United States

Department of

Agriculture

Forest Service

Pacific Northwest

Research Station

General Technical

Report

PNW-GTR-558

September 2002

Managing Young Upland Forests in

Southeast Alaska for Wood Products,

Wildlife, Aquatic Resources, and Fishes:

Problem Analysis and Study Plan

Mark S. Wipfli, Robert L. Deal, Paul E. Hennon, Adelaide C. Johnson,

Toni L. De Santo, Thomas A. Hanley, Mark E. Schultz, Mason D. Bryant,

Richard T. Edwards, Ewa H. Orlikowska, Takashi Gomi
Mark S. Wipfli is a research ecologist, Pacific Northwest Research Station, 1133 N.

Western Ave., Wenatchee, WA 98801; Robert L. Deal is a research silviculturist,

Pacific Northwest Research Station, 620 SW Main St., Suite 400, Portland, OR 97205;

Takashi Gomi is a hydrologist, Department of Forest Resources Management,

University of British Columbia, Vancouver, BC; Paul E. Hennon is a plant pathologist,

Adelaide C. Johnson is a hydrologist, Toni L. De Santo is an avian ecologist,

Thomas A. Hanley is chief wildlife biologist, Mark E. Schultz is an entomologist,

Mason D. Bryant is a research fishery biologist, Richard T. Edwards is a research

aquatic ecologist, and Ewa H. Orlikowska is a forest ecologist, Pacific Northwest

Research Station, 2770 Sherwood Lane, Juneau, AK 99801.

Authors
Abstract Wipfli, Mark S.; Deal, Robert L.; Hennon, Paul E.; Johnson, Adelaide C.;

De Santo, Toni L.; Hanley, Thomas A.; Schultz, Mark E.; Bryant, Mason D.;

Edwards, Richard T.; Orlikowska, Ewa H.; Gomi, Takashi. 2002. Managing young

upland forests in southeast Alaska for wood products, wildlife, aquatic resources, and

fishes: problem analysis and study plan. Gen. Tech. Rep. PNW-GTR-558. Portland,

OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research

Station. 64 p.

Red alder (Alnus rubra Bong.) appears to influence the productivity of young-growth

conifer forests and affect the major resources (timber, wildlife, and fisheries) of forested

ecosystems in southeast Alaska. We propose an integrated approach to understanding

how alder influences trophic links and processes in young-growth ecosystems. The

presence of red alder is expected to increase understory biomass, and aquatic, riparian,

and terrestrial invertebrate abundance, providing more food for herbivores, fish, and birds.

We predict that most red alder trees will die standing, and woody debris will be small

and mobile in streams. Nitrogen fixation by red alder in mixed stands may result in

larger, more commercially valuable conifers. Inclusion of red alder in the regenerating

stand may therefore mitigate some negative impacts of clearcutting, and may increase

total wood production from the landscape.

Keywords: Red alder, young-growth management, vegetation, wildlife, fish, invertebrates.
1 Introduction

2 Justification and Management Implications

5 Expected Science and Management Inferences

6 Background on Red Alder

6 Establishment

6 Regeneration

7 Stand Density and Tree Growth

7 Plant Associates

8 Growth Patterns

8 Succession

8 Nitrogen Fixation

9 Riparian Nitrogen Levels

9 Effect of Red Alder on Forest Growth

10 Effect of Red Alder on Stand Structure and Understory

10 Impacts of Clearcutting Coniferous Forests

11 Lifespan of Red Alder

12 Pathogens

13 Mode of Death

14 Decay

14 Fungi

15 Decay Resistance

15 Decay of Wood in Streams

16 Role of Riparian Forests in Stream Processes

16 Effect of Riparian Trees on Terrestrial Invertebrates in Stream Food Webs

18 Impacts of Changes in Riparian Forest Canopy

18 Bird Abundance Associated with Red Alder

19 Commercial Value of Red Alder

20 Perceptions of Red Alder

20 Importance of Upland Forests and Associated Red Alder for Aquatic Habitats

20 Objectives and Hypotheses

27 Quality Assurance-Quality Control Procedures

27 Documenting Measurement Error

28 Data Management and Archiving

28 English Equivalents

Contents
29 Acknowledgments

29 Literature Cited

47 Appendix 1: Methods for Sampling Stand Density, Vegetation, and

Tree Mortality

53 Appendix 2: Methods for Terrestrial Invertebrate Sampling

55 Appendix 3: Methods for Wildlife Sampling

57 Appendix 4: Methods for Aquatic and Riparian Invertebrate Sampling

60 Appendix 5: Methods for Fish Sampling

62 Appendix 6: Methods for Sampling Geomorphic Processes
1

Introduction This study proposes an integrated approach to examine the function of young-growth

ecosystems (fig. 1) and to understand how red alder (Alnus rubra Bong.) influences

trophic links and processes (figs. 2 and 3) in managed young-growth ecosystems.

Forest managers have historically regarded red alder as an undesirable species and

have attempted to remove it from riparian and upland forest stands. However, red

alder appears to have a positive influence on the productivity of young-growth conifer

forests and associated resources in southeast Alaska. Our study will:

• Assess physical disturbances associated with red alder establishment.

• Evaluate the role of headwater processes within a landscape to assess conditions

necessary for the establishment of alder-conifer stands.

• Assess history of physical disturbance processes (such as landslides) within the

landscape to evaluate possible habitat changes associated with these disturbances

in managed forests.

• Investigate whether red alder in mixed stands enhances conifer growth and wood

production.

• Contrast the mode of death of red alder trees and conifers (i.e., die standing or by

uprooting or bole snapping).

• Compare biomass of understory vegetation and forage for herbivores and

invertebrates across a range of alder-conifer mixtures (0 to 80 percent alder).

• Assess the role of alder across a range of alder-conifer mixtures on the abundance

of aquatic, riparian, and terrestrial invertebrates that provide food for fish and birds.

• Determine whether red alder influences stream nitrogen levels.

• Assess critical processes for maintaining downstream fish habitat including the

supply, storage, and transport of woody debris and sediment.

• Assess red alder’s impact on woody debris and sediment movement in streams.

• Measure fish abundance in relation to red alder.

• Determine the tradeoffs and compatibilities between the amount of alder in a

stand, production of understory vegetation, growth and yield of Sitka spruce

(Picea sitchensis (Bong.) Carr.) and western hemlock (Tsuga heterophylla (Raf.)

Sarg.), woody debris, and fish food resources.

• Address the potential of alder as a “tool” for restoring important ecosystem

functions in regenerating forests of southeast Alaska.

• Synthesize findings by describing the simultaneous beneficial and detrimental

influences of red alder to several key resources across a range of its occurrence

in mixed-species stands.

We hypothesize that management of red alder can enhance the major resources

(timber, wildlife, and fishes) of forested ecosystems, and that the presence of red

alder in young-growth stands may mitigate the impacts of clearcutting in areas where

pure conifer regrowth would compromise these resources.
2

There are three major integrating themes in this study:

1. Influence of alder in the ecosystem: vegetation, birds, invertebrates,

wood, fishes (fig. 1).

2. Flow of wood among habitats (fig. 2).

3. Influence of alder on ecosystem links and processes (fig. 3).

Since the 1950s, clearcutting has been the dominant timber management practice in

southeast Alaska forests. The dense, uniform, even-aged stands that develop after

clearcutting have many negative consequences for wildlife and fish (Dellasala et al.

1996; Hanley 1993; Schoen et al. 1981,1988; Thedinga et al. 1989; Wallmo and

Schoen 1980). Canopy closure generally occurs 25 to 35 years after cutting and is followed by a nearly complete elimination of understory vegetation for 100 years or longer

(Alaback 1982, 1984b; Tappeiner and Alaback 1989). After the canopy closes above

small streams, trophic status changes from autotrophic (dominated by organisms that

manufacture their own energy) to heterotrophic (dominated by consumers) (Hetrick et

al. 1998a, Sedell and Swanson 1984) and may affect overall aquatic productivity. The

Figure 1—Major links among ecosystem components in young-growth conifer and mixed red alder-conifer forests.

Justification and

Management

Implications

Woody debris Aquatic arthropods Water Sediment Nutrients 

Water and fishes

Wood products

Arthropods

(terrestrial and riparian)

Dead trees

Forest stand dynamics and productivity

Conifer-dominated Alder-dominated

Wildlife

Downstream transport of

Natural physical

disturbance

processes and

human impacts

Natural physical

disturbance

processes and

human impacts

Habitat

Habitat

Habitat Food

Food

Understory vegetation biomass and composition

Habitat

Habitat

Habitat Harvest

Food

Food 

Habitat

Food

Food
3

removal of streamside timber also reduces the amount and size of large wood in

a stream, with a subsequent loss of bird1 and fish habitat (Bisson et al. 1987, Bryant

1985). Changes in forest structure result in an alteration of supply, storage, and transport of woody debris and sediment through processes such as mass wasting, windsnap

and blowdown, and bank erosion (Johnson et al. 2000, Smith et al. 1993). Subsequently, there is increasing interest in developing forest management practices that

maintain or enhance biodiversity and assure long-term sustainability of forest products,

wildlife, and aquatic resources.

Recent studies of young-growth stands of red alder mixed with conifers indicate that the

presence of alder may mitigate some of the impacts of clearcutting in southeast Alaska.

Mixed alder-conifer stands have species-rich, highly productive understory vegetation

with biomass similar to that of old-growth stands of the region (Deal 1997, Hanley and

Barnard 1998, Hanley and Hoel 1996). Habitat quality for small mammals in even-aged

alder-conifer stands may be equal to that of old-growth forests (Hanley 1996; Hanley

1 De Santo, T.L. 1996. Unpublished data. On file with: Forestry

Sciences Laboratory, 2770 Sherwood Lane, Suite 2A, Juneau,

AK 99801.

Figure 2—Flow and function of wood among habitats.

Live trees (uplands)

Habitat and food for 

arthropods and wildlife

Dead trees

Habitat and food for 

arthropods and wildlife

Wood products

Habitat for humans

from forest

ecosystem

Tree mortality Timber harvesting

Forest floor

woody debris

Habitat and food

Riparian

woody debris

Habitat and food

Headwater stream

woody debris

Habitat and food

Breakage, windthrow, rotting

Mass wasting, floods

Woody debris in anadromous streams (lowlands)

Fish habitat, invertebrate food and habitat, bank stabilization, sediment routing and storage

Removed
4

and Barnard 1999a, 1999b). Although inclusion of alder will not mitigate all wildlifehabitat problems (e.g., lack of snow interception for deer winter range), it may provide

more benefits than would thinning of even-aged conifer stands. Attempts to reestablish

understory herbs and shrubs through thinning young-growth conifer stands have led to

conifer regeneration with little new herbaceous colonization (Deal and Farr 1994). Red

alder may convey further benefits in riparian forests. Riparian forests with some red

alder appear to produce more prey for fishes than conifer riparian forests (Piccolo and

Wipfli 2002, Wipfli 1997). This is significant because over half of the prey biomass ingested by juvenile salmonids in southeast Alaska is terrestrial and originates from adjacent riparian vegetation. If similar processes occur in upland forests, the presence of

red alder may increase invertebrate production, providing more food for animals such

as birds, bats, small mammals, and fish, in turn affecting their abundance and production.

This is an integrated study (fig.1) that compares mixed alder-conifer stands with pure

conifer stands to assess differences in physical disturbance history associated with tree

composition; stand structure and tree mortality; understory vegetation; invertebrate,

bird, and bat communities; and food resources for fish and birds (fig. 3). This study will

take the first step toward establishing an integrated research program to address ecological processes critical for the establishment and growth of alder-conifer forests.

Figure 3—Proposed influence of red alder on ecosystem links and processes.

Overstory species

Alder Conifer

Birds

Species and abundance

Forest

arthropods

Species and abundance

Headwater

stream

invertebrates

Species and abundance

Fish and fish habitat

Fish species and abundance,

quality and quantity of habitat

Understory

Species composition and abundance
5

In southeast Alaska, red alder is expected to increase the abundance of terrestrial invertebrates, providing a greater food resource for insectivores, such as birds (figs. 1

and 3). In riparian forests, increased production of invertebrates is expected to produce

more food for fish. A greater biomass of understory vegetation is expected to occur in

young forests with red alder; thus, habitat for birds and forage available to herbivores,

such as deer, is expected to increase. Red alder is expected to produce rapid early

growth but experience higher mortality than conifers as young forests develop. Most

trees are expected to die standing regardless of treatment; thus, woody debris produced is expected to be small, partially decomposed, and devoid of large root systems.

Such woody debris would be mobile and transient in streams, allowing sediment to

move unimpeded downstream (fig. 2). Reduced conifer wood production is expected to

occur with the presence of red alder, but individual conifers may be larger and more

commercially valuable in mixed stands. Diversity of wood products is expected to be

greater in mixed stands, and alder may be a valuable wood product in the future. Inclusion of red alder in a regenerating stand may allow for clearcutting in areas where

purely even-aged conifer stands would compromise other resources (e.g., fish and

wildlife). Inclusion of red alder also may increase total wood production from the landscape.

A considerable body of literature exists on the growth of red alder in forests of Oregon,

Washington, and British Columbia. Some of this literature deals with mixed red alder/

Douglas-fir forests (Pseudotsuga menziesii (Mirb.) Franco). We expect to observe

some similar general patterns of red alder growth, competition with conifers, and mortality in southeast Alaska. The distribution of red alder regeneration within the landscape may be different in southeast Alaska; red alder requires more soil disturbance in

Alaska to colonize sites after timber harvesting. Processes such as tree growth and

development of forests may be somewhat slower in Alaska. The age at which red alder

reaches senescence and gives way to competition with conifers may be considerably

different in Alaska. Plants that have a dominant influence on understory conditions in

alder forests to the south (e.g., salmonberry (Rubus spectabilis Pursh)) may be less

aggressive colonizers in Alaska, and may be replaced by understory plants preferred

for browse by Sitka black-tailed deer (Odocoileus hemionus sitkensis). The decay of all

types of wood is expected to be slower in Alaska owing to cooler average air and water

temperatures. Slower decay is expected to allow for a greater persistence of both

legacy and newly recruited wood for physical and biological functions in streams. We

expect the influence of red alder on streams to differ with landscape location, or topographic position, which will dictate natural large woody debris patterns (storage and

transport trends). Once these patterns of disturbance are understood, the effects of

alteration of forests (conversion from old- to young-growth forests dominated with alder

and conifer) on aquatic habitats can be assessed.

Although this study centers on young-growth forests in Alaska, the potential exists to

expand our understanding of the occurrence and role of alder in unmanaged ecosystems in southeast Alaska and other ecosystems across the landscape. Southeast

Alaska contains more extensive tracts of virgin, old-growth forest than does any other

part of the United States. Alaska offers unique opportunities to study ecological interactions among alder, site conditions, and animals in an evolutionary context. Standards

for ecological reference sets of “natural” conditions can be established more accurately

in Alaska than elsewhere. Practical applications of study results are immediate in

Alaska, and potential opportunities for other outcomes (e.g., ecological reference sets)

Expected Science

and Management

Inferences
6

are greatest in Alaska. Differences owing to latitude need to be quantified before research results from a particular geographic area are applied elsewhere; however, our

approach of integrating disparate disciplines can be used as a framework for other

studies on the manipulation of vegetation to achieve complex resource objectives.

