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BIRD COMMUNITIES IN MIXED CONIFER FORESTS
OF THE INTERIOR NORTHWEST
H. Reed Sanderson
Range Scientist
Pacific Northwest Forest and Range Experiment
Station, USDA Forest Service, La Grande, Oregon
Evelyn L. Bull
Wildlife Biologist
Pacific Northwest Forest and Range Experiment
Station, USDA Forest Service, La Grande, Oregon
Paul J. Edgerton
Wildlife Biologist
Pacific Northwest Forest and Range Experiment
Station, USDA Forest Service, La Grande, Oregon
ABSTRACT
Forest management practices adjust the direction and pace
of plant succession. The species composition and st~ucture
are altered, and, in turn, the avian species. Forest
management must include wildlife as an integral part of the
management decision. A wildlife biologist must provide sound
biological alternatives for the land manager's consideration.
We present a discussion of ecological concepts that ~ildlife
biologists can use to predict the response of bird alterations
in the interior Northwest mixed conifer forest type.
KEYWORDS:
habitat.
mixed conifer forest, silviculture, birds, nongame
Society's demand for products is the primary driving force for the
management of our natural resources. Whatever that management is, it also
affects wildlife populations whenever habitat is altered. As the human population
increases in number and affluence, the demand for products will also increase
(Maser 1979). Although timber harvest, livestock grazing, and wildlife harvest
provide products, they also " ••• stir man's economic interest and, in the longer
term, protective interest" (Maser and Thomas 1978:2). Economic demands must now
be balanced by ecological consequences because of such laws as the Environmental
224
Policy Act, Federal Land Policy and Management Act, and the Forest Policy Act.
Specifically, a land manager is to be held responsible for the consequences of
his decisions and their resulting impact on the resource, land, and environment.
A wildlife biologist's role is to provide a land manager with a set of management
alternatives and their respective consequences to habitats and attendant wildlife
species. The responsibility of wildlife biologists is to provide biologically
sound data. We can no longer shirk our responsibility with the too often heard
wildlife biologists' rationalization that, "We do not have enough information."
Our objective is to provide wildlife biologists with some ecological concepts
to assist in predicting the generalized responses of both plant communities and
birds to habitat alterations in the interior Northwest mixed conifer forests.
VEGETATION
The interior Northwest is characterized by a wide range of physical features
that create a variety of habitats for different kinds of biotic communities.
Diversity in these communities results not only from physiography, soils, and
climate, but also from fire, insects, disease, and management activities such
as timber harvesting and livestock grazing.
Foresters and ecologists have studied the vegetation of the mixed conifer
forests that occupy about 10.5 million hectares of the interior Northwest.
Kuchler (1964) broadly mapped this area as Douglas-fir (Pseudotsuga menziesii)-11
in the northern Rocky Mountains and Washington, cedar-hemlock-pine (Thuja-TsugaPinus) forests in the northern Rocky Mountains, and grand fir (Abies grandis)
- Douglas-fir forests in central Idaho, eastern Oregon, and southeastern
Washington. Regional ecologists have refined these forest descripti_ons to provide
an ecologically based system of land stratification for use by local resource
planners and land managers. They include Franklin and Dyrness (1973), Oregon
and Washington; Hall (1973), the Blue Mountains of eastern Oregon and southeastern
Washingon; Daubenmire and Daubenmire (1968), northern Idaho and adjacent
Washington; Steele et al. (Being prepared), central Idaho; and Pfister et al.
(1977) for Montana.
Interior Northwest coniferous forests occur along a predictable environmental
gradient. Climax Douglas-fir associations are usually found at mid-elevations
where they intergrade with the upper limits of more xeric ponderosa pine (Pinus
ponderosa) forests. In some areas, such as north-central Washington and the east
slopes of the northern Rocky Mountains, however, climax ponderosa pine may be
absent and Douglas-fir forests may border grasslands or shrub-steppe vegetation.
