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The adverse impacts of roadbuilding and logging on wildlife and ecological values -- surveyed by NRDC's End of the Road report -- are among the best-studied topics in the natural sciences. This supplement to our report provides additional detail on the specific effects of logging on North American wildlife, as studied and recorded in independently peer-reviewed scientific publications.


TABLE OF CONTENTS

MAMMALS
American Marten
Fisher
Wolverine
Alexander Archipelago Wolf
Brown (Grizzly) Bear
Black Bear
Sitka Black-tailed Deer
Woodland Caribou
Small Mammals
Bats
Other Mammal Studies

BIRDS
Pileated Woodpecker
Vaux's Swift
Black-Backed Woodpecker
Three-Toed Woodpecker
Red Crossbill
Marbled Murrelet
Northern Goshawk
Flammulated Owls
Forest Interior Songbirds
Other Avian Species

AMPHIBIANS


MAMMALS

American Marten
Martes americana

Bull, E. L., and T. W. Heater. In press. Resting and denning sites of American marten in northeastern Oregon. Northwest Science.

The authors radiotracked marten in the Blue Mountains of northeastern Oregon to determine what habitat structures were important for the species. Large-diameter live trees, large snags, and large hollow logs were important resting and denning sites. Natal and post-natal dens were in cavities in large trees (averaging 83 cm dbh), in large hollow logs (averaging 73 cm in diameter), underground, or in slash piles (only 6 percent). Resting sites were in trees with platforms (such as from dwarf mistletoe), in trees with cavities, in undersnow spaces, and in hollow logs or slash piles. During winter, marten primarily rested in subnivean (undersnow) spaces formed by coarse woody debris.

Flynn, R. W., and T. V. Schumacher. 2000. Ecology of marten in southeast Alaska. Research progress report. Juneau, Alaska: Federal Aid in Wildlife Restoration, Alaska Department of Fish and Game.

The authors collected data on marten ecology for nine years on northeast Chichagof Island in southeastern Alaska. For resting and denning, marten used large-diameter live trees with decay (mean dbh of 83 cm), large snags (mean dbh of 88 cm), and large logs (86 cm in diameter), selecting the largest diameter structures available in the study area. Clearcuts had low use as resting and denning sites and were never used for resting in winter. In terms of overall habitat use, investigated by radiotelemetry, marten primarily selected forested habitat, in particular multistoried forest dominated by large trees (> 82 cm dbh) or medium trees (59-81.9 cm dbh). They showed the lowest preference for recent clearcuts and other nonforest habitat types.

Bull, E. L., and A. K. Blumton. 1999. Effect of fuels reduction on American marten and their prey. Research note, PNW-RN-539. Portland, Oregon: USDA Forest Service, Pacific Northwest Research Station.

The impact of forest fuel reduction on primary marten prey was studied in lodgepole pine and mixed-conifer stands in northeastern Oregon. This study was conducted because of the strong association of red-backed voles and red squirrels with down logs and undersnow spaces. Forest fuel reduction was done by logging as well as by removing downed wood. Results generally showed that red-backed vole, red squirrel, and snowshoe hare numbers declined after fuel treatments.

Ruggiero, L. F., D. E. Pearson, and S. E. Henry. 1998. Characteristics of American marten den sites in Wyoming. Journal of Wildlife Management 62:663-73.

The authors radiotracked female marten in the Medicine Bow National Forest, Wyoming, in order to characterize their dens and den sites. Natal and maternal dens occurred in snags, logs, red squirrel middens (frequently associated with logs and snags), as well as rock crevices. Marten selected sites with late-successional characteristics -- more large-diameter (> 41 cm) logs, more middens, and greater canopy cover than random for their natal dens, and more large-diameter logs and large- (> 41 cm) and medium- (20-40 cm) diameter snags than random for maternal dens.

Chapin, T. G., D. J. Harrison, and D. D. Katnik. 1998. Influence of landscape pattern on habitat use by American marten in an industrial forest. Conservation Biology 12:1327-37.

The authors radiotracked marten to evaluate the impact of logging and forest fragmentation on marten populations in a commercial forest in Maine. Home ranges were associated with large and contiguous forest patches, and there was little or no use of the smallest forest patches and the most isolated patches.

Chapin, T. G., D. J. Harrison, and D. M. Phillips. 1997. Seasonal habitat selection by marten in an untrapped forest preserve. Journal of Wildlife Management 61:707-17.

Marten were radiotracked in a Maine preserve to investigate their use of different forest types on the stand scale (mature coniferous, deciduous, or mixed forest). Stands with severe spruce-budworm mortality had the highest selection in summer; no other differences in selection were detected. The authors believe that the vertical and horizontal structure provided in the stand with budworm mortality (no budworm management had occurred), are important habitat components for the marten.

Buskirk, S. W., and L. F. Ruggiero. 1994. American marten. In The scientific basis for conserving forest carnivores: American marten, fisher, lynx, and wolverine in the western United States, edited by L. F. Ruggiero, K. B. Aubry, S. W. Buskirk, L. J. Lyon, and W. J. Zielinski. General Technical Report, RM-254. Fort Collins, Colorado: USDA Forest Service, Rocky Mountain Forest and Range Experiment Station.

This chapter provides a review of published and unpublished information on the American marten and its life history. Research on habitat demonstrates use of old-growth components for dens, coarse woody debris for access to undersnow (subnivean) spaces, coarse woody debris for protective thermal environments during winter, and avoidance of areas that lack overhead cover. Important prey species -- red-backed voles and pine squirrels -- are associated with old-growth and mature forest. Middens of red squirrels and Douglas' squirrels are used as resting sites; these species are also associated with late successional forest.

Martin, S. K. 1994. Feeding ecology of American marten and fishers. In Marten, sables, and fishers: Biology and conservation, edited by S. W. Buskirk, A. S. Harestad, M. G. Raphael, and R. A. Powell. Ithaca, New York: Cornell University Press.

The author reviews studies on marten diet throughout the species' range and through different seasons. Red-backed voles, an important prey species, are associated with forest interiors.

Thompson, I. D., and A. S. Harestad. 1994. Effects of logging on American martens, and models for habitat management. In Martens, sables, and fishers: Biology and conservation, edited by S. W. Buskirk, A. S. Harestad, M. G. Raphael, and R. A. Powell. Ithaca, New York: Cornell University Press.

This review includes research on the effects of logging on marten and a proposed model of stand development and carrying capacity. Adverse effects of logging, through the loss of habitat, are categorized as either short-term or long-term. Over the short term, studies reviewed show that marten avoid recent clearcuts. Over the longer term, forests may become too fragmented on the landscape scale to provide suitable habitat, and if logged mixed-wood forests regenerate to deciduous forests, they provide insufficient prey and unsuitable habitat during winter.

Corn, J. G., and M. G. Raphael. 1992. Habitat characteristics at marten subnivean access sites. Journal of Wildlife Management 56:442-48.

Marten use of undersnow spaces was studied in southern Wyoming. Coarse woody debris was found to be an important component of marten winter habitat by providing access to undersnow spaces, whether for resting or for getting to prey. The sites used by marten had a greater volume of coarse woody debris and a greater volume of undecayed and moderately decayed logs than surrounding stands.

Buskirk, S. W., S. C. Forrest, M. G. Raphael, and H. J. Harlow. 1989. Winter resting site ecology of marten in the central Rocky Mountains. Journal of Wildlife Management 53:191-96.

Marten were radiotracked to investigate their preferred resting sites during winter in southeastern Wyoming. Resting sites most important to individuals were situated in and around coarse woody debris, particularly during the coldest weather.

Spencer, W. D. 1987. Seasonal rest-site preferences of pine martens in the northern Sierra Nevada. Journal of Wildlife Management 51:616-21.

The author investigated marten preference for resting sites on the east side of the Sierra Nevada, California. For above-snow nesting, snags used were the most relative to their availability and marten selected for the largest snags available (mean diameter of 102 cm). Large live trees (mean diameter of 62 cm) were also important resting sites. During periods of high snow cover, marten rested in undersnow spaces created by coarse woody debris, snags, and stumps.

Buskirk, S. W. 1984. Seasonal use of resting sites by marten in south-central Alaska. Journal of Wildlife Management 48:950-53.

Marten were radiotracked during winter to determine characteristics of their resting sites. The majority of their resting sites were red squirrel middens, primarily located in old-growth stands of white spruce or mixed spruce/birch stands with greater than 60 percent canopy closure.

Hargis, C. D., and D. R. McCullough. 1984. Winter diet and habitat selection of marten in Yosemite National Park. Journal of Wildlife Management 48:140-46.

Habitat selection of marten was studied over two winters in Yosemite National Park, California. Marten avoided areas without canopy cover and preferred areas with 100 percent canopy cover and with cover low in height (less than 3 m above the snow). Logs, melt zones around large trees, and sapling branches provided marten with access to undersnow spaces. Because of variations in snowpack depth from year to year, the authors believe mixed-aged forests to be more suitable for marten needs than even-aged stands.

Wynne, K. M., and J. A. Sherburne. 1984. Summer home range use by adult marten in northwestern Maine. Canadian Journal of Zoology 62: 941-43.

Home ranges, resting sites, and dens used by female marten were characterized by radiotracking adult marten in northwestern Maine. Both males and females avoided nonforested areas and hardwood stands. Females showed a preference for dense, mature softwoods and used large-diameter logs or standing trees for dens.

Soutiere, E. C. 1979. Effects of timber harvesting on marten in Maine. Journal of Wildlife Management 43: 850-60.

Marten occurrence was compared across clearcut, partially cut, and uncut forest in north-central Maine. Uncut forest had the heaviest use, activity was slightly reduced in partially cut stands, and 0- to 15-year-old clearcuts were rarely used. Within clearcut areas, marten activity during winter was higher in the islands of uncut and lightly cut stands than in the cutover areas.

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Fisher
Martes pennanti

Carroll, C., W. J. Zielinksi, and R. F. Noss. 1999. Using presence-absence data to build and test spatial habitat models for the fisher in the Klamath Region, U.S.A. Conservation Biology 13: 1344-59.

The authors developed a model to predict fisher distribution, using data from surveys and vegetational maps in the Klamath region of northwestern California and southwestern Oregon and the northern California coast. Fisher were strongly associated with landscapes having high levels of tree canopy closure. On a finer, plot scale, their presence was significantly related to large-diameter hardwoods. The authors attribute this to the importance of large hardwoods as resting and denning sites and as a source of mast for fisher prey species.

Truex, R. L., W. J. Zielinksi, R. T. Golightly, R. H. Barrett, and S. M. Wisely. 1998. A meta-analysis of regional variation in fisher morphology, demography, and habitat ecology in California. Draft report. California Department of Fish and Game.

Data were compared from three studies on the ecology of the Pacific fisher (Martes pennanti pacifica) in California -- in the eastern Klamath Mountains (Shasta-Trinity National Forest), in the North Coast ranges (Six Rivers National Forest), and in the southern Sierra Nevada (Sequoia National Forest). Habitat used for denning had high canopy closure (70-100 percent). Five natal dens were located; all were in cavities of large-diameter trees (3 white fir averaging 114 cm dbh and 2 black oak averaging 87.5 cm dbh). Most maternal dens were also in large-diameter trees. Resting sites also occurred in habitat with high canopy closure (88-94 percent). Cavities and broken tops of large-diameter trees and snags were important resting sites for fisher in the southern Sierra and North Coast study sites (conifers > 105 cm dbh; hardwoods > 64 cm dbh). In the eastern Klamath study area, fisher used smaller-diameter trees for resting, and used more nest and mistletoe broom platforms. Fisher density was lower in the latter study area; the authors hypothesize that habitat quality in this area is lower (due to heavier logging and fewer large-diameter trees and snags) and that the area may be a population sink for fisher.

