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Appendix A-List of Common and Scientific

Names Used in This Volumea

Common name Scientific name

INSECTS

Western spruce budworm Choristoneura occidentalis

REPTILES

Lizards

Northern alligator lizard

Sagebrush lizard

Southern alligator lizard

Western fence lizard

Western skink

Elgaria coerulea

Sceloporus graciosus

Elgaria multicarinata

Sceloporus occidentalis

Eumeces skiltonianus

Snakes

California mountain kingsnake

Common garter snake

Northwestern garter snake

Racer

Ringneck snake

Rubber boa

Sharptail snake

Western aquatic garter snake

Western rattlesnake

Western terrestrial garter snake

Lampropeltis zonata

Thamnophis sirtalis

Thamnophis ordinoides

Coluber constrictor

Diadophis punctatus

Charina bottae

Contia tenuis

Thamnophis couchii

Crotalus viridis

Thamnophis elegans

Turtles

Western pond turtle

Footnotes on page 521.

Clemmys marmorata

513
Common name Scientific name

AMPHIBIANS

Mole Salamanders

Long-toed salamander

Northwestern salamander

Giant and Olympic Salamanders

Cope’s giant salamander

Olympic salamander

Pacific giant salamander

Rocky Mountain giant salamander

Lungless Salamanders

Arboreal salamander

Black salamander

California slender salamander

Clouded salamander

Del Norte salamander

Dunn’s salamander

Ensatina

Larch Mountain salamander

Oregon slender salamander

Plethodontid salamanders

Redback salamander

Siskiyou Mountains salamander

Van Dyke’s salamander

Western redback salamander

Woodland salamanders

Newts

Roughskin newt

Frogs

Cascades frog

Foothill yellow-legged frog

Pacific treefrog

Red-legged frog

Tailed frog

Toads

Western toad

MAMMALS

Shrews

Marsh shrew

Masked shrew

Montane shrew

Pacific shrew

Trowbridge’s shrew

Vagrant shrew

Water shrew

Footnotes on page 521.

514

Ambystoma macrodactylum

Ambystoma gracile

Dicamptodon copei

Rhyacotriton olympicus

Dicamptodon tenebrosusb

Dicamptodon atterimusb

Aneides lugubris

Aneides flavipunctatus

Batrachoceps attenuatus

Aneides ferreus

Plethodon elongatus

Plethodon dunni

Ensatina eschscholtzii

Plethodon larselli

Batrachoceps wrightii

(lungless or woodland salamanders in the family Plethodontidae, including

Aneides spp., Batrachoceps spp., Ensatina eschscholtzii, and Plethodon spp.)

Plethodon cinereus

Plethodon stormi

Plethodon vandykei

Plethodon vehiculum

(see plethodontid salamanders)

Taricha granulosa

Rana cascadae

Rana boylii

Hyla regilla

Rana aurora

Ascaphus truei

Bufo boreas

Sorex bendiriiC

Sorex cinereus

Sorex monticolusd

Sorex pacificus

Sorex trowbridgii

Sorex vagrans

Sorex palustris
Common name Scientific name

Moles

Coast mole

Shrew-mole

Townsend’s mole

Scapanus orarius

Neurotrichus gibbsii

Scapanus townsendii

Bats

Big brown bat

California myotis

European pipistrelle

Fringed myotis

Hoary bat

Keen’s myotis

Little brown myotis

Long-eared myotis

Long-legged myotis

Silver-haired bat

Townsend’s big-eared bat

Western small-footed myotis

Yuma myotis

Eptesicus fuscus

Myotis californicus

Pipistrellus pipistrellus

Myotis thysanodes

Lasiurus cinereus

Myotis keenii

Myotis lucifugus

Myotis evotis

Myotis volans

Lasionycterus noctivagans

Plecotus townsendii

Myotis ciliolabrum

Myotis yumanensis

Rabbits and Pikas

Pika

Squirrels and Chipmunks

Douglas’ squirrel

Northern flying squirrel

Townsend’s chipmunk

Western gray squirrel

Yellow-pine chipmunk

Pocket Gophers

Botta’s pocket gopher

Northern pocket gopher

Western pocket gopher

Ochotona princeps

Tamiasciurus douglasii

Glaucomys sabrinus

Tamias townsendiie

Sciurus griseus

Tamias amoenus

Thomomys bottae

Thomomys talpoides

Thomomys mazama

Mice, Rats, and Voles

Black rat

Bushy-tailed woodrat

California vole

Creeping vole

Deer mouse

Dusky-footed woodrat

Forest deer mouse

Heather vole

Long-tailed vole

Montane vole

Pinon mouse

Red-backed voles

Red tree vole

Southern red-backed vole

Townsend’s vole

Rattus rattus

Neotoma cinerea

Microtus calijornicus

Microtus oregoni

Peromyscus maniculatus

Neotoma fuscipes

Peromyscus oreasf

Phenacomys intermedius

Microtus longicaudus

Microtus montanus

Peromyscus truei

Clethrionomys spp.

Arborimus longicaudus8

Clethrionomys gapperi

Microtus townsendii

Footnotes on page 521.
Common name Scientific name

Mice, Rats, and Voles--continued

Water vole

Western red-backed vole

White-footed mouse

Jumping Mice

Pacific jumping mouse

Western jumping mouse

Carnivores

Coyote

Ermine

Fisher

Grizzly bear

Marten

Wolf

Ungulates

Elk

Mountain goat

Mule deer

White-tailed deer

BIRDS

Swans, Geese, and Ducks

Bufflehead

Barrow’s goldeneye

Common merganser

Hooded merganser

Wood duck

Hawks and Vultures

Bald eagle

Cooper’s hawk

Northern goshawk

Red-shouldered hawk

Sharp-shinned hawk

Turkey vulture

Grouse and Quail

Blue grouse

California quail

Capercaillie

Mountain quail

Ruffed grouse

Murrelets

Marbled murrelet

Pigeons and Doves

Band-tailed pigeon

Mourning dove

Footnotes on page 521.