In Idaho and the Blue Mountains of Oregon and Washington, climax Douglas-fir
forest is less common. Instead, it is an important component of mixed conifer
communities transitional from ponderosa pine to grand fir. Douglas-fir and grand
fir generally dominate climax stands, but Engelmann spruce (Picea engelmannii) may
be locally abundant on moist sites, and subalpine fir (Abies lasiocarpa) becomes
an important component at higher elevations.
Fire has played a major role in determining the composition and stucture
of mixed conifer forests. Ponderosa pine, lodgepole pine (Pinus contorta),
western white pine (Pinus monticola), or western larch (Larix occidentalis)
dominate seral stands because they are better adapted to severe disturbance,
especially fire, than are the climax species. Ponderosa pine or lodgepole pine
1/
Nomenclature follows that of Garrison et al. (1976).
225
may persist on harsher sites as a fire climax. On the other hand, grand fir and
Douglas~fir regenerate abundantly in either mature, undisturbed stands, or seral
stands. In the latter situation, they gradually assume dominance as the stand
develops.
Composition and structure of the associated understory vegetation is diverse
and depends on interactions of site, plant community, fire, and forest management
activities. On drier sites dominated by Douglas-fir or mixed Douglas-fir and
ponderosa pine, grasses mixed with scattered low shrubs and forbs characterize
the understory. Dense, multilayered understories of grasses, sedges, forbs, and
tall shrubs occur on moist sites where Douglas-fir dominates the overstory. Some
characteristic species are pinegrass (Calamagrostis rubescens), elk sedge (Carex
geyeri), arnica (Arnica spp.), ninebark (Physocarpus malvaceus), and
snowberry (Symphoricarpos albus). The understory of mature or old-growth mixed
conifer forest dominated by grand fir is often characterized by low growing plants
such as American twinflower (Linnaea borealis), queencup beadlily (Clintonia
uniflora), and princespine (Chimaphila spp.). Wild rose (Rosa spp.),
huckleberries (Vaccinium spp.), yew (Taxus sp.), and other shrubs are abundant
in some communities.
Fire can be an important influence in understory development. Intense
heat generated by either wildfire or prescribed burns can destroy understory
vegetation and favor the germination and establishment of seral shrubs. Dense
fields of shrubs, such as snowbrush ceanothus (Ceanothus velutinus) and ninebark
may dominate disturbed Douglas-fir sites for 25 years or longer while the seral
forest develops. On the other hand, periodic, light underburning once maintained
open stands of Douglas-fir mixed with ponderosa pine (Hall 1977).
BIRDS OF THE _MIXED CONIFER FOREST
More than 90 species of birds use the mixed conifer forests in the interior
Northwest (Thomas 1979, Sundstrom 1978). None of these birds, however, restrict
their feeding and reproductive activities to a single forest type or to a
particular tree species. Because interior Northwest forests tend to be a mosaic
of forest types instead of large continuous blocks, management objectives
generally are not restricted to a particular forest type. Also, birds apparently
respond more to vegetative structure than they do to plant species composition per
se (Verner 1975). Consequently, management of bird communities should not be
considered by forest type, but rather by the overall impact of management on
forest structure.
Vegetative structure can be broadly equated to forest succession. As
succession progresses, plant species diversity and biomass increase; vegetative
structure becomes more complex, which in turn, creates more available niches that
result in increased bird species diversity (Meslow 1978) (Fig. 1). We have
characterized the mixed conifer forest type with six successional stages, and
have listed the birds that feed or reproduce in each successional stage
(Appendix 1).
Although we may not have all the specific information about habitat
requirements for all birds, we can fairly well predict the impact of various
management schemes on vegetative structure and plant succession and, consequently,
on bird species. Forest managers may wish to maintain as many naturally
occurring habitats as possible so future generations can have the same management
options we have today (Balda 1976). "Wildlife Habitats in Managed Forests"
226
(Thomas 1979) and "A Holistic Approach to Wildlife and Fish Habitat Management"
(Sundstrom 1978) are two publications that can be used to predict impacts of
forest management decisions on birds.