Powell, R. A., and W. J. Zielinksi. 1994. Fisher. In The scientific basis for conserving forest carnivores: American marten, fisher, lynx, and wolverine in the western United States, edited by L. F. Ruggiero, K. B. Aubry, S. W. Buskirk, L. J. Lyon, and W. J. Zielinski. General technical report, RM-254. Fort Collins, Colorado: USDA Forest Service, Rocky Mountain Forest and Range Experiment Station.

The authors review published and unpublished literature as of 1994 on the fisher's life history, the impacts of trapping and logging, and management recommendations. Fisher have been found to be more selective for denning and resting habitat than for foraging habitat. Studies on habitat relationships indicate that throughout its range, the fisher prefers habitat with overhead cover and avoids open areas, including recent clearcuts. In the western United States, fisher prefer mature and old-growth coniferous forest and riparian areas over younger successional stages. The few available studies on dens in the West show that fisher use large logs and snags. Resting sites occur primarily in closed canopy stands and include large trees, logs, and snags. The interception of snowfall by overhead canopy is hypothesized to be another factor influencing their distribution.

Buskirk, S. W., and R. A. Powell. 1994. Habitat ecology of fishers and American martens. In Martens, sables, and fishers: Biology and conservation, edited by S. W. Buskirk, A. S. Harestad, M. G. Raphael, and R. A. Powell. Ithaca, New York: Cornell University Press.

This chapter provides a review of research on habitat use by fisher. Multiple studies showed an avoidance of open areas throughout the species' range. In the Pacific Northwest, fisher are associated with low-elevation late-successional Douglas-fir forests; this is partly attributed to the association of their preferred prey species -- snowshoe hares -- with this habitat. Snowshoe hares were shown to be common in habitat with dense physical structure near the ground, including coarse woody debris, rocks, small trees, and low branches.

Jones, J. L., and E. O. Garton. 1994. Selection of successional stages by fishers in north-central Idaho. In Martens, sables, and fishers: Biology and conservation, edited by S. W. Buskirk, A. S. Harestad, M. G. Raphael, and R. A. Powell. Ithaca, New York: Cornell University Press.

The authors investigated use of five forest successional stages by radiocollared fisher in Nez Perce National Forest, Idaho. During the summer, 90 percent of fisher observations occurred in mature or old-growth forest, and individuals avoided nonforest, pole-sapling, and young forest successional stages. During the winter, fisher used mature, old-growth, and young forest, and preferred the young forest relative to nonforest or pole-sapling stands.

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Wolverine
Gulo gulo

Krebs, J. A., and D. Lewis. In press. Wolverine ecology and habitat use in the North Columbia Mountains: Progress report. Nelson, British Columbia: Columbia Basin Fish and Wildlife Compensation Program. June 15, 1999. In Species at Risk conference proceedings, February. 1999.

This report presents preliminary results from an ongoing study of a wolverine population in the Kootenay Region of British Columbia. All female natal dens were found within roadless tributary valleys, and were under woody debris or a combination of woody debris and large boulders. Protected areas (National Parks) and unroaded wilderness appeared to be refugia, having the highest use by wolverine and used in proportions greater than their availability.

Copeland, J. P. 1996. Biology of the wolverine in central Idaho. Master's thesis, University of Idaho.

Wolverine populations were studied by radiotelemetry in roadless and wilderness areas of the Sawtooth, Challis, and Boise National Forests in central Idaho. Natal dens used by females were in secluded high-elevation cirque basins. Human disturbance at maternal dens resulted in abandonment of the dens.

Banci, V. 1994. Wolverine. In The scientific basis for conserving forest carnivores: American marten, fisher, lynx, and wolverine in the western United States, edited by Leonard F. Ruggiero, K. B. Aubry, S. W. Buskirk, L. J. Lyon, and W. J. Zielinski. General technical report, RM-254. Fort Collins, Colorado: USDA Forest Service, Rocky Mountain Forest and Range Experiment Station.

This is a review of the literature as of 1994 on wolverine. Reports from throughout the species' range indicate that wolverine occupy inaccessible wilderness areas, remote from human activity and development.

Hornocker, M. G., and H. S. Hash. 1981. Ecology of the wolverine in northwestern Montana. Canadian Journal of Zoology 59:1286-1301.

A wolverine population was studied over five years in the Flathead National Forest in northwestern Montana. Wolverine avoided clearcuts. The majority of wolverine locations were in mature forest, with individuals rarely located in areas of dense, young timber.

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Alexander Archipelago Wolf
Canis lupus ligoni

Person, D. K., M. Kirchhoff, V. V. Ballenberghe, G. C. Iverson, and E. Grossman. 1996. The Alexander Archipelago wolf: A conservation assessment. General technical report, PNW-GTR-384. Portland, Oregon: USDA Forest Service, Pacific Northwest Research Station.

The authors summarize available information on the Alexander Archipelago wolf in Tongass National Forest. All documented wolf dens were in old-growth forest; one was under a large log, the others were in cavities beneath the roots of large trees (mean diameter, > 90 cm). Core-use areas of wolves studied in Prince of Wales Island were in the least densely roaded portion of their home range. Mortality due to hunting and trapping was higher in areas with higher road densities, and the authors note that a much higher percentage of wolf hunting is now from roads rather than boats. Wolf populations are closely tied to that of their primary prey, Sitka black-tailed deer. Intact productive old-growth forest is critical for these deer during the winter.

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Brown (Grizzly) Bear
Ursus arctos

Schoen, J. W., R. W. Flynn, L. H. Suring, K. Titus, and L. R. Beier. 1994. Habitat-capability model for brown bear in southeast Alaska. International Conference on Bear Research and Management 9:327-37.

The authors used data from radiocollared brown bears on Admiralty and Chichagof Islands in southeast Alaska to assess bear habitat preferences and to develop a habitat capability model. The model predicted reduced habitat capability in areas of human activity, for reasons including low habitat suitability of clearcuts and young second-growth forest and increased human-caused mortality near roads and other developments.

Kasworm, W. F., and T. L. Manley. 1991. Road and trail influences on grizzly bears and black bears in northwest Montana. International Conference on Bear Research and Management 8: 79-84.

Grizzly bears and black bears were radiocollared and monitored in the Cabinet Mountains of northwest Montana during spring and fall. Grizzly bears avoided habitat within 914 m of open roads and within 122 m of closed roads and trails. When a seasonally closed road was opened, grizzly bear locations were, on average, farther from the road than when it was closed.

Schoen, J., and L. Beier. 1990. Brown bear habitat preferences and brown bear logging and mining relationships in Southeast Alaska. Research final report. Juneau, Alaska: Alaska Department of Fish and Game, Federal Aid in Wildlife Restoration Project.

This final report summarizes data collected from 1981 to 1989 on radiocollared brown bears in southeast Alaska, on Admiralty and Chichagof Islands. Although bears used a variety of habitat through the year, upland old-growth and riparian old-growth were particularly important at certain times of the year and used in proportions much greater than their availability. Clearcuts were avoided. The mortality due to human hunting more than doubled during the time period in which most road building and logging occurred; the authors attribute this to easier human access.

Mattson, D. J., R. R. Knight, and B. M. Blanchard. 1987. The effects of developments and primary roads on grizzly bear habitat use in Yellowstone National Park, Wyoming. International Conference on Bear Research and Management 7:259-73.

Radiocollared grizzly bears were studied in Yellowstone National Park. Roads and developments displaced bears, with lower occupancy and disrupted foraging in habitat near them. Roads and developments had a disproportionately greater adverse impact on adult females and subadult males, which were displaced by adult males into habitat nearer roads and development. The authors conclude that this probably resulted in higher mortality and lower fecundity of the adult females.

Schoen, J. W., L. R. Beier, J. W. Lentfer, and A. J. Johnson. 1987. Denning ecology of brown bears on Admiralty and Chichagof Islands. International Conference on Bear Research and Management 7:293-304.

Characteristics of brown bear winter dens in the Tongass National Forest were studied by radiotracking bears on Admiralty and Chichagof Islands in southeastern Alaska. Bears used old-growth forest most frequently and generally denned at high elevations and on steep slopes. In more forested and less mountainous areas, as on Chichagof Island and in coastal areas of Admiralty Island, the bears preferred den sites in mid-volume, old-growth forest. Large-diameter Sitka spruce trees (> 75 cm dbh) and large snags with heart rot were used for excavating dens.

Peek, J. M., M. R. Pelton, H. D. Picton, J. W. Schoen, and P. Zager. 1987. Grizzly bear conservation and management: A review. Wildlife Society Bulletin 15:160-69.

This article is a review of grizzly bear conservation issues in North America. Regarding forest management, the authors note that increased human access due to forest development results in increased disturbance of bears and increased human-caused mortality.

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Black Bear
Ursus americanus

Bull, E. L., J. J. Akenson, and M. G. Henjum. In press. Characteristics of black bear dens in trees and logs in northeastern Oregon. Northwestern Naturalist.

This study investigated the characteristics of black bear den sites used by radiocollared individual bears in the Blue Mountains of northeastern Oregon. The majority of denning trees were in moist, late-seral stands of grand fir. Forty-one percent of the dens were in large-diameter hollow trees or logs. Mean diameter of den logs was 108 cm. Mean diameter of den trees was 108 cm for base-entry trees and 114 cm for top-entry trees.

Oli, M. K., H. A. Jacobson, and B. D. Leopold. 1997. Denning ecology of black bears in the White River National Wildlife Refuge, Arkansas. Journal of Wildlife Management 61:700-706.

Black bear denning was studied in a bottomland hardwood forest in Arkansas. The majority of dens (90.2 percent) were in elevated tree cavities, with the average diameter of den trees 117.5 cm.

Kasbohm, J. W., M. R. Vaughan, and J. G. Kraus. 1996. Black bear denning during a gypsy moth infestation. Wildlife Society Bulletin 24:62-70.

The authors' primary objective was to determine the impact of a gypsy moth infestation in Shenandoah National Park, Virginia, on the timing of den entry and emergence and on the mortality of den trees. Dens used by radiocollared bears were described as part of the study. Dens were predominantly cavities in large-diameter live oaks and snags (mean tree diameter, 96.8 cm). These tree cavities were particularly important for female bears -- 84 percent of pregnant females denned in tree cavities.

Kasworm, W. F., and T. J. Thier. 1994. Adult black bear reproduction, survival, and mortality sources in northwest Montana. International Conference on Bear Research and Management 9:223-30.

Black bear habitat use and population characteristics were studied for ten years in the Cabinet Mountains and Yaak River of northwest Montana. Hunting and illegal kills were the highest causes of mortality for radiocollared bears during the study period. Radiotelemetry showed that bears killed had used habitat significantly closer to roads than had bears that survived.

Unsworth, J. W., J. J. Beecham, and L. R. Irby. 1989. Female black bear habitat use in west-central Idaho. Journal of Wildlife Management 6:668-73.

Activities of radiocollared female black bears were monitored in a study area in west-central Idaho that included clearcuts, selection cuts, riparian areas, meadows, and shrubfields. Uncut forested sites were important bedding sites, with bears using them in proportions greater than their availability.

Mollohan, C. M. 1987. Characteristics of adult female black bear daybeds in northern Arizona. International Conference on Bear Research and Management 7:145-49.

