested volumes from SS stands ranged from 104 to 154 MBF

per stand; FH net volumes ranged from 192 to 222 MBF per stand. For Star and

Stellar, the harvested volumes ranged from 5.4 to 8.1 MBF per acre; Farley and Hill

harvested volumes ranged from 8.7 to 11.1 MBF per acre.

Small Mammal

Trapping Grid and

Trap Placement

Cavity and Nestbox

Placement

Silvicultural Treatment
24

The locations of skid trails and landings were planned to minimize disturbance in

thinned stands. In Star and Stellar blocks, skid trails from the 1937 harvest were used

to the greatest extent possible; otherwise, new skid trails were located in the heavy

thins, root rot thins, and along the margins of Phellinus infection centers. In Farley and

Hill blocks, skid trails and landings from previous thinning entries were used.

We employed one of the standard slash disposal treatments that the Fort Lewis

Forestry Division uses during their thinning operations. After trees were felled, they

were pulled to the landing with crowns intact. At the landing, trees were bucked into

logs and their branches trimmed; the accumulated debris was piled and burned after

harvest. By doing so, a relatively constant amount of coarse woody debris (CWD) was

maintained in the treated stands, with the net intended result being that no additional

CWD would accumulate from the thinning operations.

Western redcedar, western white pine, and grand fir were planted at equal densities

within each root rot and heavy thin subtreatment cell. Planting took place in May and

June 1994. Conifer seedlings were planted on a 9 by 9 matrix with 3.7-m spacing, a

density equal to about 510 trees/ha.21 Three planters, each with a different species,

planted three sets of three parallel rows. Thus, three lines of nine seedlings (27 total

per subtreatment cell) were planted for each conifer species (for a total of 81 conifer

seedlings per cell). The 20 red alder seedlings were planted every 11 m, between the

lines created by the planted conifers. Because substantial natural alder regeneration

was expected, red alder was planted only in the RT cells, and was planted at lower

densities than the conifers—20 red alder seedlings per grid cell vs. 27 per conifer

species per grid cell. Planting guidelines for all species were as follows:

• To avoid the relatively higher shade along the edge of the treatment cells, planting

lines began about 7 m from the treatment boundary. This produced a small buffer

around the planted seedlings and afforded space to plant seedlings not planted on a

line because of their planting location falling within 2.5 m of an existing overstory

tree.

• If a planting site fell within 2.5 m of an existing tree, the seedling was placed at the

end of that planting row. As many as two additional seedlings could be planted at

the end of any planting row. If three planting sites were displaced by existing trees,

the third seedling could be planted 3.7 m in from the treatment boundary on the next

line. The 7-m buffer around seedlings allowed space for planting trees that were

displaced by existing trees. Planting at the end or beginning of a planting line

allowed the planting of an equal number of seedlings in each planted grid cell.

Fungal inoculum was placed above cavities in selected Douglas-fir cavity trees. The

insertion points were holes drilled to a depth of 10 to 15 cm. The holes were cleaned

of chips and an inoculated dowel placed within. A PVC pipe was then inserted in the

hole at a horizontal or slightly downturned angle to prevent water entry and to prevent

closure of the hole. Five to eight cm of pipe was left exposed from the side of the tree

Underplanting

21 These figures refer to planting densities at the grid cell level. This

resulted in a stand level planting density of about 210 conifer

seedlings/ha.

Top Rot Inoculation
25

to mark the location of the inoculum. Tree d.b.h., aspect of inoculum hole, and any

tree defects (broken, forked, dead tops) were noted, as were color and texture of chips

removed from the drill hole (to assess presence of preexisting decay).

A systematic subset of six of the artificial cavity trees in each of the eight den-augmented stands was selected for inoculation. Inoculation was performed in summer

1994. Locations of the inoculation holes and specific treatments of the inoculated

trees were as follows (fig. 4):22

Grid points B4 and G5—Phellinus pini at 25 cm above created cavities

Grid points C5 and F4—Phellinus pini at 25 cm above and Fomitopsis cajanderi at

3 m above created cavities

Grid point D6—Phellinus pini at 3 m above created cavities

Grid point E3—Fomitopsis cajanderi at 3 m above created cavities

Thus, each den-augmented stand contained six inoculated trees with a total of eight

inoculation holes. Evaluation of the success of fungal inoculation will be made by

observing bird excavation and use at or near the inoculation site and sampling to

attempt to reisolate the original fungal strains.

Pretreatment tree and understory woody plant abundance—Fifteen interior

gridpoints, a minimum of 56 m apart in each of the 16 stands (240 points total), were

selected systematically as reference points for measurement of trees and understory

woody plants before the thinning treatments. Alternating N-S grid lines had two or

three sampling plots, respectively. Tree sampling was performed in fall and winter

1991. Understory vegetation was sampled in June-August 1992.

At each selected grid point, a variable-radius prism plot count (basal area factor 30

[English]) was made. Species and d.b.h. were determined for each tallied tree as well

as whether the tree originated with the current or previous stand. For the two dominant trees (site trees) nearest to plot center, species, d.b.h., height, height to live

crown, crown width, and current-year horizontal and terminal growth were recorded.

Crown width was determined by measuring the extent of the crown or dripline of the

foliage for each measured tree. Estimates of lateral growth were made on lower or

mid-crown branches; an average of the past 3 years of growth was recorded.

Woody shrub and other understory vegetation cover were sampled on 2.82-m-radius

circular plots (25-m2

), offset 5 m at a randomly determined bearing from the grid point.

We recorded (1) lifeform of plant species (e.g., evergreen shrub; deciduous shrub; fern

or forb; grass, sedge, or rush; moss; or lichen); (2) vascular plant species; (3) percentage of cover by species; and (4) mean height of shrub species. In addition, moss and

lichen species and moss cover were noted. We defined “cover” as that portion of the

ground covered by the vertical projection onto the ground of the vegetation of interest.

The incidence and areal extent of root rot pockets (particulary laminated root rot) was

mapped by using grid lines as line-intercepts. When root rot pockets were intersected,

length and width of the disease pocket were recorded.

Sampling Methods

Vegetation

22 The exact positions of the inocula and descriptions of the host

trees are on file with: Pacific Northwest Research Station, Forestry

Sciences Laboratory, 3625 93d Ave SW, Olympia, WA 98512.
26

Posttreatment stand conditions—We used a stratified random sample to assess the

silvicultural prescription. Initially, six grid cells per target subtreatment per stand were

randomly selected and sampled with fixed-radius plots. Within each thinned stand

were the three target subtreatments:

1. RT cells and Phellinus weirii pockets with underplanting

2. HT cells with underplanting

3. LT cells without underplanting

After the initial fixed-plot sampling, we sampled every additional grid cell with variableradius plots. Sample plots were located in the center of grid cells to prevent overlap

with adjacent cells and to minimize edge effects. Fixed-plot radii ranged from 4 to 15 m,

according to the variable being measured (tree, stump, or seedling density) and the

subtreatment.

In each fixed-radius plot, we counted standing trees (live and dead) by species,

measured d.b.h. (cm) outside the bark of all trees > 10 cm d.b.h., and ranked tree

health by using a modification of the Pacific Northwest Region stand exam procedure

(USDA Forest Service 1989). For stumps, we measured diameter outside bark (cm),

height (m), and presence or absence of callus formation (presence of callus was used

as an indicator of root grafting). Stump diameters were converted to tree d.b.h. after

forest-specific regression equations were developed. The equations are as follows:

Star/Stellar: Y = 0.81X + 1.43 (r2

 = 0.94) (1)

Farley/Hill: Y = 0.88X + 0.99 (r2 = 0.96) (2)

where Y = d.b.h. in cm, and

X = stump diameter at 30-cm height.

We measured density of the three underplanted conifer species and density of natural

tree regeneration by species and height class (table 4).

Table 4—Height class categories and descriptions for natural tree regeneration

Height class Approximate age Description

Centimeters Years

0-10 1 or less Emergent seedling

11-50 1-2 Established seedling

51-100 ≥ 2 Well-established seedling,

emerging above ground

vegetation

> 100 5 or more Well-established sapling,

(and < 10 cm d.b.h.) with most of crown above

ground vegetation
27

In the variable-radius plots, trees were counted with a wedge prism from the center of

each cell. A basal-area-factor (BAF) 20 (English) prism was used in SS, and BAF 30

(English) in FH. Regression equations calculated from fixed-radius plots were used to

estimate tree density from basal area:

Star/Stellar: Y = 9.23 X + 19.06 (r2 = 0.74) (3)

Farley/Hill: Y = 3.70 X + 11.41 (r2 = 0.73) (4)

where Y = residual tree density (N/ha), and

X = residual basal area (m2/ha).

Data collected from both fixed- and variable-radius plots reflected current (1995-96)

tree diameters for live trees and prethin (1992) diameters for stumps. Due to tree

growth during that 3- to 4-year period, current live tree density of trees > 20-cm d.b.h.

was greater than that of 1992 (assuming minimal tree mortality), thus resulting in a

slight overestimation of the ratio of current tree density to target tree density. Because

no growth and yield data were available for these sites, the current assessment made

no correction for growth and mortality during this period.

Posttreatment vegetation permanent plots—After the experimental thinnings,

permanent vegetation plots were established in the centers of 191 randomly selected

grid cells (fig. 6; appendix 3). During summer 1994, six permanent plots were established in each target subtreatment in each stand23 and in each control and denaugmentation-only stand by using a stratified random sampling design. In each stand

of 49 cells, 100 percent of the root rot thin cells were sampled; about 50 percent of the

heavy thin cells were sampled; and about 25 percent of the light thin cells were sampled.

About 12 percent (6 of 49) of the cells in control stands and den-augmentation-only

stands24 were sampled. Twelve permanent plots were established in each target

subtreatment in each block and 12 total in the unthinned stands in each block.

Plots were marked with 1-m lengths of 1-3/4-in diameter, schedule 40, white PVC pipe

inserted 66 cm into the ground at plot center. At each plot we measured slope and

aspect. In August 1995, each plot was photodocumented with a 24-mm wide-angle

lens on a 35-mm camera (84-degree field of view) from a point 1.5 m high and 8.4 m

south of the plot center (thus providing full lateral coverage of the 5.6-m-radius plot). A

2-m-tall photo board with alternating 0.25-m white and black sections was placed at

plot center for reference. Plots will be rephotographed every 3 to 5 years.

23 The RT subtreatment in stand 103 contained only five cells of

sufficient size to contain permanent plots, resulting in 191 rather

than 192 permanent plots.

24 Because no data are presented for experimental responses to

the den-augmentation-only treatment, we also refer to them as

control stands unless it is necessary to differentiate between the

two unthinned stands (control and den-augmentation-only) per

block.
28

Vegetation was sampled by using nested circular plots of 5.6-m radius (100 m2)

 and

12.6-m radius (500 m2). Understory composition, cover, and forest floor cover were

measured on 5.6-m-radius plots centered on the PVC pipe. For cover values, we used

octave scale categories modified from Gauch (1982).25 Within 5.6 m, we described the

forest floor cover of bare ground, moss, fine litter (needles, leaves, twigs < 2 cm

diameter), coarse litter (2 to 10 cm diameter), fine woody debris (10 to 30 cm diameter), coarse woody debris > 30 cm diameter, and all vascular plant species. Computer-generated maps depicting a range of cover values were used to train observers;

training also served to ensure consistency among observers in regard to species

identifications and cover estimates. Identification of all vascular plant species (including grasses, sedges, and rushes) was facilitated because of involvement by the

authors in a previous project that produced a vouchered floristic inventory of the Fort

Lewis Military Reservation (Thomas and others 1994). Because of this earlier effort,

vouchers from the FES were collected only for previously uncollected species.

Vouchers from the current project will be deposited in the University of Washington

Herbarium, Seattle.

Overstory composition, cover, and structure, and understory vegetation groups were

described on 12.6-m-radius plots also centered on the PVC pipe at plot center. We

described cover of forbs and ferns < 0.5 m tall, low shrubs 0.5-2 m, midstory shrubs

and understory trees > 2 m, and overstory trees. We counted live and dead trees

(> 2 m tall) by species and by size class within 12.6 m. Size classes were 5 to 20 cm

d.b.h., >20 to 40 cm d.b.h., >40 to 60 cm d.b.h., and >60 cm d.b.h. Within the 12.6-mradius plots, we used a four-category subset of the octave scale (where 1 = 0 to 12

percent; 2 = > 12 to 24 percent; 3 = > 24 to 48 percent; 4 = > 48 to 100 percent) to

estimate the percentage of volume filled by forbs, ferns, low shrubs, midstory shrubs,

and trees in cylinders 0 to 0.5 m, 0.5 to 2 m, and 2 to 10 m in height.

Vegetation plots were established and variables first measured during summer 1994,

16 to 18 months after completion of the experimental thinnings. Permanent plots

were revisited in summer 1996 and will be sampled every 3 to 5 years thereafter for

the duration of the study.

Tree age and growth response—The largest tree26 within 5 m of each grid point was

described: tree age, height, crown width, crown ratio, d.b.h., and species were recorded. In 1993 and 1994, all 64 grid-point trees in each of the 16 grids (1024 total;

985 live trees) were cored with an increment borer at breast height. Tree height and

height to live crown were measured with a tree laser (Laser Technology Inc., Criterion

400). Tree increments were used to assess the age distribution of the dominant trees

in the stands. These dominant and codominant trees were used to characterize the

site class of each forest and stand and will be used to assess growth of the retained

overstory trees during the life of the experiment.

25 The following scale was used for cover values: 1 = 0-1 percent

cover; 2 = >1-3 percent; 4 = >3-6 percent; 8 = >6-12 percent; 16 =

>12-24 percent; 32 = >24-48 percent; 64 = >48-100 percent.

26 These are the same trees on which arboreal mammal traps were

placed.
29

Supplemental nestboxes and cavities were checked for use twice per year beginning

in winter 1993. Arboreal mammals were captured and processed as per Carey and

others (1991a). Animals were handled minimally and immediately placed back in the

box when all data had been obtained. In addition, nest materials, food items, or den

use by nontarget animal species were noted. Summer den checks were conducted to

roughly coincide with the presence of northern flying squirrel young of the year. If

young were found that did not have fur developed, they were noted, but left untouched.

Pretreatment epigeous fungi—Epigeous fungi were initially sampled during fall 1992

(November 8-24) to assess effects of previous forest management practices on

epigeous fungal diversity and to establish preliminary fungal biodiversity data. No

attempt was made at that time to assess fungal biomass production. In each of the

eight stands designated as not-to-be-thinned, six 4-m2 plots were selected for fungal

sampling. Plots were 5 m northwest of randomly selected grid station points. Within

sample plots, all exposed fungi were removed, tagged, and placed in wax paper bags

for transport to the laboratory, where specimens were identified.

Posttreatment epigeous fungi—A randomly selected subset (n = 128) of the 191

permanent plots was sampled yearly to evaluate effects of experimental thinning on

species composition and biomass production of sporocarps of epigeous fungi. All

subtreatments and nonthinned stands were sampled with equal intensity (four plots

per subtreatment per stand and four plots per unthinned stand per session). Species

composition and biomass of all macrofungi (> 1 cm diameter) were recorded on the

100-m2 plots during the peak of the fall fungal seasons in 1994, 1995, and 1996. We

identified fungi in the field or collected specimens of all unknown taxa for later identification; identification was with fresh specimens where possible. Voucher specimens

were kept for most species; characteristics of fresh sporocarps, including color, odor,

staining, etc., were noted, as was the substrate from which they grew. Voucher

specimens will be deposited at the Oregon State University Herbarium, Corvallis.

Epigeous fungal sporocarp biomass production was estimated by developing fungal

cap diameter to dry weight regression equations for groups of fungi (e.g., groups of

similar life form fungi such as gilled, stalked mushrooms, Russulas only, or chanterelles).

In this way fungal biomass production was estimated without repeated destructive

sampling. We anticipate this study will continue yearly. When possible, a second

yearly sampling will take place during spring; however, pilot studies have indicated

that epigeous sporocarp production during spring is minimal compared to fall.

Identification of many species of epigeous fungi to the species level is difficult, if not

impossible, especially for complex genera such as Cortinarius, Inocybe, and Russula.

We report trends in diversity of fungal taxa; we used genus and species names from

regional guides, including Arora (1986), Phillips (1991), and Tylutki (1979, 1987).

Because of the uncertainty surrounding species concepts in fungi27 and the preliminary

Den Use

Fungi and Soil

Food Webs

27 For those fungal taxa listed in table C-3 of USDA and USDI

(1994) (e.g., “survey and manage” species), much of the taxonomic

and nomenclatural uncertainty has recently been clarified by

Castellano and O’Dell (1997). For other taxa, questions of

synonomy and nomenclature will require further investigation and

will be reported in the future.
30

nature of our report, we do not purport to present all synonyms for, or the complete

nomenclatural history of, all epigeous fungal taxa included in this report. In general,

fungal authorities for epigeous fungi follow Phillips (1991) and Tylutki (1979, 1987).

Additional authorities and some commonly encountered synonyms were provided by

Dr. J.M. Trappe of Oregon State University. For fungal taxa found in table C-3 of

USDA and USDI (1994), nomenclature follows Castellano and O’Dell (1997). The list

in appendix 1 is a preliminary summary of epigeous fungal diversity on the study sites.

Future research will clarify many species concepts and nomenclatural questions; if

possible, we will amend our findings as these results are published.

We distinguished between two groups of epigeous fungi: saprobic and mycorrhizal.28

We assigned mycorrhizal status to genera for which mycorrhizal potential has been

reported in, Miller (1983), Singer (1986), and Trappe (1962). We classified as mycorrhizal all species of the genera Amanita, Boletus, Cantharellus, Clavaria, Cortinarius,

Dentinum, Entoloma sensu stricto, Gomphidius, Hebeloma, Hygrophorus, Inocybe,

Laccaria, Lactarius, Lycoperdon, Russula, Suillus, Thelephora, and Tricholoma. We

report average ectomycorrhizal species richness per plot and total ectomycorrhizal

species richness for experimentally thinned and unthinned sites in SS and FH for the

fall sampling sessions of 1994-96.

Hypogeous fungi (truffles)—

Truffles (hypogeous sporocarps

of the fungal subdivisions

Zygomycotina, Ascomycotina,

and Basidiomycotina) were

sampled every 5 to 7 weeks from

April 1993 (immediately after the

thinnings) until December 1995.

Truffles were collected from

each of 10 circular 4-m2 plots

systematically located 10 m

apart along randomly placed

transects in each thinning

subtreatment and in nonthinned

stands (160 truffle plots per

sampling period). Stands with and stands without supplemental dens were sampled in

alternating sessions. At each truffle plot, vegetation was removed, plots were raked to

28 Mycorrhizal fungal types can be described in relation to two axes

or gradients: from lack of penetration of root cortical cells to

penetration, and from root enclosure by hyphae to nonenclosure

(Allen 1991). Root enclosure by fungal hyphae and lack of root

cortical cell penetration characterize the ectomycorrhizae, and

nonenclosure and cortical cell penetration characterize the

endomycorrhizae, of which the vesicular-arbuscular (VA)

mycorrhizae are a common form. The family Pinaceae, and

Douglas-fir in particular, are characterized by a dominance of

ectomycorrhizal fungi—the hypogeous truffles and epigeous

mushrooms. Where we refer to mycorrhizal fungi, we imply

ectomycorrhizal fungi. Otherwise, we refer to endomycorrhizal or

VA fungi where we need to distinguish between them and

ectomycorrhizae.

© Wes Colgan III. Reprinted with permission.
31

mineral soil depth (about 5 cm), and all fungal sporocarps were collected, identified to

species, and dried for biomass determinations. Truffle species identifications were

provided by Dr. J.M. Trappe of Oregon State University. Voucher specimens for all

hypogeous species were deposited at the Oregon State University Herbarium,

Corvallis.

Soil food webs—Soil food web samples were taken in spring and fall 1994 and 1995

from a subset (n = 100) of 4-m2 plots offset from permanent plot centers (to avoid

disturbance of permanent plots). The percentage of cover of recognizable ectomycorrhizal mat types (Piloderma sp., Hysterangium sp., Gautieria sp.) was measured

and mapped, and about 100 g of soil was collected from each plot. Soil was placed in

plastic bags and stored in an insulated cooler during travel. Samples were then stored

at 4 °C and microbial activity analyzed within 5 days of collection. The following

parameters were evaluated: (1) grams dry weight (gdw) per gram of soil; (2) active

and total bacterial and fungal biomass (μg/gdw); (3) total nematodes (no./ gdw); and

(4) percent fungal-feeding, percent bacterial-feeding, percent plant-feeding, and

percent predatory nematodes. Active and total bacterial and fungal biomass (AB, AF,

TB, TF) were determined as described by Babiuk and Paul (1970) and Ingham and

Klein (1984) and were used to compute the ratios, TF/TB and AF/AB. Nematodes

were extracted as described by Anderson and others (1978) and were separated into

trophic groups based on morphological characteristics used in taxonomic classification

(Bongers 1989).

Pretreatment coarse woody debris and snags—Grid lines in each of the 16 stands

were used as a template to sample CWD cover by using a line-intercept method (Van

Wagner 1968, Van Wagner and Wilson 1976). Snags were sampled in every other

grid cell in all stands. For sampling fallen

trees, 4 to 6 interior lines in each stand

(lines B through G, rows 2 through 7) were

sampled, with a total of 800 to 1200 m of

line sampled per stand. Fallen trees were

classified by origin (originating with the

previous or current stand), size class (trees

with a minimum large end diameter of 15 cm

were recorded), and decay class. For

fallen trees greater than 15 cm d.b.h., the

length of transect covered by the fallen tree

was recorded—if the line intercepted the

log on a diagonal, the diagonal measurement was recorded. If numerous logs were

piled on each other, only the top log was

recorded. For large fallen trees (greater

than 30 cm diameter), the following information was recorded: species, diameter in

centimeters at each end, length (meters),

decay class, and origin (originating from

the previous or current stand). For large

Coarse Woody Debris,

Snags, and Potential

Dens
32

fallen trees with roots still attached, the large-end diameter was recorded at a height

above the base equal to breast height. We used a three-decay-class system for fallen

logs, modified from the five-decay-class classification system developed by Fogel and

others (1973) and modified by Sollins (1982). Decay classes 1 and 2 of Fogel and

others (1973) were treated as class I; classes 3 and 4 were treated as class II; and

decay class 5 was treated as class III.

All snags greater than 15 cm d.b.h. and taller than 1.4 m were inventoried in alternating grid cells in each stand. All standing trees 15 to 30 cm d.b.h. were tallied by using

a simple dot-count method. Detailed measurements were recorded for standing dead

trees greater than 30 cm diameter. For larger snags, we recorded species, d.b.h.,

height, and decay class from the five-class system of Cline and others (1980).

Postthinning coarse woody debris—Grid lines used in the prethinning CWD survey

were resampled after thinning to assess effects of thinning operations on CWD cover

and distribution. The three-decay-class classification system was used, and the same

size of CWD was sampled as during the prethinning sampling. Grid lines were assigned one of six designations, depending on which type of thinning subtreatments the

lines separated. The six categories were (1) light thin-light thin; (2) light thin-heavy

thin; (3) light thin-root rot thin; (4) heavy thin-heavy thin; (5) heavy thin-root rot thin;

and (6) root rot thin-root rot thin.

Den availability—After completion of the experimental thinnings, we inventoried 100

percent of all 16 stands (plus a surrounding 20-m buffer) for residual snags, residual

stumps, residual live trees, conifers with sticknests, conifer snags with cavities, conifer

snags without cavities, deciduous snags with cavities, and deciduous snags without

cavities. Residual stumps were those with sapwood in an advanced state of decay

and were ≥ 1 m high and ≥ 25 cm diameter. Snags were ≥ 2 m high, ≥ 25 cm diameter, and characterized as residual, current, or deciduous. Stands were sampled in

summer 1993.

Arboreal rodents—Arboreal rodents were trapped on all 16 grids twice yearly (fall

1991, and spring and fall 1992-96). We trapped for two consecutive weeks in each

session, opening and baiting traps on Monday morning, checking traps each of the

following four mornings (Tuesday-Friday), and closing traps on Fridays. Arboreal

rodent trapping data will be reported here only for the baseline period (fall 1991 and

spring and fall 1992). We summed captures and counted individuals for the three

arboreal species (flying squirrel, Townsend’s chipmunk, and Douglas’ squirrel) for

1991-92. Chipmunks were not sampled in 1991. We estimated mean densities of

flying squirrels (three seasons) and chipmunks (two seasons) per stand (Carey and

others 1991a), combining the Chapman modification of the Lincoln-Peterson Index

with grouping captures by week and estimating area sampled with site-specific mean

maximum distances moved. Forest means were calculated from stand means. All

reported data are from prethinning sampling and do not consider the effects of supplemental dens.

Forest Floor and

Arboreal Mammals
33

During trapping, flying squirrels were radio-collared to

document movement patterns

and den use; telemetry and

den site data will be reported

elsewhere. Den site characteristics and abundance are

reported in Carey and others

(1997). Concurrent with flying

squirrel trapping, extensive

descriptive notes were taken

on captured animals and were

used to assess flying squirrel

age and reproductive status

(Villa and others 1999) as well

as foraging activity and use of

nontruffle foods (Thysell and others 1997). During trapping, fecal pellets were collected

and placed in ethanol and later were analyzed for dietary components. The protocol for

trapping is outlined in Carey and others (1991a); the protocol for fecal pellet analysis

follows McIntire and Carey (1989) and Colgan and others (1997).

Forest floor small mammals—Forest floor small mammals were trapped for 2-week

sessions during summers of 1992, 1993, and 1994 in the eight randomly selected,

not-to-be-thinned stands. In summer 1996 and 1997, all 16 stands (thinned and

unthinned) were trapped for forest floor small mammals. We report small mammal

trapping data from 1992 and 1993 only. Trapping effort was 6,400 trap nights in each

forest (FH and SS) in each reported year. Small mammal abundance was calculated

as captures per 100 corrected trap nights (CPUE) (Nelson and Clarke 1973). Abundance was calculated for each of the four trapped stands per forest per year; we report

the mean CPUE ± SE for each forest in each year based on these four stand values.

For both arboreal and forest floor mammals, we made extensive use of necropsies to

evaluate field identification of species, sex, age, and reproductive condition. Additionally, species identifications were confirmed by the Burke Museum, University of

Washington, and voucher specimens were deposited in that museum.