Microtus richardsoni

Clethrionomys californicus

Arborimus albipesi

Zapus trinotatus

Zapus princeps

Canis latrans

Mustela erminea

Martes pennanti

Ursus arctos

Martes americana

Canis lupus

Cervus elaphus

Oreamnos americanus

Odocoileus hemionus

Odocoileus virginianus

Bucephala albeola

Bucephala islandica

Mergus merganser

Lophodytes cucullatus

Aix sponsa

Haliaeetus leucocephalus

Accipiter cooperii

Accipiter gentilis

Buteo linneatus

Accipiter striatus

Cathartes aura

Dendragapus obscura

Callipepla californica

Tetrao urogallus

Oreortyx pictus

Bonasa umbellus

Brachyramphus marmoratus

Columba fasciata

Zenaida macroura

516
Common name Scientific name

Owls

Barred owl

Flammulated owl

Northern pygmy-owl

Northern saw-whet owl

Northern spotted owl

Western screech-owl

Hummingbirds and Swifts

Allen’s hummingbird

Anna’s hummingbird

Rufous hummingbird

Vaux’s swift

Kingfishers

Belted kingfisher

Woodpeckers

Acorn woodpecker

Downy woodpecker

Hairy woodpecker

Ivory-billed woodpecker

Northern flicker

Pileated woodpecker

Red-breasted sapsucker

Red-cockaded woodpecker

White-headed woodpecker

Flycatchers

Ash-throated flycatcher

Dusky flycatcher

Hammond’s flycatcher

Olive-sided flycatcher

Western flycatcher

Western wood-pewee

Jays, Crows, and Ravens

American crow

Common raven

Gray jay

Scrub jay

S teller’s jay

Chickadees, Nuthatches, and Creepers

Black-capped chickadee

Brown creeper

Brown-headed nuthatch

Chestnut-backed chickadee

Mountain chickadee

Red-breasted nuthatch

White-breasted nuthatch

Strix varia

Otus flammeolus

Glaucidium gnoma

Aegolius acadicus

Strix occidentalis caurina

Otus kennicottii

Selasphorus sasin

Calypte anna

Selasphorus rufus

Chaetura vauxi

Ceryle alcyon

Melanerpes formicivorus

Picoides pubescens

Picoides villosus

Campephilus principalis

Colaptes auratus

Dryocopus pileatus

Sphyrapicus ruber

Picoides borealis

Picoides albolarvatus

Myiarchus cinerascens

Empidonax oberholseri

Empidonax hammondi

Contopus borealis

Epidonax difficilis

Contopus sordidulus

Corvus brachyrhynchos

Corvus corax

Perisorius canadensis

Aphelocoma coerulescens

Cyanocitta stelleri

Parus atricapillus

Certhia americana

Sitta pusilla

Parus rufescens

Parus gambeli

Sitta canadensis

Sitta carolinensis
Common name Scientific name

Wrens

Bewick’s wren

House wren

Winter wren

Kinglets

Golden-crowned kinglet

Thryomanes bewickii

Troglodytes aedon

Troglodytes troglodytes

Regulus satrapa

Ruby-crowned kinglet Regulus calendula

Thrushes

American robin

Hermit thrush

Swainson’s thrush

Townsend’s solitaire

Varied thrush

Bushtits and Wrentits

Bushtit

Wrentit

Vireos

Hutton’s vireo

Solitary vireo

Warbling vireo

Wood Warblers

Bachman’s warbler

Black-throated gray warbler

Golden-winged warbler

Hermit warbler

McGillivray’s warbler

Nashville warbler

Orange-crowned warbler

Swainson’s warbler

Townsend’s warbler

Wilson’s warbler

Yellow-rumped warbler

Tanagers

Western tanager

Sparrows

Bachman’s sparrow

Chipping sparrow

Dark-eyed junco

Fox sparrow

Lazuli bunting

Rufous-sided towhee

Song sparrow

Footnotes on page 521.

Turdus migratorius

Catharus guttatus

Catharus ustulatus

Myadestes townsendi

Ixoreus naevius

Psaltriparus minimus

Chamaea fasciata

Vireo huttoni

Vireo solitarius

Vireo gilvus

Vermivora bachmanii

Dendroica nigrescens

Vermivora chrysoptera

Dendroica occidentalis

Oporornis tolmiei

Vermivora ruficapilla

Vermivora celata

Lymnothlypis swainsonii

Dendroica townsendi

Wilsonia pusilla

Dendroica coronata

Piranga ludoviciana

Aimophila aestivalis

Spizella passerina

Junco hyemalis

Passerella iliaca

Passerina amoena

Pipilo erythrothalmus

Melospiza melodia

518
Common name Scientific name

Blackbirds, Cowbirds, and Orioles

Brown-headed cowbird

Northern oriole

Red-winged blackbird

White-winged crossbill

Finches

Black-headed grosbeak

Evening grosbeak

Pine siskin

Purple finch

Red crossbill

White-winged crossbill

CONIFEROUS TREES (SOFTWOODS)

Yews

Pacific yew

Pines

Jeffrey pine

Knobcone pine

Lodgepole pine

Pinyon pine

Ponderosa pine

Sugar pine

Western white pine

Spruces

Black spruce

Engelmann spruce

Sitka spruce

White spruce

Hemlocks

Mountain hemlock

Western hemlock

True Firs

California red fir

Grand fir

Noble fir

Pacific silver fir

Subalpine fir

True firs

White fir

Douglas-firs

Douglas-fir

Redwoods

Redwood

Footnotes on page 521.

Molothrus ater

Icterus galbula

Agelaius phoeniceus

Loxia leucoptera

Pheucticus melanocephalus

Coccothraustes vespertinus

Carduelis pinus

Carpodacus purpureus

Loxia curvirostra

Loxia leucoptera

Taxus brevifolia

Pinus jeffreyi

Pinus attenuata

Pinus contorta

Pinus edulis

Pinus ponderosa

Pinus lambertiana

Pinus monticola

Picea mariana

Picea engelmannii

Picea sitchensis

Picea glauca

Tsuga mertensiana

Tsuga heterophylla

Abies magnijiica

Abies grandis

Abies procera

Abies amabilis

Abies lasiocarpa

Abies spp.

Abies concolor

Pseudotsuga menziesii

Sequoia sempervirens
Common name Scientific name

Cedars

Alaska-cedar

Incense-cedar

Port-Orford-cedar

Western redcedar

FLOWERING TREES (HARDWOODS)

Bigleaf maple

Bitter cherry

Black cottonwood

California black oak

California-laurel

Canyon live oak

Cascara buckthom

Golden chinkapin

Oregon ash

Oregon white oak

Pacific dogwood

Pacific madrone

Red alder

Scouler willow

Tanoak

Vine maple

SHRUBS AND HERBS

Alaska huckleberry

Blackberry

California hazel

Canadian thistle

Common pipsissiwa

Dwarf rose

Fireweed

Hedge-nettle

Huckleberry

Little pipsissiwa

Ocean-spray

Oregongrape

Oregon oxalis

Pacific rhododendron

Pinesap

Red huckleberry

Salal

Salmonberry

Thimbleberry

Thin-leaved huckleberry

Twinflower

FERNS

Bracken fern

Swordfem

Footnotes on page 521.

Chamaecyparis nootkatensis

Libocedrus decurrens

Chamaecyparis lawsoniana

Thuja plicata

Acer macrophyllum

Prunus emarginata

Populus trichocarpa

Quercus kelloggii

Umbellularia californica

Quercus chrysolepis

Rhamnus purshiana

Castanopsis chrysophylla

Fraxinus lattfolia

Quercus garryana

Cornus nuttallii

Arbutus menziesii

Alnus rubra

Salix scouleri

Lithocarpus densiflorus

Acer circinatum

Vaccinium alaskaense

Rubus spp.

Corylus cornuta

Cirsium arvense

Chimaphila umbellata

Rosa gymnocarpa

Epilobium spp.

Stachys cooleyae

Vaccinium spp.

Chimaphila menziesii

Holodiscus discolor

Berberis nervosa

Oxalis oregana

Rhododendron macrophyllum

Hypopitys monotropa

Vaccinium parvifolium

Gaultheria shallon

Rubus spectabilis

Rubus parviflorus

Vaccinium membranaceum

Linnaea borealis

Pteridium aquilinum

Polystichum munitum

520
Common name Scientific name

FUNGI

Red-brown butt rot

White pine blister rust

Phaeolus schweinitzii

Cronartium ribicola

a

 Except as noted below, common and scientific names follow Collins 1990

for amphibians and reptiles, Jones and others 1986 for mammals, American

Ornithologists’ Union 1983 for birds, and Hitchcock and Cronquist 1973

plants.

b

Formerly D. ensatus; species’ revisions based on Good (1989).

c

 Also known as Pacific water shrew.

d

 Also known as dusky shrew.

e

 Proposed revision of Townsend’s chipmunk into four species including the

Siskiyou chipmunk (Tamias siskiyou) is not recognized.

f

 Also known as Columbian mouse.

g

 Inclusion of the red tree vole and white-footed vole in the genus Phenacomys

is not recognized.