Red alder, the most common hardwood tree in the Pacific Northwest, is a relatively

short-lived, shade-intolerant pioneer with rapid juvenile height growth (Harrington

1990). Red alder is the only commercial tree species west of the Rocky Mountains

with the capacity to fix atmospheric nitrogen, and alder is an important species for the

hardwood industry in the Pacific Northwest (Heebner and Bergener 1983, Hibbs et al.

1994). Its range extends from southern California to southeast Alaska (Harrington

1990). In Alaska, red alder is found throughout the southeastern panhandle northwest

to Yakutat Bay in riparian areas, along roadsides and avalanche slopes, on stream

bottoms with rocky, moist soils, and along beaches where creeks enter seas (Hulten

1968, Viereck and Little 1972).

Little information is available about red alder and its ecological role in Alaska. Basic

information on silvics, stand growth and yield, tree species mixtures, succession, tree

mortality and decay, and understory vegetation is lacking, and most information about

alder in Alaska is speculative or based on research from other regions.

Low winter temperatures and lack of precipitation during the growing season appear

to be the primary limits to the range of red alder (Harrington et al. 1994). It grows in

climates from humid to superhumid with annual precipitation from 400 to 5600 mm.

Red alder grows in a wide range of soils, from well-drained gravels or sands to clays

and organic soils. It grows primarily on soils of the orders Inceptisols and Entisols but

also occurs on some Alfisols, Ultisols, and Histosols (Harrington 1990). Alder can tolerate poor drainage and some flooding during the growing season. It is not commonly

found on droughty soils. The most productive stands are found on deep alluvial soils in

river and stream flood plains; however, some productive stands also are found on upland sites on residual or colluvial soils (Harrington et al. 1994). Tree development is

best in elevations below 450 m in British Columbia, Washington, and northern Oregon.

In Alaska, red alder generally occurs near sea level.

As a pioneer species, red alder requires high light levels and is common on exposed

mineral soil. Its ability to fix atmospheric nitrogen permits establishment on geologically

young or disturbed sites with little soil nitrogen. It can form pure stands on alluvium,

avalanche paths, or other disturbed sites. Red alder is an aggressive pioneer on avalanche paths, roadsides, log landings, skid trails, and other areas where mineral soil

has been freshly exposed to seedfall (Harrington et al. 1994, Harris and Farr 1974). In

Alaska, red alder is commonly found along beaches and streams, on avalanche tracks

and landslides, and as a pioneer species with Sitka spruce, black cottonwood (Populus

trichocarpa Torr. and Gray) and willows (Salix spp.) on land recently exposed by glacial

retreat or land uplift (Harris and Farr 1974, Ruth and Harris 1979). In southeast Alaska,

intensive logging since the 1950s has increased the amount of alder, particularly in

upland areas where heavy soil disturbance resulted from tractor logging and high-lead

cable operations.

Red alder can reproduce by seed or can sprout vigorously from the stump when

young. Alder seeds have little or no dormancy, and germination of stored seeds is

not improved by stratification (Berry and Torrey 1985, Radwan and DeBell 1981).

Background on

Red Alder

Establishment

Regeneration
7

Alder germination is controlled by light quality (Bormann 1983, Haeussler and

Tappeiner 1993), and germination is reduced to varying degrees by the presence of

understory vegetation. Alder requires more light than its associated tree species, and

is classed as intolerant of shade (Minore 1979). Haeussler and Tappeiner (1993) concluded that buried seedbank is not important in alder. Red alder sprouts vigorously

from the stump when young and can be coppiced on short cycles (Harrington and

DeBell 1984). Older trees rarely sprout, and coppice regeneration cannot be expected

after saw-log size material is harvested (Harrington 1984). Because of reduced sprouting vigor, manual cutting of older alder trees can be an effective method of alder control

(DeBell and Turpin 1989), but results from cuts made at different times during summer

can differ (Pendl and D’Anjou 1990).

Density in natural stands can be as high as 124,000 seedlings per hectare at age 5

(DeBell 1972) and 1,665 trees per hectare at age 20 (Worthington et al. 1960). Live

crown ratios in crowded pure stands are very low, and trees often form narrow, domelike crowns. Branch retention and crown shape are strongly related to light levels in the

canopy. Trees grown in low densities develop large lower branches that live longer and

take longer to fall off than do branches that develop in more dense stands (Harrington

et al. 1994). Red alder self-prunes very well when grown in dense stands. Shaded

lower branches rapidly die and fall off, leaving clear boles. Epicormic branching has

been reported after thinning, especially after late or heavy thinning (Berntsen 1961b,

Smith 1978, Warrack 1964).

Red alder is relatively short-lived, maturing at about 60 to 70 years and reaching a

maximum age of about 100 years (Worthington et al. 1962). On favorable growing

sites, trees can be 30 to 40 m tall and 55 to 75 cm in diameter. Red alder has rapid

juvenile height growth and on favorable sites can grow 1 m or more the first year,

and on all but the poorest sites seedlings surpass breast height (1.3 m) the second

year (Harrington and Curtis 1986, Smith 1968). On favorable sites, alder trees may

be 9 m tall at age 5, 16 m at age 10, and 24 m at age 20 (Harrington 1990). In western

Oregon, Zavitkovski and Stevens (1972) reported net primary productivity in red alder

stands during the years of maximum growth (10 to 15 years old) averaged 26 t/ha per

year. In pure stands on favorable sites, red alder can achieve annual growth rates

of 21 m3/ha in pulpwood rotations of 10 to 12 years, and 14 m3/ha in saw-log rotations

of 30 to 32 years (DeBell et al. 1978). Radial growth generally begins in mid-April

and continues until mid-September in the Puget Sound area of Washington state

(Reukema 1965). Growth slows after the juvenile stage. Stand maximum cubic volume

(about 500 m3/ha) is attained at age 50 to 70 (Chambers 1983, DeBell et al. 1978,

Skinner 1959, Worthington et al. 1960). Associated conifers have slower juvenile

growth, but they sustain height growth longer than alder. On an average upland site,

Douglas-fir and red alder can attain the same height at about age 45 (Williamon 1968).

Red alder grows in both pure and mixed stands, with pure stands typically confined

to stream bottoms and lower slopes (Harrington et al. 1994). It is more common in

mixed stands and occurs as a minor component in most north Pacific cover types

(Eyre 1980). Tree associates include Douglas-fir, western hemlock, western redcedar (Thuja plicata Donn ex D. Don), Sitka spruce, black cottonwood, and willow.

Shrub associates include salmonberry, red elderberry (Sambucus racemosa L.), salal

(Gaultheria shallon Pursh) and devil’s club (Oplopanax horridus Miq.). Herbaceous

Stand Density and

Tree Growth

Plant Associates
8

plant associates include skunkcabbage (Lysichiton americanus Hulten & St. John),

western swordfern (Polystichum munitum (Kaul fuss) K. Presl), and ladyfern (Athyrium

filix-femina (L.) Roth).

Many alder stands in western Oregon have few associated conifers, and these stands

may ultimately be replaced by shrub communities (Carlton 1990, Newton et al. 1968).

Red alder is common on previously logged sites in the Pacific Northwest (Hibbs et al.

1994), where understories tend to be dominated by swordfern or salmonberry or both,

often to the near-term exclusion of conifer regeneration (Newton et al. 1968, Tappeiner

and Zasada 1993, Tappeiner et al. 1991). Salmonberry in the coastal Pacific Northwest

is a particularly aggressive shrub that often forms dense shrub canopies. This makes it

difficult for conifers to become established from seed and can create a serious problem

for tree regeneration and growth (Howard and Newton 1984, Knapp et al. 1984, O’Dea

1992, Tappeiner and Zasada 1993, Tappeiner et al. 1991, Walstad and Kuch 1987,

Walstad et al. 1986, Zasada et al. 1992). The abundant shrub layer also may explain

the scarcity of tree regeneration in streamside areas (Hibbs and Giordano 1996). Neither Henderson (1970) nor Carlton (1990) found many tree seedlings in their studies of

riparian and upland alder stands.

Although most of the research in mixed alder-conifer stands has been descriptive

(as will be this study), some general growth trends are apparent. Alder must remain

in the upper canopy to survive in mixed stands. Reaction of alder to competition is

influenced by many factors, including the size, species composition, and density of the

competing vegetation in the upper canopy and in the understory. Stand density strongly

influences survival and growth of alder. The growth of closely spaced dominant alder

was reported to decrease with increasing density of subordinate Douglas-fir (Shainsky

and Radosevich 1991). Turnbull (1963) provides a scholarly treatise on assessing

growth of mixed stands, including problems with quantitative comparisons of pure and

mixed stands and their species components.

Successional sequences in mixed alder-conifer stands are not well understood. Observation of mature forests in the Pacific Northwest suggests that alder is replaced by

longer lived, more shade-tolerant conifers that sustain growth rates for longer periods.

Alder’s rapid early growth and high stem densities make it difficult for other shadeintolerant species to regenerate and grow if they do not become established at the

same time alder invades a disturbed area. Douglas-fir can be eliminated in dense,

young alder stands whereas more shade-tolerant species such as western hemlock,

Sitka spruce, and western redcedar can survive, grow into the canopy, and ultimately

dominate the site (Harrington et al. 1994).

The ability of red alder to fix atmospheric nitrogen can increase both nitrogen content

and nitrogen availability in soil. Nitrogen fixed in the nodules is added to the soil in four

ways: direct excretion from living roots and nodules, decomposition of dead roots and

nodules, leaching from foliage, and decomposition of litter rich in nitrogen. Maximum

annual fixation rates of 320 kg/ha (Trappe et al. 1968) in pure stands and 130 kg/ha in

mixed stands (Binkley 1981) have been reported. Red alder increases the organic content of soils (Tarrant and Miller 1963, Trappe et al. 1968), and even if alder is shaded

out following canopy closure in mixed stands, it may make substantial contribution to

soil nitrogen prior to that time (Berg and Doerksen 1975). Soil structure and organic

matter are thought to be major drivers of ecosystem productivity (Powers 1991), and

Growth Patterns

Succession

Nitrogen Fixation
9

alder increases organic matter in the soil by producing large quantities of rapidly decomposable biomass. Alder produces more litter than do conifers (Zavitkovski and

Newton 1971), and the litter of other trees is more rapidly decomposed and incorporated into soil organic matter when mixed with nutrient-rich alder litter (Bormann and

Sidle 1990).

A well-documented study of mixed alder-conifer stands at the Wind River research

site in southwestern Washington compared a nitrogen-deficient stand of mixed Douglasfir and red alder with adjacent pure stands of Douglas-fir (Miller and Murray 1978,

Tarrant 1961, Tarrant and Miller 1963). After 48 years, mixed alder-conifer stands

showed slightly increased growth of Douglas-fir and significant growth of alder, with

overall wood production nearly double that of adjacent pure Douglas-fir stands.

Douglas-fir site index (50 year) was increased by an average of 6.4 m. Improved soil

fertility was suggested to be largely responsible for the improved growth of these stands

(Bormann et al. 1994). Other authors have indicated that on poor sites, alder may increase production in Douglas-fir plantations (Atkinson et al. 1979, Tarrant et al. 1983).

The high rainfall and rapid discharge response to storms characteristic of Pacific Northwest streams rapidly leach minerals from watersheds. Consequently, streams in the

region generally contain low concentrations of limiting plant nutrients, particularly nitrogen and phosphorus (Gregory et al. 1987). These low nutrient levels, combined with

low light levels beneath dense riparian canopies, limit primary production within streams.

In areas with significant anadromous fish runs, low nutrient inputs from catchments are

partially offset by the input of nitrogen and phosphorus from spawning fish (Minikawa

and Gara 1999, Rice and Bailey 1980). Within the upper reaches, however, anadromous nutrient inputs are indirect and small, and autochthonous production is probably

limited by nitrogen or phosphorus availability.

Logging can increase primary production by reducing light limitation, and by temporarily increasing nutrient inputs from decomposing logging debris. Once the flush of

postharvest nutrients moves through the system, however, concentrations return to

preharvest levels in the absence of other factors. Where regrowth of nitrogen-fixing

species such as red alder occurs, nitrogen inputs to streams may be elevated for decades after harvest. Red alder stands show significantly increased soil nitrogen stocks

from a few years after harvest until the stand senesces and is replaced by conifers

(Luken and Fonda 1983). Groundwater and hyporheic flows move alder-derived nitrogen into adjacent streams and can dramatically increase nutrient stocks and algal

standing stocks in streams with sufficient light levels (Fevold 1998).

Little research has been done to quantify the impact of red alder on riparian nitrogen

levels and stream production, or to assess how its significance varies with location and

stream type. The net effect of red alder on stream nutrient stocks presumably depends

on its location (riparian versus upland), soil characteristics, channel morphology, and

hydrologic response.

The Wind River site also has been used to study the effect of mixed alder-conifer

stands on stand growth and yield (Binkley and Greene 1983), mensuration (Miller and

Murray 1978, Tarrant 1961), soil chemistry (Tarrant and Miller 1963), Douglas-fir nutrition (Reid 1983, Tarrant 1961), economics (Atkinson et al. 1979, Cole et al. 1978), and

biogeochemistry (Binkley and Sollins 1990, Binkley et al. 1992). Researchers also investigated the effect of distance on the influence of red alder’s nitrogen fixation. Miller

et al. (1993) found that some positive influence of nitrogen-fixing red alder extended

Riparian Nitrogen Levels

Effect of Red Alder on

Forest Growth
10

about 15 m beyond the edge of a mixed alder-conifer stand, indicating that the influence of alder may extend beyond immediately adjacent trees. These results may have

implications for areas where red alder is mixed in stands through skid roads, along

streams, or irregularly distributed across the landscape.

Research at a mixed alder-conifer stand at Cascade Head in coastal Oregon has

yielded different results in productivity. The Cascade Head stand is much more productive than the Wind River stand. At Cascade Head, researchers found no appreciable

difference in productivity between the mixed alder-conifer stand and an adjacent pure

conifer stand. Both biomass and net primary productivity were higher in the pure conifer

stand than in the mixed stand (Binkley et al. 1992). On nitrogen-deficient soils, where

the rate of nitrogen supply limits forest productivity, the addition of alders to conifer

stands increased productivity, nitrogen levels, and nitrogen availability without decreasing soil pH. On fertile sites where growth was less limited by nitrogen availability of

nitrogen also increased, but with no corresponding increase in productivity (Binkley

et al. 1992).

The mixed alder-conifer site at Cascade Head also has been used to investigate effects of alder on stand structure and understory vegetation (Franklin and Pechanec

1968, Pechanec and Franklin 1968), tree growth (Berntsen 1961a), nutrient cycling

(Binkley et al. 1992, Bollen et al. 1967, Tarrant et al. 1968) and mensuration (Miller

and Murray 1978). Franklin and Pechanec found that the species richness and cover

of herbaceous plants was greatest under pure alder and least under pure conifer, with

mixed stands intermediate. Ground-dwelling cryptogams were more common and better developed in the mixed alder-conifer and pure conifer stands. Some suppressed

Sitka spruce saplings were present in the mixed and pure alder stands, but new conifer

seedlings were found only in the pure conifer stand.