80
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Feeding
Reproduction
£1)
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20
GrassShrubForb
Seed I i ng
PoleSapling
Young
Mature
Old
Growth
FOREST SUCCESSIONAL STAGES
Figure 1.--Enumeration of bird species orientation to forest successional stages
in the mixed conifer forest type of the interior Northwest.
FOREST MANAGEMENT
Forest management is the dominant land management activity in the interior
Northwest forests. Forest management is bird habitat management and can achieve
habitat management goals with attentive planning and execution (Thomas 1979).
A forest manager is limited in the selection of silvicultural options because
of stand conditions due to past logging practices, insect and disease problems,
control of fire, and so on. Therefore the selection of a silvicultural system
227
must be made on a stand by stand basis. Generally healthy, mixed-aged stands
are suitable for uneven-aged management, but care must be taken to prevent a shift
in species composition, especially in the mixed conifer forests of eastern Oregon
and Washington. But, an even-age management system is usually recommended to
control dwarf mistletoe (Arceuthobium spp.) and western spruce budworm
(Choristoneura occidentalis Freeman), that are prevalent throughout the area
(Seidel 1973, Wellner and Ryker 1973). Open stands favor larch and pine, and
closed stands favor the shade tolerant firs. Douglas-fir is a shade requiring
species in the interior Northwest, and seedling establishment is best in partial
shade, but growth is best in full sunlight (Seidel 1973). Because each
silvicultural system has a specific impact on habitats, wildlife biologists must
have a basic understanding of these systems to predict the consequences of their
application.
There are four generalized forest management systems that adjust the
direction and pace of plant succession and, in turn, determine the avian species
associated with the various successsional stages.
1.
Even-aged management produces a monoculture of trees approximately the
same size and height. A stand has an identified establishment period,
and the entire stand is generally removed at maturity (Franklin and
DeBell 1973, USDA Forest Service 1973).
Even-aged management reduces vertical vegetative complexity and
results in a decrease in bird species diversity (MacArthur and MacArthur
1961).
Horizontal vegetative complexity (spacing) is increased by creating
different successional stages between the various stands or cutting
units. Edges are also created where different successional stages meet,
thereby enhancing bird species richness (Thomas et al. 1978).
All guilds (a group of species that use the habitat in a similar
way; Root 1967) could be represented through several even-aged stands,
but this depends on the successional stages present within a particular
time.
2.
In contrast, uneven-aged management develops vertical vegetative
complexity, but eliminates horizontal complexity by harvesting only
mature trees, by not cutting the entire stand, and by maintaining trees
in a variety of size classes (Franklin 1977, Hann and Bare 1979).
Edges and early successional stages are minimized, as well as,
the characteristics of old-growth stands. Bird species characteristic
of the related plant communities would also be reduced. For example,
aerial-searchers and ground-brush foragers would decrease, while bark
and foliage gleaners and drillers would increase.
3.
Intensive forest management shortens early successional stages and
eliminates the final stages by emphasizing stand regeneration, growth,
and harvest (Edgerton and Thomas 1978). Silviculture practices may
include brush control, tree planting, fertilization, and thinning--all
of which tend to accelerate tree establishment and growth and reduce
plant species diversity and structural complexity.
228
Intensive forest management potentially decreases bird species
diversity. Succession is accelerated; maturity is brief.
Harvest of climax old-growth stands eliminates the associated
specialized bird species, such as the brown creeper (Certhia
familiaris), pileated woodpecker (Dryocopus pileatus), and great gray
owl (Strix nebulosa).
4.
Salvage and sanitation logging and debris disposal activities remove
snag recruits and snags and reduce the amount of dead and downed woody
material that provides feeding and nesting sites for drilling and barkgleaning guilds (Maser et al. 1979, Thomas et al. 1979). Snags also
are needed for nest sites for a wide variety of birds. In addition,
snags and broken-topped trees are needed for nesting and perching sites
for such large birds as eagles (Aquila and Haliaeetus spp.) and
osprey (Pandion haliaetus).