Owl survey—In spring 1992, we conducted an owl survey to establish baseline data

on owl diversity and abundance. In each forest (FH and SS) we surveyed for owls in

an area extending about 0.5 mile out from the experimental stands. We established a

network of calling stations within each forest—stations that allowed an appreciable

amount of audible overlap between adjacent stations when hooting conditions were

fair or better. Calling stations were visited three or four times during April and May

1992. At these times, we broadcast calls for the great-horned owl, barred owl, northern spotted owl, barn owl, northern pygmy owl, saw-whet owl, and screech owl.

Since the spring 1992 formal owl survey, we also have noted the presence of owls on

the study site by recording the observations of night telemetry personnel. We include

a list of owl and other bird species observed on the study site (appendix 1).

Predators and Birds

© Wes Colgan III. Reprinted with permission.
34

Mammals—Though the FES does not incorporate a formal mammalian predator

sampling component, we have recorded presence and numbers of mustelids caught

while we were trapping for arboreal mammals. To date, we have radio-collared two

weasels caught during trapping sessions. We report mammalian predators and all other

mammalian, amphibian, and reptilian species observed on the study site (appendix 1). A

report on short- and long-tailed weasels, drawn from experiences at the FES site and

other Pacific Northwest arboreal mammal study sites, can be found in Wilson and

Carey (1996).

Winter bird survey—Beginning in winter 1996, surveys were initiated to determine the

effects of the experimental thinnings on winter bird populations. Four experimentally

thinned and four nonthinned stands were selected for a study designed to determine

the feasibility of using standard bird census techniques to assess experimental silvicultural prescriptions. The study used a simple point-count method (Manuwal and Carey

1991), with nine point-count stations at least 80 m apart and at least 80 m from any

habitat edge in each stand. Point-count stations were located at existing grid stations

along established trails to facilitate replication and movement between stations by field

observers. The method yielded 27 point counts per stand, totaling 108 counts each in

thinned and unthinned stands. It is anticipated that the winter bird study will continue

yearly for the next 3 to 5 years. A list of all bird species observed on the study site is

presented (appendix 1).

Because data collection from the FES is in progress, and because most analyses are

preliminary, we present primarily prethinning, between-forest comparisons and summaries for those response variables discussed. In other cases (i.e., coarse woody

debris, vascular plant species richness), we present preliminary, postthinning response

data where relevant. Because we do not yet consider the effects of supplemental

dens on the reported responses, we refer to both den-augmentation-only stands and

Baseline

Conditions
35

control stands as controls unless it is necessary to distinguish between the two. Thus,

contrasts are presented between experimentally thinned and unthinned stands without

regard to supplemental dens. We present a comprehensive checklist of all vascular

plants, fungi, birds, mammals, reptiles, and amphibians observed on FES stands to

date (appendix 1).

Overstory—The overstory vegetation for all stands was dominated by Douglas-fir

(table 2). Tree species other than Douglas-fir were rarely encountered in either the

prism plots or permanent plots or during the postthinning stand assessment. Stands in

the SS blocks were similar to one another in tree size and understory vegetation, but

differed from the FH stands, which were also similar to one another (table 3). The

average diameter of all sampled Douglas-fir prism plot trees was significantly greater

in FH than in SS, and the dominant (site tree) prism-plot Douglas-fir trees in FH were

significantly larger in d.b.h., height, and in height to live crown than those in SS (tables

3 and 5). The same between-forest differences were evident in the larger sample of

permanently marked trees at gridpoints (table 6).

Prethin relative densities (Curtis 1982) in Star and Stellar29 closely matched normal

(see footnote 11) forest stocking; tree densities were high and basal areas were low to

moderate. Star/Stellar values were typical for single cohort, closed canopy stands in

the stem-exclusion phase of forest development (Oliver and Larson 1990). However,

FH stands exhibited conditions more typical of stands managed intensively for wood

production; individual tree d.b.h. was greater as a result of thinning-induced reduction

in tree density. Farley/Hill exhibited pretreatment and control stand RD values of about

7 metric (50 English)—suggested by Curtis (1982) as a maximum RD to avoid

Prethinning and

Control Stand

Conditions

29 Stellar 204 was an exception, probably because the stand

contained a number of small Phellinus weirii openings.

Table 5—Mean d.b.h. in centimeters for all Douglas-fir trees; mean d.b.h. in

centimeters, mean height in meters, and mean height to live crown in meters

(± S.E.) for Douglas-fir site trees from pretreatment prism plots, by forest

Variable Star/Stellar Farley/Hilla

All Douglas-fir prism plot trees:

Diameter at breast height

[d.b.h. in centimeters (sample size)] 33.7 ± 0.5 53.4 ± 0.6

(763) (600)

Douglas-fir site trees:

Diameter at breast height

[d.b.h. in centimeters (sample size)] 42.8 ± 0.7 60.5 ± 1.0

(236) (230)

Height (m) 33.7 ± 0.4 41.8 ± 0.4

Height to live crown (m) 18.8 ± 0.3 21.2 ± 0.4

a Bold numerals indicate significant difference between forests for that variable (Mann-Whitney U-test

[Norus ̆is 1993]; p < 0.01).
36

Table 6—Characteristics of trap-placement trees by blocka

Age at

breast Crown

Block Site Site height Height D.b.h. ratio

name Douglas-fir indexb classc ± S.E. ± S.E. ± S.E. ± S.E.

Feet CentiPercentd (meters) Years Meters meters Percent

Star 99.2 (250) 119 Low 46.9 33.6 40.5 39.0

(36.3) II ± 0.3 ± 0.2 ± 0.6 ± 0.7

Stellar 100 (248) 118 Low 47.4 33.2 40.2 41.7

(36.0) II ± 0.3 ± 0.3 ± 0.6 ± 0.7

Farley 98.8 (245) 130 High 59.6 42.6 58.3 37.4

(39.6) II ± 0.4 ± 0.2 ± 0.7 ± 0.7

Hill 97.5 (242) 133 High 59.6 42.0 58.5 41.1

(40.5) II ± 0.4 ± 0.3 ± 0.9 ± 0.7

a Site Index, tree age, tree height, tree d.b.h., and tree crown ratio are for Douglas-fir trees only.

b The site index calculations are based on 50-year site curves developed by King (1966).

c The site classes are from King (1966).

d The number in parentheses is the total number of live, sampled, trap-placement trees per block. The figure preceeding it

is the percentage of these live trees that were Douglas-fir.

significant tree mortality from suppression and restriction of crown development.

Carey and others (1996a) also suggest RD 7 as a target at age 30 years (given

precommercial thinning at 15 years) when managing for biodiversity. Stands would

then receive a commercial thinning.

The SS pretreatment tree d.b.h. distribution curve (fig. 7a) indicated a skewed-normal

distribution typical of naturally regenerating, single cohort, even-aged forests (Oliver

and Larson 1990). The FH stands (fig. 7b) showed a normal d.b.h. distribution characteristic of intensively managed stands although the distribution is somewhat bimodal.

The subdominant curve contained smaller diameter class (10-25 cm d.b.h.), younger

trees that have established in the more open FH stand conditions. The bimodal

distribution illustrates, in part, stand regenerative processes that occur as a result of

thinnings or canopy gaps (Oliver and Larson 1990).

Trap-placement trees (table 6) were similar in species, age, and size distribution to

those measured in the prism-plots (see tables 2 and 5 for a comparison). Ninety-nine

to 100 percent of SS trap trees were Douglas-fir; 97-99 percent of the FH trap trees

were Douglas-fir. Average age of nonresidual Douglas-fir tagged trees was 54 ± 0.3

years in SS and 67 ± 0.4 years in FH,30 indicating essentially even-aged cohorts of

trees with stand initiation occurring 3 to 5 years after previous harvests.

30 Total age for site index II trees was determined by adding 7 to the

age at breast height (King 1966). We counted an average of 47

rings and 60 rings at breast height, respectively, for trap trees in SS

and FH (table 6).
37

Figure 7—Distribution of d.b.h. classes of prethin, postthin, and logged trees over 7.5 cm d.b.h. in the Star/

Stellar stands (A) and Farley/Hill stands (B). D.b.h. classes are midpoints of 5-cm ranges (e.g., the 40-cm

class contains trees from 37.5 to 42.49 cm d.b.h.) Number of trees are the average number per hectare for

each d.b.h. class and category.

10

15

20

25

30

35

40

45

50

55

60

65

70

75

85

D.b.h. class midpoint (cm)

0

50

100

150

200

250

Trees per hectare

Prethin Postthin Logged

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

95

100

D.b.h. class midpoint (cm)

0

5

10

15

20

25

30

35

Trees per hectare

Prethin Postthin Logged

A

B
38

Prior to experimental thinning, western hemlock regeneration occurred infrequently on

all study stands; Douglas-fir regeneration dominated in both forests. The relatively dry

climate and soils of the region permit regeneration of Douglas-fir in the understory.

Given adequate time and absence of disturbance, the stands may develop western

hemlock associations.31 The study stands closely resemble the Tsuga heterophylla/

Gaultheria shallon/Polystichum munitum and Pseudotsuga menziesii/Gaultheria

shallon associations described by Henderson and others (1989), and the Pseudotsuga

menziesii/Tsuga heterophylla and Pseudotsuga types of the Olympic montane forests

described by Fonda and Bliss (1969). Puget lowland forests are, however, distinctly

different from forests of the Olympic Peninsula and eventually may be recognized as

such (Franklin and Dyrness 1973).

Understory—Pretreatment understory vegetation at SS was highly variable and

reflected the size of overstory openings. Star/Stellar stands generally had a very

dense overstory and, subsequently, had minimal understory development. Ground

mosses were abundant, averaging over 50 percent cover in these stands, with dominant mosses being Kindbergia oregana (Sull.) Ochyra (= Eurhynchium oreganum) and

Hylocomium splendens (Hedw.) B.S.G. Other mosses occurring in these stands

31 Nonetheless, the stumps remaining from the clearcut entries in

the 1920s and 1930s are almost entirely Douglas-fir, suggesting

presettlement history of fires or other disturbances that maintained

Douglas-fir as the dominant tree species.

Table 7—Prethin and postthin vascular plant species richness per plot by post

facto relative density (RD) class in 1994, 1 year after thinning

Postthinb

Post facto RD classd

All

Forest Prethina Control thinnedc RD8 RD6 RD4 RD2

----------- Mean number of species per plot ± S.E. (sample size)--------------

Star/Stellar 8.6 14.8 24.1 23.7 19.8 24.1 29.7

± 0.3 ± 0.6 ± 0.9* ± 1.5 ± 1.3 ± 1.5 ± 1.4

(120) (24) (71) (6) (27) (17) (21)

Farley/Hill 15.1e 18.5 34.6 29.4 31.4 26.9 44.5

± 0.4 ± 1.2 ± 1.2* ± 2.1 ± 1.5 ± 2.0 ± 1.4

(120) (24) (72) (12) (22) (14) (24)

a Plot size = 25 m2.

b Plot size = 100 m2

.

c Asterisks(*) indicate significant difference between all thinned plots and CT plots for that forest (t-test;

p < 0.01).

d Classes are defined as follows: RD2 = retained RD of < 3.25 (metric); RD4 = retained RD of 3.25 ≤ RD

< 4.75; RD6 = retained RD of 4.75 ≤ RD ≤ 6.75; RD8 = retained RD > 6.75.

e Bold numerals indicate significant difference between SS and FH for that treatment (Mann-Whitney U-test

(Norus ̆is 1993); p < 0.05).
39

Table 8—Mean cover of vascular plant subcanopy layers by stratum by post facto

relative density (RD) class for each forest type in 1994, 1 year after thinning

Farley/Hill stratumb c Star/Stellar stratum

Treatment Sample Sample

classa size 1 2 3 size 1 2 3

--Percent cover ± S.E.-- --Percent cover ± S.E.--

Control 24 28.3 48.3 10.0 24 17.7 25.8 6.3

± 4.3 ± 4.7 ± 2.0 ± 4.6 ± 4.1 ± 1.9

RD8 12 61.3 25.3 8.3 6 5.3 18.0 0.7

± 2.7 ± 6.2 ± 2.6 ± 0.8 ± 4.8 ± 0.4

RD6 22 49.5 36.6 7.2 27 8.4 11.3 2.4

± 3.9 ± 4.6 ± 1.0 ± 2.5 ± 2.0 ± 0.7

RD4 14 48.0 24.7 15.9 17 10.9 15.1 2.9

± 5.3 ± 4.6 ± 3.1 ± 1.8 ± 1.9 ± 1.0

RD2 24 62.7 14.5 5.9 21 26.5 21.4 3.3

± 1.3 ± 1.8 ± 1.5 ± 4.0 ± 4.0 ± 1.5

a Classes are defined as follows: RD2 = retained RD of < 3.25 (metric); RD4 = retained RD of 3.25 ≤ RD

< 4.75; RD6 = retained RD of 4.75 ≤ RD ≤ 6.75; RD8 = retained RD > 6.75.

b Stratum 1 included forbs and ferns < 0.5 m tall; stratum 2 included low shrubs 0.5 to 2.0 m tall; stratum 3

included midstory shrubs and understory trees > 2.0 m tall.

c Bold numerals indicate significant difference between Farley/Hill and Star/Stellar for a particular treatment

and RD class (Mann-Whitney U-test [Norus ̆is 1993]; p < 0.05).

included Rhytidiadelphus loreus (Hedw.) Warnst, R. triquetrus (Hedw.) Warnst,

Plagiothecium undulatum (Hedw.) B.S.G., Plagiomnium insigne (Mitt.) Kop.,

Leucolepis acanthoneuron (Schwaegr.) Lindb. (= L. menziesii), Isothecium stoloniferum Brid., and Dicranum scoparium Hedw. (see footnote 7). Where openings

occurred, understory vegetation was dominated by salal, Oregongrape, swordfern,

hazel, Scouler’s willow, and oceanspray.

Pretreatment understory vegetation of FH was more diverse than at SS. The former

stands were more open, incident light was higher in the understory, and species

richness was significantly greater. Of 120 pretreatment vegetation plots in each forest,

FH plots averaged 15.1 understory vascular plant species (range = 5 to 31) per 25-m2

plot, compared to an average of 8.6 species in SS (range = 3 to 16) (table 7). Control

and den-augmentation-only stand permanent plots in FH also had significantly higher

vascular plant species richness than did those in SS (table 7). Individual plants were

often more robust in FH, and dominants (salal, swordfern, Oregongrape, oceanspray,

hazel, and brackenfern) had higher cover values than at SS (table 8). Moss cover was

substantially less in FH than in SS, probably due to the extensive development of

deciduous understory shrubs and forbs. Altogether we found about 90 species of

vascular plants on FES stands during our pretreatment and control stand sampling

(table 9).
40

Table 9—Vascular plant species found in control permanent plots and in

prethinning understory vegetation plots, by block

Farley Hill Star Stellar

(species (species (species (species

richness richness richness richness

Plant taxona = 60) = 73) = 38) = 37)

Acer macrophyllum • •

Achlys triphylla • • ••

Actaea rubra •

Adenocaulon bicolor • •

Alnus rubra • •

Amelanchier alnifolia • • ••

Anemone deltoidea • •

Anaphalis margaritacea •

Arbutus menziesii •

Asarum caudatum •

Athyrium filix-femina •

Blechnum spicant • •

Bromus vulgaris • •

Carex deweyana • •

Campanula scouleri •• •

Chimaphila menziesii •

Chimaphila umbellata • •

Corylus cornuta var. californica • • ••

Collomia heterophylla •

Corallorhiza maculata • ••

Cornus nuttallii • •

Crepis capillaris •

Cytisus scoparius •

Disporum hookeri • •

Dryopteris campyloptera • •

Elymus glaucus var. glaucus • •

Epilobium angustifolium •

Festuca occidentalis • • ••

Festuca subulata •

Festuca subuliflora • •

Fragaria virginiana var. platypetala •

Frangula purshiana • • ••
41

Table 9—Vascular plant species found in control permanent plots and in

prethinning understory vegetation plots, by block (continued)

Farley Hill Star Stellar

(species (species (species (species

richness richness richness richness

Plant taxona = 60) = 73) = 38) = 37)

Galium aparine • • ••

Gaultheria shallon • • ••

Geum macrophyllum var. macrophyllum •

Goodyera oblongifolia • • ••

Hieracium albiflorum •

Holodiscus discolor •••

Holcus lanatus •

Hypochaeris radicata • •

Ilex aquifolium • • ••

Lathyrus polyphyllus •••

Leucanthemum vulgare • •

Linnaea borealis • • ••

Listera caurina •• •

Lonicera ciliosa • • ••

Lotus micranthus •

Luzula parviflora •

Mahonia aquilinum •

Mahonia nervosa • • ••

Maianthemum stellatum •

Melica subulata • •

Moehringia macrophylla •

Monotropa uniflora •

Mycelis muralis • •

Nemophila parviflora • •

Oemlaria cerasiformis •••

Osmorhiza berteroi • •

Polypodium glycyrrhiza • •

Polystichum munitum • • ••

Prunella vulgaris •

Pseudotsuga menziesii • • ••

Pteridium aquilinum • • ••

Pyrola aphylla • •
42

Table 9—Vascular plant species found in control permanent plots and in

prethinning understory vegetation plots, by block (continued)

Farley Hill Star Stellar

(species (species (species (species

richness richness richness richness

Plant taxona = 60) = 73) = 38) = 37)

Pyrola asarifolia • •

Ribes sanguineum •

Rosa gymnocarpa •• •

Rubus discolor •

Rubus laciniata •

Rubus leucodermis • •

Rubus parviflora • •

Rubus spectabilis • •

Rubus ursinus • • ••

Sambucus racemosa •••

Salix scouleriana • •

Senecio jacobea • •

Senecio sylvaticus •

Stellaria crispa •• •

Symphoricarpos alba •• •

Symphoricarpos hesperius • • ••

Taxus brevifolia •

Thuja plicata •

Tiarella trifoliata • •

Trientalis borealis ssp. latifolia • • ••

Trillium ovatum • • ••

Tsuga heterophylla •

Urtica dioica •

Vancouveria hexandra • • ••

Vaccinium ovatum •••

Vaccinium parvifolium • • ••

Vicia gigantea •

Viola sempervirens • • ••

a Species found in all 4 blocks are in bold. Introduced species are underlined. Authorities and common

names are found in appendix 1.
43

The fallen tree components of the two forests differed markedly. Star/Stellar stands

averaged 7 to 8 percent CWD cover (over 15 cm diameter large end), compared to 2

to 3 percent cover in the eight FH stands (table 10). The subset of stands (101, 103,

201, 202, 302, 303, 401, 403) selected for thinning differed similarly (table 11).

Some large trees were retained during the 1937 regeneration harvest in SS. Although

SS had not been thinned or harvested since that entry, about one-third of residual

standing trees were felled in 1986, because they were thought to pose a hazard to

low-flying helicopters. Thus, SS contained three distinct populations of fallen trees: (1)

small-diameter fallen trees from the current stand (trees that died from suppression or

root rot); (2) large-diameter and well-decayed fallen trees from the previous stand; and

(3) larger diameter, relatively undecayed trees from felling operations of 1986.

Farley/Hill stands contained minimal coarse woody debris; little remained in the stands

except for what had fallen since the last thinning entry (table 10). At the time of the

last thinnings, the entire, harvested trees were yarded to landings and bucked into

logs; tree crowns and cull material were burned on site. Debris in the stand was piled

and burned.

Total standing dead tree (snag) density was significantly greater in SS than in FH

(table 12). On average, there were 34 snags/ha (> 15 cm d.b.h.) in SS vs. 19 snags/ha

in FH. Standing dead trees also had three distinct size and decay class distributions:

(1) small current-stand material (e.g., part of the cohort of trees currently in place);

(2) large-diameter and tall residual standing dead trees from the previous stand (e.g.,

trees not cut during the harvest entries of the 1920s and 1930s); and (3) well-decayed

and short (< 2 m), standing dead stumps remaining from the harvest of the 1920s or

1930s. Density of large snags (> 30 cm d.b.h.) was greater in SS than in FH, but not

significantly so. Large snags in SS were significantly larger in diameter than in FH, but

FH large snags were significantly taller than those in SS. Snags retained from the

preceding stand (bark sloughed, stems well decayed, all but the largest branches

completely missing) in SS were similar in diameter to, but significantly taller than,

those in FH. Residual snags also were significantly more abundant in SS than in FH.

Snags from the current cohort of trees were significantly larger and significantly taller

in FH than in SS. Density of current snags was greater in FH than in SS (3.6 snags/ha

vs. 1.6 snags/ha); however, this difference was not statistically significant (table 12).

Arboreal rodent communities in both forests were composed of the northern flying

squirrel, Townsend’s chipmunk, and Douglas’ squirrel. Northern flying squirrels were

more abundant in SS than in FH, and the Townsend’s chipmunk was more abundant in

the FH. Douglas’ squirrels were caught infrequently in both forests (table 13).

Forest-floor small mammal communities in both forests included two shrew species

(Trowbridge’s and montane), the deer mouse, the Oregon creeping vole, the shrewmole, and the southern red-backed vole. Two additional species, the Columbian deer

mouse and the vagrant shrew, occurred occasionally in both forests. Other infrequently caught species included the Pacific jumping mouse, the coast mole, and the

Pacific water shrew. Although species richness was similar in both forests, overall

abundance of forest-floor small mammals was significantly higher in FH than in SS in

1992 and in 1993 (table 14).

Prethinning Coarse

Woody Debris

Standing Dead Trees

Arboreal Rodents

Forest-Floor Small

Mammals
44

Table 10—Prethinning and postthinning coarse woody debris cover

Prethinninga b

Stands to

Forest All stands be thinned Postthinningc

Mean percent cover ± S.E. (sample sized)

Star/Stellar 7.4 ± 0.5 7.1 ± 0.5 5.2 ± 0.3

(32) (16) (24)

Farley/Hill 2.5 ± 0.2 2.9 ± 0.4 1.6 ± 0.1

(48) (24) (24)

a Data are from all 16 stands and for the 8 stands designated as to-be-thinned (e.g., 101, 103, 201, 202, 302,

303, 401, 403).

b Bold figures indicate a significant difference between forests for that category or subset (Mann-Whitney

U-test [Norus ̆is 1993]; p < 0.01).

c Data are from thinned stands only (e.g., 101, 103, 201, 202, 302, 303, 401, 403).

d Number of sampled 200-m lines.

Table 11—Coarse woody debris (CWD) cover before and after thinning by decay

class by forest, thinned stands only

Star/Stellarb Farley/Hillb

(sample size = 16c ) (sample size = 24c)

CWD decay classa Prethin Postthin Prethin Postthin

-------------------Mean percent cover ± S.E. -----------------

Decay class I 0.4 ±0.1 0.6 ± 0.1 0.6 ± 0.1 0.8 ± 0.1

Decay class II 3.9 ± 0.4 4.1 ± 0.4 1.4 ± 0.2 0.7 ± 0.1

Decay class III 2.8 ± 0.4 0.5 ± 0.2 0.9 ± 0.3 0.2 ± 0.1

 Total CWD 7.1 ± 0.5 5.2 ± 0.4 2.9 ± 0.4 1.6 ± 0.1

a Our decay classes are modified from the 5-decay-class system of Fogel and others (1973) and modified by

Sollins (1982). Decay classes 1 and 2 of Fogel and others (1973) were treated as class I; classes 3 and 4

were treated as class II; and decay class 5 was treated as class III.

b Bold figures indicate significant difference between prethinning and postthinning cover for that category

(Wilcoxon matched-pairs signed-rank [Norus ̆is 1993]; p < 0.01).

c Number of sampled 200-m lines.
45

Table 12—Snag characteristics (mean ± S.E), by forest (prethinning conditions)

Snag categorya

All snags All snags Residual Current

>15 cm >30 cm >30 cm >30 cm

Forest and attribute d.b.h.b d.b.h.c d.b.h.c d d.b.h.c d

Star/Stellar:

 Density (no./ha ± S.E.) 34.1 ± 2.4 5.6 ± 0.8 3.9 ± 0.5 1.6 ± 0.4

 D.b.h. (cm) ± S.E. - 70.7 ± 2.8 83.5 ± 3.2 38.2 ± 1.1

 Height (m) ± S.E. - 8.7 ± 0.7 6.0 ± 0.6 15.0 ± 1.6

Farley/Hill:

 Density (no./ha ± S.E.) 18.9 ± 4.5 4.6 ± 1.3 1.0 ± 0.5 3.6 ± 1.0

 D.b.h. (cm) ± S.E. - 54.8 ± 2.3 87.3 ± 6.7 45.4 ± 1.4

 Height (m) ± S.E. - 20.6 ± 1.2 4.6 ± 1.0 25.3 ± 1.2

- = not applicable; d.b.h. and height were not measured on snags < 30 cm d.b.h.

a Bold figures indicate significant difference between forests for that attribute (Mann-Whitney U-test [Norus ̆is

1993]; p < 0.05).

b Densities of all snags > 15 cm are based on complete snag counts in every other 40- by 40-m grid cell in

each of the 8 stands per forest (hence 196 grid cells per forest).

c For all snags > 30 cm d.b.h, sample size = 170 in SS and 141 in FH; for residual snags, sample size = 120

in SS and 31 in FH; for current snags, sample size = 50 in SS and 110 in FH.

d Residual snags show evidence of well-decayed sapwood (and heartwood), typically have lost their bark,

and have lost most or all of their branches. Current snags have most of their bark intact, are less decayed,

and retain more branches (see Cline and others 1980 for a discussion of snag decay).

Table 13—Total summed captures, total captured individuals, mean density, and

range of density of arboreal mammals, by foresta

Farley/Hill Star/Stellar

Total captures/ Total captures/

Species total individuals Density total individuals Density

No./ha ± S.E. No./ha ± S.E.