Several studies suggest that red alder can inhibit Douglas-fir growth. In a reconstruction study of red alder and Douglas-fir in the Pacific Northwest, Newton et al. (1968)

assessed height growth of red alder and Douglas-fir in 39 nearly pure red alder stands

in western Oregon. They determined that red alder remains dominant for 25 to 35 years

on most sites, and 40 years or more on wetter sites. Emergence of Douglas-fir from

beneath red alder depends on its ability to grow at open-grown rates while being suppressed. Newton et al. (1968) concluded that Douglas-fir would be unable to maintain

height growth while being suppressed, but more shade-tolerant species might be able

to do so. In another study, Miller and Murray (1978) assessed growth in four mixed red

alder/Douglas-fir stands. Conifers showed reduced growth in heavily stocked mixed

alder/Douglas-fir stands on higher quality sites. Douglas-fir generally emerged from the

alder canopy and attained dominance in mixed stands by 25 to 35 years on drier sites

and about 10 years later on wetter sites. The researchers suggested that to attain full

stocking of Douglas-fir, the stocking and distribution of red alder would need to be controlled either by thinning or by using herbicides to kill the alder. They recommended a

stocking of 20 to 40 red alder trees per acre (50 to 100 per ha), uniformly distributed

throughout the stand.

Clearcut logging has negative consequences for wildlife habitat in southeastern Alaska

(Hanley 1993, Samon et al. 1989; Schoen et al. 1981, 1988; Wallmo and Schoen

1980). The principal problem is that dense conifer regeneration and canopy closure

result in a depauperate understory from about 25 to 150 years stand age (Alaback

1982, 1984a, 1984b). Silvicultural thinning of regenerating stands promotes tree growth

Effect of Red Alder on

Stand Structure and

Understory

Impacts of Clearcutting

Coniferous Forests
11

but has insignificant effects on understory, with widely spaced thinnings resulting in

a second layer of western hemlock regeneration (Deal and Farr 1994). Even-aged

stands with depauperate understories have become common since the widespread

use of high-lead clearcut logging, where logs are transported through the air and soil

disturbance is minimized. Earlier clearcut logging techniques involved considerable

soil disturbance and resulted in more red alder establishment and dominance on

logged sites (Ruth and Harris 1979). Until recently, those sites were largely ignored

in analyses of secondary succession in southeastern Alaska. Alaback (1982, 1984a,

1984b) deliberately excluded red alder patches from his chronosequence studies because modern logging methods minimize the presence of red alder.

Mixed red alder-conifer even-aged stands do not appear to have the same consequences for wildlife as pure conifer even-aged stands in southeastern Alaska. Understory of red alder-dominated, even-aged second-growth stands appears to be quite

different from that of pure conifer stands. Hanley and Hoel (1996) found no significant

differences in total understory biomass between 40-year-old red alder riparian forests

and old-growth upland and old-growth riparian forests. Deal (1997) found total understory canopy coverage in even-aged, red alder-conifer stands more than six times that

of a nearby even-aged, spruce-hemlock stand. Hanley and Barnard (1998) studied

within-stand variation in understory species composition and biomass in 16 evenaged stands of mixed red alder-Sitka spruce-western hemlock. The sites were upland

stands 28 to 39 years old. Understory biomass averaged 175.4 ± 54.6 kg/ha (“mixed”

microsites) to 570.0 ± 111.0 kg/ha (“alder” microsites) and was comparable to oldgrowth forest in species richness. Forbs and ferns were more abundant under alder

overstories than under conifer overstories. A calculation of food value of the understory for black-tailed deer illustrates the ecological significance of these differences:

202 deer-days per ha in the “alder” microsites, 73 deer-days per ha in the “mixed”

alder-conifer microsites, and 45 deer-days per ha in the “conifer” micro-sites. Hanley

(1996) earlier found no significant differences in the abundance and growth rates of

deermice (Peromyscus keeni), long-tailed voles (Microtus longicaudus littoralis), and

common shrews (Sorex cinereus streatori) in mixed red alder-conifer even-aged

stands compared with nearby old-growth stands. Similar results have been reported

for small mammals, amphibians, and birds in the Pacific Northwest (McComb 1994,

McComb et al. 1993).

Although red alder-dominated and mixed alder-conifer stands in the Pacific Northwest

have been studied from a timber perspective (e.g., Hibbs et al. 1994), few studies of

the biomass or successional dynamics of their understories have been conducted

(Carlton 1990, Henderson 1970, Newton and Cole 1994). Whereas salmonberry dominates the shrub layer in much of the Pacific Northwest, devil’s club is the dominant

understory shrub in southeast Alaska and creates different understory conditions (Deal

1997, Hanley and Barnard 1998, Hanley and Hoel 1996, Newton and Cole 1994). The

abundance of herbs in Alaska understories appears to be a major difference from the

Pacific Northwest. More information is needed about ecological interactions between

red alder, site factors (including disturbance), and understory vegetation before we can

understand cause-and-effect or design optimal silvicultural applications.

Red alder is relatively short-lived compared to its associated trees species.

Worthington et al. (1962) give the maximum age for red alder at about 100 years.

Smith (1968) reports that by 50 years of age, pure alder stands on highly productive

Lifespan of Red Alder
12

Douglas-fir sites in British Columbia were “breaking up.” Individual trees showed dieback and other disease symptoms as early as 40 years of age. On less productive

sites, disease symptoms and mortality occurred at stand age 60 to 70, with few red

alder stands remaining intact beyond 100 years. In Oregon, Washington, and British

Columbia, mortality of red alder increases rapidly in stands over 90 years old, and little

alder remains by the age of 130 years (Newton and Cole 1994).

Red alder has rapid early height growth in the Northwest (DeBell and Giordano 1994;

DeBell and Wilson 1978; Hoyer et al. 1978; Smith 1968, 1978; Smith and DeBell 1974),

an ecological strategy common in pioneering plants of disturbed sites. With few resources allocated for defense, red alder is attacked by a number of biotic agents. Newton (in Hoyer et al. 1978) suggests that pure alder stands show little stratification, that

alder needs continual dominance, and that tree mortality occurs soon after a tree drops

to codominant status. Red alder is replaced by longer lived, more tolerant conifers that

sustain growth rates longer. The time required for this transition has not been well

documented (Harrington et al. 1994). It is unclear whether the senescence and mortality of red alder that occurs as young-growth forests develop in Alaska results from

pathogenic factors or from red alder’s inability to maintain rapid height growth in competition with conifers.

Many pathogens occur on red alder in the Pacific Northwest and Alaska, but the extent

to which they cause tree mortality is unknown (Driver 1978). Much of the literature

provides lists of fungi on red alder. Funk (1981) lists 12 fungi that are pathogens or

weak pathogens on red alder in British Columbia. The ecology of these diseases is

not addressed, but Funk’s book gives microscopic characteristics that are helpful in

identifying fungi. Hepting (1971) provides a general review of the pathology of red

alder. Many fungi attack the leaves of red alder, but none are considered serious diseases. Taphrina japonica Kusano causes a noticeable leaf blister and curl in some

areas (Mix 1949). Microsphaera alni (Wallr.:Fr.) G. Wint. is abundant on alder leaves,

but it attacks late in the season just before leaves senesce and fall anyway (Reed

1913). Alder catkins are attacked and distorted by two fungi, Taphrina occidentalis

W.W. Ray and T. alni (Berk. and Boome) Gjaerum (Mix 1949).

In a review of the diseases of red alder, Harrington et al. (1994) suggest that several

stem canker pathogens may cause tree damage. The canker fungus Didymosphaeria

oregonensis Good. is particularly abundant on red alder, causing infection bands of

rough bark on living stems (Boyce 1961, Funk 1981). We have observed this disease

in abundance on red alder in young-growth stands in southeast Alaska. Physiological

damage may be negligible, as the fungus infects periderm, leaving the phloem relatively unaffected. Species of the Ascomycete Nectria have been reported on red alder;

N. galligena Bres. can form target-shaped cankers where phloem and cambial tissues

are killed (Ashcroft 1934, Brandt 1964). The decay fungus Hymenochaete agglutinans

Ellis is also listed as a canker-causing pathogen (Harrington et al. 1994).

Both pathogenic and saprophytic species of the root fungus Armillaria occur on red

alder. Rhizomorphs of Armillaria species are frequently observed under the bark of

recently killed red alder in southeast Alaska. Management activities can elevate levels

of Armillaria species. Centuries of coppice management in old, mixed-species forests

in England likely favors the development of several pathogenic Armillaria species on

European alder (Alnus glutinosa (L.) Gaertner (Rishbeth 1991). Although intentional

coppice is not practiced in Alaska, repeated short rotations of red alder or natural

events such as landslides that remove cover could increase Armillaria populations,

Pathogens
13

as was found in England. Phytophthora cambivora (Petri) Buisman attacks the root

collar of Alnus glutinosa in Germany (Hartmann 1995), but it is not known on red

alder. The possibility of planting red alder as a form of crop rotation to mitigate the

serious root pathogen Phellinus weirii (Murrill) Gilbertson of Douglas-fir in the Pacific

Northwest has spurred research on red alder and its influence on the rhizosphere

(Hansen 1975, Li et al. 1968, Neal et al. 1968, Nelson 1975, Wallis 1968). The root

disease form of Phellinus weirii does not occur in Alaska, however, and this form of

disease management is unnecessary. Sieber (1990) studied commensal fungi in living

red alder leaves and twigs to evaluate potential candidate mycoherbicides. Gnomonia

setacea (Pers.:Fr.) Ces. & DeNot, Gnomoniella tubaeformis Groves, and Phomopsis

sp. were the most common fungi isolated from leaves; Winterella (Ophiovalsa) suffusa

(Fr.:Fr.) Kuntze and Phomopsis sp. were the most common from twigs.

The most important disease agents of Sitka spruce and western hemlock have received more study than alder disease agents in Alaska forests. Tree pathogens can

be found at all stand ages, or stages of stand development (Oliver and Larson 1990),

but disease generally is ecologically and economically important in older stands

(Hennon 2000, Hennon and McClellan 1999). Tait et al. (1985) recorded a number

of insects and pathogens on young-growth western hemlock and Sitka spruce, but

all were considered minor in importance, and none initiated tree mortality. Porcupines

(Erethizon dorsatum) are the exception, as they kill dominant conifer trees in younggrowth stands (Eglitis and Hennon 1997).

Heart rot and hemlock dwarf mistletoe (Arceuthobium tsugense (Rosendhal) G.N.

Jones) (the latter is mainly restricted to western hemlock) are important diseases of

trees in stands 150 years or older. Dwarf mistletoe is largely absent or is eventually

eliminated through stand development in young-growth stands after clearcutting. Heart

rot can occur earlier in young-growth stands if tree wounding by animals or logging

activities is common (Hennon and DeMars 1997). Heart rot causes devastating timber

losses (Buckland et al. 1949; Farr et al. 1976; Kimmey 1956, 1964; Weir and Huber

1918) and is a common agent in small-scale disturbance (Hennon 1995) in old-growth

forests of southeast Alaska.

Hennon (1995) argues that the mode of tree death (i.e., uprooting, broken bole, standing) should be considered in studies of tree mortality because of the ecosystem processes it influences. It may be more important ecologically to determine how a tree dies

than what kills it. Several studies have classified the type of tree death in old-growth

forests (Hennon and McClellan 1999, Hocker 1990, Nowacki and Kramer 1998, Ott

1997). Quantifying types of tree death has not been attempted in young-growth forests

with red alder, except that Steinblum et al. (1984) and Keim et al. (2000) note more

uprooting of alder along streams than in forests away from streams in Oregon. Red

alder is more windfirm than Sitka spruce where they are both planted in England

(Gordon and Hall 1978). Windthrow may be uncommon in red alder because of the

intermingling of roots and branches, the absence of leaves during winter storm when

soils can be waterlogged, and the relatively deep-rooting habit of the species on welldrained soils (Harrington et al. 1994). Most red alder tree mortality is assumed to result

in dead standing trees unless landslides or bank erosion near streams occurs.

In young-growth stands, high tree density may lead to mortality for alder. Mortality related to tree density in red alder occurs when densities exceed 45 to 50 percent of that

required for crown closure (Hibbs and Carlton 1989, Puettmann et al. 1992). DeBell

and Giordano (1994) observed that most red alder trees die by suppression in winter

Mode of Death
14

or spring after other trees have leafed out; presumably energy supplies are finally exhausted. In pure stands in British Columbia, DeBell (1975) and Smith (1968) found that

alder density declines from about 120,000 stems per acre at age 1 to 2 years to 14,000

stems per acre at age 8 to 11 years. Puettmann (1994) lists a number of studies on

tree densities in red alder forests. Hibbs (1987) and Hibbs and Carlton (1989) describe

thinning to certain densities relative to crown closure to exercise some control over this

natural mortality.

Every important tree species in North America is invaded and decayed by one or more

species of heart-rot fungi (Gilbertson and Ryvarden 1986). More species of fungi, the

most important of which are Basidiomycetes, attack and deteriorate the wood of dead

trees than live trees. Wood decay by fungi occurs when enzymes degrade the major

constituents of wood—cellulose and lignin. There are two pathways for the many species of wood decay fungi (Boyce 1961, Hartig 1874): “brown rot fungi” degrade only

cellulose, and “white rot fungi” degrade both cellulose and lignin (Blanchette 1980,

Cowling 1961). Both result in greatly reduced strength of wood, but they generate

different woody debris structures and thus have different ecosystem functions (Hennon

2000). Brown rot leaves a residue, partially modified lignin, in which the strength and

much biomass of wood has been lost, but the volume is largely left intact (Forest Products Laboratory 1958). The residue from brown rot can contribute a large annual biomass to the soil and provide a zone of high microbial activity (Harvey et al. 1976).

White rot can consume all the wood, leaving trees or logs with hollows. Otjen and

Blanchette (1986) further classify types of white rot based on different micromorphological patterns of decay.

Decomposition of live and dead trees can be different. Decay of living trees in southeast Alaska is primarily by white rot fungi for western hemlock and by brown rot fungi

for Sitka spruce (Kimmey 1956). Fomitopsis pinicola (Sw.:Fr.) P. Karst., a fungus that

causes a brown cubical rot, appears to be the dominant decomposer of dead hemlock

and spruce trees in old-growth forests (Hennon 2000, Hennon and Loopstra 1991).

Edmonds (1999) reports that decay of young-growth conifers in western Washington

was mainly by white rot.

Various fungi attack the wood of alder (Hepting 1971). Farr et al. (1989) published

the most exhaustive list of fungi on the genus Alnus. Most of the several hundred taxa

listed are found on leaves, twigs, or wood of live or dead trees. Volk et al. (1994, see

page 35) report 33 fungal species on Alnus in a survey of wood-inhabiting fungi in

Alaska. Volk et al. (1994) also provide separate lists of fungi found on the wood of

Sitka spruce and western hemlock. Most of the fungi that inhabit alder are found on

dead trees. Harrington et al. (1994) list the most important heart rot fungi to red alder

in British Columbia: Heterobasidion annosum (Fr.:Fr.) Bref., Sistotrema brinkmannii

(Bres.) J. Eriksson, Pholiota adipsoa (Fr.: Fr.) P. Kumm., Trametes sp., and Meruliopsis

corium (Fr.:Fr.) Ginns. The most important wood-decay fungus of live red alder is probably Phellinus igniarius (L.:Fr.) Quel. (Driver 1978, Worthington 1957), which causes a

white rot. Interestingly, we do not see P. igniarius in red alder in southeast Alaska and

neither did Allen2 in British Columbia.

Decay

Fungi

2 Allen, E.A. 1992. Unpublished data. On file with: Pacific Forestry

Centre, 506 West Burnside Road, Victoria, BC V8Z 1M5.
15

Allen found that the number of living red alder trees with heart rot on Vancouver Island

increased with age (to 94 percent for trees >75 years), but decay volumes were not

correlated with tree age. For trees 60 to 80 years old, volume losses to decay were only

about 3.5 percent. Allen indicates that most of the decay in red alder is associated with

wounds such as scars, forks, and broken tops. The upward-growing, sprawling habit of

red alder branches and its relatively weak wood probably contribute to broken branches

and tops. Decay is limited, however, because red alder is able to compartmentalize the

decay to tissues near the wound.