521
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Appendix B-Abstracts of Poster Presentations

at the Symposium on “Old-Growth Douglas-Fir

Forests: Wildlife Communities, and Habitat

Relationships,” Held in Portland, Oregon, on

29-31 March 1989

Old-Growth Ecology and Management

Structure and Dynamics of Old-Growth Temperate

Rainforests in Coastal Alaska

Paul Alaback, USDA Forest Service

The cool temperate forests in coastal Alaska share many

species in common with both the montane and coastal floras

of the Pacific Northwest, but they have evolved under extreme conditions of cool temperatures and heavy year-round

rainfall. In south-central Alaska, hemlock-dominated forests

have been in the area for only a few thousand years and tend

to have more abundant spruce seedlings than do upland

forests of southeastern Alaska. Compared to the Northwest,

the growing season climate is much shorter, cooler, and

wetter. This climate leads to rapid soil formation, leaching,

and, in the absence of disturbance, bog formation. Larger

canopy openings are required to give the same degree of

release for hemlock seedlings. Huckleberry dominates the

understory of most upland sites because of high shade

tolerance and heavy seed production. The abundance of other

herbs is related to canopy disturbances-usually windthrow.

Dense growth of young trees after catastrophic disturbance

shades out understory plants for as much as 100 to 150

years. In the Pacific Northwest, by contrast, this lasts 50

years or less, perhaps because of greater radiation through

the forest canopy. Many wildlife species are thought to

depend on the open, structurally diverse old-growth forests.

The mosaic of forest and soil conditions and recent glacial

history makes the south coast of Alaska ideal for studies of

forest development in the face of rapid climatic change and

of landscape patterns in relation to wildlife populations.

Growth and Mortality in a 450-Year-Old Forest: Trends

During Four Decades for Douglas-fir, Western Hemlock,

and Associated Species

Dean S. DeBell, USDA Forest Service, and Jerry F. Franklin,

University of Washington

Resource managers are faced with decisions about the fate of

existing old-growth stands and the future role of old growth

in long-term plans for managing forests. Information on what

has happened since 1947 in a 450-year-old stand at Thornton

T. Munger Research Natural Area is useful background for

such decisions. Growth and mortality were measured at

6-year intervals from 1947 to 1983 in a 478-ha old-growth

stand in southwestern Washington. Principal tree species were

523
Douglas-fir, western hemlock, Pacific silver fir, western redcedar, and western white pine. They comprised 59,27,6,6,

and 1 percent, respectively, of the total cubic volume (372

m

3

) in 1947. Gross volume growth averaged 6.2 m3 per

hectare per year, and mortality averaged 6.0 m per hectare

per year. Net growth was therefore minimal, and total stand

volume remained nearly constant for 36 years. Changes are

occurring, however, in stand characteristics and species

importance. Douglas-fir, which accounted for only one-third

of the gross growth and nearly one-half of the mortality, is

losing dominance to western hemlock, which provided nearly

one-half the gross growth and only 28 percent of the mortality. Pacific silver fir increased in importance in the lower

canopy and was 60 percent of the ingrowth. Although such

changes may be relatively slow, they indicate that characteristics or functions of old-growth stands cannot be guaranteed

in perpetuity by simply preserving existing tracts. Information

is needed on how silvicultural practices might enhance development of desired attributes in younger, managed forests.

Patterns of Variation in Snag Populations on the Mt.

Hood National Forest

Christopher Topik, Nancy M. Diaz, and Timothy J. Brady,

USDA Forest Service

The functional roles of snags in the forest ecosystem depend

on their decay condition, size, and abundance. Effective

management of the snag resource requires knowledge of both

the general and habitat-specific patterns of variation in these

attributes. Survey data for plots from unmanaged westside

(western hemlock zone) and eastside (ponderosa pine,

Douglas-fir, and grand fir zones) stands greater than 60 years

old were compared, to characterize the patterns of variation

in snag populations in the Mount Hood National Forest.

Two general statements emerge from the data: the patterns of

variation in condition, size, and abundance of snag populations are, at most, only weakly correlated with successional

status; and the density of large snags (> 50 cm in d.b.h.,

> 15 m height) is essentially uniform throughout the habitats

sampled. The results suggest several habitat-specific observations as well: eastside stands contain substantially lower

snag densities than those west of the Cascade Crest: and although no trend in snag densities is discernible among decayclasses for westside stands, the distribution for eastside stands

is biased towards a greater representation of hard snags. The

data presented here suggest that snag-management policy

should remain flexible so that it can be modified in light of

habitat-specific characteristics of snag populations.

The Status of Old-Growth Forests in Six National Forests

of the Pacific Northwest

Peter H. Morrison, The Wilderness Society

The amount of old-growth forest remaining in the Pacific

Northwest has been the subject of increasing controversy. In

1988, The Wilderness Society undertook a project to determine the amount of old-growth forest remaining on six National Forests: Mount Baker-Snoqualmie, Olympic, Gifford

Pinchot, Mount Hood, Willamette, and Siskiyou. The study

used the interim definition established by the Old-Growth

Definition Task Group as a basis for the inventory. USDA

Forest Service timber inventories were analyzed to determine

forest characteristics. Inventory-plot status was updated using

Forest Service records and aerial overflights. The acreage of

old-growth forests and other mature forest types was determined by use of plot expansion factors. The results of the

study indicate that considerably less old-growth forest exists

than the acreage reported for old-growth forest in National

Forest management plans. The Wilderness Society analysis

indicated 461 538 ha of old-growth forest remain on the six

Forests studied. This compares to 1 029 555 ha reported in

Forest Service draft plans. The major reason for this discrepancy is the inclusion of all forest stands with trees over 53

cm in d.b.h. in the Forest Service estimate and the more

stringent criteria of the interim old-growth definition. Some

of the discrepancy is due to more recent updating of the

status of old-growth forests in The Wilderness Society study.

The landscape context of the remaining old growth was also

assessed in this study. Over 38 percent of the remaining old

growth exists in stands that are less than 32 ha or within 120

m of a road or clearcut. Old-growth fragmentation is particularly severe on some National Forests, such as the Gifford

Pinchot where over 68 percent of the remaining old growth

is within 120 m of a road or clearcut or in stands less than 32

ha. At current harvest rates, unfragmented old growth will

virtually disappear in a few years on such Forests.

Rain Chemistry and Buffering in an Old-Growth

Rainforest

Ted B. Thomas, Jonathon J. Rhodes, and Robert L. Edmonds,

University of Washington

The Olympic Peninsula and the forest lands of the Hoh River

Valley provide an ideal setting for characterizing wet deposition into clean ecosystems. Long-term ecosystem monitoring

provides an indication of environmental change from inputs

of atmospheric pollutants. An old-growth (650 years) forested

watershed at West Twin Creek in the Hoh Valley of Olympic

National Park was chosen by the National Acid Precipitation

Assessment Program for long-term intensive watershed study.

The objective of the program is to investigate rainfall chemistry and watershed processes that may affect surface-water

chemistry. The abundant rainfall (3500 mm) of the Hoh

524
Valley comes directly off the Pacific Ocean and is virtually

unimpacted by industrial pollution. Rainfall chemistry of the

Hoh Valley has been examined on a gradient from the Pacific

Ocean to 30 km inland. H+, S04

2

-, and NO3- ion concentration varied significantly over the 30-km transect; pH increased and S04

2

- and NO3- concentration decreased. The

long-term mean pH for precipitation in the Hoh Valley is

5.3. Interception of rainfall by old-growth forest canopies

and tree boles may acidify solutions to as low as pH = 4.0

(throughfall and stemflow ranged from 4.0-5.1). Stemflow

and throughfall solutions were more concentrated for H+,

P04

3

-, and cations than rain, Organic acids expressed as

dissolved organic carbon were most concentrated in stemflow and highly correlated with charge balance deficit. Soil

and bedrock weathering reactions buffered soil solutions and

appear to be the dominant controls on streamflow chemistry.

West Twin Creek surface waters have been consistently greater than pH = 7.0, and dominated by HC03

-

, S04

2-

, Ca2+,

and Mg2+ . Monitoring rain chemistry and characterizing ecosystem processes that affect ion transfers within old-growth

forests are important. These studies provide a baseline to

evaluate changes in forest health and surface water quality.

