United States Department of Agriculture

Forest 

Service

Pacific Northwest 

Research Station

General Technical Report

PNW-GTR-909 August 

2016

Annotated Bibliography of the Red

Tree Vole (Arborimus longicaudus),

Sonoma Tree Vole (A. pomo), and 

White-Footed Vole (A. albipes)
In accordance with Federal civil rights law and U.S. Department of Agriculture 

(USDA) civil rights regulations and policies, the USDA, its Agencies, offices, and 

employees, and institutions participating in or administering USDA programs are 

prohibited from discriminating based on race, color, national origin, religion, sex, 

gender identity (including gender expression), sexual orientation, disability, age, 

marital status, family/parental status, income derived from a public assistance 

program, political beliefs, or reprisal or retaliation for prior civil rights activity, in any 

program or activity conducted or funded by USDA (not all bases apply to all programs). Remedies and complaint filing deadlines vary by program or incident. 

Persons with disabilities who require alternative means of communication for 

program information (e.g., Braille, large print, audiotape, American Sign Language, 

etc.) should contact the responsible Agency or USDA’s TARGET Center at (202) 

720-2600 (voice and TTY) or contact USDA through the Federal Relay Service 

at (800) 877-8339. Additionally, program information may be made available in 

languages other than English. 

To file a program discrimination complaint, complete the USDA Program Discrimination Complaint Form, AD-3027, found online at http://www.ascr.usda.gov/complaint_filing_cust.html and at any USDA office or write a letter addressed to USDA 

and provide in the letter all of the information requested in the form. To request a 

copy of the complaint form, call (866) 632-9992. Submit your completed form or 

letter to USDA by: (1) mail: U.S. Department of Agriculture, Office of the Assistant 

Secretary for Civil Rights, 1400 Independence Avenue, SW, Washington, D.C. 

20250-9410; (2) fax: (202) 690-7442; or (3) email: program.intake@usda.gov. 

USDA is an equal opportunity provider, employer, and lender.

Compilers

James K. Swingle is a faculty research assistant, Department of Fisheries and 

Wildlife, Oregon State University, Corvallis, OR 97330 and Eric D. Forsman is 

a research wildlife biologist (emeritus), U.S. Department of Agriculture, Forest 

Service, Pacific Northwest Research Station, Forestry Sciences Laboratory,

3200 SW Jefferson Way, Corvallis, OR 97331.

Cover: Captive tree vole by Eric Forsman
Abstract

Swingle, James K.; Forsman, Eric D., comps. 2016. Annotated bibliography 

of the red tree vole (Arborimus longicaudus), Sonoma tree vole (A. pomo), and 

white-footed vole (A. albipes). Gen. Tech. Rep. PNW-GTR-909. Portland, OR: 

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

Station. 179 p.

This annotated bibliography contains all citations that we could find on tree voles 

(Arborimus longicaudus, A. pomo) and white-footed voles (A. albipes), including 

many unpublished sources in museum archives, court proceedings, and agency 

documents. Some documents not readily available in published form or museum 

archives are available upon request from the compilers.

Keywords: Red tree vole, Sonoma tree vole, white-footed vole, Arborimus 

longicaudus, Arborimus pomo, Arborimus albipes, Arvicolinae, Muridae.
1

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Introduction

Because of their unique adaptations for living in the forest canopy and feeding on 

needles and twigs of conifers, red tree voles (Arborimus longicaudus) and Sonoma 

tree voles (A. pomo) have long intrigued mammalogists. While conducting studies of tree voles, we have come across a wealth of information on these enigmatic 

mammals, much of which is unpublished or buried in obscure sources that are 

not easily obtainable. Because much of this information was difficult to find, we 

synthesized it into a single bibliography with annotations describing the type of 

information presented in each source. We have also included information that we 

found on the closely related white-footed vole (A. albipes), which occupies a geographic range similar to the tree vole’s, and which is also at least partially arboreal. 

Our hope is that this bibliography will be of use to others who are interested in 

these unique animals. 

We included many unpublished references, including field notes and specimen 

catalogs that we found in museums or other locations where they were difficult 

to locate. We found those obscure sources to be extremely helpful in tracing the 

history of research and management of Arborimus voles, and we think that future 

researchers will find those sources useful as well. We have also included many 

published papers on the evolutionary history of arvicoline voles because those 

papers helped us understand the evolutionary history of tree voles and white-footed 

voles, even if those species were not specifically mentioned in the published works. 

We also included a few papers on old-growth forests and conifer needle chemistry 

because we found them useful for understanding the unique relationship between 

tree voles and the structural and chemical attributes of the trees in which they live. 

Because of the long and contentious debate over whether or not to protect tree 

voles on federal lands in Oregon and California, there is much gray literature on 

tree voles that has been produced in federal management reports and court cases. 

The many documents related to court cases are particularly interesting because they 

illustrate the difficulty of finding middle ground in the debate over management of 

forests in which tree voles and many other species of obscure plants and animals 

occur. We included as many of these management documents and court documents 

as we could find, although we undoubtedly overlooked some of them. 

Although we tried to include a short annotation for every citation in the bibliography, we made no attempt to describe all of the details that were presented 

in each source. Our main objective was to provide a complete list of data sources, 

with a brief account of what was in each citation. To help readers find references 

by author or subject area, there is a subject index and author index at the end of the 

bibliography. A few of the unpublished references in the bibliography are in our 
2

GENERAL TECHNICAL REPORT PNW-GTR-909

possession and are not available from other sources; These references are available 

upon request from the compilers.We also included an appendix in which we list 

citations and numbers of tree voles and white-footed voles captured in trapping 

studies in western Oregon and northwestern California. The latter appendix can be 

used to quickly peruse the results from a large number of trapping studies, without 

reading every citation. 

Tree voles and white-footed voles belong to the subfamily Arvicolinae (voles, 

lemmings, and muskrats) in the family Muridae (Musser and Carleton 1993). Other 

subfamilies of murid rodents that occur within the range of the tree voles include 

the Murinae (old world rats and mice) and Sigmodontinae (new world rats and mice; 

Verts and Carraway 1998). Generic and specific taxonomy of tree voles and whitefooted voles has changed over time. Based on the type specimen of the red tree 

vole, which was collected near Coos Bay, Oregon in 1890, True (1890) named the 

species Phenacomys longicaudus. Taylor (1915) proposed that Phenacomys longicaudus and Phenacomys albipes be assigned to a new subgenus (Arborimus). Based 

on a single specimen from Tillamook, Oregon, Howell (1921) concluded that there 

was another species of tree vole in northwest Oregon, which he called P. silvicolus. 

Howell suggested that P. silvicolus might co-occur with P. longicaudus. Miller 

(1924) referred to this new species as the dusky tree vole and shortened the name 

to P. silvicola. In a subsequent monograph on Phenacomys voles, Howell (1926) 

continued to maintain that there were two species of tree voles in northwest Oregon, 

which he referred to as the “Red Tree Mouse” and “Forest Tree Mouse.” Contrary 

to Taylor (1915), Howell (1926) concluded that placement of P. longicaudus and P. 

albipes in the subgenus Arborimus was based on such minor morphological differences that it was unwarranted. This was the last word on taxonomy of tree voles for 

many years, until Johnson (1968) conducted the first blood protein electrophoresis 

studies of tree voles and concluded that P. silvicola should be relegated to a subspecies (P. l. silvicola). In a later paper, Johnson (1973) proposed that Arborimus should 

be elevated to full generic rank and that all named forms of tree voles should be 

included in a single species (A. longicaudus: red tree vole). Johnson and Maser 

(1982) subsequently determined that the genus of the white-footed vole should also 

be changed from Phenacomys to Arborimus. Based on captive breeding studies, 

karyotypic analyses, and morphometric comparisons, Johnson and George (1991) 

proposed that tree voles in California should be placed in a separate species (A. 

pomo), which was later given the common name Sonoma tree vole. Murray (1995) 

subsequently concluded that the dividing line between A. pomo and A. longicaudus

was the Klamath River in northern California. In this bibliography, we will use the 

Klamath River as the dividing line between A. pomo and A. longicaudus, regardless 
3

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

of specific names used in the original papers. Recent genetic and morphological 

studies have detected genetic differences in tree voles in northern and southern 

Oregon but have also found so much overlap that those differences are useless for 

taxonomic differentiation (Miller et al. 2006). Because there is uncertainty regarding the subspecific taxonomy of tree voles, we refer to all tree voles from Oregon as 

A. longicaudus, without regard to subspecific taxonomy. 

1.   Abramson, N.I.; Lebedev, V.S.; Tesakov, A.S.; Bannikova, A.A. 

2009. Supraspecies relationships in the subfamily Arvicolinae (Rodentia, 

Cricetidae): an unexpected result of nuclear gene analysis. Molecular 

Biology. 43: 834–846.

There is only one brief reference to Phenacomys in this paper, but it is a 

useful reference for anyone interested in the evolution of the Arvicolinae.

2.   Adam, M.D.; Hayes, J.P. 1998. Arborimus pomo. Mammalian Species.

593: 1–5.

This is an excellent summary of the distribution, morphology, ontogeny, 

reproduction, ecology, behavior, and genetics of the Sonoma tree vole. We 

found one minor error; the authors incorrectly cited Forsman et al. (1984) 

by stating that tree voles made up 50 percent of the diet of northern spotted 

owls (Strix occidentalis caurina) in Oregon. Forsman et al. (1984: 43) actually reported that the percentage of red tree voles in spotted owl diets in 

Oregon was highly variable among regions, ranging from 0 to 49 percent.

3.   Aiken, A. 2013. Cascadia Wildlands, The Center for Biological Diversity, 

and Benton Forest Coalition v. Bureau of Land Management and Seneca 

Sawmill Company, opinion and order. U.S. District Court for the District 

of Oregon, Eugene. Case No. 6:12-cv-01739-AA.

In this case in which the plaintiffs challenged a Bureau of Land management timber sale, Judge Aiken dismissed the case and allowed the Rickard 

Creek timber sale to proceed. The judge ruled that the Bureau of Land 

Management did not violate the Federal Land Policy and Land Management Act by classifying Rickard Creek as a “non-high priority” tree vole 

site.

4.   Alexander, C.P. 1967. The crane flies of California. Bulletin of the 

California Insect Survey. 8: 1–269.

Alexander reported crane fly larvae from a Sonoma tree vole nest at 

Stewarts Point, Sonoma County, California. The species was identified as 

Limonia nubeculosa sciophila after the larvae hatched in captivity.
4

GENERAL TECHNICAL REPORT PNW-GTR-909

5.   Alexander, L.F.; Verts, B.J.; Farrell, T.P. 1994. Diet of ringtails 

(Bassariscus astutus) in Oregon. Northwestern Naturalist. 75: 97–101.

The authors found red tree vole remains in ringtail (Bassariscus astutus) 

scat in southwest Oregon.

6.   Andrews, L.S. 1995. A survey of red tree vole utilized structures in 

the west fork of the Oak Creek sub-basin; McDonald-Dunn Research 

Forest. Unpublished report. Corvallis, OR: Oregon State University. 17 p. 

(Available upon request from the compilers).

Andrews located 87 potential trees with red tree vole nests during line 

transect surveys on McDonald Forest in Benton County, Oregon, in 

1994–1995. Of these, five trees had tree vole nest material located on the 

ground and seven were documented to be tree vole nests when they were 

climbed. The status of the other trees with potential nests was not verified, 

as the trees were not climbed.

7.   Anthony, H.E. 1928. Field book of North American mammals. New 

York: G.P. Putnam’s Sons. 628 p. 

In this general species account, Anthony discussed the tree vole and the 

white-footed vole. He said that tree voles were rare in museums and that 

little was known of their life history. He reported that average nest height 

was 9 m but did not acknowledge that this could have been biased by the 

fact that most early naturalists could not climb big trees. He repeated 

previous unsubstantiated speculation that male tree voles might be almost 

entirely terrestrial. 

8.   Anthony, R.G.; Forsman, E.D.; Green, G.A.; Witmer, G.; Nelson, 

S.K. 1987. Small mammal populations in riparian zones of different-aged 

coniferous forests. Murrelet. 68: 94–102.

The authors used museum special traps and rat traps to sample small 

mammals in riparian forest habitats in the Cascades near Blue River and 

McKenzie Bridge, Lane County, Oregon. They sampled five old-growth 

stands, five mature stands, and two young stands. In 10,800 trap-nights, 

they captured nine white-footed voles but did not capture any tree voles. 

They concluded that additional information was needed to describe 

specific habitat associations of white-footed voles. 

9.   Apple, M.; Tiekotter, K.; Snow, M.; Young, J.; Soeldner, A.; Phillips, 

D.; Tingey, D.; Bond, B.J. 2002. Needle anatomy changes with increasing tree age in Douglas-fir. Tree Physiology. 22: 129–136. 
5

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

This paper contained nothing specific about tree voles; however, anyone 

interested in the diet and feeding behavior of tree voles might find it 

useful because it is a good reference on needle structure in Douglas-fir 

(Pseudotsuga menziesii), especially with regard to the development of resin 

ducts. Compared to needles of old-growth trees, needles of young Douglasfir saplings were longer, had smaller vascular cylinders, larger resin ducts, 

and fewer hypodermal cells. These results suggest that tree age could play 

a role in the nutritional value or digestibility of Douglas-fir needles for tree 

voles, but this has not been investigated. 

10.   Ash, S.; Schmelzer, S. 2005. Let’s go voling. Audubon Society of 

Portland, OR. Warbler. 69: 9.

Ash and Schmelzer described a Portland Audubon Society field trip on

30 July 2005, during which researchers climbed to a nest and captured a 

red tree vole in the Oregon Coast Ranges a few kilometers west of Monroe, 

Benton County.

11.   Aubry, K.B.; Crites, M.J.; West, S.D. 1991. Regional patterns of 

small mammal abundance and community composition in Oregon and 

Washington. In: Ruggiero, L.F.; Aubry, K.B.; Carey, A.B.; Huff, M.H., 

tech. coords. Wildlife and vegetation of unmanaged Douglas-fir forests. Gen. Tech. Rep. PNW-GTR-285. Portland, OR: U.S. Department of 

Agriculture, Forest Service, Pacific Northwest Research Station: 285–294.

This report was a synthesis based on one study in Washington and two 

studies in Oregon (Corn and Bury 1991, Gilbert and Allwine 1991). Pitfall 

traps were used to sample small mammals. In total, 8,661 small mammals 

were captured in the three study areas. Based on a log-linear analysis of 

stand age vs. geographic province, the authors reported that red tree voles 

were closely associated with old-growth forests. In support of this conclusion, they cited two other studies in which there was some evidence for 

an association between tree voles and old-growth forest (Bury 1986, Corn 

et al. 1988). The authors concluded that “Although the data are minimal, 

existing information indicates that red tree voles are largely restricted 

to old-growth forests; younger forests apparently do not provide suitable 

habitat for this species.” This rather strong statement suggests either that 

the authors were unaware that many tree voles had been captured in young 

forests by specimen collectors, or that the authors considered the earlier 
6

GENERAL TECHNICAL REPORT PNW-GTR-909

collections biased because many specimen collectors focused their sampling efforts in young forests. The authors also suggested that extensive 

loss or fragmentation of old-growth forests could eliminate tree voles from 

forested landscapes.

12.   Aubry, K.B.; Hayes, J.P.; Biswell, B.L.; Marcot, B.G. 2003. The ecological role of tree-dwelling mammals in western coniferous forests. In: 

Zabel, C.J.; Anthony, R.G., eds. Mammal community dynamics: management and conservation in the coniferous forests of western North 

America. Cambridge, United Kingdom: Cambridge University Press: 

405–443. Chapter 12.

In this literature review, the authors concluded that tree voles were the 

most arboreal mammal in North America, were closely associated with 

large trees in Douglas-fir forests, and were most abundant in latesuccessional forests where canopies of large trees were interconnected.

13.   Bailey, V.O. 1914. Unpublished field notes. On file with: The Smithsonian 

Institution manuscript collection, Fish & Wildlife, Record Unit 7176. 

Washington, DC: U.S. Department of the Interior Field Reports, 1860 

(1887–1961), Series 1. Field reports submitted by naturalists conducting 

field research in the United States, Oregon. Box 84. Folder 21.

Vernon Bailey described three days of searching for tree voles with Alfred 

Shelton at Spencer Butte in Lane County, Oregon, in 1914. After considerable effort, they finally captured a single vole, as described in the following narrative: “With a pair of climbing irons we took turns in climbing the 

trees. The next morning the third tree fell to my turn. It was about 2 feet 

in diameter at the bottom and the nest about 80 feet up on the first green 

branches. It was a hard climb but the large flat nest at once showed fresh 

signs and I dug in with great care. Not until I had dug out the last of the 

nest did the occupant leave the house and run out on one of the branches 

a few feet below. It was only about 20 feet from me so with a fine shot 

pistol I shot at one edge of it and brought it to the ground. The stomach and 

intestines are filled with bright green semi-fluid material, evidently the 

inner pulp of spruce [Douglas-fir] leaves. The feces are small and excreted 

in bundles of green pellets.” He concluded by stating, “The treetop mouse 

is not a rare species but probably the most inaccessible of our small mammals.” This field trip was later described in a full length article in the 

Oregon Historical Quarterly (Jobanek 1988).
7

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

14.   Bailey, V. 1915. Discovery of the tree mouse (Phenacomys longicaudus—

True). The Oregon Sportsman. 3: 147–149.

Bailey described several of the early records of red tree voles in Oregon 

and recounts his trip to collect tree voles on Spencer Butte, Lane County, 

Oregon, in 1914. Bailey said that he often set traps in hopes of catching 

tree mice, without success. In 1907, William Bebb showed him several 

tree vole specimens that were collected by loggers. Bebb never published 

this information, but we found two of his specimens in the collection at the 

Bean Life Science Museum at Brigham Young University, both collected 

in 1905 near Marmot in Clackamas County, Oregon. Bailey also reported 

that Alfred Shelton collected two juvenile tree voles in 1914 at Spencer 

Butte, Lane County, Oregon. He described the day in 1914 when he and 

Shelton returned to Spencer Butte and captured an adult vole by climbing 

several trees until they chased a vole out of a nest and shot it with a collecting pistol. He described nests with green cuttings on top and says that 

trees had from one to five nests. Most nests were greater than 24 m above 

ground. He wrote, “The feces were deposited around the outside of the 

nest and helped to build up its solid walls and render it weatherproof.” He 

says that nests were built of “...fine midribs of a great number of spruce 

[Douglas-fir] needles, the sides of which had served as food.” This is one 

time when Bailey apparently got it wrong. When feeding on Douglas-fir 

needles, tree voles remove the resin ducts from the edges of the needle and 

consume the central part of the needle. Thus, we assume he must not have 

looked closely at the discarded remains.

15.   Bailey, V.O. 1936. The mammals and life zones of Oregon. North 

American Fauna. 55: 1–416.

Bailey was one of the first to discuss tree vole distribution in a spatial 

context. He stated that tree voles only occur in areas of extensive forest or 

areas of recently isolated forest. He pointed out that tree voles sometimes 

build their nests on top of old hawk or squirrel nests. He reported that 

Harry Wilder located several tree vole nests in burrows under logs or 

woodpiles near Douglas-fir trees that had low-hanging branches that were 

easily accessed from the ground. Bailey tried unsuccessfully to capture 

tree voles with snap-traps on logs or low branches. He reported that Harry 

Wilder and Irvine Clay thought that the main breeding season for tree 

voles was early spring to late summer but that tree voles breed throughout 

the year. He postulated that slow development of the young was an adaption for living in an arboreal environment. He also reported that tree voles 
8

GENERAL TECHNICAL REPORT PNW-GTR-909

did not consume water in captivity except when fed desiccated needles.

He speculated that American marten (Martes americana) and fisher

(M. [= Pekania] pennanti) were potential predators of tree voles.

16.   Bailey, V. 1939. Obituary notices: William Bebb. Journal of Mammalogy. 

20: 132–133.

In this obituary, Bailey described his friend William Bebb, dental surgeon, 

ardent naturalist, and specimen collector. In 1907, Bailey and Clint Merriam visited Bebb at his home in Pasadena, California, where they were 

stunned to find several red tree vole skins of good quality. They were curious how Bebb had collected these specimens because they had never seen 

a live tree vole and were only aware of the one mangled type specimen 

that was found dead in the road near Coos Bay, Oregon, in 1890. Bebb told 

them that he collected tree voles from Douglas-fir trees that were felled by 

loggers near Marmot, Clackamas County, Oregon. 

17.   Bake, D.B. 2003. Unpublished letter to Eric Forsman (Available upon 

request from the compilers).

In this short letter to Eric Forsman, Doug Bake confirmed a date (21 July 

2003) when he could meet and discuss his career as a logger and tree vole 

collector in the north Coast Ranges of Oregon. Doug said that he started 

collecting tree voles after another logger (Peter Walker) told him that 

museum collectors were eager to obtain specimens. Doug eventually collected 98 red tree voles, most of which are in the Slater Museum of Natural 

History at the University of Puget Sound. We found that on some of those 

specimens, his name was incorrectly spelled “D. Bates.” For more details 

on Doug Bake’s collecting efforts see Forsman and Swingle (2010). 

18.   Baker, D.; Palmer, C.; Tolle, T. 2005. Northwest Forest Plan: the first 

ten years (1994–2003) implementation monitoring: accomplishments 

and compliance with plan requirements. http://www.reo.gov/monitoring/

reports/10yr-report/implementation/ImplementationI.pdf. (August 2014).

The authors reported that requirements to “survey and manage” wildlife 

species on federal forest lands were resulting in a reduction in harvest. 

They cited an example in which predisturbance surveys for red tree voles 

resulted in portions of the proposed harvest area being protected from logging. When managers were faced with numerous tree vole nest structures 

in a sale area, they had to weigh the potential cost of climbing nest trees to 

determine tree vole activity status and reconfiguring the proposed sale unit 

versus simply dropping the sale and moving to another area. In addition, 
9

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

the need to protect numerous tree vole nest areas within a harvest unit 

sometimes made timber sales operationally unfeasible, meaning that trees 

could not be logged without damaging protected areas.

19.   Baker, R.H. 1967. Distribution of recent mammals along the Pacific 

coastal lowlands of the western hemisphere. Systematic Zoology. 16: 

28–37.

In table 4, Baker lists the red tree vole and white-footed vole as only occurring in lowland conifer forests of the Pacific coast. 

20.   Barbour, R.J.; Johnston, S.; Hayes, J.P.; Tucker, G.F. 1997. Simulated 

stand characteristics and wood product yields from Douglas-fir plantations managed for ecosystem objectives. Forest Ecology and Management. 

91: 205–219.

Based on model simulations of forest growth, the authors suggested that 

red tree voles may benefit from thinning in young forests because thinning 

produces trees with larger crowns. They provided no data on the actual 

response of tree vole populations to thinning. 

21.   Barnard, J. 2006. Ruling: BLM violated vole protections to allow logging. Associated Press. 7 November 2006.

In this newspaper article, Barnard reported that the 9th U.S. Circuit Court 

of Appeals ruled that the Bureau of Land Management failed to conduct a 

public review before dropping the red tree vole from the list of species that 

require formal surveys in proposed timber sales.

22.   Barnosky, A.D.; Hadly, E.A.; Bell, C.J. 2003. Mammalian response to 

global warming on varied temporal scales. Journal of Mammalogy. 84: 

354–368.

Based on fossil evidence, the authors suggested that Phenacomys and

Clethrionomys [= Myodes] that occupied closed canopy forests decreased 

during warm periods of the past, and that Phenacomys became progressively rarer during the last glacial-interglacial transition as the climate 

became warmer.

23.   Barrows, C. 1980. Feeding ecology of the spotted owl in California. 

Journal of Raptor Research. 14: 73–78.

Barrows collected 375 prey items from four pairs of northern spotted owls 

in three widely scattered locations in northwestern California. The sample 

included 64 Sonoma tree voles (17.1 percent of total prey items). He did not 
10

GENERAL TECHNICAL REPORT PNW-GTR-909

say how diets differed among areas, which included two locations in Marin 

County that were outside the range of the tree vole (Point Reyes and San 

Geronimo), and one location within the range in Mendocino County. 

24.   Barrows, C.W. 1985. Breeding success relative to fluctuations in diets 

for spotted owls in California. In: Gutiérrez, R.J.; Carey, A.B., tech. eds. 

Ecology and management of the spotted owl in the Pacific Northwest. 

Gen. Tech. Rep. PNW-185. Portland, OR: U.S. Department of Agriculture, 

Forest Service, Pacific Northwest Forest and Range Experiment Station: 

50–54.

Barrows reported that diets of northern spotted owls in northwestern 

California in 1977–1983 included 9 to 50 percent tree voles.

25.   Behle, W.H. 1933. Unpublished field notes. On file with: the University 

of California, Berkeley, Museum of Vertebrate Zoology. 

Behle reported that he collected an adult female tree vole that was run 

over by a car near Coyote Peak on Bald Hills Road, Humboldt County, 

California.

26.   Bell, C.J.; Jass, C.N. 2011. Polyphyly, paraphyly, provinciality, and 

the promise of intercontinental correlation: Charles Repenning’s contributions to the study of arvicoline rodent evolution and biochronology. 

Palaeontologia Electronica. 14: 18A. 15 p.

The authors reviewed the career of Charles Repenning, a paleontologist 

who was a leader in the effort to reconstruct the evolutionary history of 

arvicoline rodents. Repenning was a mentor to many, whose long and 

productive career was cut short by a burglar who shot him in his home. 

Repenning and many of his colleagues used the shape of the lower first 

molar as the primary structural feature for assessing evolutionary lineages 

of arvicolines. Based on a sample of six fossil individuals that included two 

lower M1 molars, Repenning (1987) proposed the new species Phenacomys brachyodus. Repenning et al. (1987) suggested that Phenacomys was 

derived from an unknown form of Cromeromys. Repenning (1998, 2001) 

suggested that Phenacomys were immigrants to North America from Asia 

via the Bering Land Bridge during the Pleistocene ice ages. 

27.   Bellinger, M.R.; Haig, S.M.; Forsman, E.D.; Mullins, T.D. 2005. 

Taxonomic relationships among Phenacomys voles as inferred by cytochrome b. Journal of Mammalogy. 86: 201–210.
11

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

The authors used genetic base pairs to compare taxonomic relationships 

among red tree voles, dusky tree voles, Sonoma tree voles, white-footed 

voles, and heather voles (Phenacomys intermedius). There were clear 

taxonomic differences in all pairings except for the red tree vole and its 

putative subspecies, the dusky tree vole. The authors concluded that red 

tree voles and white-footed voles were closely related, supporting their 

inclusion in the same genus (Arborimus). The authors also suggested that 

their results indicated a Pleistocene radiation of the Arborimus clade, 

concordant with pulses of diversification observed in other murid rodents. 

The authors concluded that the elevation to full generic rank of Arborimus 

from Phenacomys was open to debate, as their data did not clearly support 

a decision one way or the other. 

28.   Benson, S.B. 1930–1956. Unpublished field notes. On file with: 

University of California, Berkeley, Museum of Vertebrate Zoology. 

Seth Benson’s voluminous notes described numerous field trips during 

which he and his wife Emma, along with students and colleagues, collected a total of 49 Sonoma tree voles in Humboldt and Sonoma Counties, 

California. Benson and his graduate students kept many of these voles in 

captivity for periods of 2 to 30 days. He and Emma located 6 to 8 clouded 

salamanders (Aneides ferreus) in a single Sonoma tree vole nest. Like 

many naturalists who studied tree voles in California, Benson traveled to 

the Wilder Ranch near Carlotta, Humboldt County, where Harry Wilder 

showed him tree vole nests and freely dispensed his knowledge of tree vole 

life history. Wilder told Benson that tree voles had become scarce around 

his ranch because the forest behind the ranch had been logged and the 

population had been reduced by collectors. 

29.   Benson, S.B.; Borell, A.E. 1931. Notes on the life history of the red tree 

mouse Phenacomys longicaudus. Journal of Mammalogy. 12: 226–233.

Although the title states that this paper is about the red tree vole, it was 

actually based on observations of Sonoma tree voles in California, which 

had not been designated as a separate species at the time of publication. 

The authors described nests, young, diet, and behavior of tree voles based 

on observations of voles in the wild and kept 27 tree voles in captivity. 

They captured Sonoma tree voles by climbing trees near Carlotta, Kneeland, Bridgeville, and the Van Duzen River in Humboldt County, and near 
12

GENERAL TECHNICAL REPORT PNW-GTR-909

Monte Rio, Camp Meeker, Occidental, and Bohemian Grove in Sonoma 

County. They searched for, but did not locate tree voles in Marin and Santa 

Cruz Counties.

They noted that tree vole nests were placed on firm supports that were 

close to a food source of green needles. Some tree vole nests were built 

on top of old bird or squirrel nests. Most nests were on branches close to 

the trunk, but some were as much as 9 m out on limbs. Most nests were 

in Douglas-fir or grand fir (Abies grandis), but a few were in redwoods 

(Sequoia sempervirens). The authors did not say what the voles that nested 

in redwoods were feeding on, but they did say that nests in redwoods were 

always located near Douglas-fir trees. The authors reported that captive 

tree voles ate rolled oats, fresh apples, fresh carrots, fresh squirrel meat, 

dried pears, dried apples, raisins, Cedrus deodara, and Abies venusta, even 

when supplied with fresh Douglas-fir and grand fir. This is interesting 

because we and others (e.g., Clifton 1960) found that captive tree voles 

generally refused exotic foods except for occasional individuals that would 

eat apple.

The authors reported that reproduction occurred throughout the year, but 

they did not discuss seasonal variation in reproduction. They observed a 

one-month-old captive eating fir needles while still suckling. They corrected Howell (1926) and agreed with Taylor (1915) by stating that tree 

voles ate the midrib of Douglas-fir needles after removing the lateral edges 

of the needle, which contained the resin ducts. They described the feeding 

behavior of a captive tree vole as “almost machine-like.” The vole would 

rapidly pass a needle through its incisors from right to left and immediately flip the needle around to chew along the opposite side of the needle. 

Not all needles had the resin ducts removed in the same manner. In some 

cases, only part of the resin duct was removed. They also observed captives eating young needles without removing the resin ducts. They pointed 

out that resin ducts of Douglas-fir were easy to identify because one side 

was smooth and the other side had unique scallop-shaped marks that were 

left by the vole’s incisors.

Unlike most other collectors who have captured tree voles by climbing nest 

trees, Benson and Borell captured about the same number of females and 

males. They reported that males and females usually did not inhabit the 

same nests except for one nest in which they found an adult male with a 

subadult female. They also found one tree with three occupied nests.
13

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

They reported that tree voles were easily kept in captivity as long as they 

were provided with fresh food. They also reported keeping six tree voles 

in a bathtub filled with sawdust for 4 days. The six voles slept in the 

same nest in the tub. After removing four voles from the bathtub, the two 

remaining voles lived together for 1 month. They did not report if these 

voles were related or not, and they did not discuss the long-term survival of 

these normally solitary voles kept in such unusual conditions. The captive 

voles drank considerable water, and the authors noticed that voles kept in 

warm rooms drank more water than those kept in cool rooms.

They reported that some tree voles were infected by the small mite, 

Atricholaelaps californicus. They also found an arboreal salamander 

(Aneides lugubris) in a nest at Bohemian Grove, Monte Rio, and what they 

thought was a black salamander (A. flavipunctatus) in an abandoned tree 

vole nest near Occidental, Sonoma County. We are not aware of any other 

reports of black salamanders in tree vole nests.

30.   Bingham, B.B.; Sawyer, J.O., Jr. 1991. Canopy structure and tree condition of young, mature and old-growth Douglas-fir/hardwood forests. In: 

Kerner, H.M., ed. Proceedings of the symposium on biodiversity of northwestern California. Wildland Resources Center Report No. 29. Berkeley 

CA: University of California, Berkeley: 141–149.

The authors found that there were more broken crowns and cavities in 

old-growth stands of Douglas-fir than in young or mature forests. While 

not particularly surprising, these findings are germane to the ecology of 

tree voles because we and others have found that many tree vole nests are 

located in cavities or broken tops of old trees.

31.   Biswell, B.L. 1994. Intertree movement and spatial distribution of nest 

trees occupied by individual red tree voles: implications for conservation 

[Poster]. In: 75th anniversary meeting of the Society of Mammalogists, 

Washington, DC (Available upon request from the compilers).

Biswell studied the movement patterns of nine adult and four subadult 

red tree voles that were radio-collared in western Oregon in 1992–1993. 

Adults were followed for 35 to 106 days, and analysis of nest site usage 

was based on four to five daytime locations per week to assure independence between observations. Individual voles used two to seven nest trees 

each, and the greatest straight-line distance between two nests that were 

occupied consecutively by the same individual was 76 m. When moving to 
14

GENERAL TECHNICAL REPORT PNW-GTR-909

a new tree, adult voles reoccupied previously constructed nest structures 

at least 68 percent of the time. Of the 39 nest trees located via telemetry, 

36 percent contained more than a single nest, and one tree contained seven 

nest structures. Subadults were followed for 23 to 40 days during which 

time two dispersed from their natal nest trees. One subadult was located 

in five different trees and reached a maximum straight line distance from 

its natal nest of 340 m. Biswell concluded that dispersing subadults were 

successful in crossing small forest roads, small streams, and 30-m-wide 

canopy gaps while traveling between nest sites, which required movement 

on the ground.

32.   Biswell, B.L. 2002. Variation in the abundance of the Oregon red tree 

voles in five watersheds in western Oregon [Abstract]. Northwestern 

Naturalist. 83: 64.

This abstract from the 2002 annual meeting of the Society for Northwestern Vertebrate Biology included a more complete analysis of the data 

described by Biswell and Forsman (1999). Biswell studied red tree vole 

nest abundance in five watersheds containing various levels of old forest 

and tested for differences in abundance across watersheds and between 

serial stages. He conducted approximately 46.7 km of line transect surveys 

in 96 randomly selected stands in the Coast Ranges and interior valley 

foothills in western Oregon. Average densities of recently occupied or 

occupied vole nest trees in four watersheds were 0.2 to 3.3 nest trees/hectare). Densities of occupied or recently occupied nest trees varied substantially within and among forest age-classes, and a few “hot spots” strongly 

influenced overall density estimates in four of the watersheds. Biswell 

concluded that “...cautious management is needed to ensure sufficient 

numbers of sites to maintain appropriate levels of biological function and 

species interactions within this region.”

33.   Biswell, B.; Blow, M.; Breckel, R.; Finley, L.; Lint, J. [2002]. Survey 

protocol for the red tree vole, Arborimus longicaudus (= Phenacomys longicaudus in the Record of Decision of the Northwest Forest Plan). Version 

2.1. Portland, OR: U.S. Department of Agriculture, Forest Service and 

U.S. Department of the Interior, Bureau of Land Management. 13 p.

This was the third of four versions of the survey protocol for red tree voles 

on federal forest lands (see also Biswell et al. [2000], Huff et al. 2012, 

USDA FS and USDI BLM 1996). It included a literature review and a revision of the previous protocol. 
15

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

34.   Biswell, B.; Blow, M.; Finley, L.; Madsen, S.; Schmidt, K. [2000]. 

Survey protocol for the red tree vole, Arborimus longicaudus (= 

Phenacomys longicaudus in the Record of Decision of the Northwest 

Forest Plan). Version 2.0. Portland, OR: U.S. Department of Agriculture, 

Forest Service and U.S. Department of the Interior, Bureau of Land 

Management. 32 p.

This was the second of four versions of the survey protocol for red tree 

voles on federal forest lands (see also Biswell et al. 2002, Huff et al. 2012; 

USDA FS and USDI BLM 1996). The first part of the protocol included a 

literature review on taxonomy, distribution, biology, home range, dispersal, 

abundance, habitat, and nests of red tree voles. The second half described 

the survey protocol. This protocol was designed primarily to determine if 

tree voles were present in areas proposed for harvest.

35.   Biswell, B.L.; Finley, L.; Jones, J.M.; Schmalenberger, F. 2004. Survey 

and Manage Program strategic survey report: red tree vole strategic survey results for the 2000 Klamath RTV study in Siskiyou County, California. 

94 p. Unpublished report. U.S. Department of Agriculture, Forest Service 

and U.S. Department of the Interior, Bureau of Land Management, 

Portland, OR.