The white rot fungus Hypoxylon fuscum (Pers.:Fr.) Fr. (an Ascomycete, not a Basidiomycete) is exceedingly common as a decomposer of dead alder wood (Rogers 1968).

Miller (1961) observes that the “fungus is on almost every limb of Alnus that has recently died anywhere in the North Temperate Zone.” Rogers (1968) presents some

evidence, based on the apparent speed of colonization after tissue death, that this

fungus may live as a commensal in live alder. It thus would have a competitive advantage over other fungi to quickly and more fully colonize recently killed alder tissue.

Wood from different tree species differs in its decay resistance. This variation is due

mainly to the deposition of fungitoxic substances in cell walls during their transformation from sapwood to heartwood (Eslyn and Highley 1976). Hardwood trees contain

more sugars, starches, proteins, and nutrients in the inner bark and the sapwood than

do conifers (Harmon et al. 1986). The decay resistance (sometimes called durability)

of Alaska tree species has been tested along with many other tree species to determine how long different woods can be used when in contact with water and or soil. The

heartwood from red alder, Sitka spruce, and western hemlock are all rated in the same

broad group as “slightly or nonresistant” (Forest Products Laboratory 1987), indicating

that they decay rapidly. The sapwood of red alder, Sitka spruce, and western hemlock

were tested in vitro for resistance against white rot, brown rot, and soft rot organism

(Eslyn and Highley 1976). Red alder was susceptible to brown rot and soft rot agents

and very susceptible to white rot; western hemlock and Sitka spruce had relatively

good resistance to decay except for brown rot fungi, which caused significant decay. In

another test, the sapwood of different species was placed outdoors in Mississippi and

rated for decay: red alder was classed as “nonresistant,” decaying in less than 7 years;

Sitka spruce and western hemlock were “moderately resistant,” lasting 8 to 13 years

(Eslyn et al. 1985).

Decay of wood in streams is receiving more recent attention from researchers. Saturation with water has been shown to decrease oxygen concentrations in wood to levels

low enough to prevent penetration by fungal hyphae (Savory 1954). Bacteria are responsible for most decomposition of saturated wood (Crawford and Sutherland 1979)

and operate only on the surface where oxygen concentrations are adequate. Bilby et al.

(1999) report lower decay rates in submerged logs of all trees studied (including red

alder and western hemlock) than in logs of similar size and species in the terrestrial

environment. Cederholm et al. (1997) found that partially submerged red alder logs had

significant deterioration in only 3 years in Washington. Micro-organism and insect larvae were responsible for the rapid decomposition of Alnus glutinosa twigs in a river in

France (Chergui and Pattee 1991). The authors measured a 60-percent weight reduction in 2.5 years and expected full decomposition of the twigs in 4 to 5 years. Alder

trees mechanically pulled over into Oregon streams had more wood decay and broke

Decay Resistance

Decay of Wood in

Streams
16

more frequently than did Douglas-fir (Keim et al. 2000). Red alder with attached roots

was effective in trapping coarse woody debris, but after 3 years, the alders were losing

structural integrity owing to advancing decay and breakage. Anderson et al. (1978)

report on several aquatic invertebrates that contribute to the degradation of red alder

and western hemlock wood in Oregon streams. They found that the activity of most

invertebrates does not involve deep boring or penetration into the wood, and the wood

supported a smaller biomass of invertebrates than did leaf litter.

Riparian zones play a major role in regulating nutrient and energy flow in low-order

streams (Cummins et al. 1989, Wallace et al. 1997, Webster and Benfield 1986,

Webster et al. 1992). Leaf litter and wood from riparian vegetation enters the lotic

system and affects microbial and benthic macroinvertebrate communities that consume and process this organic material (Anderson et al. 1978, Bisson and Bilby 1998,

Cummins et al. 1989, Hax and Golladay 1993, Sedell et al. 1975, Vannote et al. 1980).

This allochthonous (plant and invertebrate) material typically constitutes the energy

base in low-order forested streams (Fisher and Likens 1973, Minshall 1967, Triska

and Sedell 1976).

Riparian and stream productivity are essential for sustaining salmonid fisheries in

southeast Alaska. These fisheries are critical to the economy and quality of life of

peoples of this region, as well as other regions of the North Pacific Rim. To maintain

a natural flow of sediment, woody debris, and water from headwater channels to the

receiving fish channels, the Tongass National Forest has implemented a policy of

providing riparian buffers to the class III to IV channels that lack anadromous fish

(USDA FS 1997).

Riparian buffer effectiveness in maintaining a natural flow of sediment, woody debris,

and water appears to be site specific. A buffer region that includes a natural source of

large woody debris (whole trees) and sediment is more likely to be effective than a

buffer of the same size that does not encompass a natural source of both large woody

debris and sediment. Mechanisms that introduce woody debris and sediment include

landslides, blowdown, bank erosion, and flooding (fig. 4).

Although it is understood that stream ecosystems are closely linked to and dependent

on associated riparian zones for organic matter and nutrient inputs, little attention has

focused on the relation between riparian vegetation and terrestrial invertebrate communities. Also not well understood is the influence of riparian vegetation on the way terrestrial invertebrates enter streams (e.g., as prey for fishes, or as allochthonous material

contributing nutrients and organic matter). Some riparian tree species contribute more

terrestrial invertebrate mass to streams than others (Mason and Macdonald 1982).

Southwood (1961) reported that deciduous trees generally have more associated

invertebrate species than do conifers. Plant community successional stage also influences invertebrate abundance and species assemblage (Mispagel and Rose 1978).

Schowalter et al. (1981) reported lower invertebrate mass 1 year after clearcutting but

higher mass 2 years after clearcutting, relative to an uncut forest in the Pacific Northwest. Mundie (1974) indicated that aerial insects were more abundant in a deforested

than forested setting in coastal British Columbia, Canada, and that terrestrial insects

may be seasonally important salmonid food.

Role of Riparian Forests

in Stream Processes

Effect of Riparian

Trees on Terrestrial

Invertebrates in

Stream Food Webs
17

7 7 

5 m

1 m

50 m

6 6 

1 1 

Stream

5 5 

4 4 

3 3 

2 2 

2.5 

m

10 m

4 m 4 m

1 m

Streamflow

Upper 300-m reach

Lower 300-m reach

Viewing tube

measurements taken

Stream pipe reference

Large fixed-area plot

Trees with d.b.h. >15 cm

Medium fixed-area plot

Trees with d.b.h. >3 cm

Shrub cover plot

Small fixed-area plot

Trees with d.b.h. >0 cm

Legend

10 m

1 

Figure 4—Riparian sampling layout. Lower 300-m reach (not shown) was sampled by using the same design as the upper 300-m reach.
18

Riparian management activities that alter plant communities may change food

resources within and surrounding a stream. Riparian plant communities govern

allochthonous inputs (Cummins et al. 1989, Webster and Benfield 1986) and

autochthonous production (instream primary production) (Cummins et al. 1989,

Fisher and Likens 1973). In southeast Alaska, clearcutting in riparian zones has

given rise to even-aged young-growth riparian communities at various successional

stages. Alder often regenerates along stream margins along with variable amounts of

Sitka spruce, western hemlock, western redcedar, Alaska yellow cedar Chamaecyparis

nootkatensis (D. Don) Spach), black cottonwood, and willow. Little is known about the

influence of these developing riparian communities on upland stream productivity and

downstream fishes. Alder could have a variety of impacts on streams, such as changes

in solar penetration and allochthonous inputs. These effects could lead to changes in

autochthonous production (Cummins et al. 1989; Fisher and Likens 1973; Hetrick et al.

1998a, 1998b) and quality and quantity of plant litter inputs (Cummins et al. 1989,

Webster and Benfield 1986) and terrestrial invertebrate inputs (Wipfli 1997). These

factors could all impact the amount of food in downstream fish-bearing reaches.

Changes in forest canopy may have both positive and negative impacts on aquatic

communities. On the positive side, riparian forests with some alder have been shown

to produce more prey for salmon than have conifer forests (Wipfli 1997). However, this

benefit may be outweighed by a decrease in large woody debris input (Andrus et al.

1988, Grette 1985, McDade et al. 1990), decrease in wood longevity (Anderson et al.

1978, Benda and Cundy 1990, Keim et al. 2000), and increases in sediment loading

(Megahan 1982); factors resulting in loss of fish habitat in systems dominated by alder

(Bilby and Ward 1991, Hicks 1989, Ralph et al. 1994). An understanding of changes

associated with forest alteration may be obtained by estimating processes associated

with natural woody debris loading (Potts and Anderson 1990) (or defined as woody

debris budgets (Benda and Sias 1998)). In addition, an understanding of the locations

where alder occurs both naturally and as a result of management may give insight into

the morphological, functional, and community-level changes that occur in streams as

succession from alder to conifer occurs. With this knowledge, it may be possible to link

upslope processes and associated aquatic habitats (i.e., pools) (Andrus et al. 1988,

Bilby and Bisson 1998).

Of particular interest to land managers are questions concerning the upper limit of fish

habitat, conditions that regulate the use of the upper sections of watersheds, and the

impacts of upslope forest management. The intensity of interactions between upslope

processes and fish populations is likely related to proximity. Salmonid populations occupying habitats at or near the upper limit of habitat (“transition zones”) (fig. 4) are more

likely to be affected by upslope forest processes. Transition zones generally occur in

reaches with gradients between 5 and 20 percent, tend to be sparsely populated, and

may be occupied only seasonally in southeast Alaska (Bryant 1983). Fish in these

zones are closely linked to—and are likely part of—downstream populations.

Bird abundance has been found to be low in young-growth stands of dense conifer

(Schwab 1979 as cited by Easton and Martin 1998). A positive association between bird

abundance and the density of live deciduous trees has been found in young-growth

forests (reviewed by Huff and Raley 1991, McComb 1994, Ruggiero et al. 1991) and

regenerating clearcuts (Morrison 1981, Santillo et al. 1989). The role of deciduous

Impacts of Changes in

Riparian Forest Canopy

Bird Abundance

Associated with

Red Alder
19

trees, however, is not understood, and little is known about the interactions between

alder and birds in the Pacific Northwest (reviewed by McComb 1994), particularly in

southeast Alaska.

The importance of alder to southeast Alaska birds has not been well studied (Gende

and Willson 2001), but it is generally thought to be of value (Sidle 1985). Alder may

provide advantages in term of food and nesting sites. Some aerial insectivores (e.g.,

flycatchers) have shown direct relations with alder, with increased numbers of individuals in alder stands (Gilbert and Allwine 1991). Because insects from foliage can be a

primary food source for insectivorous birds (Airola 1979, Holmes 1990) and insect

biomass is greater in deciduous vegetation (Stiles 1980, Willson and Comet 1996),

it follows that birds in southeast Alaska may forage for insects in alder trees.

Alder within second-growth stands may improve the quality of nesting habitat for some

bird species. Dense conifer young-growth stands typically lack a well-developed understory. Consequently, the number of nest sites for birds may be limited, and nest concealment, which is associated with successful nesting for some open-cup nesting

species (reviewed by Kelly 1993, Martin 1993; but see D’Eon 1997), may be lower in

conifer stands. Furthermore, some nest predators (e.g., red squirrels (Tamiasciurus

hudsonicus)) have been found to be more abundant in coniferous forests.3 Birds nesting in mixed alder-conifer stands may have more nest sites to choose from, safer nest

sites to choose from, and a reduced chance of encountering some types of nest predators.

Coniferous trees in southeast Alaska typically have low numbers of leaf-chewing

arthropods (Mask 1992). Red alder and, to a lesser extent, Sitka alder (Alnus sinuata

(Regel) Rydb.) probably provide a critical source of protein throughout the summer.

Little is known about the life cycle of insects on alder foliage in southeast Alaska.

The wood of red alder is pale reddish brown, moderately dense and uniformly textured,

with an attractive grain well suited for cabinet work. It is used in the production of wood

products such as furniture, cabinets, and pallets (Resch 1980), composite products

such as plywood and flakeboard (Bollen et al. 1967), and fiber-based products such as

tissues and writing paper. Alder is a common fuelwood both for home use and for commercial drying (Briggs et al. 1978, Resch 1980). In Alaska, red alder is used for smoking fish and carving but otherwise is not used commercially.

Although the commercial value of red alder has traditionally been lower than that of

associated conifers, red alder is the major hardwood tree species in the Pacific Northwest, and its value has increased substantially in recent years. In 1991, the harvest of

red alder in the Pacific Northwest Region was over 480 million board feet (Western

Hardwood Association 1992), and Gedney (1990) reported a supply of over 9 billion

board feet of red alder sawtimber in western Oregon. Alder exports to Asia and Europe

grew to over 65 million board feet in 1992, and in 1991 alder accounted for 10 percent

of volume of all U.S. hardwood lumber exports to Europe and Asia (Tarrant et al. 1994).

The scarcity of young red alder is now a concern to a wood products industry that continues to grow (Hibbs et al. 1994).

Commercial Value of

Red Alder

3 De Santo, T.L. 2000. Unpublished data. On file with: Forestry

Sciences Laboratory, 2770 Sherwood Lane, Suite 2A, Juneau,

AK 99801.
20

Traditionally, many forest managers have tried to eliminate alder from forest stands.

The value and use of alder has increased (Plank and Willits 1994, Plank et al. 1990,

Waggener 1978), and recent research has shown that alder log values compare

favorably with those of Douglas-fir on a net scale basis (Plank et al. 1990). Economic

comparisons between alder and Douglas-fir show alder to be a viable alternative to

softwoods (Tarrant et al. 1983). The market performance of red alder is strong (Western Hardwood Association 1992), and alder prices have increased steadily since 1980.

Interest in managing red alder is high, and research on red alder over the past 50 years

is providing information on its potential uses. Red alder research is being strengthened

through the work of the Hardwood Research Cooperative and with previous and current

research at the Forest Service Pacific Northwest Research Station (DeBell et al. 1978;

Harrington 1984, 1986, 1990; Harrington and Curtis 1986; Harrington and DeBell 1984;

Trappe et al. 1968; Worthington et al. 1960).

Because total area of upland forests and associated headwaters comprises 70 to 80

percent of watersheds, they are potentially important sources of sediment, water, nutrients, and organic matter for downstream habitats. Transport of organic and inorganic

material from upland forest and headwater streams may affect physical and biological

processes in downstream reaches. For instance, sediment produced in headwater

systems moves through channel networks and alters channel morphology (Benda and

Dunne 1997). Sediment transported from headwater tributaries creates various channel

environments and modifies patterns of riparian structure and hyporheic exchange in

downstream reaches (Gregory et al. 1991, Swanson et al. 1998). Movement of detrital

material and invertebrates from headwaters to downstream food webs may alter productivity, population density, and community structure of stream biota in downstream

reaches (Wipfli and Gregovich 2002). The occurrence of red alder in these uplands,

including in riparian zones of headwater streams, may have a substantial influence on

the physical and biological processes operating in these systems. Because red alder

decays faster than conifers and is a more desirable food source for invertebrates, it

likely affects the sediment, detritus, and invertebrates exported from headwaters to

downstream habitats.