The main management tool available to a wildlife biologist is to direct the
size, shape, location, and timing of silviculture practices.
Size of a treatment area has a direct relationship to the number of species
present (Galli et al. 1976). Thomas et al. (1978) estimated that bird species
richness is optimized at about 34 ha in the Blue Mountains. Such area figures
must be applied with caution, however, because they tend to become policy. Verner
(1975) suggested that a better approach would be to use the territory size of
large raptors, such as hawks and owls, because they could also serve as indicators
of the bird population vitality.
The shape of an area is related to the amount of edge produced--the more
irregular the shape, the greater the edge. Irregular shapes are also more
pleasing to a viewer and provide a more natural condition.
Location of a treatment area refers to its relationship to other forest
communities. Edges differ in their degree of contrast. For example, a sapling
stage against pole stage has a low degree of contrast, whereas a grass stage
against a mature forest stage has a high degree of contrast. Considering six
generalized successional stages, there is a possible combination of 15 edges,
all with a varying degree of contrast (Thomas et al. 1978). The juxtaposition
of various treatments can be used to achieve habitat diversity.
The final variable is time--time in relation to season of the year, and time
in relation to rotation age or number of years from tree establishment to
cutting. For example, timing of a prescribed burn may be critical to ground and
shrub nesting birds if it is done in the spring of the year, but the distribution
of silvicultural practices over years and ages of a stand affects both the pace
and direction of succession.
In summary, we paraphrase a portion of The Wildlife Society's position
statement on "Wildlife Needs in Forest Management" as adopted on March 24, 1979:
Forest management practices alter species
composition and structure of plant communities and
thereby affect attendant wildlife. Wildlife species
may increase, or decline, or be unaffected. Species
with a narrow range of tolerance for habitat change may
require special consideration.
229
Forest and wildlife management objectives can be
coordinated by maintaining diversity of plant species,
age classes, and stand densities; by retaining snags
and dead and down woody materials; and by varying the
size, shape, age, and juxtaposition of stands.
Management plans must be flexible but also must be
specific enough to meet local conditions. Management
practices must be prescribed according to site
conditions, plant and animal species involved,
successional relationships, and such local factors that
ensure a diversity of wildlife species.
Wildlife should be an intentional product of forest
management. It is a wildlife biologist's responsibility
to provide a manager with a set of alternatives. It
is a land manager's responsibility to review the
consequences of these alternatives and the trade-offs
on wildlife and their habitats (The Wildlife Society
1979).
We submit that this is our professional charge.
LITERATURE CITED
American Ornithologists' Union.
1957. Check-list of North American birds.
Inc., Baltimore, MD.
5th ed.
691 p.
Port City Press,
American Ornithologists' Union.
1973a. Thirty-second supplement to the American Ornithologists' Union
check-list of North American birds. Auk 90(2):411-419.
American Ornithologists' Union.
1973b. Corrections and additions to the "Thirty-second supplement to the
American Ornithologists' Union check-list of North American birds." Auk
90(4):887.
American Ornithologists' Union.
1976. Thirty-third supplement to the American Ornithologists' Union check-list
of North American birds. Auk 90(4):875-879.
Balda, Russell P.
1976. Vegetation structure and breeding bird diversity. In Symposium on
management of forest and range habitats for nongame birdS: Dixie R. Smith,
Tech. Coord. p. 59-80. USDA For. Serv. Gen. Tech. Rep. W0-1. U.S. Gov.
Print. Off., Wash., D.C.
Daubenmire, R., and Jean B. Daubenmire.
1968. Forest vegetation of eastern Washington and northern Idaho.
Agric. Exp. Stn. Tech. Bull. 60. 104 p.
Wash.
Edgerton, Paul J., and Jack Ward Thomas.