In 2000, contractors were hired to conduct surveys of red tree voles north 

of the Klamath River on the Klamath National Forest, Siskiyou County, 

California. They surveyed 47 km of transect in 77 mature forest plots and 

detected potential tree vole nests in 173 trees. They climbed to all potential 

nests and found that 18 (10.4 percent) were red tree vole nests. Based on a 

distance sampling analysis, the authors estimated that the effective sampling distance (mean ± SE) for all 173 potential nest structures was 21.0 ± 

0.6 m. They did not provide a separate estimate for red tree vole nests.

36.   Biswell, B.L.; Forsman, E.D. 1999. The effects of landscape character 

and stand structure on the distribution and abundance of the red tree vole 

[Abstract]. Northwestern Naturalist. 80: 119.

In this abstract from the 1999 annual meetings of the Society for Northwestern Vertebrate Biology and the California North Coast Chapter of 

the Wildlife Society, Biswell and Forsman stated they conducted 157 

line transect surveys, totaling 47.5 km, in 96 randomly selected stands 

distributed between five watersheds in western Oregon. Stands ranged 

from young regenerating forests on clearcuts to late-successional forest. 
16

GENERAL TECHNICAL REPORT PNW-GTR-909

Watersheds were about 5 000 ha and contained from 5 to 55 percent latesuccessional forest. Surveyors detected 703 nest structures of which 84 (12 

percent) were tree vole nests. The other 619 nests (88 percent) were built 

by other arboreal rodents or birds. While tree vole nests were detected in 

all age classes surveyed, the number of nests increased with stand age and 

with the amount of late-successional forest occurring within the watershed. 

The authors concluded that the specialized habitat requirements and low 

dispersal capabilities of tree voles could lead to extirpation of the species 

in heavily managed landscapes dominated by young forests.

37.   Biswell, B.; Forsman, E.; Swingle, J.; Chamblin, D.; Leach, R.; 

Lundsten, S.; Lewis, M. 1998. The effects of landscape character and 

stand structure on the distribution and abundance of the red tree voles 

(Arborimus longicaudus) in western Oregon. 2 p. Unpublished report. 

On file with: U.S. Department of Agriculture, Forest Service, Pacific 

Northwest Research Station, Forestry Sciences Laboratory, 3625 93rd

Ave. SW, Olympia, WA. (Available upon request from the compilers).

The goals of this study were to expand upon what was known of red tree 

vole distribution and abundance in different vegetation communities and to 

determine the geographic boundary between the red tree vole and Sonoma 

tree vole. The results were briefly described in abstracts at two different 

meetings (Biswell 2002, Biswell and Forsman 1999).

38.   Black, H.C.; Hooven, E.H. [sic]. 1974. Responses of small-mammal 

communities to habitat changes in western Oregon. In: Black, H.C., ed. 

Wildlife and forest management in the Pacific Northwest. Corvallis, OR: 

Oregon State University, School of Forestry: 177–186.

This paper mentions a red tree vole that was captured in a live-trap near 

Cottage Grove, Lane County, Oregon. See Borrecco (1973) for details. 

Hooven’s initials were actually E.F.

39.   Blois, J.L. 2005. A multilocus assessment of the genetic structure and 

diversity within an endemic, arboreal vole, Arborimus pomo, in northern 

California. Arcata, CA: Humboldt State University. 82 p. M.A. thesis.

Blois used amplified fragment length polymorphisms (ALFP) and 

mitochondrial DNA (mtDNA) analyses to address diversity and population structure in the Sonoma tree vole. The mtDNA analysis indicated 

that there was a north and south clade of the Sonoma tree vole, but the 

AFLP analysis indicated that all samples belonged to a single panmictic 
17

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

population. Blois concluded that, for A. pomo, “...the combined AFLP and 

mtDNA data support the recognition of two separate Management Units, 

but only a single Evolutionarily Significant Unit within the species.” She 

also reported capturing three tree voles at the Humboldt State University 

Schatz Demonstration Tree Farm, 1.6 km north of Maple Creek, Humboldt 

County. 

40.   Blois, J.L.; Arbogast, B.S. 2006. Conservation genetics of the Sonoma 

tree vole (Arborimus pomo) based on mitochondrial and amplified fragment length polymorphism markers. Journal of Mammalogy. 87: 950–960.

This was the published version of Blois’ thesis. The mtDNA analysis indicated that there was a north and south clade of the Sonoma tree vole, but 

the AFLP analysis indicated a single panmictic Sonoma tree vole population. Based on the mtDNA results, the authors concluded that tree voles 

north of the Klamath River in California should be included in the red tree 

vole clade identified by Miller et al. (2006) and Murray (1995). 

41.   Blois, J.L.; McGuire, J.L.; Hadly, E.A. 2010. Small mammal diversity 

loss in response to late-Pleistocene climate change. Nature. 465: 771–774 

plus 1 p. supplement.

The authors compared small mammal fossil remains located in a cave 

north of Redding, Shasta County, California. They reported that the taxa 

located in the cave currently exist in northern California, but the relative abundance changed over time. The fossil samples indicated that the 

white-footed vole was persistently present during late Pleistocene but only 

intermittently present during the Holocene. The cave was east of the current distribution of the white-footed vole in California. 

42.   Bochkov, A.V. 2011. Mites of the subgenus Microtimyobia (Acariformes: 

Myobiidae: Radfordia) with their host-parasite relationships with the cricetid rodents (Cricetidae). Zootaxa. 2954: 1–86.

In this literature review, Bochkov revised the myobid mites associated 

with voles. This included two mites associated with tree voles, Radfordia 

(Graphiurobia) arborimus (Fain and Whitaker 1975) and R. (Microtimyobia) arborimus (Fain and Lukoschus 1977). 

43.   Booth, E.P. 1950. How to know the mammals. Dubuque, IA: William C. 

Brown and Company. 206 p.

On pages 142–144, Booth gave a brief description of white-footed voles 

and tree voles. He said that the tree vole was one of the most peculiar voles 

in the world owing to its arboreal nature and its diet of conifer needles. He 
18

GENERAL TECHNICAL REPORT PNW-GTR-909

stated that the putative subspecies of the red tree mouse, the dusky tree 

mouse, lives mainly in Sitka spruce (Picea sitchensis) and western hemlock (Tsuga heterophylla) trees rather than Douglas-fir.

44.   Borrecco, J.E. 1973. The response of animals to herbicide-induced habitat changes. Corvallis, OR: Oregon State University. 92 p. M.S. thesis.

In 1971, Borrecco captured a single red tree vole in a Sherman live-trap at 

Carlson Creek, 25.8 km south of Cottage Grove (T23S, R04W, Section 13, 

E1/2) in Lane County, Oregon. Vegetation on the study plot was a stand of 

young Douglas-fir growing on a 19-ha area that was clearcut and burned in 

1959 and then planted in 1960, 1964, and 1970.

45.   Brock, E.M. 1954–1956. Unpublished field notes. On file with: 

University of California, Berkeley, Museum of Vertebrate Zoology.

Brock collected two adult and three juvenile Sonoma tree voles near 

Capetown and Petrolia, Humboldt County and Freestone, Sonoma County, 

California.

46.   Brown, E.R., tech. ed. 1985. Management of wildlife and fish habitats 

in forests of western Oregon and Washington. Part 1: Chapter narratives. 

Publication No. R6-F&WL-192-1985. Portland, OR: U.S. Department of 

Agriculture, Forest Service and U.S. Department of the Interior, Bureau of 

Land Management. 332 p.

Brown’s report included a chapter by Forsman et al. on pages 259–267 in 

which the authors say that tree voles comprised 13 percent of spotted owl 

prey in temperate coniferous forests and 5 percent of spotted owl prey in 

mixed-conifer forests in western Oregon.

47.   Brown, E.R. 1985. Management of wildlife and fish habitats in forests 

of western Oregon and Washington. Part 2: Appendices. Publication No. 

R6-F&WL-192-1985. Portland, OR: U.S. Department of Agriculture, 

Forest Service and U.S. Department of the Interior, Bureau of Land 

Management. 229 p.

Brown provides no new data on tree voles, but on page 171 in appendix 8, 

he suggested that tree vole home ranges include at least one tree. 

48.   Brown, L.N. 1964. Breeding records and notes on Phenacomys silvicola

in Oregon. Journal of Mammalogy. 45: 647–648.

Brown said that he found a “colony” of dusky tree voles on Oregon State 

University’s McDonald Forest, 13 km north of Corvallis, Benton County, 

Oregon, in 1963. He captured an adult male in one nest and four half-
19

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

grown juveniles in another nest, both about 9 m above ground. Apparently 

he did not see the adult female. On 15 May 1963, he captured one adult 

female and two newborns that weighed 2.56 and 2.60 g, respectively, in 

a nest about 0.4 km south of the first area. The female was kept alive for 

15 days and then killed. She had three embryos in her uterus with crownrump length averaging 4.5 mm. Brown concluded that this indicated that 

tree voles experienced “...postpartum ovulation and pregnancy upon 

occasion.” Brown claimed that his specimens were sent to the University 

of Wyoming, but the specimens are not at that institution. Like Vernon 

Bailey, he said that nests and sleeping chambers were lined with midribs of 

Douglas-fir needles, but it is obvious that he did not look closely at the nest 

material, or he would have realized they were the edges of the needles. He 

also described a tree vole nest that was occupied by two deer mice (Peromyscus maniculatus).

49.   Brylski, P. 1999. White-footed vole. Sacramento, CA: California Wildlife 

Habitat Relationships System, California Department of Fish and Game, 

California Interagency Wildlife Task Group. 2 p.

Brylski described the distribution, habitat, and reproduction of the whitefooted vole. He emphasized that there was little information on the basic 

life history and predators of this species in California.

50.   Brylski, P.; Harris, J. 1999. California red tree vole. Sacramento, CA: 

California Wildlife Habitat Relationships System, California Department 

of Fish and Game, California Interagency Wildlife Task Group. 2 p.

In this species account, the authors described the distribution, habitat, 

nests, and taxonomy of the Sonoma tree vole. The information is somewhat 

dated.

51.   Burt, W.H.; Grossenheider, R.P. 1980. A field guide to the mammals, 

North America north of Mexico. 3rd ed. Boston, MA: Houghton Mifflin 

Co. 289 p.

This field guide provides brief descriptions of the habitats and life-histories

of the heather vole, white-footed vole, and red tree vole.

52.   Burton, K. 2005. Personal communication in email to Eric Forsman

(Available upon request from the compilers).

On 11 July 2005, wildlife biologist Ken Burton saw a Steller’s jay (Cyanocitta stelleri) trying to kill a Sonoma tree vole by pecking it. The jay 

flew off as Burton drove up, leaving the tree vole alive but stunned on the 

Bald Hills Road east of Arcata, Humboldt County, California.
20

GENERAL TECHNICAL REPORT PNW-GTR-909

53.   Bury, R.B.; Corn, P.S. 1987. Evaluation of pitfall trapping in northwestern forests: trap arrays with drift fences. Journal of Wildlife Management. 

51: 112–119. 

The small mammal data used in this analysis was the same information 

described in Corn and Bury (1986, 1988) and Corn et al. (1988). These data 

were collected in 1983 at H.J. Andrews Experimental Forest, Oregon, and 

Wind River Experimental Forest, Washington. Pitfall traps were opened 

in May 1983 and monitored for 180 days. The authors compared traps with 

short drift fences to traps without fences. They concluded that drift fences 

did not improve capture rates of small mammals. In 38,880 trap-nights, 

they captured 2,029 mammals, including 17 red tree voles at H.J. Andrews. 

The tally of numbers in this report is slightly different than in their other 

papers (2,029 vs. 2,104; Corn and Bury 1986, 1988). We suspect this may 

have been because they did not include mammals captured on an experimental grid. The capture rate of tree voles was 0.044 tree voles per 100 

trap-nights. They concluded that tree voles were “...extremely difficult to 

trap by any method.” 

54.   Buzan, E.V.; Krystufek, B.; Hänfling, B.; Hutchinson, W.F. 2008. 

Mitochondrial phylogeny of Arvicolinae using comprehensive taxonomic 

sampling yields new insights. Biological Journal of the Linnean Society.

94: 825–835.

The authors concluded that Phenacomys was the only genus in the tribe 

Phenacomyini. Microtus was noted as the closest living relative to Phenacomys. Because these authors apparently did not recognize the genus 

Arborimus as distinct from Phenacomys, we assume they would also have 

included Arborimus in the same tribe.

55.   Buzzio, O.L.; Koninckx, A.; Carreno, N.B.; Castro-Vazquez, A. 2001. 

Embryo implantation during the short luteal phase of the corn mouse, 

Calomys musculinus, and the apparent lack of a lactational diapause in 

South American murid rodents. Reproduction. 121: 815–823. 

Based on information in Hamilton (1962), the authors suggested that 

the red tree vole may have lactational embryonic diapause or lactational 

delayed implantation.

56.   Cahalane, V.H. 1947. Mammals of North America. New York: 

Macmillan Co. 682 p.

In his section on chisel-toothed mammals on pages 517–519, Cahalane 

made several unsupported or incorrect statements about tree voles. He 
21

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

said that tree voles may live on the ground, with the male more likely to 

do so, but he provided no data to support this, and he ignored the evidence 

that ground nesting by tree voles was actually extremely rare. Like Bailey 

(1915), Howell (1926) and Brown (1964), he incorrectly stated that tree 

voles ate the edges of Douglas-fir needles and discarded the central part 

of the needle. And then he topped it off by saying that tree voles occurred 

from eastern Canada to east-central California. 

57.   California Academy of Sciences. 2006. Manuscripts and special collections. On file with: California Academy of Sciences, Academy Library, 

San Francisco, CA. http://researcharchive.calacademy.org/research/library/

special/BOXLISTS/o&m_fieldnotes.htm. (August 2014).

The California Academy of Sciences library has field notes for William 

Hamilton, III (1961), Chester Lamb (1921), Joseph Mailliard (1920–1936), 

and Harry Wilder (1911), all of whom worked on Sonoma tree voles. We 

examined Mailliard’s notes (see reference 258) but did not have time to 

examine the others, so we do not know if they pertain to tree voles or not.

58.   California Fish and Game. 1986. Mammalian species of special concern 

in California, red tree vole. Sacramento, CA: California Department of 

Fish and Game. 2 p.

This species account included information on the distribution, population 

status, habitat, and nests of the Sonoma tree vole. The anonymous author(s) 

suggested that what was most needed was information on population status 

and distribution. They also suggested that where tree voles are present, 

clearcutting should be replaced with selective logging.

59.   California Fish and Game. 1986. Mammalian species of special concern 

in California, white-footed vole. Sacramento, CA: California Department 

of Fish and Game. 2 p. 

This general species account included information on the distribution, 

population status, habitat, taxonomy, and management of the white-footed 

vole. The author(s) emphasized that there were only 12 specimen records of 

white-footed voles from California and that virtually nothing was known 

about the status of the species.

60.   Camp, C.L. 1913a. Unpublished field notes. On file with: University of 

California, Berkeley, Museum of Vertebrate Zoology.

Camp described the 1913 Museum of Vertebrate Zoology North Coast 

Counties Expedition, during which 17 adult and 12 juvenile Sonoma tree 

voles were collected in Mendocino and Humboldt Counties, California. 
22

GENERAL TECHNICAL REPORT PNW-GTR-909

This expedition resulted in three published papers on Sonoma tree voles 

(Taylor 1915a, 1915b, 1961).

61.   Camp, C.L. 1913b. Unpublished pictures. On file with: University of 

California, Berkeley, Museum of Vertebrate Zoology. Available from:

http://calphotos.berkeley.edu/cgi/img_query?enla

rge=5555+5555+0907+3771 and http://calphotos.berkeley.edu/cgi/img_que

ry?enlarge=5555+5555+0907+3800. (August 2014).

Camp took these pictures of the “Phenacomys grove” and “Phenacomys

colony in grand firs” near Mendocino City, California, during the 1913 

North Coast Counties Expedition. The expedition members collected 

Sonoma tree voles at this site after some local boys showed them how to 

capture tree voles by climbing nest trees.

62.   Carey, A.B. 1989. Wildlife associated with old-growth forests in the 

Pacific Northwest. Natural Areas Journal. 9: 151–162.

In this short synthesis of previous and ongoing studies, Carey presented 

a table in which he displayed tree vole capture data from pitfall studies 

conducted by Corn and Bury (1986, 1988, 1991). Based on Corn and Bury’s 

data, he concluded that tree voles were more abundant in old-growth than 

in mature and young forests. He stated that “...I have observed red tree 

vole colonies in some young stands. But my experience indicates that 

the vole is not widespread in young and mature forests; ongoing research 

shows vole nests can be very abundant in old-growth forests.” He did not 

define what he meant by a “colony” and presented no data to support the 

notion that tree voles are a colonial species. He also suggested that tree 

voles were weak dispersers, incapable of dispersing through clearcuts, 

clearings, or areas of blow-down.

63.   Carey, A.B. 1991. The biology of arboreal rodents in Douglas-fir forests. Gen. Tech. Rep. PNW-GTR-276. Portland, OR: U.S. Department of 

Agriculture, Forest Service, Pacific Northwest Research Station. 46 p.

In this literature review, Carey stated that “Red tree voles are rare in 

closed sapling-pole-sawtimber and large sawtimber stands, but relatively 

abundant in old growth in the Cascades and Coast Ranges of Oregon.” He 

did not cite any evidence for this statement. He suggested that tree voles 

rarely traveled on the ground but provided no data regarding the frequency 

of terrestrial movements. In discussing limiting factors, he suggested that 

old-growth forests are ideal habitat for tree voles because (1) old forests 

are more stable than young forests, (2) primary production of needles 
23

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

is higher in old forests, and (3) old-growth canopies have higher waterholding capacity and intercept fog that voles use as a source of free water 

by licking the needles. In table 4, he classified forests of small sawtimber 

and large sawtimber as population sinks, but provided no data to support 

this hypothesis.

64.   Carey, A.B. 1996. Interactions of northwest forest canopies and arboreal 

mammals. Northwest Science 70. (Special Issue): 72–78.

Carey concluded this literature review with the oft-repeated observation that the red tree vole is the most arboreal mammal in the Pacific 

Northwest. He stated that tree voles have the narrowest niche of Pacific 

Northwest arboreal mammals, require free water in the form of fog or 

rain droplets, and are most abundant in old growth. He speculated that the 

interconnected limbs of old-growth trees facilitate access to other voles 

and serve as escape routes from predators. 

65.   Carey, A.B. 1999. Red tree vole/Arborimus longicaudus. In: Wilson, D.E.; 

Ruff, S. eds. Washington, DC: The Smithsonian book of North American 

mammals. Smithsonian Institution Press: 620–622.

Carey provided a general overview of the ecology and habitat relationships 

of red tree voles. He suggested that old-growth forest is optimum habitat 

for tree voles and that tree voles are declining as a result of logging.

66.   Carey, A.B.; Biswell, B.L.; Witt, J.W. 1991. Methods for measuring populations of arboreal rodents. Gen. Tech. Rep. PNW-GTR-273. 

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

Northwest Research Station. 24 p.

Most of this report is dedicated to sampling methods for northern flying 

squirrels (Glaucomys sabrinus). In two paragraphs that refer to tree voles, 

the authors state that tree voles are hard to study because they can only 

be consistently captured by climbing trees. They suggest that searching 

for nests in felled trees may be a more effective method for sampling tree 

voles than ground-based transects. 

67.   Carleton, M.D.; Musser, G.G. 1984. Muroid rodents. In: Anderson, S.; 

Jones, J.K., Jr., eds. Orders and families of recent mammals of the world. 

New York: John Wiley and Sons: 289–388.

In this book section, the authors agreed with Hall (1981) who argued that 

the scientific name of the red tree vole should be Phenacomys longicaudus. 

This was in contrast to Johnson (1973) and Johnson and Maser (1982), who 

proposed the use of Arborimus longicaudus. 
24

GENERAL TECHNICAL REPORT PNW-GTR-909

68.   Carleton, M.D.; Musser, G.G. 2005. Order Rodentia. In: Wilson, D.E.; 

Reeder, D.M., eds. Mammal species of the world: a taxonomic and geographic reference. 3rd ed. Baltimore, MD: John Hopkins University Press: 

963 p.

Carleton and Musser reversed their 1984 position and classified the red 

tree vole and Sonoma tree vole as Arborimus longicaudus and A. pomo, 

respectively. 

69.   Carraway, L.N.; Verts, B.J. 1982. A bibliography of Oregon mammalogy. Corvallis, OR: Oregon State University, Agricultural Experiment 

Station. 47 p.

This bibliography included many citations for red tree voles and whitefooted voles, all of which we include in the current bibliography.

70.   Carroll, C.; Johnson, D.S.; Dunk, J.R.; Zielinski, W.J. 2010. 

Hierarchical Bayesian spatial models for multispecies conservation planning and monitoring. Conservation Biology. 24: 1538–1548.

The authors compared the relative performance of nonspatial and hierarchical Bayesian spatial models for predicting the probability of occurrence 

of red tree voles and two other species. The best spatial model for red tree 

voles included quadratic terms for percentage slope, tree basal area, maximum tree diameter at breast height (dbh), and standard deviation of conifer 

dbh The best nonspatial models were the same except that the maximum 

tree dbh was a linear function.

71.   Center for Biological Diversity; Cascadia Wildlands; Benton Forest 

Coalition. 2012. Lawsuit filed to protect threatened north Oregon 

coast red tree vole: suit aims to halt plans to clearcut 92 acres of federal 

lands. Press release. 1 p. http//www.biologicaldiversity.org/news/press_

releases/2012/red-tree-vole-09-26-2012.html. (August 2014).

This press release described a legal challenge to the Rickard Creek Timber Sale on the Salem District of the Bureau of Land Management. The 

plaintiffs argued that the sale would harm tree voles that were present 

within the sale area, which included 37 ha of potential clearcut and 8 ha of 

thinning in old forest.

72.   Center for Biological Diversity; Oregon Chapter of the Sierra Club; 

Audubon Society of Portland; Cascadia Wildlands Project; Oregon 

Wild. 2007. Petition to list the dusky tree vole (Arborimus longicaudus 

silvicola) as threatened or endangered under the Endangered Species Act. 
25

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Submitted to the U.S. Fish and Wildlife Service, June 18, 2007.

58 p. http://www.biologicaldiversity.org/species/mammals/dusky_tree_

vole/pdfs/DTV-petition.pdf. (August 2014).

The petitioners argued that U.S. Fish and Wildlife Service should do one 

of three things: (1) list the dusky tree vole as a threatened or endangered 

subspecies, (2) list the north coast population of the red tree vole as a 

threatened or endangered distinct population segment, or (3) list the red 

tree vole as threatened or endangered throughout its range. 

73.   Chaline, J.; Brunet-Lecomte, P.; Montuire, S.; Viriot, L.; Courant, F. 

1999. Anatomy of the arvicoline radiation (Rodentia): palaeogeographical, palaeoecological history and evolutionary data. Annales Zoologici 

Fennici. 36: 239–267. 

In this paper on phylogenetic relationships of the Arvicolinae, the authors 

speculated that the red tree vole evolved from Phenacomys deeringensis, 

an extinct species that occurred in Alaska.

74.   Chaline, J.; Graf, J-D. 1988. Phylogeny of the Arvicolidae (Rodentia): 

biochemical and paleontological evidence. Journal of Mammalogy. 69: 

22–33. 

The authors used genetic distances and dental morphology to compare the 

phylogeny of arvicolids as inferred from the fossil record. In a phenogram 

analysis, the red tree vole was isolated from three other congeners in 

North America, the Sonoma tree vole, white-footed vole, and heather vole. 

Hinton (1926) considered Phenacomys (= Arborimus) as a primitive genus 

and Repenning’s (1983) paleontological data suggested that Phenacomys

was isolated from other voles about 4 million years ago. 

75.   Chavez, A.S.; Kenagy, G.J. 2010. Historical biogeography of western 

heather voles (Phenacomys intermedius) in montane systems of the Pacific 

Northwest. Journal of Mammalogy. 91: 874–885.

The authors used the white-footed vole as one of the outgroups in their 

phylogenetic analysis. They estimated that Phenacomys and Arborimus

diverged approximately 2.45 million years ago.

76.   Chinnici, S.J.; Bigger, D.; Johnson, E. 2012. Sonoma tree vole habitat on managed redwood and Douglas-fir forest lands in north coastal 

California. In: Standiford, R.B.; Weller, T.J.; Piirto, D.D.; Stuart, 

J.D., tech. cords. Proceedings of coast redwood forests in a changing 

California: a symposium for scientists and managers. Gen. Tech. Rep. 
26

GENERAL TECHNICAL REPORT PNW-GTR-909

PSW-GTR-238. Albany, CA: U.S. Department of Agriculture, Forest 

Service, Pacific Southwest Research Station: 389–397.

The authors used line-transect surveys to sample nests of Sonoma tree 

voles in different forest age classes in Humboldt County, California, in 

2001–2005. Because 33 percent of nests were found in unharvested or 

partially harvested old-growth stands, the authors suggested that forest 

managers should retain mature Douglas-fir trees in harvest areas. However, they did not provide specifics regarding the distribution or density of 

leave trees. 

77.   Christensen, J.R.; Larrison, E.J. 1982. Mammals of the Pacific 

Northwest: a pictorial introduction. Moscow, ID: University Press of 

Idaho. 166 p.

On page 90, the authors provided a brief description of the biology and 

distribution of red tree voles and white-footed voles.

78.   Clifton, P.L.; Goodwin, G.G.; Van Gelder, R.G. 1957–1960. 

Unpublished letters. On file with: the American Museum of Natural 

History, New York. (Available upon request from the compilers).

From 1957 through 1960, Percy Clifton sold the American Museum of 

Natural History many specimens including 21 red tree voles, 12 Sonoma 

tree voles, and 3 white-footed voles. We found 36 letters between Clifton 

and the museum curators (George G. Goodwin and Richard G. Van Gelder) 

in the museum archives. Goodwin offered Clifton $10 per tree vole after 

receiving Clifton’s initial inquiry. Clifton responded that he thought $10 

per tree vole was too low because of the effort required to collect them 

(a mild understatement!). In his counter-offer, Goodwin refused to budge 

on the price per tree vole, but agreed to pay for other small mammals, up 

to $100 for all. Upon receiving the specimens, Goodwin sent a letter to 

Clifton complimenting him on the quality of the skins and skulls. After 

receiving the initial shipment, Goodwin apparently decided he had enough 

tree voles because he sent Clifton a letter telling him that he would only 

pay $5 per tree vole in the future, and that what he really needed was more 

white-footed voles. In his final letter, Clifton wrote Goodwin that he had 

collected three white-footed voles near Tillamook, Oregon, after several 

hundred trap-nights. 

79.   Clifton, P.L.; Reeder, W.; Ghiselin, J. 1959–1960. Unpublished letters. 

On file with: University of Wisconsin Zoological Museum. 

(Available upon request from the compilers).
27

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

We found five letters at the University of Wisconsin Zoological Museum 

that documented specimen transactions between Percy Clifton and 

museum curators (W. Reeder and Jon Ghiselin) in 1958–1960. Percy sold 

them nine red tree voles and one Sonoma tree vole. Some comments from 

Reeder and Ghiselin indicated that they were not happy because some 

of the specimens were captive-bred, the skulls and skeletons were not 

cleaned, and data were “scant.” 

80.   Clifton, P.L. 1960. Biology and life history of the dusky tree mouse 

Phenacomys silvicola (Howell). Walla Walla, WA: Walla Walla College. 

140 p. M.A. thesis. 

In this thesis, Clifton produced an amazing amount of quantitative data on 

the reproductive biology, natal development, and behavior of tree voles. 

Although it was produced in 1960, this thesis was apparently unknown to 

all mammalogists who worked on tree voles during the next four decades, 

because it was never cited until Clifton brought it to the attention of Jerry 

Mires at the Roseburg District of the Bureau of Land Management in 2001. 

By that time, Clifton had recently retired from his job as a medical doctor 

and was living near Sutherlin, Oregon. 

In his introduction, Clifton stated that tree voles were not colonial in the 

traditional sense, but were usually found in proximity to each other. He 

captured about 33 adult tree voles and several young tree voles from a 

study area near the town of Gaston in the Chehalem Mountains of Oregon 

in 1956–1959. He kept the voles in captivity in wire cages constructed of 

1/2-inch hardware cloth. His captive voles produced 25 litters in captivity, 

including three litters from females that were pregnant when captured, 

and 22 litters from females that were bred in captivity. Clifton also collected tree voles from several other locations, including Roseburg, Douglas 

County, Oregon, Cottage Grove, Lane County, Oregon, and Albion, Mendocino County, California. Based on his observations of captive voles, he 

estimated that the average gestation period of red tree voles was 28 days, 

14 hours (n = 6), and that the average gestation of hybrid crosses between 

red tree voles and Sonoma tree voles was 26 days, 21 hours (n = 3). Some 

of his wild-caught adult voles lived over 2 years in captivity. 

Clifton searched for voles primarily in young forests of Douglas-fir. He 

captured voles by grabbing them as they fled the nest or by having an 

assistant grab the voles when they jumped from the nest. He stated that 

even if a vole escaped on the first capture attempt, he could usually return 
28

GENERAL TECHNICAL REPORT PNW-GTR-909

the next day and capture it from the same nest. He fed voles needles on 

capture but gradually weaned many of them onto a diet of rolled oats and 

apple as did Benson and Borell (1931). He provided measurements of 33 

dusky tree voles and described how their rusty brown pelage differed from 

the red pelage of red tree voles. He said that compared to dusky tree voles, 

red tree voles were more inclined to jump from the nest, were swifter runners, and built larger nests. He also said that dusky tree voles had smaller 

litters (usually two) than red tree voles (usually two or more). He compared 

cranial characteristics between the subspecies.

Clifton captured 16 of his adult specimens 6.4 km east of Gaston, Yamhill 

County, Oregon, on the northern end of Ribbon Ridge about 3.2 km north 

of the Laurelwood Academy. He described the habitat as young stands of 

dense Douglas-fir that were 12 to 24 m tall, with dense canopies and little 

undergrowth. He captured another 12 adult specimens approximately 6.4 

km north of Newburg, directly below the gravel road that runs along the 

top of Ribbon Ridge, about 9.7 km southeast of Bald Peak State Park. He 

suggested that tree voles were at one time present the full length of the 

ridge, but at the time of his study, voles were only present in a few remnant 

stands of young trees that had not been clearcut. He suggested that many 

tree voles were killed during logging operations, but he provided no direct 

evidence except that he could find no tree voles or nests in the few remaining trees in areas that had been logged. 

He suggested that tree voles preferred dense forests because they provided 

protection from predators and had interconnecting branches that allowed 

the voles to travel from tree to tree instead of descending to the ground. 

Regarding the young forests in which he collected most of his voles he 

says, “They are easily found in second- or third-growth trees, but they may 

be present in old-growth. This so-called third-growth is characterized by 

dense, small, fir trees on which the limbs often come to the ground.”

Regarding predators, Clifton speculated that northern flying squirrels may 

prey upon tree voles. He described tree vole nests torn apart by unknown 

predators and said that he found tree vole skulls below hawk and owl nests. 

He also reported the top of a tree vole nest torn off and flattened by a raccoon (Procyon lotor) that he saw sitting on the nest. 

He provided a detailed description of how tree voles removed resin ducts 

from needles of Douglas-fir and how they ate twig bark. He said that a 

male vole always passed the needle from left to right through its mouth 
29

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

when removing the resin ducts. He also used a microscope to determine 

that voles discarded the vascular bundle or midrib of western hemlock 

needles after eating the outer edges. He said that voles collected in Douglas-fir near Gaston would eat either Douglas-fir or western hemlock and 

seemed to have no preference for one or the other. 

He conducted feeding trials with captive voles and estimated that when a 

vole was fed only needles, its daily intake was 22.9 g per day. Foods that 

were refused by his captive tree voles were white fir (Abies concolor), 

Pacific yew (Taxus brevifolia), Engelmann spruce (Picea engelmannii), 

lodgepole pine (Pinus contorta), fresh meat, carrots, lettuce, blackberries, 

wild rose fruits, hazelnuts, lichens, mushrooms, moss, canned fruit, and 

many others. 

He said that stomachs of wild-caught tree voles contained only finely 

pulverized needles and little bits of twig bark. He found no lichens in vole 

stomachs, in contrast to the single stomach examined by Wight (1925). 

He observed that voles often tried to escape by crossing from tree to tree 

on interlocking limbs, and he suggested that they probably used the same 

limbs to forage in adjacent trees. 

He said that he observed captive voles lapping up drops of water adhering 

to conifer needles. He also reported that voles that were fed fresh needles 

had wet fecal pellets, whereas voles that were fed dry needles had comparatively dry fecal pellets. 

He stated that most tree vole nests near Gaston were built in or on deserted 

nests of squirrels, hawks, or birds, and that the majority of nests were 

against the trunk of the tree. He found no preference regarding slope aspect 

or which side of the bole the nest was located. He never found more than 

one occupied nest in the same tree.

He described numerous parasites, including mites, fleas, ticks, and tapeworms. He also mentioned, but did not describe, large numbers of arthropods found in nests. He observed a disease that killed many captive voles 

that had fluke-like parasites in their nasal passages. He also mentioned 

hemoflagellates and a contagious fungal infection.

He said that temperament varied considerably among individual voles. In 

most cases, tree voles placed in the same cage did not tolerate each other. 

Females briefly tolerated males in their cages when they were ready to 

breed but would attack and drive off males on most other occasions. On 
30

GENERAL TECHNICAL REPORT PNW-GTR-909

one occasion, a female attacked and killed a male that was left with her 

for 2 to 3 hours. Males were also intolerant of each other. However, female 

siblings were sometimes kept together for long periods of time with no 

evidence of aggression. 

Based on a female that had five litters in a year, Clifton suggested that six 

to seven litters per year may be possible. He described a female born in 

late February that mated on 27 May and produced young on 26 June, when 

she was about 4 months old. He suggested that the breeding season starts 

as early as the first of January and ends in August, with most tree voles 

producing no young in September through December. He found that the 

sex ratio at birth was about 50:50 and suggested that the preponderance 

of females captured by tree climbers was because female nests were more 

obvious. He found no evidence that males were more likely to flee the nest 

than were females. 

81.   Clothier, C. 1960. Pen and ink drawings. On file with: University of 

Washington Burke Museum, Seattle, WA.

In the 1960s, Carol Clothier produced a series of unique pen-and-ink drawings of red tree voles for Murray Johnson. We found the drawings among 

Murray’s archived papers at the Burke Museum. Clothier’s models for the 

drawings were captive tree voles collected by Chris Maser. 

82.   Cole, E.C.; McComb, W.C.; Newton, M.; Leeming, J.P.; Chambers, 

C.L. 1998. Response of small mammals to clearcutting, burning, and 

glyphosate application in the Oregon Coast Range. Journal of Wildlife 

Management. 62: 1207–1216.

The authors used pitfall traps and Sherman live-traps to sample small 

mammals in the Oregon Coast Ranges in 1988–1992. In 50,256, trapnights they captured one red tree vole and one white-footed vole. They did 

not say whether they caught the voles in pitfall traps or live-traps. 

83.   Collins, P.W. 1998. White-footed vole, Arborimus albipes. In: Bolster, 

B.C., ed. Terrestrial mammal species of special concern in California. 

California Fish and Game. Available from: http://www.dfg.ca.gov/wildlife/

nongame/ssc//1998mssc.html. (August 2014).

In this species account, Collins described the taxonomy, distribution, life 

history, habitat, and conservation status of the white-footed vole in California and included management recommendations for the species.
31

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

84.   Conroy, C.J.; Cook, J.A. 1999. MtDNA evidence for repeated pulses of 

speciation within arvicoline and murid rodents. Journal of Mammalian 

Evolution. 6: 221–245.

The authors suggested that the Arvicolinae were monophyletic, first 

appeared in the late Miocene, and then rapidly diversified 3 to 5 million 

years ago. They used the heather vole as their Phenacomys sample and did 

not mention tree voles or white-footed voles.

85.   Coriell, T.R. 1974. A comparison of the climbing behavior of Arborimus 

longicaudus and Microtus californicus. Arcata, CA: Humboldt State 

University. 24 p. M.A. thesis. 