There are three main themes of this research: (1) influence of alder on specific ecosystem components, i.e., on vegetation, birds, invertebrates, wood, and fishes (fig. 1);

(2) flow of wood from riparian forests to adjacent streams, and from upland to lowland

habitats (fig. 2); and (3) influence of alder on ecosystem links and processes (fig. 3).

In southeast Alaska, red alder may increase the abundance and diversity of understory

plants, in turn influencing aquatic and terrestrial invertebrate communities and the bird

and fish communities that feed on these prey (figs. 2 and 3). Alder also may decay

more quickly than conifers, affecting the residence time of wood in terrestrial and

aquatic habitats, and its movement and distribution in streams (fig. 2).

Study sites will represent a continuum of alder-conifer mixtures. To reduce the range

of site variability, we will select study areas in the Maybeso and Harris watersheds on

Prince of Wales Island, Alaska, using composition of tree species as a primary site

selection criterion. These watersheds have large areas of relatively uniform stand age

and productivity, and a wide range of alder-conifer mixtures. Sites will be selected and

sampled during the 2000 and 2001 field seasons. The objectives for each component

of the study require specific selection criteria, but we will seek to maximize sites that

can be used for all resources, including timber, wildlife, and fish. See table 1 for a

complete listing of sites.

Importance of Upland

Forests and Associated

Red Alder for Aquatic

Habitats

Objectives and

Hypotheses

Perceptions of Red Alder
21

Table 1—Common study sites, site description, drainage, and components studied at each site in Maybeso and Harris watershedWales Island, Alaska

Preliminary Components studied

assessment

of red alder Stream Woody Riparian Upland Aquatic 

Study site abundance Drainage Geomorphology nitrogen debris vegetation vegetation ecology Fish Bird

Alluvial Fan High Maybeso X X X X Big Spruce Medium Maybeso X X X X X X X X Broken Bridge East High Maybeso X X X X XBroken Bridge West Medium Maybeso X X X X XBrushy High Maybeso X X X X X XCedar 1 Low Maybeso X X X X X X X XCedar 2 Low Maybeso X X X X X X XCotton Medium Maybeso X X X X X XCotton Tributary High Maybeso X X XCreature Creek Low Maybeso X X X X X XGomi Medium Maybeso X X X X X XLost Bob Low Maybeso X X X XLower Broken Bridge High Maybeso X XLower Good Example Low Maybeso X X X XLower Morning Medium Maybeso X X XUpper Good Example Low Maybeso X X X X X X X Upper Morning Low Maybeso X X X X X XGroup Photo Medium Harris X X X Mile 22 High Harris X X X X X X X X 
22

To assess the effect of alder in young-growth forests, and to evaluate the potential

tradeoffs in alder-dominated and conifer-dominated forests, we will investigate interactions in six areas: wood production, vegetation, and tree disease; terrestrial invertebrate

ecology; wildlife ecology; aquatic and riparian ecology; fish ecology; and geomorphic

processes. Objectives and hypotheses are summarized by discipline on the following

pages. See appendices for detailed description of methods.

Stand density, vegetation, and tree mortality (app. 1)—

Objective 1: Compare total stand density across a range of alder-conifer mixtures.

Hypothesis: Mixed alder-conifer young-growth stands have lower total stand basal

area and tree stocking than do pure conifer stands, and both stand density and tree

stocking decline with increasing proportion of alder in stand.

Rationale: Stand growth and wood production are maintained much longer for conifers

than for alders, and at the stand age of this study (40 to 50 years), it is expected that

spruce and hemlock wood production will be greater than for red alder. In nitrogenlimited sites in the Pacific Northwest (Miller and Murray 1978, Tarrant 1961), however,

mixed alder-conifer stands accumulated more wood than did pure conifer stands. In

contrast, more productive sites generally show less wood production, biomass, and net

primary productivity (Binkley et al. 1992) in red alder compared with pure conifer

stands. Little is known about the productivity of alder in Alaska, but if highly productive

conifer sites in southeast Alaska follow patterns in the Pacific Northwest, then growth

and wood production of mixed alder-conifer stands should be less than in pure conifer

stands.

Objective 2: Compare understory plant diversity and biomass in stands across a range

of alder-conifer mixtures.

Hypothesis: Mixed alder-conifer young-growth stands have greater understory plant

diversity and biomass than do pure conifer young-growth stands.

Rationale: Dense conifer regeneration and canopy closure in stands 25 to 150 years

old result in greatly reduced understory plant diversity and abundance (Alaback 1982,

1984b). Red alder stands, however, appear to have significantly greater understory

biomass (Hanley and Hoel 1996) and greater canopy coverage (Deal 1997) than do

conifer stands. It is expected that increasing proportions of alder in these mixed alderconifer stands will provide greater understory plant diversity and abundance.

Objective 3: Compare stand growth and regeneration in stands across a range of alder-conifer mixtures.

Hypothesis: Pure conifer young-growth stands have more tree regeneration than do

mixed alder-conifer stands, and regeneration decreases with increasing proportions of

alder in stands. Tree diameter growth is greater for overstory conifers (dominant and

codominant trees) in mixed alder-conifer young-growth stands than in pure conifer

stands, and tree diameter growth increases with increasing proportions of alder in

stand.

Rationale: Stands 40 to 50 years old are in the stem-exclusion stage of stand development in southeast Alaska (Alaback 1982, Deal et al. 1991), and new tree regeneration

will be limited. Red alder generally colonizes heavily disturbed sites with exposed mineral soil. Conifers regenerate on logs and organic material (Harmon and Franklin 1989,

Yount 1997), and the lack of rooting medium in alder stands is expected to reduce tree
23

regeneration. Red alder stands are less dense and provide more growing space for

trees than do unthinned natural conifer stands. It is expected that dominant conifers

in mixed alder-conifer stands will show increases in diameter growth compared with

pure conifer stands.

Objective 4: Compare frequency and type of tree mortality (mode of tree death) in

stands across a range of alder-conifer mixtures.

Hypothesis: The greatest frequency of tree mortality occurs in trees of the smallest

size class. More trees die standing than by bole breakage or uprooting. The amount

of white and brown rot wood decay produced in these stands differs by species of

dead tree.

Rationale: Most woody debris that reaches the forest floor or streams is the result of

tree mortality. This pattern of tree mortality differs from that of old-growth forests

wherein some dominant trees die and produce various structures (e.g., hollow logs

from broken boles or attached root systems from uprooted trees). The pathway of

wood decomposition (brown rot or white rot) also may influence physical and biological

functions.

Objective 5: Compare total stand density, tree diameter distribution, and size-specific

tree mortality across a range of riparian alder-conifer mixtures.

Hypothesis: Young-growth riparian stands with a greater alder component will have

lower total stand density and tree mortality than will stands with less alder.

Rationale: A greater alder composition in young-growth riparian stands will result in

widely spaced stands and less competition for light. This decreased density will result

in lower mortality among trees, and most dead trees will be small in diameter. Conifers

will have larger diameter than in upland stands owing to light-enhanced diameter

growth, and will create a good source of large woody debris for habitat improvement

of adjacent streams.

Objective 6: Compare the spatial distribution and stocking of alder and conifer trees

between upland and riparian stands across a range of alder-conifer mixtures.

Hypothesis: Riparian stands will have less alder, and the alder will be aggregated

close to stream margins, whereas alder in upland stands will be more densely stocked

and randomly distributed.

Rationale: Frequent surface soil disturbance in riparian areas, owing to hydrological

processes such as floods, tends to favor aggregations of alder. Surface soil disturbance in upland areas is expected to be more randomly distributed, associated with

events such as landslides or forest blowdown.

Objective 7: Measure size and condition of woody debris entering the forest floor

and streams, including type of decay, species of wood, and presence or absence of

attached root system.

Hypothesis: The amount of white and brown rot wood decay produced in these stands

differs by the abundance and species of dead trees.

Rationale: Wood decay type (i.e., white rot and brown rot) and extent of decay in

dead trees will influence abundance of invertebrates in dead stems, wildlife use of

dead stems, persistence of dead standing trees, woody debris on the forest floor,

and mobility of woody debris in streams.
24

Terrestrial invertebrate ecology (app. 2)—

Objective 8: Compare terrestrial invertebrate abundance and species richness in foliage, branches, and stems of woody vegetation as well as in leaf litter in stands across

a range of alder-conifer mixtures.

Hypothesis: Terrestrial invertebrate abundance and species richness are greater in

mixed alder-conifer young-growth stands than in conifer young-growth stands.

Rationale: Invertebrates are larger and more abundant in mixed alder-conifer younggrowth stands than in conifer young-growth stands because red alder increases the

food value of neighboring plants (Mason et al. 1992).

Wildlife ecology (app. 3)—

Objective 9: Compare bird density and species richness in mixed alder-conifer and

conifer stands.

Hypothesis: Bird density and species richness are greater in mixed alder-conifer

young-growth stands than in conifer young-growth stands.

Rationale: Young-growth stands dominated by conifers have little understory vegetation. Sparse understory vegetation probably results in reduced nesting and foraging

substrate for understory bird species. Alder in young-growth stands may provide resources for foraging and nesting that are unavailable in conifer stands.

Objective 10: Conduct behavioral observations of foraging birds in mixed alder-conifer

young-growth stands.

Hypothesis: Insectivorous birds forage on alder more often than on conifer in mixed

alder-conifer young-growth stands.

Rationale: Nearly all birds are insectivorous during some stage of their life cycle, and

all shrub and forest birds in southeast Alaska feed insects to their young. Insect populations are generally greater in deciduous vegetation. If this holds true in southeast

Alaska, and if we find that avian species diversity and abundance are higher in younggrowth stands containing alder, then birds may be selecting these areas to take advantage of the insect resources these stands provide. Understory vegetation also provides

other types of food resources (primarily berries) for some understory birds (e.g.,

thrushes (Turdidae)) in southeast Alaska.

Objective 11: Compare nesting success in mixed alder-conifer young-growth stands

and in conifer-dominated young-growth stands.

Hypothesis: Nest predation is lower in mixed alder-conifer young-growth stands than

in conifer-dominated young-growth stands.

Rationale: Young-growth forests without understory vegetation provide little nesting

substrate or nest cover. Nest predation is the leading cause of nest failure in southeast

Alaska.

Objective 12: Compare summer food value for deer in young-growth stands differing

in their proportion of alder component.

Hypothesis: Understory plant species richness, canopy coverage, biomass, and food

value for deer increase with increasing proportion of alder in young-growth stands.
25

Rationale: Dense conifer regeneration and canopy closure in conifer stands 25 to

150 years old results in greatly reduced understory and plant diversity and abundance

(Alaback 1982, 1984b). Understory of red alder stands appears to have significantly

greater biomass (Hanley and Hoel 1996) and greater canopy coverage (Deal 1997).

Food value of understory for deer depends on plant species composition as well as

biomass, and the greater abundance of forbs under alder than conifer is expected to

benefit food value for deer (Hanley and Barnard 1998).

Aquatic and riparian ecology (app. 4)—

Objective 13: Compare the amount of export of invertebrates and detritus from

fishless headwater streams to downstream fish-bearing habitats across a gradient

of riparian alder abundance.

Hypothesis: Headwater streams with more alder in their riparian zones will deliver

more invertebrates (aquatic and terrestrial) and organic detritus via fluvial transport

to downstream habitats than will streams with less alder.

Rationale: Alder is more nutritious than conifers and therefore will support more riparian and aquatic invertebrates, which in turn will get carried downstream to fish habitats

via fluvial transport.

Objective 14: Compare invertebrate abundance in and along headwater streams,

including on woody debris at various stages of decay, in stands with different alderconifer mixtures.

Hypothesis: Invertebrate abundance and species richness are greater in younggrowth riparian stands and associated headwater streams that contain more alder,

and are greater on decaying alder wood than conifer wood.

Rationale: Alder is more nutritious than conifers and therefore will support greater

densities and more species of invertebrates.

Objective 15: Compare nutrient concentrations among streams that differ in the

amount of upstream red alder and in discharge and disturbance history.

Hypothesis: The concentration of nitrate in stream water is directly proportional to the

amount of alder in the catchment and inversely proportional to the discharge rate.

Rationale: Soil nitrogen concentrations in alder-dominated soils are higher than in soils

dominated by conifer species. Alder-derived nitrate moves from upland soils to streams

via groundwater flows and hyporheic exchange. Where past logging practices have

increased the amount of alder in small catchments, stream nutrient concentrations and

productivity may vary with alder coverage.

Fish ecology (app. 5)—

Objective 16: Compare salmonid densities in stream reaches below and within coniferdominated and alder-dominated upslope riparian forests.

Hypothesis: Salmonid densities will be higher in stream reaches below and in coniferdominated upslope riparian forests than in reaches with alder-dominated upslope riparian forests.
26

Rationale: High-gradient first- and second-order stream channels in heavily managed

sites are prone to landslides and debris flows. As a result of such disturbances, these

sites have been colonized and vegetated with alder. Debris flows can affect fish habitat

by filling in pools, straightening channels, removing large woody debris, and adding fine

sediment to spawning gravel. There will be a higher quantity and quality of habitat to

support a higher diversity and density of salmonids in transition reaches not affected

by landslides. Therefore, we expect salmonid densities to be higher in upper stream

reaches (transition zone) not affected by debris flows (i.e., upslope conifer forest).

Objective 17: Assess seasonal variation and relative species abundance of salmonids

in upper high-gradient reaches (transitions zones) (fig. 4).

Hypothesis a: Salmonid densities will be higher in upper high-gradient reaches

(transition zones) during spring and fall than during summer.

Rationale a: Upper sections of small tributaries are more likely to be used for spawning by adult fish such as coho salmon (Oncorhynchus kisutch) and Dolly Varden

char (Salvelinus malma) during fall, and by cutthroat trout (Oncorhynchus clarki clarki)

during spring. Upper sections have smaller drainage area and are more likely to become intermittent during low flows that occur in summer. Upper reaches may be

affected by upslope disturbances such as landslides that can scour channels, removing large wood and pool habitat, and can deposit substantial amounts of sediment.

Hypothesis b: The relative abundance of salmonid species will differ between downstream reaches and transition reaches. Downstream abundance will be coho salmon >

Dolly Varden char > cutthroat trout. Transition reach abundance will be Dolly Varden

char > cutthroat trout > coho salmon.

Rationale b: Steeper gradient results in decreased habitat for juvenile coho salmon.

Upstream effects will result in a reduction of large wood and pools in the upper reaches

of fish-bearing streams. Downstream deposition of large wood by landslides may result

in greater habitat complexity, which is generally favored by juvenile coho salmon.

Objective 18: Assess the relation between the abundance of alder in a riparian forest

and stream inputs and processes.

Hypothesis: Large woody debris volume, riffle substrate size, and number of pools

will decrease as the ratio of alder to conifer increases.

Rationale: The presence of alder in upslope riparian reaches indicates greater and

more recent disturbance activity than is typical in conifer-dominated riparian forests.

Disturbance activities in these reaches tend to be landslides and debris flows that

remove or bury large woody debris, add fine sediment, and fill in pools.

Geomorphic processes (app. 6)—

Objective 19: Describe geomorphic processes and land use history at study sites

selected by Wood Compatibility Initiative colleagues.

Rationale: Dominant disturbance processes are related to landform type and type of

forest cover. This analysis will provide (1) an increased understanding of the role of

headwater disturbance processes in the formation of aquatic habitats in a watershed

context and (2) a means to assess the relevance of habitat changes (both positive and
27

negative) that occur as a result of alteration in land use (conversion from old-growth to

second-growth forest with stands of alder and conifer). No hypothesis will be tested for

this objective.