(range) (range)

Flying squirrel 207/126 0.3 ± 0.1 384/214 0.6 ± 0.1

(0.18 - 0.46) (0.28 - 1.04)

Townsend’s 641/312 0.7 ± 0.1 100/34 0.1 ± 0.02

chipmunk (0.29 - 1.43) (0.0 - 0.28)

Douglas’ squirrel 29/20 - 21/20 -

- = not calculated.

a Data are summaries (captures, individuals) or means (density) of fall 1991 and spring and fall 1992

trapping sessions, and represent prethinning conditions only.
46

Table 14—Mean captures per 100 corrected trap nights (± S.E.) for all forest floor

small mammal species combined, and for selected species, by foresta

1992 1993

Species Farley/Hillb Star/Stellar Farley/Hillb Star/Stellar

All species 13.9 ± 1.1** 9.5 ± 0.3 11.4 ± 1.5* 7.8 ± 0.7

Peromyscus maniculatus

(deer mouse) 2.5 ± 0.4*** 0.6 ± 0.2 0.7 ± 0.1** 0.15±0.03

Microtus oregoni

(Oregon creeping vole) 2.1 ± 0.6** 0.2 ± 0.1 1.8 ± 0.4*** 0.07±0.04

Sorex monticolus and

S. trowbridgii 3.1 ± 0.4 2.6 ± 0.4 3.5 ± 0.3 2.7 ± 0.4

a Data are from unthinned (control) stands only (102, 104, 203, 204, 301, 304, 402, 404).

b FH captures in a given year differ significantly from SS captures (t - test): * = p < 0.10; ** = p < 0.05;

*** = p < 0.01.

The deer mouse and Oregon creeping vole accounted for most of the between-forest

difference in overall small mammal abundance in both years (table 14). In 1992, deer

mice and Oregon creeping voles were significantly more abundant in FH than in SS.

In 1993, both taxa were again significantly more abundant in FH than in SS. Other

taxa were trapped consistently and frequently (e.g., southern red-backed vole,

Trowbridge’s and montane shrews, shrew-mole), or infrequently (e.g., Columbian deer

mouse, vagrant shrew) in both forests in both years, and did not greatly contribute to

differences in overall abundance between forests. Trowbridge’s shrew and the montane shrew were numerically the most abundant forest-floor small mammals in both

forests in both years (table 14).

All descriptions provided for stand and subtreatment forest parameters considered

standing trees, both live and dead, with a d.b.h. > 20 cm (the exception being forestlevel d.b.h. distributions where smaller trees were also measured—see below). Stand

and forest-level descriptions are based on weighted means derived from the actual

distribution of post facto thinning classes for each stand. Subtreatment-level descriptions are based on forestwide averages for each post facto thinning class.

Stand and forest level stocking conditions—Thinning operations reduced the

heavily stocked SS stands from an average of 454 stems/ha to 291 stems/ha and

removed about 30 percent of the basal area from SS stands. This represented a

reduction from a forestwide average RD of 7.4 before thinning, to RD 4.8 after thinning

(table 15). In FH stands, stocking levels were reduced from an average of 203 stems/

ha to 147 stems/ha; basal area was reduced by about 24 percent. Relative density in

FH treated stands was reduced from an average of 7.9 to a residual mean of 4.7

(table 15).

Postthinning

Conditions

Silvicultural

Conditions
47

Table 15—Prethinning and postthinning silvicultural conditions by forest and post facto relative

density (RD) class

Prethinning Posttreatmentc

Relative Relative

Treatment Sample Basal Tree density Basal Tree density

Forest classa sizeb area density (metric) area density (metric)

m2/ha No./ha m2/ha No./ha

Star/Stellar RD2 36 31.5 351.1 5.6 11.4 108.9 1.9

± 2.2 ± 26.3 ± 0.4 ± 0.9 ± 9.6 ± 0.1

RD4 68 37.3 410.1 6.4 25.3 263.0 4.1

± 1.2 ± 17.7 ± 0.2 ± 0.3 ± 6.6 ± 0.1

RD6 66 49.1 503.2 8.3 35.9 353.6 5.8

± 1.0 ± 13.5 ± 0.2 ± 0.5 ± 6.3 ± 0.1

RD8 26 60.4 589.4 10.2 49.1 457.6 7.9

± 1.9 ± 24.3 ± 0.3 ± 1.3 ± 14.9 ± 0.2

All thinned 196 43.2 454.4 7.4 29.5 291.0 4.8

± 1.0 ± 11.0 ± 0.2 ± 0.9 ± 8.7 ± 0.1

Control 14 42.7 515.1 7.5 42.7 515.1 7.5

± 5.1 ± 60.3 ± 0.9 ± 5.1 ± 60.3 ± 0.9

Farley/Hill RD2 42 43.2 199.4 7.7 14.7 59.1 2.0

± 3.1 ± 15.9 ± 0.6 ± 0.9 ± 4.8 ± 0.1

RD4 50 35.6 152.4 5.4 30.9 130.2 4.0

± 1.1 ± 6.8 ± 0.3 ± 0.5 ± 3.8 ± 0.1

RD6 77 52.8 216.9 8.5 44.1 178.4 5.6

± 1.1 ± 5.6 ± 0.2 ± 0.6 ± 3.4 ± 0.1

RD8 27 67.6 259.2 10.8 59.7 223.5 7.8

± 1.6 ± 8.2 ± 0.4 ± 0.9 ± 6.3 ± 0.1

All thinned 196 48.4 202.5 7.8 36.6 146.8 4.7

± 1.1 ± 5.1 ± 0.2 ± 1.1 ± 4.4 ± 0.1

Control 12 44.3 259.7 6.5 44.3 259.7 6.5

± 2.5 ± 15.7 ± 0.3 ± 2.5 ± 15.7 ± 0.3

a Classes are defined as follows: RD2 = retained RD of < 3.25 (metric); RD4 = retained RD of 3.25 ≤ RD < 4.75; RD6 = retained

RD of 4.75 ≤ RD ≤ 6.75; RD8 = retained RD > 6.75.

b Number of sampled grid cells.

c Bold figures indicate significant difference between posttreatment and control stand conditions for that variable and forest

(t-test; p < 0.001).
48

Residual stems per hectare and basal area in thinned stands in both forests differed

significantly from those of their respective unthinned stands (table 15). In SS, basal

area in thinned stands was 29.5 ± 0.9 m2/ha vs. 42.7 ± 5.1 m2/ha in SS control stands

(t-test, p < 0.001). Stem density in SS thinned stands was 291.0 ± 8.7 trees/ha vs.

515.0 ± 60.3 trees/ha in SS control stands (t-test, p < 0.01). In FH, basal area in

thinned stands was 36.6 ± 1.1 m2/ha in thinned stands vs. 44.3 ± 2.5 m2/ha in control

stands (t-test, p < 0.01). Stem density in FH thinned stands was 146.8 ± 4.4 trees/ha

vs. 259.7 ± 15.7 trees/ha in control stands (t-test, p < 0.001).

Forest-level d.b.h.-class distributions—Owing to disparate management histories in

SS and FH, experimental thinnings had different effects on the resulting d.b.h.-class

distributions in the two forests. In SS, with its pretreatment skewed-normal distribution

typical of a naturally regenerated stand (Oliver and Larson 1990), thinnings were

clearly weighted towards removal of smaller d.b.h. classes (fig. 7a). In contrast,

treatment of the previously thinned FH forest achieved a more equitable removal of

trees from all d.b.h. classes (fig. 7b). However, some large-diameter trees (> 70 cm

d.b.h.) were removed in FH, even though the prescription stipulated thinning from

below and leaving large, dominant trees. These large trees probably were removed

from RT thinning cells in FH, where only the largest diameter trees or hardwoods were

selected for retention. Such trees also may have exhibited low vigor (e.g., thin

crowns) or may have been in severe competition with other leave trees at the time of

marking.

Subtreatment level descriptions—The postthinning stand assessment was performed to document within-stand distribution of cells of known stocking levels. Having

adopted Curtis’ (1982) relative density as an index of stand density, and having

determined relative density within each treated cell, we propose four post facto relative

density classes to describe and encompass the range of variability found in the postVDT stands. The post facto classes are presented to characterize the actual variability

resulting from our thinning regime and to facilitate the use of RD as a covariate in the

analysis of other experimental variables. Stated ranges are provisional and will be

further refined during the course of this study. Post facto RD classes were delineated

and designated as follows, with designations representing the approximate midpoint of

the suggested range of relative densities:

1. RD 2: retained relative density of < 3.25. These were the most heavily thinned cells

and correspond to RT target areas.

2. RD 4: retained relative density of 3.25 ≤ RD < 4.75. Areas of relatively heavy

thinning, cells of this class correspond to target HT areas.

3. RD 6: retained relative density of 4.75 ≤ RD ≤ 6.75. Areas of relatively light

thinning, cells of this class correspond to target LT cells.

4. RD 8: retained relative density > 6.75. These cells had insufficient trees removed

during VDT for them to substantially depart from suppression mortality conditions.

Such cells had some trees removed during the experimental thinnings, but

remained in an overstocked, closed-canopy state.
49

Distribution of cells by post facto class, and average pretreatment and posttreatment

basal area, tree density, and relative density are summarized by forest in table 15.

Distribution of post facto thinning cells by stand and forest is summarized in table 16.

Descriptive statistics of each post facto RD class for each treated stand and a cell-bycell summary of forest conditions are provided in appendix 4. Stand maps of each

treated stand are found in appendix 5.

Root grafting—Of 1,224 stumps with a d.b.h. equivalent of > 20 cm, 7 percent

(n = 83) had some callus growth on the cut surface.32 The mean d.b.h. equivalent

of callused stumps was 34 cm (of those > 20 cm d.b.h.). Incidence of callusing was

slightly higher in Star (8 percent, or 34 of 413) and Hill (9 percent, or 13 of 150) than

in Stellar (5 percent, or 25 of 472) or Farley (6 percent or 11 of 189).

Forest health and logging damage—A total of 128 of 1,417 sampled trees (excluding the 65 dead trees) had some visual damage or reduced vigor. Logging damage

was the most commonly observed type of damage, but it had low overall incidence

(7 percent) among the total tree population. Typically, logging damage occurred at or

near ground level, suggesting that it was the result of skidding operations. At the time

of sampling, most damaged trees (98 percent) showed no outward signs of reduced

vigor. Other damage or diseases (including Phellinus weirii) were present at very low

levels (table 17).

Logging error—A total of 1,482 trees > 20 cm d.b.h. were sampled on all stands;

these included 65 dead trees and 9 standing live trees that had been marked to be cut.

Among sampled trees > 10 cm d.b.h. (n = 2,597), 32 standing trees were marked as

cut-trees but remained uncut.

Underplanting—Although four tree species were planted (Thuja plicata, Pinus

monticola, Abies grandis, and Alnus rubra), only the three conifer species were

sampled because of difficulty in determining whether red alder seedlings and saplings

had been planted or had resulted from natural regeneration. Because of difficulty in

locating many dead (and live) underplanted seedlings, we report current observed

seedling density rather than percentage of survival.

All blocks demonstrated a general trend of decreasing seedling presence with increasing tree cover. Seedling density ranged from stand averages of 189 to 263 seedlings/

ha in the RD2 cells and from 120 to 225 seedlings/ha in other thinned cells. Average

density of grand fir in RD2 cells in both forests was substantially greater than density

of the other two conifer species. Grand fir densities also were higher in RD2 cells than

in any other thinned cells. In other thinned cells, approximately even numbers of each

of the three underplanted species were found in both forests. The average numbers

per hectare of each underplanted conifer species in RD2 and other-thinned cells in

each forest are shown in figure 8.33

32 Callus growth is used to indicate root-grafting.

33 Underplanting initially was stratified by target subtreatments, with

only RT and HT cells to be planted. However, following the

thinning assessment, we determined that all four post facto thinning

classes (RD2, RD4, RD6, RD8) contained cells that had been

underplanted. Because few cells of the RD6 and RD8 post facto

classes contained planted trees, we present RD2 seedling density

in contrast to all other thinned cells that had been underplanted

(RD4, RD6, and RD8).
50

Table 16—Distribution of post facto relative density (RD) classes by stand and

forest, thinned stands only

Post facto thinning classa

Stand RD2 RD4 RD6 RD8

---------------- Number of cells (percentage of total) ---------------

Star/Stellar:

 101b 13 (26) 15 (31) 13 (27) 8 (16)

 103 6 (12) 17 (35) 25 (51) 1 (2)

 201 8 (16) 18 (37) 12 (24) 11 (22)

 202 9 (18) 18 (37) 16 (33) 6 (12)

 Total, Star/Stellar 36 (18) 68 (35) 66 (34) 26 (13)

Farley/Hill:

 302 11 (22) 5 (10) 26 (53) 7 (14)

 303 13 (26) 14 (29) 17 (35) 5 (10)

 401 8 (16) 20 (41) 10 (20) 11 (22)

 403 10 (20) 11 (22) 24 (49) 4 (8)

 Total, Farley/Hill 42 (21) 50 (26) 77 (39) 27 (14)

a Classes are defined as follows: RD2 = retained RD of < 3.25 (metric); RD4 = retained RD of 3.25 ≤ RD <

4.75; RD6 = retained RD of 4.75 ≤ RD ≤ 6.75; RD8 = retained RD > 6.75.

b 49 cells per stand; 196 per forest.

Table 17—Percentage of sampled trees showing evidence of damage, by stand

Stand

Damage type 101 103 201 202 302 303 401 402 403

Animal damage

 (species unknown) 0 0.2 0 0 0 0 0 0 0

Bark beetle

 (species unknown) 0 0.3 0 0.1 0 0 0.4 2.7 0

Douglas-fir beetlea 0 0 0 0 3.5 0.8 3.0 4.1 0

Laminated root rot 0 0 0.2 0.1 0 0 0.2 0 0

Logging damage 7.3 1.3 2.3 5.9 7.5 4.5 4.0 0 4.8

Wind 0 0 0.1 0 0 0 0 0 0

Other damage

 (unspecified) 1.4 0.4 0 0.1 0.4 0 1.2 2.7 0.6

a Dendroctonus pseudotsugae Hopkins (name from Furniss and Carolin 1977).
51

Natural tree regeneration—Seven tree species were commonly found regenerating

at the time of sampling: Douglas-fir, western hemlock, western redcedar, Scouler’s

willow, black cottonwood, red alder, and bigleaf maple. In addition, Quercus garryana

and Ilex aquifolium were found infrequently. Summaries of regeneration data indicated

the sample size was too small to draw any significant inferences. Only 1.24 percent of

the area of any treated stand was sampled for tree regeneration, with many plots

containing no regenerating trees.

Natural regeneration in all stands was dominated by Douglas-fir, which comprised 60

to 100 percent of regenerating trees. Substantial numbers (25 to 40 percent of total

seedlings) of Scouler’s willow and red alder seedlings were found in FH thinned

stands. About 30 percent of seedlings were western hemlock in the Star 104 control

stand, and about 20 percent of seedlings in the Star 101 thinned stand were western

redcedar. No further sampling or analysis was performed to examine within-stand,

between-stand, or between-treatment variation across blocks or forests.

Most young, naturally regenerating trees belonged to the lowest height class (fig. 9).

In SS, up to 90 percent of natural regeneration consisted of young (less than 1 year

old), newly emergent Douglas-fir seedlings. Many more small seedlings were seen in

RD2 cells than in other thinned cells, with a total of 6,000 to 7,000 seedlings/ha (< 50

cm tall) found in some SS RD2 cells. Very few seedlings of the larger height classes

were found in thinned SS cells (fig. 9a).

The FH treated stands contained fewer total naturally regenerating seedlings and

saplings than SS treated stands, but exhibited a more even distribution of size classes

(fig. 9b). More small (< 50 cm) seedlings were found in FH RD2 cells than in other

treated cells; the larger two regeneration height classes were found in approximately

Figure 8—Density of underplanted conifer species by forest for post facto

root rot thinned cells and all other thinned cells that had been underplanted.

Figures are averages of the four treated stands per forest.

104.5 103.7

74.3 56.7

72.6 65.4

65.4

56.6

74.3

59.2

71.8

68.5

SS RD2 FH RD2 SS other thin FH other thin

Forest and thinning category

0

50

100

150

200

250

300

Trees per hectare

Grand fir

White pine

Western redcedar

104.5 103.7

74.3 56.7
52

equal numbers in all FH thinned cells. Very few naturally regenerating seedlings were

found in SS control stands; of those found in FH control stands, most were in the 10-

to 50-cm height class (fig. 9).

Permanent vegetation plots were initially stratified by target subtreatments. After the

silvicultural assessment, retained relative density values for each grid cell (and consequently each permanent plot location) were computed; vegetation summaries are

presented in relation to the post facto RD classes based on retained RD and described

above. Locations of all vegetation permanent plots are provided in appendix 3, along

with retained relative density values and post facto RD class designations for each

plot. One year after experimental thinnings, FH stands had significantly more plant

species per plot than SS in both control and thinned stands. Thinning resulted in

significant increases in species richness in both forests, with the greatest increases in

species richness occurring in areas of lowest retained RD (table 7). Of the 146

Figure 9—Densities of naturally regenerating seedlings by size class by

post facto thinning class for Star/Stellar (A) and Farley/Hill (B) forests.

Figures are averages of the four treated stands per forest or from the two

sampled control stands per forest.

Postthinning

Understory Vegetation

RD2 RD4 RD6 RD8 CT

Star/Stellar thinning class

0

500

1000

1500

2000

2500

3000

3500

4000

Average trees per hectare

RD2 RD4 RD6 RD8 CT

Farley/Hill thinning class

0

500

1000

1500

2000

2500

Average trees per hectare

Tree height (cm)

0.1-10 10.1-50

50.1-100 100+

A 

B 
53

vascular plant species found during permanent plot sampling in 1994, 138 were found

in all treatments combined in FH (only 69 species in control stands), and 101 species

were found in all treatments combined in SS (only 44 species in control stands). A

complete list of all vascular plant species found during the study is given in appendix 1.

Cover values in FH were significantly higher than in SS for forbs-ferns and tall shrubsunderstory trees in all RD classes. Though cover of low shrubs in RD2 plots in FH

was less than in SS, differences were not statistically significant. The RD2 treatment

cover for low shrubs was less in FH than in SS, probably because some SS RD2

areas were in actual root rot pockets (table 8).

Of 34 vascular plant species unique to FH control plots (e.g., not found in SS controls),

12 were alien (nonnative) species (none was a graminoid), 4 were native graminoids,

8 were native woody understory or midstory species, and 10 were native forbs and

ferns. All 9 species unique to SS controls were native forb, fern, or shade-tolerant tree

species. Of 44 species found only in FH (all treatments combined), 14 were aliens (4

were graminoids), 7 were native graminoids, 7 were native woody understory or

midstory species, and 12 were native forbs and ferns. No species was unique to SS

(all treatments combined).

By summer 1994, average per plot species richness in both forests had increased by 7

to 26 species in thinned areas compared to controls (table 7). The increase was most

evident in the forb-fern layer, where about 40 percent of the new species in both

forests (all treatments combined) were aliens. In both forests, weed species were

mostly composites (family Asteraceae), grasses, and legumes. Of 65 plant species

found only in thinned areas (not in control plots), 33 (51 percent) were alien species (7

were grasses), and 14 were graminoids. Of the 20 plant species found only in RD2

permanent plots, 16 (80 percent) were aliens (5 were grasses). All 20 species were

found infrequently and at low cover values.

Weeds had a low percentage of cover in both forests, except for occasional thick

patches of wall lettuce, bull thistle, and clovers that were mostly confined to RD2

treatments. Farley/Hill also had higher understory and midstory woody species

richness and cover values than SS, where the lengthy stem-exclusion phase had

precluded understory development (tables 7 and 8).

After thinning, forestwide averages of CWD cover in SS remained significantly greater

than in FH (table 10), with all experimentally thinned stands having suffered reductions

in CWD cover. These reductions were statistically significant in both forests (table 11).

In SS, prethinning and postthinning cover of decay classes I and II were not significantly different, but decay class III cover was significantly reduced after thinning. In

FH, overall CWD cover was significantly reduced after thinning; cover of decay classes

II and III was also significantly reduced. Cover of decay class I increased significantly,

though the difference did not appear to be biologically significant (table 11).

Postthinning Coarse

Woody Debris
54

Stands differed in availability of dens for northern flying squirrels. Densities of residual

stumps and live conifers with top rot were significantly less in the SS thinned stands

than in SS unthinned stands (table 18). Other den categories did not differ significantly

between experimentally thinned and unthinned stands in either forest (data not

shown). Of den types not differing significantly in abundance between thinned and

unthinned stands (within a forest), residual snags, residual live trees, and conifers with

stick nests were significantly more abundant in SS than in FH; coniferous snags

without cavities and deciduous snags without cavities were significantly more abundant in FH than in SS. Other den types did not differ significantly in abundance

between forests (table 18). Thinning subtreatments within forests (FH, SS) did not

differ in abundance of dens.

In both forests, use of supplemental nestboxes and cavities (defined by the presence

of nest material) steadily increased after their installation (fig. 10). By the fifth nestbox

check in March 1995, 80 percent (102 of 128) of all nestboxes exhibited use; three

stands had ≥ 93 percent use of nestboxes (all in FH), one of which (303) had 100

percent use. During March 1996 (after the addition of 32 nestboxes per forest in

December 1995), both nestboxes (70 percent) and cavities (35 percent) were more

frequently used in FH than in SS (45 percent and 25 percent respectively). Nest

materials in both forests were predominantly moss (primarily Isothecium stoloniferum

Den Site Availability

and Snags

Supplemental Den Use

Table 18—Mean densities per hectare (± S.E.) of den site types by forest and in

thinned and unthinned stands within forestsa

Den site type Star/Stellar Farley/Hill

Residual snags 3.5 ± 0.7b 0.4 ± 0.1

Residual live trees 2.7 ± 0.8 0.4 ± 0.4

Conifers with sticknest 1.7 ± 0.3 0.6 ± 0.1

Conifer snags with cavity 0.4 ± 0.1 0.4 ± 0.1

Conifer snags without cavity 1.7 ± 0.5 4.1 ± 1.0

Deciduous snags with cavity 0.0 ± 0.0 0.2 ± 0.1

Deciduous snags without cavity 0.1 ± 0.1 2.1 ± 0.8

Unthinned Thinned Unthinned Thinned

stands stands stands stands

Residual stumps 62.6 ± 2.0 33.9 ± 4.4 21.9 ± 3.1 20.5 ± 3.8

Live conifers with top rot 1.3 ± 0.2 0.5 ± 0.1 0.9 ± 0.4 0.6 ± 0.1

a Figures are means from inventories of all grid cells in every stand (n = 8 stands per forest for betweenforest comparisons; n = 4 stands each for thinned or unthinned stands).

b Bold figures indicate significant difference between forests or between thinned and unthinned stands in

Star/Stellar for that attribute (Mann-Whitney U-test [Norus ̆is 1993]; p < 0.05).
55

Fungi

Figure 10—Seasonal changes in percentage of use of supplemental den

sites, by forest, from winter (W) 1993 through summer (S) 1996. Dens

were cleaned of all nest material during winter 1995. An additional eight

nestboxes per den-augmented stand (32 boxes per forest) were added in

December 1995. Figures represent percentage of use of 64 possible (SS

and FH cavities and nestboxes, W93-S95, and SS/FH cavities, W96 and

S96) or 96 possible (SS and FH nestboxes, W96 and S96).

and Kindbergia oregana), with some shredded cambium, fronds of brackenfern, and

fruticose lichen (Usnea sp.). By March 1996, 214 sciurids had been found in supplemental dens--209 in nestboxes and 5 in cavities. Of these, 72 percent (154) were in

FH, and 28 percent (60) were in SS. Flying squirrels (94 in FH vs. 30 in SS), Douglas’

squirrels (57 in FH vs. 30 in SS), and Townsend’s chipmunks (3 in FH vs. 0 in SS)

were all found more frequently in FH than in SS.

Epigeous fungi—Sixty-four mushroom species were found during sampling of SS and

FH forests in October and November 1992; 37 in SS (19 mycorrhizal) and 44 in FH (15

mycorrhizal). Over 260 species34 of fungi (124 mycorrhizal) were observed on permanent

plots during fall sampling periods in 1994, 1995, and 1996 (appendix 1). Epigeous

fungal communities in both forests were dominated by the ectomycorrhizal genera

Russula, Cantharellus, Cortinarius, Inocybe, Laccaria, Lactarius, Boletus, and

Amanita, and the saprobic genera Clitocybe, Collybia, Hypholoma, and Mycena.

34 Some of what we refer to as species are actually groups

containing one or more similar taxa. Further investigation into

these groups (including such difficult genera as Inocybe,

Cortinarius, and Russula) will undoubtedly expand the number of

well-circumscribed taxa. As such, our current estimate of epigeous

fungal species richness is considered to be conservative.

W93 S93 W94 S94 W95 S95 W96 S96

Season and year

0

16

32

48

64

80

96

Dens used (percent)

SS Cavities FH Cavities

SS Nestboxes FH Nestboxes
56

Notwithstanding substantial year-to-year variation in ectomycorrhizal species richness,

average ectomycorrhizal species richness per plot was significantly greater in SS than

in FH for all years and categories except for control plots in 1994. Total ectomycorrhizal

species richness was greater in SS than in FH for each year and for all years combined (table 19). Average ectomycorrhizal species richness per plot was significantly

greater in control plots than in thinned plots in each forest and year except in SS in

1994.35 Of 78 ectomycorrhizal taxa found in all years combined in FH, 16 were not

found in SS. Of 108 ectomycorrhizal taxa found during all years combined in SS, 45

were not found in FH. Of those 45, 28 were in the genus Cortinarius. On the FES

study sites, the genus Cortinarius was represented by 40 or more distinct taxa—about

one-third of the total ectomycorrhizal species richness we have observed to date.

Table 19—Mean mycorrhizal mushroom species per permanent vegetation plot

by forest and year for all plots, control plots, and all thinned plots, and control

stand and total mycorrhizal species richness by forest and year

Star/Stellara b Farley/Hill

Species Species

richness richness

total/ All Control Thinned total/ All Control Thinned

Year controlc plots plotsd plots controlc plots plotsd plots

No. No. species ± S.E. No. No. species ± S.E.

species -------(sample size)------- species -------(sample size)-------

1994 37/26 2.7b 4.0 2.3 30/20 1.3 2.1 1.1

± 0.4 ± 1.1 ± 0.3 ± 0.2 ± 0.4** ± 0.2

(63) (15) (48) (64) (16) (48)

1995 65/49 4.8 7.9 3.8 55/39 3.3 4.8 2.9

± 0.5 ± 1.4** ± 0.4 ± 0.4 ± 0.6** ± 0.4

(63) (16) (47) (64) (16) (48)

1996 86/65 8.9 12.7 7.6 56/45 5.3 8.0 4.3

± 0.6 ± 0.9* ± 0.6 ± 0.4 ± 0.7* ± 0.4

(79) (20) (59) (81) (22) (59)

All 108/89 5.7 8.6 4.8 78/65 3.5 5.3 2.8

± 0.3 ± 0.8* ± 0.3 ± 0.2 ± 0.5* ± 0.2

(205) (51) (154) (210) (54) (156)

a Bold figures indicate significant differences between forests (t-test, p < 0.05).

b Underlined figures indicate significant differences between forests (t-test, p < 0.10).

c The number of mycorrhizal species found in all sampled plots (total) or control-only plots.

d Control and thinned plots for a forest and year or years differ significantly (t-test): * = p < 0.01;

** = p < 0.05).

35 1994 was a low fungal production year. Though average species

richness values in SS CT plots were substantially greater than for

CT plots in FH and for SS thinned plots, the inherent variability and

relatively small sample size precluded detection of significant

differences.
57

Hypogeous fungi: in situ and in fecal pellets—Overall species richness and number

of species unique to a particular forest (SS or FH) were similar in each forest. Thirtyseven species were found in FH; 16 of those were found only in FH. Thirty-five

species were found in SS; 16 only there. Of the 51 identified species of hypogeous

fungi, 4 to 6 species and 2 genera were undescribed previously (appendix 1). Truffles

were found on 15 percent (550 of 3,680) of truffle plots (Colgan 1997). The most

frequently encountered genera were Rhizopogon, Melanogaster, Hysterangium,

Endogone, and Leucogaster.