1978. Silviculture options and habitat values in coniferous forests. In
Proceedings of Workshop on nongame bird habitat management in the coniferous
forests of the western United States. Richard M. DeGraaf, Tech. Coord.
p. 56-65. USDA For. Serv. Gen. Tech. Rep. PNW-64. Pac. Northwest For. and
Range Exp. Stn., Portland, Oreg.
230
Franklin, Jerry F.
1977. Effects of uneven-aged management on species composition. In Uneven-aged
silviculture and management in the western United States. Proc. In-Service
Workshop, Oct. 19-21, 1976. Redding, Calif.
Franklin, Jerry F., and DeanS. DeBell.
1973. Effects of various harvesting on forest regeneration. In Even-aged
management. Richard K. Hermann and Denis P. Lavender, Compilers and Editors.
Pap. 848, p. 29-57. Sch. For., Oreg. State Univ., Corvallis.
Franklin, Jerry F., and C. T. Dyrness.
1973. Natural vegetation of Oregon and Washington. USDA For. Serv. Gen. Tech.
Rep. PNW-8. 417 p. Pac. Northwest For. and Range Exp. Stn., Portland, .Oreg.
Galli, Anne E., Charles F. Leek, and RichardT. T. Forman.
1976. Avian distribution patterns in forest islands of different sizes in
central New Jersey. Auk 93(2):356-364.
Garrison, G. A., J. M. Skovlin, C. E. Poulton, and A. H. Winward.
1976. Northwest plant names and symbols for ecosystem inventory and analysis.
4th ed. USDA For. Serv. Gen. Tech. Rep. PNW-46. 263 p. Pac. Northwest For.
and Range Exp. Stn., Portland, Oreg.
Hall, Frederick C.
1973. Plant communities of the Blue Mountains in eastern Oregon and Washington.
USDA For. Serv., Reg. 6 Area Guide 3-1. 62 p. Portland, Oreg.
Hall, Frederick C.
1977. Ecology of natural underburning in the Blue Mountains of Oregon.
For. Serv. Reg. 6, R6-ECOL-79-001. 11 p. Portland, Oreg.
USDA
Hann, David W., and B. Bruce Bare.
1979. Uneven-aged forest management: state of the art (or science?). USDA
For. Serv. Gen. Tech. Rep. INT-58. 18 p. Intermt. For. and Range Exp. Stn.,
Ogden, Utah.
Kuchler, A. W.
1964. Potential natural vegetation of the conterminous United States.
Geogr. Soc. Spec. Pub!. 36. 116 p. New York.
Am.
MacArthur, Robert H., and John W. MacArthur.
1961. On bird species diversity. Ecology 45(3):594-598.
Maser, Chris.
1979. Holistic management--can we achieve it?
Portland, Oreg. (abstract)
30th Conf. Wild!. Soc.,
Maser, Chris, Ralph G. Anderson, Kermit Cromack, Jr., Jerry T. Williams, and
Robert E. Martin.
1979. Dead and down woody material. In Wildlife habitats in managed forests-the Blue Mountains of Oregon and Washington. Jack Ward Thomas, Tech. Ed.
p. 78-95. USDA For. Serv. Agric. Handb. 553. U.S. Gov. Print. Off., Wash.,
D.C.
231
Maser, Chris, and Jack Ward Thomas.
1978. Ecosystems, habitats, wildlife, and management. In Proceedings of
Workshop on nongame bird habitat management in the coniferous forests of the
western United States. Richard M. DeGraaf, Tech. Coord. p. 1-4. USDA For.
Serv. Gen. Tech. Rep. PNW-64. Pac. Northwest For. and Range Exp. Stn.,
Portland, Oreg.
Meslow, E. Charles.
1978. The relationship of birds to habitat, plant communities, and
successional stages. In Proceedings of Workshop on nongame bird habitat
management in the coniferous forests of the western United States.
Richard M. DeGraaf, Tech. Coord. p. 12-18. USDA For. Serv. Gen. Tech. Rep.