Objective 20: Assess processes that deliver sediment and woody debris to streams

from alder-dominated and conifer-dominated forests, and compare delivery rates

(fig. 4).

Hypothesis: Sediment and woody debris input to headwater channels is higher from

alder stands than from conifer stands.

Rationale: Woody debris and sediment are delivered to stream systems through the

processes of blowdown, bank erosion, forest mortality, and landsliding. On steep terrain, landsliding is the dominant source of woody debris and sediment (Gomi 2002).

Given that the presence of alder often indicates a history of landsliding, we expect

more sediment and woody debris delivery in these regions than in conifer-dominated

regions.

Objective 21: Estimate differences in sediment storage capacity in headwater streams

within conifer-dominated and alder-dominated forests.

Hypothesis: Sediment storage capacity in upland streams (above fish habitat) is

greater in forests dominated by conifers than in alder-dominated forests.

Rationale: The presence of alder in steep ecosystems often indicates that landslides

have occurred. Landslides typically erode woody debris and sediment from steep regions above fish habitat; thus we expect more woody debris in reaches where there

has been no history of landsliding.

An essential element of this project’s study plan is documenting the types and amount

of measurement error. Measurement error can occur in many ways but typically has at

least two fundamental components. The first is “observer bias” or error that occurs

among individuals, i.e., when more than one individual obtains values for the same

variable, such as diameter at breast height (d.b.h.). This occurs because no two individuals will consistently obtain the same values when recording observations. This is

especially true for categorical data that entail subjective assessments (e.g., vegetative

cover estimated to the nearest 10 percent), but may frequently occur with measurements of continuous variables (e.g., d.b.h.).

A second source of measurement error is inconsistency in measuring the same variables at multiple sites. At least three factors contribute to this inconsistency. The first is

natural variation that precludes applying techniques in a similar fashion at all locations.

For example, d.b.h. values may be influenced by tree taper, ground slope, microrelief,

or many other factors. The second component is related to the resolution at which observations are recorded. Generally, the finer the resolution, the greater the potential for

measurement error. For example, it is easier to estimate vegetative cover consistently

to the nearest 25 percent than to the nearest 5 percent. Estimating the weight of small

birds or mammals to the nearest 0.1 g on a 1-g scale will almost certainly contribute

substantial measurement error to the data, whereas accurate estimates to the nearest

0.5 g may be achievable. The third component is the human element: humans cannot

repeat a process without some variation in the way the technique is applied.

Quality AssuranceQuality Control

Procedures

Documenting

Measurement Error
28

We will document measurement error by repeat sampling or simultaneous sampling of

a given variable. For repeat sampling, the time interval between successive measurements will be relatively short to reduce the influence of natural variation over time on

our estimates of measurement error. For both types of documentation, a randomly

selected subset of previously sampled units representing 5 to 10 percent of the total

sample units will either be revisited (e.g., vegetation plots) or simultaneously sampled

by two individuals (e.g., bird surveys). Measurement error will be recorded as the difference between the two measurements of the same variable. In circumstances where

there exists potential observer bias, different recorders will visit the same site.

Standard protocols for data management will ensure long-term integrity and availability

of project data. Field observations will be recorded primarily on handheld electronic

data recorders with additional information written in field notebooks. The data will be

downloaded to floppy disk daily and backed up to minimize the chance of data loss.

Upon returning to the office, raw data files will be inspected by the field crew leader,

formatted in spreadsheets, and added to the master database by the database manager. A spreadsheet on the database computer tracks the data through these steps.

The database is written to a compact disc (CD) monthly, and the raw and formatted

data files are archived on a data library CD that is backed up biannually. Backup copies

of each CD and handwritten notes are stored in a fireproof cabinet at the Juneau Forestry Sciences Laboratory and at an offsite location. Data will be proofread for errors

independently of the data collection and entry process.

Below is the expected timeline for completing the project:

1999 2000 2001 2002 2003 2004

Study plan X X

Fieldwork X X

Manuscript writing X X

Publishing X X

Data Management and

Archiving

English Equivalents When you know: Multiply by: To find:

Meters (m) 3.28 Feet

Centimeters (cm) 0.39 Inches

Micrometers (μm) 3.9 × 10-5 Inches

Millimeters (mm) 0.039 Inches

Square meters (m2) 1.20 Square yards

Cubic meters (m3) 35.3 Cubic feet

Kilograms (kg) 2.21 Pounds

Grams (g) 0.035 Ounces

Hectares (ha) 2.47 Acres

Kilograms per hectare (kg/ha) 0.89 Pounds per acre

Metric tons per hectare (t/ha) 890 Pounds per acre

Cubic meters per hectare (m3/ha) 0.011 Cubic feet per acre

Milliliters (mL) 0.034 Fluid ounces

Celsius (°C) 1.8 and add 32 Fahrenheit
29

We thank Rebecca Braun for her diligence and hard work with editing and making this

study plan a much more cohesive and readable product. We thank Alison Cooney for

field, laboratory, editorial, and technical assistance. We acknowledge Richard Haynes,

Dave Hibbs, Winnie Kessler, Charley Peterson, Deanna Stouder, and Rick Woodsmith

for their comments on earlier versions of this document. We thank Richard Haynes and

Deanna Stouder for their encouragement and constructive suggestions throughout the

writing process. Thanks to Christian LeSage and Ryan Kimbirauskas for contributing to

the stream and riparian research components as part of their graduate degree work.

Finally, we acknowledge funding of this study through the Wood Compatibility Initiative,

Pacific Northwest Research Station, Portland, OR.

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47

Appendix 1: Methods for Sampling Stand Density, Vegetation, and Tree Mortality

To evaluate potential tradeoffs in stand density and understory plant diversity and

biomass between alder-dominated and conifer-dominated forests, we will sample

across a range of alder-conifer mixtures. The proportion of alder basal area is the

major criterion for stand selection, with stands representing a continuum from pure

conifer to predominantly alder. To reduce variability among stands we will limit sites to

stands 40 to 50 years old in the Maybeso and Harris watersheds on Prince of Wales

Island. These watersheds contain large areas that were logged during the 1950s and

have stands of relatively similar age and site productivity, with a wide range of alderconifer composition.

Study areas were selected from a pool of potential sites identified by using aerial photographs and Forest Service district files. Field visits were used to identify stands and

select study areas based on tree species composition. To control or reduce variability

among sites, the following selection criteria were used: (1) stands that were logged and

naturally regenerated, (2) stands that had no intermediate management activities such

as tree thinning or alder girdling, (3) stand size of generally 5 to 10 ha with a minimum

size of 2 ha, (4) stand elevation of less than 150 m, (5) stand slopes of 10 to 40 percent, and (6) no unusual site conditions within stands such as gravel borrow pits, recent

logging roads, or beaver ponds. Nine stands were selected for use.

A total of 13 headwater streams in the Maybeso and Harris watersheds were chosen as

study sites; 6 with 600-m reaches and 7 with 300-m reaches. “Reach location” refers to

the uppermost portion of the fish-transition zone, marked by a white mesh net and plastic pipe and sand bags along the streambank (used for sampling transport of prey and

detritus downstream). The upper 300-m reach starts at that stream sampling station

and extends 300 m upstream, and the lower 300-m reach starts 50 m downstream

from the stream sampling station and extends 350 m downstream.

Compare total stand density across a range of alder-conifer mixtures.

Compare understory plant diversity and biomass in stands across a range of alderconifer mixtures.

Compare stand growth and regeneration in stands across a range of alder-conifer mixtures.

Compare frequency and type of tree mortality (mode of tree death) in stands across a

range of alder-conifer mixtures.

Compare total stand density, tree diameter distribution, and size-specific tree mortality

across a range of riparian alder-conifer mixtures.

Compare the spatial distribution and stocking of alder and conifer trees between upland

and riparian stands across a range of alder-conifer mixtures.

Measure size and condition of woody debris entering the forest floor and streams,

including type of decay, species of wood, and presence or absence of attached root

systems.

Objective 1

Objective 2

Objective 3

Objective 4

Objective 5

Objective 6

Objective 7

Study Sites
48

Overstory vegetation data methods—

Data collection—Variable-radius plots will be used to assess stand density, the proportion of alder basal area in each stand, patterns of alder frequency and distribution

throughout the stand, and understory vegetation. Larger fixed-area plots will be used

to determine stand and tree growth, tree mortality, tree height growth patterns, and

the effect of stand density on understory vegetation.

The size and shape of each stand will be delineated, and a grid of 20 variable-radius

plots will be systematically installed throughout the stand. We will use a metric 4-basalarea-factor (BAF) prism plot (about 20-BAF, English units) to sample live and dead

trees in the expected tree diameter distribution of this study (Dillworth and Bell 1967). In

addition, we will install one 0.0025-ha fixed-area plot (radius 2.82 m) at each variableradius plot to measure live and dead trees at least 3 cm in diameter at breast height

(d.b.h.). The 3-cm d.b.h. tree threshold is the minimum diameter for the class of woody

debris known as “fine woody debris.”

Fixed-area plots will be installed at five randomly selected variable-radius plot locations

in each stand. Each fixed-area plot will contain nested circular plots to sample trees in

different size classes (Avery and Burkhart 1994): one 0.05-ha plot (radius 12.62 m),

and three 0.0025-ha plots (radius 2.82 m). One of the smaller plots will be located at

plot center, and two will be located 8 m from plot center in randomly selected directions.

All live and dead trees greater than 20 cm d.b.h. will be measured in the 0.05-ha plot.

All live and dead trees greater than 3 cm d.b.h. will be measured in the 0.0025-ha plots.

Two 1-m2 vegetation quadrats (quads) and two 2-m-radius shrub plots will be established at each variable-radius overstory plot. The quads and shrub plots will be located

4 m north and 4 m south of plot center, and the shrub plots will be centered on the vegetation quads. Vegetation cover for all vascular plants will be estimated, and tree regeneration will be tallied on the quadrat and shrub plots. One quadrat at each overstory plot

will be randomly selected for biomass sampling.

Within each 0.05-ha overstory plot, eight 1-m2 vegetation quads and eight 2-m-radius

shrub plots will be established. Four of the quads will be located 4 m from the plot center, one in each of the four cardinal directions (N, E, S, W), and will be used for biomass sampling. The other four quads will be 8 m from the plot center along the NE,

NW, SE, and SW directions, and will be used for vegetation cover estimates.

In the riparian stands, nested fixed-area plots will be used to assess stand density, the

proportion of alder basal area, alder stocking and distribution in the riparian zone, and

type of tree mortality. Seven pairs of nested fixed-area plots (one of each pair on the

left bank and one on the right) will be established along each 300-m reach of each

stream. Each fixed-area plot will contain a small (5 m2), a medium (25 m2), and a large

(50 m2) plot. The small plot will be 5 m along the bank of the stream by 1 m into the

riparian forest. The medium plot will be located 5 m parallel to the stream by 5 m into

the forest. The large plot will be established 5 m parallel to the streambank by 10 m into

the riparian forest (fig. 4). Plots will be sampled as one walks upstream, beginning at

the stream-pipe sampling reference point. Sites will be named according to their location relative to the reference point: upper or lower stream reach, plot location, distance,

and streambank. Left and right bank will be named while one looks upstream. The first

pair of plots of the upper 300-m reach will be located at the sampling pipe. Each successive pair will be 50 m from the beginning of the previous one, so that all seven pairs
49

of plots will be spaced evenly across the 300-m stream reach. The plots of the lower

300-m stream reach will start 50 m downstream from the stream sampling station and

extend 350 m downstream.

Canopy coverage measurements will be taken from a point in the center of the stream

2.5 m from the edge of the fixed-area plot and 2.5 m from the sampling pipe. Measuring points will be 50 m apart (fig. 4). There should be seven canopy coverage measurements within each 300-m stream reach.

Methods for woody debris data collection are detailed in appendix 4.

On each variable-radius plot, tree species, d.b.h., and crown position will be measured

for all tallied live trees within the prism plot. Species, d.b.h., and type of tree mortality

(uprooted, broken, dead standing, dead down) will be recorded for all dead trees. On

the 0.0025-ha fixed-area plot, tree species, d.b.h., and crown position will be measured

for all live trees at least 3 cm d.b.h.; species, d.b.h., and type of tree mortality will be

measured or noted for dead trees at least 3 cm d.b.h.

For each fixed-area nested plot, tree species, crown class, d.b.h., height, percentage

of live crown, and tree condition data such as top damage will be measured for all live

trees to provide current stand structural information. The same information listed above

will be collected for dead trees to provide information on tree mortality. The deterioration class (1= at least half the bark retained, 2 = less than half the bark retained) and

the type of wood decay (white rot or brown rot) will be noted for all dead trees over 3

cm d.b.h. on fixed-area plots. Tree crown width will be measured for two randomly selected trees from each species and crown class to determine tree crown dimensions.

Tree increment cores or stem sections will be taken from two randomly selected trees

from each species and crown class for tree-ring analysis to determine tree age and

changes in diameter growth. In addition, one dominant alder and one dominant conifer

tree will be selected from each 0.05-ha plot for tree age and height growth analysis.

These trees will be destructively sampled with stem sections collected at base, 1.3 m,

3 m, and every subsequent 2 m up the bole to the top of the tree.

On each fixed-area plot in the riparian stands, tree species, tree status (live or dead),

and d.b.h. will be measured for all tallied trees. For dead trees, the type of mortality

(uprooted, broken, dead standing, dead down, leaning over the stream) will be recorded. Within each small fixed-area nested plot all trees will be recorded; trees at

least 1 m tall or at least 3 cm d.b.h. will be tallied; and all trees at least 3 cm d.b.h. will

be measured. Within each medium fixed-area nested plot, all trees at least 3 cm d.b.h.

will be measured. Within each large fixed-area nested plot, all trees at least 15 cm

d.b.h. will be measured. Percentage of canopy coverage will be estimated by looking

through a 30-cm long, 10 cm (inside diameter) plastic tube. The observer will stand in

the center of the stream, hold the tube 90 degrees from level, look up at the forest

canopy, and estimate the percentage of alder, conifer, and open canopy (Piccolo and

Wipfli 2002).

Shrub cover will be estimated on small fixed-area plots located along streambanks

(fig. 4).
50

Data analysis—

• Overstory data

Tree data for the 20 variable-radius plots will be combined by stand to assess stand

density, the proportion of alder basal area in each stand, and patterns of alder frequency and distribution within each stand. Tree data for the five large fixed-area plots

will be combined by stand to determine stand and tree growth, tree mortality, tree

height and diameter distribution, and tree regeneration. Regression analyses will be

used to determine the relation between the proportion of alder basal area and the

following variables: stand density, wood production, stand growth, tree height and

diameter growth, and tree regeneration.

• Riparian stand data

Riparian tree data for all 13 streams will be combined by stream reach to assess

stand density, the proportion of alder basal area in each stand, species composition,

alder distribution and frequency, tree mortality, and tree diameter distribution. Regression analyses will be used to determine the relation between the proportion of

alder basal area and the following variables: stand density, wood production, and

diameter growth. Results will be compared to upland stands.