The characteristics of black bear bedding sites were investigated by radiotracking female bears in northern Arizona (Apache-Sitgreaves, Coconino, and Tonto National Forests). The majority of daybeds (74 percent) were located on the uphill side of the largest tree on the site (average diameter, 73 cm). Other bedding sites were associated with logs, large rocks, or dense thickets of regeneration.

Brody, A. J., and J. N. Stone. 1987. Timber harvest and black bear population dynamics in a southern Appalachian forest. International Conference on Bear Research and Management 7:243-50.

The authors used data from research in the Pisgah National Forest of North Carolina to develop a model that simulated the effect of different logging operations on black bear populations. The model included vulnerability to hunting, which increased with road density. The model indicated that any benefit to habitat carrying capacity from logging was greatly outweighed by the increase in vulnerability to hunting due to increased road access.

Young, D. D., and J. J. Beecham. 1986. Black bear habitat use at Priest Lake, Idaho. International Conference on Bear Research and Management 6:73-80.

The authors studied habitat use by black bears in the Selkirk Mountains of Idaho. Bears used a variety of habitat but avoided clearcuts during all seasons. Female bears avoided roads.

Wathen, W. J., K. G. Johnson, and M. R. Pelton. 1986. Characteristics of black bear dens in the southern Appalachian region. International Conference on Bear Research and Management 6:119-27.

The authors analyzed characteristics of winter dens used by black bears in Cherokee National Forest and the Great Smoky Mountains National Park. Large trees were important for both tree dens and ground dens. A majority of dens (55.8%) were in tree cavities high above the ground; average diameter of these trees was 100.7 cm. Ground dens included cavities at the base of live trees (average diameter, 102.7 cm) and cavities under roots of wind-tilted trees.

Manville, A. M., II. 1983. Human impact on the black bear in Michigan's Lower Peninsula. International Conference on Bear Research and Management 5:20-33.

Bears were radiocollared to investigate the impact of a variety of human activities, including oil well service roads. Hunting pressure on bears was heavier in areas that had extensive road systems.

Beecham, J. J., D. G. Reynolds, and M. G. Hornocker. 1983. Black bear denning activities and den characteristics in west-central Idaho. International Conference on Bear Research and Management 5:79-86.

Black bear dens were characterized as part of a study in west-central Idaho. Bears used a variety of sites for denning, including hollow logs and large hollow trees; trees used averaged 111.5 cm in diameter.

Johnson, K. G., and M. R. Pelton. 1981. Selection and availability of dens for black bears in Tennessee. Journal of Wildlife Management 45:111-19.

Radio-telemetry data were collected on black bears in the Great Smoky Mountains National Park, Tennessee. Bears preferred tree cavities high above ground as winter dens (averaging 11.2 m above ground) rather than ground dens. Cavities used were in large trees and snags; average diameter of trees was 95 cm. Bears also used dens in cavities under root systems of wind-tilted trees; average diameter of these trees was 72 cm. Adult females (who use dens for giving birth and lactation) selected tree cavities more frequently than did adult males.

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Sitka Black-tailed Deer
Odocoileus hemionus sitkensis

Kirchhoff, M. D., and S. R. G. Thomson. 1998. Effects of selective logging on deer habitat in southeast Alaska: A retrospective study. Research final report. Juneau, Alaska: Alaska Department of Fish and Game, Federal Aid in Wildlife Restoration Project.

The authors investigated forest composition, structure, and growth in stands with different intensities of logging conducted 50 or more years prior. The plots sampled had 20 to 80 percent of their basal area removed. The biomass of winter deer forage species (primarily blueberry) significantly decreased as logging intensity increased.

Person, D. K., M. Kirchhoff, V. V. Ballenberghe, G. C. Iverson, and E. Grossman. 1996. The Alexander Archipelago wolf: A conservation assessment. General technical report, PNW-GTR-384. Portland, Oregon: USDA Forest Service, Pacific Northwest Research Station.

The habitat needs of Sitka black-tailed deer are summarized as part of this conservation assessment on the Alexander Archipelago wolf. The research reviewed indicates that old-growth forest stands are important to deer during the winter and that clearcuts and even-aged second-growth stands provide poor habitat. In addition, fragmentation of old-growth due to clearcut logging results in deer being confined to isolated stands during the winter and suffering higher rates of mortality from malnutrition or predation.

Hanley, T. A. 1991. Old-growth forests and biological conservation in temperate rainforests of North America. In Wildlife conservation: Present trends and perspectives for the 21st century, edited by N. Maruyama, B. Bobek, Y. Ono, W. Regelin, L. Bartos, and P. Ratcliffe. Proceedings of the International Symposium on Wildlife Conservation in Tsukuba and Yokohama, Japan (August 21-25, 1990). Tokyo: Japan Wildlife Research Center.

A brief review of old-growth temperate rainforest ecology, focusing on southeastern Alaska and natural disturbance regimes versus clearcut disturbance. In particular, the authors address the impact on herbivores such as deer because of differences in plant species and plant chemical composition between old-growth and even-aged managed forests. Leaves growing in shaded understories have two times greater concentrations of digestible proteins than leaves (of the same species) growing in open clearcuts. Even-aged management and silvicultural thinnings also result in lower forage quality because herbs -- the most nutritiously valuable forage -- are quickly shaded out by a dense shrub layer and the natural, small, and frequent disturbance regime of gap formation in old forests is no longer present.

Schoen, J. W., and M. D. Kirchhoff. 1990. Seasonal habitat use by Sitka black-tailed deer on Admiralty Island, Alaska. Journal of Wildlife Management 54:371-78.

The authors radiotracked Sitka black-tailed deer to investigate their habitat use during different seasons. During winter and spring, the deer used old growth almost exclusively, with particularly high usage of high-volume old growth, as occurs at low elevations.

Hanley, T. A., C. T. Robbins, and D. E. Spalinger. 1989. Forest habitats and the nutritional ecology of Sitka black-tailed deer: A research synthesis with implications for forest management. General technical report, PNW-GTR-230. Portland, Oregon: USDA Forest Service, Pacific Northwest Research Station.

A review of research on habitat and food resources used by the Sitka black-tailed deer in western hemlock and Sitka spruce forests of southeastern Alaska. The authors conclude that it is important to retain old-growth forests, as they provide key food resources during the winter months, when they have lower snow accumulation, and during the summer, when shaded leaves provide higher-protein food.

Hanley, T. A., R. G. Cates, B. V. Horne, and J. D. McKendrick. 1987. Forest stand-age-related differences in apparent nutritional quality of forage for deer in southeastern Alaska. Proceedings -- Symposium on Plant-Herbivore Interactions. General technical report, INT-222. USDA Forest Service, Intermountain Research Station.

The authors conducted several studies to evaluate the quality of forage for Sitka black-tailed deer in southeastern Alaska. In particular, they analyzed the chemical composition of two preferred understory species: Alaska blueberry and bunchberry dogwood. A study comparing five stands of different ages showed that forage quality was higher (higher concentrations of nutrients and lower concentrations of phenolics and astringents) in older stands with well-developed canopies than in young clearcuts. A second study showed that concentrations of digestible protein were twice as great in leaves from an old-growth forest stand compared to an 8-year-old clearcut. And a comparison of palatability with captive deer showed that they preferred blueberry leaves from the forest to those from the clearcut.

Kirchhoff, M. D., and J. W. Schoen. 1987. Forest cover and snow: Implications for deer habitat in Southeast Alaska. Journal of Wildlife Management 51:28-33.

The authors investigated the relationship between snow depth and forest stand characteristics in order to identify valuable winter habitat for Sitka black-tailed deer. The lowest snow depths occurred in high-volume old-growth stands.

Rose, C. L. 1984. Response of deer to forest succession on Annette Island, southeastern Alaska. In Fish and wildlife relationships in old-growth forests, edited by W. R. Meehan, T. R. Merrell, Jr. and T. A. Hanley. Morehead City, North Carolina: American Institute of Fishery Research Biologists.

Deer use of old-growth forests during winter was compared to their use of clearcuts and second-growth forest (1- to 270-year-old stands) on Annette Island at the southern tip of the Alexander Archipelago. Deer showed a strong preference for old-growth forest in the winter, using old-growth stands almost seven times more than clearcuts and second-growth.

Wallmo, O. C., and J. W. Schoen. 1980. Response of deer to secondary forest succession in southeast Alaska. Forest Science 26:448-62.

The authors compared deer use of old-growth stands to their use of young, even-aged stands ranging in age from new clearcuts to 147-year-old second growth. Old-growth stands were used more intensively by Sitka black-tailed deer than were younger stands, regardless of the latter's age post-logging. Old-growth stands had a more diverse and abundant understory and therefore more deer forage than young-growth stands.

Bloom, A. M. 1978. Sitka black-tailed deer winter range in the Kadashan Bay area, southeast Alaska. Journal of Wildlife Management 42:108-12.

Deer use of mature forest during the late winter was evaluated. Deer use was greatest in low-elevation forests that were heavily stocked old growth. These forests also had the lowest snow accumulation.

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Woodland Caribou
Rangifer tarandus caribou

Rominger, E. M., C. T. Robbins, and M. A. Evans. 1996. Winter foraging ecology of woodland caribou in northeastern Washington. Journal of Wildlife Management 60:719-28.

Woodland caribou lichen foraging behavior during late winter was documented in experimental field trials. Trees were classified by the biomass of lichen present; higher biomass occurred on trees older than 100 to 120 years.

Servheen, G., and L. J. Lyon. 1989. Habitat use by woodland caribou in the Selkirk Mountains. Journal of Wildlife Management 53:230-37.

The authors radiocollared individual woodland caribou to investigate habitat use in the Selkirk Mountains. During early winter, caribou relied on old-growth western hemlock forest that had high basal area and closed canopy. Snow interception was greatest in these forests. Lichens (slow-growing and associated with older trees) were the primary forage during winter and the calving season.

Rominger, E. M., and J. L. Oldemeyer. 1989. Early-winter habitat of woodland caribou, Selkirk Mountains, British Columbia. Journal of Wildlife Management 53:238-43.

Radiocollared woodland caribou were monitored to investigate their use of habitat during early winter. Caribou used lower-elevation old-growth stands of western hemlock and western red cedar. Recently windthrown trees were an important source of lichens.

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Small Mammals

Carey, A. B. 1995. Small mammals in managed, naturally young, and old-growth forests. Ecological Applications 5:336-52.

Forest floor small-mammal communities were compared in old-growth forest (300-400 years old) and young managed stands (44-67 years old) on the Olympic Peninsula. Old growth had 1.5 times more individuals and biomass than managed forest. Sorex trowbridgii, Peromyscus oreas, Sorex monticolus, and Neurotrichus gibbsii were significantly more abundant in old growth than in managed stands.

Tallmon, D., and L. S. Mills. 1994. Use of logs within home ranges of California red-backed voles on a remnant of forest. Journal of Mammalogy 75:97-101.

California red-backed voles (Clethrionomys californicus) were radiotracked in an old-growth forest remnant in the Klamath Mountains of Siskiyou National Forest. Voles used only the forest remnant and not the surrounding clearcut. The great majority of their locations (98 percent) coincided with downed logs, with voles using logs in later stages of decay more than logs in earlier stages of decay.

Clarkson, D. A., and L. S. Mills. 1994. Hypogeous sporocarps in forest remnants and clearcuts in southwest Oregon. Northwest Science 68:259-65.