Coriell compared the climbing behavior of tree voles and California 

voles (Microtus californicus). His sample of tree voles included 3 that he 

captured, 19 captive-bred and 3 wild-caught voles that he obtained from 

Murray Johnson. He concluded that tree voles were better adapted for 

climbing than were terrestrial voles. No data were presented on the actual 

mechanics of climbing. 

86.   Corn, P.S.; Bury, R.B. 1984–1985. Oregon old-growth project pitfalltrap locality order. Unpublished document. On file with: the U.S. National 

Museum, Washington, DC.

This document describes the locations of the pitfall trapping grids in which 

Corn and Bury sampled small mammals in Oregon. From 1984 through 

1985, they trapped 31 red tree voles and two white-footed voles in the 

central Coast Ranges, Oregon. 

87.   Corn, P.S.; Bury, R.B. 1986. Habitat use and terrestrial activity by red 

tree voles (Arborimus longicaudus) in Oregon. Journal of Mammalogy. 

67: 404–406.

Corn and Bury reported that capture rates of tree voles in pitfall traps were 

highest in old-growth forests. These were the same data reported in Bury 

and Corn (1987) and Corn and Bury (1988), but with much more information on the locations of captured voles. Of 2,104 mammals captured, 18 

(0.86 percent) were red tree voles. However, the authors only used 17 of 

the tree voles in their analysis of habitat associations, because one vole 

was captured in a test grid that was not part of the main study. All tree 

voles captured in old-growth were in mesic or wet sites. Elevations of tree 

vole capture locations ranged from 510 to 1,040 m. The total number of 

trap-nights was 38,880, and the number of tree voles captured per 100 
32

GENERAL TECHNICAL REPORT PNW-GTR-909

trap-nights was 0.046. One tree vole was captured in a clearcut 200 to 300 

m from the nearest forest edge and the authors speculated that it may have 

been a dispersing individual. They concluded that, “Although the sample 

size is small, our data support the contention that, A. longicaudus finds 

optimum habitat in wet and mesic old-growth forests.” Most of the voles 

were captured in the late spring and late autumn. Relatively few were 

captured during mid-summer. Of the 18 tree voles captured, 78 percent 

were males. The authors speculated that the disproportionate sex ratio 

of captured voles could have been due to (1) males using terrestrial nests 

more often than females, (2) males having larger home ranges, or (3) males 

dispersing farther than females. They did not consider the alternative 

hypothesis that males were spending more time on the ground than females 

because they were frequently traveling between trees to seek out females.

88.   Corn, P.S.; Bury, R.B. 1988. Distribution of the voles Arborimus longicaudus and Phenacomys intermedius in the central Oregon Cascades. 

Journal of Mammalogy. 69: 427–429.

This paper was based on the same data described in Corn and Bury (1986) 

but included additional information on capture locations. Based on the 

proximity of captures of red tree voles and heather voles in the Cascade 

Range, the authors suggested some degree of spatial overlap between 

the two species. They suggested a scenario in which “...A. longicaudus

originally occurred in old-growth forests along the river drainages at lower 

elevations, but its distribution probably was reduced by extensive logging 

of this habitat. The original primary habitats of P. intermedius were above 

timber line and high-elevation meadows, but this species is extending its 

range west along the higher ridges as they are clearcut. Occasionally, there 

is a small amount of overlap, but, in general, P. intermedius and A. longicaudus would have interdigitated parapatric distributions along the Cascade Range south from the Columbia River to the area between McKenzie 

Pass and Willamette Pass.” 

89.   Corn, P.S.; Bury, R.B. 1991. Small mammal communities in the Oregon 

Coast Range. In: Ruggiero, L.F.; Aubry, K.B.; Carey, A.B.; Huff, M.H., 

tech. coords. Wildlife and vegetation of unmanaged Douglas-fir forests. Gen. Tech. Rep. PNW-GTR-285. Portland, OR: U.S. Department of 

Agriculture, Forest Service, Pacific Northwest Research Station: 241–254.
33

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

The authors used pitfall traps to sample 48 sites in the Oregon Coast 

Ranges, including 3 clearcuts, 8 young stands (40 to 75 years), 10 mature 

stands (80 to 120 years) and 27 old-growth stands (150 to 525 years). 

They opened traps in October and sampled continuously for 50 days in 

1984 and 30 days in 1985. They captured 3,047 mammals, including 13 

(0.43 percent) red tree voles. All but one of the tree voles were captured 

in old-growth. In their discussion, they said that, “The 17 red tree voles 

we trapped in the Coast Ranges constitute a small sample, but since 1983, 

46 red tree voles have been trapped in the Coast and Cascade Ranges 

of Oregon by old-growth researchers (this study; Corn and Bury 1986; 

Gilbert and Allwine 1991). Most (n = 39) were captured in old-growth 

stands and because the trapping effort was about evenly split between 

old-growth stands and younger stands, a significant excess of captures 

were in old-growth.” They also reported that Raphael (1988) captured an 

additional 19 Sonoma tree voles in pitfall traps in Douglas-fir forests in 

northwestern California. Although the difference in relative abundance 

among stand age-classes was not significant, most Sonoma tree voles 

captured by Raphael were in stands older than 100 years. They also stated 

that Meiselman and Doyle (1996) found that nests of Sonoma tree voles in 

northwestern California were more abundant in old-growth forests than 

in younger forests. Based on these collective observations, they concluded 

that “These results indicate that red tree voles are strongly associated with 

old-growth Douglas-fir forests.”

90.   Corn, P.S.; Bury, R.B.; Spies, T.A. 1988. Douglas-fir forests in the 

Cascade Mountains of Oregon and Washington: is the abundance of small 

mammals related to stand age and moisture? In: Szaro, R.C.; Severson, 

K.E.; Patton, D.R., tech. cords. Management of amphibians, reptiles and 

small mammals in North America. Gen. Tech. Rep. RM-166. Fort Collins, 

CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain 

Forest and Range Experiment Station: 340–352.

This paper was based on the same data as Bury and Corn (1987), Corn and 

Bury (1986), and Corn et al. (1988). They sampled 18 different stands on 

the H.J. Andrews Experimental Forest, including 8 old-growth stands (195 

to 450 years), 3 mature stands (80 to 195 years), 4 young stand (30 to 80 

years), and 3 recent clearcuts. They captured 17 red tree voles, including 

12 in old-growth forest, 3 in mature forest, 1 in young forest, and 1 in a 
34

GENERAL TECHNICAL REPORT PNW-GTR-909

clearcut. They stated that tree voles were associated with old-growth forest 

and speculated that, “...the extensive logging of low-elevation old-growth 

forests in Oregon has probably eliminated much of the habitat of red tree 

voles. The giant Douglas-fir trees, which seem to be preferred as nest sites, 

will not occur in managed forests.”

91.   Coughenour, J.C. 2009. Conservation Northwest; Gifford Pinchot 

Task Force; Environmental Protection Information Center; Klamath 

Forest Alliance; Umpqua Watersheds, Inc.; Siskiyou Regional Education 

Project; Klamath-Siskiyous Wildlands Center; Oregon Wild American 

Lands Alliance; Center for Biological Diversity; and Nothcoast [sic] 

Environmental Center v. Mark E. Rey; C. Stephen Allred; Synthia U. 

Barry; U.S. Forest Service; U.S. Bureau of Land Management; U.S. Fish 

and Wildlife Service, and D.R. Johnson Lumber Company. U.S. District 

Court, Western District of Washington at Seattle. Case No. C08-1067-

JCC. 36 p.

In this court case, the plaintiffs challenged the 2004 supplemental environmental impact statement (EIS; USDA FS and USDI BLM 2004) and the 

2007 final EIS (FEIS; USDA FS and USDI BLM 2007) in which the Forest 

Service and Bureau of Land Management decided to remove the “survey 

and manage” requirements from the Northwest Forest Plan. The goal of the 

2007 FEIS was to reduce cost, time, and effort required to conserve rare 

and little-known species. In the 2007 FEIS, the agencies concluded that 

survey and manage, “...has unexpectedly limited the Agency’s ability to 

accomplish the goals and objectives of the Northwest Forest Plan,” and that 

the best remedy would be to cease to implement the survey and manage 

requirements entirely. The court did not issue a final verdict and directed 

the parties to confer about appropriate case management.

92.   Coughenour, J.C. 2011. Conservation Northwest et al. v. Harris Sherman 

et al. and D.R. Johnson Lumber Company. Parties’ stipulation and proposed order. U.S. District Court, Western District at Seattle. Case No. 

C08-1067-JCC. 3 p.

In this settlement agreement, the plaintiffs and the defendants agreed that 

the 2001 Record of Decision and Standards and Guidelines for survey and 

manage species would remain in effect. This ruling meant that the U.S. 

Forest Service and Bureau of Land Management had to continue conducting predisturbance surveys for red tree voles. However, the court also 
35

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

accepted a long list of exemptions for situations in which surveys were 

not required, including the exemptions agreed to in a previous settlement 

between Northwest Ecosystem Alliance v. Rey (see Pechman 2006).

93.   Courtney, S.P.; Blakesley, J.A.; Bigley, R.E.; Cody, M.L.; Dumbacher, 

J.P.; Fleischer, R.C.; Franklin, A.B.; Franklin, J.F.; Gutiérrez, R.J.; 

Marzluff, J.M.; Sztukowski, L. 2004. Scientific evaluation of the status

of the northern spotted owl. Portland, OR: Sustainable Ecosystems 

Institute. 508 p.

This broad-ranging review of the biology of the northern spotted owl, 

included a section (appendix pages 3-14 to 3-15) in which the authors 

described the habitat associations and distribution of red tree voles. They 

relied primarily on data from pitfall trapping studies and ground-based 

survey transects to reach the conclusion that tree voles were most abundant 

in old-growth forest. However, they also included a personal communication from Eric Forsman, who suggested that pitfall traps might be a biased 

estimator of tree vole abundance in different forest types. They stated 

that searching of felled trees (Gillesberg and Carey 1991) may be the most 

cost-effective method to survey large areas but did not present any kind 

of cost-benefit comparison between that approach and other approaches. 

They also stated that tree voles were colonial but presented no evidence to 

document that assertion. 

94.   Csuti, B.; Kimerling, A.J.; O’Neil, T.A.: Shaughnessy, M.M.; Gaines, 

E.P.; Huso, M.M.P. 1997. Red tree vole (Phenacomys longicaudus). Atlas 

of Oregon wildlife: distribution, habitat, and natural history. Corvallis, 

OR: Oregon State University Press. 403 p.

The authors presented a red tree vole distribution map for western Oregon. 

In the text, they described the range, habitat, reproduction, food habits, and 

ecology of tree voles. They incorrectly stated that the range of the Sonoma 

tree vole extended south to San Francisco Bay, when it actually ends near 

Freestone and Bodega in Sonoma County, well north of San Francisco Bay. 

Their range map also included areas where recent surveys suggest that tree 

voles do not occur, including most of Clatsop County and all of Columbia 

County, and most of western Multnomah County. They also stated that tree 

voles feed on true firs (Abies spp.), which is only partly true, as tree voles 

have only been documented to eat one species of true fir, grand fir. They 

concluded by saying that the distribution is discontinuous and that tree 

voles usually occur in old forest.
36

GENERAL TECHNICAL REPORT PNW-GTR-909

95.   Csuti, B.; Kimerling, A.J.; O’Neil, T.A.; Shaughnessy, M.M.; Gaines, 

E.P.; Huso, M.M.P. 1997. Atlas of Oregon wildlife: distribution, habitat, 

and natural history. Corvallis, OR: Oregon State University Press: 402.

This species account described the range, habitat, reproduction, food 

habits, ecology, and conservation status of the white-footed vole in Oregon. 

They also presented a range map based on locations and habitat modeling.

96.   Dalquest, W.W. 1943–1944. Unpublished field notes. On file with: 

University of California, Berkeley, Museum of Vertebrate Zoology.

Dalquest’s notes from 1943 through 1944 recorded that he, his wife Peggy, 

and Bob Storer collected nine Sonoma tree voles at Camp Meeker and 

Monte Rio in Sonoma County, California. Like many before him, he mistakenly concluded that discarded resin ducts were midribs of Douglas-fir 

needles. He noted postpartum mating after capturing an adult female that 

had one embryo and was nursing. He documented an occupied Sonoma 

tree vole nest that also contained two arboreal salamanders.

97.   Dash, J.A.; Jenness, R. 1985. Ascorbate content of foliage of eucalypts 

and conifers utilized by some Australian and North American mammals. 

Experientia. 41: 952–955.

The authors cited Johnson (1973) to support their conclusion that the 

red tree vole is one of the most obligate folivorous mammals in North 

America. They compared ascorbate levels in eucalypts and conifers, 

including Douglas-fir and suggested that “...high ascorbate intakes by 

animals feeding on evergreen foliage may facilitate the metabolism or 

excretion of undesirable compounds (e.g., phenols, terpenes) that are also 

in the foliage.”

98.   Davis, W.B. 1936–1940. Unpublished field notes. On file with: University 

of California, Berkeley, Museum of Vertebrate Zoology.

In his notes from 1936 to 1940, William Davis described the capture of 

seven Sonoma tree voles near Occidental, Sonoma County, California. 

Some of these voles were kept alive in captivity, but all died within 2 to 6 

days. 

99.   Dearden, L.C. 1955. The genus Lagurus, anatomy and taxonomy. Salt 

Lake City, UT: University of Utah. 195 p. Ph.D. dissertation.

Dearden used red tree vole specimens from the U.S. National Museum in 

his analysis of taxonomy and anatomy of voles in the genus Lagurus.
37

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

100.  Dearden, L.C. 1958. The baculum in Lagurus and related microtines. 

Journal of Mammalogy. 39: 541–553.

In this published version of his dissertation, Dearden examined the bacula 

of three red tree voles. The bacula that he examined were markedly different from the tree vole baculum pictured in Hamilton (1946: 370), and 

Dearden goes on at great length to describe the differences. 

101.   Denison, W.C. 1973. Life in the tall trees. Scientific American. 228: 

74–80.

With regard to the foliose lichen Lobaria spp, Denison said that “Certain 

vertebrates, such as the rodent called the red tree vole, supplement their 

diet with this lichen, among other plants.” He did not say how he came to 

this conclusion or cite a reference that tree voles eat lichens. Wight (1925) 

is the only article that we are aware of to claim that the stomach of a tree 

vole contained lichen. 

102.  Dizney, L.; Jones, P.D.; Ruedas, L.A. 2008. Efficacy of three types of 

live traps used for surveying small mammals in the Pacific Northwest. 

Northwestern Naturalist. 89: 171–180.

The authors compared capture and mortality rates among Sherman traps, 

custom-made steel-mesh traps, and pitfall traps. The traps were used 

around the Portland metropolitan area at Forest Park, Tryon Creek State 

Park, Powell Butte Nature Park, Oxbow Regional Park, and Tualatin River 

National Wildlife Refuge. In 65,600 trap-nights, they captured 2,710 small 

mammals, none of which were red tree voles or white-footed voles.

103.  Doyle, A.T. 1985. Small mammal micro- and macrohabitat in streamside ecosystems. Corvallis, OR: Oregon State University. 211 p. Ph.D. 

dissertation. 

In 1981–1983, Doyle used Sherman live-traps to study small mammal 

abundance in different forest habitats on the H.J. Andrews Experimental 

Forest near Blue River, Lane County, Oregon (40,152 trap-nights). She also 

attempted to evaluate competitive interactions between small mammals 

by removing deer mice from some sampling grids (52,752 trap-nights). In 

92,904 trap-nights, she captured one white-footed vole and no tree voles.

104.  Doyle, A.T. 1990. Use of riparian and upland habitats by small mammals. 

Journal of Mammalogy. 71: 14–23. 

This was the published version of the habitat association section of Doyle’s 

dissertation. She used Sherman live-traps to sample small mammals on the 
38

GENERAL TECHNICAL REPORT PNW-GTR-909

H.J. Andrews Experimental Forest and did not capture any red tree voles 

or white-footed voles during 40,152 trap-nights in 1981–1983. 

105.  Dunk, J.R.; Hawley, J.J.V.G. 2009. Red-tree [sic] vole habitat suitability 

modeling: implications for conservation and management. Forest Ecology 

and Management. 258: 626–634.

This paper was based on the random-grid strategic survey conducted by 

the U.S. Forest Service and Bureau of Land Management Survey and 

Manage Program (Rittenhouse et al. 2002). The authors modeled red 

tree vole habitat associations based on presence or absence of tree vole 

nests in a stratified random sample of Forest Inventory and Analysis and 

Current Vegetation Survey plots. Plots were stratified into four groups 

based on forest age and land use allocations in the Northwest Forest Plan. 

The percentage of plots in each category was 60 percent in reserve/latesuccessional old growth (LSOG), 20 percent in reserve/non-LSOG, 10 

percent in nonreserve/LSOG, and 10 percent in nonreserve/non-LSOG. 

The best models included spatial coordinates, percentage slope, basal area 

of trees 45 to 90 cm diameter breast height (dbh), maximum tree dbh, and 

standard deviation of conifer dbh. The authors concluded that presence of 

red tree vole nests was strongly associated with old-growth/late-seral forest 

conditions. Of the 365 1-ha plots in the sample, 96 (26 percent) had one or 

more trees with a tree vole nest. The estimated rate of false negatives (nests 

present but undetected during ground surveys) was 6 percent.

106.  Eisenberg, J.F. 1978. The evolution of arboreal herbivores in the class 

Mammalia. In: Montgomery, G.G., ed. The ecology of arboreal folivores. 

Washington, DC: Smithsonian Institution Press: 135–152.

In table 2, Eisenberg rated the red tree vole as highly arboreal and highly 

herbivorous. He concluded with, “The surprising thing to me is that in 

the Holarctic temperate zone, more species of rodents have not evolved to 

arboreal, folivorous niches utilizing conifers in the manner that Arborimus 

longicaudus has in the northwest coast of North America.”

107.   Fain, A.; Whitaker, J.O., Jr. 1975. Two new species of Myobiidae from 

North American mammals (Acarina). Bulletin et Annales de la Societe 

Royale d’ Entomologie de Belgique. 111: 57–65. 

The authors described a new species of mite (Radfordia arborimus) found 

on a red tree vole collected by Chris Maser.
39

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

108.  Fisher, H.I. 1942. Unpublished field notes. On file with: University of 

California, Berkeley, Museum of Vertebrate Zoology.

In his field notes for 8 September 1942, Fisher said that he caught a male 

tree vole in a snap-trap set in a brush pile beneath a Douglas-fir that had an 

apparent tree vole nest.

109.   Fleming, T.H.; Rauscher, R.J. 1978. On the evolution of litter size in 

Peromyscus leucopus. Evolution. 32: 45–55. 

The authors found that birth weight in the white-footed mouse (Peromyscus leucopus) declined with larger litter size. They pointed out that 

Hamilton (1962) found that birth weights also declined with litter size in 

the Sonoma tree vole.

110.   Fontaine, J.B. 2007. Influences of high severity fire and postfire logging 

on avian and small mammal communities of Siskiyou Mountains, Oregon, 

USA. Corvallis, OR: Oregon State University. 152 p. Ph.D. dissertation. 

In 2004–2005, Fontaine used Sherman live-traps to sample small mammals in the Siskiyou Mountains after the Biscuit Fire, which occurred 

in 2002. In 8,900 trap-nights he captured one tree vole in an unburned 

mature stand (0.01 tree vole captures per 100 trap-nights).

111.   Forsman, E.D. 1975. A preliminary investigation of the spotted owl in 

Oregon. Corvallis, OR: Oregon State University. 127 p. M.S. thesis. 

Forsman described diets of northern spotted owls in different regions of 

western Oregon based on a sample of 2,647 prey items found in regurgitated pellets collected from 42 pairs of owls. He found that red tree voles 

comprised 30.6 percent of the diet in the Coast Ranges, 12.5 percent of the 

diet in the western Cascades, 12.7 percent of the diet at Oregon Caves, and 

8.5 percent of the diet in the eastern Siskiyou Mountains. He concluded 

that spotted owls were an important predator on tree voles.

112.  Forsman, E.D. 1980a. Habitat utilization by spotted owls in the westcentral Cascades of Oregon. Corvallis, OR: Oregon State University. 95 p. 

Ph.D. dissertation. 

Forsman summarized annual and seasonal variation in diets of northern 

spotted owls on the H.J. Andrews Experimental Forest in Lane County, 

Oregon. Of 753 prey identified in owl pellets, 97 (12.9 percent) were red 

tree voles. Northern flying squirrels and tree voles were the most common 

prey items in the diet during winter, when other sources of food were less 
40

GENERAL TECHNICAL REPORT PNW-GTR-909

available. Based on tree voles in pellets, Forsman estimated that a nesting 

pair of spotted owls on the H.J. Andrews study area would consume 184 

tree voles per year. 

113.  Forsman, E.D. 1980b. Habitat utilization by spotted owls on the 

Eugene District of the Bureau of Land Management. Eugene, OR: U.S. 

Department of the Interior, Bureau of Land Management. 62 p. (Available 

upon request from the compilers).

Forsman identified 256 prey items in pellets of three radio-marked pairs of 

northern spotted owls in 1980. Of the 256 prey items, 24 (9.4 percent) were 

red tree voles.

114.   Forsman, E.D.; Anthony, R.G.; Meslow, E.C.; Zabel, C.J. 2004a. 

Regional, local, and annual variation in diets of spotted owls in Oregon 

[Abstract]. In: 39th annual meeting of the Oregon Chapter of the Wildlife 

Society. Bend, OR. (Available upon request from the compilers).

Of the 23,847 prey items identified in northern spotted owl pellets, 54 

percent were arboreal mammals, including red tree voles. In addition, the 

authors found five white-footed voles in spotted owl pellets. The authors 

suggested that forest management practices that were beneficial to arboreal 

and scansorial mammals would also be beneficial to spotted owls.

115.  Forsman, E.D.; Anthony, R.G.; Meslow, E.C.; Zabel, C.J. 2004b. Diets 

and foraging behavior of northern spotted owls in Oregon. Journal of 

Raptor Research. 38: 214–230.

Of 23,847 prey items identified in pellets of northern spotted owls collected from 1970 through 2001, 2,954 (12.3 percent) were red tree voles, 

and 5 (0.02 percent) were white-footed voles. 

116.   Forsman, E.D.; Anthony, R.G.; Zabel, C.J. 2004c. Distribution and 

abundance of red tree voles in Oregon based on occurrence in pellets of 

northern spotted owls. Northwest Science. 78: 294–302.

The authors identified 24,497 prey items from 1,118 northern spotted owl 

territories in western Oregon. Of these, 2,954 (12.3 percent) were red tree 

voles. The analysis indicated that tree voles were widely distributed in 

western Oregon but were most common in the south and central Coast 

Range and central Cascades. Tree voles were uncommon in diets of owls 

that occupied areas above 1200 m in the central Cascades.

117.   Forsman, E.D.; Horn, K.M.; Mires, G.W. 1985. Northern spotted owls. 

In: Brown, E.R., tech. ed. Management of wildlife and fish habitats in 
41

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

forests of western Oregon and Washington. Part 1: Chapter narratives. 

Publication No. R6-F&WL-192-1985. Portland, OR: U.S. Department of 

Agriculture, Forest Service: 259–267. 

The authors reported that a sample of 817 prey items from northern spotted 

owl pellets in the western Cascades of Oregon contained 13 percent red 

tree voles. A collection of 654 prey items from spotted owl pellets in 

mixed-conifer forest contained 5 percent tree voles.

118.  Forsman, E.D.; Maser, C. 1970. Saw-whet owl preys on red tree mice. 

Murrelet. 51: 10.

The authors described prey remains identified in 36 pellets from northern saw-whet owls (Aegolius acadicus) in a 30- to 50-year-old stand of 

Douglas-fir 4.0 km north of Bellfountain, Benton County, Oregon. Of 42 

prey identified, 3 (7 percent) were tree voles.

119.   Forsman, E.D.; Meslow, E.C.; Wight, H.M. 1984. Distribution and

biology of the spotted owl in Oregon. Wildlife Monographs. 87: 3–64.

The authors described the diets of northern spotted owls in western Oregon 

based on 4,527 prey items. They reported that the mean percentage of red 

tree voles in the diet varied from 5 to 49 percent in different regions of 

western Oregon.

120.  Forsman, E.D.; Mowdy, J.S.; Price, A.L.; Swingle, J.K. 2012. Survey 

of red tree voles on the Weyerhaeuser Millicoma Tree Farm in Coos 

and Douglas Counties, Oregon, 2011. 16 p. Unpublished report. On file 

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

Research Station, 3200 SW Jefferson Way, Corvallis, OR 97331. 

(Available upon request from the compilers).

In 2011, the authors searched for tree vole nests while driving or walking 

along 176 km of logging roads on the Weyerhaeuser Millicoma Tree Farm 

in Coos and Douglas County, Oregon. Of 132 potential nests detected, 52 

percent were climbed to see if they were built by tree voles. Only three 

occupied nests were found, all in young stands that were 30 to 36 years 

old. This survey revealed that tree voles were uncommon in the young 

forests that dominated the tree farm, but were still present in a few areas.

121.   Forsman, E.D.; Otto, I.A. 2006. Healed fractures and other abnormalities in bones of small mammals. Northwestern Naturalist. 87: 143–146.

Of 21,721 small mammals identified in pellets of northern spotted owls, 

15,455 were arboreal or scansorial, 6,199 were primarily terrestrial, and 
42

GENERAL TECHNICAL REPORT PNW-GTR-909

67 were bats. Of 2,953 tree voles in the sample, only 2 (0.07 percent) had 

healed fractures. The authors concluded that tree voles were so small and 

light that they rarely broke bones when they fell or jumped from trees. 

However, the authors did not consider that tree voles may be unable to 

climb or feed after breaking legs, so the frequency of injuries from falls 

may be underestimated based on the frequency of healed fractures in owl 

pellets.

122.  Forsman, E.D.; Price, A.L. 2011. Water consumption by red tree voles 

(Arborimus longicaudus). Northwestern Naturalist. 92: 116–119.

Forsman and Price measured water consumption by red tree voles in a 

laboratory setting where the voles were fed a natural diet of well-hydrated 

conifer needles. Under these conditions, the voles drank little free water 

and sometimes went for weeks or months without drinking any water. 

The authors concluded that tree voles obtained nearly all their water from 

their food. Although observations of tree voles licking water from needles 

in laboratory conditions have led many authors to suggest that tree voles 

require free water in the form of dew or rain in order to survive (Benson 

and Borell 1931; Carey 1991, 1996, 1999; Clifton 1960; Maser 1966; 

Taylor 1915), there is actually no evidence that tree voles in the wild need 

significant amounts of water other than what they obtain from their food. 

Tree voles and other species of voles that drink copious amounts of water 

in laboratory settings may be responding to conditions in which they are 

maintained in unnatural conditions with low humidity and food that is low 

in water content.

123.  Forsman, E.D.; Swingle, J.K. 2004. Percy Clifton interview, 13 

December 2004. 3 p. Unpublished notes. On file with: U.S. Department 

of Agriculture, Forest Service, Pacific Northwest Research Station, 3200 

SW Jefferson Way, Corvallis, OR 97331. (Available upon request from the 

compilers).

These notes described an interview conducted on 13 December 2004 

with Percy Clifton at his residence on Calapooya Creek east of Sutherlin, 

Douglas County, Oregon. The notes describe how Percy first became 

interested in tree voles and eventually ended up completing a master’s 

thesis on tree voles at Walla Walla College in Washington (Clifton 1960). 

Clifton grew up near Gaston, Washington County, Oregon, where as a 

child he liked climbing trees to capture northern flying squirrels and tree 

voles. In a biology class at Walla Walla College, professor Ernest Booth 
43

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

showed the students a picture of a tree vole and said that it was an interesting species that they would likely never see. After class, Clifton approached 

Booth and told him that not only had he seen tree voles, but he had captured 

them and kept them as pets. Booth was skeptical until Percy went home on 

break and returned to school with a tree vole. This eventually led Booth to 

accept Percy as a graduate student. For his master’s project, Percy captured 

tree voles at various locations in Oregon and California and raised them 

in captivity. His wide-ranging thesis is probably the most detailed paper 

ever written on the reproductive biology and behavior of tree voles. After 

graduating from college, Clifton made a living for a few years collecting 

specimens in Mexico for Raymond Hall at the University of Kansas. He 

then went back to school and became a medical doctor. He retired in 1999 

and settled on Calapooya Creek. As described earlier (see Clifton 1960), his 

thesis was never published in a professional journal, and remained largely 

unknown until Percy brought it to the attention of Jerry Mires in 2001. 

124.  Forsman, E.D.; Swingle, J.K. 2005a. Interview with Kenneth and 

Charlotte Walker. 2 p. Unpublished notes. On file with: U.S. Department 

of Agriculture, Forest Service, Pacific Northwest Research Station, 3200 

SW Jefferson Way, Corvallis, OR 97331. (Available upon request from the 

compilers).

The authors interviewed retired biology professor Kenneth (Ken) Walker 

and his wife Charlotte at their home in Monmouth, Oregon, on 4 April 

2005. Ken was the son of Alex Walker, who was one of the earliest naturalists to collect tree voles in Oregon. During 1916–1967, Alex, his wife 

Rosaline, his brother Peter, and other family members collected 30 tree 

voles on the north coast of Oregon, primarily around Tillamook. Ken said 

that to collect tree voles, his father would climb to the nest and his mother 

would stand below with a shotgun to shoot tree voles if they could not be 

captured by hand. As a kid, Ken said he also climbed trees to capture tree 

voles. Ken said that he remembered climbing for tree voles at Oceanside 

and Short Beach on Cape Meares and at Cape Lookout, west of Tillamook. 

Alex also collected tree voles with his brother Peter near Cape Lookout 

(see Walker 1930). Ken said that Peter’s son Robert collected six red tree 

voles while logging around Tillamook in 1960. 

125.  Forsman, E.D.; Swingle, J.K. 2005b. Retrospective surveys of red tree 

voles in northwest Oregon: going, going, gone? [Abstract] Northwestern 

Naturalist. 86: 95. 
44

GENERAL TECHNICAL REPORT PNW-GTR-909

The authors found that tree voles were rare in the northern Coast Ranges 

of Oregon and suggested that patches of old forest may serve as refugia for 

tree voles in areas where most forests have been logged and converted to 

young forest.

126.  Forsman, E.D.; Swingle, J.K. 2006a. Interview with William (Bill) J. 

Hamilton, III. 2 p. Unpublished notes. On file with: USDA Forest Service, 

Pacific Northwest Research Station, 3200 SW Jefferson Way, Corvallis, 

OR 97331. (Available upon request from the compilers).

In this 2 March 2006 interview at his home near Winters, California, Bill 

Hamilton recounted the years when he and Don Roberts captured or raised 

in captivity over 258 Sonoma tree voles and 1 red tree vole. Most of the voles 

they captured were collected from sites near Jenner and the lower Russian 

River, in Sonoma County, California. Most of their specimens were eventually donated to the Slater Museum of Natural History at the University of 

Puget Sound.

127.   Forsman, E.D.; Swingle, J.K. 2006b. White-footed voles living in arboreal nests. Northwest Science. 80: 308–310.

The authors described the first confirmed arboreal nests of white-footed 

voles. They suggested that use of arboreal nests by white-footed voles may 

be a seasonal response to the presence of the leaves of red alder (Alnus 

rubra) during spring and summer.

128.  Forsman, E.D.; Swingle, J.K. 2007. Use of arboreal nests of tree voles 

(Arborimus spp.) by amphibians. Herpetological Conservation and 

Biology. 2: 113–118.

The authors described four species of amphibians that occurred in tree vole 

nests, including clouded salamanders, arboreal salamanders, Pacific tree 

frogs (Pseudacris regilla), and a California giant salamander (Dicamptodon ensatus). They suggested that arboreal activity by amphibians may be 

a more common phenomenon than is generally known. They hypothesized 

that tree vole nests may attract arboreal amphibians because they are warm 

and damp inside, and often have high concentrations of invertebrates that 

are potential food for salamanders.

129.   Forsman, E.D.; Swingle, J.K. 2010. Doug Bake and red tree voles: a 

tale of discovery in the Pacific Northwest. Northwestern Naturalist. 91: 

102–105.
45

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

In this historical account, Forsman and Swingle described the life of Doug 

Bake, a logger who collected 98 tree voles in the region around Tillamook, 

Oregon. Most of the specimens that Doug collected were given to Alex 

Walker or Murray Johnson and ended up in the Slater Museum of Nautral 

History at the University of Puget Sound. 

130.  Forsman, E.D.; Swingle, J.K.; Biswell, B.L.; Davis, R.J.; Andrews, 

L.S. [In press]. Distribution, habitat, diet, and management of the red tree 

vole and Sonoma tree vole. Manuscript in preparation. On file with: U.S. 

Department of Agriculture, Forest Service, Pacific Northwest Research 

Station, 3200 SW Jefferson Way, Corvallis, OR 97331.

The authors described past and current distribution of red tree voles and 

Sonoma tree voles based on data from owl pellets, specimen collections, 

and field surveys. They also modeled red tree vole habitat using presenceonly data and described regional variation in diet of tree voles. In a 

concluding chapter, they used historical and recent forest habitat maps to 

describe long-term changes in tree vole habitat. 

131.   Forsman, E.D.; Swingle, J.K.; Hatch, N.R. 2009. Behavior of red tree 

voles (Arborimus longicaudus) based on continuous video monitoring of 

nests. Northwest Science. 83: 262–272.

The authors used video cameras to record activity patterns and behavior at 

nests of three female red tree voles near Corvallis, Benton County, Oregon. 

All three voles were primarily nocturnal but occasionally foraged for brief 

periods during the day when they had large young in the nest. The median 

amount of time spent outside the nest per night was 27 minutes (range = 0 

to 712 minutes), and most of this time was spent foraging. Nocturnal activity consisted primarily of long periods inside the nest interspersed with 

short periods of intensive foraging, during which voles ran out of the nest, 

harvested cuttings and hauled them back to the nest. Cuttings were stored 

inside the nest or on top of the nest. The mean ± SE number of foraging 

bouts per night was 3.4 ± 0.1 (range = 1 to 13), and the mean number of 

cuttings harvested per night was 20.4 ± 3.5 (range = 0 to 75). Mean length 

of cuttings was 13.7 ± 3.5 cm (range = 1 to 37). Females harvested more 

cuttings when they had large young in the nest, and there was no evidence 

that they ate anything but the needles and bark of the cuttings that they 

brought to their nests. Juveniles first began to explore outside the nest 

when they were 30 to 35 days old and dispersed when they were 47 to 54 

days old. During the last 2 weeks, before they dispersed, juveniles spent 
46

GENERAL TECHNICAL REPORT PNW-GTR-909

considerable time outside the nest at night, becoming increasingly adept 

climbers and gradually beginning to harvest their own food. Dispersal of 

siblings occurred on the same night, and in at least some cases, appeared to 

be precipitated by female aggression towards the young.

132.  Forsman, E.D.; Swingle, J.K.; McDonald, M.A.; Graham, S.A.; 

Hatch, N.R. 2009. Red tree voles in the Columbia River Gorge and Hood 

River basin, Oregon. Northwestern Naturalist. 90: 227–232.

In 2003–2008, the authors conducted surveys to document the eastern and 

northern range limits of the range of red tree voles in the Columbia River 

Gorge and Hood River basin, Oregon. They concluded that the northern 

limits of the range of the vole included the forested bluffs and canyons 

bordering the Columbia River Gorge from a few kilometers east of Troutdale, east to within a few kilometers of Hood River. They also found tree 

vole nests in the headwaters of Lake Branch of Hood River on the east 

slope of the Cascades. This is the only region where tree voles occur on the 

east slope of the Cascades. Of 70 tree vole nests located, 74 percent were 

in forests dominated by mature or old-growth trees, and 26 percent were in 

forests that were 25 to 60 years old.

133.  Forsman, E.D.; Swingle, J.K.; Mowdy, J.S.; Price, A.L. 2012. 

Distance sampling surveys of red tree voles (Arborimus longicaudus) on 

Tillamook and Clatsop State Forests, Oregon, 2011–2012: progress report 

2 May 2012. 17 p. Unpublished report. On file with: U.S. Department of 

Agriculture, Forest Service, Pacific Northwest Research Station, 3200 

SW Jefferson Way, Corvallis, OR 97331. (Available upon request from the 

compilers). 