• Mortality data

Regression analyses will be used to explore how the proportion of alder in younggrowth stands influences woody debris recruitment, in total (i.e., number of stems

and volume), by tree species, by decay type, and with unique structures (e.g., attached root system). Mortality rates will be calculated from the number of live and

dead trees (extrapolated from nested plot data to per-hectare data) in each stand for

both alder and conifers. Mortality for each species also will be expressed as basal

area and percentage of basal area. Two-factor analysis of variance (ANOVA) will be

used to contrast the mean diameter of live and dead trees by tree species (alder or

conifer—first factor) and by site (second factor) to describe whether large or small

trees are dying (and giving rise to woody debris) in these stands. By using trees dissected for stem analysis, the amount of decay in live alder (percentage of infected

trees and mean percentage of volume affected per tree) will be contrasted by stand

age (40-year-old stands versus 100-year-old stands) by using Student’s t-test.

For the riparian stands, regression analyses will be used to assess how the proportion of alder may impact recruitment of woody debris in total and by tree species.

Mortality rates of alder and conifers will be calculated from the number of live and

dead trees in each stand. Mortality for each species also will be expressed as basal

area and percentage of basal area. Two-factor analysis of variance (ANOVA) will be

used to contrast the mean diameter of live and dead trees by tree species (alder or

conifer—first factor) and by site (second factor) to describe whether large or small

trees are dying (and giving rise to woody debris) in these stands. Differences in size

of woody debris recruited (i.e., diameter of dead trees) will be contrasted by site and

tree species by using analysis of variance.
51

Understory vegetation data methods—

Data collection—

• Ground vegetation cover

We will use cover classes (Daubenmire 1959) to visually estimate the percentage of

cover of each species rooted in the quad: conifer seedlings 3 to 10 cm tall, all ferns,

herbaceous plants, club mosses, common mosses and liverworts, and lichens (terrestrial and fallen epiphytics). Cover will be recorded for recently downed logs (trees that

have died and fallen since logging date). Estimates will be recorded with the accepted

abbreviations in the plant code list in field notebooks.

• Seedlings

We will tally and record the number of alder and conifer seedlings in the following

height classes:

(0.3) 0.1 to .5 m

(1) 0.5 to 1.5 m

(2) 1.5 to 2.5 m

(3) 2.5 to 3.5 m (<3.0 cm d.b.h.)

• Shrub cover

Cover of shrubs and conifer regeneration (10 cm tall and up to 3 cm d.b.h.) will be estimated by species in the 2-m-diameter shrub plot, whether or not they are rooted in the

plot. Cover of recently downed logs in the shrub plot will be recorded. The average

height of each shrub species in the shrub plot will be recorded. For the riparian stands,

on each small fixed-area plot, each shrub species and its relative abundance will be

recorded as percentage of cover.

• Biomass of clipped species

Biomass will be estimated for shrubs, conifer seedlings, ferns, and herbaceous plants

rooted within the quad. Generally, if a plant is more than 50 percent dead, it will not be

included in the samples. However, in the case of woody plants, dead branches will be

removed, and the rest of the plant will be included in the biomass.

Herbs: Collect subsamples of each species, including ferns, horsetails, and Lycopodium spp.

Shrubs: Separate each species into three components: leaves (including flowers), twigs

(growth of current year), and stems (prior years’ growth).

Conifer seedlings: Separate into two components: new (growth of current year) and old

(prior years’ growth).

Data analysis—Biomass data for the 20 vegetation quadrats at the variable-radius plot

locations will be combined by stand. Regression analyses will be used to determine

how the proportion of alder basal area affects species richness and biomass of vascular plants. Correlations between overstory and understory variables will be determined
52

from each alder-conifer stand sampled in this study. Overstory variables include biomass and canopy coverage of vascular plant species. Regression equations will quantify tradeoffs between stand structure and understory variables. Understory species

composition and biomass will be quantified in terms of food value for deer (deer-days

per ha) with a nutritional model for deer habitat (Hanley and Rogers 1989) described in

appendix 3. Biomass and cover data for the 0.2-ha overstory plots will be analyzed by

using regression equations to determine the effect of stand density and tree species

composition on biomass.

For the riparian stands, data for shrub percentage of cover will be combined by stream

reach, and regression analyses will be used to assess the relation between alder basal

area and species richness.
53

Appendix 2: Methods for Terrestrial Invertebrate Sampling

Objective 8

The terrestrial invertebrate study will use the same eight second-growth sites and the

same four old-growth sites selected for the wildlife studies (app. 3). Of the secondgrowth sites, four are alder-conifer mixed stands and four are conifer-dominated. The

four old-growth sites do not have alder.

Compare terrestrial invertebrate abundance and species richness in foliage,

branches, and stems of woody vegetation as well as in leaf litter in stands across a

range of alder-conifer mixtures.

Methods—

Data collection—

• Migrating arthropods

In each stand, traps will be permanently installed around the boles of three trees of

each tree species, by using methods described by Hanula and New (1996). Traps will

be located 1.5 to 2 m from the ground and will have a detachable cup that contains

70-percent ethanol solution. Invertebrates crawl up the tree into the trap and fall into

the collecting cup. Sampling will occur from May through September. Invertebrates

collected in the traps will be measured (Hodar 1996) and identified at least to family

(alder-conifer mixed stands: 22 sample times × 4 stands × 3 tree species [red alder,

western hemlock, Sitka spruce] × 3 bole traps per species = 792 samples; conifer

stands: 22 sample times × 4 stands × 2 tree species [western hemlock, Sitka spruce]

× 3 bole traps per species = 528 samples).

• Flying invertebrates

Malaise traps will be used to collect flying and some crawling invertebrates. The traps

will be set up for 48 hours. A sample will be collected after the first 24 hours and another before the traps are taken down. There will be one trap per stand, and each

stand will be sampled during the same period. Sampling will occur from May through

September. Invertebrates collected in the traps will be measured (Hodar 1996) and

identified at least to family (8 sampling times × 8 stands × 2 samples per stand = 128

samples).

• Foliage-feeding invertebrate sampling

Branch clippings will be sampled from red alder, western hemlock, and Sitka spruce

(Johnson 2000). Clippings will be taken from trees of various sizes and in areas near

openings and streams. If the crowns of Sitka spruce cannot be reached at a particular

site, Sitka spruce will not be sampled there. Sampling will occur from May through

September.

Once a week, three branch tips from three trees of each species will be clipped for

leaf-chewing invertebrates. Invertebrates will be shaken from a branch onto a white

bag, dispensed into a jar containing 70-percent ethanol solution, measured (Hodar

1996), and identified. The number of invertebrates per unit of foliage will be used to

determine the density of invertebrates (Werner 1969).

Study Sites
54

The ratio of invertebrate weight to foliage weight will be determined for each sample.

For western hemlock and Sitka spruce, the length of primary and secondary branches

will be recorded and used to estimate biomass of the needles. For red alder, leaf biomass will be determined by stripping the leaves into a paper bag, recording the weight

of the “wet” leaves and then subtracting the oven-dry weight. Invertebrate weight and

weight per unit weight of foliage will be estimated by using formulas based on body

length for each invertebrate order (Ganihar 1997) (alder-conifer mixed stands: 22

sample times × 4 stands × 3 tree species [red alder, western hemlock, Sitka spruce] ×

3 bole traps per species = 792 samples; conifer stands: 22 sample times x 4 stands ×

2 tree species [western hemlock, Sitka spruce] × 3 bole traps per species = 528

samples).

• Leaf-litter invertebrates

A fixed area of leaf litter will be randomly collected from study sites. Berlese funnels will

be used to separate the insects from the leaf litter. Four leaf litter samples will be collected per study site. This sampling will be done once in summer and once in fall. Invertebrates collected will be measured (Hodar 1996) and identified.

Data analysis—The number and morphological species description will be determined

for each tree species and study site. An average of the number and weight of each

species or grouping of species will be calculated for each sample date. Stand-to-stand

differences will be determined by using t-tests of the average number and weight of

invertebrates for each tree species. Frequency diagram will be produced for the number of individuals per date for each tree species.
55

Appendix 3: Methods for Wildlife Sampling

Objective 9

Study Sites We will select 12 young-growth forest stands (approximately 1 ha each) consisting of

at least 20 percent alder and 12 young-growth stands consisting almost exclusively of

conifer from stands within the Harris and Maybeso watersheds on Prince of Wales

Island, Alaska.

Compare bird density and species richness in alder-conifer mixed and conifer stands.

Bird species diversity and abundance methods—

Data collection—Point-count censuses (Bibby et al. 1992): Limited radius (40-m)

censuses of 8-minute duration will be conducted three times at each station during

the breeding season (early May until early to mid-July). Surveys will not be conducted

during heavy rain or high wind.

Data analysis—Compare estimated bird densities between stand types by using a

t-test. Bird density data from point-count censuses will be compared to measures of

understory vegetation and forest structure as reported in the vegetation component

of this study.

Conduct behavioral observations of foraging birds in mixed alder-conifer young-growth

stands.

Foraging data methods—

Data collection—Within stands of different habitat types, observe individual birds

during foraging, noting foraging substrate used and prey taken.

Data analysis—In these approaches, the bird is the experimental unit. Bird foraging

data are contrasted with substrate availability.

Compare nesting success in mixed alder-conifer young-growth stands and in coniferdominated young-growth stands.

Nesting success methods—

Data collection—

• Artificial nest predation (Bayne et al. 1997)

Make artificial thrush-sized eggs (23 × 17 mm) out of porcelain clay dipped in plastic

polymer (Plastidip1) and artificial thrush-sized nests out of moss (6- to 7-cm-diameter

nest cup, 10- to 15-cm outer diameter). Let nests and eggs air for 2 days to let odors

dissipate, and wear rubber gloves and boots during experiment to minimize scent trails

to nests. In early to mid-June, place two handmade moss nests containing two artificial

eggs at 15-m intervals along 150-m transects (nest density is 0.13 nests per meter or

1 nest every 7.5 m, for a total of 20 nests per transect). Place nests in “thrush-like”

places (i.e., up to 2.5 m aboveground on a stump or log or next to a tree trunk). Check

Objective 10

Objective 11

1 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.
56

nests 3 and 12 days after experiment begins. The day the eggs are placed is day 1.

Record all types of disturbance to nest and eggs, inspecting eggs for “bite” or peck

marks to determine predator.

• Natural nest monitoring

Natural nests will be located and monitored every 1 to 3 days until the young fledges

or the nest fails. Nest success will be calculated by using the Mayfield method

(Hensler and Nichols 1981; Mayfield 1961, 1975) if sample sizes permit (i.e., n > 15)

or by simple percentages if sample sizes are smaller. Once a nest is no longer active

(after failure or fledging of all nesting attempts for the season within that nest site), the

following physical characteristics of the nest site will be recorded: height above ground,

support structure, and estimated percentage of concealment of nest in four cardinal

directions and from above and below.

Data analysis—Angular transformed percentage of data of proportion of nests

depredated will be analyzed by using a Wilcoxon two-sample test. Mayfield daily

nest survival rates will be analyzed with the computer program CONTRAST, which

uses a chi-square analysis with multiple comparisons (Hines and Sauer 1989, Sauer

and William 1989). Angular transformed percentage of data of simple percentage

nesting success will be analyzed with a Wilcoxon two-sample test. Differences in nestsite characteristics of safe and unsafe nest sites and between nests in habitat types

(mixed alder-conifer and conifer) will be analyzed with a Wilcoxon two-sample test.

Compare summer food value for deer in young-growth stands differing in their

proportion of alder component.

Summer food resources for black-tailed deer methods—

Data collection—Data will be collected as described in the understory vegetation

component in appendix 1.

Data analysis—Understory species composition and biomass will be quantified in

terms of food value for deer (deer-days per ha) by use of a nutritional model for deer

habitat evaluation (Hanley and Rogers 1989). The Hanley and Rogers model converts

a multivariate data set of biomass, digestible energy, and digestible protein for the full

list of plant species into one measure of food value for deer—the number of deer-days

that could be supported by that food at a specified level of nutritional requirement.

Values of digestible energy and digestible protein for each species will come from a

composite, regional database of nutritional data for forages in southeast Alaska.2

Objective 12

2 Hanley, T.A. 2002. Unpublished data. On file with: Forestry

Sciences Laboratory, 2770 Sherwood Lane, Suite 2A, Juneau,

AK 99801.
57

Appendix 4: Methods for Aquatic and Riparian Invertebrate Sampling

Objective 13

Study Sites We will select a subset of study sites used for all study components, as well as additional riparian stands that meet criteria particular to aquatic ecology study (riparian

stands with alder blend, fish transition zones, class 3 and 4 streams), for a total of

13 study streams.

Compare the amount of export of invertebrates and detritus from fishless headwater

streams to downstream fish-bearing habitats across a gradient of riparian alder abundance.

Invertebrate transport data methods—

Data collection—We will measure transport of invertebrates and detritus from fishless habitats to fish-bearing transition zones across a range of alder-conifer mixtures.

Sampling protocols will be similar to those described in Allan (1995), Hauer and

Lamberti (1996), and Wipfli and Gregovich (2002).

We will select 13 streams within vegetation plots with riparian habitats that represent

a range of alder gradient. Transport of invertebrates and detritus will be sampled with

250-μm mesh nets attached to 10-cm plastic tubes anchored in the stream with sandbags. Samples will be collected three times a year (April, June, and August) at all sites.

Nets will be placed in streams on the same day for all streams, and contents will be

collected one day later (1 sample × 13 streams × 3 seasons = 39 drift samples per

year). Invertebrates will be placed in ethanol, and later sorted, counted, and identified

to the lowest reliable taxon (likely genus) for analysis.

Immediately after the June 2000 sampling, we will select five of the streams for use in

testing the hypothesis that the transport rate of invertebrates (number of invertebrates

per unit volume of water) is equal along the stream reach. We will place one net in

each of the five streams (following the protocol described above) 150 m upstream of

the first point sampled (see above) for a 24-hour period.

Data analysis—Data will be natural logarithm-transformed and analyzed by using

least squares linear regression, with percentage of alder as the independent variable

and invertebrate abundance (count density) as the dependent variable (SAS 1989).

Data for the five-stream experiment will be analyzed with a t-test, and streams will

be the replicates (n = 5).

Compare invertebrate abundance in and along headwater streams, including on woody

debris at various stages of decay, in stands with different alder-conifer mixtures.

Habitat use by invertebrates methods—

Data collection—Sampling protocols used in similar studies in other regions

(Anderson et al. 1978, Cummins et al. 1989, Hax and Golladay 1993, Sedell et al.

1975, Vannote et al. 1980) will be followed and modified where appropriate.

This component looks at decay of woody debris and processing of leaf litter in stream

and riparian habitats. We are interested in comparing the decay processes and associated fungal and macroinvertebrate communities that break down alder and spruce

Objective 14
58

woody debris and leaf litter in streams and associated riparian habitats in forested

uplands in southeast Alaska. The study will compare the various pathways of decay

and the macroinvertebrates colonizing woody debris in streams and riparian zones.

• Stream habitat

We will compare invertebrate communities (e.g., abundance, species composition)

on spruce and alder woody debris, and compare loss of wood mass over time. Boles

from alder and spruce will be collected, cut into small pieces (5 cm diameter by 20 cm

length), dried, weighed, and sorted into four categories: early decay alder, late decay

alder, early decay spruce, and late decay spruce. Pieces will be arranged in random

order and secured with cable locks to rigid plastic rafts so that each raft has one piece

from each category. Rafts will be strengthened by a gutter nail affixed perpendicular to

the wood pieces, and also labeled with a steel tag. Nine rafts will be placed at seven

sites in six streams in forests with a range of alder component (9 rafts × 7 sites = 63

rafts × 4 pieces per raft = 252 total pieces). Gutter nails and steel leaders will be used

to anchor rafts to the streambed. Placements will be in pools, runs, and riffles. Rafts

will be set in June 2000.