The authors studied California red-backed voles (C. californicus) and truffle production (a major food source for this species) in the Klamath Mountains. Late-successional forest remnants had significantly more truffles than did the clearcuts surrounding them. Within the forest remnants, truffles were associated with coarse woody debris. Red-backed voles were more likely to occur in areas with truffles than in those lacking them.

Nordyke, K. A., and S. W. Buskirk. 1991. Southern red-backed vole, Clethrionomys gapperi, populations in relation to stand succession and old-growth character in the central Rocky Mountains. Canadian Field-Naturalist 105:330-34.

Research was conducted on the southern red-backed vole (C. gapperi) in the Medicine Bow Mountains of southeastern Wyoming. Vole abundances were compared in different forest successional stages and forest types. Vole abundance was greatest and body condition best in old-growth spruce-fir.

Carey, A. B. 1991. The biology of arboreal rodents in Douglas-fir forests. General technical report, PNW-GTR-276. USDA Forest Service, Pacific Northwest Research Station.

The authors summarized the life history characteristics and habitat relations of tree rodent species closely associated with old growth in the Pacific Northwest west-side forests, including red tree voles, flying squirrels, Douglas' squirrels, woodrats, and Townsend's chipmunks. They report that the tree rodent community has the highest diversity and abundance in old-growth forests and all species are sensitive to logging because of loss of habitat and/or barriers to dispersal.

Buchanan, J. D., R. W. Lundquist, and K. B. Aubry. 1990. Winter populations of Douglas' squirrels in different-aged Douglas-fir forests. Journal of Wildlife Management 54:577-81.

The relative abundance of Douglas' squirrels (Tamiasciurus douglasii) during winter were compared in old growth and naturally regenerated young stands (< 165 years old) in the western Cascades of southern Washington. Squirrel populations were higher in old-growth forests. Squirrel populations were also closely tied to conifer cone production cycles and the authors note that old-growth forests likely provide greater and more reliable quantities of conifer seeds.

Hayes, J. P., and S. P. Cross. 1987. Characteristics of logs used by western red-backed voles, Clethrionomys californicus, and deer mice, Peromyscus maniculatus. Canadian Field-Naturalist 101:543-46.

The authors studied the association of red-backed voles (C. californicus) with different log characteristics in an old-growth mixed-conifer forest in the Oregon Cascades. Red-backed vole numbers were positively correlated with log diameter and with log overhang area, i.e. with larger logs.

Patton, D. R., and J. R. Vahle. 1986. Cache and nest characteristics of the red squirrel in an Arizona mixed-conifer forest. Western Journal of Applied Forestry 1:48-51.

The authors studied habitat characteristics of food cache sites and nesting sites of red squirrels (Tamiasciurus hudsonicus), a species closely associated with old-growth and mature forests. Primary caches of cones were in sites with high canopy cover, had an average of three large dominant conifers (which would produce the most cones) nearby, and were in stands with mixed-aged trees.

Vahle, J. R., and D. R. Patton. 1983. Red squirrel cover requirements in Arizona mixed- conifer forests. Journal of Forestry 81:1-15, 22.

The authors inventoried red squirrel food caches in a mixed-conifer forest. Caches were located in dense groups of trees providing shade and having one or more large, dominant conifers. Large snags and large downed logs were important sites for cone storage.

Ramirez, P., Jr., and M. Hornocker. 1981. Small mammal populations in different-aged clearcuts in northwestern Montana. Journal of Mammalogy 62:400-403.

The authors trapped small mammals in uncut subalpine fir-type forests and in clearcuts of varying ages. Densities of southern red-backed voles (C. gapperi) were significantly greater in the uncut forests than in the clearcuts.

Gashwiler, J. S. 1970. Plant and mammal changes on a clearcut in west-central Oregon. Ecology 51:1018-26.

Small mammal populations were compared in virgin old-growth forest and a clearcut on the west slope of the Cascade Mountains in Oregon. Red-backed voles (Clethrionomys occidentalis), Douglas' squirrels (Tamiasciurus douglasii), and northern flying squirrels (Glaucomys sabrinus) were not found on the clearcut but were present in old growth.

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Bats

Crampton, L. H., and R. M. R. Barclay. 1998. Selection of roosting and foraging habitat by bats in different-aged aspen mixed-wood stands. Conservation Biology 12:1347-58.

The relative abundance and foraging activity of bats were compared in old (> 120 years old), mature (50-65 years old), and young (20-30 years old) stands of aspen mixed wood in Alberta. Bat occurrence was highest in old stands. All roost trees used by females bats of the two species radiotracked -- little brown bats (Myotis lucifugus) and silver-haired bats (Lasionycteris noctivagans) -- were in old stands and were primarily in deep tree cavities. Roost trees were larger and more likely to have rot than random trees. Potential roost trees were scarce in young and mature forest.

Parker, D. I., J. A. Cook, and S. W. Lewis. 1996. Effects of timber harvest on bat activity in southeastern Alaska's temperate rainforests. In Bats and forests symposium, edited by M. R. Barclay and R. M. Brigham (held October 19-21, 1995). Victoria, British Columbia.

The authors investigated bat activity on Prince of Wales and Revillagigedo Islands in southeastern Alaska. Bat activity during the summer was studied in old-growth forest, riparian areas within old growth, second growth (25-70 years old), and young clearcuts (5-17 years old). Bat activity was higher in old growth than in second growth or clearcuts, and bat calls were heard in old growth during 76 percent of the nights monitored (versus 28 percent in clearcuts and 3 percent in second growth). The authors believe that old growth provides important summer roosting habitat. Riparian habitat was the most important foraging habitat.

Perkins, J. M., and S. P. Cross. 1988. Differential use of some coniferous forest habitat by hoary and silver-haired bats in Oregon. Murrelet 69:21-24.

The habitat association of hoary bats (Lasiurus cinereus) and silver-haired bats (Lasionycteris noctivagans) was investigated in Oregon coniferous forests. Both species showed a preference for old-growth Douglas-fir forest, with all hoary bats being captured in old-growth stands and 94 percent of silver-haired bats being captured in old growth. The authors attribute this to large old trees meeting these species' roosting requirements -- crevices in bark and limbs and foliage high above the ground.

Thomas, D. W. 1988. The distribution of bats in different ages of Douglas-fir forests. Journal of Wildlife Management 52:619-26.

The author compared bat distribution in young, mature, and old-growth Douglas-fir stands in the southern Washington Cascades and in the Oregon Coast Range. Activity detection rates in the early evening indicated that bats were 3-10 times more abundant in old-growth than in younger stands. The authors attribute this to higher roost availability in old growth.

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Other Mammal Studies

Hundertmark, K. J., W. L. Eberhardt, and R. E. Ball. 1990. Winter habitat use by moose in southeastern Alaska: Implications for forest management. Alces 26:108-14.

Habitat use by moose (Alces alces) in the Chilkat River watershed of southeastern Alaska was monitored using radiotelemetry. Moose use of forest stands during a low-snow winter and high-snow winter was compared. During the high-snow winter, moose used clearcuts, open areas, and deciduous stands much less than they did in the low-snow winter. The authors attribute this to the better shelter provided by uncut forest and by coniferous stands.

Schoen, J. W., M. D. Kirchhoff, and J. H. Hughes. 1988. Wildlife and old-growth forests in southeastern Alaska. Natural Areas Journal 8:138-45.

This article is an overview of wildlife species associated with old-growth forests in southeast Alaska and their habitat requirements. Even-aged second-growth stands that regenerate after clearcutting have been shown to be unsuitable for many species and remain so for more than one hundred years.

Rosenberg, K. V., and M. G. Raphael. 1986. Effects of forest fragmentation on vertebrates in Douglas-fir forests. In Wildlife 2000: Modeling habitat relationships of terrestrial vertebrates, edited by J. Verner, M. L. Morrison, and C. J. Ralph. Madison: The University of Wisconsin Press.

Vertebrate populations were inventoried in forty-six forest stands of varying sizes (bordered by clearcuts) in the Six Rivers, Klamath, and Shasta-Trinity National Forests in northwestern California. The authors tested for several different measures of fragmentation. Species that showed a negative response to forest fragmentation included the fisher, gray fox, spotted owl, pileated woodpecker, ringtail, northern flying squirrel, sharp-shinned hawk, blue grouse, and Pacific giant salamander.

Sigman, M. J. 1985. The impacts of clearcut logging on the wildlife resources of southeast Alaska. Technical report, 85-3, Part I. Juneau, Alaska: Alaska Department of Fish and Game.

This is a review of the literature documenting the impacts of clearcut logging on forest succession and old growth associated wildlife in southeast Alaska. It includes published and unpublished research and provides useful summaries of many of the studies' results. Species adversely affected by logging include the Sitka black-tailed deer, marten, land otter, mountain goat, osprey, brown bear, bald eagle, and a number of breeding bird species. Clearcut logging has been shown to eliminate habitat for species that depend on old-growth forest for all or part of the year. The benefits of increased forage in clearcuts for some species are negated by these areas being unusable during periods of deep snow, and winter habitat carrying capacity being much lower as a result. The benefits of increased understory production (after logging) are also short-term and lost within 20-30 years as the clearcut reaches the pole stand stage of succession. These cutover areas remain low in forage production for 75 percent of a 100-year rotation.

Schoen, J. W., and M. D. Kirchhoff. 1982. Habitat use by mountain goats in southeast Alaska. Final report. Juneau, Alaska: Alaska Department of Fish and Game, Federal Aid in Wildlife Restoration Project.

In their study of mountain goat (Oreamnus americanus) habitat use in southeast Alaska, the authors found that during winter goats on the coast used old growth near steep, rocky cliffs. Goats were found to be vulnerable to increased hunting if access by humans due to roads increased.

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BIRDS

Pileated Woodpecker
Dryocopus pileatus

Bull, E. L., and J. A. Jackson. 1995. Pileated woodpecker. In The birds of North America, No. 148, edited by A. Poole and F. Gills. Philadelphia: The Academy of Natural Sciences; Washington, DC: The American Ornithologists' Union.

This is a review of published and unpublished research available as of 1995, on the pileated woodpecker's life history. The authors cover information on habitat use throughout the bird's range, including the pileated woodpecker's preference for mature or old forest during the breeding season; its use of large hollow trees for roosting and tree cavities in large snags for nesting; and the reliance on snags and down logs when foraging for insects and beetle larvae. Conservation and management needs are also reviewed, with logging pinpointed as having the most adverse impact on pileated woodpecker habitat

Torgersen, T. R., and E. L. Bull. 1995. Down logs as habitat for forest-dwelling ants -- the primary prey of pileated woodpeckers in northeastern Oregon. Northwest Science 69:294-303.

The authors' research provides information on the foraging habitat of pileated woodpeckers in the Blue Mountains of northeastern Oregon. The authors were specifically interested in wood-dwelling ants, which are the primary prey of pileated woodpeckers. Study plots were located within woodpecker home ranges. Thirteen species of ants used a complex range of tree species and size and decay stage of logs. Carpenter ants (Camponotus spp.), a preferred prey species, occurred more frequently in large logs (51-120 cm in diameter) than in small logs.

Bull, E. L., and R. S. Holthausen. 1993. Habitat use and management of pileated woodpeckers in northeastern Oregon. Journal of Wildlife Management 57:335-45.