Northeast Cascades Recreation Strategy Research Needs

E.L. Burr, Mazama, Washington

The objective of wilderness management to maintain the natural forest renewal processes requires baseline data related to

fire, avalanche, grazing, browsing, endangered species, spraying, fuelwood gathering, logging, and interagency boundary

discontinuities. Research funding is too often narrowly focused and has been largely reactive, forced by crises with

western spruce budworms, mountain goats, mule deer, grizzly bears, ski development, and spotted owls. Research that

anticipates management needs by probability and priority is

necessary to minimize social-resource management conflicts.

The first research priority for the east side should be the

definition of old-growth forests, then the ecological effects of

fire and spotted owl management. Questions about grizzly

bears as an old-growth-dependent species need to be addressed. Court injunctions threaten to shackle forest management unless research becomes more relevent.

Forest Fragmentation and Landscape Ecology

A Comparison of Three Methods of Estimating Fractal

Dimension as a Measure of Fragmentation of

Old-Growth Habitat Using a GIS

Kenneth R. Dixon, Timothy A. Young, K. Lea Knutson,

James R. Eby, and Harriet L. Allen, Washington Department

of Wildlife

Landsat MSS data were used to classify spotted owl

old-growth habitat in western Washington. Habitat

fragmentation was measured using the fractal dimension of

Mandelbrot. Three different methods of calculation (length

of trail, relation of area to perimeter, and Korcak empirical

relation for islands) produced three different estimates. Each

method may have advantages over the others, depending on

scale and application.

Minimum Fragmentation Landscape Design

Miles Hemstrom, Steve Eubanks, and Karen Geary, USDA

Forest Service

The spatial placement of harvest units across a landscape

greatly influences the resulting kinds and qualities of stand

conditions present. Because the distribution of stand ageclasses developed by timber harvest influences harvest options beyond one rotation, the pattern imposed on the landscape during the initial rotation tends to be self perpetuating.

The staggered-setting approach, typically used to disperse

harvest impacts across the landscape, results in the maximum rate of fragmentation of remaining unharvested stands.

When the landscape is half cut over, the remaining patches

are the same size as the harvest units. Edge effects, which

extend into remaining stands for some distance, may render

all the remaining unharvested stands unsuitable as habitat for

interior species.

An alternative harvest regime, based on maintaining large

unharvested stands and associated travel corridors as long as

possible, was developed for an area of the Blue River Ranger

District. This minimum fragmentation alternative is based on

the same rate of decadal timber harvest as the staggered-setting approach and can be implemented given existing management regulations. Both alternatives were analyzed in a

geographic information system. Minimum fragmentation

designs are being considered for most new timber sales on

the Blue River Ranger District.

Monitoring Biodiversity With Permanent PlotsLandscape, Stand Structure, and Understory Species

Janet L. Ohmann and Charles L. Bolsinger, USDA Forest

Service

Permanent plots are useful for monitoring biological diversity and wildlife habitat across the forested landscape. Forest

Inventory and Analysis conducts a continuing multiresource

inventory of non-Federal forest lands in Washington, Oregon,

and California. Plots, established on a systematic grid, consist

of five sample points over about 20 ha. Live and standing

dead trees and understory vegetation are measured, and area

descriptors such as aspect, slope, elevation, and disturbance

history are recorded. Plot data in western Oregon indicate

that 82 percent of the non-Federal timberland is in a midsuccessional stage, and only 1 percent qualifies as old growth.

Estimates of old-growth area vary by definition: in western

Oregon, only 8 percent of stands over 100 years old meet an

525
ecological definition of old growth, while several stands

younger than 100 years qualify. Snag density increases with

forest succession, from 1 ha in very young stands to 7.1 in

old growth. In southwest Washington, permanent plots show

that within 10 to 25 years after clearcutting, Douglas-fir and

western hemlock make up a slightly larger portion of new

stands, western redcedar has been reduced markedly, and

other tree species have changed little. Average shrub cover

decreased slightly after logging, but number of shrub species

increased. Groundcover in forbs increased slightly after logging, but number of species nearly doubled, including several

pioneer species such as Canadian thistle, fireweed, and hedgenettle. In conclusion, Forest Inventory data show that intensive management of non-Federal timberland has reduced

structural diversity across the landscape as well as on specific

sites, but has increased diversity as measured by number of

plant species (although species proportions have changed).

These changes have implications for diversity of wildlife

communities as well as for long-term management for other

resources. This information should be useful to planners,

policy makers, and others concerned about the effects of land

management on both public and private lands.

Spotted Owl

Banding as a Management Tool: The Case of the

Northern Spotted Owl

Robin R. Bown and Joseph B. Lint, USDI Bureau of Land

Management

Since 1985, biologists have banded 216 adult, 37 subadult,

and 107 juvenile spotted owls with USDI Fish and Wildlife

Service colored leg bands on the Roseburg District of BLM.

Intensive efforts to observe and band birds each year allow

the biologists to define stable sites, detect changes in occupancy over time, and document movements of the nest areas.

By identifying individual owls, the biologists gather demographic information including site and mate fidelity, survival,

replacement, and movements. Birds banded as subadults or

juveniles provide age-specific information on survival,

reproduction, site fidelity, and movements. Of 116 adults

banded on 68 stable territories, 74 percent were observed

after 1 year, compared to 45 percent of 11 subadults. To

date, 28 banded birds have been replaced by new birds,

including three complete pairs. Three adult females have

moved from existing sites and paired with an established

male on another site. Six of the 69 juveniles banded in 1986

and 1987 have been encountered. Long-term banding

information will permit us to calculate the demographic

values for population analyses and viability assessment, as

well as to determine a relative measure of habitat quality

based on the occupation and reproductive history of sites.

Spotted Owl Monitoring in Washington and Oregon

Kathleen A. O’Halloran, A. Grant Gunderson, and Richard

S. Holthausen, USDA Forest Service

The goal of Pacific Northwest Region spotted owl habitat

management is to ensure the viability of spotted owls on

Forest Service lands as mandated by the regulations implementing the National Forest Management Act of 1976

(36 CFR 219). To determine if spotted owl populations in

spotted owl habitat areas (SOHAs) are stable, 11 Forests

conducted monitoring with standardized procedures. In 1987,

405 ha SOHAs were monitored. The regional average for

nonreserved SOHAs with any owls (singles and pairs) was

89 percent, and 80 percent of the reserved SOHAs had owls;

46 percent of the nonreserved SOHAs had pairs, and 30

percent of the reserved SOHAs had pairs. In 1988, SOHAs

were designated primarily on nonreserved lands, and size

varied from 405 to 1093 ha, depending on the physiographic

province. The 1988 regional average for SOHAs with any

owls was 91 percent, with 56 percent having pairs. To examine general population trends, random sample areas (RSAs)

were monitored for spotted owls. In 1987, 116 RSAs of 1822

ha were monitored; 73 percent had owls, with 26 percent

having owl pairs. Because of interpretation and implementation problems, RSAs were reduced to 405 ha. In 1988 also,

RSAs were established and monitored on reserved lands; 57

percent of the nonreserved RSAs had owls and 15 percent

had owl pairs. For reserved RSAs, 38 percent had owls and

17 percent had owl pairs. Monitoring of these areas will

continue on a stratified basis. Within 5 years, trend information from the SOHA and RSA monitoring will be evaluated

to determine if the Forest Service’s spotted owl management

strategy is working.

Ecology of Spotted Owls on the Olympic Peninsula,

Washington

Eric D. Forsman, Stan G. Sovern, and Duane Aubuchon,

USDA Forest Service

Twenty-one spotted owls were radio-tagged on the west side

of the Olympic Peninsula in 1987-88 and observed for periods ranging from 7 to 19 months. Home ranges of individuals

varied from 1367 to 13 475 ha. Home ranges expanded considerably during winter, with few individuals wandering

extensively into areas up to 22 km from the center of summer

activity. Color banding of the 21 radio-tagged owls and an

additional sample of 46 owls that were not radio-tagged

indicated an approximate adult survival rate of 0.94 during

the first year of the study, Reproduction during 1987 was

poor, with only 9 percent of the population producing young.