Three randomly selected rafts will be collected from each of seven sites in July 2001,

and another three in July 2002. Rafts will be pulled from the stream and transferred to

aluminum trays. Each piece will be transferred to a 5-gallon (19-liter) bucket, rinsed

with a portable pressurized sprayer, and decay will be measured with a penetrometer

(seven measurements per piece). The penetrometer will be modified for woody debris

by affixing a size-14 sewing needle to the otherwise blunt tip. Contents of the bucket

will be strained through a 250-μm sieve, and the invertebrates will be washed into

sealed, 250-mL plastic bags with 80-percent ethanol solution. All pieces of wood will

be dried and weighed to compare loss of wood mass over time, and invertebrates will

be counted and identified to lowest reliable taxon.

We will compare invertebrate communities (percentage colonization) on woody debris

from two distinct decay classes (early and late decay) of alder and spruce. Naturally

occurring pieces (roughly 5 cm diameter by 25 cm length) of alder and spruce will be

collected from 13 streams in forests with a range of alder component (2 early decay ×

2 late decay × 2 tree species × 13 streams = 104 total pieces of woody debris). Sampling will be conducted during May and July of 2001 and 2002. Selected pieces will be

removed from the stream (with a D-frame aquatic net placed downstream when appropriate). Decay and invertebrates will be measured as described above. Surface area

will be figured for each piece of wood to calculate percentage colonization.

• Riparian habitat

We will compare invertebrate abundance and species composition on conifer and

alder woody debris in riparian areas along streams. Methods mirror those described

above for the stream component. We will collect old-decay and recent-decay conifer

and alder wood debris from the riparian areas along 11 streams in forests with a range

of alder component. Stream locations are separated into five 30-m sections, for a total

of 150 m. Wood debris samples will be randomly selected from four of the five sections

(4 samples × 11 streams = 44 total woody debris samples). A saw will be used to cut

pieces to fit into 1-gallon (3.8-liter) plastic bags. Invertebrates will be washed from

wood samples into a 5-gallon bucket with a pressurized backpack sprayer. Each wood

piece will be dissected to remove all invertebrates that cannot be removed by pressure

washing. Each wood piece will be measured so that biomass may be estimated. A
59

250-μm sieve will be used to separate excess water from the sample, and the sample

will be placed into a sealed, 250-mL plastic bag, preserved with 80-percent ethanol

solution, and returned to the lab for processing. Invertebrates will be collected from

each wood sample, identified to the lowest reliable taxon, and counted.

We will compare invertebrate communities in alder and conifer leaf packs in riparian

habitats along headwater streams. We will collect three litter samples of natural detritus

from riparian habitats along each of the 11 streams (3 samples per habitat x 11 streams

= 33 detritus samples). The three samples will be randomly chosen from three of the

five stream sections. A section of 6-in (15.2-cm)-diameter stovepipe will be dropped at

random within 1 m of the stream. Litter and woody debris contained within the stovepipe will be placed in sealed plastic bags. Invertebrates will be separated from the litter

by using a Berlese funnel. Samples will be hand picked to assure all invertebrates have

been separated from the litter. Samples will be transferred into a scintillation vial, preserved in 80-percent ethanol solution, and returned to the lab for processing. Each

litter sample will be sorted into the following groups: alder, conifer, and woody debris.

Leaves and woody debris will be dried in an oven at approximately 42 °C for 24 hours.

Each component will be weighed individually so that alder and conifer composition can

be estimated. Invertebrates will be collected from detritus, identified to the lowest reliable taxon, and counted.

Data analysis—Data for the 11-stream experiment will be log-transformed and analyzed with least squares linear regression, with percentage of alder as the independent

variable and invertebrate abundance (count density) as the dependent variable (SAS

version 8.0). Data for the five-stream experiment will be analyzed by using a t-test with

streams as the replicates (n = 5).

Compare nutrient concentrations among 13 streams that differ in the amount of

upstream red alder and in discharge and disturbance history.

Stream nutrient data methods—

Thirteen small permanent streams (about 10- to 20-ha subbasins) across the range

of 0 to 39.5 m2 basal area alder coverage will be sampled for concentrations of various

form of nitrogen and phosphorus during the 2001 growth season. Regressions of

various nutrient parameters against the aerial alder coverage will be used to detect

the effect of alder on stream nutrient concentrations. Because other factors such

as hydrologic contributing area and disturbance history also may influence nutrient

dynamics through their effect on runoff patterns and retention, we will partition the

data into categories suggested by the geomorphic study and explore their effects

with multiple regression.

Objective 15
60

Appendix 5: Methods for Fish Sampling

Objective 16

Study Sites Thirteen catchments in the Maybeso and Harris watersheds on Prince of Wales Island

will be selected for the entire set of studies in the Wood Compatibility Initiative. All

streams will be surveyed for fish distribution. We will select a subset of these for an

intensive analysis of fish populations.

Stream selection criteria will be perennial flow allowing connection to downstream

main-stem rivers, presence of spawning gravel, and absence of active beaver ponds

or lakes. Study reaches will be designated on streams with headwaters having mixed

alder-conifer and conifer-dominated riparian zones. A transition reach that extends to

the upper limit of salmonid presence (gradient >7 percent) will be designated on every

stream, while a moderate gradient reach (gradient 4 to 7 percent) and flood-plain reach

(gradient <4 percent) will be designated on streams where these conditions exist and

sampling was practical. Each reach will be identified and marked to serve as a permanent site for the duration of the study.

Compare salmonid densities in stream reaches below and within conifer-dominated

and alder-dominated upslope riparian forests.

Assess seasonal variation and relative species abundance of salmonids in upper

high-gradient reaches (transition zones).

Assess the relation between the abundance of alder in a riparian forest and stream

inputs and processes.

Salmonid density and distribution and habitat data methods—

Data collection—

• Fish populations

For the initial fish-sampling period (summer 2000), up to three stream reaches (one in

each gradient zone) in each study stream will be isolated with nets or natural barriers

and sampled for salmonid populations. A target length of 100 m will be used to designate study reaches, although the characteristics of individual streams may force the

use of shorter sections.

Each sample reach will be divided into 10-m subsections with beach seines or natural

barriers to reduce potential fish movement, and fish will be captured with minnow

traps or by electrofishing. Population estimates in each reach will be made with the

generalized removal method (three-pass depletion) (White et al. 1982). Fish will be

identified, counted, and measured, and a subsample of the catch weighed. In subsequent sampling periods, streams will be intensively sampled by using electrofishing

and the three-pass depletion population estimate technique. To determine the point at

which fish reach their upper limit in streams, we will electrofish in upstream reaches

(>10 percent gradient) moving upstream until fish are no longer detected.

Objective 17

Objective 18
61

• Habitat measurements

Each fish sampling reach is surveyed by using a modified tier III habitat survey

(USDA FS 2001). Key measurements include size, number, and volume of large

wood pieces, pool size and depth, width-to-depth ratio, substrate composition, and

sediment retention. A more intensive large woody debris (LWD) survey also will be

used in a stratified sample at all streams. Channel conditions above study sites will be

surveyed to determine landslide history. Comparison of key variables (e.g., LWD and

sediment retention) will be made between sites with and without recent (<20 years)

landslides.

Data analysis—Upslope effects on fish population will be measured by using the ratio

of conifer to alder (X1) and landslide frequency (X2) above the transition zones of each

stream. The relation between these two variables and the density of resident 60- to

180-mm Dolly Varden (Y1) and 60- to 180-mm cutthroat trout (Y2) will be examined

with a linear model and tested for interaction between X1 and X2:

YI = a = b1X1 + b2X2 + b3X1X2 .

The model will be applied to fish in the transition zone for three separate periods:

spring, summer, and fall.

Longitudinal effects on fish density will be compared across seasons for the transition

zone, moderate zone, and flood plain. Separate comparisons will be made for Dolly

Varden and cutthroat trout by using a two-way ANOVA. Juvenile coho salmon are not

expected to occur in most transition zones and will be excluded. If insufficient samples

occur with cutthroat trout, then only Dolly Varden will be used in the analysis. Species

distribution (i.e., proportion of species in each sample unit) will be compared through a

chi-square test by using the same comparison used for longitudinal distribution described below.

The following tabulation shows the statistical design for comparison of longitudinal

distribution of salmonid populations (Xi,k where X = fish density, i = reach, y = season).

Reach Spring Summer Fall

Transition X1,1 X1,2 X1,3

Moderate X2,1 X2,2 X2,3

Flood plain X3,1 X3,2 X3,3

A stepwise regression will be used to measure habitat response to upslope riparian

forest (Y1 = conifer to alder ratio) and disturbance history (Y2 = landslide frequency).

Habitat variables are number of pieces of large wood (X1), pool volume (X2), and

substrate composition (X3).
62

Appendix 6: Methods for Geomorphic Processes

Objective 19

Study Sites The goals of the Wood Compatibility Initiative (WCI) are to improve our understanding

of young-growth ecosystem function and to understand how alder influences trophic

links and processes in managed landscapes. Meeting these goals depends on an understanding of both natural and human-caused distributions of disturbance processes

in a watershed context. To achieve these goals, study sites are selected at two scales:

landscape and individual headwater channel. The landscape-scale analysis is conducted to describe dominant disturbance pattern (both natural and human caused)

within entire watersheds (i.e., Maybeso and Karta). Individual headwater channels (20

sites total: 14 in second growth and 6 in old growth) are studied to identify dominant

processes (landslides, mortality, bank erosion, and catastrophic blowdown).

Describe geomorphic processes and land use history at study sites selected by

WCI colleagues.

Landscape analysis methods—

Data collection—Using a model of blowdown (Kramer et al. 2001), a model estimating

bank erosion (Hooke 1980), estimates of forest mortality,1 and a model of landslide

occurrence and deposition (assumption that landslides initiate on slopes over 30° and

deposit on slopes between 5° and 15°) (Johnson et al. 2000), we will produce a map

indicating general process occurrence within watersheds.

We will estimate the relative magnitude of each disturbance process within the

Maybeso and Karta watersheds. We also will review literature and examine land

use history by using aerial photographs and timber harvest records.

Data analysis—We will describe natural disturbance history and land use history

within Maybeso and Karta. We also will determine the processes (by the means described in a previous section) that introduce the majority of woody debris deposits.

Assess processes that deliver sediment and woody debris to streams from alderdominated and conifer-dominated forests, and compare delivery rates.

Headwater channel process analysis methods—

Data collection—We will select sites having variation in upslope topography and

variation in percentage cover of alder and conifer, and then delineate forest cover

for headwater regions contributing to each site by using infrared aerial photographs

and thematic mapper. Estimates will be made of:

• Landslide rate2

• Bank erosion (Hooke 1980)

Objective 20

1 Hennon, P.E. 1997. Unpublished data. On file with:

Forestry Sciences Laboratory, 2770 Sherwood Lane,

Suite 2A, Juneau, AK 99801.

2 Helmers, A.E. 1961–1985. Unpublished landslide

inventory. On file with: Forestry Sciences Laboratory,

2770 Sherwood Lane, Suite 2A, Juneau, AK 99801.
63

• Catastrophic blowdown (Kramer et al. 2001)

• Forest mortality (Hennon, collaborator in WCI).

We will classify the mountain slope landform (USDA FS 1997) contributing to deposition sites by using a measuring tape and clinometer to assess longitudinal profiles

above, through, and below transition areas from steep slopes (>15°, 25 percent) to

low-gradient slopes (<4°, 7 percent). We will plot longitudinal profiles and categorize

the variation as follows:

• Gradual slope change

• Moderate slope change

• Rapid slope change (sharply concave).

We will compare channel characteristics (channel substrate size) by using pebble

counts (Wolman 1954) and channel widths directly above and below transition areas.

We will estimate area contributing to each transition area by using field and map measurements.

Data analysis—We will create a series of tables to describe topographic characteristics of WCI study sites including average slope gradient of uplands contributing to

streams, longitudinal gradients, substrate size above and below transition areas, vegetation type, landform type, landslide history, land use history, and an estimate of contributing area. We will describe dominant processes and evaluate the relation between

the continuous variables percentage alder and landslide rate. We will determine the

degree of variance explained by the regression and examine the F-statistics (overall

relation linear trend). We will then examine t-statistics for each parameter and assess

the power of the test. We will examine adherence of the assumptions of regression by

using residual tables. Differences will be judged to be significant at a = 0.05 (p = 0.10).

Estimate differences in woody debris loading and sediment storage capacity in streams

within conifer-dominated and alder-dominated forests.

Sediment storage capacity methods—

Data collection—Distributions and accumulations of organic debris along headwater

channels will be measured. Organic debris in streams will be classified as woody debris

or fine organic debris (FOD). Woody debris will be further classified as large woody

debris (LWD) (pieces >0.5 m in length and ≥0.1 m in diameter) or fine woody debris

(FWD) (pieces ≥0.5 m in length and 0.03 to 0.1 m in diameter). The LWD pieces recruited before logging activities will be identified as “legacy” based on decay process

and the absence of a cut edge. To quantify the distribution and accumulation of woody

debris at each site, the following properties of LWD will be measured: in-channel,

bankfull, and total lengths; diameter; position; and orientation. In-channel length is the

length of a LWD piece located within the wet channel width that significantly dissipates

flow energy and affects sediment transport. Bankfull length is the length of a LWD

piece within the banks of the channel. Total length is the length of the entire piece of

Objective 21
64

LWD, including terrestrial portions. Diameter at the middle of each woody debris piece

will be recorded. Volume (V) of LWD will be calculated as follows for the in-channel,

bankfull, and total volume components of LWD:

V = π × (D/2)2 × L ,

where D is the mid-log diameter and L is length.

All LWD pieces will be classified as functional (interacting directly with streams),

transitional (suspended just above streams and decomposed enough to interact

with streams in the near future), and nonfunctional (no interaction with streams or

suspended well above channels). Orientation of LWD will be measured in relation to a

line parallel to the channel axis to determine the degree of interaction of LWD pieces

with streams. Both left-hand (+) and right-hand (-) orientations of LWD from 0° to 90°

were recorded in 5° intervals.

The FWD will be surveyed for channel position and number of pieces. Volume of

FOD, such as accumulations of leaves, branches, and fine logging slash, will be categorized as small (<0.01 m3), medium (0.01 to 0.1 m3), and large (≥0.1 m3) volumes

where FOD accumulations contributed to or formed a sediment wedge.

Sediment storage behind woody debris and other obstructions (e.g., boulders and

bedrock) will be measured in these headwater streams based on the geometry of

sediment wedge (width (w), length of the wedge (Lw), and average depth at the front

of the wedge (d)). Average depth of the sediment wedge will be measured by using a

sediment probe at several points. The cause of sediment deposition will be categorized

according to the formation elements of debris dam: LWD, FWD, FOD, rocks, and bedrock. The volume of sediment stored behind woody debris and other obstructions will

be computed based on a rectilinear pyramid:

Sediment volume = (w × Lw × d)/3 ,

The approximation of a pyramid-shaped wedge is appropriate because the upstream

end of stored sediment typically converges to a point in these small channels.

Data analysis—We will evaluate the relation between the continuous variables

percentage basal area alder and sediment storage: We will determine the degree

of variance explained by the regression and examine the F-statistics (overall relation

linear trend). We then will examine t-statistics for each parameter and assess the

power of the test. We will examine adherence to the assumptions of regression by

using residual tables. A significance level of α = 0.05 will be used for all statistical

analyses.
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