In order to evaluate Forest Service guidelines on snag density in pileated woodpecker management areas, the authors investigated population densities and habitat use in the Blue Mountains of Oregon. The density of large snags (> 51 cm dbh) was the best predictor of the density of pileated woodpeckers present. Radiotagging revealed that pileated woodpeckers used stands with old growth, stands with more than 60 percent canopy closure, grand fir, and no logging more than expected based on availability. Observations of foraging behavior indicated that 38 percent of observations were on logs, 38 percent on snags, 18 percent on live trees, and 6 percent on stumps. The woodpeckers showed a preference for logs greater than 38 cm in diameter and for logs with the greatest decay.

Renken, R. B., and E. P. Wiggers. 1993. Habitat characteristics related to pileated woodpecker densities in Missouri. Wilson Bulletin 105:77-83.

The authors examined the relationship between habitat and pileated woodpecker densities in oak-hickory forests of Missouri. Large trees and snags were important features in woodpecker habitat, with the authors' analysis showing that pileated woodpecker abundance was positively correlated with the density of trees larger than 30 cm in diameter, the density of snags larger than 54 cm in diameter, and the percent of area covered with bottomland forest. Woodpecker abundance was negatively correlated with percent of area covered with pole timber (15-25 cm in diameter).

Bull, E. L., R. S. Holthausen, and M. G. Henjum. 1992. Roost trees used by pileated woodpeckers in northeastern Oregon. Journal of Wildlife Management 56:786-93.

The authors studied the characteristics of roost trees and nest trees used by pileated woodpeckers in the Blue Mountains of northeastern Oregon. Roost trees were typically large-diameter live or dead trees (mean tree dbh 70.6 cm). The hollow interior of these trees was created primarily by decay, such as by Indian paint fungus. All nests were in large dead trees (mean tree dbh 80 cm). Old-growth grand fir stands with more than 60 percent canopy closure and little or no logging were preferred for roosting and nesting sites.

Bull, E. L. 1987. Ecology of the pileated woodpecker in northeastern Oregon. Journal of Wildlife Management 51:472-81.

The author presents results of an 11-year study on pileated woodpeckers in the Wallowa-Whitman National Forest in the Blue Mountains of Oregon. Pileated woodpeckers nested primarily in large-diameter dead trees, averaging 84 cm in diameter. Dead ponderosa pine and dead western larch were preferred (73 percent and 25 percent respectively). The majority of nest sites were in mature stands (54 percent) or in old-growth stands (21 percent). Woodpeckers also roosted in large-diameter dead trees, which were similar to nesting trees but had been dead longer. The birds foraged in large-diameter logs, snags, and live trees.

Bull, E. L., S. R. Peterson, and J. W. Thomas. 1986. Resource partitioning among woodpeckers in northeastern Oregon. Research note, PNW-444. Portland, Oregon: USDA Forest Service, Pacific Northwest Research Station.

The authors studied nesting and foraging habits of eight species of woodpeckers that coexist in the Blue Mountains of northeastern Oregon. Pileated woodpeckers foraged in logs, snags, and live trees, with a preference for larger-diameter material of each. Snags and live trees greater than 50 cm in diameter were preferred for foraging, as were Douglas-fir and western larch logs with diameters greater than 25 cm and lengths greater than 15 m. These woodpeckers nested in ponderosa pine and western larch snags, selecting the largest tree available. Habitat surrounding the nest tree had more large (> 30 cm dbh) live and dead trees present than other plots, and canopy cover was generally more than 75 percent.

Conner, R. N., R. G. Hopper, H. S. Crawford, and H. S. Mosby. 1975. Woodpecker nesting habitat in cut and uncut woodlands in Virginia. Journal of Wildlife Management 39:144-50.

The authors conducted research on the habitat used by four species of woodpeckers in the Jefferson National Forest in southwestern Virginia. Nests of pileated woodpeckers were typically in dead trees (13 out of 18 nests) or in dead sections of live trees. Nest trees were 33-91 cm in diameter, averaging 54.6 cm.

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Vaux's Swift
Chaetura vauxi

Bull, E. L., and C. T. Collins. 1993. Vaux's swift (Chaetura vauxi). In The Birds of North America, No. 77, edited by A. Poole and F. Gills. Philadelphia: The Academy of Natural Sciences; Washington, DC: The American Ornithologists' Union.

The authors summarize the literature available on the Vaux's swift as of 1993. In its breeding range (western North America), this species is more common in old-growth forest than in younger stands and uses large-diameter, hollow trees for nesting and for roosting.

Bull, E. L., and J. E. Hohmann. 1993. The association between Vaux's swifts and old- growth forests in northeastern Oregon. Western Birds 24:38-42.

The authors surveyed old-growth and logged stands for Vaux's swifts in the Umatilla National Forest and the Wallowa-Whitman National Forest in northeastern Oregon. Swifts occurred in 41 percent of the old-growth stands, but only 8 percent of the logged stands had swifts. The number of swifts was significantly higher in old-growth stands than in logged stands. All swift nests were in large, hollow grand firs with an average diameter of 83 cm.

Bull, E. L., and H. D. Cooper. 1991. Vaux's swift nests in hollow trees. Western Birds 22:85-91.

The authors located Vaux's swift nests in northeastern Oregon. Eighty-five percent of the nest trees were in old-growth forest. All nests were in large-diameter live or dead grand fir (45-96 cm dbh). Trees had been hollowed out by Indian paint fungus.

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Black-Backed Woodpecker
Picoides arcticus

Marshall, D. B. 1992. Status of the black-backed woodpecker in Oregon and Washington. Portland, Oregon: Audubon Society of Portland.

The author's report summarizes information on the black-backed woodpecker in Oregon and Washington from research in the Wallowa-Whitman and Deschutes National Forests of Oregon and the Gifford Pinchot and Mt. Hood National Forests of Washington. The birds almost exclusively use mature and overmature forests of lodgepole pine or mixed conifer, foraging on trees that are infested with bark beetles -- their primary food source. Nest cavities are constructed in live or recently dead trees with heart rot. The author believes that this woodpecker will be adversely affected by salvage logging of infested trees and fire-killed trees and by logging of trees before they are old enough to be most susceptible to beetle outbreaks.

Goggans, R., R. D. Dixon, and L. C. Seminara. 1989. Habitat use by three-toed and black-backed woodpeckers, Deschutes National Forest, Oregon. Non-game project number 87-3-02. Oregon Department of Fish and Wildlife.

The authors conducted their study in the Deschutes National Forest, on the east slope of the Cascade Mountains in Oregon. This forest had experienced a bark beetle epidemic about ten years earlier and the Forest Service had instituted an intensive pest management program of salvage logging and thinning. Habitat use by the black-backed woodpecker in lodgepole pine forest was investigated. Mature and overmature stands were selected for home ranges, foraging and roosting; younger forest stands and logged areas were avoided. The majority of foraging observations was on dead lodgepole pine trees, infested with mountain pine beetles. Nests were excavated in live and dead lodgepole pine trees with heartrot.

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Three-Toed Woodpecker
Picoides tridactylus

Goggans, R., R. D. Dixon, and L. C. Seminara. 1989. Habitat use by three-toed and black-backed woodpeckers, Deschutes National Forest, Oregon. Non-game project number 87-3-02. Oregon Department of Fish and Wildlife.

The authors conducted their study in the Deschutes National Forest, on the east slope of the Cascade Mountains in Oregon. This forest had experienced a bark beetle epidemic about ten years earlier and the Forest Service had instituted an intensive pest management program of salvage logging and thinning. Habitat use by the three-toed woodpecker was investigated in lodgepole pine forest and in mixed conifer. Mature and overmature forest stands were selected for home ranges, for foraging, and for roosting, while younger stands and logged areas were avoided. Most foraging occurred in dead trees. Nests occurred in logged and unlogged areas proportionally to their availability. Nests were primarily in dead trees; all nests were excavated in lodgepole pine with heartrot.

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Red Crossbill
Loxia curvirostra

Holimon, W. C., C. W. Benkman, and M. F. Willson. 1998. The importance of mature conifers to red crossbills in southeast Alaska. Forest Ecology and Management 102:167-72.

Surveys for red crossbills (Loxia curvirostra), which specialize in feeding on conifer seeds, and surveys of tree cone production were conducted in mature (> 150 years old) and young (20-30 years old) stands in southeast Alaska. Red crossbills foraged only in the mature stands. Cone production increased with tree size and was greater in mature stands than in young stands.

Benkman, C. W. 1993. Logging, conifers, and the conservation of crossbills. Conservation Biology 7:473-79.

A review of the literature on red crossbills (Loxia curvirostra) and the impact of logging on production of cones and seeds, their primary food source. Cone and seed production is highest and most consistent in mature and older forests. Logging, by increasing the preponderance of younger trees, greatly reduces cone and seed production. Studies reviewed on crossbill abundance show that red crossbills were more abundant in older than in younger forests. During a year of poor seed crops in Washington, crossbills were thirty times more abundant in an old-growth forest compared to a younger forest (65-140 years old).

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Marbled Murrelet
Brachyramphus marmoratus

U.S. Fish and Wildlife Service. 1997. Recovery plan for the threatened marbled murrelet (Brachyramphus marmoratus) in Washington, Oregon, and California. Portland, Oregon.

This recovery plan was prepared after federal listing in 1992 of the marbled murrelet as a threatened species in California, Washington, and Oregon. The primary reason given for its listing is the loss of old-forest nesting habitat through heavy logging and degradation of habitat through fragmentation. The recovery plan summarizes research findings on murrelet use of old-growth and mature forests for nesting.

deGange, A. R. 1996. A conservation assessment for the marbled murrelet in southeast Alaska. General technical report, PNW-GTR-388. Portland, Oregon: USDA Forest Service, Pacific Northwest Research Station.

This conservation assessment was prepared in conjunction with the Tongass National Forest land management plan revision. The author summarizes available information on the murrelet in southeast Alaska, most of which was in the form of unpublished data and involved extensive personal communication with various scientists. Given the evidence available, the murrelet is concluded to be reliant on structurally complex old-growth forest for nesting and extremely vulnerable to logging of these stands. All nests found, both ground and tree nests, were in old growth.

Ralph, C. J., G. L. Hunt, Jr., M. G. Raphael, J. F. Piatt, tech. eds. 1995. Ecology and conservation of the marbled murrelet. General technical report, PSW-GTR-152. Albany, California: USDA Forest Service, Pacific Southwest Research Station.

s report was compiled by the Marbled Murrelet Conservation Assessment Team, including both scientists and managers. Chapters include original research as well as literature reviews on the marbled murrelet's biology, habitat, and conservation needs in Washington, Oregon, California, British Columbia, and Alaska. The majority of nesting sites are reported to be in late-successional and old-growth forests within 60 km of the coast. All nests are in old-growth trees, usually greater than 81 cm in diameter, on large limbs. The editors of this report speculate that old-growth stands may provide lower temperatures and more protection from weather and predation, and they also provide the stand structure and large trees needed for nesting. They conclude that the fate of the marbled murrelet is tied to the fate of its reproductive habitat -- old-growth forest and forest with older trees.

Hamer, T. E., and S. K. Nelson. 1995. Characteristics of marbled murrelet nest trees and nesting stands. In Ecology and conservation of the marbled murrelet, edited by J. C. Ralph, G. L. Hunt, Jr., M. G. Raphael, and J. F. Piatt. General technical report, PSW-GTR-152. Albany, California: USDA Forest Service, Pacific Southwest Research Station.

The authors summarize information from field biologists on nest and nesting stand characteristics of marbled murrelet nests located between 1974 and 1993 in California, Oregon, Washington, and British Columbia. All nests were in low-elevation old-growth or mature coniferous forests, in stands with multi-layered canopies. Nests were usually in the largest trees available in a stand, with a mean nest tree diameter of 211 cm and a mean nest branch diameter of 32 cm.