PNW-64. Pac. Northwest For. and Range Exp. Stn., Portland, Oreg.
Pfister, Robert D., Bernard L. Kovalchik, Stephen F. Arno, and Richard C. Presby.
1977. Forest habitat types of Montana. USDA For. Serv. Gen. Tech. Rep. INT-34.
174 p. Intermt. For. and Range Exp. Stn., Ogden, Utah.
Root, Richard B.
1967. The niche exploitation pattern of the blue-gray gnatcatcher.
Monogr. 37(4):317-350.
Ecol.
Seidel, K. W.
1973. Mixed pine-fir of eastern Oregon and Washington. In Silvicultural
systems for major forest types of the United States. p-.-15-17. USDA For.
Serv. Agric. Handb. 445. U.S. Gov. Print. Off., Wash., D.C.
Steele, Robert, Robert D. Pfister, Russell A. Ryker, and Jay A. Kitams.
Being prepared. Forest habitat types of central Idaho. U.S. Dep. Agric. For.
Serv. Intermt. For. and Range Exp. Stn., Odgen, Utah.
Sundstrom, Charles.
1978. A holistic approach to wildlife and fish habitat management.
Beaverhead National Forest, Dillon, Mont.
135 ?·
The Wildlife Society.
1979. Position statement of The Wildlife Society on recognition of wildlife
needs in forest management. Oreg. Chap. Newsletter, Nov. 1979. 8 p.
Thomas, Jack Ward. (Tech. Ed.).
1979. Wildlife habitats in managed forests--the Blue Mountains of Oregon and
Washington. USDA For. Serv. Agric. Handb. 553. 512 p. U.S. Gov. Print.
Off., Wash., D.C.
Thomas, Jack Ward, Ralph G. Anderson, Chris Maser, and Evelyn Bull.
1979. Snags. In Wildlife habitats in managed forests--the Blue Mountains of
Oregon and Washington. Jack Ward Thomas, Tech. Ed. p. 60-77. USDA ~or.
Serv. Agric. Handb. 553. U.S. Gov. Print. Off., Wash., D.C.
Thomas, Jack Ward, Chris Maser, and Jon E. Rodiek.
1978. Edges--their interspersion, resulting diversity, and its measurement.
In Proceedings of Workshop on nongame bird habitat management in the
coniferous forests of the western United States. Richard M. DeGraaf, Tech.
Coord. p. 91-100. USDA For. Serv. Gen. Tech. Rep. PNW-64. Pac. Northwest
For. and Range Exp. Stn., Portland, Oreg.
232
United States Department of Agriculture, Forest Service.
1973. Silvicultural systems for major forest types of the United States.
Dep. Agric. Handb. 445. 114 p. U.S. Gov. Print. Off., Wash., D.C.
U.S.
Verner, Jared.
1975. Avian behavior and habitat management. In Symposium on management of
forest and range habitats for nongame birds. Dixie R. Smith, Tech. Coord.
p. 39-58. USDA For. Serv. Gen. Tech. Rep. W0-1. U.S. Gov. Print. Off.,
Wash., D.C.
Wellner, Charles A., and Russell A. Ryker.
1973. Ponderosa pine and Rocky Mountain Douglas-fir. In Silvicultural systems
for major forest types of the United States. p. 35-37: USDA For. Serv.
Agric. Handb. 445. U.S. Gov. Print. Off., Wash., D.C.