Analysis of flying squirrel fecal pellets collected during the 1991-94 arboreal trapping

sessions showed that the genera Rhizopogon, Gautieria, Leucogaster, and Melanogaster

were common in squirrel diets in both forests, with Rhizopogon sp. being the most

frequently encountered spore.36 Gautieria sp. was more commonly consumed in the

SS forest, and Melanogaster spp. were more commonly consumed in the FH forest.

Fecal pellets from both forests also commonly contained spores of 10 to 12 other

truffle genera, as well as amorphous plant material (not identified), and fungal spores

of the mushroom genera Russula and Peziza and the families Agaricaceae and

Boletaceae. All genera identified in fecal pellets also were collected during truffle

sampling. Truffles were the primary food for flying squirrels throughout the year in the

study area, with seasonal variation in taxa consumed.

The FES has two aspects: retrospective and prospective or experimental. The retrospective portion determines how, and to what extent, forest vertical and horizontal

structure and plant, animal, and fungal species composition differ between two forests

with widely different management histories. The experimental component seeks to

answer questions about how such forests will respond to thinning treatments designed

to accelerate forest development. Here, we summarize our preliminary, tentative

conclusions from these two views.

Vegetation—Both forests were established after large-scale clearcutting in the 1920s

and 1930s. Their subsequent management practices have differed substantially,

though, with FH having intensive management similar to many other managed forests

in western Washington and Oregon, and SS having retained legacies but receiving

little or no subsequent intervention. Thus the differences we observed between the

two forests may illustrate long-term, but poorly understood, management-induced

changes relevant to other managed west-side forests.

The Farley/Hill forest developed a substantially richer and more complex vascular

plant community than SS, with more species and greater abundances (cover and

volume) in all subcanopy vegetation layers (e.g., forbs and ferns, low shrubs, and

midstory shrubs and low trees). Control stands in FH commonly contained weed

species not found in SS (table 9), with introduced species such as Hypochaeris

Retrospective Aspects

Discussion

36 This spore type also includes the closely related genera Alpova

and Truncocolumella, spores of which are indistinguishable from

Rhizopogon (Castellano and others 1989). In western Washington,

however, the common species of Alpova is associated only with

alder, and Truncocolumella has been collected only in late summer

and fall, and only in SS. Thus, we refer to all these spore types as

Rhizopogon sp.
58

radicata, Ilex aquifolium, and Rubus laciniata well established in these systematically

thinned stands. Other alien species, including Rubus discolor, Mycelis muralis,

Senecio jacobea, and S. sylvaticus also were found in FH, but not SS,37 control stands.

Graminoids were almost completely absent from the closed-canopy SS control stands

(two native grass taxa, three occurrences on 24 plots), but the more open conditions in

FH control stands supported a graminoid flora of nine species (eight of which were

native) occurring 28 times.

Another group, the mycotrophic vascular plants (including members of the families

Orchidaceae and Ericaceae sensu lato), were represented by more species and were

more frequently encountered in SS than in FH. Members of the genera Chimaphila,

Coralorrhiza, Goodyera, Monotropa, and Pyrola were rarely encountered in FH control

stands (3 species, 6 occurrences) compared to 6 species and 23 occurrences in SS

control stands.

Salal was the dominant understory woody component in both forests; low shrub cover

in control stands was about 50 percent in FH (60 percent of that salal), and 25 percent

in SS (80 percent salal). Farley/Hill also had higher species richness in low-shrub

and mid-shrub-understory tree layers (Carey and others 1996a). Low stocking and

thinnings in FH stands also produced significantly larger diameter trees than in SS,

where suppression was uniformly evident.

In contrast to the uniform canopy (and understory) that are produced by light and

systematic thinnings, VDT generated a mosaic of overstory and understory conditions,

thus creating an admixture of habitats. In so doing, VDT increased stand structural

and biological diversity by ensuring habitat availability for most plant species and

species groups.

Coarse woody debris and snags—Intensive management of FH reduced CWD to

exceptionally low levels. Retention of legacies provided moderate levels of CWD in

SS. This unplanned experiment and previous research are helping us understand the

role of CWD.

Coarse woody debris is an important correlate of small mammal abundance in the

Pacific Northwest (Carey and Johnson 1995), mycorrhizal colonization and truffle

abundance (Amaranthus and others 1994), and diversity of mycorrhizal fungi in flying

squirrel diets (Carey, unpublished data). Vertebrates and invertebrates that inhabit

and consume coarse woody debris and snags carry decomposer bacteria and fungi as

well as mycorrhizal fungi (Harmon and others 1986, Li and others 1986). Other

organisms seek shelter, nests, or forage in these woody structures. Wood-boring

beetles enter dead wood and while doing so, inoculate wood with a rich assemblage of

fungi, bacteria, protozoans, and nematodes; different functional groups of insects also

carry different assemblages of decomposer organisms, many of which are symbiotic

37 The only weed species found in SS control stands was Ilex

aquifolium. Ilex is bird dispersed, very shade tolerant, and widely

established in the understory of western Washington forests

(Thysell 1997).
59

with their specific insect host (Schowalter and others 1992). Numerous other predators, parasites, and fungal-feeding insects can then enter CWD via insect galleries,

and in doing so, disperse their own complement of fungi, bacteria, protozoans, nematodes, and microarthropods (Harmon and others 1986, Schowalter and others 1992).

Small mammals and birds also are drawn to such woody structures for protection or

for foraging (for insects, fungi, and seeds), and while there deposit fecal pellets

containing mycorrhizal fungal spores (Harmon and others 1986; Maser and others

1985, 1986), other microorganisms (Li and others 1986), or seeds. Vascular plants,

such as red huckleberry and western hemlock, mosses, liverworts and lichens (e.g.,

Tetraphis pellucida Hedw., Lepidozia reptans (L.) Dum., Cladonia spp.38), and mycorrhizal and decomposer fungi (e.g., Piloderma fallax, Inocybe lanuginosa, Pleurotus

ostreatus), establish preferentially or are found nearly exclusively on woody debris

(Arora 1986, Christy and Mack 1984,39 Hitchcock and Cronquist 1973, Maser and

Trappe 1984, Vitt and others 1988). Thus, CWD plays important, though incompletely

understood, roles in the maintenance of biodiversity40 in managed forests. Preliminary

results from the FES are suggesting a degree of substitutability of understory vegetation for CWD in the arthropod (Burgess 1997), fungal, and forest floor mammal communities, but not for flying squirrels; for example, understory vegetation provides cover

from predation, seasonal contributions of organic matter from the foliage, and cooler

microclimate through shading the forest floor. Fruits and seeds also may substitute for

CWD-inhabiting fungi in small mammal diets.

Both large and small snags were more abundant in SS than in FH; abundant smalld.b.h. snags in SS were a result of stem exclusion and of root rot infestations. Many

small d.b.h. trees in SS were suppressed and dying; these structures, however, are of

little value to cavity-nesting wildlife (Carey and others 1997). Residual large snags in

SS were a legacy left from the preceding stand. Residual snags and other den sites of

northern flying squirrels (especially live trees) have been suggested as a limiting factor

for flying squirrels and other arboreal mammals in managed stands (Carey 1995).

These snags contribute not only to abundance of dens for arboreal rodents and cavity

38 Names and authorities are from Vitt and others (1988) (lichen,

liverwort), and Schofield (1992) (moss).

39 They found 98 percent of western hemlock seedlings in an oldgrowth Douglas-fir/western hemlock forest were rooted on rotten

wood, which covered only 6 percent of the forest floor.

40 Fungi and insects are two of the most species-rich groups of

organisms in the world, with new insect and fungal taxa being

regularly found in the conifer-dominated forests of the Pacific

Northwest. Hundreds of species of insects are associated with

fallen Douglas-fir, many of which may be undescribed (Deyrup

1981, Marra and Edmonds 1995); numerous undescribed truffle

taxa have been found during the course of the FES project. Many

more undescribed truffles and other mycorrhizal fungi also have

been collected in the Pacific Northwest (Trappe 1996). Insects

frequently harbor obligate mutualist fungal symbionts, parasites,

or commensals; these fungal groups are very poorly studied

(Hawksworth 1991). Thus insects and fungi, many found on or

in standing or down coarse woody debris in temperate forests,

contribute greatly to overall biodiversity as well as to crucial forest

processes (Parks and Shaw 1996).
60

nesting birds but also to roost sites for raptors, recruitment of large CWD, and to

increased biological, structural, and functional diversity within the forest (Parks and

Shaw 1996). Many residual snags in our study area, however, were well-decayed,

short structures of marginal value to cavity-using wildlife (Carey and others 1991b,

1997).

Arboreal and forest floor mammals—Small mammal communities in our study were

composed of the same characteristic species but had a different structure than those

reported by Carey and Johnson (1995) in their analysis of managed and old-growth

forests of the Olympic Peninsula. In particular, FH and SS communities were dominated by fewer species, showed a reversal of relative abundances of Columbian mice

and deer mice, and SS contained fewer creeping voles when compared to managed

stands on the Olympic Peninsula.

Carey and Johnson (1995) found prevalence of herbaceous cover and shrub abundance (with CWD) to be the best predictors, respectively, of creeping vole and deer

mouse abundance in managed stands, with Columbian mice increasing with total

understory vegetation in managed stands. Deer mice and creeping voles accounted

for most of the differences in abundance between FH and SS (table 14). Creeping

voles are primarily foliage eaters, and deer mice are primarily seed and fruit eaters but

will consume large quantities of insects; both are also known to consume truffles and

mushrooms (Carraway and Verts 1985, Maser and others 1978, Rhoades 1986, Van

Horne 1982). As such, both benefit from the increased vascular plant species richness

and cover observed in FH, which are attributable to FH management history. Thus, we

believe the greater abundance of deer mice and Oregon voles in FH to be a further

validation of Carey and Johnson’s (1995) predictors. These same species may be

limited by the paucity of understory plant species richness and cover that characterizes SS and other closed canopy, stem-exclusion phase stands. The rarity of the

Columbian mouse in SS and FH may be related to lack of tall understory development

and lack of tree species diversity. Thus, plant and small mammal biodiversity seem

more likely to be preserved if understory vegetation enhancement and avoidance of

stem-exclusion conditions are explicit goals in managed forests.

The arboreal rodent community, represented by the same three species in both

forests, exhibits taxon-specific differences in abundance that we believe are attributable, in part, to past management practices. The northern flying squirrel is strictly

nocturnal and relies almost entirely on truffles for food in our study sites. We identified

16 truffle genera from squirrel fecal pellets out of the 21 genera identified during truffle

sampling (these 16 genera contain 40 of the 50+ truffle species we have identified to

date), validating the important role flying squirrels have in truffle dispersal. Unlike in

other portions of their range (Maser and others 1985), flying squirrels did not use

lichens as a food source.
61

In SS with its larger component of residual trees and snags (potential den sites), CWD,

and an absence of human disturbance since stand origin, we found a greater abundance of flying squirrels than in FH. Recently, Carey and others (1997) confirmed the

significantly higher availability of potential natural den sites (large trees, snags) in SS

than in FH and a substantially greater use of supplemental dens in FH, thus suggesting a need to proactively manage for such structures if maintenance of flying squirrel

and other sciurid populations is desired.

In contrast, the Townsend’s chipmunk is diurnal and is more general in its diet and use

of dens (Carey 1991a, Maser and others 1981). Townsend’s chipmunks eat seeds

and fruits as well as fungi (truffles and mushrooms) and insects (Maser and others

1981). Food is thought to be the most limiting factor for Townsend’s chipmunks, with

annual differences in conifer seed and fungi production leading to large year-to-year

variance in population density. Their dens are widely variable, being found both

underground in burrows and in trees (Carey 1991a). The greater abundance of

Townsend’s chipmunks in FH is, we believe, primarily a response to the better developed and more diverse understory (Carey 1995) with its greater variety of fruits, nuts,

and seeds, with den availability being less constraining for this species.

Fungi—Overall truffle species richness was similar in FH and SS control stands;

further analysis of truffle data is currently underway by collaborators and will not be

reported herein. Average epigeous mycorrhizal species richness per plot and total

epigeous mycorrhizal species richness were higher in SS control stands than in FH in

every sampled year and for all years combined. In spite of substantial year-to-year

variation in overall and per-plot richness, the consistently higher species richness in

SS suggests that different management practices could be partly responsible for such

between-forest differences. Mycorrhizal and nonmycorrhizal fungi may be strongly

associated with CWD, which is also much less abundant in FH. Many weedy plant

species found in FH are also nonectomycorrhizal; thus, the combination of less

substrate for some fungi (e.g., woody debris) and an abundance of nonectomycorrhizal

host plant species could reduce ectomycorrhizal diversity in such stands.

Vegetation—There was a rapid increase in vascular plant species richness after

experimental thinnings and as a function of thinning intensity within stands. The

increase in species richness after thinning was evident in both forests, with most new

species being found in the lowest stratum (ferns, forbs, trailing vines, herbaceous

annuals and perennials). After thinning, rapidly dispersing and colonizing composites

(family Asteraceae), grasses, and legumes were encountered in both forests, especially in RD2 cells. Increased species richness also resulted from growth of residual

species and of other native forest floor species having the capacity to rapidly establish

by runners, suckering, or by germination from a soil seed bank. Extensive or longterm weed colonization does not appear to be a problem because of the intentionally

small scale of the thinnings and because of our effort to capitalize on preexisting shrub

patches.

Prospective and

Experimental Aspects
62

Conversely, small areas of root rot or of RT treatment in closed-canopy forests provide

sites for ruderals, shade-intolerant species, and other early- to mid-successional

organisms, and so increase forestwide diversity. Similar responses are seen with

other localized tree mortality events and canopy gap formation processes (Spies and

others 1990) that are a part of natural forest vegetative development.

Seedlings of species that contribute to development of understory and subcanopy

layers (e.g., willow, alder, maple, salal, huckleberry, hazelnut, and cascara) were found

in most of the permanent plots in thinned cells (93 percent of plots had one or more

species; 65 percent had three or more species). Gaps such as those created by VDT

may become long-lasting shrub or herb patches, or they may fill with trees (Spies and

others 1990)—either naturally regenerating Douglas-fir or shade-tolerant species, or

planted grand fir, western white pine, western redcedar (of which about 225 total

planted conifer seedlings/ha survive in planted cells), or red alder. In any case,

architectural and spatial complexity has the potential to quickly develop on thinned

sites, resulting in an increase in community-level diversity in variably thinned stands.

Though shrub cover in each forest was substantially less in thinned than in control

stands one year after FES thinnings (table 8), we believe this to be a short-duration

response to VDT thinning disturbances. Recent work by Klinka and others (1996) has

quantified the inverse relation between canopy cover and cover of salal and other

shrubs in similar forests on Vancouver Island. Their work suggests that salal and other

shrub cover on FES sites should increase dramatically in response to thinnings.

Salal is one of a number of woody plants that may form mycorrhizae with some of the

same fungal symbionts as Douglas-fir (Amaranthus and Perry 1994, Largent and

others 1980, Molina and Trappe 1982, Smith and others 199541). If so, salal and other

hosts could facilitate the transition from thinned to restocked stands by stimulating

rapid mycorrhizal recolonization and establishment on Douglas-fir (and other

Pinaceae) in thinned stands where salal is present. The presence of salal, other

potential mycorrhizal host species, and CWD (Amaranthus and others 1994) in thinned

sites may be important because nonmycorrhizal weeds and annuals (Trappe 1987)

could gain competitive advantages if mycorrhizal fungi and other essential rhizosphere

species decline in abundance (Amaranthus and Perry 1994, Perry and others 1989)—

changes that could alter soil food webs and make developing a native plant community

difficult (Amaranthus and Perry 1994, Ingham and Thies 1996). By capitalizing on

preexisting openings with intentionally small-scale thinning units and by retaining

substantial cover of overstory trees, practices such as the VDT thinnings have the

potential to maintain and even accelerate natural forest processes and interactions,

while minimizing the establishment of introduced and noxious weeds.

41 In greenhouse studies, Smith and others (1995) found limited

ectomycorrhizal development on salal of two ectomycorrhizal types

also commonly found on Douglas-fir, but they point out that the

degree to which salal forms ectomycorrhizae in the field is not

known. For some other members of Betulaceae, Ericaceae,

Fagaceae, and Salicaceae, the sharing of ectomycorrhizae with

Douglas-fir and other Pinaceae is clearer (Amaranthus and Perry

1994, Borchers and Perry 1990, Molina and Trappe 1982).
63

Prethinning and early postexperimental thinning vegetative conditions have affected

abundance and size of naturally regenerating Douglas-fir seedlings and saplings in

both forests. In SS control stands, the understory was depauperate in species and

cover. Following VDT operations, tree seedlings (almost all Douglas-fir) quickly

emerged; many seedlings (mostly < 50 cm height) were present in RD2 to RD8 cells.

Emergent tree seedlings in SS experienced much less competition from herbaceous

and shrub species than in FH. The RD6 and RD8 overstories in SS were still very

heavily stocked after thinning and it is unlikely that shade-intolerant Douglas-fir would

survive for long under such dense canopies, as evidenced by the near absence of tall

seedlings.

Although FH stands contained fewer Douglas-fir seedlings and saplings than did SS,

FH had a more equitable distribution of seedlings over the four size classes. This

pattern is consistent with the management history of FH; previous thinning entries in

FH provided opportunities not only for increased herbaceous and shrub growth, but

also for Douglas-fir seedling emergence (hence the bimodal d.b.h. distribution evident

in fig. 7b). Consequently, understory development, especially the abundant shrub

cover, resulted in reduced survivorship of newly emergent Douglas-fir seedlings.

Because of the near absence of appropriate seed sources in both forests, naturally

regenerating shade-tolerant tree seedlings were infrequent.

Because SS RD2 subtreatments were applied to actual root rot areas, shrub cover

there was greater than in the RD2 cells of FH as a result of previous tree mortality and

increased incident light. The presence of such root rot areas (and associated understory) in stands to be managed for both commodities and biodiversity may provide an

opportunity to capitalize on preexisting spatial, structural, and species diversity in stem

exclusion stands.

Fungi—Preliminary data on hypogeous fungi support the importance of maintaining

salal and other native species in place during thinning operations. The two forests

were similar in patterns of truffle species richness and standing crop production;

community structural differences have not yet been evaluated. Intensive management

in FH did not change diversity and abundance of truffles compared to SS. In both

forests, the effect of VDT on the productivity and biodiversity of truffles was limited

temporally and spatially, with even the most aggressive treatments showing only shortterm reduction in truffle standing crop (Colgan and others 1994, Colgan 1997). We

believe this to be a result of the tempering effect of the deliberately small spatial scale

of variable-density thinnings. Such reductions may have been longer lasting if entire

blocks had received RD2 and RD4 treatments. In addition, systematic, heavy

thinnings could have resulted in more widespread and persistent weed populations

and in difficulty in tree regeneration, because of alterations to the soil microbial community (Ingham and Thies 1996, Perry and others 1989).

The FES sites contain substantial ectomycorrhizal species richness, both hypogeous

and epigeous. Trappe (1977) reports Douglas-fir as having 2,000 or more fungal

associates throughout its range, but little guidance is available for predicting total

ectomycorrhizal biodiversity within a stand or more limited geographical region.

Our evidence of 175 or more ectomycorrhizal symbionts in a relatively small area
64

(ca. 200 ha) of even-aged Douglas-fir forests lends credence to the extreme complexity

within ectomycorrhizal guild structure and function (Molina and others 1992), while

also demonstrating the rather remarkable diversity42 to be found in just one organismal

group within a small area of even-aged, managed forest.

Our work indicates at least a moderate-term (10-year) reduction in epigeous

ectomycorrhizal species richness (as evidenced by sporocarp production) after thinning. Species richness values in FH control plots were significantly less than SS CT

values and were about the same as SS thinned stand values. Species richness also

was significantly less in FH thinned plots than in FH CT plots, thereby suggesting the

possibility of cumulative deleterious effects of repeated disturbance on ectomycorrhizal

communities.

In contrast, abundant endomycorrhizal plant host species in heavily thinned sites

should increase the abundance of mycobionts such as Glomus, sporocarps of which

are widely consumed by forest-floor small mammals (Maser and others 1978, McIntire

1984), as well as being used by arboreal rodents (Colgan 1997; Maser and others

1985, 1986). Thus, changes in plant species composition will alter mycorrhizal communities in several ways. Increased abundance of herbaceous vegetation, maples,

western redcedars, and other endomycorrhizal hosts will support Glomus and other VA

mycorrhizae. An increase in the abundance, diversity, and age-class distribution of

ectomycorrhizal hosts, such as willow, poplar, hazelnut, white pine, and grand fir, can

augment ectomycorrhizal diversity within a community otherwise dominated by a

single host species and age class (e.g., Douglas-fir).

Our results tend to support the hypothesis that “disturbing the forest to remove commodities does not necessarily conflict with maintaining spatial and temporal linkages

between plants via ectomycorrhizal fungi” (Amaranthus and Perry 1994). The intentionally small size of our thinning subtreatments (0.16 ha), coupled with the root

grafting evident in these and other (Eis 1972) essentially monospecific Douglas-fir

stands and the protective effect salal and other understory shrubs may have on

ectomycorrhizal fungi (Amaranthus and Perry 1994, Largent and others 1980, Perry

and others 1989, Smith and others 1995), should temper and ameliorate soil microbial

community disturbances that accompany VDT. Thus, VDT should not produce the

long-term alterations in soil structure and function known to be deleterious to tree

establishment and which have occurred in some large clearcuts that were burned or

had herbicides applied before replanting (Ingham and Thies 1996, Perry and others

1989). Indeed, our results suggest that VDT is a management activity that “can mimic

natural disturbance regimes and help assure that ectomycorrhizal linkages among

plants are not severed and that trees and soil ecosystem continue to operate as a

coherent and dynamic unit” (Amaranthus and Perry 1994).

42 Sporocarp diversity typically underestimates total ectomycorrhizal

diversity as seen on the actual root tips, and sporocarp abundance

or dry weight may not accurately reflect actual hyphal biomass

dominance (Molina 1997). As such, ectomycorrhizal diversity is

probably higher than indicated by sporocarp diversity.
65

The FES is the first application of VDT and the first deliberate attempt to alter the

trajectory of managed second-growth stands from homogeneity, simplification, and

wood production to heterogeneity, complexity, and biodiversity, as well as production of

economic goods and services. Because the FES was a first attempt, little guidance

was available on thinning targets or on operational application of VDT. Although

simple in concept, VDT is complex in application. First, it is a departure from traditional thinning in spacing (even vs. varying between patches), residual density (VDT

removes more trees than traditional thinning), and marking (root rot infestations and

existing understory development are capitalized on). Second, because the objective is

to grow large trees and a diverse understory concurrently, care must be exercised to

avoid creating salal brushfields separated from a high canopy (an undesirable condition). Shade-tolerant regeneration must be promoted to fill canopy-to-shrub layer

gaps; thus, judgment must temper density or RD targets. Underplanting may be

necessary to reach forest structure targets. Third, many opportunities exist now in

40- to 70-year-old stands that have an accumulated history of management or lack

thereof. These stands may have risk factors (susceptibility to wind throw, root rot

infestation, insect infestations) that younger stands with consistent density and spacing management do not.

In designing prescriptions for accelerating forest development in second-growth

stands, theoretical targets based on retrospective studies (Carey 1995, Carey and

Johnson 1995), simulations using growth and yield models (Carey and others 1996a),

and models derived from experimental silvicultural plots (Curtis 1970, 1982; Curtis and

Carey 1996) must be adjusted to take into account management history and existing

site conditions. Factors such as tree vigor (e.g., as determined by the ratio of live

crown to total tree height), risk of high winds and degree of wind resistance, presence

of disease (e.g., root rot or insect infestations), and site productivity, especially as it

influences understory development and potential natural vegetation, must be considered. In FH, only light thinning was necessary to further understory development;

limited heavy thinning was necessary to promote shade-tolerant regeneration and

spatial heterogeneity in the stands. Because shade-tolerant natural regeneration

was lacking, underplanting also was necessary. In SS, even heavy thinning was

inadequate to achieve the desired theoretical spacing; however, heavier thinning in

SS would have substantially increased the risk of windthrow. In both forests, multiple

entries will be needed to obtain the desired future condition.

In our application of VDT, we met with Fort Lewis foresters and discussed our goals

and approach in general terms by using density targets conceptually, but stressed the

development of patchy understory. Because we were about to treat stands with

disparate management histories, we asked these foresters, with their substantial

experience (> 15 years) with traditional thinnings on Fort Lewis, to exercise their

professional judgment. Their task was formidable, given the constraint of thinning from

below (subordinate trees), very high stocking levels, obvious root rot infestations, and

lack of understory in SS, and the moderate stocking levels, inconspicuous root rot

Variable-Density

Thinning
66

infestations, and well-developed, low (< 2 m tall) understory in FH. The root rot

treatments, an addition to the VDT concept, would result in substantial addition of light

to adjacent grid cells and create open overstory that could potentially “trap” high winds

during storms.

Though specific cell-by-cell thinning targets (trees per hectare) were not always

reached, the overall goal of creating a mosaic of variably stocked cells while retaining

wind firmness was achieved—few trees blew down during an extreme wind storm in

winter 1995. Many SS cells remained overstocked in relation to target densities, and

some cells in FH were understocked relative to targets. Nonetheless, wide variation in

tree density and basal area was achieved at the cell level in both forests, resulting in

substantial within-stand diversity in habitat conditions (see appendices 4 and 5).