Nelson, S. K. and T. E. Hamer. 1995. Nest success and the effects of predation on marbled murrelets. In Ecology and conservation of the marbled murrelet, edited by J. C. Ralph, G. L. Hunt, Jr., M. G. Raphael, and J. F. Piatt. General technical report, PSW-GTR-152. Albany, California: USDA Forest Service, Pacific Southwest Research Station.

Records on marbled murrelet nest success and failure were analyzed to determine patterns of survival in the Pacific Northwest and Alaska. Predation was the major cause of nest failure. Successful nests were significantly farther from edges and had better concealment (through canopy cover) than unsuccessful nests. Although limited data were available, the authors suggest that increased forest edge because of roads or clearcuts could result in increased nest predation rates and decreased reproductive success.

Hamer, T. E. 1995. Inland habitat associations of marbled murrelets in western Washington. In Ecology and conservation of the marbled murrelet, edited by J. C. Ralph, G. L. Hunt, Jr., M. G. Raphael, and J. F. Piatt. General technical report, PSW-GTR-152. Albany, California: USDA Forest Service, Pacific Southwest Research Station.

Murrelet surveys and vegetational data from old-growth stands in western Washington were used to characterize nesting stands. The author's model indicated that the mean diameter of western hemlock and the stem density of dominant trees (> 81 cm dbh) were important predictors of the probability of occupancy of an old-growth stand by a marbled murrelet.

Raphael, M. G., J. A. Young, and B. M. Galleher. 1995. A landscape-level analysis of marbled murrelet habitat in western Washington. In Ecology and conservation of the marbled murrelet, edited by J. C. Ralph, G. L. Hunt, Jr., M. G. Raphael, and J. F. Piatt. General technical report, PSW-GTR-152. Albany, California: USDA Forest Service, Pacific Southwest Research Station.

Characteristics of murrelet nesting habitat at the landscape scale were analyzed using murrelet survey information from the Washington Department of Fish and Wildlife. Sites occupied had higher proportions of old growth and large sawtimber than sites where murrelets were not detected. Occupied sites had greater contiguous cover, with murrelets occurring in the larger patch sizes of old-growth and mature timber available.

Grenier, J. J., and S. K. Nelson. 1995. Marbled murrelet habitat associations in Oregon. In Ecology and conservation of the marbled murrelet, edited by J. C. Ralph, G. L. Hunt, Jr., M. G. Raphael, and J. F. Piatt. General technical report, PSW-GTR-152. Albany, California: USDA Forest Service, Pacific Southwest Research Station.

A murrelet survey was conducted in the Coast Range and Klamath/Siskiyou Mountains of Oregon. Murrelets used mature or old-growth forests. Sites occupied by marbled murrelets were older, had larger midstory trees, and included larger and greater densities of dominant trees than random sites. Nests were in large-diameter Douglas-fir, Sitka spruce, and western hemlock trees. All nests were on large limbs, located in trees greater than 127 cm in diameter.

Miller, S. L., and C. J. Ralph. 1995. Relationships of marbled murrelets with habitat characteristics at inland sites in California. In Ecology and conservation of the marbled murrelet, edited by J. C. Ralph, G. L. Hunt, Jr., M. G. Raphael, and J. F. Piatt. General technical report, PSW-GTR-152. Albany, California: USDA Forest Service, Pacific Southwest Research Station.

The authors surveyed for marbled murrelets in old-growth stands of California. The density of old-growth trees and the dominance of redwood trees were the most important factors positively related to murrelet occupation of stands.

Mendenhall, V. M. 1992. Distribution, breeding records, and conservation problems of the marbled murrelet in Alaska. In Status and Conservation of the Marbled Murrelet in North America, edited by H. R. Carter and M. L. Morrison. Proceedings of the Western Foundation of Vertebrate Zoology, 5:5-16.

The author summarizes published and unpublished observations of marbled murrelets in Alaska. The highest numbers were concentrated in southeastern Alaska and coincided with the largest areas of old-growth forest.

Paton, P. W., C. J. Ralph, and R. A. Erickson. 1992. Use of an inland site in northwestern California by marbled murrelets. In Status and Conservation of the Marbled Murrelet in North America, edited by H. R. Carter and M. L. Morrison. Proceedings of the Western Foundation of Vertebrate Zoology, 5:109-116.

Marbled murrelet activity was censused at stations in old-growth redwood, partially logged stands, and clearcut stands at Redwood Experimental Forest in northwestern California. During the breeding season, murrelets were detected more often in old-growth stands than in partially logged stands or in clearcuts.

Carter, H. R., and S. G. Sealy. 1987. Inland records of downy young and fledgling marbled murrelets in North America. The Murrelet 68.

The authors compiled records on observations of downy young and fledglings of the marbled murrelet in inland habitat along the western coast. The majority of downy young observations (80 percent) were from old-growth forest. The authors hypothesize that since downy young are not usually mobile, the birds must have been in or near their nests. From the fledgling records, it was found that more than half (20 out of 31) were in old growth. The remaining downy young and fledglings were recorded near old-growth forest.

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Northern Goshawk
Accipiter gentiles

Squires, J. R., and L. F. Ruggiero. 1996. Nest-site preference of northern goshawks in southcentral Wyoming. Journal of Wildlife Management 60:170-77.

The authors investigated the habitat preferences of goshawks nesting in the lodgepole pine forests of Medicine Bow National Forest, Wyoming. Goshawks showed a preference for mature lodgepole pine stands. They nested in larger and taller trees with greater canopy height compared to trees available. Areas around nest trees had a higher density of large trees, greater basal area, and taller trees than random sites.

Iverson, G. C., G. D. Hayward, K. Titus, E. DeGayner, R. E. Lowell, D. C. Crocker-Bedford, P. F. Schempf, and J. Lindell. 1996. Conservation assessment for the northern goshawk in southeast Alaska. General technical report, PNW-GTR-387. Portland, Oregon: USDA Forest Service, Pacific Northwest Research Station.

A summary of the information available on northern goshawks (A. gentilis laingi) and their habitat use in the Tongass National Forest in southeast Alaska. Research showed that goshawks preferred old-growth forest and avoided clearcuts, nonforest, and alpine cover types. Nest trees were primarily located in old-growth forest stands. Studies of 30-acre plots around the nest tree showed that forest cover was around 98 percent at nest sites. Most of the principal prey species also occurred at higher densities in productive old-growth forest.

Crocker-Bedford, D. C. 1995. Northern goshawk reproduction relative to selection harvest in Arizona. Abstract in Journal of Raptor Research 29:42-43.

The author compared goshawks' occupancy of nest clusters to the degree of selective logging in their home range that had occurred in prior years. Nest occupancy rates declined as the percent area with selective logging increased.

Hargis, C. D., C. McCarthy, and R. D. Perloff. 1994. Home ranges and habitats of northern goshawks in eastern California. Studies in Avian Biology 16:66-74.

Goshawk home ranges, including nest sites and areas used after young had fledged, were studied using radiotelemetry in the Inyo National Forest in the eastern Sierra Nevada. Nest sites and perching sites were in locations with greater basal area and greater canopy cover than the average available. Nests were in large-diameter trees with a mean dbh of 87.2 cm, and within the largest-diameter class of trees in the study.

Woodbridge, B., and P. J. Detrich. 1994. Territory occupancy and habitat patch size of northern goshawks in the southern Cascades of California. Studies in Avian Biology 16:83-87.

The authors surveyed for goshawk territories over a number of years in part of the Klamath National Forest in the southern Cascades. The area had experienced extensive logging and forest fragmentation and consisted of mature unlogged forest interspersed with younger successional stages. Goshawk territories were mostly associated with the larger remaining patches of mature forest, and territory occupancy was higher in larger stands compared to smaller stands.

Bosakowski, T., and R. Speiser. 1994. Macrohabitat selection by nesting northern goshawks: Implications for managing eastern forests. Studies in Avian Biology 16:46-49.

Goshawk choice of macrohabitat (landscape characteristics) was analyzed using nest site data from the New York-New Jersey Highlands. The authors' model indicated that the distance to paved roads and to human habitation were important variables in habitat selection and that goshawks preferred remote areas and large blocks of forested habitat.

Crocker-Bedford, D. C. 1990. Goshawk reproduction and forest management. Wildlife Society Bulletin 18:262-69.

In the Kaibab Plateau of Arizona, the author compared goshawks' occupancy and re-occupancy of nest trees and nesting stands in large unlogged stands to their occupancy of logged stands with varying sizes of nest-tree buffer zones. Re-use of nests was highest in the unlogged territories. In logged territories, occupancy of nesting stands dropped by 80 percent, even with large buffers. In addition, open-forest raptor species replaced goshawks in most logged territories but not in control areas.

Hayward, G. D., and R. E. Escano. 1989. Goshawk nest-site characteristics in western Montana and northern Idaho. Condor 91:476-79.

The authors described goshawk nest sites in two climate zones of the northern Rockies -- the Columbian Highlands and Rocky Mountain forest. The majority of nest sites were in old or mature forest with closed canopies. Nest trees were large, dominant or codominant trees in the stand.

Crocker-Bedford, D. C., and B. Chaney. 1988. Characteristics of goshawk nesting stands. In Proceedings of the southwest raptor management symposium and workshop, edited by R. L. Glinski, et al. Washington, DC: National Wildlife Federation.

Northern goshawk nesting habitat was examined in northern Arizona. Goshawks nested in the densest stands available. They did not nest in stands with less than 60 percent canopy cover. Most individual goshawks preferred stands with more than 80 percent canopy cover.

Speiser, R., and T. Bosakowski. 1987. Nest site selection by northern goshawks in northern New Jersey and southeastern New York. Condor 89:387-94.

Goshawk nest sites were compared to randomly selected forest plots in the New York-New Jersey Highlands. The majority of goshawk nests were in stands of mature or old-growth forest in large tracts of forest. Nest sites had more large trees and greater basal area than random sites. Nest trees selected by the goshawks were large in diameter (mean diameter, 38.3 cm) compared to those available.

Reynolds, R. T., E. C. Meslow, and H. M. Wight. 1982. Nesting habitat of coexisting Accipiter in Oregon. Journal of Wildlife Management 46:124-38.

Different forest types throughout Oregon were surveyed for nests of three Accipiter species. Northern goshawks nested in dense stands of mature or old-growth conifers rather than in younger, second-growth stands. Nests were located on large limbs or at a branch, primarily in ponderosa pine and Douglas fir, with a mean tree diameter of 82.3 cm dbh.

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Flammulated Owls
Otus flammeolus

Linkhart, B. D., R. T. Reynolds, and R. A. Ryder. 1998. Home range and habitat of breeding flammulated owls in Colorado. Wilson Bulletin 110:342-51.

This paper presents data from radiotracking flammulated owls as part of the authors' long-term study in central Colorado's Manitou Experimental Forest. Owls established home ranges each spring in areas with more old ponderosa pine/Douglas-fir forest (200-400 years old) than other overstory types available. Males preferred foraging in old ponderosa pine/Douglas-fir forest and used large, old trees for foraging, territorial singing, and day-roosting.

Linkhart, B. D., and R. T. Reynolds. 1997. Territories of flammulated owls (Otus flammeolus): is occupancy a measure of habitat quality? In Biology and conservation of owls of the Northern Hemisphere, edited by J.R. Duncan, D. H. Johnson, and T. H. Nichols. General technical report, NC-190. USDA Forest Service.