During winter of 2011–2012, the authors searched for tree vole nests while 

walking linear transects in 33 randomly selected plots on the Clatsop and 

Tillamook State Forests in the northern Coast Ranges of Oregon. Each plot 

was sampled with 500 m of transect. Potential nests detected while walking transects were climbed to determine if they were tree vole nests. Tree 

vole nests were detected in only one of 33 plots sampled. That plot was in 

125-year-old forest on the western edge of the Tillamook State Forest. 

134.  Forsman, E.D.; Swingle, J.K.; Price, W. 2008. Red tree vole (Arborimus 

longicaudus) surveys at Oswald West, Fort Stevens, and Ecola State 

Parks, 26–28 April 2008. 8 p. Unpublished report. On file with: U.S. 

Department of Agriculture, Forest Service, Pacific Northwest Research 

Station, 3200 SW Jefferson Way, Corvallis, OR 97331. (Available upon 

request from the compilers).
47

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

The authors climbed trees and searched for tree vole nests in old forests in 

state parks in the northern Coast Ranges of Oregon. They found recently 

occupied tree vole nests at Oswald West and Ecola State Parks. These were 

the first confirmed nests at these locations. All nests were in Sitka spruce 

or western hemlock trees.

135.  Foster, J.B. 1961. Life history of the Phenacomys vole. Journal of 

Mammalogy. 42: 181–198. 

This was primarily a paper on the eastern heather vole (Phenacomys 

ungava), but Foster concluded that red tree voles, dusky tree voles, and 

white-footed voles were clearly distinct when compared to heather voles.

136.  Franklin, J.F.; Cromack, K., Jr.; Denison, W.; McKee, A.; Maser, C.; 

Sedell, J.; Swanson, F.; Juday, G. 1981. Ecological characteristics of oldgrowth Douglas-fir forests. Gen. Tech. Rep. PNW-118. Portland, OR: U.S. 

Department of Agriculture, Forest Service, Pacific Northwest Forest and 

Range Experiment Station. 48 p.

The authors pointed out that large, old-growth Douglas-fir trees were 

individualistic and commonly had deep crowns with irregularly arranged, 

large, coarse branch systems. They suggested that this structure may 

provide ideal habitat for specialized vertebrates, such as the red tree vole, 

northern spotted owl, and northern flying squirrel, as well as nitrogenfixing lichens. They cited Meslow et al. (1981) in support of the hypothesis 

that red tree voles find optimum habitat for foraging and nesting in oldgrowth forests of Douglas-fir and western hemlock. They concluded that 

old-growth trees provide nest sites, food, and cover for tree voles, and they 

speculated that many generations of tree voles can live (sequentially) in 

the same tree. They emphasized that the large branch systems and organic 

accumulations on limbs of old-growth trees provide habitat for many 

species.

137.   Franklin, J.F.; Spies, T.A. 1986. The ecology of old-growth Douglas-fir 

forests. Oregon Birds. 12: 79–90.

This paper provided no new data on tree voles, but the authors did report 

that many species occupy old-growth forests, including the red tree vole. 

They emphasized that managers should not overlook the important roles 

that large branch systems and organic accumulations play in providing 

habitat for species like tree voles. 

138.  Gannon, W.T. 1981–1984. Unpublished field notes. On file with: 

Humboldt State University, Vertebrate Museum, Arcata, CA.
48

GENERAL TECHNICAL REPORT PNW-GTR-909

These field notes included multiple references to Sonoma tree voles 

collected in 1981−1984 by William Gannon, Mike Webb, and others from 

areas east of Arcata, Humboldt County, California. We found most of these 

specimens in museum collections. 

139.   George, S.B. 1999. Sonoma tree vole Arborimus pomo. In: Wilson, 

D.E.; Ruff, S., eds. The Smithsonian book of North American Mammals. 

Washington, DC: Smithsonian Institution Press: 620–622.

This species account included information on the distribution, habitat, 

food, nest characteristics, reproduction, chromosome number, and fossil record of the Sonoma tree vole. Like many before her, George gets it 

wrong when she says that tree voles eat the edge of Douglas-fir needles 

and discard the central vascular bundle.

140.  George, S.B.; Johnson, M.L. 1999. Morphometric reassessment of 

systematic relationships in the genus Arborimus [Abstract]. Fifty-ninth 

annual meeting of the American Society of Mammalogists: Corvallis, OR: 

Oregon State University. (Available upon request from the compilers).

The authors used a discriminant function analysis to compare morphometric characteristics of tree voles in three different regions. They concluded 

that populations of tree voles in California, which were then referred to as 

A. longicaudus, actually represented a distinct, as yet unnamed species. 

They concluded that populations of Arborimus in northern Oregon were 

subspecifically distinct from populations of A. longicaudus in the rest of 

Oregon. They proposed the subspecific name A. l. silvicola for tree voles in 

northern Oregon and A. l. longicaudus for tree voles in the rest of Oregon. 

141.   Gilbert, F.F.; Allwine, R. 1991. Small mammal communities in the 

Oregon Cascade Range. In: Ruggiero, L.F.; Aubry, K.B.; Carey, A.B.; 

Huff, M.H., tech. coords. Wildlife and vegetation of unmanaged Douglasfir forests. Portland, OR: U.S. Department of Agriculture, Forest Service, 

Pacific Northwest Research Station: 257–267.

The authors used pitfall traps and snap-traps to sample small mammals in 

three different regions in the western Cascades in Oregon. They caught no 

red tree voles in snap-traps. They did catch tree voles in their pitfall traps, 

but their results were a bit confusing because the numbers in their tables 

did not always agree. In their summary of pitfall results in table 3, they 

say they caught two tree voles in mature forests and six tree voles in old 

forests, for a total of eight voles. In the same table, they say they caught six 

tree voles in mesic sites and three tree voles in wet sites for a total of nine 

tree voles. Why the two sets of numbers don’t add up to the same total is 
49

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

unclear because the authors stated that all of the tree voles they captured 

were in mature and old forests. Regardless of this minor inconsistency, 

the upshot was that they caught voles in mature and old forests, but not in 

young forest, which was the same pattern observed in other pitfall studies 

conducted by Corn and Bury (1986). The authors qualified their results 

by saying that they caught so few tree voles they could not do meaningful 

statistical comparisons among forest age classes or moisture classes. 

142.   Gillesberg, A-M.; Carey, A.B. 1986. Red tree voles (Arborimus longicaudus) in the Oregon Coast Range: pilot study plan. 43 p. Unpublished 

report. U.S. Department of Agriculture, Forest Service, Pacific Northwest 

Research Station, 3625 93rd Ave. SW, Olympia, WA 98512. 43 p. 

(Available upon request from the compilers).

This study plan described a sampling method in which the authors proposed 

to search for red tree vole nests in trees that were felled by a Bureau of Land 

Management sampling crew. They suggested that this method was the only 

expedient and cost-efficient method available to study tree vole nests because 

they felt that tree climbing was unsafe and impractical on a large scale. 

143.   Gillesberg, A-M.; Carey, A.B. 1987. Red tree voles (Arborimus longicaudus) in the Oregon Coast Range. Progress report 1987. 13 p. Unpublished 

report. U.S. Department of Agriculture, Forest Service, Pacific Northwest 

Research Station, 3625 93rd Ave. SW, Olympia, WA 98512. (Available upon 

request from the compilers).

In this progress report, the authors reported that they changed their search 

methods from their original study plan after they found that they could 

only survey two felled trees per day using their original methods. Instead 

of doing an intense search of each tree, they started doing a less intense 

examination in which they walked along each felled tree “...pausing at 

intervals to look carefully amongst the foliage on either side of the trunk 

for red tree vole nests/sign.” They sampled a total of 93 trees, including 

50 trees with the original intensive sampling method and 43 trees with 

the less intense method. Their searches were conducted 3 to 4 weeks after 

the trees had been felled. The proportion of trees with vole nests was 62 

percent in the intensive sampling group and 44 percent in the less-intensive 

sampling group. Over 80 percent of the nest material found was collected 

during the intensive searches. The authors did not mention whether they 

could determine if nest fragments were from occupied or old vole nests, 

and they never saw any tree voles. They spent 39 days sampling 93 trees 

and collecting vegetation data. They did not discuss the potential impact of 

destructive sampling on vole nesting habitat. 
50

GENERAL TECHNICAL REPORT PNW-GTR-909

144.  Gillesberg, A-M.; Carey, A.B. 1991a. Red tree voles in the Oregon Coast 

Ranges [Abstract]. In: Ruggiero, L.F.; Aubry, K.B.; Carey, A.B.; Huff, 

M.H., tech. coords. Wildlife and vegetation of unmanaged Douglas-fir forests. Gen. Tech. Rep. PNW-GTR-285. Portland, OR: U.S. Department of 

Agriculture, Forest Service, Pacific Northwest Research Station: 528.

Gillesberg and Carey argued that counting red tree vole nest fragments in 

felled trees was a more efficient way to document the presence of tree voles 

than other methods such as trapping or climbing trees. They examined 93 

felled trees and found 117 pieces of tree vole nest material. They found 

that smaller nests usually consisted solely of resin ducts and fecal material, 

whereas larger nests contained resin ducts, conifer needles, twigs, lichens, 

feces, and other organic matter. Nests were found in a variety of locations, 

including hollow limbs, broken tops, and resting on branches. Nests were 

most often found in the lower one-third of the canopy, and larger trees (>100 

cm diameter at breast height) were preferred over smaller ones. 

145.  Gillesberg, A-M.; Carey, A.B. 1991b. Arboreal nests of Phenacomys 

longicaudus in Oregon. Journal of Mammalogy. 72: 784–787.

Gillesberg and Carey concluded that searching felled trees was an effective 

method for finding nests of red tree voles. They examined 82 felled trees 

for evidence of tree vole nests in a 35.2-ha plot near Yoncalla, Douglas 

County, Oregon. All trees were in old-growth stands. A total of 117 nests 

or nest fragments were found in 50 of the 82 trees examined. Thirty-five 

trees contained 1 to 2 nests, and only 2 trees had more than 5 nests (9 and 

11 nests, respectively). The actual number of nests represented by these 

fragments was unknown. Of the 117 nests or nest fragments located, 59

(50 percent) were in the lower third of the live crown, 34 (29 percent) were 

in the middle one-third, and 24 (21 percent) were in the upper one-third. Of 

27 nests located in situ, 15 were out on branches and 12 were against the 

trunk. Two of four cavities examined had nests in them, one in a broken 

top and one in a hollow branch. Trees with nests tended to be taller and 

larger in diameter. 

The authors did not examine felled trees in young stands but did review 

other literature in which researchers sampled multiple age classes. They 

concluded (pages 786–787) that “...live old-growth trees provide optimum 

habitat for the red tree vole because primary production is high and leaves 

are concentrated in old-growth, resulting in maximum food availability.” 
51

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

They also stated (page 787) that the “...moist old-growth canopy functions 

as a climatic buffer and its high water-holding capacity maintains fresh 

foliage and provides free water to be licked from needles.” 

The authors also said that they tried many other techniques for sampling 

tree voles, including visual searches from ground transects, Sherman livetraps, and pitfall traps, but did not catch any voles and only found nests in 

4 of 77 stands in which they visually searched for nests on ground transects. However, they also said that “From searches in the stands in which 

we captured red tree voles, we did not locate any nests, despite the use of 

binoculars and hillside vantage points.” This is confusing because they 

also said that they never captured or observed any tree voles. We suspect 

they were referring to stands in which Corn and Bury (1991) captured tree 

voles in pitfall traps. 

146.  Gitzen, R.A.; West, S.D.; Maguire, C.C.; Manning, T.; Halpern, C.B. 

2007. Response of terrestrial small mammals to varying amounts and patterns of green-tree retention in Pacific Northwest forests. Forest Ecology 

and Management. 251: 142–155.

The authors analyzed effects of variable-retention harvest treatments on 

small mammals in the western Cascades, Lane County, Oregon. Their 

sample of 6,208 small mammals captured in pitfall traps included one red 

tree vole and one white-footed vole in 256,896 trap-nights. The authors 

said that the number of captures was too small to evaluate treatment effects 

on tree voles and white-footed voles, but they also speculated that tree 

voles require retention of large trees. 

147.   Golightly, R.T.; Penland, T.F.; Zielinski, W.J.; Higley, J.M. 2006. 

Fisher diet in the Klamath/north coast bioregion. Arcata, CA: Humboldt 

State University; final report to U.S. Fish and Wildlife Service; contract 

113334J025 and Scotia Pacific contract 8355. 56 p. http://www.fws.gov/

yreka/Fisher/Literature/Golightly%20et%20al%202006%20Fisher%20

Diet.pdf. (August 2014).

The authors found Arborimus remains in fisher scats in two of four study 

areas in northern California. In their Six Rivers Study Area in Humboldt 

and Trinity Counties, 14 of 152 scats (9 percent) contained remains of 

Arborimus. In their Hoopa Study Area in Humboldt County, 1 of 64 scats 

(2 percent) contained Arborimus remains. The authors did not differentiate 

between remains of tree voles and white-footed voles.
52

GENERAL TECHNICAL REPORT PNW-GTR-909

148.  Gomez, D.M. 1992. Small mammal and herpetofauna abundance in riparian and upslope areas of five forest conditions. Corvallis, OR: Oregon 

State University. 118 p. M.S. thesis. 

Gomez used pitfall traps to sample 15 different plots in five watersheds in 

the Coast Ranges of western Oregon in 1989–1990. He sampled three plots 

in each of five forest types (three shrub communities, three pole stands, 

three mature stands, three old-growth stands, and three deciduous stands). 

Each plot was sampled with an array of 60 pitfall traps. He captured 9,723 

mammals and 917 amphibians/reptiles in 100,800 trap-nights. He caught 

20 species of mammals, including 23 red tree voles and 59 white-footed 

voles (0.023 tree voles and 0.059 white-footed voles per 100 trap-nights). 

He provided legal coordinates of his pitfall grids but did not say how many 

tree voles he captured at each grid. Of the 23 tree voles captured, 16 were 

in old-growth forests, 4 were in “large-saw timber,” 1 was in pole-size 

forest, 2 were in shrub vegetation, and none were captured in deciduous 

forest. Gomez concluded that “...tree voles were captured more frequently 

(P < 0.01) in old-growth than in each of the other forest types.” The number of tree voles captured in riparian transects versus upslope transects did 

not differ. More white-footed voles were captured in riparian (82 percent) 

than upslope (18 percent) habitat. White-footed voles were captured most 

frequently in areas of deciduous trees and shrubs.

149.   Gomez, D.M.; Anthony, R.G. 1998. Small mammal abundance in riparian and upland areas of five seral stages in western Oregon. Northwest 

Science. 72: 293–302. 

In this published version of Doug Gomez’s thesis, the authors reported that 

they used pitfall traps to sample 15 plots in five watersheds in the Oregon 

Coast Ranges. They captured 9,764 small mammals and 917 amphibians/

reptiles in 100,800 trap-nights. They caught 20 species of mammals, 

including 23 red tree voles and 59 white-footed voles. Capture rates for 

tree voles and white-footed voles were 0.023 and 0.059 per 100 trapnights, respectively. They captured more tree voles in old-growth forest 

than in any other seral stages and concluded that red tree voles were most 

abundant in old forest. They reported capturing about equal numbers of

tree voles in upland and riparian habitat. They suggested that white-footed 

voles were dependent on deciduous vegetation and not restricted to riparian 

areas as some have stated. This conclusion was in agreement with Voth et 

al. (1983) who also suggested that white-footed voles were closely associated with deciduous habitat.
53

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

150.  Gomez, D.M.; Anthony, R.G.; Lowe, R.W. 1997. Small mammal and 

amphibian abundance at Cape Meares National Wildlife Refuge. 34 p. 

Unpublished report. On file with: Oregon Cooperative Wildlife Research 

Unit, Oregon State University, Corvallis, OR 97331. (Available upon 

request from the compilers).

This study was conducted in 1996 at Cape Meares National Wildlife Refuge, Tillamook County, Oregon. In 7,290 trap-nights using a combination 

of pitfall traps and Sherman live-traps, the authors captured four whitefooted voles in young or wind-damaged forests and three red tree voles 

in old-growth forests. The capture rate per 100 trap-nights was 0.04 for 

tree voles and 0.06 for white-footed voles. The specimens were apparently 

discarded, as we could not find them in museums.

151.   Gordon, K.L. 1966. Mammals and the influence of the Columbia River 

Gorge on their distribution. Northwest Science. 40: 142–146.

Gordon mentioned that the Columbia River was the northern limit of the 

ranges of the red tree vole and white-footed vole. 

152.  Gould, G.I. 1987. Nongame wildlife investigations: forest mammal survey and inventory. 25 p. Unpublished report. Project Number 

CA W-065-R04/JOB IV-11. California Fish and Game, 1416 9th Street, 

Sacramento, CA 95814. (Available upon request from the compilers).

This was a short progress report on Nina Meiselman’s thesis on Sonoma 

tree vole nest surveys (see Meiselman 1992, Meiselman and Doyle 1996). 

153.  Graham, S.A. 2012 Diet composition, niche and geographic characteristics, and prey size preference of barred owls (Strix varia) in the Pacific 

Northwest. Boise, ID: Boise State University. 147 p. M.S. thesis. 

Of 3,686 prey items identified in barred owl (Strix varia) pellets collected 

west of Eugene, Lane County, Oregon, 2,387 were mammals, including 

147 (4.0 percent) red tree voles.

154.  Graham, S.A.; Mires, G.W. 2005. Predation on red tree voles by owls 

and diurnal raptors. Northwestern Naturalist. 86: 38–40.

The authors described several cases in which red tree voles were captured 

by owls or hawks in Douglas County, Oregon. In one case, Graham 

observed a female northern spotted owl as she captured a tree vole that 

was climbing down a tree trunk. Mires observed a northern pygmy-owl 

(Glaucidium gnoma) flying with a tree vole in its talons and also watched 

a red-tailed hawk (Buteo jamaicensis) as it tore into a tree vole nest and 

captured a tree vole. 
54

GENERAL TECHNICAL REPORT PNW-GTR-909

155.  Green Diamond Resource Company. 2006. Exhibit A: Green 

Diamond Resource Company northern spotted owl conservation 

plan, phase one comprehensive review. http://www.fws.gov/arcata/

es/reports/1992GrnDiamond_NSOHCP/DraftEAS/Exhibit%20A_

Comprehensive%20%20Review.pdf. (August 2014).

Although there were no authors listed on this report, the sections on tree 

voles were written by Lowell Diller, Keith Hamm, Dave Lamphear, and 

Trent McDonald at the Korbel office of Green Diamond Resource Company 

(Lowell Diller, wildlife biologist, Green Diamond Resource Company, 1165 

Maple Creek Road, Korbel, CA 95550). They identified 3,056 prey items in 

pellets collected from northern spotted owls in northwestern California in 

1989–2004. The sample included 497 tree voles (16.3 percent). 

The authors also used line transects to estimate the distribution and 

abundance of nests of Sonoma tree voles on Green Diamond lands from 

2001–2004. They sampled 10-ha plots in four different forest age classes 

(20 to 29 years, 30 to 39 years, 40 to 50 years, and >50 years). They did 

not climb to verify nests but used time constraint sampling by searching 

for resin ducts on the ground below all nests detected. Tree vole nests were 

detected in 32 of 68 plots sampled (47 percent). The authors located a total 

of 129 tree vole nests (range = 0 to 19 per plot). Abundance of nests was 

positively correlated with forest age. 

156.  Greenway, G. 2005. Location of the historical Lierly Ranch in California. 

Email to Eric Forsman. (13 September). (Available upon request from the 

compilers).

In this email, Greg Greenway, an archeologist on the Mendocino National 

Forest, described the location of the historical Lierly Ranch on a 1917 map 

as T19 N R12W section 31 NE1/4 of the NW1/4. This is the location near 

Mount Sanhedrin where two Sonoma tree voles were collected in 1899 and 

1913, respectively. However, Greenway conceded that the exact location 

could be problematic because the Lierly Ranch was also labeled as the 

Ascherman Ranch in section 30. Harry Lierly lived at the ranch until he 

died in 1924. 

157.   Grinnell, J. 1913. A distributional list of the mammals of California. 

Proceedings of the California Academy of Sciences (Series 4). 3: 316. 

At the time he wrote this report, Grinnell noted that there was only one 

“long-tailed lemming mouse” record for California. This was a tree vole 

found dead in the road near Mount Sanhedrin in 1899, as described by 

Witmer Stone (1904).
55

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

158.  Grinnell, J. 1933. Review of the recent mammal fauna of California. 

University of California Publications in Zoology. 40: 71–234.

Grinnell described the range and habitat of the tree vole and the “Pacific 

coast lemming mouse” (white-footed vole) in California.

159.   Grosvenor, M.B. 1966. Wild animals of North America. Washington, 

DC: National Geographic Book Service. 400 p.

In this romanticized and misleading account of tree vole biology, Grosvenor said that male tree voles were primarily terrestrial and were usually 

“...content to rummage under the debris on the forest floor.” He also said 

that juvenile tree voles “...take about a month to mature.” None of this is 

supported by the literature, and most of it is simply wrong. 

160.  Guilday, J.E.; Parmalee, P.W. 1972. Quaternary periglacial records 

of voles of the genus Phenacomys Merriam (Cricetidae: Rodentia). 

Quaternary Research. 2: 170–175.

The authors concluded that Phenacomys could not be used as an index 

fossil but was a good indicator of climate because of its association with 

the modern Hudsonian Life Zones of North America. They stated that 

Pliophenacomys Hibbard was the ancestral genus of modern Phenacomys

and was recovered from late Pliocene deposits of the Great Plains. They 

emphasized that fossil Phenacomys voles were excellent indicators of 

boreal conditions because their molars were easy to identify. They stated 

that inter- and intraspecific variation in dentition was too great to allow 

identification of Phenacomys fossils to species. Thus all fossil forms had 

been identified as Phenacomys intermedius. They reported that remains 

of Phenacomys had been located at ≥20 Pleistocene to early recent sites in 

the United States, from the east coast to south central Idaho. Some of their 

taxonomy was a bit out of date. For example, they considered the dusky 

tree vole as a unique species.

161.   Hafner, D.J. 1998. California red tree vole (Arborimus longicaudus), 

white-footed vole (A. albipes). In: Hafner, D.J.; Yensen, E.; Kirkland, 

G.L., Jr., eds. Status survey and conservation action plan: North American 

rodents. Gland, Switzerland: The World Conservation Union: 86–87.

The authors concluded that the World Conservation Union Red List 

Category was “data deficient” for the red tree vole and white-footed vole. 

They recommended conducting studies on the ecology of these species to 

determine if they were truly rare and threatened.
56

GENERAL TECHNICAL REPORT PNW-GTR-909

162.  Hagar, J.C. 2007. Wildlife species associated with non-coniferous vegetation in Pacific Northwest conifer forests: a review. Forest Ecology and 

Management. 246: 108–122.

In table 1 on page 115, Hagar listed the white-footed vole as a species 

that forages on shrubs, understory vegetation, California hazel (Corylus 

cornuta), and red alder.

163.  Hagele, T.; Kaufman, B.; Whitaker, J.O., Jr.; Klompen, H. 2005. The 

genus Euryparasitus in North America (Mesostigmata: Euryparasitidae). 

Zootaxa. 1036: 1–20.

In this paper on Euryparasitus mites of North America, the authors 

reported that mites were collected from eight white-footed voles in 

Lincoln, Benton, Curry, and Linn Counties, Oregon. They included the 

geographic coordinates for each collection.

164.  Hall, E.R. 1981. The mammals of North America. 2nd ed. New York: 

John Wiley and Sons. 1181 p. Vol. 2.

In this revised edition of the Mammals of North America, Hall concluded 

that the subspecific status of the dusky tree vole was provisional owing to 

uncertainty about its relationship with the red tree vole.

165.  Hall, E.R.; Cockrum, E.L. 1953. A synopsis of the North American 

microtine rodents. University of Kansas Publications, Museum of Natural 

History. 5: 373–498.

In addition to a key to the Phenacomys, Hall and Cockrum provided range 

maps, specimen measurements, and skull drawings for the heather vole, 

red tree vole, dusky tree vole, and white-footed vole.

166.  Hall, E.R.; Kelson, K.R. 1959. The mammals of North America. New 

York: The Ronald Press Company. 1083 p. Vol. 2.

On pages 720–721, the authors provided external measurements, descriptions, and distributions for the white-footed vole, dusky tree vole, and red 

tree vole.

167.   Hamilton, W.J., Jr. 1939. American mammals: their lives, habitats, and 

economic relations. New York: McGraw-Hill Book Company. 434 p.

In this general species account, Hamilton described the red tree vole as a 

specialized feeder that lacks the ability to change food and will starve in 

the absence of their specialized foods (conifer needles). However, he also 

reported that Benson and Borell (1931) found that tree voles would sometimes eat other foods in captivity. He repeats much of the conventional 
57

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

wisdom regarding tree voles, including (1) when they feed on needles 

of Douglas-fir they discard the resin ducts before eating the medullary 

portion of the needle, (2) they seldom come to the ground except to reach 

some other tree, (3) many nests cannot be seen from the ground because 

they are hidden in dense foliage, (4) females build bigger nests than males, 

and (5) tree voles occasionally build their nests on top of abandoned squirrel nests.

168.  Hamilton, W.J., Jr. 1946. A study of the baculum in some North 

American Microtinae. Journal of Mammalogy. 27: 378–387.

Based on a single specimen, Hamilton concluded that the shape of the 

baculum of the tree vole was distinctive compared to all other genera of 

Arvicolinae. 

169.   Hamilton, W.J., III. 1962. Reproductive adaptations of the red tree 

mouse. Journal of Mammalogy. 43: 486–504.

Hamilton described the estrous cycle, ovulation, mating behavior, gestation 

period, postpartum estrus, growth rates, and feeding behavior of adult and 

juvenile tree voles. He and Don Roberts used over 50 adult animals in their 

laboratory experiments. Most of his findings were identical to the observations reported 2 years earlier by Clifton (1960). He reported that gestation 

of nonlactating females was 27.0 to 28.5 days. He presented data from a 

captive female with two young that bred when her young were 2 days old 

and produced a second litter 41.8 days later. He also suggested that adult 

females that produced litters 40 days after having previous litters was 

evidence of delayed implantation. He cited many papers indicating that 

postpartum estrus was common in voles. However, his tests of postpartum 

estrus were only successful in one of three cases in which males were 

introduced into cages with females that had just had litters. In two cases, 

the females were unreceptive. In the third case, the female mated with the 

male 27 to 28 hours after parturition but was not successful in weaning 

the young from the first litter. Hamilton said that the young were normally 

weaned at 30 to 40 days of age when they weighed approximately 17 g. 

He found that eyes opened on about the 21st day, at which point there was 

a spurt in daily weight gain as the young began to eat solid food. He said 

that Douglas-fir cells were found in fecal pellets of young in some nests 

as early as 17 days of age, 3 to 4 days before their eyes opened. He suggested this was due to ingestion of the mother’s fecal pellets (coprophagy), 

which was first witnessed on the 17th day. He said that, “When held in 
58

GENERAL TECHNICAL REPORT PNW-GTR-909

the hand on this date the young avidly consumed all the feces they came 

upon.” This included voles eating their own fecal pellets. He suggested 

that coprophagy was necessary to inoculate the gut with micro-organisms 

needed for digestion of Douglas-fir needles. Juveniles were first seen 

eating needles when they were 25 days old, and by 27 days were removing 

resin ducts and eating needles like adults. He described a juvenile that 

escaped from its cage at 32 days of age and was found 3 days later in a nest 

that it had built in a pile of Douglas-fir limbs that were kept in the lab. It 

had continued to gain weight, indicating that it was able to survive on its 

own at this age. There is an excellent discussion of induced ovulation and 

adaptations to a low-energy diet, including a longer period of intrauterine 

development, small litter size, slow juvenile growth rates, and long period 

of female care. He suggested that these reproductive traits are primarily 

an adaptive response to a low-energy diet of conifer needles as opposed to 

adaptations for living in an arboreal environment. He suggested that tree 

voles were not able to live at high elevations because their arboreal nests 

were too small and exposed to keep them warm during winter.

170.   Hamilton, W.J., III; Roberts, D.R. 1957–1961. Unpublished lab and 

field notes. On file with: University of Puget Sound, Slater Museum of 

Natural History, Tacoma, WA.

We obtained these original notes from Don Roberts when we interviewed 

him at his home in Santa Cruz, California, in 2006. After transcribing the 

notes, we donated the originals to the Slater Museum of Natural History at 

the University of Puget Sound. These notes described the capture locations 

and laboratory histories of 248 tree voles that Roberts and Bill Hamilton 

used in their captive-breeding study of Sonoma tree voles from 1957 

through 1961. Most of the specimens that they captured or bred in captivity 

are at the Slater Museum. Most of these specimens were captured near the 

Russian River in Sonoma County, California, but a few were collected in 

Mendocino and Humboldt Counties, California.

171.   Hammer, W.; Maser, C. 1967. Unpublished field notes. On file with: 

University of Puget Sound, Slater Museum of Natural History, Tacoma, WA.

These notes included drawings that Wayne Hammer recorded when he 

captured a red tree vole near Vida, Lane County, Oregon, on 13 September 

1967. Hammer was a government trapper who helped Chris Maser and 

Murray Johnson collect many tree voles in western Oregon.
59

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

172.   Harris, L.D. 1984. The fragmented forest: island biogeography theory 

and the preservation of biotic diversity. Chicago, IL: The University of 

Chicago Press. 199 p.

Harris described and discussed tree vole biology in several sections of his 

book, citing Maser (1966), Johnson (1973), and Maser et al. (1981) as his 

primary sources of data. He described the loss of leader dominance in trees 

and resulting upturned branches that support nests of tree voles. He said 

that tree voles spend their entire life in the canopy and that many individuals may inhabit the same old-growth tree during the span of centuries. He 

said that tree voles are primarily restricted to mature Douglas-fir forests, 

although they do occur in other tree species and in younger successional 

stands. He describes the diet as restricted almost entirely to Douglas-fir 

needles. He repeated the oft-stated fact that the red tree vole is the most 

highly specialized vole in the world and the most arboreal mammal in 

North America. He also said that the white-footed vole is the smallest 

mammalian browser and perhaps the rarest vole in North America. He 

speculated that red tree vole home ranges average about 0.082 ha. 

173.   Haukisalmi, V.; Rausch, R.L.; Henttonen, H. 2005. Morphological 

characterisation of Paranoplocephala bairdi (Schad, 1954) (Cestoda: 

Anoplocephalidae) in heather voles Phenacomys spp., and tree voles 

Arborimus spp., and related species in voles and lemmings (Muridae: 

Arvicolinae). Folia Parasitologica. 52: 311–321.

The authors assessed the taxonomic status of Paranoplocephala bairdi, 

a cestode flatworm found in heather voles and tree voles. Their results 

supported the independent status and conspecificity of specimens from 

heather voles and tree voles. They suggested that P. bairdi was a hostspecialist parasite of tree voles and heather voles, with a wide geographical 

distribution in North America.

174.   Haverkost, T.R.; Gardner, S.L.; Peterson, A.T. 2010. Predicting the 

distribution of a parasite using the ecological niche model, GARP. Revista 

Mexicana de Biodiversidad. 81: 895–902.

The authors modeled the ecological niche of tapeworms of voles in North 

America, including Andrya neotomae found in white-footed voles and 

Anoplocephaloides cf. dentata found in red tree voles.

175.   Hayes, J.P. 1995. Unpublished field notes. On file with: U.S. Department 

of Agriculture, Forest Service, Pacific Northwest Research Station, 3200 

SW Jefferson Way, Corvallis, OR 97331 (Available upon request from the 

compilers).
60

GENERAL TECHNICAL REPORT PNW-GTR-909

These field notes described a field trip in 1995 during which John Hayes, 

Mike Murray, and others spent 2 days climbing trees to capture red tree 

voles northeast of Waldport, Lincoln County, Oregon. Of the six tree voles 

captured, two were melanistic (Hayes 1996). The six live specimens were 

sent to Milo Richmond at the Cornell Cooperative Wildlife Research Unit 

for captive breeding, where they all died. These were the first confirmed 

cases of melanism in the tree vole. Unfortunately, the specimens were lost 

or discarded. 

176.   Hayes, J.P. 1996. Arborimus longicaudus. Mammalian Species. 532: 1–5.

In this species account, Hayes summarized knowledge of red tree voles 

as of 1996, including a reference to melanistic specimens that he collected in 1995. He pointed out that skulls of voles in the genus Arborimus

could be distinguished from those of other arvicolines in the range of A. 

longicaudus by the shape of the deep reentrant angles of the lower molars. 

He described the rootless molariform teeth of juveniles that gradually 

develop roots as tree voles mature (Hooper and Hart 1962, Howell 1926). 

Rooted molars generally are considered to be an ancestral character for 

the subfamily. He reported that Johnson (1973) speculated that the genus 

Arborimus evolved in the Pacific Northwest during the Pliocene, but that 

this could not be confirmed because there are no known tree vole fossils. 

177.   Hayes, J.P. 2005. Email to Jim Swingle. (28 March). (Available upon 

request from the compilers).

John Hayes said that he sent six red tree voles (four normal, two melanistic) 

that he captured northeast of Waldport, Lincoln County, Oregon, to Milo 

(Mike) Richmond, the leader of the Cooperative Wildlife Research Unit at 

Cornell University. When contacted by Forsman, Richmond said he kept 

all six tree voles in the same cage and that they were all females because 

they never bred. He said that the voles lived about a year, but he did not 

remember what became of the specimens.

178.  Hayes, J.P.; Adam, M.D.; Anthony, R.G.; Witt, J.W. 1996. Comparison 

of the effectiveness of Sherman and modified Fitch live-traps for capture 

of small mammals. Northwestern Naturalist. 77: 40–43.

The authors reported that they caught an 18.9-g adult male white-footed 

vole in a Fitch live-trap in riparian habitat in the Oregon Coast Range. 

They did not specify where they captured the vole, but their four study 

sites were in Benton, Lincoln, and Douglas Counties.
61

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

179.   Hernandez, P.A.; Graham, C.H.; Master, L.L.; Albert, D.L. 2006. The 

effect of sample size and species characteristics on performance of different species distribution modeling methods. Ecography. 29: 773–785. 

One of the species they used in their distribution and habitat modeling 

exercise was the Sonoma tree vole.

180.  Hooper, E.T. 1936. Unpublished field notes. On file with: University of 

California, Berkeley, Museum of Vertebrate Zoology.

In his notes, Hooper says that he prepared five Sonoma tree voles that 

were captured by Seth Benson and his wife Emma, and David Johnson 

between Monte Rio and Camp Meeker in Sonoma County, California. The 

two adult females and three juvenile females all died after a few days in 

captivity.

181.   Hooper, E.T.; Hart, B.S. 1962. A synopsis of recent North American 

microtine rodents. Miscellaneous Publications, Museum of Zoology, 

University of Michigan. 20: 5–68.

The authors described the glans penis of arvicolines, including tree voles. 

They viewed tree voles as a divergent form of the heather vole.

182.  Hooven, E.F.; Black, H.C. 1976. Effects of some clearcutting practices 

on small-mammal populations in western Oregon. Northwest Science. 50: 

189–208.

The authors used Sherman live-traps to sample small mammals along 

the South Fork of the McKenzie River in eastern Lane County, Oregon, 

1968–1970. Of 3,630 small mammals captured, 3 were white-footed voles.

183.  Hopkins, D.S. 2013. Exhibit 6. Declaration of Scott Hopkins in the case 

of Cascadia Wildland, The Center for Biological Diversity, and Benton 

Forest Coalition v. Bureau of Land Management and Seneca Sawmill 

Co. U.S. District Court, District of Oregon, Eugene Division. Case No. 

6:12-cv-1739-AA. 7 p.