The ranges of RDs in post facto classes were quite large, with root rot cells having the

least variable range of relative density values. Cells selected as root rot subtreatment

areas in the original prescription largely coincided with the lowest tree density (RD2)

cells shown on the stand maps. Remaining cells included a wide variation in retained

tree densities and relative densities (RD3-RD10 in SS; RD2-RD9 in FH). After thinning, each treated stand nonetheless contained an admixture of densities sufficient to

stimulate variable, cell-specific responses in understory and tree development. Variability in vegetation response is both a central goal of the FES and, we believe, a

direct result of initiating the creation of a mosaic canopy in these managed stands.

This variability is already manifested by increases in species richness and cover

values in thinned stands; existing variability will be further capitalized on in future

entries.

Random assignment of subtreatments provided the ability to examine subtreatment

effects (for example, on fruiting of ectomycorrhizal fungi and on soil food webs) but

was not necessary for stand-level analyses. However, irregular patterns of subtreatment

assignment may have made marking more difficult and may have led to difficulty in

interpretation of cell prescriptions.

Several steps could be taken to refine application of VDT. Application could be systematic (fig. 3, c and d), and a repeating pattern (light-light-heavy) could be applied to

consecutive 0.5-acre squares or to strips (but only if strips were perpendicular to

prevailing winds). A stand density measure such as Curtis’ (1982) RD is easy to use

and is perhaps the best guide for applying thinnings; RD is most directly applicable

when density is controlled beginning with stand initiation. Interventions to alter the

developmental pathway of existing stands must consider existing conditions of understory development and forest health in addition to stocking. Application of RD is most

straightforward with regular spacing between trees. Biodiversity pathways43 for forest

management, however, require developing heterogeneity in spacing between small

43 The theoretical and empirical background of such a management

approach is provided in Carey and Curtis (1996) and Carey and

others (1996a). A biodiversity pathway for forest management

involves specific steps intended to conserve biological legacies,

minimize area and time in the competitive-exclusion stage, make

use of extended rotations, minimize site preparation, and make use

of alternative types of thinnings such as VDT.

Operational Aspects

of Variable-Density

Thinning
67

patches (ca. 0.16 ha). Wide spacing in small patches (heavy thins and root rot thins)

changes light conditions (admits more light) in adjacent patches, introducing a confounding effect of variable-density thinning; thus, RD calculated as a weighted average

could be conservative (underestimate reduction of competition among trees) due to

effects of sun-angle (a relatively small proportion of light is received perpendicular to

the canopy). Thus, professional judgment about the many factors potentially influencing stand development must be used in choosing exact RD targets.

Some general rules for multiple-entry VDTs might be (1) thin 30- to 40-year-old stands

(with current RD 8.5 to 10.0 [metric]) to RD 7 by using patches of RD 8.5 and RD 5.75;

(2) thin previously thinned stands to RD 4.3 to 5.0 by using a combination of RD 5.75

and RD 4.3; (3) in all cases, incorporate site-specific information on risk of windthrow,

disease, site quality, vegetation series, stocking, understory vegetation, and advanced

regeneration of shade-tolerant species, while also taking advantage of naturally

occurring openings, groups of snags, or other forest features.

Tree marking is applied to individual trees and is therefore operationally a density

control. But when there was substantial cell-to-cell variation, we found markers tended

to undermark or mark based on a perceived average stand condition (thus overmarking

sparse cells and undermarking the dense cells). Particularly in the more dense cells,

markers tended to undermark generally, probably due to a perception that too many

trees were being marked. Thus, tree marking in heavily stocked stands, especially

with abundant suppressed trees, may be best achieved by conspicuously marking

leave trees, and by basing marking guidelines on relative density as opposed to other

forest parameters. Then, RD can be checked easily by using a wedge prism. In

lesser stocked stands, trees to be cut or trees to be left could be marked. In either

case, consideration of the stand d.b.h. distribution is important, so that understory

trees not a part of the main cohort are not removed or damaged.

Our initial application of VDT has set the trajectory of the stands toward development

of late-seral forest. Star/Stellar will move from competitive exclusion to understory

reinitiation and FH will move from understory reinitiation, to developed understory. The

second VDT entry will be tailored specifically to conditions found in the two forests. In

SS, heavy VDT will be used to further accelerate understory development. In FH,

lighter VDT and coarse woody debris-augmentation will move FH toward the niche

diversification stage (Carey and others 1996a). Both groups of stands will require

additional underplanting of shade-tolerant conifers and, perhaps, seeding of bigleaf

maple to add to midstory development.

As an integrated, multidisciplinary study of accelerating forest development of Douglas-fir

forests in western Washington, the FES seeks to evaluate the extent to which, and the

time frame within which, the development of late-seral attributes can be hastened

through management of young, even-aged stands. By dividing the study between two

forests with different management histories, the study also seeks to evaluate effects of

past forest practices on forest communities, structures, and processes in forests not

far removed temporally from pre-European-settlement conditions. Our studies indicate

substantial differences between the two forests in terms of their overall biodiversity,

CWD cover and snag density, tree growth, invasive species establishment, and

Summary
68

vegetative and mammalian community composition. Our studies also suggest that

both forests contain much of the documented fungal, plant, and small mammal diversity found in later-seral stands (Carey 1989, 1995; Carey and Johnson 1995; Ruggiero

and others 1991; Thomas and others 1993). Variable-density thinnings that aim to

recreate the patchiness of late-seral forests when applied to stem-exclusion forests

are an opportunity to manage for biodiversity, prevent stand degradation, and accelerate forest development (Carey and others 1996a, Carey and Johnson 1995).

Our choices of experimental treatments and response variables were developed to

evaluate ecosystem processes that include photosynthetically and fungally supported

trophic pathways. Mycorrhizal fungi comprise a large portion of the fungal community

in forest soils (Allen and Allen 1992) and assimilate large amounts of net primary

productivity in Douglas-fir forests (Fogel and Hunt 1983), while enhancing primary

productivity by augmenting plant uptake of water and minerals, improving resistance to

drought, and protecting against pathogens (Molina and Amaranthus 1991). Fungi

(both mycorrhizal and saprobic) are food for a large diversity of consumers including

arthropods, nematodes, and mammals (Perry and others 1989; Thysell and others

1997). Fungal sporocarps are the primary food for northern flying squirrels (Carey

1995; Carey and others 1992, 1996b) and are widely consumed by numerous other

small mammals as well (Carey 1991a, Carey and Johnson 1995, Maser and others

1981). These small mammals, in turn, are important prey for predators, including the

northern spotted owl, the northern goshawk, mustelids, and other animals at the

highest trophic level.

The coniferous overstory species and other ectomycorrhizal understory species are

hypothesized to “preserve ectomycorrhizal fungi during periods of rapidly changing

above ground community structure and that mycorrhizal links between hardwoods and

conifers facilitate conifer establishment by providing a ready source of inoculum,

nutrients, and water” (Amaranthus and Perry 1994). Natural ecosystems, driven by

photosynthetic and fungal processes and symbioses, maintain a “balanced, stable

disequilibrium through the buffering capacity of biodiversity” (Perry and others 1989).

We now recognize that a great deal of biodiversity is to be found within the plant,

fungal, microbial, and animal communities and that these communities are highly

interdependent. The various communities are associated with one another via numerous functional connections and pathways that include crucial forest processes such as

(1) response to changing soil arthropod populations by forest insectivores; (2) diaspore

dispersal by sciurids, forest floor mammals, and arthropods; (3) decay and recycling of

recalcitrant organic compounds; (4) mutualisms resulting in enhanced mineral and

water use, carbon storage, and ultimately, fitness; and (5) fungal- and photosyntheticbased trophic pathways supporting large numbers of herbivores, fungivores, and their

predators (Carey and others 1996b). The species-, physiological-, and geneticdiversity of the plant-fungal-microbial mutualisms (along with their predators and

consumers), and the redundancy in shared rhizosphere flora, should stabilize the

plant-soil system (Perry and others 1990), and provide numerous benefits to plants,

soil microorganisms, and soil structure (Molina and Amaranthus 1991). Retention of
69

such linkages and mutualisms (Amaranthus and Perry 1994, Molina and Amaranthus

1991) and protection of soil through the maintenance of vigorous tree and shrub cover

(Perry and others 1990) should be an explicit goal during any planned management

disturbance.

We believe these functions, relations, and links can be maintained and enhanced in

managed forests, and that “many of these functions can be served simultaneously as

well as sustainably” (Myers 1995) through proactive and innovative management

approaches, such as those we are testing with the Forest Ecosystem Study. By taking

such approaches, we believe long-term site productivity can be protected; plant,

fungal, and small mammal and predator populations can be maintained; establishment

of invasive or noxious weeds can be minimized; threatened species conflicts can be

avoided; development of late-seral forest and landscape conditions can be hastened;

and future management options can be preserved.

This research was funded by the Pacific Northwest Research Station, U.S. Department

of Agriculture Forest Service; the Environmental and Natural Resources Division,

Department of Army, Fort Lewis; and the U.S. Department of Agriculture National

Research Initiative. We thank the U.S. Department of Army, Fort Lewis Military

Reservation, for assistance in selection of the study sites and long-term commitment

to the study. We thank personnel of the Environmental and Natural Resources Division, and past and present members of the Ecological Foundations of Biodiversity

Team of the Pacific Northwest Research Station, for invaluable assistance in the study.

We acknowledge the following major contributors to the Forest Ecosystem Study,

without whom specific projects would have been impossible: K. Berdinka (coarse

woody debris and epigeous fungi); W. Colgan (truffles); K. Glos (flying squirrel reproductive biology); M. Holmes (soil food webs); E. Ingham (soil food webs); C. Parks

(top rot fungi inoculation); T. Thomas (FES project planning and silvicultural prescription);

J. Trappe (truffles); S. Wilson (small mammals); T. Wilson (arboreal mammals,

telemetry); and A. Wolff (den site inventory). We also thank The Evergreen State

College, Olympia, Washington, for their continued and ongoing support in the form

of more than 130 interns, volunteers, and supportive faculty members. We thank

R. Molina, J. M. Trappe, and L. Bednar for their valuable review comments.

1 inch (in) = 2.54 centimeters

1 foot (ft) = 0.3048 meter

1 square foot (ft2

) = 0.09290 square meter

1 acre = 0.40469 hectare

1 square foot per acre (ft2/acre) = 0.22957 square meters per acre

1 mile (mi) = 1.6093 kilometer

1,000 board feet (MBF) per acre = 5.808 cubic meters per hectare

1 meter (m) = 3.28084 feet

1 square meter (m2) = 10.764 square feet

Acknowledgments

Metric and English

Conversions
70

1 hectare (ha) = 2.471 acres

1 board foot (BF) = 0.00236 cubic meter

1 kilometer (km) = 0.6214 mile

1 millimeter (mm) = 0.04 inch

1 centimeter (cm) = 0.39 inch

Celsius (°C) = (°F - 32) ÷ 1.8

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80

Scientific name1 Code2 Common name3

Vascular Plants:

Abies grandis (Dougl. Ex D. Don) Lindl. ABGR Grand fir

Acer circinatum Pursh ACCI Vine maple

Acer macrophyllum Pursh ACMA3 Big-leafed maple

Achlys triphylla (Sm.) DC.4 ACTR Vanillaleaf

Actaea rubra (Ait.) Willd. ACRU2 Baneberry

Adenocaulon bicolor Hook.4 ADBI Trail-plant

Adiantum aleuticum (Rupr.) Paris ADAL Maidenhair fern

 (=A. pedatum L.)4

Agrostis capillaris L. (= A. tenuis Sibth.) AGCA5 Colonial bentgrass

Agrostis exarata Trin. AGEX Spike bentgrass

Aira caryophyllea L. AICA Silver hairgrass

Aira praecox L. AIPR Little hairgrass

Alnus rubra Bong. ALRU2 Red alder

Amelanchier alnifolia (Nutt.) Nutt. ex AMAL2 Western serviceberry

 M. Roemer

Anaphalis margaritacea (L.) Benth. & ANMA Pearly-everlasting

 Hook. f.

Anemone deltoidea Hook.4 ANDE3 Western anemone

Anthoxanthum odoratum L. ANOD Sweet vernalgrass

Arbutus menziesii Pursh ARME Madrone

Arctostaphyllos uva-ursi (L.) Spreng. ARUV Kinninnick

Arrhenatherum elatius (L.) J. & K. Presl AREL3 Tall oatgrass

Artemisia suksdorfii Piper ARSU4 Suksdorf’s sagebrush

Asarum caudatum Lindl.4 ASCA2 Wild ginger

Athyrium filix-femina (L.) Roth ATFI Lady-fern

Blechnum spicant (L.) Roth BLSP Deer-fern

Bromus vulgaris (Hook.) Shear BRVU Columbia brome

Campanula scouleri Hook. ex A. DC. CASC7 Scouler’s harebell

Cardamine pensylvanica Muhl. ex Willd. CAPE3 Pacific bittercress

Carex deweyana Schwein. CADE9 Dewey’s sedge

Appendix 1

Forest Ecosystem

Study Species List

Footnotes are on p. 102.
81

Scientific name1 Code2 Common name3

Carex pachystachya Cham. ex Steud. CAPA14 Thick-headed sedge

Cerastium fontanum Baumg. ssp. CEFOV2 Common chickweed

 vulgare (Hartman) Greuter & Burdet

 (= C. vulgatum L.)

Chimaphila menziesii (R. Br. ex D. Don) CHME Pipsissewa

 Spreng.4

Chimaphila umbellata (L.) W. Bart.4 CHUM Common pipsissewa

Circaea alpina L. CIAL Enchanter’s nightshade

Cirsium arvense (L.) Scop. CIAR4 Canada thistle

Cirsium brevistylum Cronq. CIBR2 Short-styled thistle

Cirsium vulgare (Savi) Ten. CIVU Bull thistle

Claytonia perfoliata Donn ex Willd. ssp. CLPEP Miner’s lettuce

 perfoliata (= Montia perfoliata (Donn.)

 Howell)

Claytonia sibirica L. var. sibirica

 (= Montia siberica (L.) Howell) CLSIS Candyflower

Collomia heterophylla Dougl. ex Hook. COHE2 Varied-leafed collomia

Conyza canadensis (L.) Cronq. COCA5 Canada fleabane

Corallorhiza maculata (Raf.) Raf.4 COMA4 Spotted coral-root

Cornus nuttallii Audubon ex Torr. & Gray CONU4 Pacific dogwood

Corylus cornuta Marsh. var. californica COCOC California hazel

 (A. DC.) Sharp

Crepis capillaris (L.) Wallr. CRCA3 Smooth hawksbeard

Cytisus scoparius (L.) Link CYSC4 Scot’s broom

Deschampsia elongata (Hook.) Munro DEEL Slender hairgrass

Dicentra formosa (Andr.) Walp. DIFO Pacific bleedingheart

Digitalis purpurea L. DIPU Foxglove

Disporum hookeri (Torr.) Nichols.4 DIHO3 Hooker’s fairy-bell

Dryopteris campyloptera Clarkson4 DRCA3 Spreading woodfern

 (= D. austriaca (Jacq.) Woynar)

Elymus glaucus Buckl. ssp. glaucus ELGLG Blue wild-rye

Epilobium angustifolium L. EPAN2 Fireweed
82

Scientific name1 Code2 Common name3

Epilobium glaberrimum Barbey ssp. EPGLG Smooth willow-herb

 glaberrimum

Epilobium minutum Lindl. ex Lehm. EPMI Small-flowered willow-herb

Erechtites minima (Poir.) DC. ERMI6 Toothed coast burnweed

Festuca occidentalis Hook. FEOC Western fescue

Festuca subulata Trin. FESU Bearded fescue

Festuca subuliflora Scribn. FESU2 Coast Range fescue

Fragaria crinita Rydb. (= F. vesca L.var. FRCR Pacific strawberry

 crinita (Rydb.) C.L. Hitchc.)

Fragaria virginiana Duchesne ssp. FRVIP2 Broadpetal strawberry

 platypetala (Rydb.) Staudt

Frangula purshiana (DC.) Cooper FRPU7 Pursh’s buckthorn

 (= Rhamnus purshiana DC.)

Galium aparine L. GAAP2 Cleavers

Galium triflorum Michx. GATR3 Sweetscented bedstraw

Gamochaeta purpurea (L.) Cabrera GAPU3 Spoonleaf purple

 (= Gnaphalium purpureum L.) everlasting

Gaultheria shallon Pursh GASH Salal

Geranium robertianum L. GERO Herb Robert

Geum macrophyllum Willd.var. GEMAM Oregon avens

 macrophyllum

Gnaphalium microcephalum Nutt. GNMI Slender cudweed

Goodyera oblongifolia Raf.4 GOOB2 Rattlesnake-plantain

Hieracium albiflorum Hook. HIAL2 White-flowered hawkweed

Holcus lanatus L. HOLA Common velvet-grass

Holodiscus discolor (Pursh) Maxim. HODI Oceanspray

Hypericum perforatum L. HYPE Klamath weed

Hypochaeris radicata L. HYRA3 Spotted cats-ear

Ilex aquifolium L. ILAQ80 English holly

Juncus bufonius L. JUBU Toad rush

Juncus effusus L. var. gracilis Hook. JUEFG Common rush

Juncus tenuis Willd. JUTE Slender rush
83

Scientific name1 Code2 Common name3

Lactuca biennis (Moench) Fern. LABI Tall blue lettuce

Lapsana communis L. LACO3 Nipplewort

Lathyrus polyphyllus Nutt.4 LAPO3 Leafy peavine

Leucanthemum vulgare Lam. LEVU Oxeye-daisy

 (= Chrysanthemum leucanthemum L.)

Lilium columbianum hort. ex Baker LICO Tiger lilly

Linnaea borealis L. LIBO3 Twinflower

Listera caurina Piper 4 LICA10 Western twayblade

Lonicera ciliosa (Pursh) Poir. ex DC. LOCI3 Trumpet honeysuckle

Lonicera hispidula (Lindl.) Dougl. LOHI2 Hairy honeysuckle

 ex Torr. & Gray

Lotus corniculatus L. LOCO6 Birdsfoot-trefoil

Lotus micranthus Benth. LOMI Small-flowered deervetch

Lupinus rivularis Dougl. ex Lindl. LURI Stream lupine

Luzula campestris (L.) DC. LUCA2 Woodrush

Luzula parviflora (Ehrh.) Desv. LUPA4 Small-flowered woodrush

Mahonia aquifolium (Pursh) Nutt. MAAQ2 Tall Oregongrape

 (= Berberis aquifolium Pursh)

Mahonia nervosa (Pursh) Nutt. MANE2 Cascade Oregongrape

 (= Berberis nervosa Pursh)

Maianthemum stellatum (L.) Link4 MAST4 Starry false Solomon’s seal

 (= Smilacina stellata (L.) Desf.)

Melica subulata (Griseb.) Scribn.4 MESU Alaska oniongrass

Mentha canadensis L. MECA7 Field mint

Moehringia macrophylla (Hook.) Fenzl MOMA3 Big-leafed sandwort

 (= Arenaria macrophylla Hook.)

Monotropa uniflora L.4 MOUN3 Indian-pipe

Mycelis muralis (L.) Dumort. MYMU Wall lettuce

 (= Lactuca muralis (L.) Fresen.)

Nemophila parviflora Dougl. ex Benth. NEPA Small-flowered nemophila

Oemlaria cerasiformis OECE Indian plum

 (Torr. & Gray ex Hook. & Arn.) Landon
84

Scientific name1 Code2 Common name3

Osmorhiza berteroi DC. OSBE Sweet-cicely

 (= O. chilensis H. & A.)

Petasites frigidus (L.) Fries PEFRP Coltsfoot

 var. palmatus (Ait.) Cronq.

Phalaris arundinacea L. PHAR3 Reed canarygrass

Philadelphus lewisii Pursh PHPR3 Mockorange

Phleum pratense L. PHLE3 Timothy

Pinus monticola Dougl. ex D. Don PIMO3 White pine

Pityopus californicus (Eastw.) H.F. Copel.4 5 PICA9 California pine-foot

Plantago lanceolata L. PLLA English plantain

Plantago major L. PLMA2 Common plantain

Polypodium glycyrrhiza D.C. Eat. POGL8 Licorice fern

Polystichum munitum (Kaulfuss) K. Presl POMU Swordfern

Populus balsamifera L. ssp. trichocarpa POBAT Black cottonwood

 (Torr. & Gray ex Hook.) Brayshaw

 (= P. trichocarpa T. & G.)

Prunella vulgaris L. PRVU Self-heal

Pseudotsuga menziesii PSMEM Douglas-fir

 (Mirbel) Franco var. menziesii

Pteridium aquilinum (L.) Kuhn. PTAQ Brackenfern

Pyrola asarifolia Michx.4 PYAS Common pink wintergreen

Pyrola picta Sm.4 (includes P. aphylla) PYPI2 White vein pyrola

Quercus garryana Dougl. ex Hook. QUGA4 Oregon white oak

Ranunculus occidentalis Nutt. RAOC Western buttercup

Ranunculus repens L. var. repens RARER Creeping buttercup

Ranunculus uncinatus D. Don ex G. Don RAUNP Little buttercup

 var. parviflorus (Torr.) L. Benson

Ribes sanguineum Pursh RISA Red currant

Rosa gymnocarpa Nutt. ROGY Baldhip rose

Rubus discolor Weihe & Nees RUDI2 Himalayan blackberry

Rubus laciniatus Willd. RULA Evergreen blackberry
85

Scientific name1 Code2 Common name3

Rubus leucodermis RULE Blackcap

 Dougl. ex Torr. & Gray

Rubus parviflorus Nutt. RUPA Thimbleberry

Rubus spectabilis Pursh RUSP Salmonberry

Rubus ursinus Cham. & Schlecht. RUUR California blackberry

Rumex acetosella L. RUAC3 Sheep sorel

Rumex crispus L. RUCR Curly dock

Salix scouleriana Barratt ex Hook. SASC Scouler willow

Sambucus racemosa L. SARA2 Red elderberry

Satureja douglasii (Benth.) Briq.4 SADO5 Yerba buena

Senecio jacobaea L. SEJA Tansy ragwort

Senecio sylvaticus L. SESY Wood groundsel

Solanum dulcamera L. SODU Bittersweet

Solidago canadensis L. var. salebrosa SOCAS Canadian goldenrod

 (Piper) M. E. Jones

Sonchus asper (L.) Hill SOAS Prickly sow-thistle

Sonchus oleraceus L. SOOL Common sow-thistle

Spiraea douglasii Hook. var. douglasii SPDOD Douglas’ spiraea

Stellaria crispa Cham. & Schlecht. STCR2 Crisped starwort

Stellaria longipes Goldie STLO2 Longstalk starwort

Symphoricarpos albus (L.) Blake SYAL Common snowberry

Symphoricarpos hesperius G. N. Jones SYHE Creeping snowberry

 (= S. mollis Nutt. ssp. hesperius

 (G.N. Jones) Abrams ex Ferris)

Taxus brevifolia Nutt.4 TABR2 Pacific yew

Thuja plicata Donn ex D. Don 4 THPL Western redcedar

Tiarella trifoliata L.4 TITR Foamflower

Trientalis borealis Raf. ssp. latifolia TRBOL Western starflower

 (Hook.) Hulten (= T. latifolia Hook.)

Trifolium campestre Schreb. TRCA5 Hop clover

 (= T. procumbens L.)

Trifolium pratense L. TRPR2 Red clover
86

Scientific name1 Code2 Common name3

Trifolium repens L. TRRE3 White clover

Trillium ovatum Pursh4 TROV2 Western wake-robin

Trisetum cernuum Trin. TRCE2 Nodding trisetum

Tsuga heterophylla (Raf.) Sarg. TSHE Western hemlock

Urtica dioica L. URDI Stinging nettle

Vaccinium ovatum Pursh VAOV2 Evergreen huckleberry

Vaccinium parvifolium Sm.4 VAPA Red huckleberry

Vancouveria hexandra (Hook.) VAHE Inside-out flower

 Morr. & Dcne.4

Vicia americana Muhl. ex Willd. VIAM American vetch

Vicia nigricans Hook. & Arn. ssp. VINIG Giant vetch

 gigantea (Hook.) Lassetter & Gunn.

 (= V. gigantea Hook.)