Appendix 1.--Bird species feeding (F) or reproducing (R) in the mixed conifer
forest successional stages of the interior Northwest
Forest Successional Stage
Species
1/
Wood duckAix sponsa
Barrow's goldeneye
Bucephala islandica
Bufflehead
Bucephala albeola
Harlequin duck
Histrionicus histrionicus
Hooded merganser
Lophodytes cucullatus
Turkey vulture
Cathartes aura
Goshawk
Accipiter gentilis
Sharp-shinned hawk
Accipiter striatus
Cooper's hawk
Accipiter cooperii
Red-tailed hawk
Buteo jamaicensis
Golden eagle
Aquila chrysaetos
Bald eagle
Haliaeetus leucocephalus
Osprey
Pandion haliaetus
Grassforb
Shrubseeding
Polesapling
Young
Mature
Oldgrowth
F
F
F
F
RF
RF
F
F
F
F
RF
RF
F
F
F
F
RF
RF
F
F
F
RF
RF
RF
F
F
F
F
RF
RF
RF
RF
RF
F
F
RF
RF
F
RF
RF
RF
F
F
F
F
RF
RF
F
F
F
F
RF
RF
RF
RF
RF
RF
RF
RF
RF
·.
F
F
F
F
F
F
1/
F
F
F
F
Nomenclature follows that of American Ornithologists' Union (1957,
1973a, 1973b, 1976).
233
~:
~
.
Appendix 1.--Continued.
Forest Successional Stage
Species
Peregrine falcon
Falco peregrinus
Merlin
Falco columbarius
American kestrel
Falco sparverius
Blue grouse
Dendragapus obscurus
Franklin's grouse
Canachites canadensis
Ruffed grouse
Bonasa umbellus
Turkey
Meleagris gallopavo
Barn owl
~ alba
Flammulated owl
Otus flammeolus
Great horned owl
Bubo virginianus
Pygmy owl
Glaucidium gnoma
Barred owl
Strix varia
Great gray owl
Strix nebulosa
Long-eared owl
Asio otus
Saw-whet owl
Aegolius acadicus
Vaux's swift
Chaetura vauxi
Black-chinned hummingbird
Archilochus alexandri
Rufous hummingbird
Selasphorus rufus
Calliope hummingbird
Stellula calliope
Belted kingfisher
Megaceryle alcyon
Common flicker
Colaptes auratus
Pileated woodpecker
Dryocopus pileatus
Lewis' woodpecker
Melanerpes lewis
Grassforb
Shrubseeding
Polesapling
Young
Mature
Oldgrowth
RF
RF
RF
RF
RF
RF
F
F
F
F
RF
RF
F
F
R
RF
F
RF
RF
F
F
F
F
RF
RF
RF
F
F
RF
RF
F
F
RF
RF
R
RF
RF
R
RF
R
F
RF
R
F
F
F
F
F
F
F
RF
RF
RF
F
F
F
RF
RF
RF
F
F
RF
RF
RF
F
F
RF
RF
RF
R
RF
RF
RF
RF
F
F
R
F
F
F
F
F
RF
RF
F
RF
R
F
RF
RF
F
F
F
RF
F
F
F
F
F
F
F
F
RF
RF
RF
RF
RF
RF
RF
R
R
234
F
RF
R
RF
R
Appendix 1.--Continued.
Forest Successional Stage
Species
Williamson's sapsucker
Sphyrapicus thyroideus
Hairy woodpecker
Picoides villosus
White-headed woodpecker
Picoides albolarvatus
Black-backed three-toed
woodpecker
Picoides arcticus
Alder flycatcher
Empidonax alnorum
Willow flycatcher
Empidonax traillii
Hammond's flycatcher
Empidonax hammondii
Dusky flycatcher
Empidonax oberholseri
Western flycatcher
Empidonax difficilis
Western wood pewee
Contopus sordidulus
Olive-sided flycatcher
Nuttalornis borealis
Tree swallow
Iridoprocne bicolor
Gray jay
Perisoreus canadensis
Steller's jay
Cyanocitta stelleri
Black-billed magpie
Pica pica
Common raven
Corvus corax
Common crow
Corvus brachirhinchos
Clark's nutcracker
Nucifraga columbiana
Mountain chickadee
Parus gambeli
Chestnut-backed chickadee
Parus rufescens
White-breasted nuthatch
Sitta carolinensis
Red-breasted nuthatch
Sitta canadensis
Grassforb
Shrubseeding
Polesapling
Young
RF
RF
Mature
Oldgrowth
RF
RF
RF
RF
RF
RF
RF
RF
F
RF
RF
F
RF
RF
F
F
F
RF
RF
RF
RF
RF
RF
RF
F
F
RF
RF
RF
F
F
RF
RF
RF
F
F
RF
RF
RF
RF
F
F
R
R
RF
F
F
RF
RF
RF
F
F
F
RF
RF
RF
RF
F
RF
RF
RF
{i
F
RF
RF
RF
F
F
RF
F
RF
RF
RF
F
RF
RF
RF
RF
RF
RF
RF
RF
RF
RF
F
RF
RF
RF
R
RF
235
Appendix 1.--Continued.