In 2013, the Bureau of Land Management (BLM) hired contractors to 

climb 206 trees in 10 remnant stands of old-growth and mature forest in 

the Marys Peak Resource Area of the Salem District of BLM. Trees were 

selected on the basis of visible nests or unique structure, such as broken 

tops, forks, or cavities. Climbers found evidence of tree vole presence in 

132 of 206 trees climbed (64 percent), including 77 trees (37 percent) with 

evidence of recent activity. On the basis of this nonrandom sample, Hopkins concluded that tree voles were common in, “...every suitable patch 

of habitat that was surveyed, and some survey units showed abundant 
62

GENERAL TECHNICAL REPORT PNW-GTR-909

evidence of vole presence regardless of habitat patch size.” He went on to 

infer that all of the late-successional forests in reserved allocations in the 

resource area (about 2171 ha) would have similar levels of vole presence. 

He concluded that tree voles were “...very common and likely abundant 

within all suitable habitat in the watershed.” And finally, he concluded that 

the proposed BLM timber sale would have “...a negligible effect on the 

vole population within the watershed...” There was no discussion regarding the validity of drawing general conclusions based on a nonrandom 

sample of trees selected specifically because they had visible nests or 

unique structure. There was also no discussion regarding the validity of 

population estimates that were based on nest counts as opposed to actual 

documentation of live animals. 

184.  Howell, A.B. 1920. A second record of Phenacomys albipes in California, 

with a discussion on the species. Journal of Mammalogy. 1: 242–243. 

A white-footed vole collected near Orick, Humboldt County, California, 

was compared to five other white-footed vole specimens. Howell concluded that “After studying the California species of Phenacomys, I can see 

nothing to be gained by adopting the subgenus Arborimus Taylor. Most of 

the characters on which it is based are either very slight, or inconsistent. 

It seems to me that the genus is an unusually well-defined one, and that 

nothing whatever can be gained by further division. However, if a subgenus must be recognized, let it contain only longicaudus, and be based upon 

the arboreal habits and hairy tail of that species, rather than on ill-defined 

characters that can be segregated only with considerable difficulty.”

185.  Howell, A.B. 1921. Description of a new species of Phenacomys from 

Oregon. Journal of Mammalogy. 2: 98–100.

Based on a single specimen collected near Tillamook, Oregon, Howell 

proposed a new species of tree vole (Forest lemming-mouse; Phenacomys 

silvicolus) that was darker and had smaller ears than the red tree vole. 

He concluded that the skull of this new species was longer with a narrower braincase that had more pronounced temporal ridges, and that the 

molariform teeth were structurally different (much heavier with a different 

occlusal pattern). Miller (1925) subsequently shortened P. silvicolus to P. 

silvicola.
63

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

186.  Howell, A.B. 1926. Voles of the genus Phenacomys. I. Revision of the 

genus Phenacomys. II. Life history of the red tree mouse (Phenacomys 

longicaudus). North American Fauna. 48: 1–66.

Howell concluded that Phenacomys longicaudus and P. silvicola were 

definitely but not exclusively arboreal and probably had fewer predatory 

enemies to deal with when compared to terrestrial voles. He called these 

two species the “red tree mouse” and “forest tree mouse,” respectively. He 

suggested that the distribution of tree voles was discontinuous throughout 

the humid coastal belt of Oregon and California. He incorrectly speculated 

that tree voles could occur in Washington. He reported that tree voles 

descended to the ground to reach other trees and that the few ground 

nests that had been discovered had all been occupied by males. He stated 

that tree voles fed on the fleshy part of needles and on tender twig bark 

from green twigs. He said that tree voles were seldom found in isolated 

patches of forest and speculated that open expanses of nonforest may 

be barriers to dispersal. He incorrectly said that tree voles discarded the 

midrib of Douglas-fir needles. He estimated that a captive subadult tree 

vole ate 100 Douglas-fir needles an hour, day and night. He also noted 

that what appears to be good habitat can contain no tree voles. He stated 

that he usually selected forests of small (young) trees when searching for 

tree voles because nests were more accessible in small trees. He said that 

“Throughout their range these mice usually frequent the taller trees, and 

for this reason it is extremely difficult to estimate their numbers, for an 

immense Douglas-fir might harbor a dozen nests that cannot be recognized 

as such from the ground.” He reported that Harry Wilder found this to be 

the case when felling large trees on his Carlotta ranch. He claimed that on 

two consecutive mornings, Wilder and Irvine Clay captured 50 tree voles. 

We think the latter claim was an exaggeration on someone’s part, because 

museum specimens from Wilder and Clay never exceeded more than a few 

specimens in a day. Howell’s efforts to trap tree voles were unsuccessful, 

“As it is probably impossible to offer any bait that would attract the mice.” 

Like most tree climbers, he captured more females than males. He suggested that this was because males had smaller nests and eluded capture 

better than females. He stated, “Any opinion regarding the abundance of 

tree mice in a given area is little more than a haphazard guess.” He refuted 
64

GENERAL TECHNICAL REPORT PNW-GTR-909

the claim that tree voles were colonial by stating, “One who is familiar 

with the tree mouse is prone to refer to an aggregation of individuals in 

any well-defined area as a colony; but the animals really have no colonizing tendency and are not at all sociable in the strict sense of the word.” He 

went on to say that, “In reality the mice are of a markedly solitary disposition, the females usually being truculent and intolerant of other individuals.” Howell also recounted an episode in which Wilder observed a Steller’s 

jay demolishing a tree vole nest.

187.   Howell, A.B. 1928. The food and habitat preferences of Phenacomys albipes. Journal of Mammalogy. 9: 153–154.

Howell sent three white-footed vole stomachs to C.C. Sperry at the U.S. Biological Survey division of food habits research. The stomach contents were 

100 percent pulverized roots of herbaceous plants. This is in contrast to other 

reports in which white-footed voles fed on leaves of herbaceous trees or 

shrubs (Forsman and Swingle 2006, Manning et al. 2003, Voth et al. 1983). 

188.  Howell, A.B. 1936. Harry Edward Wilder. Journal of Mammalogy. 17: 

313–314.

In this obituary, Howell described the life of Harry Wilder, who collected 

many tree voles at his ranch from 1913 to 1927. Wilder was born in 1865 

in New Hampshire. He developed an interest in nature though his main 

income was as a beekeeper. In 1912, he and his wife Charlotte moved 

to a ranch near Carlotta, Humboldt County, California, where Wilder 

subsequently collected at least 46 Sonoma tree voles. Wilder collected an 

additional five Sonoma tree voles near Bridgeville in Humboldt County, 

and one Sonoma tree vole near Ruth in Trinity County. During 1913–1930, 

an additional 26 Sonoma tree vole museum specimens were collected at 

Wilder’s Ranch at Carlotta. At this time, just about everyone who was trying to collect tree voles in California stopped by to see Wilder, including 

Walter Taylor (1915), Brazier Howell (1926), Seth Benson (1931), Laurence 

Huey (1926), and Joseph Mailliard (1923).

189.   Hsu, T.C.; Benirschke, K. 1974. Phenacomys longicaudus (tree mouse). 

An Atlas of Mammalian Chromosomes. 8(373): 1–2. 

The authors examined karyotypes of red tree voles collected from Coos 

and Benton Counties, Oregon. They reported a complex situation in which 

the diploid numbers ranged from 40 to 52 depending on locality.
65

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

190.  Hubbard, C.A. 1940. A check list of the fleas of the Pacific Northwest 

(Washington, Oregon, northern California, and northwestern Nevada) 

with notes from southern California. Pacific University Bulletin. 37: 1–4.

Hubbard reported that he collected a flea (Megabothris quirini Rothschild 

1905) from a tree vole near Newberg, Yamhill County, Oregon, in 1935. We 

suspect this is one of the three tree vole specimens that he sent to the U.S. 

National Museum, but none of those specimens has a 1935 collection date.

191.   Hubbard, C.A. 1941. The fleas of rare western mice. Pacific University 

Bulletin. 37(9A): 1–4.

Hubbard identified one flea, Megabothris quirini Rothschild 1905, from 

two tree voles collected in 1938 in the Chehalem Mountains, northeast 

of Newberg, Oregon. These tree vole specimens are in the U.S. National 

Museum.

192.  Huey, L.M. 1926. Unpublished field notes. On file with: San Diego 

Natural History Museum, CA. 

These notes described a collecting trip in which Laurence Huey, a naturalist at the San Diego Natural History Museum, and Brazier Howell, from 

the Smithsonian Institute, drove up Highway 101 to the Trinidad whaling 

station in 1926 (http://www.sdnhm.org/about-us/our-museum/history/trinidad/). On this trip, they spent a day at the Wilder Ranch, near Carlotta, 

Humboldt County, California, where they collected one adult male and two 

juvenile Sonoma tree voles. In addition, Huey said that Howell collected 

six white-footed voles at Trinidad, Humboldt County, California.

193.  Huff, M.H.; Holthausen, R.S.; Aubry, K.B. 1992. Habitat management 

for red tree voles in Douglas-fir forests. Gen. Tech. Rep. PNW-GTR-302. 

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

Northwest Research Station. 16 p.

In this literature review, the authors cited Carey (1991) to support their 

contention that young forests were population sinks for tree voles. They 

did not mention the large number of specimens (including many breeding 

females) collected in young forests by previous researchers. They stated 

that red tree voles occurred almost exclusively in Douglas-fir forests 

but did not mention that tree voles along the coast in northwest Oregon 

occurred primarily in forests of western hemlock and Sitka spruce, as 

documented by previous researchers (Clifton 1960; Jewett 1920, 1930; 

Walker 1928, 1930). They stated that the only data available to describe 

habitat occupied by tree voles were data from pitfall studies conducted by 
66

GENERAL TECHNICAL REPORT PNW-GTR-909

the U.S. Forest Service Pacific Northwest Research Station and their contractors from 1984 to 1986. They did not acknowledge the wealth of information on tree vole habitat in the earlier literature and in the unpublished 

notes of Murray Johnson (1957–1985), William Hamilton, III, and Don 

Roberts (1957–1961), Vernon Bailey (1914), Chris Maser (1967–1974), Seth 

Benson (1930–1956), Walter Taylor (1913), Walter Dalquest (1943–1944), 

and others. They suggested that suitable habitat for tree voles was mesic

forests and that tree voles avoided dry environments. They discussed 

elevational limits based on data from pitfall studies and reported that they 

found no difference in the amount of old forest around stands in which 

tree voles were captured in pitfalls and stands in which no tree voles were 

captured in pitfalls. They proposed a model for evaluating habitat quality 

for tree voles based on basal area of Douglas-fir, density of large trees, 

height of live trees, stand age and size, site moisture class and elevation.

They suggested that tree voles were vulnerable to local extirpations from 

fragmentation and loss of habitat, and that vole populations had undergone 

significant declines owing to cutting of old-growth forests.

194.  Huff, M.; Mellen, T.K.; Hagestedt, R. 2001. A model to assess potential 

vertebrate habitat at landscape scales: HABSCAPES. In: Johnson, D.H.; 

O’Neil, T.A., managing directors. Wildlife: habitat relationships in Oregon 

and Washington. Corvallis, OR: Oregon State University Press: 544–550.

The authors said that red tree voles are closely associated with latesuccessional forests, are almost exclusively arboreal, and require habitat 

connectivity because they have limited dispersal capability. They modeled 

the amount and distribution of tree vole habitat on Mount Hood National 

Forest based on Huff et al. (1992). 

195.  Huff, R.; Van Norman, K.; Hughes, C.; Davis, R.; Mellen-McLean, 

K. 2012. Survey protocol for the red tree vole, Arborimus longicaudus (= 

Phenacomys longicaudus in the record of decision of the Northwest Forest 

Plan). Portland, OR: U.S. Department of Agriculture, Forest Service and 

U.S. Department of the Interior, Bureau of Land Management. 52 p.

This was the fourth edition of the survey protocol used by the U.S. Forest 

Service and Bureau of Land Management to document presence of tree 

voles in proposed project areas. The first half of the report included a 

review of tree vole biology and distribution. The second half was a revised 

protocol for surveys. 
67

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

196.   Humboldt Redwood Company. 2009a. 2008 Humboldt Redwood Company 

(HRC) habitat conservation plan (HCP), Sonoma tree vole annual report 

(HCP 6.9). Humboldt Redwood Company, Forest Science Department. 

http://mrc.com/pdf/HRC%20Terrestrial%20Wildlife%20Reports%20

2008/2008%20Sonoma%20Tree%20Vole%20Report.pdf. (August 2014). 

The authors conducted surveys of Sonoma tree vole nests in 64 stands that 

were subdivided into three forest size classes, including pole (15 to 28 cm 

diameter at breast height [dbh]), young (29 to 60 cm dbh), and old (>61 

cm dbh) forest. Each stand was surveyed with 500-m transects. Surveyors 

located 441 tree vole nests (215 “active,” 226 “inactive”). They located 

at least one tree vole nest in 89 percent of the stands surveyed, including 

19 of 24 pole stands, 10 of 12 young forest stands, and 26 of 28 old-forest 

stands. Most (n = 302) of the nests were in old forest. The authors did not 

report if any of the young or pole stands had remnant old trees, but they 

did mention that old trees appeared to maintain suitable tree vole habitat in 

young stands. They did not report how many nests were identified based 

on tree climbing as opposed to visual observation from the ground.

197.   Humboldt Redwood Company. 2009b. Humboldt Redwood Company, 

LLC management plan draft September 2009. http://hrcllc.com/pdf/HRCManagementPlan.pdf. (August 2014).

The authors stated that Sonoma tree voles were relatively common on their 

forested lands. Their conservation strategy was a habitat-based approach 

in which they proposed to sustain viable populations of tree voles in each 

watershed by retaining some large old conifers. They suggested riparian 

management zones would provide dispersal corridors for tree voles.

198.  Humboldt State University Vertebrate Museum. 1982–1984. Specimen 

catalog. Arcata, CA.

This specimen catalog included notes describing one Sonoma tree vole 

specimen traded to the University of Illinois and one Sonoma tree vole 

and one red tree vole traded to the U.S. Department of Agriculture, Forest 

Service, Pacific Northwest Research Station, Arcata, California.

199.  Hunter, J.E.; Bond, M.L. 2001. Residual trees: wildlife associations and 

recommendations. Wildlife Society Bulletin. 29: 995–999.

In the abstract, the authors stated that red tree voles were normally associated with older forest. They suggested that leaving old Douglas-fir trees 

in harvest areas might provide habitat for tree voles in young forests that 

reestablish on cutover areas. 
68

GENERAL TECHNICAL REPORT PNW-GTR-909

200.  Ingles, L.G. 1947. Mammals of California. Stanford, CA: Stanford 

University Press: 166–170.

In this short species account, Ingles described nests of tree voles and accurately described how they removed resin ducts from Douglas-fir needles 

before eating the rest. He said that tree voles breed throughout the year and 

suggested that weasels, martens, small hawks, and owls were important 

predators of tree voles. 

201.  Ingles, L.G. 1965. Mammals of the Pacific states. Stanford, CA: Stanford 

University Press: 284–300.

Ingles provided brief species accounts of red tree voles and white-footed 

voles. Most of the information presented is correct, except that Ingles errs 

when he says that tree vole nests include the mid-ribs and resin ducts of 

Douglas-fir needles. Mid-ribs of Douglas-fir needles are never found in 

tree vole nests, because they are consumed. 

202.  Institute for Natural Resources. 2011. Final report: science review of 

the Oregon Department of Forestry’s proposed species of concern strategy 

and the Board of Forestry’s state forests performance measures. Corvallis, 

OR: Oregon State University. 96 p. http://oregonstate.edu/inr/sites/default/

files/documents_reports/FINAL_Report_ODF_28Feb.pdf. (August 2014).

In this analysis, the Institute of Natural Resources science advisory 

team criticized the Oregon Department of Forestry (ODF) analysis of 

the effects of management on red tree voles, northern spotted owls, and 

aquatic amphibians. They argued that (1) tree voles required a more robust 

spatially explicit management strategy than was proposed by ODF, and 

(2) that the ODF analysis failed to account for the distinctive conservation 

challenge presented by the dusky tree vole. They proposed an alternative 

approach for analyzing the effects of management on tree voles. 

203.  Irwin, L.L.; Clark, L.A.; Rock, D.C.; Rock, S.L. 2007. Modeling foraging habitat of California spotted owls. Journal of Wildlife Management. 

71: 1183–1191.

In this paper on foraging behavior of spotted owls the authors incorrectly 

listed the red tree vole as a major prey item of California spotted owls 

(Strix occidentalis occidentalis) in the Sierra Nevada, where tree voles do 

not occur.
69

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

204.  Ishii, H.; Ford, E.D.; Boscolo, M.E.; Manriquez, A.C.; Wilson, M.E.; 

Hinckley, T.M. 2002. Variation in specific needle area of old-growth 

Douglas-fir in relation to needle age, within-crown position and epicormic 

shoot production. Tree Physiology. 22: 31–40.

There was nothing specific about tree voles in this paper, but the information about factors that influence needle growth in old-growth Douglas-fir 

may be important for understanding relationships between tree voles and 

their habitat. 

205.  Ishii, H.; Ford, E.D.; Dinnie, C.E. 2002. The role of epicormic shoot 

production in maintaining foliage in old Pseudotsuga menziesii (Douglasfir) trees II. Basal reiteration from older branch axes. Canadian Journal of 

Botany. 80: 916–926.

There was nothing specific about tree voles in this paper, but the description of epicormic branching in Douglas-fir will be of interest to anyone 

interested in the ecology or management of tree voles.

206.  Jaros, G. 2000. A grasp on elusive red voles [sic]: biologists study the 

tree top mammals’ nest sites, DNA to determine habitat needs. Roseburg 

News-Review. 13 September. http://www.fs.fed.us/treeclimbing/news/redvoles/. (August 2014).

This newspaper article described a field study in which researchers were 

collecting tissue samples for a genetic study of tree voles in western 

Oregon in 2000.

207.   Jewett, S.G. 1913–1955. Unpublished specimen catalog. Photocopy of 

original catalog. On file with: University of Puget Sound, Slater Museum 

of Natural History, Tacoma, WA.

This photocopy of Jewett’s unpublished specimen catalog lists 21 tree voles 

and 3 white-footed voles collected by Jewett from 1917 through 1936. He 

also listed six additional red tree voles that he received from other collectors. The original copy of this specimen catalog and Jewett’s original field 

notes were sent to the San Diego Museum of Natural History, where they 

were either lost or discarded.

208.  Jewett, S.G. 1915. Discovery of a rare rodent. The Oregon Sportsman. 3: 

37–38.

Jewett described the second white-footed vole known to science and the 

first in Oregon. He captured the vole in a mouse trap baited with oatmeal 

on 18 May 1914. The location was a riparian area of dense salmonberry 
70

GENERAL TECHNICAL REPORT PNW-GTR-909

(Rubus spectabilis) and sword fern (Polystichum munitum) near the

McKenzie River about 3.2 km downstream from Vida, Lane County. 

209.   Jewett, S.G. 1920. Notes on two species of Phenacomys in Oregon. 

Journal of Mammalogy. 1: 164–168.

Jewett introduced this paper with a somewhat poetic sentence, “In the 

dense forests of the Pacific Northwest, where the sun seldom penetrates the 

heavy growth of firs and spruces to mother earth, live two species of the 

most interesting of our small mammals, Phenacomys longicaudus in the 

tree tops, and Phenacomys albipes on the moss-covered floor of the forest.” He then provided a wealth of information on the habitat and distribution of the tree vole. He described lots of nests, and suggested that, “...the 

lower Rogue River watershed in Curry County is the center of abundance 

of these little tree mice.” He obviously was aware that tree vole nests can 

be confused with nests of woodrats and squirrels and made it clear that the 

nests he observed were indeed tree vole nests. He also mentioned that some 

of the larger nests that he observed were, “...evidently built by woodrats 

and later occupied by these mice.” He said that most nests, “...are located 

in Douglas-fir, but several have been noted in Sitka spruce near the coast.” 

He described two large nests “...in the barn yard...” at the John Adams 

Ranch in Curry County that were about 3 m above ground and 8 m apart 

where he caught a male and female, respectively. He said that, “Mr. Adams 

told me the nests had been there ten years to his knowledge, and as far as 

he could remember were the same as when he located his buildings on the 

ground.” He discussed escape behavior, including running out on limbs, 

climbing down trees head-first, and jumping and free-falling to the ground. 

He also described a case where a vole hid among the limbs when a small 

boy was induced to climb a likely looking nest tree at the Lowry Ranch 

on the Rogue River. He described a number of nests in young forests and 

talked about young boys who kept tree voles as pets. He also mentioned 

that tree vole nests had been found near Bonneville in Multnomah County, 

and Forest Grove in Washington County, but did not provide details. He 

described feeding behavior but didn’t quite get it right in terms of how the 

voles removed resin ducts from Douglas-fir.

210.   Jewett, S.G. 1923. A breeding record of Phenacomys longicaudus. 

Journal of Mammalogy. 4: 125.

Jewett provided a second-hand account of a case in which Elmer Williams 

captured a female tree vole and her three newborn young near Agness, 
71

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Curry County, Oregon, on 22 May 1921. The nest was about 12 m up 

in a Douglas-fir. Jewett said that Williams did not measure or save the 

specimens. 

211.   Jewett, S.G. 1930. A nest of Phenacomys silvicola in Oregon. Journal of 

Mammalogy. 11: 81–83.

Jewett and his family found a tree vole nest 5 m up in a Douglas-fir tree 

while walking through the woods near the beach at Netarts, Tillamook 

County, Oregon, on 14 August 1929. He had his 12-year-old son climb 

to the nest, at which point a vole ran out and jumped to the ground and 

escaped despite a search by the entire Jewett family. They returned the 

next morning and Jewett sent his son up the tree again, at which point the 

vole began to climb down the bole headfirst and was shot. The vole was 

an adult female that had recently given birth, but they did not find young 

in the nest. He said that she was a slow and cautious climber. He provided 

measurements of the type specimen of dusky tree vole collected by Alex 

Walker at Tillamook in 1916 and concluded that the species was indeed 

arboreal, as was previously documented by Howell 1926. He described the 

center of the nest as a, “...well matted mass of dissected leaves of fir.” 

212.   Jobanek, G.A. 1988. Searching for the tree vole: an episode in the 1914 

Biological Survey of Oregon. Oregon Historical Quarterly. 89: 369–400.

Jobanek provided a detailed account of the 1914 expedition in which 

Vernon Bailey travelled to Oregon to try to capture a tree vole. Bailey met 

with Alfred Shelton in Eugene and they spent several days climbing trees 

on Spencer Butte before finally catching a tree vole. Jobanek provided 

many interesting details about personalities, correspondence between 

people, campsites, methods, maladies, and modes of travel in 1914. This is 

an excellent read for anyone interested in the early history of mammalogy 

in Oregon.

213.   Johnson, D.H. 1936a. Aneides lugubris. Specimen catalog. On file 

with: University of California, Berkeley, Museum of Vertebrate Zoology 

Herpetology Collection. 

Johnson’s catalog described a case in which an arboreal salamander 

was collected in a Sonoma tree vole nest 4.8 km east of Duncans Mills, 

Sonoma County, California, on 1 March 1936. The specimen number for 

the arboreal salamander is MVZ19144.

214.   Johnson, D.H. 1936b. Unpublished field notes. On file with: University 

of California, Berkeley, Museum of Vertebrate Zoology.
72

GENERAL TECHNICAL REPORT PNW-GTR-909

Johnson observed a case in which a female Sonoma tree vole gave birth 

in captivity shortly after being captured with two large young. He did not 

take note of the fact, but this was another record that documented postpartum mating. He described the general appearance of the two neonates 

and used a microscope to observe five distinct mystacial rows of vibrissae. 

Johnson did not say so in his notes, but the voles (n = 10) he described were 

collected on field trips with Seth and Emma Benson.

215.   Johnson, K.N.; Franklin, J.F. 2013. Recommendations for future implementation of ecological forestry projects on BLM western Oregon forests. 

Unpublished report. 45 p. http://fes.forestry.oregonstate.edu/sites/fes.forestry.oregonstate.edu/files/PDFs/BLMJohnson_Franklin_APRIL16.pdf. 

(August 2014).

In this white paper on forest management on lands administered by the 

Bureau of Land Management, Johnson and Franklin proposed that monitoring of red tree voles be replaced by assessments using experimental 

treatments of vole habitat. They also suggested that the survey and manage 

mitigation standards for tree voles in the Northwest Forest Plan be replaced 

by designating the tree vole as a sensitive species. Johnson and Franklin 

did not say so, but if adopted, their recommendations would eliminate 

requirements to protect tree voles in areas proposed for harvest.

216.   Johnson, M.L. 1955–1984. Specimen catalog. On file with: University of 

Washington, Burke Museum, Seattle, WA.

Murray Johnson’s catalog included a list of tree voles and white-footed 

voles that he collected or that were born in his captive breeding colony. It 

also included many specimens that he obtained from others. 

217.   Johnson, M.L. 1957–1985. Unpublished field and lab notes. On file with: 

University of Washington, Burke Museum, Seattle, WA.

Murray Johnson was a biologist and thoracic surgeon whose avocation was 

searching for and studying tree voles. He published only a few papers, but 

his field notes, lab notes, and letters fill several file drawers and a large 

stack of three-ring binders. These notes described extensive field trips 

to collect tree voles in Oregon and California, as well as many pages of 

lab notes on tree voles that he kept in captivity. These notes document 

the history of 367 tree voles (Arborimus longicaudus, A. pomo, and their 

hybrids) that Johnson collected or bred in captivity as well as 169 tree voles 

that he obtained from others, including Doug Bake, Bill Rowland, Ricky 
73

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Brown, Chris Maser, Wayne Hammer, and Harry Schoenborn. A warning to anyone wanting to look at these notes—Murray’s handwriting was 

terribly difficult to read, and his lab notes were not well organized. But, 

hidden away in these notes are many interesting stories, including trials 

and tribulations involved in keeping tree voles in captivity, field trips when 

he was chased out of trees by bees, interactions with hostile landowners, 

bouts with poison oak (Toxicodendron diversilobum), and mysterious 

stomach ailments. 

218.   Johnson, M.L. 1965–1986. Unpublished captive breeding records. On 

file with: University of Puget Sound, Slater Museum of Natural History, 

Tacoma, WA.

These unpublished notes consist of detailed histories of the individual tree 

voles that Murray kept in captivity, including records of breeding attempts, 

birth and death dates, and outcomes of hybrid crosses between red tree 

voles and Sonoma tree voles. 

219.   Johnson, M.L. 1967–1969. Unpublished letters to Dorland Hammer. On 

file with: University of Washington, Burke Museum, Seattle, WA.

Murray Johnson wrote to Dorland Hammer on 15 March 1967 notifying 

him that all three red tree voles collected on their field trip a few weeks 

previous to Bills Peak, Coos County, Oregon, were males. Murray wrote 

Dorland on 7 March 1969 asking him to keep an eye out for tree voles in 

Coos County, Oregon, as there were few data from that area.

220.  Johnson, M.L. 1968. Application of blood protein electrophoretic studies 

to problems in mammalian taxonomy. Systematic Zoology. 17: 23–30.

Johnson used electrophoretic analyses of proteins in sera and hemoglobin 

to compare two heather voles and 16 red tree voles. The albumin migration was slightly faster for Arborimus when compared to Phenacomys. 

Based on these results, Johnson proposed combining the named forms of 

Arborimus longicaudus into a single species, and separating the genera 

Arborimus and Phenacomys. 

221.   Johnson, M.L. 1973. Characters of the heather vole, Phenacomys, and the 

red tree vole, Arborimus. Journal of Mammalogy. 54: 239–244.

Based on differences in juvenile development, numbers of mammary 

glands, bacula, phallus, testes, dentition, skulls, and skeletal morphology, 

Johnson suggested that tree voles should be placed in the genus Arborimus, 

whereas the heather vole should be retained in the genus Phenacomys. 

He suggested that differences in many morphological and physiological 
74

GENERAL TECHNICAL REPORT PNW-GTR-909

attributes indicated a major evolutionary divergence between Arborimus

and Phenacomys. He also speculated that the genus Arborimus evolved in 

the Klamath Mountains during the Pliocene.

222.  Johnson, M.L. 1974. Mammals. In: Wright, C.A., ed. Biochemical and 

immunological taxonomy of animals. New York: Academic Press: 1–87. 

Chapter 1.

In this book chapter, Murray described the utility of biochemistry and 

immunological data analysis as a taxonomic tool. He cited his previous 

analysis (Johnson 1968) as evidence that the subgenus Arborimus should 

be separated from the genus Phenacomys. 

223.  Johnson, M.L. 1988–1989. Unpublished letters to Doug Bake. On file 

with: University of Washington, Burke Museum, Seattle, WA.

In a letter dated 30 August 1988, Murray thanked Doug Bake for his interest, opinion, and data on tree voles in Sitka spruce and western hemlock 

forests. As described in Forsman and Swingle (2010), Doug provided 

Murray with many tree vole specimens from northwest Oregon. In another 

letter, Murray sent a memo to the other authors of the “Tree voles of 

Douglas-fir forests” manuscript stating that Martin Raphael had been 

asked to be a third technical editor. He also mentioned that an additional 

chapter on the heather vole was being added to the manuscript as a companion to the chapter on the white-footed vole.

224.  Johnson, M.L.; George, S.B. 1991. Species limits within the Arborimus 

longicaudus species-complex (Mammalia: Rodentia) with a description of 

a new species from California. Contributions in Science, Natural History 

Museum of Los Angeles County. 429: 1–16.

Based on karyotypic and anatomical differences, Johnson and George proposed splitting the tree vole into two species, the red tree vole in Oregon 

and the Sonoma tree vole in California. They suggested that low reproductive success of crosses between red tree voles and Sonoma tree voles 

indicated that there was a “strong reproductive barrier between Oregon 

and California populations.” The reproductive crosses referred to in this 

manuscript were all based on captive breeding experiments conducted by 

Johnson from 1955 through 1986. Unfortunately, the details are buried in 

his voluminous lab notes and were never published.

225.  Johnson, M.L.; Hammer, E.W. 1966–1968. Unpublished letters between 

Murray Johnson and Ellis “Wayne” Hammer. On file with: University of 

Washington, Burke Museum, Seattle, WA.
75

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

In a letter dated 10 February 1966, Wayne Hammer told Murray Johnson 

that he just got back from gathering data on an underground tree vole nest. 

There is a color picture of this nest in Murray’s files. To our knowledge, 

this is the only ground nest of a tree vole ever discovered in Oregon. In 

another letter, Wayne stated he had two captive tree voles.

226.  Johnson, M.L.; Johnson, S. 1982. Voles, Microtus species. In: Chapman, 

J.A.; Feldhamer, G.A., eds. Wild mammals of North America: biology, management, and economics. Baltimore, MD: The John Hopkins 

University Press: 326–354. Chapter 16.

Murray Johnson and his wife Sherry briefly mentioned red tree voles and 

white-footed voles in this excellent species account of North American 

Microtus. 

227.   Johnson, M.L.; Kesner, M.H. 1982. Unpublished letters. On file with: 

University of Washington, Burke Museum, Seattle, WA.

Mike Kesner sent a letter to Murray Johnson thanking him for providing 

two tree voles that Kesner used for his papers on myology and manus 

musculature of microtines (Kesner 1980, 1986). Kesner was planning on 

conducting field work on tree voles and asked Johnson where he should go 

to obtain specimens. Johnson responded by providing a long list of sites 

where he had collected tree voles. He also warned Kesner to beware of 

poison oak when climbing tree vole nest trees and said that he had to be 

especially careful about poison oak “because of my own sensitivities.” 

228.  Johnson, M.L.; Lidicker, W.Z., Jr. 1977. Unpublished letters. On file 

with: University of Washington, Burke Museum, Seattle, WA.

This exchange of letters reflected a difference of opinion between Murray 

Johnson and Bill Lidicker over who should be in possession of the Sonoma 

tree voles collected and raised in captivity by Don Roberts and Bill Hamilton, III when they were graduate students at University of California, 

Berkeley. Lidicker wanted the specimens housed at the University of 

California, Berkeley, Museum of Vertebrate Zoology (MVZ) where the 

work was done, whereas Murray felt they should stay at the Slater Museum 

of Natural History in Tacoma, Washington, which was where Hamilton 

and Roberts donated them. In a letter dated 22 September 1977, Johnson 

acknowledged that the work was done while Roberts and Hamilton were 

students at Berkeley, but he disagreed with Lidicker that the specimens 

should be housed at MVZ because the study that Roberts and Hamilton 

conducted on tree voles was independent of their graduate work. Lidicker 
76

GENERAL TECHNICAL REPORT PNW-GTR-909

did not argue further but did respond in a letter on 26 October 1977 in 

which he mentioned that an additional 16 tree voles that were from Roberts 

and Hamilton at the MVZ were sent to the Slater Museum. We believe 

these were the 16 uncataloged specimens that we found at the Slater 

Museum when we visited the museum in 2011.

229.   Johnson, M.L.; Maser, C. 1982. Generic relationships of Phenacomys 

albipes. Northwest Science. 56: 17–19.

Johnson and Maser proposed changing the genus of the white-footed vole 

from Phenacomys to Arborimus. They based this recommendation on 

differences in morphological characteristics and the number of diploid 

chromosomes.

230.  Johnson, M.L.; Maser, C. 1989. Unpublished letter. On file with: 

University of Washington, Burke Museum, Seattle, WA.

In this letter, Murray Johnson and Chris Maser updated their coauthors 

on the tree vole book that they were working on. They discussed possible funding from the U.S. Forest Service and mentioned that they had 

approached Martin Raphael about becoming a technical editor.

231.   Johnson, M.L.; Maser, C.; Roberts, D.R. 1989. Tree voles of the 

Douglas-fir forest. Unpublished manuscript. On file with: University of 

Washington, Burke Museum, Seattle, WA.

This working draft of a book on tree voles was never completed. It 

included chapters on fossil history, taxonomy, anatomy, distribution, forest 

ecosystem, field methods, habitat, nests, behavior, food and water requirements, reproduction, juvenile development, demography, parasites, captives, white-footed voles, and heather voles.

232.  Johnson, M.L.; Maser, C.; Roberts, D.; Hammer, W. 1987. The red tree 

mouse and relatives (genus Arborimus): a study in extreme specialization. 

Unpublished document. On file with: University of Washington, Burke 

Museum, Seattle, WA.

This outline was developed for a book that Murray and his coauthors 

wanted to write. There were at least three versions of the outline. The book 

was started but never finished.

233.  Johnston, A.N. 2006. Relative abundance and habitat associations of 

small mammals in two forest types in southern Oregon. Corvallis, OR: 

Oregon State University. 151 p. M.S. thesis.
77

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

In 17,600 trap-nights, Aaron Johnston did not capture a single red tree vole 

or white-footed vole in live-traps on the Cascade-Siskiyou National Monument east of Ashland, Jackson County, Oregon.

234.  Jollie, M. 1945. Unpublished field notes and specimen catalog. On file 

with: University of California, Berkeley, Museum of Vertebrate Zoology.

In his field notes and specimen catalog, Malcolm Jollie reported that he 

collected four adult and six juvenile tree voles from Jenner and Anchor 

Bay in Sonoma County, and Laytonville in Mendocino County, California. 

Two of the juveniles were from a site 1.6 km east of Jenner. We suspect 

that the latter site was the “Poison Oak Knoll” location on the Russian 

River, where Seth Benson, Don Roberts, William Hamilton III, Murray 

Johnson, Alan Ziegler, and others (including us) collected tree voles. Jollie 

said that he caught a female tree vole in a snap-trap 12.9 km southwest 

of Laytonville. The snap-trap was set on a log between two Douglas-fir 

trees. He also said that he looked for tree vole nests farther inland between 

Willets and Ukiah in Mendocino County, without success.

235.  Jones, C.; Hoffmann, R.S.; Rice, D.W.; Engstrom, M.D.; Bradley, 

R.D.; Schmidly, D.J.; Jones, C.A.; Baker, R.J. 1997. Revised checklist 

of North American mammals north of Mexico, 1997. Museum of Texas 

Tech University Occasional Papers. 173: 1–20.

The authors reported that Johnson and George (1991) applied the generic 

name Arborimus to the white-footed vole and red tree vole, and that 

Johnson and George (1991) further proposed that the Sonoma tree vole was 

distinct from the red tree vole. The authors did not say whether they agreed 

or disagreed with these changes.