Viola glabella Nutt.4 VIGL Stream violet

Viola howellii Gray VIHO Howell’s violet

Viola sempervirens Greene VISE3 Evergreen violet

Amphibians and reptiles:

Mole salamanders—

Ambystoma gracile (Baird, 1859) Northwestern salamander

Ambystoma macrodactylum Baird, 1849 Long-toed salamander

Lungless salamanders—

Ensatina escholtzii Gray, 1850 Ensatina

Plethodon vehiculum (Cooper, 1860) Western red-backed

 salamander

Frogs and toads—

Bufo boreas Baird & Girard, 1852 Western toad

Pseudacris regilla Baird & Girard, 1852 Pacific tree frog

Rana aurora Baird & Girard, 1852 Red-legged frog

Rana catesbeiana Shaw, 1802 Bullfrog

Newt—

Taricha granulosa (Skilton, 1849) Rough-skinned newt
87

Scientific name1 Common name3

Lizard—

Gerrhonotus coeruleus Wiegmann, 1828 Northern alligator lizard

Snakes—

Charina bottae (Blainville, 1835) Rubber boa

Thamnophis ordinoides (Baird & Girard, 1852) Northwestern garter snake

Mammals:

Marsupial—

Didelphis virginiana Kerr, 1792 Virginia opossum

Insectivores—

Neurotrichus gibbsii (Baird, 1858) Shrew mole

Scapanus orarius True, 1896 Coast mole

Scapanus townsendii (Bachman, 1839) Townsend’s mole

Sorex bendirii (Merriam, 1884) Pacific marsh shrew

Sorex monticolus Merriam, 1890 Montane shrew

Sorex trowbridgii Baird, 1858 Trowbridge’s shrew

Sorex vagrans Baird, 1858 Vagrant shrew

Bats—

Myotis californicus (Audubon & Bachman, 1842) California myotis

Plecotus townsendii Cooper, 1837 Townsend’s big-eared bat

Rabbits—

Lepus americanus Erxleben, 1777 Snowshoe hare

Sylvilagus floridanus (J.A. Allen, 1890) Eastern cottontail

Rodents—

Aplodonia rufa (Rafinesque, 1817) Mountain beaver

Erethizon dorsatum (Linnaeus, 1758) Porcupine

Squirrels

Glaucomys sabrinus (Shaw, 1801) Northern flying squirrel

Sciurus carolinensis Gmelin, 1788 Eastern gray squirrel
88

Scientific name1 Common name3

Tamias townsendii Bachman, 1839 Townsend’s chipmunk

Tamiasciurus douglasii (Bachman, 1839) Douglas’ squirrel

Mice and voles

Clethrionomys gapperi (Vigors, 1830) Southern red-backed vole

Microtus longicaudus (Merriam, 1888) Long-tailed vole

Microtus oregoni (Bachman, 1839) Oregon creeping vole

Mus musculus Linnaeus, 1758 House mouse

Peromyscus maniculatus (Wagner, 1845) Deer mouse

Peromyscus oreas Bangs, 1898 Columbian deer mouse

Zapus trinotatus Rhoads, 1895 Pacific jumping mouse

Carnivores—

Canis latrans Say, 1823 Coyote

Mustela erminea Linnaeus, 1758 Short-tailed weasel

Mustela frenata Lichtenstein, 1831 Long-tailed weasel

Procyon lotor (Linnaeus, 1758) Racoon

Ursus americanus Pallas, 1780 Black bear

Vulpes vulpes (Linnaeus, 1758) Red fox

Ungulate—

Odocoileus hemionus (Rafinesque, 1817) Black-tailed deer

Birds:

Heron—

Ardea herodias Linnaeus, 1758 Great blue heron

Ducks and geese—

Anas platyrhynchos Linnaeus, 1758 Mallard

Branta canadensis (Linnaeus, 1758) Canada goose

Hawks and vultures—

Accipiter cooperii (Bonaparte, 1828) Cooper’s hawk

Accipiter gentilis (Linnaeus, 1758) Northern goshawk

Accipiter striatus (Vieillot, 1807) Sharp-shinned hawk
89

Scientific name1 Common name3

Buteo jamaicensis (Gmelin, 1788) Red-tailed hawk

Cathartes aura (Linnaeus, 1758) Turkey vulture

Haliaeetus leucocephalus (Linnaeus, 1766) Bald eagle

Grouse and quail—

Bonasa umbellus (Linnaeus, 1766) Ruffed grouse

Dendragapus obscurus (Say, 1823) Blue grouse

Pigeons and doves—

Columba fasciata Say, 1823 Band-tailed pigeon

Zenaida macroura (Linnaeus, 1758) Mourning dove

Owls—

Aegolius acadicus (Gmelin, 1788) Northern saw-whet owl

Bubo virginianus (Gmelin, 1788) Great-horned owl

Glaucidium gnoma Wagler, 1832 Northern pygmy owl

Otus kennicotti (Elliot, 1867) Western screech owl

Strix varia Barton, 1799 Barred owl

Tyto alba (Scopoli, 1769) Barn owl

Goatsucker—

Chordeiles minor (Forster, 1771) Common nighthawk

Hummingbird—

Selasphorus rufus (Gmelin, 1788) Rufous hummingbird

Woodpeckers and allies—

Colaptes auratus (Linnaeus, 1758) Northern flicker

Dryocopus pileatus (Linnaeus, 1758) Pileated woodpecker

Picoides pubescens (Linnaeus, 1766) Downy woodpecker

Picoides villosus (Linnaeus, 1766) Hairy woodpecker

Sphyrapicus ruber (Gmelin, 1788) Red-breasted sapsucker

Flycatchers—

Contopus borealis (Swainson, 1832) Olive-sided flycatcher

Empidonax difficilis (Baird, 1858) Pacific slope flycatcher

Empidonax traillii (Audubon, 1828) Willow flycatcher
90

Scientific name1 Common name3

Swallows—

Hirundo rustica Linnaeus, 1758 Barn swallow

Tachycineta bicolor (Vieillot, 1808) Tree swallow

Jays, crows, and ravens—

Corvus brachyrhynchos Brehm, 1822 American crow

Corvus corax Linnaeus, 1758 Common raven

Cyanocitta stelleri (Gmelin, 1788) Steller’s jay

Perisoreus canadensis Linnaeus, 1766 Gray jay

Chickadees, nuthatches, and creepers—

Certhia americana Bonaparte, 1838 Brown creeper

Parus atricapillus Linnaeus, 1766 Black-capped chickadee

Parus rufescens Townsend, 1837 Chestnut-backed chickadee

Psaltriparus minimus (Townsend, 1837) Common bushtit

Sitta canadensis Linnaeus, 1766 Red-breasted nuthatch

Wren—

Troglodytes troglodytes (Linnaeus, 1758) Winter wren

Kinglets—

Regulus calendula (Linnaeus, 1766) Ruby-crowned kinglet

Regulus saltrapa Lichtenstein, 1823 Golden-crowned kinglet

Thrushes—

Catharus guttatus (Pallas, 1811) Hermit thrush

Catharus ustulatus (Nuttall, 1840) Swainson’s thrush

Ixoreus naevius (Gmelin, 1789) Varied thrush

Myadestes townsendi (Audubon, 1838) Townsend’s solitaire

Sialia mexicana Swainson, 1832 Western bluebird

Turdus migratorius Linnaeus, 1766 American robin

Waxwing—

Bombycilla cedrorum Vieillot, 1808 Cedar waxwing
91

Scientific name1 Common name3

Vireos—

Vireo gilvus (Vieillot, 1808) Warbling vireo

Vireo huttoni (Cassin, 1851) Hutton’s vireo

Vireo solitarius (Wilson, 1810) Solitary vireo

Warblers—

Dendroica coronata (Linnaeus, 1766) Yellow-rumped warbler

Dendroica nigrescens (Townsend, 1837) Black-throated gray warbler

Dendroica occidentalis (Townsend, 1837) Hermit warbler

Dendroica townsendi (Townsend, 1837) Townsend’s warbler

Oporornis tolmiei (Townsend, 1837) MacGillivray’s warbler

Vermivora celata (Say, 1823) Orange-crowned warbler

Wilsonia pusilla (Wilson, 1811) Wilson’s warbler

Tanager and grosbeak—

Pheucticus melanocephalus (Swainson, 1827) Black-headed grosbeak

Piranga ludoviciana (Wilson, 1811) Western tanager

Sparrows—

Junco hyemalis (Linnaeus, 1758) Dark-eyed junco

Melospiza melodia (Wilson, 1810) Song sparrow

Pipilo erythrophthalmus (Linnaeus, 1758) Rufous-sided towhee

Spizella passerina (Bechstein, 1798) Chipping sparrow

Zonotrichia leucophrys (Forster, 1772) White-crowned sparrow

Blackbirds—

Agelaius phoeniceus (Linnaeus, 1766) Red-winged blackbird

Molothrus ater (Boddaert, 1783) Brown-headed cowbird

Finches—

Carduelis pinus (Wilson, 1810) Pine siskin

Carpodacus purpureus (Gmelin, 1789) Purple finch

Loxia curvirostra Linnaeus, 1758 Red crossbill
92

Scientific name1

Hypogeous fungi (truffles):

Alpova diplophloeus (Zeller & Dodge) Trappe & Smith

Elaphomyces granulatus Fr.4

Elaphomyces muricatus Fr.4

Endogone lactiflua Bk. & Bk.

Endogone pisiformis Link:Fr.

Gautieria monticola Harkn.

Gautieria sp. nov.6

Genabea cerebriformis (Harkn.) Trappe

Genea harknessii Gilkey

Genea intermedia Gilkey

Glomus sp. nov.6

Glomus macrocarpum Tul. & Tul.

Glomus microcarpum Tul. & Tul.

Glomus mosseae (Nicol. & Gerd.) Gerd. & Trappe

Hydnotrya variiformis Gilkey

Hymenogaster sublilacinus Smith

Hymenogaster sp. (to be identified)

Hysterangium coriaceum Hesse

Hysterangium crassirhachis Zeller & Dodge

Hysterangium setchellii Fischer

Leucangium carthusianum (Tul. & Tul.) Paoletti

Leucogaster candidus (Harkn.) Fogel comb. ined.

Leucogaster citrinus (Harkn.) Zeller & Dodge 4 7

Leucogaster gelatinosus Fogel nom. ined.

Leucogaster rubescens Zeller & Dodge

Leucogaster sp. nov.

Leucophleps magnata Harkn.

Leucophleps spinispora Fogel

Melanogaster ambiguus (Vitt.) Tul. & Tul.
93

Scientific name1

Melanogaster euryspermus (Zeller & Dodge) Zeller

Melanogaster natsii Wang, Trappe, & Castellano sp. nov.

Melanogaster thiersii Wang, Trappe, & Castellano sp. nov.

Melanogaster trappei Wang sp. nov.

Melanogaster tuberiformis Corda in Sturm

Melanogaster variegatus (Vitt.) Tul. & Tul.

Pachyphloeus thysellii Colgan & Castellano, nom. ined.6

Piloderma fallax (Lib.) Stalp.

Radiigera fuscogleba Zeller

Rhizopogon hawkerae Smith

Rhizopogon rogersii Smith

Rhizopogon subareolatus Smith

Rhizopogon villosulus Zeller

Rhizopogon vinicolor Smith

Rhizopogon vulgaris (Vittad.) M. Lange

Scleroderma hypogaeum Zeller

Truncocolumella citrina Zeller

Tuber anniae Colgan & Trappe

Tuber gibbosum Harkn.

Tuber monticola Harkn.

Genus & species nov. # 16

Genus & species nov. # 26

Epigeous fungi (mushrooms):

Agaricus albolutescens Zeller

Agaricus diminutivus Pk.

Agaricus micromegathus Pk.

Agaricus silvicola (Vitt.) Pk.

Agrocybe praecox (Fr.) Fayod

Albatrellus pescaprae (Fr.) Pouzar
94

Scientific name1

Aleuria aurantia (Fr.) Fuckel

Amanita gemmata var. gemmata (Fr.) Bert.4

Amanita gemmata var. exannulata Lange

Amanita pantherina (DC:Fr.) Schum.4

Amanita porphyria (A.& S. ex Fr.) Secr.4

Amanita silvicola Kauffm.

Amanita smithiana Bas.4

Armillaria mellea (Vahl ex Fr.) Karsten grp.8

Auriscalpium vulgare S.F. Gray

Bolbitius sp.

Boletus chrysenteron Fr.4

Boletus zelleri Murr.4

Cantharellus formosus Corner4 7 9

Cantharellus subalbidus Smith & Morse4 7

Cantharellus tubaeformis Fr.4 7

Clavaria sp.

Clavulina cinerea (Fr.) Schroet.4 7

Clavulina cristata (Fr.) Schroet.4 7

Clavulinopsis corniculata (Schaeff.:Fr.) Corner (= Clavaria corniculata Fr.)

Clavulinopsis laeticolor (Berk. & Curt.) R.H. Petersen4

Clitocybe coniferophila H.E. Bigelow grp.8

Clitocybe deceptiva H.E. Bigelow

Clitocybe dilatata Pers. ex Karsten

Clitocybe inversa (Fr.) Gill.

Collybia acervata (Fr.) Kummer4

Collybia alcalivirens Singer

Collybia butyracea (Fr.) Quel.4

Collybia confluens (Pers. ex Fr.) Kummer

Collybia dryophila (Bull. ex Fr.) Kummer

Collybia fuscopurpurea (Pers.:Fr.) Kumm. grp.8
95

Scientific name1

Collybia oregonensis A.H. Smith

Collybia racemosa (Pers.:Fr.) Quél. 4 7

Coltricia cinnamomea (Jacq.:Pers.) Murr.

Conocybe cyanopus (Atk.) Kühner

Conocybe tenera (Schaeff. ex Fr.) Kühner

Coprinus micaceus (Bull. ex Fr.) Fr.

Coprinus plicatilis (Curt.:Fr.) Fr.

Coprinus sylvaticus Pk.

Cortinarius acutus Fr. grp.4 8

Cortinarius alboviolaceus (Pers. ex Fr.) Fr.

Cortinarius angulosus Fr.

Cortinarius brunneus Fr.

Cortinarius californicus A.H. Smith

Cortinarius cinnamomeus (Fr.) Fr. grp.8

Cortinarius corrugatus Peck

Cortinarius cotoneus Fr. grp.8

Cortinarius crassus Fr.4

Cortinarius crystallinus Fr.

Cortinarius decipiens Fr.

Cortinarius duracinus Fr.

Cortinarius evernius Fr.4

Cortinarius glaucopus (Schaeff.:Fr.) Fr.4

Cortinarius hemitrichus Fr.

Cortinarius infractus (Pers.:Fr.) Fr.4

Cortinarius laniger Fr. grp.4 8

Cortinarius malachius Fr.

Cortinarius mucosus (Bull.:Fr.) Fr.

Cortinarius multiformis (Fr.) Fr.

Cortinarius nigrocuspidatus

Cortinarius obtusus Fr. grp.4 8
96

Scientific name1

Cortinarius olympianus A.H. Smith 4 7

Cortinarius paleaceus Fr.4

Cortinarius percomis Fr.

Cortinarius phoeniceus (Bull.) R. Maire var. occidentalis Smith.4

Cortinarius pinetorum (Fr.) Kauffman4

Cortinarius plumiger (Fr.) Fr.

Cortinarius prasinus Fr.

Cortinarius pseudobolaris Maire sensu Smith

Cortinarius rubripes Kauffman

Cortinarius sanguineus (Fr.) Fr.4

Cortinarius scaurus Fr. grp.8

Cortinarius semisanguineus (Fr.) Gillet 4

Cortinarius subfoetidus A.H. Smith

Cortinarius subg. bulbopodium

Cortinarius subg. phlegmacium

Cortinarius subg. sericeocybe

Cortinarius subg. telemonia

Cortinarius superbus A.H. Smith

Cortinarius triformis Fr.

Cortinarius uliginosus Berk.

Cortinarius uraceus Fr.

Cortinarius vibratilis Fr.4

Crepidotus herbarum (Pk.) Sacc.

Crepidotus mollis (Fr.) Stde.

Crucibulum laeve (Huds.) Kamb.

Cystoderma amianthinum (Scop.:Fr.) Fr.

Cystoderma fallax Smith & Singer

Cystoderma granulosum (Fr.) Fayod

Cudonia monticola Mains 4 7

Dacrymyces palmatus (Schw.) Bres.
97

Scientific name1

Entoloma nidorosum (Fr.) Quél. grp.8

Entoloma rhodopolium (Fr.) Kumm. grp.8

Fomitopsis cajanderi (Karsten) Kotlaba & Pouz.

Fomitopsis officinalis (Fr.) Bond. & Sing.

Fomitopsis pinicola (Fr.) Karst.

Galerina autumnalis (Pk.) Smith & Singer

Galerina heterocystis (Atk.) A.H. Smith 4 7

Galerina marginata (Fr.) Kühner

Gomphidius glutenosus (Fr.) Fr.

Gomphidius oregonensis Peck

Gomphidius smithii Miller

Gomphidius subroseus Kauffman

Gomphus clavatus (Fr.) S.F. Gray 4 7

Guepiniopsis alpinus (Tracy & Earle) Bres.

Gymnopilus bellulus (Peck) Murr.4

Gymnopilus liquiritiae (Pers.:Fr.) Karst.

Gymnopilus penetrans (Fr. ex Fr.) Murr.

Gymnopilus sapineus (Fr.) Maire

Hebeloma crustuliniforme (Bull. ex St. Amams) Quel.4

Hebeloma sinapizans (Paulet:Fr.) Gill.

Helvella crispa Scop. ex Fr.

Helvella elastica Bulliard:Fr.4 7

Helvella lacunosa Afz. ex Fr.

Hydnum repandum L. ex Fr. (= Dentinum repandum (Fr.) S.F. Gray)4 7

Hydnum umbilicatum Pk. (= Dentinum umbilicatum (Pk.) Pouzar)4 7

Hygrocybe miniata (Fr.) Kumm.

Hygrophoropsis aurantiaca (Wulf. ex Fr.) Maire4

Hygrophorus calophyllus Karst.

Hygrophorus conicus (Fr.) Fr. (= Hygrocybe conica Fr.)4

Hygrophorus glutinosus (Schff.:Fr.) Fr.
98

Scientific name1

Hypholoma capnoides (Fr. ex Fr.) Kummer (= Naematoloma capnoides (Fr.)

 Karsten)4

Hypholoma dispersum (Fr.) Quel. (= Naematoloma dispersum (Fr.) Karsten)4

Hypholoma fasciculare (Huds. ex Fr.) Kummer (= Naematoloma fasciculare

 (Huds. ex Fr.) Karsten)

Inocybe albodisca Pk.

Inocybe calamistrata (Fr.) Gillet4

Inocybe cincinnatula Kühner

Inocybe cookei Bres.

Inocybe eutheles Berk. & Br.

Inocybe fastigiata (Schaeff. ex Fr.) Quel.

Inocybe fuscodisca (Peck) Massae4

Inocybe geophylla (Sow. ex Fr.) Kummer

Inocybe lanatodisca Kauffman

Inocybe lanuginosa (Bull. ex Fr.) Kummer4

Inocybe lilacina (Boud.) Kauffman

Inocybe maculata Boud.

Inocybe mixtilis Britz.

Inocybe olympiana A.H. Smith

Inocybe pudica Kühner

Inocybe sororia Kauffman4

Inocybe subcarpta Kühner & Boursier

Laccaria amethysteo-occidentalis Mueller4

Laccaria laccata (Scop. ex Fr.) Cke.4

Lactarius affinis Pk.

Lactarius deliciosus (Fr.) S. F. Gray4

Lactarius fragilis var. rubidus Hels. & Smith

Lactarius pseudomucidus Smith & Hesler4

Lactarius rubrilacteus Smith & Hesler

Lactarius subflammeus Smith & Hesler

Lactarius uvidus Fr.
99

Scientific name1

Laetiporus sulphureus (Bull. ex Fr.) Murr.4

Lentinus sulcatus Berk.

Leotia lubrica Fr.

Lepiota clypeolaria (Bull. ex Fr.) Kummer

Lepiota rhacodes Pilat

Leptonia gracilipes Peck

Leptonia parva Peck

Leptonia serrulata (Fr.:Fr.) Kumm.

Leptonia undulatella (Peck) Sacc.

Limacella glioderma (Fr.) R. Maire4

Lycoperdon foetidum Bon.

Lycoperdon perlatum Pers.

Lycoperdon pyrifome Schaeff.:Pers.4

Lyophyllum decastes (Fr.) Singer

Marasmius candidus (Bolt.:Fr.) Fr.

Marasmius copelandii Peck

Marasmius umbilicatus Kauffman

Melanoleuca melaleuca (Pers. ex Fr.) Murr.

Mycena acicula (Schaeff. ex Fr.) Kummer

Mycena amabilissima (Peck) Sacc.4

Mycena atroalboides (Peck) Sacc.

Mycena aurantiomarginata (Fr.) Quél.4

Mycena capillaris (Schum.:Fr.) Kumm

Mycena citrinomarginata Gillet

Mycena delicatella (Pk.) Smith

Mycena elegantula Peck4

Mycena epipterygia (Fr.) S. F. Gray4

Mycena maculata Karst.4

Mycena murina Murrill grp.8

Mycena occidentalis Murrill
100

Scientific name1

Mycena oregonensis Smith

Mycena pura (Pers. ex Fr.) Kummer

Mycena purpureofusca (Peck) Sacc.

Mycena rorida (Fr.) Quél.

Mycena scabripes (Murrill) Singer

Mycena subcana A.H. Smith

Nidula candida (Pk.) White

Nidula niveotomentosa (Henn.) Lloyd

Nolanea mammosa (L.) Quél. grp.8

Omphalina luteicolor Murrill

Otidea leporina var. leporina (Batsch:Fr.) Fuckel 4 7

Paxillus atrotomentosus (Fr.) Fr.4

Peziza badia Pers.

Peziza sp.

Phaeolus schweinitzii (Fr.) Pat.

Phellinus pini (Fr.) Ames

Pholiota astragalina (Fr.) Singer4

Pholiota decorata (Murr.) Smith & Kessler4

Pholiota mutabilis (Schaeff. ex Fr.) Kummer

Pholiota terrestris Overholts

Pleurocybella porrigens (Pers. ex Fr.) Singer (= Pleurotus porrigens (Pers. ex

 Fr.) Kummer4

Pleurotus ostreatus (Jacq. ex Fr.) Kummer

Pluteus cervinus (Fr.) Kummer

Polyporus badius (S.F. Gray) Schw.

Polyporus hirtus Quél. (=Jahnoporus hirtus (Quél.) Nuss.)4

Polyporus volvatus Peck (=Cryptoporus volvatus (Peck) Shear)

Psathyrella gracilis (Fr.) Quél.

Psathyrella hydrophila (Fr.) Maire

Psathyrella longistriata (Murre) Smith
101

Scientific name1

Pseudohydnum gelatinosum (Scop. ex Fr.) Karsten

Pseudoplectania melaena (Fr.) Boud.

Ramaria stricta (Fr.) Quél.

Ramariopsis kunzei (Fr.) Donk.

Rhodocybe hirneola (Fr.) Orton

Russula aeruginea Lindl.4

Russula albonigra (Krombh.) Fr.4

Russula alutacea (Pers.:Fr.) Fr.

Russula bicolor Burl.4

Russula brevipes Pk.

Russula cremoricolor Earle

Russula cyanoxantha (Schw.) Fr. grp.8

Russula densifolia (Secr.) Gillet

Russula emetica Fr.

Russula fragrantissima Rom.

Russula gracilis Burlingham

Russulas nigricans Fr.4

Russula placita Burl grp.8

Russula sororia (Fr.) Romell grp.8

Russula xerampelina (Schaeff. ex Secr.) Fr.4

Sparassis crispa Wulf:Fr.4 7

Stereum complicatum (Fr.) Fr.

Strobilurus trullisatus (Murr.) Lennox

Stropharia ambigua (Pk.) Zeller

Stropharia hornemannii (Fr.) Lundell4

Suillus lakei (Murr.) Smith & Thiers

Suillus ponderosus Smith & Thiers

Suillus tomentosus (Kauff.) Singer, Snell, & Dick

Thelephora americana Lloyd

Thelephora palmata Scop.:Fr.
102

Scientific name1

Thelephora terrestris Fr.

Trametes hirsuta (Wulf.:Fr.) Pilat

Trametes versicolor (L. ex Fr.) Pilat (= Coriolus versicolor (L. ex Fr.) Quél.)

Trichaptum abietinus (Fr.) Donk.

Trichoglossum hirsutum (Fr.) Boudier

Tricholoma flavovirens (Pers. ex Fr.) Lundell4

Tricholoma imbricatum (Fr.:Fr.) Kumm.4

Tricholoma sulphureum (Bull.:Fr.) Kumm.

Tricholoma terreum (Schaeff.:Fr.) Kumm.

Tricholomopsis rutilans (Schaeff. ex Fr.) Singer

Tubaria furfuracea (Pers. ex Fr.) Gillet

Tyromyces caesius (Fr.) Murr.

Tyromyces chioneus (Fr.) Karsten

Xeromphalina campanella (Bat. ex Fr.) Kuhner & Maire

Xeromphalina fulvipes (Murr.) Smith4

Xylaria hypoxylon (L. ex Hooker) Grev.

1 Vascular plant names and authorities are from Kartesz (1994), with synonomy (in parentheses) to names

found in Hitchcock and Cronquist (1973), where different. Mammal, reptile, amphibian, and bird names and

authorities are from Banks and others (1987) and Jones and others (1992). Hypogeous fungi identifications

were made and authorities assigned by J.M. Trappe of Oregon State University, Corvallis, OR. Epigeous

fungal names are from Arora (1986), Castellano and O’Dell (1997), Phillips (1991), and Tylutki (1979, 1987).

2 Alphacode abbreviations for plants are from the PLANTS database (USDA NRCS 1997).

3 Common names are provided for plants and animals. Because many fungi do not have common names,

they are not provided for any of the included fungi.

4 Species closely associated with late-successional or old-growth forests within the range of the northern

spotted owl (FEMAT 1993).

5 First record for this vascular plant in Washington (Thysell and others 1997).

6 Previously undescribed hypogeous fungal species collected during the course of the Forest Ecosystem

Study.

7 Species to be protected through survey and management guidelines as outlined in the standards and

guidelines for management of habitat for late-successional and old-growth forest related species within the

range of the northern spotted owl (USDA and USDI 1994).

8 Those fungal taxa ending in grp. have been placed in a group of related or similar species pending further

investigation.

9 This is the common yellow chantarelle of the Pacific Northwest. For a discussion of its nomenclatural and

taxonomic history, see Castellano and O’Dell (1997).
103

Appendix 2

Nestbox and Cavity

Design

Figure 11—Construction of the Flyger nestboxes (Carey and Gill 1983).

Figure 12—Construction and dimensions of the original cavities and faceplates.

3-in cut

BAFFLE

1.5- by 1.5-in 

cut

8.5 in

8 in

8.5 in

7 in

6.25 in

6.5 in

Side view

Original faceplate 

design

Bottom entrance Top entrance

7 in

Front view

Cavity dimensions 

7 by 4 by 6 in

Hole diam. = 1.75 in
104

Figure 13—Construction and dimensions of the modified cavities and faceplates.