Forest Successional Stage
Species
Pygmy nuthatch
Sitta pygmaea
Brown creeper
Certhia familiaris
Dipper
Cinclus mexicanus
House wren
Troglodytes aedon
Winter wren
Troglodytes troglodytes
Rock wren
Salpincles obsoletus
American robin
Turdus migratorius
Varied thrush
Ixoreus naevius
Hermit thrush
Catharus guttatus
Swainson's thrush
Catharus ustulatus
Western bluebird
Sialia mexicana
Mountain bluebird
Sialia currucoides
Golden-crowned kinglet
Regulus satrapa
Ruby-crowned kinglet
Regulus calendula
Solitary vireo
Vireo solitarius
Nashville warbler
Vermivora ruficapilla
Yellow-rumped warbler
Dendroica coronata
Black-throated gray warbler
Dendroica ni~rescens
Townsend's warbler
Dendroica townsendi
MacGillivray's warbler
Oporornis tolmiei
Wilson's warbler
Wilsonia pusilla
Northern waterthrush
Seiurus noveboracensis
Yellow-breasted chat
Icteria virens
Grassforb
Shrubseeding
Polesapling
Mature
Oldgrowth
RF
RF
F
RF
RF
Young
RF
RF
RF
RF
RF
RF
F
F
RF
R
R
R
RF
RF
F
RF
RF
RF
RF
F
RF
RF
RF
RF
RF
F
RF
RF
RF
RF
F
F
RF
RF
RF
RF
RF
RF
RF
F
F
R
R
R
F
F
R
R
R
F
RF
RF
RF
F
F
RF
RF
RF
RF
RF
RF
F
F
F
RF
RF
RF
RF
RF
RF
RF
F
RF
RF
F
RF
RF
RF
RF
RF
RF
RF
RF
RF
RF
RF
RF
RF
RF
RF
RF
236
F
Appendix 1.--Continued.
Forest Successional Stage
Species
Northern oriole
Icterus galbula
Brewer's blackbird
Euphagus cyanocephalus
Brown-headed cowbird
Molothrlls ater
Western tanager
Piranga ludoviciana
Black-headed grosbeak
Pheucticus melanocephalus
Evening grosbeak
Hesperiphona vespertina
Purple finch
Carpodacus purpureus
Cassin's finch
Carpodacus cassinii
House finch
Carpodacus mexicanus
Pine grosbeak
Pinicola enucleator
Pine siskin
Carduelis pinus
Red crossbill
Loxia curvirostra
Dark-eyed junco
Junco hyemalis
Chipping sparrow
Spizella passerina
Song sparrow
Melospiza melodia
Shrubseeding
Polesapling
Young
Mature
RF
RF
RF
RF
F
RF
RF
RF
R F·
RF
RF
RF
RF
RF
RF
F
F
F
RF
RF
RF
F
F
RF
RF
RF
F
F
F
RF
RF
Grassforb
Oldgrowth
F
F
RF
RF
RF
RF
F
F
F
RF
RF
RF
F
RF
RF
RF
RF
F
F
F
RF
RF
RF
RF
RF
RF
F
RF
RF
F
F
RF
RF
RF
RF
RF
RF
F
RF
RF
RF
RF
RF
R J.t·
RF
RF
RF
237