Part of the authors' long-term study on flammulated owls in central Colorado, this paper presents data from sixteen years of research on territory preference. Occupancy of territories by breeding pairs was positively correlated with the amount of old (200-400 years) ponderosa pine/Douglas fir present in the territory (as compared to mature quaking aspen/blue spruce, mature quaking aspen, and young Douglas-fir/blue spruce stands).

Powers, L. R., A. Dale, P. A. Gaede, C. Rodes, L. Nelson, J. J. Dean, and J. D. May. 1996. Nesting and food habits of the flammulated owl (Otus flammeolus) in southcentral Idaho. Journal of Raptor Research 30:15-20.

The authors documented nesting and foraging activities of flammulated owls during four nesting seasons in the Sublett Mountains of Idaho. Owls nested in cavities in Douglas fir and trembling aspen; 83 percent of nest cavities were in dead trees and 17 percent in live trees. The mean diameter of nest trees was 49.9 cm.

McCallum, D. A. 1994. Conservation status of flammulated owls in the United States. In Flammulated, boreal, and great gray owls in the United States: A technical conservation assessment, edited by G. D. Hayward and J. Verner. General technical report, RM-253. Fort Collins, Colorado: USDA Forest Service, Rocky Mountain Forest and Range Experiment Station.

The author evaluates the conservation status of the flammulated owl based on the science available and presented earlier in this technical report. He notes that the flammulated owl is a habitat specialist with low fertility and a preference for the structure provided by old forests; that research suggests it is sensitive to habitat change; and that it is likely that population numbers have decreased due to loss of habitat from logging, fire, and stand- type conversions. He notes that data for proposed benefits of restoration thinning are lacking.

Reynolds, R. T., and B. D. Linkhart. 1992. Flammulated owls in ponderosa pine: Evidence of preference for old growth. In Old-growth forests in the Southwest and Rocky Mountain regions, edited by M. R. Kaufmann, W. H. Moir, and R. L. Bassett. General technical report, RM-213. Fort Collins, Colorado: USDA Forest Service, Rocky Mountain Forest and Range Experiment Station.

This paper is a summary of Reynolds and Linkhart findings as presented in their other papers. In addition, song trees are described: males sang from "hidden positions next to tree trunks or in dense clumps of foliage."

Bull, E. L., A. L. Wright, and M. G. Henjum. 1990. Nesting habitat of flammulated owls in Oregon. Journal of Raptor Research 24:52-55.

The authors characterized nesting trees and nesting sites in northeastern Oregon's Starkey Experimental Forest. Nests were in cavities in large-diameter dead trees -- 91 percent of nests were in dead trees, with average dbh of nest trees, 72 cm. Owls showed a preference for cavities excavated by pileated woodpeckers. Owls preferred nesting in stands with large-diameter (> 50 cm dbh) ponderosa pine and Douglas fir and in stands of grand fir with ponderosa pine in the overstory.

Reynolds, R. T., and B. T. Linkhart. 1987. The nesting biology of flammulated owls in Colorado. In Biology and conservation of northern forest owls, edited by R. W. Nero, R. J. Clark, R. J. Knapton, and R. H. Hamre. General technical report, RM-142. Fort Collins, Colorado: USDA Forest Service, Rocky Mountain Forest and Range Experiment Station.

Part of the authors' long-term study on flammulated owls in central Colorado, this paper presents data from 1980-1986 on habitat use by nesting owls. Owls used tree cavities for nesting. The majority of territories were in old-growth ponderosa pine mixed with Douglas fir. Two territories also included mature quaking aspen-blue spruce stands and mature quaking aspen stands. Males, the primary food providers during the breeding season, concentrated their foraging in old-growth ponderosa pine/Douglas-fir forest.

Marcot, B. G., and R. Hill. 1980. Flammulated owls in northwestern California. Western Birds 11:141-49.

A survey for flammulated owls was conducted in northwestern California, mostly in Six Rivers National Forest. Territories included California black oak and yellow pine trees and a dense group of tall, mature trees. The owls were found calling from the dense foliage of mature trees.

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Forest Interior Songbirds

Ortega, Y. K., and D. E. Capen. 1999. Effects of forest roads on habitat quality for ovenbirds in a forested landscape. Auk 116:937-46.

The authors studied the influence of forest roads on ovenbird (Seiurus aurocapillus) densities in Green Mountain National Forest, Vermont. Territory densities were 40 percent lower in edge areas (extending 150 m from roads) than in forest interior areas. Pairing success had a lower trend within edge areas than interior areas.

Porneluzi, P. A., and J. Faaborg. 1999. Season-long fecundity, survival, and viability of ovenbirds in fragmented and unfragmented landscapes. Conservation Biology 13:1151-61.

The authors compared the breeding success of ovenbirds in fragmented forest versus unfragmented forest in Missouri. Reproductive success was higher in the continuous forest, with only 4 percent of nests parasitized compared to 72 percent in the fragmented landscape. Average density of male birds was lower in the fragmented landscape. Territorial males had lower densities near forest edges (within 100 m of an edge) than away from the edge (200 m and beyond).

Roberts, C., and C. J. Norment. 1999. Effects of plot size and habitat characteristics on breeding success of scarlet tanagers. Auk 116:73-82.

The breeding success of scarlet tanagers (Piranga olivacea) was studied in western New York. Tanagers were absent from all forest patches smaller than ten hectares. Fledging success increased with forest patch size and was highest in continuous forest.

Haney, J. C. 1999. Hierarchical comparisons of breeding birds in old-growth conifer-hardwood forest on the Appalachian Plateau. Wilson Bulletin 111:89-99.

The author compared relative abundances of breeding birds in old-growth forest (> 300 years old) on the northern Appalachian Plateau, Pennsylvania. Of fifty-six bird species recorded, about one-third were more likely to occur in old-growth conifer-hardwood forest than in the broader landscape (province and state level). This included some of the state's rarest breeding birds such as the yellow-bellied flycatcher and Swainson's thrush.

Hanners, L. A., and S. R. Patton. 1998. Worm-eating warbler (Helmitheros vermivorus). In The birds of North America, No. 367, edited by A. Poole and F. Gills. Philadelphia: The Academy of Natural Sciences; Washington, DC: The American Ornithologists' Union.

This is a summary of information available for the worm-eating warbler (Helmitheros vermivorus), a forest songbird that breeds in eastern North America. Densities of breeding birds are highest in large, continuous forest with little edge.

Gale, G. A., L. A. Hanners, and S. R. Patton. 1997. Reproductive success of worm-eating warblers in a forested landscape. Conservation Biology 11:246-50.

The authors compared the breeding success of worm-eating warblers in a large, unfragmented forest (750 ha) to their success in small forest fragments (< 56 ha) in southwestern Connecticut. The density of males on the large site was, on average, seven times higher than in the forest fragments.

Van Horn, M. A., and T. M. Donovan. 1994. Ovenbird (Seiurus aurocapillus). In The birds of North America, No. 88, edited by A. Poole and F. Gills. Philadelphia: The Academy of Natural Sciences; Washington, DC: The American Ornithologists' Union.

A summary of research on the ovenbird, a forest songbird. This warbler requires large, continuous forest for successful breeding. Male pairing success is lower in smaller forest tracts. Rates of nest predation and parasitism by the brown-headed cowbird are higher near forest edges.

Wenny, D. G., R. L. Clawson, J. Faaborg, and S. L. Sheriff. 1993. Population density, habitat selection and minimum area requirements of three forest-interior warblers in central Missouri. Condor 95:968-79.

Population densities of ovenbirds, Kentucky warblers (Oporornis formosus), and worm-eating warblers in small forest patches (300 ha) were compared to densities in a large, continuous forest. The study was located in Missouri, one site being in the Mark Twain National Forest. Breeding worm-eating warblers were only located in the large forest. Average densities of ovenbirds and Kentucky warblers were significantly higher in the large forest tract than in the smaller forests. Ovenbirds had higher densities away from the forest edge (200 m beyond) than near the edge (within 100 m).

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Other Avian Species

McIver, J. D., and L. Starr, tech. eds. 2000. Environmental effects of postfire logging: Literature review and annotated bibliography. General technical report, PNW-GTR-486. USDA Forest Service, Pacific Northwest Research Station.

The authors review the literature available on logging after wildfire. They note that there are no studies documenting a reduction in fire intensity in stands that have been logged postfire. The authors also note that the possibility of mitigating soil loss and erosion through logging (by disrupting water-repellant layers, for instance) has not been tested experimentally. Studies with cavity-nesters in the intermountain West have shown that most species decreased in abundance after post-fire logging, including the black-backed, hairy, and three-toed woodpeckers.

Bull, E. L., C. G. Parks, and T. T. Torgersen. 1997. Trees and logs important to wildlife in the Interior Columbia River Basin. General technical report, PNW-GTR-391. Portland, Oregon: USDA Forest Service, Pacific Northwest Research Station.

This report includes a review of the species that depend on snags, hollow trees, downed wood, and decayed living trees in the Interior Columbia River Basin. In terms of hollow trees, large-diameter hollow trees provide the best habitat; the authors report that these are typically found only in late- and old-seral stands of grand fir and western red cedar and are uncommon in managed landscapes. For snags, the authors note that large-diameter snags provide nest habitat for the greatest variety of cavity nesters and that these remain standing longer than small-diameter snags. For logs, the authors report that logs 15 inches or greater in diameter are particularly important for wildlife species such as pileated woodpeckers.

Dellasala, D., J. A. Hagar, K. A. Engel, W. C. McComb, R. L. Fairbanks, and E. G. Campbell. 1996. Effects of silvicultural modifications of temperate rainforest on breeding and wintering bird communities, Prince of Wales Island, southeast Alaska. Condor 98: 706-21.

The authors inventoried breeding bird species and wintering bird species in virgin old growth and in three treatments of 20-year-old young growth that had originated from clearcuts. The study was conducted on Prince of Wales Island in southeast Alaska. During the breeding season, abundance of three of the common breeding bird species -- golden-crowned kinglet (Regulus satrapa), Pacific-slope flycatcher (Empidonax difficilis), and red-breasted sapsucker (Sphyrapicus ruber) -- was significantly greater (6-14 times greater) in old growth than in young growth. During the wintering season, species richness was greater in old growth. Golden-crowned kinglets were significantly more abundant in old growth than in young growth. Brown creepers (Certhia americana) occurred only in old growth during both the breeding and the wintering season.

Newton, I. 1994. The role of nest sites in limiting the numbers of hole-nesting birds: A review. Biological Conservation 70:265-76.

The author reviews the factors influencing the availability of tree cavities for nesting and the limitations on populations of cavity-nesting bird species. The studies reviewed showed that the number of usable cavities in a forest increased with the age of the trees and that the abundance and diversity of cavity nesters increased as the forest aged and more dead wood/snags were available.

Sakai, H. F. and B. R. Noon. 1991. Nest-site characteristics of Hammond's and Pacific-slope flycatchers in northwestern California. Condor 93:563-74.

Nesting by Hammond's flycatchers (Empidonax hammondii) and Pacific-slope flycatchers (E. difficilis) was compared in young, mature, and old-growth Douglas-fir/tan oak forest stands in northwestern California. Hammond's flycatchers occurred and nested only in mature and old-growth forest (stands older than 90 years). Within these stands, they used nest trees of significantly larger diameter (mean nest-tree diameter of 104.2 cm) than what was available. Within old growth, they also selected for areas with more large (50-100 cm) tan oaks, fewer small tan oaks, and higher canopy bole height.