Star/Stellar permanent plots Farley/Hill permanent plots

Target Retained Post facto Target Retained Post facto

Stand Cell ID1 treatment RD RD class2 Stand Cell ID1 treatment RD RD class2

101 A/B 3/4 RT 1.9 RD2 301 A/B 6/7 CT -3 -

101 B/C 1/2 RT 3.4 RD4 301 B/C 4/5 CT - -

101 C/D 1/2 RT 1.1 RD2 301 E/F 3/4 CT - -

101 C/D 3/4 RT 2.5 RD2 301 E/F 4/5 CT - -

101 C/D 6/7 RT 1.2 RD2 301 F/G 1/2 CT - -

101 G/H 3/4 RT 5.1 RD6 301 G/H 5/6 CT - -

101 B/C 3/4 HT 3.0 RD2 302 B/C 3/4 RT 1.3 RD2

101 B/C 4/5 HT 6.1 RD6 302 C/D 4/5 RT 1.0 RD2

101 B/C 6/7 HT 3.8 RD4 302 D/E 6/7 RT 1.9 RD2

101 D/E 5/6 HT 7.0 RD8 302 E/F 4/5 RT .6 RD2

101 F/G 4/5 HT 4.0 RD4 302 F/G 2/3 RT 3.0 RD2

101 G/H 6/7 HT 6.5 RD6 302 G/H 5/6 RT 2.9 RD2

101 A/B 6/7 LT 6.7 RD6 302 B/C 2/3 HT 5.5 RD6

101 C/D 2/3 LT 5.3 RD6 302 B/C 5/6 HT 6.4 RD6

101 E/F 4/5 LT 5.3 RD6 302 C/D 6/7 HT 2.9 RD2

Appendix 3

Permanent

Plot

Locations

Elongated top 

entrance 

design

13 in

Elongated middle 

entrance design

Elongated 

faceplate 

design

Area to be excavated

with chainsaw

Front view Side view
105

Star/Stellar permanent plots Farley/Hill permanent plots

Target Retained Post facto Target Retained Post facto

Stand Cell ID1 treatment RD RD class2 Stand Cell ID1 treatment RD RD class2

101 F/G 3/4 LT 4.5 RD4 302 C/D 7/8 HT 4.8 RD6

101 G/H 5/6 LT 6.7 RD6 302 D/E 3/4 HT 6.4 RD6

101 G/H 7/8 LT 6.0 RD6 302 F/G 6/7 HT 7.2 RD8

102 A/B 3/4 CT - - 302 A/B 1/2 LT 5.9 RD6

102 B/C 6/7 CT - - 302 A/B 7/8 LT 6.4 RD6

102 C/D 2/3 CT - - 302 D/E 2/3 LT 5.5 RD6

102 D/E 5/6 CT - - 302 E/F 3/4 LT 8.3 RD8

102 F/G 2/3 CT - - 302 F/G 7/8 LT 5.5 RD6

102 F/G 6/7 CT - - 302 G/H 3/4 LT 6.4 RD6

103 C/D 7/8 RT4 1.7 RD2 303 B/C 2/3 RT 1.9 RD2

103 D/E 6/7 RT 2.3 RD2 303 C/D 3/4 RT 1.0 RD2

103 E/F 2/3 RT 1.7 RD2 303 E/F 3/4 RT 1.6 RD2

103 E/F 7/8 RT 3.3 RD4 303 F/G 1/2 RT 1.9 RD2

103 G/H 1/2 RT 1.5 RD2 303 F/G 5/6 RT 1.8 RD2

103 A/B 3/4 HT 6.0 RD6 303 G/H 5/6 RT 1.6 RD2

103 A/B 7/8 HT 6.0 RD6 303 A/B 3/4 HT 2.8 RD2

103 D/E 1/2 HT 5.7 RD6 303 C/D 1/2 HT 5.4 RD6

103 D/E 3/4 HT 6.2 RD6 303 C/D 4/5 HT 6.3 RD6

103 E/F 6/7 HT 6.0 RD6 303 C/D 6/7 HT 5.5 RD6

103 F/G 3/4 HT 7.6 RD8 303 E/F 6/7 HT 7.3 RD8

103 A/B 4/5 LT 4.7 RD4 303 F/G 3/4 HT 4.2 RD4

103 B/C 2/3 LT 6.3 RD6 303 A/B 6/7 LT 5.4 RD6

103 B/C 5/6 LT 6.3 RD6 303 A/B 7/8 LT 5.4 RD6

103 B/C 6/7 LT 6.0 RD6 303 C/D 7/8 LT 3.6 RD4

103 C/D 3/4 LT 6.0 RD6 303 D/E 1/2 LT 4.6 RD4

103 E/F 3/4 LT 5.1 RD6 303 E/F 2/3 LT 4.6 RD4

104 B/C 4/5 CT - - 303 F/G 2/3 LT 3.9 RD4

104 C/D 6/7 CT - - 304 A/B 3/4 CT - -

104 D/E 2/3 CT - - 304 B/C 3/4 CT - -

104 E/F 7/8 CT - - 304 B/C 7/8 CT - -
106

Star/Stellar permanent plots Farley/Hill permanent plots

Target Retained Post facto Target Retained Post facto

Stand Cell ID1 treatment RD RD class2 Stand Cell ID1 treatment RD RD class2

104 F/G 3/4 CT - - 304 C/D 5/6 CT - -

104 F/G 5/6 CT - - 304 D/E 7/8 CT - -

201 B/C 2/3 RT 2.4 RD2 304 F/G 2/3 CT - -

201 B/C 7/8 RT 1.0 RD2 401 B/C 2/3 RT 2.7 RD2

201 C/D 5/6 RT .8 RD2 401 C/D 5/6 RT 1.2 RD2

201 C/D 7/8 RT .4 RD2 401 D/E 1/2 RT 3.5 RD4

201 E/F 4/5 RT .5 RD2 401 E/F 4/5 RT 3.7 RD4

201 F/G 4/5 RT 1.8 RD2 401 E/F 7/8 RT 1.4 RD4

201 A/B 1/2 HT 4.5 RD4 401 F/G 4/5 RT 1.6 RD4

201 A/B 2/3 HT 7.0 RD8 401 A/B 4/5 HT 4.3 RD4

201 E/F 6/7 HT 4.7 RD4 401 C/D 2/3 HT 3.6 RD4

201 F/G 3/4 HT 3.7 RD4 401 C/D 3/4 HT 6.4 RD6

201 G/H 2/3 HT 4.3 RD4 401 E/F 5/6 HT 7.7 RD8

201 G/H 3/4 HT 3.7 RD4 401 F/G 7/8 HT 4.5 RD4

201 A/B 3/4 LT 6.0 RD6 401 G/H 5/6 HT 8.4 RD8

201 A/B 5/6 LT 8.4 RD8 401 A/B 3/4 LT 3.5 RD4

201 C/D 3/4 LT 3.7 RD4 401 A/B 6/7 LT 7.4 RD8

201 C/D 4/5 LT 6.0 RD6 401 B/C 1/2 LT 7.1 RD8

201 F/G 6/7 LT 5.2 RD6 401 D/E 4/5 LT 6.4 RD6

201 F/G 7/8 LT 5.2 RD6 401 E/F 3/4 LT 3.6 RD4

202 A/B 1/2 RT 1.7 RD2 401 F/G 6/7 LT 8.3 RD8

202 A/B 3/4 RT 1.4 RD2 402 A/B 5/6 CT - -

202 B/C 4/5 RT 3.7 RD4 402 B/C 3/4 CT - -

202 E/F 4/5 RT 1.2 RD2 402 C/D 7/8 CT - -

202 F/G 2/3 RT .7 RD2 402 D/E 2/3 CT - -

202 F/G 6/7 RT .8 RD2 402 D/E 5/6 CT - -

202 B/C 3/4 HT 5.9 RD6 402 F/G 4/5 CT - -

202 B/C 7/8 HT 5.9 RD6 403 A/B 6/7 RT 1.3 RD2

202 D/E 3/4 HT 4.4 RD4 403 D/E 1/2 RT 1.5 RD2

202 E/F 2/3 HT 5.2 RD6 403 E/F 2/3 RT .7 RD2
107

Star/Stellar permanent plots Farley/Hill permanent plots

Target Retained Post facto Target Retained Post facto

Stand Cell ID1 treatment RD RD class2 Stand Cell ID1 treatment RD RD class2

202 F/G 7/8 HT 3.9 RD4 403 E/F 6/7 RT 1.8 RD2

202 G/H 2/3 HT 4.4 RD4 403 F/G 3/4 RT 2.8 RD2

202 A/B 4/5 LT 7.1 RD8 403 F/G 7/8 RT 1.4 RD2

202 D/E 2/3 LT 3.1 RD2 403 A/B 2/3 HT 6.5 RD6

202 E/F 3/4 LT 4.0 RD4 403 B/C 4/5 HT 5.5 RD6

202 F/G 3/4 LT 7.1 RD8 403 B/C 6/7 HT 7.3 RD8

202 F/G 4/5 LT 5.2 RD6 403 C/D 2/3 HT 6.9 RD8

202 G/H 3/4 LT 5.9 RD6 403 C/D 5/6 HT 3.9 RD4

203 A/B 7/8 CT - - 403 D/E 3/4 HT 6.5 RD6

203 C/D 3/4 CT - - 403 B/C 1/2 LT 7.3 RD8

203 C/D 5/6 CT - - 403 C/D 4/5 LT 7.3 RD8

203 D/E 1/2 CT - - 403 D/E 2/3 LT 4.3 RD4

203 D/E 6/7 CT - - 403 F/G 4/5 LT 5.4 RD6

203 F/G 5/6 CT - - 403 F/G 6/7 LT 6.5 RD6

204 A/B 7/8 CT - - 403 G/H 4/5 LT 6.5 RD6

204 B/C 2/3 CT - - 404 A/B 3/4 CT - -

204 B/C 6/7 CT - - 404 B/C 3/4 CT - -

204 D/E 2/3 CT - - 404 D/E 2/3 CT - -

204 D/E 5/6 CT - - 404 D/E 5/6 CT - -

204 F/G 5/6 CT - - 404 F/G 4/5 CT - -

404 F/G 6/7 CT - -

1 Cell ID provides the alphanumeric identity of the four corners of the 40- by 40-m cell in the center of which is the

permanent plot.

2 Post facto RD classes are delineated as follows: RD2 = RD < 3.25; RD4 = 3.25 < RD < 4.75; RD6 = 4.75 < RD < 6.75;

RD8 = RD > 6.75.

 3 - = not applicable.

4 All target subtreatments or treatments in every stand were sampled with six plots, except for the RT subtreatments in

stand 103 which contained only five total RT cells of sufficient size in which to establish permanent plots.
108

Grid Prethin Residual Prethin Residual

cell quadratic quadratic Prethin Residual PrethinResidual relative relative

(NW mean mean basal basal tree tree density density

Stand corner) diameter1 diameter1 area area density density (metric) (metric)

----Centimeters---- -------m2/ha------- -------No./ha-------

101 A1 34.0 34.0 18.4 18.4 188.6 188.6 3.1 3.1

101 A2 34.4 36.9 41.3 36.7 400.5 358.1 7.0 6.0

101 A3 31.1 34.0 24.9 11.0 273.5 155.6 4.5 1.9

101 A4 34.4 36.9 41.6 23.7 746.7 353.7 7.1 3.9

101 A5 34.6 37.5 59.7 50.5 570.1 485.3 10.1 8.2

101 A6 34.6 37.5 59.7 41.3 570.1 400.5 10.1 6.7

101 A7 34.4 36.9 43.3 22.5 746.7 353.7 7.4 3.7

101 B1 31.1 34.0 48.3 19.8 495.1 141.5 8.7 3.4

101 B2 34.0 34.0 13.8 13.8 146.2 146.2 2.4 2.4

101 B3 31.1 34.0 22.1 17.3 243.2 176.8 4.0 3.0

101 B4 34.4 36.9 59.1 36.8 641.0 331.6 10.1 6.1

101 B5 31.1 34.0 18.4 13.8 188.6 146.2 3.3 2.4

101 B6 34.4 36.9 50.5 23.0 485.3 231.0 8.6 3.8

101 B7 34.6 37.5 55.1 50.5 527.7 485.3 9.4 8.2

101 C1 31.1 34.0 43.0 6.4 542.3 70.7 7.7 1.1

101 C2 34.4 36.9 64.3 32.1 612.4 315.7 11.0 5.3

101 C3 31.1 34.0 46.8 14.9 545.4 113.2 8.4 2.5

101 C4 34.4 36.9 43.1 26.1 508.4 221.0 7.4 4.3

101 C5 34.4 36.9 37.5 27.7 589.5 393.0 6.4 4.6

101 C6 31.1 34.0 47.2 6.7 620.9 70.7 8.5 1.2

101 C7 34.4 36.9 55.1 32.1 527.7 315.7 9.4 5.3

101 D1 34.4 36.9 45.9 27.6 442.9 273.4 7.8 4.5

101 D2 34.6 37.5 68.9 41.3 654.8 400.5 11.7 6.7

101 D3 36.9 36.9 27.6 27.6 273.4 273.4 4.5 4.5

101 D4 34.4 36.9 50.5 27.6 485.3 273.4 8.6 4.5

101 D5 34.6 37.5 61.3 42.8 795.8 486.3 10.4 7.0

101 D6 34.6 37.5 53.2 38.0 786.0 510.9 9.0 6.2

101 D7 34.4 36.9 41.3 27.6 400.5 273.4 7.0 4.5

Appendix 4

Cell-by-Cell

Estimates

of Stand

Parameters

(From

Postthinning

Assessment)

Footnote is on p. 121.
109

Grid Prethin Residual Prethin Residual

cell quadratic quadratic Prethin Residual PrethinResidual relative relative

(NW mean mean basal basal tree tree density density

Stand corner) diameter diameter area area density density (metric) (metric)

----Centimeters---- -------m2/ha------- -------No./ha-------

101 E1 34.6 37.5 73.5 64.3 697.2 612.4 12.5 10.5

101 E2 34.4 36.9 41.3 36.7 400.5 358.1 7.0 6.0

101 E3 34.6 37.5 53.9 42.9 589.5 393.0 9.2 7.0

101 E4 34.4 36.9 45.9 32.1 442.9 315.7 7.8 5.3

101 E5 36.9 36.9 23.0 23.0 231.0 231.0 3.8 3.8

101 E6 31.1 34.0 45.7 6.8 396.1 42.4 8.2 1.2

101 E7 34.0 34.0 18.4 18.4 188.6 188.6 3.1 3.1

101 F1 34.4 36.9 32.1 23.0 315.7 231.0 5.5 3.8

101 F2 31.1 34.0 36.7 18.4 358.1 188.6 6.6 3.1

101 F3 36.9 36.9 27.6 27.6 273.4 273.4 4.5 4.5

101 F4 34.4 36.9 36.6 24.4 464.2 265.3 6.2 4.0

101 F5 34.0 34.0 18.4 18.4 188.6 188.6 3.1 3.1

101 F6 34.6 37.5 50.5 45.9 485.3 442.9 8.6 7.5

101 F7 34.6 37.5 59.7 45.9 570.1 442.9 10.1 7.5

101 G1 34.0 34.0 4.6 4.6 61.4 61.4 .8 .8

101 G2 34.6 37.5 49.5 43.8 353.7 275.1 8.4 7.2

101 G3 34.4 36.9 59.3 30.7 633.5 240.5 10.1 5.1

101 G4 36.9 36.9 23.0 23.0 231.0 231.0 3.8 3.8

101 G5 34.6 37.5 55.1 41.3 527.7 400.5 9.4 6.7

101 G6 34.6 37.5 45.8 39.7 486.3 353.7 7.8 6.5

101 G7 36.9 36.9 36.7 36.7 358.1 358.1 6.0 6.0

103 A1 34.4 37.2 52.4 27.2 530.5 198.9 8.9 4.5

103 A2 34.4 37.2 27.2 24.1 353.7 275.1 4.6 3.9

103 A3 34.4 37.2 52.7 36.8 397.9 221.0 9.0 6.0

103 A4 34.4 37.2 28.0 28.7 393.0 353.7 4.8 4.7

103 A5 34.6 43.7 50.5 41.3 485.3 400.5 8.6 6.3

103 A6 34.6 43.7 49.8 37.5 746.7 471.6 8.5 5.7

103 A7 34.4 37.2 59.7 36.7 570.1 358.1 10.2 6.0
110

Grid Prethin Residual Prethin Residual

cell quadratic quadratic Prethin Residual PrethinResidual relative relative

(NW mean mean basal basal tree tree density density

Stand corner) diameter diameter area area density density (metric) (metric)

----Centimeters---- -------m2/ha------- -------No./ha-------

103 B1 34.4 37.2 59.7 36.7 570.1 358.1 10.2 6.0

103 B2 34.4 37.2 36.7 32.1 358.1 315.7 6.3 5.3

103 B2 34.6 43.7 50.5 41.3 485.3 400.5 8.6 6.3

103 B4 34.4 37.2 36.7 27.6 358.1 273.4 6.3 4.5

103 B5 34.6 43.7 64.3 41.3 612.4 400.5 10.9 6.3

103 B6 34.4 37.2 45.9 36.7 442.9 358.1 7.8 6.0

103 B7 34.6 43.7 50.5 41.3 485.3 400.5 8.6 6.3

103 C1 34.4 37.2 36.7 32.1 358.1 315.7 6.3 5.3

103 C2 34.4 37.2 50.5 36.7 485.3 358.1 8.6 6.0

103 C3 34.4 37.2 50.5 36.7 485.3 358.1 8.6 6.0

103 C4 34.4 37.2 50.5 27.6 485.3 273.4 8.6 4.5

103 C5 43.7 43.7 41.3 41.3 400.5 400.5 6.3 6.3

103 C6 37.2 37.2 23.0 23.0 231.0 231.0 3.8 3.8

103 C7 31.1 35.2 39.2 10.3 382.0 56.6 7.0 1.7

103 D1 34.6 43.7 50.4 37.9 508.4 331.6 8.6 5.7

103 D2 34.4 37.2 41.3 27.6 400.5 273.4 7.0 4.5

103 D3 34.6 43.7 53.2 40.9 618.9 397.9 9.0 6.2

103 D4 34.4 37.2 36.7 23.0 358.1 231.0 6.3 3.8

103 D5 34.4 37.2 32.1 27.6 315.7 273.4 5.5 4.5

103 D6 31.1 35.2 52.1 13.8 622.5 113.2 9.3 2.3

103 D7 34.4 37.2 27.6 23.0 273.4 231.0 4.7 3.8

103 E1 34.6 43.7 55.1 41.3 527.7 400.5 9.4 6.3

103 E2 31.1 35.2 40.3 10.3 466.9 56.6 7.2 1.7

103 E3 34.4 37.2 40.9 31.3 589.5 432.3 7.0 5.1

103 E4 31.1 35.2 27.6 18.4 273.4 188.6 4.9 3.1

103 E5 34.4 37.2 52.8 28.5 746.7 353.7 9.0 4.7

103 E6 34.6 43.7 53.2 39.8 464.2 265.3 9.0 6.0

103 E7 31.1 35.2 53.8 19.5 495.1 84.9 9.7 3.3
111

Grid Prethin Residual Prethin Residual

cell quadratic quadratic Prethin Residual PrethinResidual relative relative

(NW mean mean basal basal tree tree density density

Stand corner) diameter diameter area area density density (metric) (metric)

----Centimeters---- -------m2/ha------- -------No./ha-------

103 F1 34.4 37.2 41.3 32.1 400.5 315.7 7.0 5.3

103 F2 34.6 43.7 50.5 41.3 485.3 400.5 8.6 6.3

103 F3 34.6 43.7 55.1 50.5 527.7 485.3 9.4 7.6

103 F4 34.4 37.2 36.7 32.1 358.1 315.7 6.3 5.3

103 F5 34.4 37.2 32.1 23.0 315.7 231.0 5.5 3.8

103 F6 34.4 37.2 52.2 29.3 746.7 353.7 8.9 4.8

103 F7 34.4 37.2 32.1 27.6 315.7 273.4 5.5 4.5

103 G1 31.1 35.2 49.4 8.8 650.8 70.7 8.9 1.5

103 G2 31.1 35.2 25.2 19.3 397.9 287.4 4.5 3.3

103 G3 35.2 35.2 13.8 13.8 146.2 146.2 2.3 2.3

103 G4 34.4 37.2 45.9 23.0 442.9 231.0 7.8 3.8

103 G5 34.4 37.2 45.9 32.1 442.9 315.7 7.8 5.3

103 G6 34.4 37.2 36.7 27.6 358.1 273.4 6.3 4.5

103 G7 34.4 37.2 50.5 32.1 485.3 315.7 8.6 5.3

201 A1 34.4 38.0 32.1 27.6 315.7 273.4 5.5 4.5

201 A2 34.6 36.5 48.4 42.1 464.2 397.9 8.2 7.0

201 A3 34.4 38.0 45.9 36.7 442.9 358.1 7.8 6.0

201 A4 34.4 38.0 50.5 27.6 485.3 273.4 8.6 4.5

201 A5 34.6 36.5 55.1 50.5 527.7 485.3 9.4 8.4

201 A6 34.4 38.0 59.7 36.7 570.1 358.1 10.2 6.0

201 A7 34.4 38.0 36.7 27.6 358.1 273.4 6.3 4.5

201 B1 34.6 36.5 71.2 49.3 685.3 375.8 12.1 8.2

201 B2 31.1 33.1 42.5 13.6 460.7 84.9 7.6 2.4

201 B3 34.4 38.0 59.7 27.6 570.1 273.4 10.2 4.5

201 B4 34.4 38.0 45.8 32.9 668.1 432.3 7.8 5.3

201 B5 34.6 36.5 78.1 64.3 739.6 612.4 13.3 10.6

201 B6 34.6 36.5 50.5 41.3 485.3 400.5 8.6 6.8

201 B7 31.1 33.1 19.1 5.8 263.5 42.4 3.4 1.0
112

Grid Prethin Residual Prethin Residual

cell quadratic quadratic Prethin Residual PrethinResidual relative relative

(NW mean mean basal basal tree tree density density

Stand corner) diameter diameter area area density density (metric) (metric)

----Centimeters---- -------m2/ha------- -------No./ha-------

201 C1 34.4 38.0 32.1 23.0 315.7 231.0 5.5 3.7

201 C2 34.4 38.0 49.0 33.3 746.7 432.3 8.4 5.4

201 C3 38.0 38.0 23.0 23.0 231.0 231.0 3.7 3.7

201 C4 34.4 38.0 55.1 36.7 527.7 358.1 9.4 6.0

201 C5 31.1 33.1 20.8 4.6 313.9 70.7 3.7 .8

201 C6 38.0 38.0 32.1 32.1 315.7 315.7 5.2 5.2

201 C7 31.1 33.1 25.4 2.4 426.2 28.3 4.6 .4

201 D1 34.6 36.5 53.3 48.6 550.2 471.6 9.1 8.0

201 D2 31.1 33.1 41.3 18.4 400.5 188.6 7.4 3.2

201 D3 34.6 36.5 82.7 64.3 782.0 612.4 14.1 10.6

201 D4 34.4 38.0 45.9 32.1 442.9 315.7 7.8 5.2

201 D5 38.0 38.0 27.6 27.6 273.4 273.4 4.5 4.5

201 D6 34.4 38.0 32.1 23.0 315.7 231.0 5.5 3.7

201 D7 38.0 38.0 27.6 27.6 273.4 273.4 4.5 4.5

201 E1 31.1 33.1 13.8 9.2 146.2 103.8 2.5 1.6

201 E2 38.0 38.0 23.0 23.0 231.0 231.0 3.7 3.7

201 E3 34.4 38.0 42.2 35.6 530.5 397.9 7.2 5.8

201 E4 31.1 33.1 33.6 2.7 448.3 28.3 6.0 .5

201 E5 34.4 38.0 45.0 25.8 628.8 314.4 7.7 4.2

201 E6 34.4 38.0 45.4 28.7 707.4 331.6 7.7 4.7

201 E7 34.6 36.5 64.3 45.9 612.4 442.9 10.9 7.6

201 F1 34.4 38.0 41.3 23.0 400.5 231.0 7.0 3.7

201 F2 34.6 36.5 64.3 45.9 612.4 442.9 10.9 7.6

201 F3 34.4 38.0 27.6 23.0 273.4 231.0 4.7 3.7

201 F4 31.1 33.1 30.1 10.3 372.2 84.9 5.4 1.8

201 F5 34.6 36.5 68.9 50.5 654.8 485.3 11.7 8.4

201 F6 34.4 38.0 50.5 32.1 485.3 315.7 8.6 5.2

201 F7 34.4 38.0 45.9 32.1 442.9 315.7 7.8 5.2
113

Grid Prethin Residual Prethin Residual

cell quadratic quadratic Prethin Residual PrethinResidual relative relative

(NW mean mean basal basal tree tree density density

Stand corner) diameter diameter area area density density (metric) (metric)

----Centimeters---- -------m2/ha------- -------No./ha-------

201 G1 34.4 38.0 36.7 23.0 358.1 231.0 6.3 3.7

201 G2 34.4 38.0 40.1 26.3 397.9 221.0 6.8 4.3

201 G3 31.1 33.1 38.6 21.2 596.8 287.4 6.9 3.7

201 G4 34.6 36.5 52.2 45.9 550.2 393.0 8.9 7.6

201 G5 34.4 38.0 36.7 23.0 358.1 231.0 6.3 3.7

201 G6 34.4 38.0 43.8 30.9 707.4 432.3 7.5 5.0

201 G7 34.4 38.0 45.9 32.1 442.9 315.7 7.8 5.2

202 A1 31.1 35.5 39.7 9.9 353.7 99.0 7.1 1.7

202 A2 31.1 35.5 23.0 13.8 231.0 146.2 4.1 2.3

202 A3 31.1 35.5 41.3 8.3 481.0 84.9 7.4 1.4

202 A4 34.6 42.2 55.1 45.9 527.7 442.9 9.4 7.1

202 A5 34.4 38.3 32.1 27.6 315.7 273.4 5.5 4.4

202 A6 34.4 38.3 41.3 36.7 400.5 358.1 7.0 5.9

202 A7 34.6 42.2 52.7 38.2 596.8 331.6 9.0 5.9

202 B1 34.4 38.3 36.7 32.1 358.1 315.7 6.3 5.2

202 B2 34.6 42.2 50.5 45.9 485.3 442.9 8.6 7.1

202 B3 34.6 42.2 48.9 38.2 464.2 331.6 8.3 5.9

202 B4 31.1 35.5 60.3 22.1 608.3 155.6 10.8 3.7

202 B5 38.3 38.3 27.6 27.6 273.4 273.4 4.4 4.4

202 B6 34.4 38.3 36.7 27.6 358.1 273.4 6.3 4.4

202 B7 34.4 38.3 57.8 36.5 530.5 265.3 9.9 5.9

202 C1 34.4 38.3 68.2 35.2 596.8 309.5 11.6 5.7

202 C2 34.4 38.3 36.7 27.6 358.1 273.4 6.3 4.4

202 C3 34.4 38.3 41.3 32.1 400.5 315.7 7.0 5.2

202 C4 34.4 38.3 32.1 27.6 315.7 273.4 5.5 4.4

202 C5 34.4 38.3 41.3 36.7 400.5 358.1 7.0 5.9

202 C6 34.6 42.2 50.5 41.3 485.3 400.5 8.6 6.4

202 C7 34.4 38.3 36.7 27.6 358.1 273.4 6.3 4.4
114

Grid Prethin Residual Prethin Residual

cell quadratic quadratic Prethin Residual PrethinResidual relative relative

(NW mean mean basal basal tree tree density density

Stand corner) diameter diameter area area density density (metric) (metric)