236.  Jones, J.M. 2003. Habitat associations and ecology of the Sonoma tree 

vole (Arborimus pomo) in northwestern California. Arcata, CA: Humboldt 

State University. 55 p. M.S. thesis. 

Jones used distance sampling methods to compare relative abundance of 

Sonoma tree vole nests in different forest size classes. She concluded that 

tree vole nests were significantly more abundant in mature forest (>61 cm 

diameter at breast height [dbh]), than in pole forest (15 to 28 cm dbh) or 

young forest (29 to 61 cm dbh). She also modeled relationships between 

habitat variables and nest abundance at 61 sites. The best predictive models 

included percentage basal area of Douglas-fir, dbh of dominant trees, total 

basal area, percentage slope, and slope position.
78

GENERAL TECHNICAL REPORT PNW-GTR-909

237.   Jones, J.; Zabel, C.J. 2003. Habitat associations and ecology of 

the Sonoma tree vole (Arborimus pomo) in northwestern California 

[Abstract]. In: Joint annual meeting of the Society of Northwestern 

Vertebrate Biology and California Coast Chapter of the Wildlife Society. 

Northwestern Naturalist. 84: 103.

In this published abstract, the authors compared relative abundance of 

Sonoma tree vole nests in three size classes of Douglas-fir forest on lands 

owned by The Pacific Lumber Company (currently, Humboldt Redwood 

Company) in northwestern California. Vole nests were significantly 

more abundant in mature than in young or pole forests, but there were 

no significant differences between young and pole stands. The authors 

also evaluated associations between landscape-level habitat attributes and 

Sonoma tree vole abundance within Humboldt Redwoods State Park, Redwood National Park, and the Angelo Coast Range Reserve in northwestern 

California. Sixty-one sites were searched for tree vole nests using line 

transects, and nest densities were estimated using program DISTANCE. 

Nest densities were positively correlated with diameter at breast height of 

the dominant trees, percentage basal area of Douglas-fir, and slope position. Total basal area and percentage slope were also important predictors 

of nest density.

238.  Kagan, J.; Christy, J.; Vrilakas, S.; Gaines, E.; Alton, C.; Koepke 

Wise, L.; Doyle, E. 2010. Rare, threatened and endangered species of 

Oregon. Portland, OR: Oregon Biodiversity Information Center, Institute 

for Natural Resources, Portland State University. 105 p.

The authors compared conservation status rankings of red tree voles by 

four different conservation organizations. Nature Serve/Natural Heritage 

Network Ranks listed the red tree vole as rare, uncommon, or threatened 

but not immediately imperiled. Oregon Department of Fish and Wildlife 

listed the red tree vole as vulnerable in the Coast Ranges. The U.S. Fish 

and Wildlife Service listed the red tree vole as a species of concern, and 

the Oregon Biological Information Center listed the red tree vole as a species of conservation concern.

239.  Kagan, J.; Christy, J.; Vrilakas, S.; Gaines, E.; Alton, C.; Wise, L.; 

Howell, K. 2013. Rare, threatened and endangered species of Oregon. 

Portland, OR: Oregon Biodiversity Information Center, Institute for 

Natural Resources, Portland State University. 111 p.
79

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

The authors updated conservation status rankings for the red tree vole and 

the white-footed vole from their 2010 version. The rankings were identical 

except that the U.S. Fish and Wildlife Service listed the red tree vole for 

the north Oregon coast as endangered or threatened.

240.  Kays, R.W.; Wilson, D.E. 2002. Mammals of North America. Princeton, 

NJ: Princeton University Press. 240 p.

On pages 124–125, the authors provided a brief description of red tree 

voles and Sonoma tree voles.

241.  Kelsey, R.G.; Forsman, E.D.; Swingle, J.K. 2009. Terpenoid resin distribution in conifer needles with implications for red tree vole, Arborimus 

longicaudus, foraging. Canadian Field-Naturalist. 123: 12–18.

The authors found that terpenoids were located exclusively in the resin 

ducts in Douglas-fir, Sitka spruce, and western hemlock needles. They 

concluded that tree voles were able to largely avoid exposure to terpenoids 

in Douglas-fir and western hemlock by discarding the resin ducts. They 

reported that resin ducts in Sitka spruce were discontinuous and sometimes 

absent entirely. How tree voles avoided ingesting terpenoids when feeding 

on Sitka spruce was not documented.

242.  Kesner, M.H. 1978. Myology of the cranial and pectoral appendicular 

regions of rodents of the subfamily Microtinae. Amherst, MA: University 

of Massachusetts. 197 p. Ph.D. dissertation. 

Kesner dissected and compared the masticatory musculature of 12 species 

of voles, including 2 red tree voles and 2 heather voles. Based on myological differences, he suggested that Arborimus was a distinct genus from 

Phenacomys.

243.  Kesner, M.H. 1980. Functional morphology of the masticatory musculature 

of the rodent subfamily Microtinae. Journal of Morphology. 165: 205–222.

In this first publication from his dissertation, Kesner said that the tree vole 

had one of the more derived microtine digastric muscles.

244.  Kesner, M.H. 1986. The myology of the manus of microtine rodents. 

Journal of Zoology, London. (A) 210: 1–22. 

Kesner examined the structure of the forearm and hand in voles and mice. 

He concluded that the tree vole manus had a combination of primitive and 

derived characters that included the most highly developed “clasping” trait 

of the arvicolines sampled.
80

GENERAL TECHNICAL REPORT PNW-GTR-909

245.  Koffman, R. 2012. Forest Park’s wildlife mysteries: collected data show 

new species and some declining bird life and establish an information 

baseline. The Oregonian. 12 November.

This newspaper report described a talk given by Portland Parks and 

Recreation wildlife biologist John Deshler on 7 November 2012. Deshler 

suggested that the gray fox (Urocyon cinereoargenteus), dusky-footed 

woodrat (Neotoma fuscipes), and red tree vole would be good candidates 

for reintroduction to Forest Park in Portland, Oregon.

246.  Koford, M. 1949. Unpublished field notes. On file with: University of 

California, Berkeley, Museum of Vertebrate Zoology.

Mary Koford reported that she, Marietta Voge, Anita Pearson, and Lois 

Taylor searched for tree vole nests on 19 May 1949 near Fort Ross, Sonoma 

County, California. They captured an adult female that gave birth to four 

young the next day in captivity.

247.   Kritzman, E.B. 1977. Little mammals of the Pacific Northwest. Seattle, 

WA: Pacific Search Press: 20.

Kritzman provided a short life-history account of the red tree vole.

248.  Krohne, D.T. 1981. Intraspecific litter size variation in Microtus californicus: variation within populations. Journal of Mammalogy. 62: 29–40. 

Krohne used the red tree vole as an example of one of many species of 

arvicolines in which neonate birth weights were negatively correlated with 

litter size (Hamilton 1962).

249.  Lamb, C.C. 1932. Unpublished field notes. On file with: University of 

California, Berkeley, Museum of Vertebrate Zoology.

Lamb reported that he and Harley Watson captured three tree voles in 

northwest California near Mad River, Trinity County, and one tree vole 

near Bridgeville, Humboldt County.

250.  Larrison, E.J. 1976. Mammals of the northwest: Washington, Oregon, 

Idaho, and British Columbia. Seattle, WA: Seattle Audubon Society: 

111–112. 

Larrison briefly described red tree voles and white-footed voles.

251.  Lawlor, T.E. 2003. Faunal composition and distribution of mammals in 

western coniferous forests. In: Zabel, C.J.; Anthony, R.G., eds. Mammal 

community dynamics: management and conservation in the coniferous forests of western North America. Cambridge, United Kingdom: 

Cambridge University Press: 41–80. Chapter 3.
81

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

In this book chapter, Lawlor suggested that tree voles may respond differently to logging of old-growth forests in different parts of their range. To 

support this hypothesis, he cited studies in Oregon in which authors found 

that tree voles were strongly associated with old-growth forests (Aubry 

et al. 1991, Corn and Bury 1991, Gilbert and Allwine 1991) and papers in 

California in which the association with old forests was less conclusive 

(Meiselman 1992, Meiselman and Doyle 1996, Raphael 1988, Thompson 

and Diller 2002).

252.  Lehmkuhl, J.F.; Ruggiero, L.F. 1991. Forest fragmentation in the 

Pacific Northwest and its potential effects on wildlife. In: Ruggiero, L.F.; 

Aubry, K.B.; Carey, A.B.; Huff, M.H., tech. coords. Wildlife and vegetation of unmanaged Douglas-fir forests. Gen. Tech. Rep. PNW-GTR-285. 

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

Northwest Research Station: 35–46. 

The authors ranked forest amphibians, birds, and mammals according to 

their perceived risk of local extinction in western Oregon and Washington. 

Of 93 species examined, red tree voles and white-footed voles ranked near 

the top.

253.  Lewis, R.E.; Lewis, J.H.; Maser, C. 1988. The fleas of the Pacific 

Northwest. Corvallis, OR: Oregon State University Press. 296 p.

The authors listed fleas that had been found on white-footed voles, 

including Catallagia sculleni sculleni, Epitedia scapani, Hystrichopsylla 

occidentalis occidentalis, Megabothris (Amegabothris) abantis, and 

Rhadinopsylla sectilis goodi.

254.  Little, L. 1968. Alfred Brazier Howell, 1886–1961. Journal of 

Mammalogy. 49: 732–742.

In this obituary, Little said that he accompanied Howell to Alton, Humboldt County, California, to collect tree voles in 1923. He said that in late 

summer, Howell collected five tree voles at Harry Wilder’s Ranch near 

Carlotta. We found evidence from museum specimen tags that Howell kept 

three of those voles in captivity and that one of them produced one young. 

We could not find Howell’s field notes, but we did find museum specimens 

of 18 Sonoma tree voles and 5 white-footed voles that he collected.

255.  Livingston, S. 2009. Unpublished meeting notes regarding genetics of 

red tree voles, 6 February 2009, Corvallis, Oregon. On file with: U.S. 

Department of the Interior, Fish and Wildlife Service, 911 NE 11th Ave., 

Portland, OR 97232.
82

GENERAL TECHNICAL REPORT PNW-GTR-909

These notes described minutes of a meeting in which Sue Livingston of the 

U.S. Fish and Wildlife Service discussed the taxonomic status of red tree 

voles and dusky tree voles with Mark Miller, U.S. Geological Survey, Forest and Rangeland Ecosystem Science Center. Miller stated that his paper 

(Miller et al. 2006) did not address the subspecies issue and that he did not 

work on the Bellinger et al. (2005) paper, so he could not say if his paper 

enhanced or contradicted Bellinger et al. (2005). Miller et al. (2006) found 

that there were genetically discrete populations in the north Coast Ranges 

and North Cascades compared to the rest of the tree vole population farther 

south. This north-south discontinuity has also been observed in plants 

(Soltis et al. 1997; Plant Systematics and Evolution 206: 353–373) and the 

southern torrent salamander (Rhyacotriton variegatus; Miller et al. 2006 

Journal of Heredity 6: 561–570).

256.  Livingston, S. 2009. Interpretation of red tree vole genetics papers. 

Questions posed by Sue Livingston, U.S. Department of the Interior, 

Fish and Wildlife Service to Mark Miller and Sue Haig, U.S Department 

of the Interior, Geological Survey, Forest and Rangeland Ecosystem 

Science Center, Corvallis, OR. 6 p. Unpublished report. On file with: U.S. 

Department of the Interior, Fish and Wildlife Service, 911 NE 11th Ave., 

Portland, OR 97232. 

In this unpublished agency report, Livingston discussed the subspecific 

taxonomy of tree voles in Oregon based on comments she received from 

Mark Miller and Sue Haig at the U.S. Geological Survey Forest and Rangeland Ecosystem Science Center in Corvallis, Oregon. Miller and Haig 

suggested that the north/south genetic discontinuity between tree voles 

could possibly support subspecies designation.

257.   Maguire, C.C. 2002. Dead wood and the richness of small terrestrial 

vertebrates in southwestern Oregon. In: Laudenslayer, W.F., Jr.; Shea, P.J.; 

Valentine, B.E.; Weatherspoon, C.P.; Lisle, T.E., tech. coords. Proceedings 

of the symposium on the ecology and management of dead wood in western forests. Gen. Tech. Rep. PSW-GTR-181. Albany, CA: U.S. Department 

of Agriculture, Forest Service, Pacific Southwest Research Station: 

331–345.

Maguire used pitfall traps to sample small mammals at four locations in 

the central Cascades in Douglas and Lane Counties in Oregon. There were 

6 replicate trapping grids at each location, for a total of 24 sampling grids. 

Tree voles were captured at three of the four locations (four at Little River, 
83

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

three at Layng Creek, and one at Dog Prairie). White-footed voles were 

captured at only one location (Layng Creek), where they were captured in 

three of six stands sampled.

258.  Mailliard, J. 1921–1926. Unpublished field notes. On file with: 

California Academy of Sciences, San Francisco, CA.

In 1921, Joseph Mailliard, Chester Lamb, and Irvine Clay collected two 

adult and three juvenile Sonoma tree voles at Kneeland, southeast of 

Arcata, Humboldt County, California. In 1926, Mailliard and Raleigh 

Borell collected 11 Sonoma tree voles at the McClellan Ranch near Bridgeville, Humboldt, County, and one tree vole from Harry Wilder’s ranch 

near Carlotta, Humboldt County, California. One of the tree voles was collected by shooting the nest. Mailliard also said that Borell found the head 

and shoulders of a tree vole under a tree, and that Harry Wilder told him 

that he and Irvine Clay collected 50 tree voles in one day. The latter story 

was repeated by Howell (1926), but we found no evidence that it was true. 

259.  Mailliard, J. 1923. Field work among the birds and mammals of the 

northern coast of California in 1921. Proceedings of the California 

Academy of Sciences. (Series 4): 1–26.

In autumn 1921, Joseph Mailliard, Chester Lamb, and Irvine Clay conducted field work in northwestern California. Mailliard wrote, “Mr. Clay 

not only led the way to Kneeland, but gave several demonstrations along 

the road through the fir forest of how to capture the long-tailed tree mouse. 

In fact, if it had not been for this practical demonstration of just how to 

proceed in the matter, it is doubtful any mice would have been secured by 

the field party. At least in part of the springtime, the larger nests contain 

one female with (usually) two or three young, while the male has a small 

nest of his own, probably often in another tree nearby, as no nests of males 

were discovered in the small sapling fir trees in which the females’ nests 

were found. In each case, there were branches interlocking from adjacent 

trees that could furnish means of communication with the family nest. But 

few nests discovered by us were occupied, and these only in forest that had 

been extensively thinned out by wood cutting. Careful watch was kept for 

the nests of this mouse in every locality visited by us, but the only ones 

discovered were on the road between Eureka and Kneeland Post Office.” 

Later on in the paper Mailliard wrote, “A sharp watch was kept in all 

suitable forest country for nests of the long-tailed tree mouse as outlined 
84

GENERAL TECHNICAL REPORT PNW-GTR-909

elsewhere in this paper, but very few nests were found, and of those discovered in accessible situations the great majority appeared deserted.” His 

comment about occupied nests being found only in stands that had been 

heavily thinned by woodcutters is interesting because we found no other 

references in which researchers found tree voles in heavily thinned stands. 

260.  Manning, J.A. 2002. Small mammal responses to silvicultural fuels treatments in southwest Oregon. Corvallis, OR: Oregon State University. 68 p. 

M.S. thesis.

Manning used Sherman live-traps to sample small mammal populations in 

the Sterling Creek watershed 10.0 km southeast of Ruch, Jackson County, 

Oregon. Of 593 small mammals captured, none were tree voles or whitefooted voles. His study adds to the considerable body of evidence that tree 

voles do not occur east of the Applegate River.

261.   Manning, J.A.; Edge, W.D. 2004. Small mammal survival and downed 

wood at multiple scales in managed forests. Journal of Mammalogy. 85: 

87–96.

Manning and Edge used Sherman live-traps to sample small mammals 

in Salmonberry Creek and Little Lobster Creek, approximately 9.5 km 

southwest of Alsea, Lincoln County, Oregon. They captured 926 small 

mammals but did not state the number of trap-nights sampled. No red tree 

voles or white-footed voles were captured.

262.  Manning, J.A.; Edge, W.D. 2008. Small mammal responses to fine 

woody debris and forest fuel reduction in southwest Oregon. Journal of 

Wildlife Management. 73: 625–632.

This was the published version of Jeff Manning’s thesis. They used Sherman live-traps to sample small mammals in Jackson County, Oregon. They 

captured 538 deer mice and 248 red-backed voles (Myodes californicus) 

but did not state how many trap-nights or what other small mammals they 

captured. Why the numbers differ from his thesis is not clear. 

263.  Manning, T.; Maguire, C.C. 1999. A new elevation record for the 

red tree vole in Oregon: implications for National Forest management. 

American Midland Naturalist. 142: 421–423.

Manning and Maguire reported that they captured a 24-g male red tree 

vole in a pitfall trap at 1600 m elevation near Dog Prairie Creek in the 

central Cascades, Douglas County, Oregon. However, when we contacted 

Tom Manning to ask for details, he said that the elevation in the published 
85

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

note was wrong and that the elevation of the capture site was actually 1500 

m. This record represented a considerable range expansion.

264.  Manning, T.; Maguire, C.C.; Jacobs, K.M.; Luoma, D.L. 2003. 

Additional habitat, diet and range information for the white-footed vole 

(Arborimus albipes). American Midland Naturalist. 150: 115–122. 

Based on a sample of 13 white-footed voles captured in pitfall traps in the 

Oregon Cascades, the authors described a southward range extension. They 

concluded that the species was semiarboreal based on presence of hazel 

pollen from unopened catkins in fecal pellets. Capture sites were associated with brushy areas of red alder and California hazel, further supporting 

the hypothesis that this species tends to be associated with thick brush and 

shrubs. In stomach samples, the authors found that green plant tissues were 

present in 9 to 91 percent of the fields viewed under a microscope (mean = 

50 percent), and that arthropods were present in 3 of the 13 samples examined. Spores from hypogeous fungi were present in all but one sample.

265.  Marcot, B.G.; Livingston, S. 2009. Report on the red tree vole listing advisory panel held August 25–26, 2009 at the U.S. Department of 

the Interior, Fish and Wildlife Service Oregon Office. On file with: U.S. 

Department of the Interior, Fish and Wildlife Service, Portland, OR. 51 p.

On 25 and 26 August 2009, the U.S. Fish and Wildlife Service convened 

a four-person listing advisory panel workshop for the red tree vole. The 

panel listened to reports from research scientists on management status, 

genetics, biology, distribution, and habitat associations of the red tree 

vole. Using a modified Delphi scoring method, the panel then ranked 

the distinct population segment of tree voles in the northern Oregon 

Coast Ranges as threatened (67 percent), not warranted (32 percent), or 

endangered (1 percent). The panel identified the following evidence in 

their decisionmaking process: loss of area and connectivity of old-growth 

forests from harvesting and fire, low reproductive potential, high predation 

rate, lack of guidelines and management, and isolated populations.

266.  Marcot, B.G.; Molina, R. 2006. Chapter 8: conservation of other species associated with older forest conditions. In: Haynes, R.W.; Bormann, 

B.T.; Lee, D.C.; Martin, J.R., tech. eds. Northwest Forest Plan—the first 

10 years (1994–2003): synthesis of monitoring and research results. 

Gen. Tech. Rep. PNW-GTR-651. Portland, OR: U.S. Department of 

Agriculture, Forest Service, Pacific Northwest Research Station: 145–179.
86

GENERAL TECHNICAL REPORT PNW-GTR-909

Marcot and Molina reported that much research had been conducted on the 

distribution and biology of the red tree vole subsequent to the adoption of 

the Northwest Forest Plan and that federal land managers were frustrated 

because the requirement to protect red tree vole nests found during predisturbance surveys was interfering with timber harvest.

267.   Martin, A.C.; Zim, H.S.; Nelson, A.L. 1951. American wildlife and 

plants. New York, NY: McGraw-Hill Book Company. 500 p.

In this brief species account, the authors stated that tree vole diets contained 0.5 to 2.0 percent Douglas-fir seeds. We found no evidence supporting this statement.

268.  Martin, K.J. 1998. Habitat associations of small mammals and amphibians in the central Oregon Coast Range. Corvallis, OR: Oregon State 

University. 86 p. Ph.D. dissertation. 

Karl Martin used pitfall traps to sample small mammals in the Oregon 

Coast Ranges (Nestucca River, Lobster Creek, and Drift Creek drainages) 

from 1995 through 1996. He captured 16,892 small mammals, including 

42 red tree voles and 85 white-footed voles. The number of individuals 

captured per 100 trap-nights was 0.033 for tree voles and 0.066 for whitefooted voles. Although capture rates were highest in forests with large 

trees (>53 cm diameter at breast height), none of the differences among forest size classes were significant. Most of Martin’s tree vole specimens were 

donated to the University of Alaska, Fairbanks, Museum of the North. 

Martin captured an additional 16 tree voles in pitfall traps during a pilot 

study near Prairie Mountain, Benton County, Oregon in 1994. We found 

the skulls of those specimens in a freezer at Oregon State University; they 

are currently housed at the U.S. Department of Agriculture Forest Service, 

Pacific Northwest Research Station, Corvallis, Oregon. 

269.  Martin, K.J.; Conroy, C.; Cook, J. 1996. Unpublished letters. On file 

with: University of Alaska, Fairbanks, Museum of the North.

These museum files contained several letters and specimen lists from 

Karl Martin to Conroy and Cook regarding the capture locations of 38 red 

tree voles and 74 white-footed voles that Martin captured in Oregon and 

donated to the museum.

270.  Martin, K.J.; McComb, W.C. 2002. Small mammal habitat associations 

at patch and landscape scales in Oregon. Forest Science. 48: 255–266.

This was the published version of Karl Martin’s dissertation. The authors 

captured 42 red tree voles and 85 white-footed voles in pitfall traps. They 

concluded that tree vole capture rates were positively associated with 
87

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

mature conifer forests and were negatively affected by decreasing patch 

densities of large sawtimber (fewer individual patches).

271.  Martin, L.D. 1994. A new genus of Miocene vole possibly related to 

Phenacomys. Institute for Tertiary-Quaternary Studies: TER-QUA 

Symposium Series 2: 129–130.

Martin described Propliophenacomys parkeri, a new genus and species 

that represented the oldest known member of the lineage leading to Pliophenacomys, and possibly to Phenacomys.

272.  Martin, R.A. 1989. Arvicolid rodents of the early Pleistocene Java 

local fauna from north-central South Dakota. Journal of Vertebrate 

Paleontology. 9: 438–450.

Martin found eight arvicolid species at a site in South Dakota, including 

Pliophenacomys. He compared the teeth of these fossil voles to the teeth of 

heather voles, white-footed voles, and red tree voles.

273.  Maser, C. 1963–1966. Unpublished data. On file with: University of 

Washington, Burke Museum, Seattle, WA.

These notes included a large binder with original field data forms describing red tree vole nests that Chris Maser found in Oregon.

274.   Maser, C. 1963–1972. Unpublished specimen catalog. On file with: 

University of Puget Sound, Slater Museum of Natural History, Tacoma, WA.

Maser’s specimen catalog listed 99 red tree voles and 1 white-footed vole 

collected in Oregon from 1963 through 1972. 

275.  Maser, C. 1965a. Notes on the contents of owl pellets found in Oregon. 

Murrelet. 46: 44.

Maser reported remains of one Phenacomys silvicola and one Phenacomys 

spp. that he found in owl pellets at 610 m elevation on the north side of 

Marys Peak, Benton County, Oregon, in 1963. 

276.  Maser, C. 1965b. Spotted owl preys on dusky three [sic] mice. Murrelet. 

46: 46.

On 19 February 1965, Maser shot an adult female northern spotted owl 

with a pellet rifle on Oregon State University’s McDonald Forest, a few 

kilometers north of Corvallis, Benton County, Oregon. The stomach of the 

owl contained remains of two adult tree voles. The skulls are in the Oregon 

State University collection (can No. 256). Maser described the habitat as a 

dense stand of young Douglas-fir. The owl was roosting about 60 m southeast of a nest in which Maser found a female tree vole and two newborns. 
88

GENERAL TECHNICAL REPORT PNW-GTR-909

277.   Maser, C.O. 1966. Life histories and ecology of Phenacomys albipes, 

Phenacomys longicaudus, Phenacomys silvicola. Corvallis, OR: Oregon 

State University. 221 p. M.S. thesis.

This thesis included an extensive literature review on tree voles, whitefooted voles, and heather voles, as well as a wealth of information that 

Maser collected personally, including observations on nest attributes, 

reproduction, sex ratios, escape behavior, feeding behavior, and habitat. 

Based on a single ground nest located by Wayne Hammer, Maser speculated that tree voles might live underground during part of the year. He 

stated that stick size was not a valid criterion to determine which species 

built a nest because tree voles sometimes cut twigs that were nearly as 

large as those cut by squirrels or woodrats. He described the free-fall position that tree voles assume when jumping from their nests. He observed 

tree voles licking water off needles in captivity and suggested that tree 

voles also lick water off needles in the wild. 

In April through June 1965, Maser conducted a population density study 

of tree voles. He searched a 12.4-ha area of young forest for tree vole nests 

and dissected all nests that he detected to determine the number of voles in 

each nest. He captured 40 voles, including 12 adults, 3 subadults, and 25 

juveniles. Thus adult density was 0.96 per hectare. This was a minimum 

estimate, as he did not know how many voles went undetected. The study 

plot was located in Benton County, 7.3 km SW of Monroe.

278.  Maser, C. 1967–1974. Unpublished field notes. On file with: University 

of Washington, Burke Museum, Seattle, WA.

These field notes described Maser’s collecting activities in western Oregon 

from 1967 through 1974. He collected tree voles with Rita Maser, Jack and 

Rob Booth, Ruth and Damon Smith, Wayne and Dorland Hammer, Ron 

Nussbaum, and Eric Forsman. He also obtained four tree vole specimens 

from Bill Harvey (a logger) who collected them while clearcutting an oldgrowth forest on Prairie Mountain, Benton County, Oregon. 

279.   Maser, C. 1998. Mammals of the Pacific Northwest. Corvallis, OR: 

Oregon State University Press. 406 p.

On pages 219–225, Maser described the appearance, distribution, habitat, 

diet, feeding behavior, reproduction, dispersal, nests, predators, escape 

behavior, and crypsis of red tree voles. He described pelage color variation 
89

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

of red tree voles from north to south. He stated that young trees were not 

structurally able to fulfill nesting requirements or offer much protection 

from weather. He called the tree vole one of the most specialized arboreal 

mammals in the world. He said that young and tender conifer needles are 

sometimes eaten whole and that some twigs are split, apparently to obtain 

their pithy center. He said that tree voles typically construct their nests 

at or above the height of the first live branch, and that female tree voles 

may leave their young and disperse to a new nest once the young become 

capable of fending for themselves [definitely not true in most cases]. He 

speculated that many hundreds of generations of tree voles can live in one 

large Douglas-fir. 

Maser considered the white-footed vole (pages 218–219) one of the rarest 

voles in North America because only 72 had been captured prior to 1974. 

He said that white-footed voles feed on deciduous leaves and forbs. He 

concluded that white-footed voles nest and forage on the forest floor, but 

also forage in the forest canopy. 

280.  Maser, C. 2004. Declaration of Chris Maser. Klamath Siskiyou Wildlands 

Center, Umpqua Watersheds, and Cascadia Wildlands Project v. Lynda L. 

Boody, Dwight Fielder, and Bureau of Land Management. U.S. District 

Court for the District of Oregon. Civ. No. 03-3124-CO.

In this court deposition, Maser gives a passionate defense for the protection of old-growth forests as habitat for tree voles.

281.  Maser, C.; Franklin, J.F. 1974. Checklist of vertebrate animals of the 

Cascade Head Experimental Forest. Resour. Bull. PNW-51. Portland, OR: 

U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest 

and Range Experiment Station. 32 p.

This checklist was based on 3 months during which Chris and Rita Maser 

conducted field work at Cascade Head Experimental Forest in Tillamook 

County, Oregon, in winter 1971–1972. Maser called the white-footed vole 

the rarest member of the vole family in North America. He captured whitefooted voles primarily in riparian alder forests along small streams. Maser 

called the red tree vole the most highly specialized vole in the world. He 

speculated that tree vole populations were widely scattered and disappearing as a result of land development. He reported that tree vole nests were 

found from 2 to 47 m above ground in the live crowns of Douglas-fir, Sitka 

spruce, western hemlock, and grand fir. He stated that tree voles fed on 
90

GENERAL TECHNICAL REPORT PNW-GTR-909

conifer needles and tender bark removed from conifer twigs. He stated that 

known predators were the northern spotted owl, great horned owl (Bubo 

virginianus), long-eared owl (Asio otus), and northern saw-whet owl.

282.  Maser, C.; Johnson, M.L. 1967. Notes on the white-footed vole 

(Phenacomys albipes). Murrelet. 48: 24–27.

Maser and Johnson described morphology, distribution, habitat, and diet 

of the white-footed vole based on their field work and examination of 

museum specimens. They described the white-footed vole as blunt-nosed, 

with white feet contrasting with the overall brown pelage. They presented 

all known white-footed vole locations in the text and map figure. They 

described finding herbaceous plant material in stomachs of white-footed 

voles. One white-footed vole was captured underground in a trap set for 

mountain beaver (Aplodontia rufa). 

283.  Maser, C.; Mate, B.R.; Franklin, J.F.; Dyrness, C.T. 1981. Natural history of Oregon coast mammals. Gen. Tech. Rep. PNW-133. Portland, OR: 

U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest 

and Range Experiment Station. 496 p.

Much of the information on tree voles and white-footed voles in this book 

repeated information previously presented in Maser’s master’s thesis. 

The authors suggested that “Since red tree voles seldom inhabit trees less 

than 25 years of age, clearcut logging decimates entire populations and is 

responsible for the disappearance of tree vole populations in many areas, 

and in large measure, for their widely scattered present distribution.” 

They cited Johnson (1968) in stating that there were two subspecies of red 

tree voles in Oregon (A. longicaudus longicaudus and A. l. silvicola), and 

speculated that tree voles and white-footed voles evolved in the ancient 

Klamath Mountains of southwest Oregon. They quoted Johnson (1973: 

240), who said that “Geologic history provides data for an attractive 

hypothesis; Arborimus evolved in this region from some primitive microtine during the Pliocene, adapting to a habitat of mixed deciduous and 

coniferous forest during a time of moderate temperature and high rainfall; 

consistent climatic conditions along the Pacific Coast have maintained the 

restricted habitat.”

284.  Maser, C.; Storm, R.M. 1970. A key to Microtinae of the Pacific 

Northwest (Oregon, Washington, Idaho). Corvallis, OR: Oregon State 

University Book Stores. 162 p.
91

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

This excellent (but out-of-print) key included detailed drawings by James 

Cloutier of the skulls of all vole species that occur in Oregon and Washington, plus species accounts and distribution maps for each species.

285.  Mayer, W.V. 1952. The hair of California mammals with keys to the dorsal guard hairs of California mammals. American Midland Naturalist. 48: 

480–512. 

Mayer reported the maximum length of guard hairs for red tree voles and 

white-footed voles. He concluded that, “The most difficult genera to distinguish from one another, due to overlapping measurements in length, width, 

and medullary width, are the Phenacomys [= Arborimus], Clethrionomys 

[= Myodes], and Microtus.”

286.  Mazurek, M.J.; Zielinski, W.J. 2003. The importance of the individual 

legacy old-growth tree in the maintenance of biodiversity in commercial 

redwood forests: final report April 29, 2003. Arcata, CA: U.S. Department 

of Agriculture, Forest Service, Pacific Southwest Research Station. 40 p.

http://www.fs.fed.us/psw/publications/4251/mazurek1.pdf. (August 2014).

Mazurek and Zielinski reported that they detected a Sonoma tree vole 

at a remote camera station in Mendocino County, California, in 2002. A 

Trailmaster TM550 infrared Trail Monitor 35mm camera located at the 

base of a tree was triggered by motion and heat sensors.

287.  Mazurek, M.J.; Zielinski, W.J. 2004. Individual legacy trees influence 

vertebrate wildlife diversity in commercial forests. Forest Ecology and 

Management. 193: 321–334.

This was the published version of their final report. They set cameras at 

the base of paired legacy trees and nearby control trees in young forest. 

They detected an adult Sonoma tree vole at one of the legacy trees.

288.  McComb, W.C.; Chambers, C.L.; Newton, M. 1993a. Small mammal 

and amphibian communities and habitat associations in red alder stands, 

central Oregon Coast Range. Northwest Science. 67: 181–188.

The authors used pitfall traps to sample small mammals in three stands 

of 40- to 50-year-old red alder with scattered residual Douglas-fir and 

western redcedar (Thuja plicata) in Benton, Lincoln, and Coos Counties 

in Oregon. They did not give exact numbers but said that they captured 

<10 tree voles in 28,800 trap-nights. All tree voles were captured in upland 

stands of pole-size conifers.
92

GENERAL TECHNICAL REPORT PNW-GTR-909

289.  McComb, W.C.; McGarigal, K.; Anthony, R.G. 1993b. Small mammal 

and amphibian abundance in streamside and upslope habitats of mature 

Douglas-fir stands, western Oregon. Northwest Science. 67: 7–15.

In 1988, the authors used pitfall traps and snap-traps to sample six mature 

stands in Drift Creek, Lincoln County, Oregon. In 31,200 trap-nights, they 

captured 2,562 mammals and 288 amphibians. They captured two red tree 

voles (0.006 per 100 trap-nights) and 11 white footed voles (0.035 per 100 

trap-nights). Of the white-footed voles, 10 were captured along streams and 

1 was captured in upslope habitat.

290.  McLellan, J.E. 1894. Unpublished field notes. On file with: Smithsonian 

Institution, Washington, DC, Record Unit 7176 Smithsonian Institution 

Manuscript Collection Fish and Wildlife; U.S. Department of the Interior, 

Field Reports, 1860 (1887–1961), Series 1. Field reports submitted by 

naturalists conducting field research in the United States, Oregon. Box 87. 

Folder 2.

McLellan reported collecting several tree vole specimens and their nests 

from “cat spruce” that were growing along rivers in tidal marsh near Gardiner, Douglas County, Oregon, in 1894. The specimens were captured in 

traps that were set on limbs beside nests. We do not think these were tree 

voles for two reasons. First, McLellan says that the nests were constructed 

mostly of moss and marsh grass, and second, we did not find any tree vole 

specimens listed in McLellan’s collecting catalog.

291.  McNab, B.K. 1980. Food habits, energetics, and the population biology of 

mammals. American Naturalist. 116: 106–124.

McNab hypothesized that tree vole energetics may differ from other voles 

because tree voles have small litters, slow development of young, and a 

long gestation period.

292.  McNab, B.K. 1992. The comparative energetics of rigid endothermy: the 

Arvicolidae. Journal of Zoology, London. 227: 585–606.

McNab examined body temperature and basal metabolic rates in 24 species of arvicolid rodents, including six Sonoma tree voles. The tree voles 

eventually died because he could not obtain adequate food for them in 

Florida. He concluded that there was “...little variation in the precision or 

level at which body temperature of arvicolid rodents is regulated.” Body 

temperatures for most arvicolids ranged between 37.4 and 38.3 °C, and 
93

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

there was no evidence that any species ever went into torpor. He speculated 

that the high metabolic rate of arvicolids may allow them to breed throughout the year and produce large litters. He concluded that arvicolid rodents 

are “specialists in maximizing production, which permits them to live in 

generally harsh environments characterized by cool to cold temperatures.” 

293.  Meiselman, N. 1992. Nest-site characteristics of red tree voles in 

Douglas-fir forests of northern California. Arcata, CA: Humboldt State 

University. 60 p. M.S. thesis. 

Meiselman located 79 tree vole nests in 9 stands near Branscomb, Mendocino County, California. The majority of nests (57 percent) were in 

mesic old-growth/mature stands. She found that nests tended to be located 

on the south side of the tree on south-facing slopes. She suggested that tree 

voles may select southern exposures to take advantage of solar warming 

in winter. She discussed the error in calling this species “colonial” in a 

concise and succinct manner. She stated that nests were significantly more 

abundant in old-growth stands than mature or young stands. She also 

suggested that tree vole nests were more difficult to detect in old-growth 

forests and that this could bias comparisons among forest age classes. 