Tobalske, B. W., R. C. Shearer, and R. L. Hutto. 1991. Bird populations in logged and unlogged western larch/Douglas-fir forest in northwestern Montana. Research paper, INT-442. Ogden, Utah: USDA Forest Service, Intermountain Research Station.

Population sizes of breeding birds were compared in logged and unlogged forest stands in the Flathead National Forest, northwestern Montana. When types of foraging behavior were compared, foliage foragers and tree gleaners were less abundant in logged areas compared to unlogged areas, as were conifer tree nesters. Species significantly less abundant in the logged areas were: golden-crowned kinglets (Regulus satrapa), Swainson's thrushes (Catharus ustulatus), varied thrushes (Ixoreus naevius), and Townsend's warblers (Dendroica townsendi). In addition, varied thrush populations were significantly higher in the larger unlogged forest compared to the unlogged remnants, indicating an adverse impact of fragmentation.

Mariani, A. M., and D. A. Manuwal. 1990. Factors influencing brown creeper (Certhia americana) abundance patterns in the southern Washington Cascade range. Studies in Avian Biology 13:53-57.

The authors compared brown creeper abundance and arthropod abundance in young stands (65-80 years old), mature stands (105-130 years old), and old-growth forest (375 years old). Brown creeper abundance was positively correlated with the abundance of large Douglas-fir trees (> 100 cm dbh). Larger trees had deeper bark furrows and more arthropod prey.

Manuwal, D. A., and M. H. Huff. 1987. Spring and winter bird populations in a Douglas-fir forest sere. Journal of Wildlife Management 51:586-95.

Bird communities were compared among young, mature, and old-growth (250-500+ years) stands in Douglas-fir forests of the southern Washington Cascades. Old growth provided better winter habitat than younger forest -- bird species richness, diversity, and abundance were all higher in old growth during winter. The brown creeper (Certhia americana), red crossbill (Loxia curvirostra), red-breasted nuthatch (Sitta canadensis), and gray jay (Perisoreus canadensis) were all significantly more abundant in old-growth than in young stands.

Kessler, W. B., and T. E. Kogut. 1985. Habitat orientations of forest birds in southeastern Alaska. Northwest Science 59:58-65.

A range of forest successional stages, including old growth, recent clearcuts, sapling/shrub stages, and pole stages, were surveyed for bird species. All surveys were conducted on Kosciusko and Prince of Wales Islands in southeast Alaska. Townsend's warbler, golden-crowned kinglet, western flycatcher, Swainson's thrush, and pine siskin occurred in old growth but were absent or rare in young stands.

Mannan, R. W., and E. C. Meslow. 1984. Bird populations and vegetation characteristics in managed and old-growth forests, northeastern Oregon. Journal of Wildlife Management 48:1219-38.

The authors' objectives were to compare structural components of older managed forests (85 years old) with old growth (greater than 200 years old) and to compare bird populations in each. The study was conducted in the Wallowa-Whitman National Forest in northeastern Oregon. Mean densities of large trees, large snags, and tree height diversity (multiple canopy layers) were greater in old-growth forests. There were relatively higher numbers of cavity-nesting bird species in the old growth than the managed forest; the authors attribute this to the greater abundance of large snags in old growth.

Raphael, M. G., and M. White. 1984. Use of snags by cavity-nesting birds in the Sierra Nevada. Wildlife Monographs 86:1-66.

The authors studied nesting and foraging habitat selection by eighteen cavity-nesting bird species on the eastern side of the Sierra Nevada. Snags rather than live trees were clearly preferred as nests, with 72 percent of nests situated in snags even though only 7 percent of the available trees were snags. Compared to available trees, species preferred large-diameter trees for nesting (mean diameter, 62 cm), and they preferred sites surrounded by a larger number of snags. Density of cavity-nesting birds increased in proportion to snag density and was strongly correlated with the density of large snags (> 38 cm dbh).

Doerr, J. G., C. L. Barescu, J. M. Brighenti, Jr., and M. P. Morin. 1984. Use of clearcutting and old-growth forests by male blue grouse in central southeast Alaska. In Fish and wildlife relationships in old-growth forests, edited by W. R. Meehan, T. R. Merrell, Jr., and T. A. Hanley. Morehead City, North Carolina: American Institute of Fishery Research Biologists.

To determine preferred breeding habitat, singing male blue grouse (Dendragapus obscurus) were located on Kuiu Island, on Mitkof Island, and near Thomas Bay in southeastern Alaska. Densities of singing males were forty-five times higher in old-growth forests than in clearcuts (1-23 years old), indicating a strong preference for old-growth forest.

Lebeda, C. S., and J. T. Ratti. 1983. Reproductive biology of Vancouver Canada geese on Admiralty Island, Alaska. Journal of Wildlife Management 47:297-306.

The nesting and brood-rearing biology of Vancouver Canada geese (Branta canadensis fulva), a subspecies of Canada geese, was studied in Admiralty Island, southeast Alaska. Various habitat types -- including spruce/hemlock forest, forest edge, grass/sedge meadows, muskeg, riparian zones, and lake shorelines -- were searched for nests. The majority of nests (22 out of 26) were in the forest. Geese with broods used the forest interior as escape cover.

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AMPHIBIANS

Lydic, J. 1999. Populations of salamanders within an old and secondary growth mesic cove forest with reference to coarse woody debris. Master's thesis, Edinboro University of Pennsylvania.

The author compared salamander abundance and distribution in old-growth forest and secondary forest (60-70 years old) of the southern Appalachians. The total number of salamanders encountered was higher in old growth than in secondary growth. Numbers of Plethodon jordani, Desmognathus wrighti, and Desmognathus santeetlah were significantly higher in the old-growth forest. Higher numbers of D. wrighti in a reproductive state occurred in the old growth than in the secondary forest, indicating better environmental conditions.

Grover, M. C. 1998. Influence of cover and moisture on abundances of the terrestrial salamanders Plethodon cinereus and Plethodon glutinosus. Journal of Herpetology 32:489-97.

The author studied two species of terrestrial salamanders -- red-backed salamanders (Plethodon cinereus) and slimy salamanders (P. glutinosus) -- in the southern Appalachians, southwestern Virginia. Their requirement for moist environments and retreat/foraging sites was investigated by manipulating moisture levels and the density of cover objects. The abundance of both species increased with higher moisture levels and greater cover density. Activity levels and body mass also increased.

Welsh, H. H., Jr., and A. J. Lind. 1995. Habitat correlates of the Del Norte salamander, Plethodon elongatus (Caudata: Plethodontidae), in northwestern California. Journal of Herpetology 29:198-210.

Habitat relationships of Del Norte salamanders, a species of "special concern" in California, were sampled in clearcuts, young forest (31-99), mature forest (100-199 years old), and old-growth forest (200 + years old) in northwestern California. Del Norte salamanders were associated with older forests (> 140 years), with a closed, multi-storied canopy of conifers and hardwoods, and with cool, moist microclimates.

Petranka, J. W., M. P. Brannon, M. E. Hopey, and C. K. Smith. 1994. Effects of timber harvesting on low-elevation populations of southern Appalachian salamanders. Forest Ecology and Management 67:135-47.

Forest stands of different ages, from clearcuts less than 5 years old to mature forest stands more than 120 years old, were sampled for salamander abundance and diversity. The study was conducted in Pisgah National Forest, North Carolina. Salamander abundance and species richness was greatest in mature stands and lowest in very young stands. Lungless salamanders (Plethodon spp.) were most abundant in stands greater than 120 years old and were rare or absent in younger stands. Numbers of Desmognathus spp. were highest in stands greater than 80 years old.

Walls, S. C., A. R. Blaustein, and J. J. Beatty. 1992. Amphibian biodiversity of the Pacific Northwest with special reference to old-growth stands. Northwest Environmental Journal: 53-69.

The authors reviewed amphibian biodiversity in the Pacific Northwest (second highest in the United States) and identified species specifically associated with old-growth stands. Six species of terrestrial salamanders (lungless salamanders) were found to inhabit the forest floor of old-growth stands: Ensatina eschscholtzii, Plethodon vehiculum, P. elongatus, P. vandykei, Aneides ferreus, and Batrachoseps wrighti. Their habitat includes leaf litter, logs, and loose bark. A number of aquatic breeding species, including Pacific giant salamanders (Dicamptodon ensatus), inhabit the forest floor during their terrestrial adult stage. Logging, with the associated drying out of the forest floor and loss of woody debris, is expected to adversely impact amphibian habitat.

Aubry, K. B., and P. A. Hall. 1991. Terrestrial amphibian communities in the southern Washington Cascade range. In Wildlife and vegetation of unmanaged Douglas-fir forests, edited by L. F. Ruggiero, K. B. Aubry, A. B. Carey, and M. H. Huff. General technical report, PNW-GTR-285. Portland, Oregon: USDA Forest Service, Pacific Northwest Research Station.

The authors surveyed amphibian populations in unmanaged stands 55 to 730 years old on the western slope of the southern Cascades in Washington. Northwestern salamanders (Ambystoma gracile) and roughskin newts (Taricha granulosa) were four to five times more abundant in old-growth stands than in younger stands. Cascades frogs (Rana cascadae) were also most abundant in old-growth forest. Ensatinas (Ensatina eschscholtzi), western red-backed salamanders (Plethodon vehiculum), and red-legged frogs (Rana aurora) were closely associated with coarse woody debris, with ensatinas being most abundant in old-growth stands and young, unmanaged stands.

Corn, P. S., and R. B. Bury. 1991. Terrestrial amphibian communities in the Oregon Coast Range. In Wildlife and vegetation of unmanaged Douglas-fir forests, edited by L. F. Ruggiero, K. B. Aubry, A. B. Carey, and M. H. Huff. General technical report, PNW-GTR-285. Portland, Oregon: USDA Forest Service, Pacific Northwest Research Station.

Amphibian populations were surveyed in clearcuts, young, mature, and old-growth forest sites on the Coast Range of Oregon. Northwestern salamanders (Ambystoma gracile) and tailed frogs (Ascaphus truei) were absent from clearcuts. Olympic salamanders (Rhyacotriton olympicus) and clouded salamanders (Aneides ferreus) were captured in significantly higher numbers in old-growth forest. Clouded salamanders and ensatinas were closely associated with down wood, with preferences for different degrees of log decay. Clouded salamanders preferred large Douglas-fir logs with little decay, whereas ensatinas preferred well-decayed logs.

Welsh Jr., H. H. 1990. Relictual amphibians and old-growth forests. Conservation Biology 4:309-19.

Young, mature, and old-growth forest in northwestern California and southwestern Oregon were sampled for amphibians. Ninety-one percent of Del Norte salamanders (Plethodon elongatus) were found on old-growth sites. Olympic salamanders (Rhyacotriton olympicus) were found primarily in mature and old-growth stands rather than in young stands. Tailed frogs (Ascaphus truei) were also associated with older forest and cooler streams.

Bury, R. B. 1983. Differences in amphibian populations in logged and old-growth redwood forest. Northwest Science 57:167-78.

Amphibian populations were compared in California old-growth redwood forest and redwood forest logged 6-15 years earlier. Species composition in the two forest types was clearly different, with forest-dependent species -- Olympic salamanders (Rhyacotriton olympicus), tailed frogs (Ascaphus truei), and Pacific giant salamanders (Dicamptodon ensatus) -- present only in the old-growth plots and not in the logged plots. Ensatinas (Ensatina eschscholtzi) and California slender salamanders (Batrachoseps attenuatus) were present in both forest types, but were more abundant and had higher biomass in the old-growth plots than in the second-growth plots.

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