----Centimeters---- -------m2/ha------- -------No./ha-------

202 D1 31.1 35.5 32.1 18.4 315.7 188.6 5.8 3.1

202 D2 35.5 35.5 18.4 18.4 188.6 188.6 3.1 3.1

202 D3 34.4 38.3 50.5 27.6 485.3 273.4 8.6 4.4

202 D4 31.1 35.5 56.7 13.1 523.4 84.9 10.2 2.2

202 D5 34.4 38.3 50.5 32.1 485.3 315.7 8.6 5.2

202 D6 34.6 42.2 66.7 54.5 668.1 471.6 11.3 8.4

202 D7 34.4 38.3 32.1 27.6 315.7 273.4 5.5 4.4

202 E1 34.6 42.2 74.5 46.7 903.8 471.6 12.7 7.2

202 E2 34.4 38.3 45.9 32.1 442.9 315.7 7.8 5.2

202 E3 34.4 38.3 27.3 24.5 235.8 196.5 4.7 4.0

202 E4 31.1 35.5 47.6 7.4 339.5 42.4 8.5 1.2

202 E5 34.4 38.3 47.2 28.6 530.5 287.4 8.0 4.6

202 E6 34.6 42.2 55.1 50.5 527.7 485.3 9.4 7.8

202 E7 34.4 38.3 59.7 36.7 570.1 358.1 10.2 5.9

202 F1 34.4 38.3 41.3 23.0 400.5 231.0 7.0 3.7

202 F2 31.1 35.5 26.6 4.0 268.8 14.1 4.8 .7

202 F3 34.6 42.2 51.9 46.3 432.3 353.7 8.8 7.1

202 F4 34.4 38.3 36.7 32.1 358.1 315.7 6.3 5.2

202 F5 34.4 38.3 36.7 32.1 358.1 315.7 6.3 5.2

202 F6 31.1 35.5 39.9 5.0 622.5 70.7 7.2 .8

202 F7 34.4 38.3 50.9 24.0 840.0 331.6 8.7 3.9

202 G1 34.4 38.3 23.8 23.8 235.8 235.8 4.1 3.8

202 G2 34.4 38.3 45.9 27.6 442.9 273.4 7.8 4.4

202 G3 34.4 38.3 50.5 36.7 485.3 358.1 8.6 5.9

202 G4 34.4 38.3 27.6 23.0 273.4 231.0 4.7 3.7

202 G5 34.4 38.3 30.6 24.1 628.8 471.6 5.2 3.9

202 G6 34.4 38.3 36.7 27.6 358.1 273.4 6.3 4.4

202 G7 42.2 42.2 41.3 41.3 400.5 400.5 6.4 6.4
115

Grid Prethin Residual Prethin Residual

cell quadratic quadratic Prethin Residual PrethinResidual relative relative

(NW mean mean basal basal tree tree density density

Stand corner) diameter diameter area area density density (metric) (metric)

----Centimeters---- -------m2/ha------- -------No./ha-------

302 A1 34.6 56.3 51.8 44.1 397.9 309.5 8.8 5.9

302 A2 56.3 56.3 48.2 48.2 189.8 189.8 6.4 6.4

302 A3 34.4 51.9 55.1 34.4 215.3 138.8 9.4 4.8

302 A4 34.4 51.9 48.2 34.4 189.8 138.8 8.2 4.8

302 A5 34.6 56.3 48.4 37.5 265.3 176.8 8.2 5.0

302 A6 56.3 56.3 41.3 41.3 164.3 164.3 5.5 5.5

302 A7 56.3 56.3 48.2 48.2 189.8 189.8 6.4 6.4

302 B1 34.6 56.3 48.7 38.7 331.6 243.2 8.3 5.2

302 B2 34.6 56.3 48.2 41.3 189.8 164.3 8.2 5.5

302 B3 31.1 51.6 40.5 9.5 241.4 42.4 7.3 1.3

302 B4 34.6 56.3 55.1 41.3 215.3 164.3 9.4 5.5

302 B5 34.6 56.3 55.1 48.2 215.3 189.8 9.4 6.4

302 B6 31.1 51.6 34.6 20.7 221.0 132.6 6.2 2.9

302 B7 51.6 51.6 20.7 20.7 87.9 87.9 2.9 2.9

302 C1 56.3 56.3 55.1 55.1 215.3 215.3 7.3 7.3

302 C2 34.6 56.3 75.8 48.2 291.8 189.8 12.9 6.4

302 C3 34.6 56.3 82.7 68.9 317.2 266.3 14.1 9.2

302 C4 31.1 51.6 55.2 6.9 285.6 42.4 9.9 1.0

302 C5 51.9 51.9 27.6 27.6 113.4 113.4 3.8 3.8

302 C6 51.6 51.6 20.7 20.7 87.9 87.9 2.9 2.9

302 C7 34.4 51.9 41.3 34.4 164.3 138.8 7.0 4.8

302 D1 34.6 56.3 58.2 47.0 309.5 243.2 9.9 6.3

302 D2 34.6 56.3 62.0 41.3 240.8 164.3 10.5 5.5

302 D3 34.6 56.3 62.0 48.2 240.8 189.8 10.5 6.4

302 D4 31.1 51.6 51.5 18.4 269.7 70.7 9.2 2.6

302 D5 51.9 51.9 34.4 34.4 138.8 138.8 4.8 4.8

302 D6 31.1 51.6 30.4 13.5 130.9 42.4 5.5 1.9

302 D7 51.9 51.9 34.4 34.4 138.8 138.8 4.8 4.8
116

Grid Prethin Residual Prethin Residual

cell quadratic quadratic Prethin Residual PrethinResidual relative relative

(NW mean mean basal basal tree tree density density

Stand corner) diameter diameter area area density density (metric) (metric)

----Centimeters---- -------m2/ha------- -------No./ha-------

302 E1 34.4 51.9 34.4 27.6 138.8 113.4 5.9 3.8

302 E2 34.6 56.3 51.9 46.2 243.2 198.9 8.8 6.2

302 E3 34.6 56.3 68.9 62.0 266.3 240.8 11.7 8.3

302 E4 31.1 51.6 53.0 4.0 213.1 14.1 9.5 .6

302 E5 34.4 51.9 36.9 28.2 198.9 154.7 6.3 3.9

302 E6 34.6 56.3 75.8 55.1 291.8 215.3 12.9 7.3

302 E7 56.3 56.3 64.8 64.8 265.3 265.3 8.6 8.6

302 F1 34.4 51.9 26.3 25.3 198.9 176.8 4.5 3.5

302 F2 31.1 51.6 56.0 21.6 290.0 113.2 10.0 3.0

302 F3 56.3 56.3 62.0 62.0 240.8 240.8 8.3 8.3

302 F4 34.4 51.9 39.0 36.2 132.6 110.5 6.7 5.0

302 F5 34.6 56.3 48.2 41.3 189.8 164.3 8.2 5.5

302 F6 56.3 56.3 53.6 53.6 221.0 221.0 7.2 7.2

302 F7 56.3 56.3 41.3 41.3 164.3 164.3 5.5 5.5

302 G1 31.1 51.6 57.2 15.8 336.0 70.7 10.3 2.2

302 G2 34.4 51.9 48.2 34.4 189.8 138.8 8.2 4.8

302 G3 34.6 56.3 55.1 48.2 215.3 189.8 9.4 6.4

302 G4 34.6 56.3 51.8 43.3 176.8 154.7 8.8 5.8

302 G5 31.1 51.6 69.7 20.6 153.0 42.4 12.5 2.9

302 G6 51.9 51.9 27.6 27.6 113.4 113.4 3.8 3.8

302 G7 51.9 51.9 34.4 34.4 138.8 138.8 4.8 4.8

303 A1 57.2 57.2 27.6 27.6 113.4 113.4 3.6 3.6

303 A2 57.2 57.2 34.4 34.4 138.8 138.8 4.6 4.6

303 A3 53.0 53.0 20.7 20.7 87.9 87.9 2.8 2.8

303 A4 34.6 58.2 62.0 41.3 240.8 164.3 10.5 5.4

303 A5 58.2 58.2 41.3 41.3 164.3 164.3 5.4 5.4

303 A6 34.6 58.2 55.1 41.3 215.3 164.3 9.4 5.4

303 A7 34.6 58.2 48.2 41.3 189.8 164.3 8.2 5.4
117

Grid Prethin Residual Prethin Residual

cell quadratic quadratic Prethin Residual PrethinResidual relative relative

(NW mean mean basal basal tree tree density density

Stand corner) diameter diameter area area density density (metric) (metric)

----Centimeters---- -------m2/ha------- -------No./ha-------

303 B1 34.6 58.2 48.2 41.3 189.8 164.3 8.2 5.4

303 B2 31.1 53.0 52.8 13.5 261.7 84.9 9.5 1.9

303 B3 34.6 58.2 62.3 47.4 265.3 198.9 10.6 6.2

303 B4 34.6 58.2 82.7 62.0 317.2 240.8 14.1 8.1

303 B5 53.0 53.0 6.9 6.9 36.9 36.9 .9 .9

303 B6 57.2 57.2 27.6 27.6 113.4 113.4 3.6 3.6

303 B7 53.0 53.0 6.9 6.9 36.9 36.9 .9 .9

303 C1 34.6 58.2 62.0 41.3 240.8 164.3 10.5 5.4

303 C2 34.6 58.2 55.1 48.2 215.3 189.8 9.4 6.3

303 C3 31.1 53.0 49.8 7.1 168.9 14.1 8.9 1.0

303 C4 34.6 58.2 62.0 48.2 240.8 189.8 10.5 6.3

303 C5 57.2 57.2 34.4 34.4 138.8 138.8 4.6 4.6

303 C6 34.6 58.2 48.4 42.1 265.3 221.0 8.2 5.5

303 C7 34.4 57.2 34.4 27.6 138.8 113.4 5.9 3.6

303 D1 34.4 57.2 41.3 34.4 164.3 138.8 7.0 4.6

303 D2 31.1 53.0 27.6 20.7 113.4 87.9 4.9 2.8

303 D3 57.2 57.2 34.4 34.4 138.8 138.8 4.6 4.6

303 D4 34.6 58.2 55.3 53.2 243.2 221.0 9.4 7.0

303 D5 58.2 58.2 41.3 41.3 164.3 164.3 5.4 5.4

303 D6 34.6 58.2 62.0 48.2 240.8 189.8 10.5 6.3

303 D7 34.6 58.2 54.6 41.2 221.0 154.7 9.3 5.4

303 E1 57.2 57.2 34.4 34.4 138.8 138.8 4.6 4.6

303 E2 57.2 57.2 34.4 34.4 138.8 138.8 4.6 4.6

303 E3 31.1 53.0 54.7 11.4 307.7 42.4 9.8 1.6

303 E4 31.1 53.0 55.2 17.8 247.6 70.7 9.9 2.4

303 E5 34.6 58.2 71.8 54.4 287.4 198.9 12.2 7.1

303 E6 34.6 58.2 68.7 56.0 265.3 198.9 11.7 7.3

303 E7 57.2 57.2 35.9 35.9 154.7 154.7 4.8 4.8
118

Grid Prethin Residual Prethin Residual

cell quadratic quadratic Prethin Residual PrethinResidual relative relative

(NW mean mean basal basal tree tree density density

Stand corner) diameter diameter area area density density (metric) (metric)

----Centimeters---- -------m2/ha------- -------No./ha-------

303 F1 31.1 53.0 48.3 13.8 203.4 70.7 8.7 1.9

303 F2 57.2 57.2 29.4 29.4 110.5 110.5 3.9 3.9

303 F3 34.4 57.2 38.7 31.9 176.8 132.6 6.6 4.2

303 F4 34.6 58.2 58.9 44.0 287.4 198.9 10.0 5.8

303 F5 31.1 53.0 63.5 12.8 241.4 42.4 11.4 1.8

303 F6 34.6 58.2 52.2 44.0 265.3 176.8 8.9 5.8

303 F7 31.1 53.0 27.6 20.7 113.4 87.9 4.9 2.8

303 G1 31.1 53.0 26.8 21.2 114.9 70.7 4.8 2.9

303 G2 58.2 58.2 65.0 65.0 176.8 176.8 8.5 8.5

303 G3 34.4 57.2 53.8 31.2 309.5 198.9 9.2 4.1

303 G4 34.4 57.2 48.2 34.4 189.8 138.8 8.2 4.6

303 G5 31.1 53.0 65.7 11.9 344.0 56.6 11.8 1.6

303 G6 34.6 58.2 68.9 48.2 266.3 189.8 11.7 6.3

303 G7 34.4 57.2 34.4 27.6 138.8 113.4 5.9 3.6

401 A1 59.4 59.4 34.4 34.4 138.8 138.8 4.5 4.5

401 A2 34.6 56.2 55.9 50.8 243.2 198.9 9.5 6.8

401 A3 34.4 59.4 28.3 26.7 154.7 132.6 4.8 3.5

401 A4 34.4 59.4 37.8 33.2 265.3 221.0 6.4 4.3

401 A5 34.6 56.2 62.0 48.2 240.8 189.8 10.5 6.4

401 A6 34.6 56.2 62.0 55.1 240.8 215.3 10.5 7.4

401 A7 59.4 59.4 34.4 34.4 138.8 138.8 4.5 4.5

401 B1 34.6 56.2 62.6 53.5 287.4 221.0 10.6 7.1

401 B2 31.1 54.6 46.4 19.6 223.7 113.2 8.3 2.7

401 B3 59.4 59.4 27.6 27.6 113.4 113.4 3.6 3.6

401 B4 34.4 59.4 26.2 24.0 154.7 132.6 4.5 3.1

401 B5 59.4 59.4 27.6 27.6 113.4 113.4 3.6 3.6

401 B6 59.4 59.4 34.4 34.4 138.8 138.8 4.5 4.5

401 B7 56.2 56.2 55.1 55.1 215.3 215.3 7.4 7.4
119

Grid Prethin Residual Prethin Residual

cell quadratic quadratic Prethin Residual PrethinResidual relative relative

(NW mean mean basal basal tree tree density density

Stand corner) diameter diameter area area density density (metric) (metric)

----Centimeters---- -------m2/ha------- -------No./ha-------

401 C1 56.2 56.2 43.8 43.8 132.6 132.6 5.8 5.8

401 C2 34.4 59.4 34.4 27.6 138.8 113.4 5.9 3.6

401 C3 34.6 56.2 55.1 48.2 215.3 189.8 9.4 6.4

401 C4 54.6 54.6 13.8 13.8 62.4 62.4 1.9 1.9

401 C5 31.1 54.6 67.3 9.1 329.8 42.4 12.1 1.2

401 C6 59.4 59.4 34.4 34.4 138.8 138.8 4.5 4.5

401 C7 34.6 56.2 73.8 67.9 287.4 243.2 12.5 9.1

401 D1 59.4 59.4 27.0 27.0 141.5 141.5 3.5 3.5

401 D2 34.6 56.2 55.1 41.3 215.3 164.3 9.4 5.5

401 D3 59.4 59.4 27.6 27.6 113.4 113.4 3.6 3.6

401 D4 34.6 56.2 68.9 48.2 266.3 189.8 11.7 6.4

401 D5 31.1 54.6 83.6 13.0 410.3 56.6 15.0 1.8

401 D6 34.6 56.2 48.2 41.3 189.8 164.3 8.2 5.5

401 D7 34.4 59.4 41.3 34.4 164.3 138.8 7.0 4.5

401 E1 56.2 56.2 62.0 62.0 240.8 240.8 8.3 8.3

401 E2 34.6 56.2 48.2 41.3 189.8 164.3 8.2 5.5

401 E3 59.4 59.4 27.6 27.6 113.4 113.4 3.6 3.6

401 E4 34.4 59.4 67.2 28.2 211.3 56.6 11.5 3.7

401 E5 34.6 56.2 72.1 57.5 221.0 176.8 12.3 7.7

401 E6 56.2 56.2 62.0 62.0 240.8 240.8 8.3 8.3

401 E7 31.1 54.6 41.9 10.1 183.0 28.3 7.5 1.4

401 F1 59.4 59.4 36.1 36.1 132.6 132.6 4.7 4.7

401 F2 56.2 56.2 48.2 48.2 189.8 189.8 6.4 6.4

401 F3 54.6 54.6 6.9 6.9 36.9 36.9 .9 .9

401 F4 31.1 54.6 68.9 11.5 307.7 42.4 12.4 1.6

401 F5 34.6 56.2 72.5 65.1 287.4 243.2 12.3 8.7

401 F6 34.6 56.2 68.9 62.0 266.3 240.8 11.7 8.3

401 F7 34.4 59.4 41.3 34.4 164.3 138.8 7.0 4.5
120

Grid Prethin Residual Prethin Residual

cell quadratic quadratic Prethin Residual PrethinResidual relative relative

(NW mean mean basal basal tree tree density density

Stand corner) diameter diameter area area density density (metric) (metric)

----Centimeters---- -------m2/ha------- -------No./ha-------

401 G1 56.2 56.2 48.2 48.2 189.8 189.8 6.4 6.4

401 G2 59.4 59.4 27.6 27.6 113.4 113.4 3.6 3.6

401 G3 34.4 59.4 35.8 28.4 163.6 141.5 6.1 3.7

401 G4 34.4 59.4 41.3 34.4 164.3 138.8 7.0 4.5

401 G5 34.6 56.2 73.1 63.3 331.6 265.3 12.4 8.4

401 G6 56.2 56.2 48.2 48.2 189.8 189.8 6.4 6.4

401 G7 34.4 59.4 43.6 34.5 221.0 154.7 7.4 4.5

403 A1 72.6 72.6 48.2 48.2 189.8 189.8 5.7 5.7

403 A2 34.6 72.6 60.7 55.0 265.3 176.8 10.3 6.5

403 A3 34.6 72.6 68.9 48.2 266.3 189.8 11.7 5.7

403 A4 34.4 63.3 48.2 34.4 189.8 138.8 8.2 4.3

403 A5 63.3 63.3 27.0 27.0 117.9 117.9 3.4 3.4

403 A6 31.1 58.0 68.1 10.1 460.6 28.3 12.2 1.3

403 A7 34.6 72.6 62.0 48.2 240.8 189.8 10.5 5.7

403 B1 34.6 72.6 68.9 62.0 266.3 240.8 11.7 7.3

403 B2 72.6 72.6 41.3 41.3 164.3 164.3 4.9 4.9

403 B3 72.6 72.6 48.2 48.2 189.8 189.8 5.7 5.7

403 B4 34.6 72.6 54.2 46.8 243.2 198.9 9.2 5.5

403 B5 34.4 63.3 41.3 27.6 164.3 113.4 7.0 3.5

403 B6 34.6 72.6 75.8 62.0 291.8 240.8 12.9 7.3

403 B7 34.6 72.6 51.6 43.4 309.5 243.2 8.8 5.1

403 C1 34.6 72.6 62.0 55.1 240.8 215.3 10.5 6.5

403 C2 34.6 72.6 71.0 59.1 154.7 110.5 12.1 6.9

403 C3 34.6 72.6 68.9 41.3 266.3 164.3 11.7 4.9

403 C4 34.6 72.6 82.7 62.0 317.2 240.8 14.1 7.3

403 C5 63.3 63.3 31.1 31.1 66.3 66.3 3.9 3.9

403 C6 34.4 63.3 34.4 27.6 138.8 113.4 5.9 3.5

403 C7 34.4 63.3 42.8 30.5 198.9 132.6 7.3 3.8

403 D1 31.1 58.0 56.6 11.1 264.1 28.3 10.2 1.5
121

Grid Prethin Residual Prethin Residual

cell quadratic quadratic Prethin Residual PrethinResidual relative relative

(NW mean mean basal basal tree tree density density

Stand corner) diameter diameter area area density density (metric) (metric)

----Centimeters---- -------m2/ha------- -------No./ha-------

403 D2 63.3 63.3 34.4 34.4 138.8 138.8 4.3 4.3

403 D3 34.6 72.6 68.9 55.1 266.3 215.3 11.7 6.5

403 D4 72.6 72.6 38.6 38.6 88.4 88.4 4.5 4.5

403 D5 34.6 72.6 48.2 41.3 189.8 164.3 8.2 4.9

403 D6 63.3 63.3 27.6 27.6 113.4 113.4 3.5 3.5

403 D7 31.1 58.0 58.0 19.3 278.2 42.4 10.4 2.5

403 E1 34.6 72.6 55.1 48.2 215.3 189.8 9.4 5.7

403 E2 31.1 58.0 28.3 5.4 92.7 14.1 5.1 .7

403 E3 31.1 58.0 67.0 17.1 160.3 42.4 12.0 2.3

403 E4 34.6 72.6 72.7 52.0 221.0 154.7 12.4 6.1

403 E5 63.3 63.3 34.4 34.4 138.8 138.8 4.3 4.3

403 E6 31.1 58.0 65.8 13.4 264.1 28.3 11.8 1.8

403 E7 34.4 63.3 47.8 31.7 309.5 176.8 8.2 4.0

403 F1 31.1 58.0 27.6 20.7 113.4 87.9 4.9 2.7

403 F2 72.6 72.6 48.2 48.2 189.8 189.8 5.7 5.7

403 F3 31.1 58.0 39.8 21.1 121.0 42.4 7.1 2.8

403 F4 34.6 72.6 47.6 46.0 176.8 154.7 8.1 5.4

403 F5 34.6 72.6 55.1 41.3 215.3 164.3 9.4 4.9

403 F6 34.6 72.6 62.0 55.1 240.8 215.3 10.5 6.5

403 F7 31.1 58.0 31.2 10.5 185.5 28.3 5.6 1.4

403 G1 34.6 72.6 48.9 41.1 198.9 154.7 8.3 4.8

403 G2 72.6 72.6 41.3 41.3 164.3 164.3 4.9 4.9

403 G3 72.6 72.6 41.3 41.3 164.3 164.3 4.9 4.9

403 G4 34.6 72.6 75.8 55.1 291.8 215.3 12.9 6.5

403 G5 34.6 72.6 48.2 41.3 189.8 164.3 8.2 4.9

403 G6 34.6 72.6 55.1 48.2 215.3 189.8 9.4 5.7

403 G7 34.4 63.3 27.4 23.1 132.6 88.4 4.7 2.9

1 First, 1 of 3 treatment codes was assigned to each cell based on its

residual basal area: root rot thin (residual basal area < 22.5 m2

/ha);

heavy thin (22.5-37.5 m2/ha); or light thin (> 37.5 m2/ha). Next, dbh

was measured in a subsample of cells in each stand and a mean

quadratic mean diameter was calculated by treatment code for

each stand.
122

Appendix 5

Postthinning Stand

Maps

1

2

3

4

5

6

7

8

A B C D E F GH

Stand 101 Residual Relative Density

Legend: Curtis' (1982) Relative Density

RD > 6.75

RD 4.75 - 6.75

RD 3.25 - 4.75

RD < 3.25
123

 

A B C D E F GH

Stand 103 Residual Relative Density 1

2

3

4

5

6

7

8

Legend: Curtis' (1982) Relative Density

RD > 6.75

RD 4.75 - 6.75

RD 3.25 - 4.75

RD < 3.25
124

A B C D E F GH

Stand 201 Residual Relative Density

1

2

3

4

5

6

7

8

Legend: Curtis' (1982) Relative Density

RD > 6.75

RD 4.75 - 6.75

RD 3.25 - 4.75

RD < 3.25
125

A B C D E F GH

Stand 202 Residual Relative Density

1

2

3

4

5

6

7

8

Legend: Curtis' (1982) Relative Density

RD > 6.75

RD 4.75 - 6.75

RD 3.25 - 4.75

RD < 3.25
126

A B C D E F GH

Stand 302 Residual Relative Density

1

2

3

4

5

6

7

8

Legend: Curtis' (1982) Relative Density

RD > 6.75

RD 4.75 - 6.75

RD 3.25 - 4.75

RD < 3.25
127

A B C D E F GH

Stand 303 Residual Relative Density

1

2

3

4

5

6

7

8

Legend: Curtis' (1982) Relative Density

RD > 6.75

RD 4.75 - 6.75

RD 3.25 - 4.75

RD < 3.25
128

A B C D E F GH

Stand 401 Residual Relative Density

1

2

3

4

5

6

7

8

Legend: Curtis' (1982) Relative Density

RD > 6.75

RD 4.75 - 6.75

RD 3.25 - 4.75

RD < 3.25
129

A B C D E F GH

Stand 403 Residual Relative Density

1

2

3

4

5

6

7

8

Legend: Curtis' (1982) Relative Density

RD > 6.75

RD 4.75 - 6.75

RD 3.25 - 4.75

RD < 3.25
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Carey, Andrew B.; Thysell, David R.; Brodie, Angus W. 1999. The Forest

Ecosystem Study: background, rationale, implementation, baseline conditions,

and silvicultural assessment. Gen. Tech. Rep. PNW-GTR-457. Portland, OR:

U.S. Department of Agriculture, Forest Service, Pacific Northwest Research

Station. 129 p.

The Forest Ecosystem Study (FES) came about as an early response to the need for innovative silvicultural methods designed to stimulate development of late-successional attributes in

managed forests—a need ensuing from the exceptional and long-standing controversies over

old-growth forests and endangered species concerns in the Pacific Northwest. In 1991,

scientists with the FES applied experimental, variable-density thinning to even-aged Douglasfir (Pseudotsuga menziesii (Mirbel) Franco) forests on the Fort Lewis Military Reservation in

western Washington after having first accumulated extensive baseline data on arboreal

rodents, small mammals, trees and other vascular plants, and fungi. We present study

background, rationale, baseline conditions, and selected preliminary responses, as well as

a silvicultural assessment of the variable-density thinning.

Keywords: Variable-density thinning, Pacific Northwest, Douglas-fir, biodiversity, northern

flying squirrel, truffle, Forest Ecosystem Study, experimental silviculture.

The Forest Service of the U.S. Department of

Agriculture is dedicated to the principle of multiple

use management of the Nation’s forest resources for

sustained yields of wood, water, forage, wildlife, and

recreation. Through forestry research, cooperation with

the States and private forest owners, and management

of the National Forests and National Grasslands,

it strives—as directed by Congress—to provide

increasingly greater service to a growing Nation.

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, or

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 and TDD).

USDA is an equal opportunity provider and employer.

Pacific Northwest Research Station

333 S.W. First Avenue

P.O. Box 3890

Portland, Oregon 97208-3890
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