294.  Meiselman, N.; Doyle, A.T. 1996. Habitat and microhabitat use by the red 

tree vole (Phenacomys longicaudus). American Midland Naturalist. 135: 

33–42.

This is the published version of Meiselman’s thesis. She conducted surveys 

for tree vole nests in 100- by 1670-m strip transects in three forest age 

classes (<100 years, 100 to 200 years, >200 years) with three replicates in 

each category. Of 149 potential nests detected on transects, 79 (53 percent) 

were tree vole nests. Of the 79 tree vole nests, 39 (49 percent) were in 

>200-year-old stands, 22 (28 percent) were in 100- to 200-year-old stands, 

and 18 (23 percent) were in stands that were <100 years old.

295.  Merriam, C.H. 1889. Description of a new genus (Phenacomys) and four 

new species of Arvicolinae. North American Fauna. 2: 27–53.

Merriam described four new species, including Phenacomys intermedius, 

P. celatus, P. latimanus, and P. ungava. He noted that the molars of these 

voles were rooted, with two divergent roots. This was one year before True 

(1890) published the first description of Phenacomys longicaudus.

296.  Merriam, C.H. 1901. Two new rodents from northwestern California. 

Proceedings of the Biological Society of Washington. 14: 125–126.
94

GENERAL TECHNICAL REPORT PNW-GTR-909

Merriam described the type specimen of the white-footed vole that was 

collected on 24 May 1899 by Walter Fisher near Arcata, Humboldt County, 

California. He compared it with the two red tree vole specimens collected 

by Aurelius Todd in Oregon. He said that the white-footed vole was very 

different when compared to the red tree vole. Specifically, the tail was 

more slender and scantily furred, the skull was narrower and longer, the 

zygomata were narrower and less spreading, the braincase was narrower 

and longer, the rostrum and nasals were longer, the bullae were larger, 

and the fossa was longer and squared-off at the anterior end as opposed to 

rounded or angular.

297.  Meslow, E.C.; Maser, C.; Verner, J. 1981. Old-growth forests as wildlife habitat. Transactions of the 46th North American Wildlife and Natural 

Resources Conference. 46: 329–335. 

The authors listed the red tree vole as a species that finds its optimal 

habitat in old-growth forests, but no data were presented.

298.  Michel, A.K.; Winter, S. 2009. Tree microhabitat structures as indicators 

of biodiversity in Douglas-fir forests of different stand ages and management histories in the Pacific Northwest, U.S.A. Forest Ecology and 

Management. 257: 1453–1464.

Michel and Winter suggested that dense, misshapen branches of Douglasfir provided good cover and support for tree vole nests.

299.  Millar, J.S. 1977. Adaptive features of mammalian reproduction. 

Evolution. 31: 370–386. 

Based on data obtained from Hamilton (1962), Millar concluded that 

juvenile red tree voles were 65 percent of adult size at weaning, which is 

relatively large for an arvicoline. 

300.  Millar, J.S. 1981. Pre-partum reproductive characteristics of eutherian 

mammals. Evolution. 35: 1149–1163.

In appendix 1, Millar listed reproductive characteristics of tree voles, as 

follows: adult weight = 0.023 kg, birth weight = 0.0024 kg, gestation period 

= 27 days, fetal growth rate = 0.008/day, and litter size = 2.5. He cited his 

previous paper (Millar 1977) as the source for this information, although 

the data actually came from Hamilton (1962).

301.  Miller, A.H. 1933. The red tree-mouse preyed upon by the spotted owl. 

Journal of Mammalogy. 14: 162.
95

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Miller found a tree vole skull in the stomach of a female northern spotted 

owl collected along the Eel River, 16.1 km north of Garberville, Mendocino County, California, on 26 May 1932. He suggested that spotted 

owls might be an important predator on tree voles, a fact that has since 

been documented in many different studies.

302.  Miller, A.H. 1942. Unpublished field notes. On file with: University of 

California, Berkeley, Museum of Vertebrate Zoology. 

Miller said that he captured a white-footed vole on the bank of a stream 

on the Dollar Ranch on Maple Creek, 1.6 km north of the confluence with 

the Mad River, Humboldt County, California, on 9 September 1942. He 

and Ward Russell also visited the Wilder Ranch near Carlotta, Humboldt 

County, California. At the time of their visit on 17 September 1942, Harry 

Wilder had been dead for 7 years, but they talked with Mrs. Wilder and 

examined Wilder’s collection of  ≥300 specimens. According to Miller, 

Mrs. Wilder expressed interested in giving the specimens to a museum. 

We located 52 Sonoma tree vole specimens that Wilder collected from 

1913 to 1927 in the collections of 5 museums.

303.  Miller, G.S., Jr. 1896. Genera and subgenera of voles and lemmings. 

North American Fauna. 12: 1–84.

Miller revised the genera and subgenera of voles and lemmings based on 

morphological differences in the skull and teeth. He reported that Phenacomys voles differed from other voles in that the teeth were rooted and the 

reentrant angles on the lingual side of the tooth were much deeper than 

on the labial side. He noted no pronounced external differences between 

Phenacomys spp. and other voles, except that the tail of Phenacomys 

longicaudus was proportionally longer than in other species.

304.  Miller, G.S., Jr. 1897. Synopsis of the voles of the genus Phenacomys. 

Proceedings of the Biological Society of Washington. 11: 77–87.

Miller presented a key to the species of Phenacomys and described the status, characteristics, and distribution of six forms of Phenacomys, including 

the tree vole. He stated that nothing in the external appearance distinguished 

Phenacomys from small species of Microtus except for the long tail and 

arboreal behavior of the tree vole. He described the type specimen of the 

tree vole and also mentioned an isabelline specimen that was collected on 13 

April 1891 at Meadows, Lane County, Oregon. The latter specimen was pale 

yellowish drab on the head, back, and sides, with the venter grayish white. 

This specimen is at the U.S. National Museum (USNM030649).
96

GENERAL TECHNICAL REPORT PNW-GTR-909

305.  Miller, G.S., Jr. 1912. List of North American land mammals in the 

United States National Museum, 1911. U.S. National Museum Bulletin No. 

79. 455 p.

Miller listed 11 species of Phenacomys, including the red tree vole and 

white-footed vole.

306.  Miller, G.S., Jr. 1924. List of North American recent mammals, 1923. 

U.S. National Museum Bulletin No. 128. 673 p.

Miller listed 12 species of Phenacomys, including the red tree vole, whitefooted vole, and dusky tree vole. He shortened the species name from 

Phenacomys silvicolus (Howell) to P. silvicola, but did not say why he 

made this change.

307.  Miller, G.S., Jr.; Kellogg, R. 1955. List of North American recent mammals. U.S. National Museum Bulletin No. 205. 954 p.

Miller and Kellogg listed the species of Phenacomys in North America, 

including the red tree vole, white-footed vole, and dusky tree vole.

308.  Miller, G.S., Jr.; Rehn, J.A.G. 1902. Systematic results of the study of 

North American land mammals to the close of the year 1900. Proceedings 

of the Boston Society of Natural History. 30: 1–352.

Miller and Rehn listed nine forms of Phenacomys from North America, 

including the red tree vole and white-footed vole. They noted that two species of Phenacomys were added to the list since their 1901 paper.

309.  Miller, M.P.; Bellinger, M.R.; Forsman, E.D.; Haig, S.M. 2006. 

Effects of historical climate change, habitat connectivity, and vicariance 

on genetic structure and diversity across the range of the red tree vole 

(Phenacomys longicaudus) in the Pacific Northwestern United States. 

Molecular Ecology. 15: 145–159.

Based on an analysis of mitochondrial control region sequences in red tree 

voles, the authors concluded that there were two haplotype groups corresponding to the northern and southern regions of the range. Spatial genetic 

analyses demonstrated a primary genetic discontinuity separating northern 

and southern sampling areas, while a secondary discontinuity separated 

northern sampling areas into eastern and western groups on opposite sides 

of the Willamette Valley.

310.   Miller, M.P.; Forsman, E.D.; Swingle, J.K.; Miller, S.A.; Haig, 

S.M. 2010. Size-associated morphological variation in the red tree vole 

(Arborimus longicaudus). Northwestern Naturalist. 91: 63–73.
97

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

The authors analyzed patterns of size-associated morphological variation 

of red tree voles. They concluded that there were subtle, but significant, 

morphological differences across the boundary that had been proposed to 

delimit the ranges of the dusky tree vole and red tree vole. However, the 

morphological characters examined had virtually no diagnostic utility for 

distinguishing between the two putative subspecies. 

311.   Miller, M.P.; Haig, S.M. 2010. Identifying shared genetic structure pattern among Pacific Northwest forest taxa: insights from use of visualization tools and computer simulation. PLoS ONE 5:e13683. 10 p.

Miller and Haig found strong spatial genetic structure in red tree voles in 

Oregon, with genetic distances greater in the north. They suggested that 

habitat fragmentation during the last glacial maximum constrained dispersal, resulting in greater genetic diversity in the north than the glacial free 

south.

312.  Molina, R.; Marcot, B.G.; Lesher, R. 2006. Protecting rare, old-growth, 

forest-associated species under the Survey and Manage Program guidelines of the Northwest Forest Plan. Conservation Biology. 20: 306–318.

The authors briefly mentioned the red tree vole in their discussion of the 

Survey and Manage Program, which was adopted as part of the Northwest 

Forest Plan. 

313.  Murie, O.J. 1954. A field guide to animal tracks. Boston, MA: Houghton 

Mifflin Company. 374 p.

On pages 221–225, Murie summarized information about nests, feeding 

behavior, and distribution of red tree voles. He also told a story about 

climbing to what he thought was a hawk’s nest near Forest Grove, Washington County, Oregon, only to find that it was filled with shredded fir 

needles. He did not realize it was a tree vole nest until years later. He 

suggested that tree voles may consume young terminal needles without 

removing the resin ducts.

314.   Murray, M.A. 1993. Unpublished field notes. On file with: Humboldt 

State University, Vertebrate Museum, Arcata, CA.

These notes described Murray’s field trips to collect tree voles used in the 

genetic analysis in his master’s thesis. He collected 12 Sonoma tree voles in 

Humboldt, Mendocino, and Sonoma Counties, California. He also collected 

nine red tree voles, including five in Del Norte County, California and four 

in Lincoln County, Oregon. The four tree voles in Oregon were collected 
98

GENERAL TECHNICAL REPORT PNW-GTR-909

with the help of John Hayes at Oregon State University, at the same location 

where Hayes (1995) later captured two melanistic red tree voles.

315.  Murray, M.A. 1995. Biochemical systematics of the genus Arborimus. 

Arcata, CA: Humboldt State University. 46 p. M.A. thesis. 

Based on a comparison of restriction enzyme sites of mtDNA, Murray 

concluded that there were two main groups of tree voles in California, red 

tree voles north of the Klamath River and Sonoma tree voles south of the 

Klamath River.

316.   Musser, G.G.; Carleton, M.D. 1993. Family Muridae. In: Wilson, D.E.; 

Reeder, D.M., eds. Mammal species of the world: a taxonomic and geographic reference. Washington, DC: Smithsonian Institute Press. 1207 p. 

Musser and Carleton assigned the red tree vole, Sonoma tree vole, and 

white-footed vole to the genus Arborimus instead of Phenacomys. They 

considered the dusky tree vole to be synonymous with the red tree vole 

(pages 504–505, 533).

317.   Nawa, R.K. 2006. Re: Draft supplemental of the 2004 Final SEIS to 

remove or modify the survey and manage mitigation measure standards and guidelines. 3 p. Unpublished document. On file with: Siskiyou 

Project, 950 SW 6th Steet, Grants Pass, OR 97526. (Available upon request 

from the compilers).

In this critique of the survey and manage guidelines, Richard Nawa, staff 

ecologist for the Siskiyou Project, argued that line transect surveys were 

grossly inadequate for surveys of red tree voles because many nests went 

undetected.

318.   Nelson, D.W.; Thompson, D.R.; Paez, R.A. 2006. Klamath Siskiyou 

Wildlands Center; Umpqua Watersheds; Cascadia Wildlands Project v. 

Lynda Boody, in her official capacity as Glendale Field Manager and 

Bureau of Land Management, an agency of the U.S. Department of 

the Interior; Katrina Symons, in her official capacity as Glendale Field 

Manager; William Haigh, in his official capacity as South River Field 

Office Manager on the Roseburg District, BLM; D.R. Johnson Lumber 

Company. No. 06-35214. D.C. No CV-03-03124-JPC opinion. U.S. Court 

of Appeals for the Ninth Circuit, San Francisco, CA.

In this ruling, the Ninth Circuit Court reversed an earlier district court 

judgment and placed an injunction on the Cowcatcher and Cottonsnake 

timber sales on the Roseburg and Medford Districts of the Bureau of Land 

Management. The court ruled that the U.S. Forest Service and Bureau of 
99

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Land Management violated federal law in downgrading the survey and 

manage classification for the red tree vole.

319.   Nelson, E.W. 1918. Smaller mammals of North America. National 

Geographic Magazine. 33: 409–411.

In this popular article, Nelson apparently got most of his information on 

tree voles from Walter Taylor and Harry Wilder. He described the “tree 

mouse” as the most aberrant member of the genus Phenacomys. He briefly 

described the range, habitat, nests, diet, nocturnal habits, and litter size of 

tree voles.

320.  Nowak, R.M.; Paradiso, J.L. 1983. Rodentia: Muridae: genus 

Phenacomys Merriam 1889. In: Walker’s mammals of the world. 4th ed. 

Baltimore, MD: John Hopkins University Press: 660–661.

Nowak and Paradiso provided a brief summary of the taxonomy, habitat, 

nests, and life history of the red tree vole. Like others before them, they 

said that tree voles were slow and cautious climbers (but see Forsman et 

al. 2009). They repeated previous speculation that male tree voles lived 

primarily in burrows or debris piles, but presented no data to support this 

contention.

321.   Nussbaum, R.A.; Maser, C. 1975. Food habits of the bobcat, Lynx rufus, 

in the Coast and Cascade Ranges of western Oregon in relation to present 

management policies. Northwest Science. 49: 261–266.

Nussbaum and Maser found remains of one Arborimus spp. among 227 prey 

items in 143 scats collected from bobcats (Lynx rufus) in the Oregon Coast 

Ranges from 1970 to 1972. They were unable to determine the species.

322.  Olson, D.H.; Burnett, K.M. 2009. Design and management of linkage 

areas across headwater drainages to conserve biodiversity in forest ecosystems. Forest Ecology and Management. 258S: S117–S126.

Olson and Burnett speculated that habitat contiguity may be essential for 

red tree voles.

323.  Olson, D.H.; Van Norman, K.J.; Huff, R.D. 2007. The utility of strategic surveys for rare and little-known species under the Northwest Forest 

Plan. Gen. Tech. Rep. PNW-GTR-708. Portland, OR: U.S. Department of 

Agriculture, Forest Service, Pacific Northwest Research Station. 48 p. 

In this review, the authors reported that strategic surveys of red tree voles 

were expensive and that protection of red tree voles made it difficult to 

conduct timber sales.
100

GENERAL TECHNICAL REPORT PNW-GTR-909

324.  Olterman, J.H. 1972. Rare, endangered, and recently extirpated mammals in Oregon. Corvallis, OR: Oregon State University. 134 p. M.S. 

thesis. 

Olterman reviewed the literature and examined specimens deposited in 

museums and personal collections. He interviewed mammalogists, game 

biologists, naturalists, trappers, and outdoorsmen to ascertain current distribution and abundance of mammals. He classified the white-footed vole 

as rare based on the small number of specimens that he found in museums 

(n = 46), but conceded that they could be more common than indicated by 

specimens. Olterman concluded that the red tree vole was neither rare nor 

endangered because he located 233 specimens, most of which were collected by Murray Johnson and Chris Maser.

325.  Olterman, J.H.; Verts, B.J. 1972a. Endangered plants and animals of 

Oregon. IV. Mammals. Special Report 364. Corvallis, OR: Oregon State 

University, Agricultural Experiment Station. 47 p. 

Olterman and Verts presented a range map based exclusively on red 

tree vole museum specimens and concluded that the species occupied a 

large range in western Oregon. They did not consider tree voles rare or 

endangered. Based on the few specimens in museums, they concluded that 

white-footed voles were rare.

326.  Olterman, J.H.; Verts, B.J. 1972b. Collections containing mammals 

from Oregon. Special Report 362. Corvallis, OR: Oregon State University, 

Agricultural Experiment Station. 28 p.

Olterman and Verts found 284 red tree vole specimens in various collections and provided a detailed list of where those specimens were housed. 

This list is now out of date because many specimens have been collected 

since it was published.

327.   Oregon Department of Fish and Wildlife. 2008. Oregon Department of 

Fish and Wildlife sensitive species: frequently asked questions and sensitive species list. http://www.dfw.state.or.us/wildlife/diversity/species/docs/

SSL_by_category.pdf. (August 2014).

The Oregon Department of Fish and Wildlife listed the red tree vole as 

“vulnerable” in the Oregon Coast Ranges. They defined vulnerable as 

facing one or more threats to the population or habitat. Species on this list 

are not currently threatened with extirpation but could become so with 

continued or increased threats to their population or habitat.
101

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

328.  Oregon Department of Forestry. 2009. The influence of modeled management scenarios on habitat for species of concern. 57 p. http://www.

oregon.gov/ODF/BOARD/docs/June_3_2009/2_Att_1.pdf. (August 2014).

The Oregon Board of Forestry directed the Oregon Department of Forestry 

to identify specific strategies to maintain, enhance, and restore fish and 

wildlife habitat for species of concern on the Clatsop and Tillamook State 

Forests, while at the same time meeting timber harvest targets. In response 

to this request, the Oregon Department of Forestry modeled forest trends 

on the state forests and concluded that the rarity of complex late-successional forest habitat was a limiting factor for red tree voles and that young 

forest was a barrier to dispersal of tree voles. 

329.   Oregon Natural Heritage Information Center. 2004. Rare, threatened 

and endangered species of Oregon. Corvallis, OR: Institute of Natural 

Resources, Oregon State University. 105 p.

The Oregon Natural Heritage Information Center (ORNHIC) compared 

species status listings by different agencies in Oregon. The U.S. Fish and 

Wildlife Service listed the red tree vole, dusky tree vole, and white-footed 

vole as species of special concern. The Oregon Department of Fish and 

Wildlife listed the white-footed vole as a sensitive species. The ORNHIC 

listed the dusky tree vole as threatened throughout its range and the whitefooted vole and red tree vole as species of conservation concern.

330.  O’Neil, T.A.; Steidl, R.J.; Edge, W.D.; Csuti, B. 1995. Using wildlife 

communities to improve vegetation classification for conserving biodiversity. Conservation Biology. 9: 1482–1491.

The authors used quantitative methods to classify vegetation types into 

habitats for 420 wildlife species, including the red tree vole.

331.   Owings, D.H. 1972. Antipredator behavior of arboreal and terrestrial 

rodents. Seattle, WA: University of Washington. 82 p. Ph.D. dissertation. 

Owings used laboratory experiments to compare predator avoidance 

behavior of red tree voles and gray-tailed voles (Microtus canicaudus). 

His predator avoidance hypothesis was that arboreal rodents would rely 

on quickness and agility to escape while terrestrial rodents would freeze 

unless attacked. He also hypothesized that terrestrial rodents would 

become more aggressive when attacked. He found no difference in freezing behavior. Gray-tailed voles were more aggressive toward attackers than 

tree voles.
102

GENERAL TECHNICAL REPORT PNW-GTR-909

332.  Pacific Lumber Company. 1999. Habitat conservation plan for the properties of The Pacific Lumber Company, Scotia Pacific Holding Company, 

and Salmon Creek Corporation. 138 p. http://www.fws.gov/arcata/es/birds/

NSO/documents/Pacific_Lumber_Co_(Humboldt_Redwood_Co.)_1999_

final_HCP.pdf. (August 2014).

In their habitat conservation plan, the Pacific Lumber Company (currently 

the Humboldt Redwood Company) proposed to provide habitat for tree 

voles on their lands by managing at least 10 percent of each watershed as 

late-seral forest.

333.  Pampush, A.; Hopkins, S. 2004. Dusky tree vole distribution on federal 

forestlands within the northern Oregon Coast Range: Interagency Special 

Status Sensitive Species Program FY2005 Inventory and Conservation 

Planning. Salem, OR: U.S. Department of the Interior, Bureau of Land 

Management, Salem District. 4 p.

This strategic survey on the Salem District of the Bureau of Land Management was completed, but we are unaware of any report that followed. 

Contractors conducted targeted surveys for red tree vole nests to add 

information on distribution and abundance. Bureau of Land Management 

staff selected survey sites, but methods were not described in the proposal.

334.  Parmer, R. [N.d.] California’s treetop dwelling rodent: the red tree 

vole. Outdoor California. http://www.dfg.ca.gov/hcpb/species/sss /

sscmamml/m_a15_18.pdf. 4 p. (April 2010).

This leaflet included information on capture methods, distribution, and 

behavior of Sonoma tree voles.

335.  Pearson, O.P. 1949. Unpublished field notes. On file with: University of 

California, Berkeley, Museum of Vertebrate Zoology.

In these field notes, Pearson said that he talked with a “tough hombre” who 

recognized tree vole nests around his place near Coyote Peak, west of the 

Hoopa Valley Indian Reservation, Humboldt County, California. In our 

tree vole specimen database, there are two Sonoma tree voles that were 

collected by Pearson. One was at French Camp on 23 August 1949 near 

Coyote Peak, Humboldt County, California. The other was 11.3 km north 

of Fort Ross, Sonoma County, California, on 20 May 1949.

336.  Pechman, M.J. 2005. Northwest Ecosystem Alliance, et al. v. Mark E. 

Rey, et al. Order granting part and in denying part plaintiffs’ motion and 

denying defendants’ motion. 1 August 2005. Case No. C04-844P. U.S. 

District Court, Western District of Washington, Seattle, WA.
103

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

In her summary judgment, Judge Pechman ruled that the U.S. Forest Service and Bureau of Land Management failed to comply with the National 

Environmental Policy Act in preparing their 2004 final environmental 

impact statement to remove or modify the Northwest Forest Plan’s survey 

and manage measure. Pechman also ruled that the federal agencies failed 

to analyze the impacts of moving survey and manage species (including 

the red tree vole) to the sensitive species status program. She also ruled 

that the agencies failed to thoroughly analyze their assumption that latesuccessional reserves would protect survey and manage species.

337.   Pechman, M.J. 2006a. Northwest Ecosystem Alliance, et al. v. Mark E. 

Rey, et al. Order on plaintiffs’ motion for injunctive relief. 9 January 2005. 

Case No. 04-844P. U.S. District Court, Western District of Washington, 

Seattle, WA.

In this ruling, Judge Pechman set aside the 2004 Record of Decision to 

eliminate the survey and manage measure from the Northwest Forest Plan. 

She ordered the U.S. Forest Service and Bureau of Land Management to 

reinstate the 2001 Record of Decision for the amendment to the survey and 

measure guidelines.

338.  Pechman, M.J. 2006b. Northwest Ecosystem Alliance, et al. v. Mark E. 

Rey, et al. Stipulation and (proposed) order RE: injunction. 11 October 

2006. Case No. C04-844-P. U. S. District Court, Western District of 

Washington, Seattle, WA. 3 p.

In this court order, Judge Pechman ruled that federal timber sales or habitat disturbing activities that were proposed in the 2004 Record of Decision 

could only go forward if they were in compliance with the 2001 Record 

of Decision. This decision meant that federal agencies had to continue 

to survey for and manage red tree voles in old forests. However, Judge 

Pechman allowed several exemptions to this decision, one of which was 

that thinning projects in forests <80 years old could be conducted without 

surveying and managing for tree voles. This exemption had large implications for the conservation of red tree voles because it meant that federal 

agencies no longer had to survey or manage for red tree voles in forests 

that were <80 years old.

339.   Pechman, M.J. 2007. Conservation Northwest, et al. v. Mark E. Rey, et 

al. Order denying defendants’ motion for relief pursuant to Fed. R. Civ. 

P. 60(b). Case No. C-04-844-P. U.S. District Court, Western District of 

Washington, Seattle, WA.
104

GENERAL TECHNICAL REPORT PNW-GTR-909

In this court order, Judge Pechman denied a motion in which the U.S. Forest Service and Bureau of Land Management had requested relief from the 

injunction placed on their 2007 Record of Decision (ROD) to remove the 

survey and manage measures. Pechman concluded that the federal agencies 

failed to establish that the 2001 ROD for the amendment to the survey and 

manage standards and guidelines was unworkable, and that they failed to 

show they entered into the 2006 stipulation in good faith or made a reasonable effort to comply with it before modifying the document. She also 

ruled that the federal agencies failed to show that the 2007 ROD complied 

with all appropriate law. The final argument for relief was also denied as 

the court ruled that it did not matter how much money or time the agencies 

spent preparing the 2007 document.

340.  Pettinato, J.S.; Marshall, S.A.; Perron, B. 2013. The Bureau of 

Land Management’s cross-motion for summary judgment. Cascadia 

Wildlands et al. v. The Bureau of Land Management and Seneca Sawmill 

Company. U.S. District Court, District Court, Eugene Division. Case No. 

6:12-cv-1739-AA.

In this cross-motion, the Bureau of Land Management requested a summary judgment to allow the Rickard Creek timber sale to proceed. They 

argued that the administrative record indicated that the timber sale area 

was not a high-priority site for tree voles.

341.   Phillips, C.; Chrostowski, H. 1981. Wildlife habitats and species management relationships program: Oregon Coast Range, volume IV, mammals. Corvallis, OR: U.S. Department of Agriculture, Forest Service, 

Siuslaw National Forest: 106–109.

Phillips and Chrostowski summarized the status, distribution, habitat, 

reproduction, behavior, home range, food habits, and management recommendations for red tree voles and white-footed voles. They stated that tree 

voles require fog condensation on canopy needles as a water source and 

that male tree voles nest primarily, if not exclusively, on the ground. 

342.  Pratt, H.S. 1923. A manual of land and fresh water vertebrate animals of 

the United States (exclusive of birds). Philadelphia, PA: P. Blakiston’s Son 

& Company. 422 p.

On page 320, Pratt briefly described tree voles and white-footed voles.

343.  Price, W.W.; Landon, C.P.; Forsman, E.D. 2010. Aneides ferreus

(clouded salamander): arboreal activity. Herpetological Review. 4: 58–59.
105

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Price and Landon found two adult clouded salamanders 75 m up in a cavity 

in the top of a Douglas-fir that was 78 m tall. The cavity also contained 

copious amounts of red tree vole fecal pellets. The location was in western 

Douglas County, adjacent to the tallest Douglas-fir in Oregon, the 99-m 

Brummit Fir (since renamed “Doerner Fir”).

344.  Pugh, S.R.; Johnson, S.; Tamarin, R.H. 2003. Voles, Microtus species. In: Feldhamer, G.A.; Thompson, B.C.; Chapman, J.A., eds. Wild 

mammals of North America: biology, management, and conservation. 

Baltimore, MD: John Hopkins University Press: 349–370.

In this excellent literature review, the authors described the distribution, 

morphology, genetics, physiology, reproduction, development, ecology, 

feeding habits, behavior, mortality, aging, economic status, and management and conservation of Microtus. They only mentioned the red tree vole 

once when they reported the negative effects of hybridization between 

sibling species of Arborimus. We assume that they were referring to 

Murray Johnson’s hybrids between A. pomo and A. longicaudus. Johnson 

and George (1991: 4) stated that hybrid crosses had a much higher rate 

of pregnancy-related deaths and suggested that this was indicative of a 

reproductive barrier between the two species.

345.  Quay, W.B. 1953. Phenacomys albipes in Lincoln County, Oregon. 

Journal of Mammalogy. 34: 391.

Quay described two white-footed vole specimens collected by Harold 

Broadbooks on 17 June 1942, 1.6 km east of Newport, Lincoln County, 

Oregon. The skulls and skins of these specimens were originally misidentified as Microtus until Quay corrected this misidentification at the University of Michigan Museum of Zoology.

346.  Quay, W.B. 1954a. The meibomian glands of voles and lemmings 

(Microtinae). Miscellaneous Publications, Museum of Zoology, University 

of Michigan, No. 82. 17 p.

Quay compared the number and size of the meibomian (tarsal) glands in 

the eyelids of voles and lemmings, including the red tree vole. He suggested that the reduction in gland number and increase in gland size in 

more advanced fossorial species may be the result of a reduction in eye 

size.

347.   Quay, W.B. 1954b. The anatomy of the diastemal palate in microtine 

rodents. Miscellaneous Publications, Museum of Zoology, University of 

Michigan, No 86. 41 p.
106

GENERAL TECHNICAL REPORT PNW-GTR-909

Quay described the morphology of the diastemal palate in many different 

voles, including red tree voles. The diastemal palate is the region of the 

skull between the incisors and molars.

348.  Quay, W.B. 1965. Comparative survey of the sebaceous and sudoriferous 

glands of the oral lips and angle in rodents. Journal of Mammalogy. 46: 

23–37.

Quay compared the sebaceous and sudoriferous glands of the mouth of 

many different mammals, including red tree voles. Regarding the significance of these differences he said that “Little if anything can be offered as 

definitive evidence of the functions or functional significance of the oral 

sebaceous or sudoriferous glands.”

349.  Quay, W.B. 1968. The specialized posterolateral sebaceous glandular 

regions in microtine rodents. Journal of Mammalogy. 49: 427−445.

Quay compared the development and location of sebaceous glands in many 

different voles, including red tree voles. He discussed taxonomic differences and possible functions of sebaceous glands in communication.

350.  Ralph, C.J.; Paton, P.W.C.; Taylor, C.A. 1991. Habitat association patterns of breeding birds and small mammals in Douglas-fir/hardwood 

stands in northwestern California and southwestern Oregon. In: Ruggiero, 

L.F.; Aubry, K.B.; Carey, A.B.; Huff, M.H., tech. cords. Wildlife and vegetation of unmanaged Douglas-fir forest. Gen. Tech. Rep. PNW-GTR-285. 

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

Northwest Research Station: 379–393.

The authors used pitfall traps, live-traps, and snap-traps to sample 47 

forest stands in southwestern Oregon and northwest California from 1984 

through 1985. They captured 22 tree voles in 141,120 trap-nights (0.016 per 

100 trap-nights). All tree voles were captured in pitfalls. The samples were 

apparently too small to test relationships between capture rates and forest 

age or structure. Near Cave Junction, Oregon, they captured five red tree 

voles in 43,200 trap-nights (0.012 per 100 trap-nights). Near Willow Creek, 

California, they captured nine Sonoma tree voles in 48,960 trap-nights 

(0.018 per 100 trap-nights). Near Branscomb, California, they captured 

eight Sonoma tree voles in 48,960 trap-nights (0.016 per 100 trap-nights). 

The authors did not mention age or sex ratios of tree voles captured in 

pitfalls, but we examined 13 of their museum specimens and found that 

they were all males (5 adults, 7 subadults).
107

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

351.   Raphael, M.G. 1981–1983. Unpublished mammal and amphibian catalog. 

On file with: University of California, Berkeley, Museum of Vertebrate 

Zoology.

In his specimen catalog for 1981–1983, Raphael listed 13 Sonoma tree 

voles (4 males, 9 females) captured in pitfall traps in Humboldt County and 

western Trinity County, California. We found 11 of these specimens at the 

Museum of Vertebrate Zoology.

352.  Raphael, M.G. 1988. Long-term trends in abundance of amphibians, 

reptiles, and mammals in Douglas-fir forests of northwestern California. 

In: Szaro, R.C.; Severson, K.E.; Patton, D.R., tech. cords. Management of 

amphibians, reptiles, and small mammals in North America. Gen. Tech. 

Rep. RM-166. Flagstaff, AZ: U.S. Department of Agriculture, Forest 

Service, Rocky Mountain Research Station: 23–31.

From 1981 through 1983, Raphael used pitfall traps with drift fences to 

sample small mammals and amphibians in six seral stages of forest within 

80 km of Willow Creek, Humboldt County, California. He captured 19 

Sonoma tree voles in 899,431 trap-nights (0.002 tree voles per 100 trapnights). Relative abundance of tree voles did not differ among forest age 

classes, but the power to detect differences among age classes was low 

because of the small number of voles captured. 

353.  Raphael, M.G. 1990. Mammals of the ancient forest. In: Norse, E.A., ed. 

Ancient forests of the Pacific Northwest. Washington, DC: Island Press: 

95–100. 

Raphael’s essay included the red tree vole as an ancient forest species. He 

said that the tree vole was highly specialized because it was the only arboreal mammal in North American that fed exclusively on conifer needles.

354.  Rausch, R.L.; Johnson, M.L. 1983. Occurrence and pathogenicity of 

Heligmosomoides spp. (Nematoda: Heligmosomidae) associated with 

cecal villi in arvicolid rodents. Proceedings of the Helminthological 

Society of Washington. 50: 25–35. 

Rausch and Johnson reported that the cecal villi of the heather vole closely 

resembled the cecal villi of the red tree vole and differed only slightly from 

the cecal villi of the white-footed vole.

355.  Repenning, C.A. 1985. Unpublished letter to Murray Johnson. On file 

with: University of Washington, Burke Museum, Seattle, WA.

In this 15 January 1985 letter to Murray Johnson, Repenning said that he 

did not agree with Johnson that the red tree vole should be in the genus 
108

GENERAL TECHNICAL REPORT PNW-GTR-909

Arborimus instead of Phenacomys. Repenning’s opinion was based on his 

interpretation of paleontological data.

356.  Repenning, C.A. 2001. Beringian climate during intercontinental dispersal: a mouse eye view. Quaternary Science Review. 20: 25–40.

Repenning suggested that microtines evolved so rapidly that they could 

be correlated to specific climatic events in North America. He noted that 

current Phenacomys voles were limited to subarctic and temperate forests, 

and suggested that Phenacomys voles evolved in Beringia from a Eurasian 

ancestor.

357.   Repenning, C.A.; Brouwers, E.M.; Carter, L.D.; Marincovich, L., 

Jr.; Ager, T.A. 1987. The Beringian ancestry of Phenacomys (Rodentia: 

Cricetidae) and the beginning of the modern Arctic Ocean borderland 

biota. U.S. Geological Survey Bulletin. 1687: 1–28.

The authors discussed a recently discovered fossil of Phenacomys intermedius from the late Pliocene (2.5 million years ago) that was discovered 

in northern Alaska. The oldest P. intermedius fossil in the conterminous 

United States was 0.8 million years old and confined more or less to 

Beringia. They cited a personal communication from Walter Dalquest who 

suggested that some P. intermedius were 1.6 million years old based on 

unpublished data. They described the rooted hypsodont teeth of Phenacomys. Though they used Phenacomys instead of Arborimus for the red tree 

and white-footed vole, they said that they did so without opinion. They 

said that there were two pre-Pleistocene Phenacomys (P. deeringensis), one 

from Alaska and one from Siberia. The Alaskan P. deeringensis was more 

primitive and had very low dentine tracts compared to P. deeringensis

from Siberia. The first lower molar of the extinct P. deeringensis was 

essentially identical to living P. intermedius. Repenning also named P. 

gryci located from the northern Alaskan site.

358.  Repenning, C.A.; Grady, F. 1988. The microtine rodents of the Cheetah 

Room Fauna, Hamilton Cave, West Virginia, and the spontaneous origin 

of Synaptomys. U.S. Geological Survey Bulletin. 1853: 1–28.

Repenning and Grady named Paraphenacomys as a new subgenus that they 

said was closely related to Phenacomys and the Arborimus subgenus. They 

included P. deeringensis and P. intermedius (living) in the new subgenus 

while noting three unnamed species. In the subgenus Paraphenacomys, 

Repenning and Grady included P. albipes, P. brachyodus, and P. gryci. 

They said that the only known white-footed vole fossil was an uncataloged 
109

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

first lower molar at the University of Arizona from the 13,000-year-old 

Smith Creek Cave fauna, White Pine, Nevada. P. brachyodus was a new 

species from the Cheetah Room, West Virginia. Unfortunately, the Repenning (in press) paper that was cited several times by the authors was never 

published because it was one of the chapters in the unfinished tree vole 

book that Murray Johnson was working on when he died. 