In 1988,36 percent of the population produced young. In a

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360 km2

 intensive study area on the west side of the peninsula, density of owls was 0.064 owls per km2

, which is less

than half the density reported in studies in Oregon and

northern California.

Spotted Owl Home Ranges in Five Landscapes

Scott P. Horton, Andrew B. Carey, Janice A. Reid, and

Robert B. Horn, USDA Forest Service

Spotted owl home ranges in the Pacific Northwest are typically large and primarily composed of late-successional

forests. Home-range size and composition may vary with

landscape composition and pattern, but few data are available

for such comparisons. We obtained over 7000 relocations on

members of 23 pairs of radio-tagged spotted owls, between 1

September 1987 and 31 August 1988, in five study areas in

southwestern Oregon. Pair ranges (medians) were 1567 and

2864 ha, respectively, in old-growth dominated (five pairs)

and mosaic-patterned (four pairs) Douglas-fir study areas;

571 and 1648 ha, respectively, in old-growth dominated

(three pairs) and mosaic (five pairs) mixed-conifer areas; and

1662 ha in a mosaic, mixed-evergreen (six pairs) area. Pair

ranges were composed of an average of 65 percent and 47

percent late-successional forest in homogeneous and mosaicpatterned landscapes, respectively. All but 1 of the 46 owls

selected late-successional stands for foraging, based on their

availability within the owls’ ranges. Other types of stands

were used either in proportion to their availability or avoided.

Both landscape composition and pattern appeared to influence

home-range sizes. Pair home ranges were smaller in the

mixed-conifer/evergreen forest zone than in the Douglas-fir

zone in landscapes of similar pattern. And pair home ranges

were smaller in homogeneous, old-growth dominated landscapes than in mosaic-patterned landscapes in similar forest

zones.

Temporal Variation in Spotted Owl Home Ranges

Kevin C. Peeler, Andrew B. Carey, J.H. Guetterman, and

Peter J. Loschl, USDA Forest Service

Spotted owls are known to have large home ranges, however

little is known of how these home ranges change over time.

Our objective is to report the seasonal, yearly, and cumulative

home ranges of four adult spotted owls that were followed for

about 2.5 years, Three male and 1 female spotted owls were

radio-tagged in the spring of 1986 in the southern Oregon

Coast Range. The owls were relocated every other night and

on one day during the 5-day work week, for an average of

377 fixes per owl over 2.5 years. We estimated home ranges

using the minimum convex polygon method for each individual for two or three breeding seasons (March through

August), two nonbreeding seasons (September through

February), two l-year periods (September through August),

and for the entire 2.5-year period. Breeding season home

ranges averaged 802 ha (n = 11, SE = 118). Nonbreeding season home ranges averaged 1566 ha (n = 8, SE = 305). Yearly

home ranges averaged 1715 ha (n = 8, SE = 283). The

2.5-year cumulative home range averaged 2,454 ha (n = 4,

SE = 554). Yearly home range size averaged 73 percent of

the 2.5-year cumulative home range (n = 8, SE = 7).

Although four individuals is a small sample, spotted owl

home ranges appear to increase over time.

Diet of Spotted Owls on the Olympic Peninsula,

Washington and the Roseburg District of the Bureau of

Land Management

Eric D. Forsman, Ivy Otto, and Andrew B. Carey, USDA

Forest Service

Between May 1987 and November 1988, 884 prey were

identified from pellets of spotted owls on the Olympic

Peninsula. The northern flying squirrel was 64 percent and

42 percent of the diet on the west side of the Peninsula,

respectively. Bushy-tailed woodrats and southern red-backed

voles were important prey on the east side of the Peninsula

but were rare in the diet of owls on the west side. Deer mice

were common in the diet on all study areas. Most predation

on snowshoe hares consisted of small juveniles captured

between April and September. In both 1987 and 1988, small

juvenile flying squirrels were largely absent from the diet

except during September-November. This suggested that

flying squirrels on the Peninsula produced only one litter per

year in July or August, and that the young squirrels began to

forage outside their natal nests in September. Diets of male

and female spotted owls did not differ significantly either in

terms of composition or mean prey size. On the Roseburg

Study Area, the diet of spotted owls in five different

geographic areas was compared. The main difference noted

among areas was that woodrats tended to predominate in

drier mixed-conifer and mixed-evergreen forest types, but

flying squirrels predominated in the diet in more humid

forests of Douglas-fir.

Roosting Habits of the Spotted Owl in the Oregon Coast

Range

Christopher C. Foster, Joseph A. Bums, and Andrew B.

Carey, USDA Forest Service

Radio-telemetry studies of the spotted owl have been conducted in the Oregon Coast Range since 1986. Site characteristics have been recorded for 772 diurnal roost locations

in three vegetation zones (western hemlock, mixed conifer,

and mixed evergreen) and five stand types (sapling, pole,

young, mature, and old-growth). Old-growth stands in all

vegetation zones were preferred as roost locations. Owls

selected for northeast-facing slopes and tended to roost on

the northeast side of the tree. Slope and perch aspects did not

vary significantly with season. Owls roosted predominantly

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in Douglas-fir. Roost heights were inversely related to

ambient temperature in winter. Overhead perch protection

averaged 75 percent. Old growth offers the northern spotted

owl the variety of roost conditions necessary to survive in a

varying climate.

Spotted Owl Prey Studies

Red Tree Voles in the Oregon Coast Ranges

Anne-Marie Gillesberg and Andrew B. Carey, USDA Forest

Service

Red tree voles are of interest because of their restricted

geographical distribution, preference for old growth, and as

an important food item for the spotted owl. To manage for

viable populations of the spotted owl, we need to describe

the ecology of its prey base in greater detail. Several

techniques have been used in an attempt to estimate the

abundance of red tree voles. Because nest surveys and

trapping efforts in the Oregon Cascades and Coast Range

proved largely unsuccessful, we explored an alternative

technique: searching felled trees. The Bureau of Land

Management assesses the value of timber in old-growth

stands by felling trees and examining them for defects (a

procedure known as 3P sampling). We examined 93 felled

trees from four 3P samples for evidence of red tree voles.

We found 117 pieces of nest material and described their

composition. Smaller nests usually consisted solely of resin

ducts and fecal material, but larger nests contained, in

addition to resin ducts, conifer needles, twigs, lichen, feces,

and other organic matter. Nests were found in hollow limbs

and broken tops, resting on branches or situated against the

boles of the sample trees. Nests were most often within the

lower one-third of the canopy, and larger trees (>100 cm in

d.b.h.) were preferred over smaller trees. We concluded that

searching a sample of felled trees, in conjunction with 3P

sampling (or thinnings), can be an effective method of

determining the relative abundance of red tree vole nests and

investigating potential dependency of this species on oldgrowth forests.

Woodrat Abundance in Forests of the Southern Oregon

Coast Range

G. Scott Center, Bruce R. Casler, and Andrew B. Carey,

USDA Forest Service

Abundance patterns of dusky-footed woodrats and bushytailed woodrats were described as part of a spotted owl preybase study in southwest Oregon. Exploratory live-trapping

was conducted in Douglas-fir and mixed-conifer forests in

seral stages ranging from recently cut to old growth (>200

years). Forty-five sites were sampled: six stages in mixedconifer forests and five stages in Douglas-fir forests. Abundance of both woodrat species was higher in mixed-conifer

forests than in Douglas-fir forests. Both species were locally

abundant in mixed-conifer old growth; only the bushy-tailed

woodrat occurred in Douglas-fir old growth. Both bushytailed woodrats and dusky-footed woodrats were abundant in

riparian zones. Dusky-footed woodrats were abundant in

sapling-pole, young (<80 years), and in disturbed portions of

old stands (>80 years) where dense regrowth occurred in the

mixed-conifer forests. The percentage of ground cover of

fine and coarse litter shows a significant positive relationship

to dusky-footed woodrat abundance. Dusky-footed woodrats

built shelters (“middens”) of coarse woody debris. Bushytailed woodrats appeared to use cavities in trees or crevices

in rocks, which might explain their association with old

growth (where snags are abundant), and riparian zones, where

rocks are exposed and deciduous trees provide additional

cavities.

Flying Squirrel Abundance in Young and Old-Growth

Forests of the Pacific Northwest

Brian L. Biswell and Andrew B. Carey, USDA Forest Service

Northern flying squirrel abundance was determined in stands

representative of the home ranges of spotted owls, particularly young and old-growth Douglas-fir forests in the Oregon

Coast Range and climax western hemlock and young westem hemlock and Douglas-fir forests on the Olympic Peninsula, WA. Live-trapping was conducted during 1985-88 in

18 Coast Range sites: 8 young, 3 mature, and 7 old-growth.

Fourteen Olympic Peninsula sites were trapped twice during

1987-88: 8 young and 6 climax. Trapping was conducted

during spring and fall in the Coast Range. Trapping was

conducted only in fall in the Olympics. Flying squirrel

densities in old stands in the Coast Range were 0.1 to 4.8

(averaging 1.8) animals per hectare and 0.3 to 2.0 (averaging

0.9) animals per hectare in young stands. Fall flying squirrel

densities on the Olympic Peninsula were 0.0 to 0.8 (averaging

0.3) animals per hectare in climax stands and 0.0 to 1.2

(averaging 0.4) animals per hectare in young stands. No significant difference in flying squirrel abundance was found

between young and climax forests on the Olympic Peninsula.

Densities were 2 times greater in old stands than in young

stands in the Coast Range. Young stands averaged 2.3-times

higher densities in the Coast Range than in the Olympics and

6-times higher densities in old-growth Douglas-fir than in

climax western hemlock.

Comparison of Small Mammal Abundance in Climax and

Young Forests of the Olympic Peninsula, Washington

Brian L. Biswell, John Boulanger, and Andrew B. Carey,

USDA Forest Service

Small mammal abundances in climax western hemlock stands

on the Olympic Peninsula of Washington were compared to

young stands as part of a spotted owl prey-base study. The

focus of the study was to determine the abundance patterns

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of deer mice and southern red-backed voles. An index to

abundance was obtained by using snap-trap transects in 14

stands. Trapping was conducted during the summers of 1987

and 1988. Sampled stands were classified as climax (250

years) or young (40-90 years and regenerated after logging

or wind storms). Transects consisted of two lines of 25 stations, 15 m apart. Two Museum Special snaptraps were

placed at each station and checked for 4 nights. Young and

climax stands were paired by geographic areas for analysis.

Capture rates between paired stands were compared by using

Wilcoxon’s signed-ranks test. Twelve species of small

mammals were caught, Trowbridge’s shrew, red-backed vole,

and deer mice were most abundant. The numbers of species

trapped were nearly equal between stand types. Deer mice

were more abundant (P = 0.058) in climax stands than in

young stands. The abundance of red-backed voles did not

differ between stand types (P = 0.46). Deer mice decreased

between 1987 and 1988, but red-backed voles increased (P =

0.003). Our results document the preference of deer mice for

climax forests, and the high variability of small mammal

abundance.

Increasing the Carrying Capacity of Second-Growth

Stands for Flying Squirrels With the Use of Nest Boxes

J.W. Witt, U.S. Department of the Interior, Bureau of Land

Management

Three experimental and three control areas were established

in second-growth Douglas-fir stands in western Oregon. Nest

boxes were installed in the experimental areas at a density

of about 2.8 boxes per hectare. The mean adult-trapping success and densities of adult northern flying squirrel were estimated using an arboreal live-trapping grid. The estimates

were compared between and within their respective areas,

and compared with similar estimates from an old-growth

grid established in the Coast Range. Nest-box use and occupancy were examined in the second-growth stands and in an

old-growth nest-box grid. The mean adult density in the

experimental grids, using the boundary-strip technique, was

0.80 squirrels per ha with a range of 0.29 to 1.54; the mean

density in the control areas was 0.39 squirrels per ha, with a

range of 0.08 to 0.81. Two-way analysis of variance with

replication (ANOVA Model I) indicated a significant difference (P c 0.005) between the treatment and the control, and

the density estimates between treatment areas was also significantly different (P < 0.0005). The mean density estimates

in spring from the experimental areas were not significantly

different from the old-growth mean, but a mean density estimate in the fall derived from two of the three experimental

areas indicated a significant difference between the second

growth and the old growth (P < 0.05), with the old growth

being larger. Although the response to the nest boxes was

variable, the results are consistent in that they demonstrate

that the carrying capacity of second-growth habitat can be

increased with the use of nest boxes.

Management implications of the study: if the target species is

the northern flying squirrel, then nest boxes are an appropriate and useful management tool; as to managing the

northern spotted owl-by contributing to its prey base, next

boxes are a supplemental tool but not a surrogate for old

growth.

Flying Squirrel Live-Trapping Success

Janelle G. Corn, U.S. Department of the Interior, Fish and

Wildlife Service, and Joseph Witt, U.S. Department of the

Interior, Bureau of Land Management

The northern flying squirrel is an important component of

old-growth forests in the Pacific Northwest. It is the primary

prey of the spotted owl and may be an important part of the

forest nutrient cycle. Variations in flying squirrel livetrapping success were examined in an old-growth Douglas-fir

forest stand in Oregon’s Coast Range to determine whether

microhabitat and trap placement affect capture rates. The

results of three trapping periods over 2 years were used to

identify a subset of trap sites with good or poor squirrel capture rates. Forest structural and compositional features were

measured within 5 m of each trap, and trap-placement features were described at each trap. Variables selected for

measurement were thought to relate to proximity to squirrel

foraging or resting areas, protection for squirrels from predators, and reduced invasion of traps by other small mammals.

Few measured variables were correlated with trap success.

The few correlations between trap-site features and capture

rates indicate that flying squirrel trap-success is not influenced by conditions existing at or very near the trap, and that

these features will not confound large-scale patterns of habitat use.

Marbled Murrelet

The Marbled Murrelet in Western Oregon: A Summary

of Current Knowledge

S. Kim Nelson, Oregon State University

The marbled murrelet is a small robin-sized seabird that inhabits near-shore coastal waters and inland older-aged (80

years) coniferous forests of the Pacific Northwest. Despite

this alcid’s frequent and widespread occurrence along the

Pacific coast, little is known about their inland distribution

and specific habitat preferences. A total of 14 nests of this

species have been recorded in its entire range; 9 on the

ground or in rock cavities in tundra habitats, and 5 on mosscovered branches or broken tops of large conifer trees in

529
forested habitats. In western Oregon, the marbled murrelet is

known to occur at 42 inland locations, up to 47 km from the

ocean; however, no nests have been found. These inland

sightings occurred primarily in mature (80-200 years) and

old-growth (>200 years) forests of the central Coast Range

that included large conifer trees with moss-covered branches.

Although exact use (nesting, roosting, or incidental) of these

inland sites has not been determined, older forests with specific vegetation characteristics may be required for marbled

murrelet nesting habitat. Conflict with timber management

practices may develop because some inland sites where

murrelets were located are scheduled for harvest within the

next 10 years.

Geographic Distribution of the Marbled Murrelet in

California at Inland Sites

Peter W.C. Paton and C. John Ralph, USDA Forest Service

The marbled mm-relet is an inhabitant of the coastal oldgrowth redwood forests of the northern half of California, yet

little is known of its ecology away from the ocean. During

the summer of 1988, we conducted a series of systematic

surveys to quantify the distribution of the murrelet at inland

sites from the Oregon border south to Monterey Bay. We

identified old-growth and mature forests as potential habitat

using remote-sensing techniques developed by Larry Fox of

Humboldt State University. We then conducted transects,

quantifying the relative abundance of murrelet activity, their

behavior, and various vegetative aspects of the stands. Transects consisted of 8 to 12 stations spaced 250 m to 1 km

apart, with each station surveyed for 10 minutes. The survey

period extended from 45 minutes before to 90 minutes after

sunrise. Each transect was surveyed at least twice during the

1988 field season. A total of 283 morning censuses were conducted on 127 transects, with murrelets detected on 53 percent (66) of the transects. In addition, stationary counts at

fixed stations were conducted on 37 mornings and 31 evenings. Eighty percent of the murrelet detections occurred

from 30 minutes before to 30 minutes after sunrise. Morning

censuses had 5 to 6 times more detections than evening censuses at the same fixed station during the same 24-hour

period. About 25 percent of the detections were visual observations, the rest were only heard. Flock size was small, single

birds and pairs accounted for 80 percent of all detections.

Bird distribution was patchy and restricted to the old-growth,

redwood-dominated forests of Del Norte, Humboldt, San

Mateo, and Santa Cruz Counties. No birds were detected in

Mendocino, Sonoma, or Marin Counties, with the exception

of one possible detection in Mendocino County. Areas with

the highest activity were the Redwood State and National

Parks within 25 km of the coast, although birds were detected

39 km inland at Grizzly Creek Redwoods State Park in

Humboldt County.

Forest Birds

Does Habitat Heterogeneity Matter to Breeding Birds in

Natural Douglas-fir Forests?

Andrew J. Hansen, Oregon State University, and Barry R.

Noon and Kathryn L. Purcell, USDA Forest Service

We studied the influence of fine-scale habitat heterogeneity

on bird community structure in a natural Douglas-fir forest to

determine if habitat specialists are less common here than in

other types of forests and if the size of microhabitat patches

influences their suitability as habitat. Vegetation and birds

were sampled in spring 1987 along 80-m wide, continuousbelt transects totaling 4 km in length. The data were analyzed

at scales of resolution of 800, 1600, 3200, and 6400 m2

.

Principal components analyses revealed that 2 of 10 bird

species (20 percent) used nonrandom subsets of the habitat in

the study area. This proportion of habitat “specialists” was

slightly less than the 29 percent found in a comparable study

in a northern hardwood forest. Also, 20 percent of our species

showed significantly different densities in forest gaps and in

patches of closed forest. The comparable figure from four

studies in other biomes is 16 to 45 percent.

Relative density of one bird species was positively associated

with increasing area of suitable microhabitats. A group of

species used large patches of unsuitable habitat less than they

used small patches of unsuitable habitat.

Our findings support the hypothesis that birds specializing on

specific microhabitats are generally less common in Douglasfir forests than in other types of forest. Nonetheless, microhabitat diversity and spatial patterning are important habitat

components for some birds in Douglas-fir forests.

The Effects of Forest Management on Cavity-Nesting

Birds in Northwestern Washington

Jill E. Zarnowitz, Oregon Department of Fish and Wildlife

I studied the population characteristics and nest-site preferences of cavity-nesting birds in the Olympic National Forest

of northwestern Washington. I characterized breeding populations in four different forest successional stages, from

clearcuts to old growth, where either high or low densities of

snags occurred. I described active cavity nests for 11 bird

species. Species richness, densities, and diversities of cavitynesting birds were greater in the plots with high snag densities (snag plots) than those with low snag densities (clean

plots). Cavity-nesting bird populations increased with increasing densities of snags. Active cavity-nesters were 5 times

more numerous on snag plots than clean plots. Hairy woodpeckers, a primary cavity-nester, selected western hemlock

snags for nest sites. In contrast, broken-topped Douglas-fir

530
snags were preferred by secondary cavity-nesters. The average diameter at breast height (d.b.h.) for active nest trees was

substantially greater than the mean d.b.h. for sampled snags

in the Forest. Low snag density appears to be a limiting factor

for breeding cavity-nesting populations. Management recommendations for cavity-nesting birds in the Forest are

discussed.

Significance of Old-Growth Forest for the Capercaillie

Pekka Helle and Harto Linden, Finnish Game and Fisheries

Research Institute

The capercaillie, a Palearctic grouse species, inhabits old

conifer stands, especially during winter and lekking season.

Its well-documented population decline in Fennoscandia

during past decades is associated with modem industrial

forestry.

Two studies from Finland dealing with habitat distribution

and stand characteristics of capercaillie leks are described. In

northern Finland close to the Arctic Circle, the lekking

stands did not differ from randomly selected stands in the

1950s. In the 1980s, when forestry had considerably altered

stand structure, lekking stands were older, larger, and had a

higher timber volume than random stands. Mean stand age

and timber volume significantly decreased when moving

away from the lek center, relatively evenly up to a distance

of 4 km, to which distance adjacent areas were assessed.

A southern Finnish study based on topographic maps showed

that the proportion of forest distinctly decreased with increasing distance from the lek center, up to a distance of 1 to 1.5

km. The data suggest that, in optimum habitats, a finegrained mosaic of habitat patches is advantageous, but in

marginal habitats, a coarse-grained habitat configuration is

better. The results are discussed in terms of forestry planning,

landscape ecology, and grain-size concepts.

Habitat Use of Cavity-Nesting Birds, Oregon Coast Range

S. Kim Nelson, Oregon State University

The availability of suitable standing dead trees or snags is

known to influence cavity-nesting bird populations. Current

snag-management practices on Federal lands may not be

providing the appropriate number or distribution of snags to

meet requirements of cavity-nesting birds. This study was

undertaken to describe the nest-tree and nest-site characteristics of cavity-nesting birds in unmanaged old-growth, mature,

and young forests, and to provide management recommendations. I found 277 active nests of 9 cavity-nesting species.

No differences were found among species in their nest-tree

or nest-site preferences. In general, all species preferred to

nest in Douglas-fir snags >50 cm in diameter (d.b.h.) and

>21 m in height. Large, tall snags were used significantly

more than available in all stand ages. Cavity-nesting birds

also selected hard snags with more bark and branches compared to available snags, and nested in snag patches with

dense midstory and understory cover. Current snag management policies include retaining or creating snags, and retaining live green-tree replacements in clearcuts. These snags

and trees will provide feeding habitat for all cavity-nesting

birds, but not nesting habitat for species tied to the forest

interior. Management practices should be modified to

provide snags in all stages of managed forests, and retain or

create snag habitat islands that include large (>50 cm in

d.b.h.), tall snags, large green-tree replacements, and intact

midstory and understory vegetation layers.

Comparative Foraging Ecology of Hammond’s and

Western Flycatchers in Northwestern California

Howard F. Sakai, Barry R. Noon, and C. John Ralph,

USDA Forest Service

Stand structure and vegetation composition of old Douglasfir forests in northern California and the Pacific Northwest

are constantly being subjected to modifications from natural

and human-caused disturbances. Effects of these habitat

modifications, and conversion of old to young forests, on

foraging site selection of breeding flycatchers has not been

quantified and is relatively unknown. Foraging behaviors of

Hammond’s and western flycatchers in different-aged

Douglas-fir stands were studied in northwestern California

during the breeding seasons of 1984-85. Habitat characteristics of foraging sites were compared within species across

stand ages and between species independent of stand age.

Tree-structure characteristics, visual obstruction of perches,

and foliage density at different heights best differentiated the

two species among different foraging sites. Comparisons

between species indicated that Hammond’s differ from

western flycatchers by perching and foraging higher, and

choosing taller and larger diameter trees. Use of foraging

sites varied within a species according to structural features

associated with the different-aged stands. Comparisons

within stand age-class showed that Hammond’s foraged on

fewer plant species than western flycatchers. Both species

generally did not select forage tree species according to their

availability in a stand.

Other Wildlife

Arthropods in Old Growth: Sensitive Environmental

Indicators

Andrew R. Moldenke and John D. Lattin, Oregon State

University

In all ecosystems, the species-rich arthropods regulate numerous critical biologic transformations and confer stability on

the total system. A forest ecosystem has structure, and

animal and plant species respond to this structure spatially

531
and seasonally. This structure changes geographically as well

as with succession. Invertebrates function in the food webs

that involve every plant and animal in the forest, Current research focuses on old growth of the H.J. Andrews Experimental Forest.

Soil arthropods play important roles in the breakdown of

litter, cycling of nutrients, and dispersal of microbial inocula.

Pacific Northwest old-growth soils typically maintain nearly

200 arthropod species per square foot at any time of the year

(5- to 8-year-old clearcuts have yet to recover to one-tenth

that diversity). These species-rich assemblages respond predictably to changes in slope face and succession with strong

turnover in individual species abundance. Soil arthropod

studies indicate that fundamental ecological processes (for

example, herbivore populations, parasitoids, microbial

grazers) are differentially keyed to advancing successional

seres. The composition of an individual series of samples can

thus predict the type of forest association they were collected

from. Thus, soil arthropods demonstrate promise to be used

as “biologic probes” of biodiversity, disturbance, and edge

effects caused by habitat fragmentation.

Ecology of Marten in the Pacific Northwest: Technique

Evaluation

Lawrence L.C. Jones, Leonard F. Ruggiero, and James K.

Swingle, USDA Forest Service

The marten, once widespread and abundant in boreal forests,

has been extirpated from much of its former range by trapping and habitat alteration. Although the marten is a Management Indicator Species on many National Forests in Oregon

and Washington, published studies on the ecology of marten

in the Pacific Northwest are virtually nonexistent. The purpose of this pilot study is twofold: to identify techniques that

might prove helpful to management biologists for monitoring

marten, and to determine which techniques will be both feasible and effective for use in future studies of marten ecology.

Four census techniques are currently being evaluated: hair

traps (winter), track plates (winter, fall), track counts in snow

(winter), and live-trapping (summer, fall). The success rate

of our modified hair traps and track plates was low: 2 and 6

detections for 410 and 286 sample-nights, respectively, during

the winter! Live-trapping does not appear to be a feasible

technique in our study area because only one marten was captured for every 221 corrected trap-nights. Snow track counts

yielded 41 sets of tracks or runways in 30 search-days, making this the most successful and cost-effective technique to

determine patterns of abundance for marten in the Pacific

Northwest.

Logging on Old-Growth Mule Deer Winter Range

Harold M. Armleder and Rick J. Dawson, Ministry of Forests,

British Columbia

Douglas-fir is an important component of the timber supply

in the Cariboo Forest Region of British Columbia. A significant amount of old-growth Douglas-fir is located on mule

deer winter range. Douglas-fir is an essential component of

winter habitat providing deer with food and shelter. Common

types of logging do not preserve these winter-range values.

Eight years of research on mule deer habitat relationships has

resulted in the development of an integrated management

system designed especially for winter range. Managers apply

the system by using procedures for assessing the current status of a winter range and determining prescriptions that will

maintain habitat values. A major component of the system is

a specialized method of low-volume selective logging that

involves removing only about 20 percent of the merchantable

volume of a stand. Trained loggers select trees to cut according to criteria developed especially for winter range. The

objective is to maintain and promote a healthy stand of trees

in all size- and age-classes, including a significant component

of large, old trees. Two user-oriented publications that aid in

implementing the system are highlighted.

Historic, Present, and Future Abundance of Terrestrial

Vertebrates in Northwestern California

Martin G. Raphael and Bruce G. Marcot, USDA Forest

Service

Douglas-fir forest covers over 1 million ha in northern California and has been heavily harvested over the last 30 years.

Because of logging, old-growth stands have been reduced to

50 percent of their historic acreage and may ultimately be

reduced to 10 to 15 percent. Based on extensive field surveys,

we estimated the relative abundance of 115 terrestrial vertebrate species in clearcuts, young forest stands, and mature

stands. We also estimated historic, present, and future acreages of these seral stages. We then estimated relative population sizes of each vertebrate species at each period to predict long-term effects of timber harvest on populations. We

project that future populations of 17 species will have declined >50 percent from historic populations; an equal number will likely increase by >50 percent. Species associated

with early-seral vegetation are currently at peak abundance

and will decline, but species most strongly associated with

mature forest have declined and will continue to do so. None

of the species will be pushed to extinction, unless fragmentation effects are so severe that patch sizes of suitable habitat

are too small and too widely dispersed to support viable

populations.

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Another trend is increased abundance among species of

southern affinity and decreased abundance among species of

boreal affinity. Increasers tend to be widespread and associated with a wide variety of habitats; decreasers tend to

occupy restricted ranges and a narrow range of habitats.

Therefore, even though numbers of increasers and decreasers

are equal, the effects of old-growth reduction are not neutral.

Occurrence of Old-Growth-Related Wildlife Species in

Non-Old-Growth Habitats

Steven J. Kerns, Wildland Resource Managers, Round

Mountain, California

During the summer of 1988, I studied the occurrence of three

old-growth-related wildlife species-spotted owls, marbled

murrelets, and fishers-in habitats generally considered

unsuitable for the species. The study was conducted on lands

of the Pacific Lumber Company in Northern California’s

Humboldt County. Owls were surveyed on 24 transects using

methods developed by the USDA Forest Service. Fishers

were surveyed with sooted track stations placed in different

timber types and by monitoring tracks in the dust of logging

roads, skid trails, and landings. Marbled mm-relets were

surveyed in five major drainages on Pacific Lumber property

by sampling techniques developed by the Pacific Seabird

Group and the USDA Forest Service. Twenty-two owls were

located on 16 transects. Owls were found in 40 vegetative

complexes, of which 9 (22 percent) can be classified as old

growth. The remainder (78 percent) may be classified as

either second-growth stands or residual stands (stands containing scattered large trees and smaller second-growth trees).

Coastal weather influence and timber-growing conditions may

account for the presence of owls in a diversity of habitat

types not suitable in other regions of their range. Marbled

murrelets were found in four of the five drainages I sampled.

The common factor associated with the presence of murrelets

was large conifers (2127 cm in d.b.h.) comprising at least 25

25 percent of the overstory. Only one fisher was located-in

a second-growth redwood-Douglas-fir stand containing trees

100 cm in d.b.h.

Old-Growth Forest Management (Slide Presentation)

Barbara A. Middleton, Oregon State University

The objective of this presentation is to define old-growth

forests and describe the benefits and values derived from

them, their role in the natural system, and why their

management is so controversial. The 15-minute slide-tape

program attempts to build common ground to discuss a

critical planning and management issue in the Pacific

Northwest. With a look at the results of current ecological

research, the four structural features (snags, large live trees,

fallen trees, and multilayered forest) are discussed.

Perspectives, attitudes, values, and benefits derived from

them are discussed to share differing management objectives.

Finally, the importance of citizen input in the planning

process is highlighted.

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The Forest Service of the U.S. Department of

Agriculture is dedicated to the principle of multiple

use management of the Nation’s forest resources

for sustained yields of wood, water, forage, wildlife,

and recreation. Through forestry research,

cooperation with the States and private forest

owners, and management of the National Forests

and National Grasslands, it strives-as directed by

Congress-to provide increasingly greater service

to a growing Nation.

The U.S. Department of Agriculture is an Equal

Opportunity Employer. Applicants for all Department

programs will be given equal consideration without

regard to age, race, color, sex, religion, or national

origin.

Pacific Northwest Research Station

319 S.W. Pine St.

P.O. Box 3890

Portland, Oregon 97208-3890