359.   Reynolds, R.T. 1970. Nest observations of the long-eared owl (Asio otus) 

in Benton County, Oregon, with notes on their food habits. Murrelet. 51: 

8–9.

Reynolds collected 104 regurgitated pellets from a pair of nesting longeared owls (Asio otus) 16.1 km north of Corvallis, Benton County, Oregon. 

The pellets contained 153 prey items, one of which was a red tree vole.

360.  Richards, R.L. 1986. Late Pleistocene remains of boreal voles (Genera 

Phenacomys and Clethrionomys [= Myodes]) from southern Indiana 

Caves. Proceedings of Indiana Academy of Science. 95: 537–546.

Richards suggested that the presence of heather vole fossils indicated that 

southern Indiana was once covered by boreal forests. There was nothing 

specific about tree voles in this paper, but it is pertinent to the evolutionary 

history of the genus Phenacomys.

361.   Richardson, A.D.; Ashton, P.M.S.; Berlyn, G.P.; McGroddy, M.E.; 

Cameron, I.R. 2001. Within-crown foliar plasticity of western hemlocks, Tsuga heterophylla, in relation to stand age. Annals of Botany. 88: 

1007–1015.

In western hemlock, needles in the upper outer crown were thicker and had 

more vascular tissue than needles in the lower inner crown. The authors 

did not report whether resin ducts also varied in size depending on position 

within the crown. There was nothing specific about tree voles in this paper, 

but within-species or within-tree variation in needle structure may play an 

important role in habitat selection and foraging behavior of tree voles.

362.  Rittenhouse, B.; Breckel, R.; Alegria, J.; Max, T.; Edwards, T. 2002. 

Survey protocol for conducting red tree vole strategic surveys on randomly selected CVS/FIA plots. 26 p. Unpublished document. On file with: 

U.S. Department of Agriculture, Forest Service and U.S. Department of 

the Interior, Bureau of Land Management, 1220 SW 3rd Ave., Portland, 

OR 97204. (Available upon request from the compilers). 

This study plan described the methods used to collect data on density of 

red tree vole nests in a stratified random sample of plots in western Oregon 
110

GENERAL TECHNICAL REPORT PNW-GTR-909

and northwestern California (Dunk and Hawley 2009). The primary goal 

was to compare the relative abundance of red tree vole nest trees in different forest age classes and land use categories on federal lands within 

the range of the species. The original plan was to survey a square 1-ha 

plot at each of 400 randomly selected Current Vegetation Survey/Forest 

Inventory Assessment plots. However, an additional 1-ha plot was added 

at each location after the start of the survey to increase the probability 

of detecting nests. The second plot was installed on the north side of the 

first plot. Within each plot, contractors visually searched for nests in trees 

while walking along linear transects that were spaced 25 m apart. Potential 

nests seen from the ground were climbed to determine if they were vole 

nests and to estimate their occupancy status. In plots where no tree vole 

nests were detected during ground surveys or where the forest canopy was 

so complex that nests were hard to detect, contract climbers climbed five 

randomly selected large trees (>61 cm diameter at breast height) to see if 

they could find nests. When they analyzed the data from this study, Dunk 

and Hawley (2009) only used data from the original sample of 1-ha plots 

because they were concerned that the second plot at each location was 

not selected randomly and might be in a different cover type or land use 

allocation than the first plot.

363.  Robovský, J.; Řičánková, V.; Zrzavý, J. 2008. Phylogeny of Arvicolinae 

(Mammalia, Cricetidae): utility of morphological and molecular data sets 

in a recently radiating clade. Zoologica Scripta. 37: 571–590.

This phylogenetic analysis indicated that Dicrostonyx, Phenacomys, and 

Arborimus formed a monophyletic clade. The authors proposed this clade 

be given a new tribal classification name, Dicrostonychini Kretzoi 1955.

364.  Roest, A.I. 1949. Mammals of the Oregon Caves area. Corvallis, OR: 

Oregon State University. 95 p. M.S. thesis.

Roest described a juvenile red tree vole that was collected in the “guide 

shack” at Oregon Caves National Monument in August 1948. This specimen is in the Oregon State University Fisheries and Wildlife collection 

(OSUFW844). Roest did not say how the vole was captured.

365.  Roest, A.I. 1951. Mammals of the Oregon Caves area, Josephine County. 

Journal of Mammalogy. 32: 345–351.

In this published version of his 1949 thesis, Roest said that the juvenile red 

tree vole that was captured in the guide shack at Oregon Caves National 

Monument was killed, but he did not say how. 
111

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

366.  Rosenberg, D.K.; Swindle, K.A.; Anthony, R.G. 2003. Influence of 

prey abundance on northern spotted owl reproductive success in western 

Oregon. Canadian Journal of Zoology. 81: 1715–1725.

The authors analyzed prey remains in regurgitated pellets of northern 

spotted owls on the Blue River and McKenzie Ranger Districts on the west 

slope of the Cascade Range, Lane County, Oregon. Pellets were collected 

from 1987 through 1996. The mean number of spotted owl territories 

where pellets were collected per year was 16.6 (range = 4 to 17). The mean 

± SE percentage of tree voles in the diet was 6.7 ± 1.2 percent based on 

prey numbers, and 3.7 ± 0.8 percent based on prey biomass. The authors 

also used Sherman live-traps to sample small mammals from 1988 through 

1996. They did not report the total number of trap-nights but did report 

using 100 Sherman live-traps each year for 8 consecutive days. They 

captured 1,501 deer mice and 303 red-backed voles but did not report 

capturing any other small mammals.

367.   Ruggiero, L.F.; Jones, L.L.C.; Aubry, K.B. 1991. Plant and animal 

habitat associations in Douglas-fir forests of the Pacific Northwest: an 

overview. In: Ruggiero, L.F.; Aubry, K.B.; Carey, A.B.; Huff, M.H., 

tech. coords. Wildlife and vegetation of unmanaged Douglas-fir forests. Gen. Tech. Rep. PNW-GTR-285. Portland, OR: U.S. Department of 

Agriculture, Forest Service, Pacific Northwest Research Station: 447–462.

The authors listed the red tree vole as a species of special interest or 

concern to forest managers because the species was difficult to study and 

the patterns of abundance or occurrence were poorly understood.

368.  Russell, W.C. 1942. Unpublished field notes. On file with: University of 

California, Berkeley, Museum of Vertebrate Zoology.

In 1942, Russell collected eight Sonoma tree voles by placing snap-traps 

baited with walnuts on top of tree vole nests at Maple Creek east of Arcata, 

Humboldt County, California. He captured four adult females, two adult 

males, and two subadult males. He also collected a northern spotted owl 

at the same location. The owl had four tree vole skulls in its ventriculus. 

Russell also collected two additional Sonoma tree voles in 1968 near 

Kneeland, Humboldt County, California.

369.   Santucci, V.L.; Kenworthy, J.; Kerbo, R. 2001. An inventory of paleontological resources associated with National Park Service caves. Tech. 

Rep. NPS/NRGRD/GRDTR-01/02. Lakewood, CO: U.S. Department of 
112

GENERAL TECHNICAL REPORT PNW-GTR-909

the Interior, National Park Service, Geological Resource Division.

50 p. http://www.nature.nps.gov/geology/paleontology/pub/cavepaleo.pdf. 

(August 2014).

The authors reported that they found fossil remains of Phenacomys/

Arborimus in cave sediments at Oregon Caves National Monument. 

370.  Semken, H.A., Jr.; Wallace, S.C. 2002. Key to arvicoline (“microtine” rodents) and arvicoline-like lower first molars recovered from Late 

Wisconsinan and Holocene archaeological and paleontological sites in 

eastern North America. Journal of Archaeological Science. 29: 23–31.

Semken and Wallace described qualitative and quantitative methods used 

to identify arvicoline molars from archaeological and paleontological sites. 

They included the heather vole in their key but not tree voles or whitefooted voles. They cited Guilday and Parmalee (1972) in support of the 

assertion that the four living Phenacomys vole species cannot be distinguished by dentition alone.

371.   Sharnoff, S.; Rosentreter, R. 1998. Lichen use by wildlife in North 

America. U.S. Department of the Interior, Bureau of Land Management, 

Idaho. http://www.lichen.com/fauna.html. (August 2014).

The authors cited Denison (1973), Zentner (1977), and Gillesberg and 

Carey (1991) as sources indicating that red tree voles used lichens as nest 

material and possibly as food.

372.  Shelton, A.C. 1913a. Unpublished photo. On file with: University of 

California, Berkeley, Museum of Vertebrate Zoology (Image 1134; 

Timber near Lierley’s Ranch, Mendocino County, CA., August 1913). 

http://calphotos.berkeley.edu/cgi/img_query?query_src=MVZ&seq_

num=53797&one=T. (August 2014).

Alfred Shelton and Charles Camp collected a Sonoma tree vole at the 

Lierly Ranch in 1913. This photo provides some context, but it is unclear if 

the area in the photo was near where the vole was collected or not.

373.   Shelton, A.C. 1913b. Unpublished field notes. On file with: University of 

California, Berkeley, Museum of Vertebrate Zoology.

In his field notes for the North Coast Counties Expedition of 1913, Shelton 

concentrated on birds while fellow expedition member Charles Camp 

(1913) dealt with mammals. However, Shelton listed two Sonoma tree vole 

specimens in his field notes, a subadult male collected at Mendocino City, 

and an adult female collected at the Lierly Ranch 12.9 km north of Hearst, 

Mendocino County, California.
113

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

374.   Shelton, A.C. 1914. Unpublished field notes. On file with: University of 

Oregon, Museum of Natural and Cultural History, Eugene, OR.

Shelton’s field notes included several references to red tree voles. In February 1914, he and John Bovard captured three large juveniles 33 m up in a 

“tall fir” at Spencer Butte, south of Eugene, Lane County, Oregon. In June 

1914, he described a specimen that Vernon Bailey chased out of a nest and 

shot off a limb while tree climbing on Spencer Butte (see Bailey 1914, 1915; 

Jobanek 1988). In October 1914, he described a tree vole that eluded capture 

in a grove of large Douglas-fir on the Curry Estate where the North and 

South Forks of the Umpqua River join west of Roseburg, Douglas County, 

Oregon. He also says that he found a cluster of tree vole nests near the community of Mable in the Mohawk Valley, Lane County, Oregon.

375.   Sherrell, P.E. 1969. A new locality record for Phenacomys albipes. 

Murrelet. 50: 39.

While conducting field work for his thesis in 1968, Sherrell captured a 

male white-footed vole at Brush Creek, 1.6 km from the ocean and 12.1 

km south of Port Orford, Curry County, Oregon. He caught the vole in a 

museum special trap that was placed under a thicket of salmonberry in the 

floodplain of Brush Creek.

376.   Sherrell, P.E. 1970. An annotated check list of the land mammals of 

Curry County, Oregon. Ellensburg, WA: Central Washington State 

University. 150 p. M.S. thesis.

Sherrell captured an adult male white-footed vole in a museum special trap 

at Brush Creek (Sherrell 1969). This was only the second white-footed vole 

documented in Curry County. Sherrell did not capture any tree voles, but 

he did review the literature and corresponded with Murray Johnson and 

others to come up with locations for 19 tree voles in Curry County, most of 

which were collected by or given to Stanley Jewett.

377.   Silovsky, G.D.; Pinto, C. 1974. Forest wildlife inventories: identification of conflicts and management needs. In: Black, H.C., ed. Wildlife and 

forest management in the Pacific Northwest: proceedings of a symposium 

held September 11–12, 1973. Corvallis, OR: Oregon State University, 

School of Forestry: 53–61.

Silovsky and Pinto listed the red tree vole as having a moderate dependence on snags in their table on the native wildlife of the Siuslaw National 

Forest. They suggested that tree voles would be adversely affected by forest management methods that were prevalent in 1974 (clearcut and burn).
114

GENERAL TECHNICAL REPORT PNW-GTR-909

378.  Simberloff, D.; Cox, J. 1987. Consequences and costs of conservation 

corridors. Conservation Biology. 1: 63–71.

Simberloff and Cox cited Soulé and Simberloff (1986) to support the 

hypothesis that red tree voles would avoid riparian corridors between 

stands of mature Douglas-fir. They provided no evidence that this was 

actually the case. 

379.   Slauson, K.M.; Higley, J.M.; Matthews, S.M. 2011. Fisher (Martes 

pennanti) conservation on the Hoopa Valley Reservation, California: fallwinter diet of fishers on the Hoopa Valley Reservation. 15 p. Unpublished 

report. On file with: U.S. Department of Agriculture, Forest Service, 

Pacific Southwest Research Station, Arcata, CA. 

The authors found remains of four Sonoma tree voles in 140 scats collected 

from fisher on the Hoopa Valley Reservation, Humboldt County, California, from 2004 through 2008.

380.  Smith, W.P.; Anthony, R.G.; Waters, J.R.; Dodd, N.L.; Zabel, C.J. 

2003. Ecology and conservation of arboreal rodents of western coniferous 

forests. In: Zabel, C.J.; Anthony, R.G., eds. Mammal community dynamics: management and conservation in the coniferous forests of western 

North America. Cambridge, United Kingdom: Cambridge University 

Press: 157–206. Chapter 6.

The authors reviewed the literature on taxonomy, distribution, feeding 

habits, water consumption, reproduction, predators, nests, and habitat of 

red tree voles and Sonoma tree voles. They found no published data on 

movements, estimates of home range size, longevity, or survival of tree 

voles. They also found no published documentation of ecological differences between red tree voles and Sonoma tree voles, and discussed general 

ecological characteristics that they assumed to be common to both species.

381.  Smoluk, A. 2011. Geographic distributions of prey of the northern spotted owl in central western Cascades, Oregon, 1988–2009. Corvallis, OR: 

Oregon State University. 74 p. M.S. thesis. 

Smoluk analyzed the geographic distribution of prey items identified in 

northern spotted owl pellets on the H.J. Andrews Experimental Forest in 

Lane County, Oregon, 1988–2009. Of 4,183 prey items identified in pellets, 

272 (6.5 percent) were red tree voles. Based on their abundance in owl 

diets, Smoluk concluded that tree voles were more abundant at low elevations and in steep canyons. She reported that red tree voles were rare in 

pellets collected at owl territories above 1000 m elevation. 
115

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

382.  Snyder, K.U.; Sullivan, T.J.; Moore, D.L.; Raymond, R.B.; Gilbert, 

E.H. 2003. Trask River watershed analysis: final report, August 2003. 

122 p. Unpublished report submitted to Oregon Department of Forestry 

and U.S. Department of the Interior, Bureau of Land Management. http://

www.oregon.gov/odf/state_forests/docs/watershed/trask_watershed_analysis/101_front_material.pdf. (August 2014).

The authors reported that potential red tree vole habitat was rare in the 

Trask River watershed, except for a few stands of old-growth in the 

Upper Trask River. They stated that red tree voles required large blocks 

of contiguous habitat or connecting corridors of suitable habitat. This may 

be true, but they did not cite any papers or present any data to support this 

statement. They do cite Carey (1991) to support their claim that stands 

<100 years old cannot support viable populations of tree voles, but they do 

not acknowledge that Carey never actually monitored vital rates or population trends of tree vole populations.

383.  Sobb, N. 2006. Re: Addendum to the Anderson West Landscape 

Management Project (EA #OR110-03-17). On file with: U.S. Department 

of the Interior, Bureau of Land Management, Medford District, 3040 

Biddle Road, Medford, OR 97504. 2 p. (Available upon request from the 

compilers).

In this addendum to tree vole survey data that were sent to the Medford 

District of the Bureau of Land Management by the Northwest Ecosystem 

Survey Team, Sobb emphasized that many red tree vole nests were not 

detected during ground-based surveys.

384.  Sondenaa, A.C. 1991. The wild mammals of McDonald and Paul M. 

Dunn Forests. Corvallis, OR: Oregon State University Research Forests. 

29 p.

In this guidebook to mammals of Oregon State University’s McDonald 

Forest, Sondenaa considered the white-footed vole as rare and limited 

primarily to riparian areas. She stated that red tree voles occurred primarily in Douglas-fir forests and built their nests of twigs and discarded resin 

ducts on dense branches.

385.  Soulé, M.E.; Simberloff, D. 1986. What do genetics and ecology tell us 

about the design of nature reserves? Biological Conservation. 35: 19–40.

The authors suggested that tree voles would not use riparian corridors 

between patches of mature Douglas-fir forest. They did not explain why 

they thought this to be the case.
116

GENERAL TECHNICAL REPORT PNW-GTR-909

386.  Spade, A. 2012. Unpublished pictures of a Sonoma tree vole nest in a 

Douglas-fir. Photos On file with: University of California Museum of 

Vertebrate Zoology. http://calphotos.berkeley.edu/cgi/img_query?seq_

num=415710&one=T, http://calphotos.berkeley.edu/cgi/img_query?seq_

num=415712&one=T, and http://calphotos.berkeley.edu/cgi/img_query? 

enlarge=0000+0000+0712+0189. (August 2014).

Spade took these pictures of a Sonoma tree vole nest at Ten Mile River 

north of Fort Bragg, Mendocino County, California in 2012. The forest 

consisted of Douglas-fir and redwood. The pictures included a picture of 

the nest in the tree as well as pictures of nest materials on the ground under 

the nest.

387.   Spickler, J.C.; Sillett, S.C.; Marks, S.B.; Welsh, H.H., Jr. 2006. 

Evidence of a new niche for a North American salamander: Aneides 

vagrans residing in the canopy of old-growth redwood forest. 

Herpetological Conservation and Biology. 1: 16–26.

This paper was mainly about arboreal behavior of wandering salamanders 

(Aneides vagrans), but the authors also reported that they found clouded 

salamanders (A. ferreus) in three nests that were built by western gray 

squirrels (Sciurus griseus) in the Coast Ranges of Oregon. This report was 

confusing because the authors said that they were studying red tree voles 

and that the salamanders were found in nests that contained fecal pellets 

of rodents that had recently left the nests. This left us unsure whether the 

nests occupied by the clouded salamanders were squirrel nests or vole 

nests. The authors also incorrectly listed the species name of the red tree 

vole as Arborimus pomo.

388.  Spies, T.A.; Adams, D.M.; Harmon, M.E.; Johnson, K.N.; Reeves, 

G.H. 2004. Final report to National Commission on Science for 

Sustainable Forestry—Project A5: Assess the scientific basis for standards/practices at the stand management unit, landscape, and region 

level: Oregon Coast Range. 21 p. Unpublished report. On file with: U.S. 

Department of Agriculture, Forest Service, Pacific Northwest Research 

Station, Corvallis, OR. http://ncseonline.org/sites/default/files/A5%20

West%20Final%20report.pdf. (August 2014). 

The authors modeled consequences of forest management policies on 

biological diversity, carbon, and timber production in the Coast Ranges 

of western Oregon. Their habitat model was based on tree diameter, 

percentage of Douglas-fir, canopy closure, and canopy heterogeneity. 
117

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

They concluded that under current management policy, there would be 

an increase in areas dominated by old forest and that red tree vole habitat 

would increase by 25 percent. If thinning of plantations on federal lands 

ceased, their model predicted it would have little effect on the amount of 

red tree vole habitat.

389.  Spies, T.A.; McComb, B.C.; Kennedy, R.S.H.; McGrath, M.T.; Olsen, 

K.; Pabst, R.J. 2007. Potential effects of forest policies on terrestrial biodiversity in a multi-ownership province. Appendix D—Habitat capability 

index for red tree vole (Arborimus longicaudus). Ecological Applications. 

17: 48–65.

The authors modeled potential biodiversity in the Oregon Coast Ranges 

province under the current federal policy (Northwest Forest Plan) and 

two alternative strategies. The red tree vole was one of their focal species. 

Variables used for modeling suitable habitat for tree voles were percentage of 

Douglas-fir basal area, quadratic mean diameter, percentage canopy cover, 

and canopy heterogeneity. They included an appendix that explained the 

assumptions made in building a habitat capability index for the red tree vole.

390.  Stagnaro, B. 2008. Unpublished picture of tree vole nest material consisting of Bishop pine, Mendocino, Mendocino County, CA. On file with: 

University of California, Berkeley, Museum of Vertebrate Zoology.

http://calphotos.berkeley.edu/cgi/img_query?seq_num=264655&one=T. 

(August 2014).

The nest material in this picture contained Bishop pine (Pinus muricata) 

resin ducts from the nest of a Sonoma tree vole. This was a different record 

than the cases reported by Wooster and Town (2002).

391.   Stein, B.R. 1986. Comparative limb myology of four arvicolid rodent 

genera (Mammalia, Rodentia). Journal of Morphology. 187: 321–342.

Stein argued that the primitive digit feature that Kesner (1978) suggested 

was a derived and primitive characteristic may just be variation. Stein 

found the same finger feature in 2 of 12 Clethrionomys [= Myodes] she 

examined. 

392.  Stephens, F. 1906. California mammals. San Diego, CA: West Coast 

Publishing Company. 351 p.

On page 123, Stephens described the “redwood lemming mouse,” which 

was another name used for the recently described white-footed vole.

393.  Stone, G.E. 1913. Unpublished field notes. On file with: University of 

California, Berkeley, Museum of Vertebrate Zoology.
118

GENERAL TECHNICAL REPORT PNW-GTR-909

In his field notes, Stone said that nests of tree voles were common at the 

headwaters of Ryan Creek, 9.7 km north of Willits, Mendocino County, 

California. However, all of the nests he examined were deserted, and he 

speculated that this was due to fragmentation of the stand from logging. 

He did not say how he determined that the nests were tree vole nests.

394.  Stone, W. 1904. On a collection of birds and mammals from Mount 

Sanhedrin, California. Proceedings of the Academy of Natural Sciences of 

Philadelphia. 56: 576–585.

On page 578, Stone explained how the Academy of Natural Sciences of 

Philadelphia (currently the Academy of Natural Sciences of Drexel University) obtained a tree vole specimen collected by A.S. Bunnell at the Lierly 

Ranch, near Mount Sanhedrin, Mendocino County, California. The adult 

female specimen (ANSP11625) was found dead on a road in 1899. Stone 

described the specimen and gave measurements. He noted that the teeth 

were heavily worn, indicating that the vole was very old.

395.  Storer, J.E. 2003. The eastern Beringian vole Microtus deceitensis

(Rodentia, Muridae, Arvicolinae) in late Pliocene and early Pleistocene 

faunas of Alaska and Yukon. Quaternary Research. 60: 84–93.

Storer reported that fossil Phenacomys spp. were found at Cape Deceit, 

Alaska.

396.  Storer, R.W. 1944. Unpublished field notes. On file with: University of 

California, Berkeley, Museum of Vertebrate Zoology.

On 13 February 1944, Storer and Walter Dalquest collected an adult female 

tree vole and two fairly large juveniles from a nest 3.2 km north of Occidental, Sonoma County, California.

397.   Strong, W.D. 1930. Notes on mammals of the Labrador interior. Journal 

of Mammalogy. 11: 1–10.

Strong discussed mammals of Labrador, including information from 

Native Americans. He wrote, “The picture of the red-backed mouse (Clethrionomys) was likewise identified as a-pu-wish-ish, and that of the rufous 

tree mouse (Phenacomys) as an-i-suk-i-sash...” Strong concluded that this 

was an erroneous identification because the red tree vole only occurs in the 

Pacific Northwest.

398.  Sturges, F.W. 1955. Habits and distribution of land vertebrates on the 

Corvallis Watershed, Mary’s Peak, Benton County, Oregon. Corvallis, 

OR: Oregon State College. 78 p. M.A. thesis. 
119

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Sturges collected two white-footed voles in riparian habitat adjacent to a 

small stream at 305 m elevation at Connors Camp on Marys Peak. 

399.   Suzuki, N. 1992. Habitat classification and characteristics of small mammal and amphibian communities in beaver-pond habitats of the Oregon 

Coast Range. Oregon State University. 91 p. M.S. thesis. 

Suzuki used pitfall traps and museum special snap-traps to sample small 

mammals in riparian habitats in Drift Creek, Lincoln County, Oregon. In 

19,300 trap-nights, he captured 14 white-footed voles (0.07 captures per 

100 trap-nights) and no red tree voles.

400.  Suzuki, N. 2000. Effects of thinning on forest-floor vertebrates and analysis of habitat associations along ecological gradients in Oregon coastal 

Douglas-fir forests. Corvallis, OR: Oregon State University. 168 p. Ph.D. 

dissertation.

Suzuki used pitfall traps to sample thinned and unthinned stands. In his 

experimental study, he sampled four replicate plots of 35- to 45-year-old 

forest in the Tillamook Burn between Tillamook and Forest Grove. In 

addition, he sampled eight “observational” sites with paired plots (thinned 

vs. unthinned plots) at each site. The latter stands were the same age for 

each paired sample, with ages ranging from 52 to 100 years old. Observational plots were scattered over a large area from Tillamook County south 

to northern Coos County in the Coast Range. On the experimental sites, 

he captured 9,016 vertebrates in 75,600 trap-nights but did not capture a 

single red tree vole. He did report 0.05 white-footed vole captures per 100 

trap-nights, which we back-calculated to equal 38 individuals. In 2 years of 

sampling at the observational sites, he accumulated 64,000 trap-nights and 

captured 5,094 vertebrates. He reported that he caught less than 30 red tree 

voles and white-footed voles, but he did not provide any information on the 

age of the stands in which they were captured or whether those stands were 

thinned or unthinned. In his dissertation, there were no tables showing the 

locations of the actual sample plots or the numbers of animals captured in 

each stand. We could not find any of his specimens in museums.

401.   Suzuki, N.; Hayes, J.P. 2003. Effects of thinning on small mammals in 

Oregon coastal forests. Journal of Wildlife Management. 67: 352–371.

In this published version of Suzuki’s dissertation, the authors reported that 

they captured 38 white-footed voles on their experimental plots and five 

white-footed voles on their retrospective plots (the latter plots were called 
120

GENERAL TECHNICAL REPORT PNW-GTR-909

“observational plots” in the dissertation). They caught no tree voles in the 

experimental plots but did report capturing nine tree voles in the retrospective plots. They did not describe forest stands in which they captured tree 

voles.

402.  Swingle, J.K. 2005. Daily activity patterns, survival, and movements of 

red tree voles (Arborimus longicaudus) in western Oregon. Corvallis, OR: 

Oregon State University. 118 p. M.S. thesis. 

Swingle radio-collared 61 red tree voles at 2 study areas in Douglas 

County, Oregon. He reported home range size, number of nests used, activity patterns, estimated survival, nest attributes, and nest detectability from 

ground-based observers. He also reported on sexual differences in body 

mass and movements, nest fidelity, nest size, subadult dispersal, predators, reproduction, and terrestrial activity. Of 61 voles collared, 6 were 

melanistic.

403.  Swingle, J.K.; Forsman, E.D. 2004a. Red tree vole predation and survival [Abstract]. Bend, OR: 39th annual meeting of the Oregon Chapter of 

the Wildlife Society. (Available upon request from the compilers).

In western Oregon, data from 61 radio-collared red tree voles were used 

to estimate home range areas, behavior and survival rates. Predators of the 

radio-collared tree voles were weasels, owls, and a gopher snake (Pituophis 

catenifer). 

404.  Swingle, J.K.; Forsman, E.D. 2004b. Estimation of red tree vole 

(Arborimus longicaudus) home range from radio-telemetry [Abstract]. 

Corvallis, OR: Second annual conference on research in fisheries, wildlife, and ecology. (Available upon request from the compilers). 

The authors estimated home range size of radio-collared red tree voles. 

Estimates of home ranges ranged from one tree to 10,672 m2

.

405.  Swingle, J.K.; Forsman, E.D. 2009. Home range areas and activity 

patterns of red tree voles (Arborimus longicaudus) in western Oregon. 

Northwest Science. 83: 273–286.

The authors analyzed the movements of 15 male and 30 female red tree 

voles. Tree voles were primarily active at night. One of the individuals 

tracked, 17 used multiple nests (range = -7), and 18 occupied just single 

nest. Estimated mean home range size was 1,732 m2

 (median = 760 m2). 

Home range size did not differ between males and females except in late 
121

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

winter and spring when home ranges of males were larger than those of 

females. There was no evidence of ground nesting by any of the voles. 

406.  Swingle, J.K.; Forsman, E.D.; Anthony, R.G. 2010. Survival, mortality, and predators of red tree voles (Arborimus longicaudus). Northwest 

Science. 84: 255–265.

The authors estimated that the annual survival rate of 50 radio-collared red 

tree voles was 0.15. There were no strong effects of sex, vole age, or forest 

age, and little effect of mass at initial capture on survival. Of 25 mortalities attributed to predators, 15 were attributed to weasels (Mustela spp.). 

The majority (93 percent) of tree voles killed by weasels were females. 

Other predators included owls, a gopher snake, and a domestic dog (Canis 

familiaris).

407.   Swingle, J.K.; Forsman, E.D.; Sovern, S.G. 2004. A method for livetrapping tree voles. Northwestern Naturalist. 85: 134–135.

The authors placed 8- by 8- by 25-cm Havahart double-door live-traps on 

limbs next to 19 red tree vole nests. Based on observation of fresh tree vole 

nest material, all of the nests were either occupied or recently occupied. 

Traps were “baited” with fresh resin ducts collected from nearby tree vole 

nests, and pieces of apple were wired inside most of the traps (130 of 184 

trap-nights). Three red tree voles were captured in 184 trap-nights (1.6 

tree voles per 100 trap-nights). The number of tree voles captured per 100 

trap-nights was much higher than from published studies using pitfall 

traps and live-traps placed on the ground. In addition to the voles captured 

in live-traps, the authors captured two adult tree voles when nests were 

probed with a thin metal rod when the traps were removed. Although voles 

were captured in live-traps, the method was labor-intensive because trees 

had to be climbed frequently to check sprung traps.

408.  Taylor, W.P. 1913. Unpublished field notes. On file with: University of 

California, Berkeley, Museum of Vertebrate Zoology.

These notes described Walter Taylor’s activities while he was a member 

of the 1913 North Coast Counties Expedition, sponsored by the University 

of California, Berkeley, Museum of Vertebrate Zoology. Taylor said they 

found no sign of tree voles at the start of the expedition, but then Gordon 

Ferris talked to some boys at Mendocino City who told him they knew how 

to catch tree voles. The boys then led Taylor and Ferris out into the woods 

and showed them how to climb trees and catch tree voles. This experience 
122

GENERAL TECHNICAL REPORT PNW-GTR-909

provided background material for several subsequent papers published by 

Taylor (1915a, 1915b, 1961).

409.  Taylor, W.P. 1915a. Relative numbers in sexes of the tree mouse. The 

Oregon Sportsman. 3: 150–151.

Taylor pointed out that six of eight adult tree voles collected during the 

1913 North Coast Counties Expedition were females. In addition, they 

captured “...8 young males and 14 young females.” We assume he meant 

juveniles captured in their maternal nests. Based on the small number of 

adult males captured, he suggested that the mice were either polygamous 

or that females were easier to catch because they were less inclined to flee 

the nest when nests were torn apart. An alternative, and we think more 

likely, explanation is that females are easier to find because their nests are 

bigger and more conspicuous than male nests. Still another hypothesis is 

that male tree voles are harder to find because they nest on the ground; 

there is little evidence to support this hypothesis and much evidence to the 

contrary.

410.   Taylor, W.P. 1915b. Description of a new subgenus (Arborimus) of 

Phenacomys, with a contribution to knowledge of the habits and distribution of Phenacomys longicaudus True. Proceedings of the California 

Academy of Sciences (Series 4). 5: 111–161.

In this seminal paper, Taylor provided a wealth of information on the life 

history and behavior of tree voles. This information was gleaned from his 

own field work during the 1913 North Coast Counties Expedition, as well 

as conversations with other tree vole experts such as Harry Wilder and 

Aurelius Todd. Taylor thought tree voles were fairly abundant and widely 

distributed and that the lack of abundance in museum collections was due 

to the lack of knowledge about their habitat. He stated that all of the tree 

vole nests located near Mendocino City were in grand fir and that those 

tree voles were feeding on grand fir. He said that nests had also been found 

in Douglas-fir and Sitka spruce. He noted a Bishop pine cutting in a tree 

vole nest in a grand fir but did not speculate that tree voles were feeding 

on Bishop pine. He discussed colonial relationships, clustering of nests, 

locomotion, climbing behavior, docile behavior, home range, hibernation, 

nest height, nest construction, male and female nests, and much more. He 

said that tree voles do not have any extraordinary agility or quickness, and 

though they are not slow, they are a little clumsy. He pointed out that it is 

the midrib that is eaten from Douglas-fir and grand fir needles, and that 

some previous observers had obviously mistaken the uneaten resin ducts 
123

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

for the midrib. He said that tree voles were primarily arboreal, perhaps 

exclusively so. He did note that Harry Wilder thought that tree voles must 

come to the ground occasionally because some trees with nests did not 

have intertwining branches with other trees.

Taylor said tree voles were relatively docile when handled, but that a few 

had bitten their captors hard enough to draw blood. He said that green 

conifer cuttings were typically present in occupied nests and that tree voles 

occurred in the humid coastal zone of frequent fog and moderately heavy 

rainfall. In discussing home range, Taylor wrote, “It is not improbable that 

the tree mouse ranges freely through the foliage of several trees in the 

vicinity of his home nest tree.” 

Taylor said that the majority of tree vole nests were located against or 

around the bole on branch whorls and that tree voles sometimes built 

their nests on top of abandoned nests of western gray squirrels. Net lichen 

(Ramalina reticulata) was found mixed with resin ducts in some nests. He 

said that green cuttings found on the ground were indicative of occupied 

nests [actually, we have found that green cuttings on the ground are often 

indicative of predated nests]. Regarding the composting action that takes 

place inside tree vole nests, Taylor wrote “Below the level of the used 

portion of the nest there was usually found a mass of decaying matter, 

sweating and steaming like a pile of old manure or like green feed in a 

silo.” Taylor thought that large nests were the result of years of occupancy 

by tree voles, and that the type of supporting foundation was the most 

important factor determining the form of the nest. 

Taylor said that Aurelius Todd told him that nests were frequently changed 

or deserted, and that Harry Wilder reported that tree voles abandoned 

disturbed nests. Taylor suggested that small boys might be a major cause 

of abandonment of nests, but also acknowledged that some nests became 

uninhabitable when they were dislodged by storms. He did not seem to 

consider the obvious fact that voles are short-lived animals that usually die 

from predation, and that this is the primary cause of nest abandonment. 

Regarding reproduction, Taylor said that young may be born any time 

of the year but that most reproduction occurred in late winter, spring, 

and summer. He said that nests were typically occupied by a single adult 

female and her young or by solitary adult males and females. Taylor 

reported that female maternal nests were much larger than male nests 

and that male nests were higher in trees and thus harder to detect. He also 
124

GENERAL TECHNICAL REPORT PNW-GTR-909

reported that large maternal nests were more likely to be climbed to and 

dissected. He cited one instance in which two adult males were collected 

from the same tree but did not say if they came from the same nest.

411.   Taylor, W.P. 1961. A great naturalist and the long-tailed tree mouse. In: 

Terres, J.K., ed. Discovery: great moments in the lives of outstanding 

naturalists. Philadelphia, PA: F.B. Lippincott Company: 38–45.

Taylor discussed his relationship with Joseph Grinnell and described his 

first introduction to the tree vole by small boys in northwestern California. 

He finished with a succinct summary of his previous papers on tree voles 

(Taylor 1915a, 1915b).

412.  Tevis, L., Jr. 1956. Responses of small mammal populations to logging of 

Douglas-fir. Journal of Mammalogy. 57: 189–196.

From 1952 through 1954, Tevis sampled small mammals using snap-traps 

near Sayler, Humboldt and Trinity Counties, California. He captured 1,033 

small mammals but did not capture any tree voles or white-footed voles. 

We cite this paper because it is just one more piece of evidence that terrestrial snap-traps are ineffective for sampling tree voles and white-footed 

voles.

413.   Thomas, J.W.; Forsman, E.D.; Lint, J.B.; Meslow, E.C.; Noon, B.R.; 

Verner, J. 1990. A conservation strategy for the northern spotted owl: 

report of the Interagency Scientific Committee to address the conservation 

of the northern spotted owl. Portland, OR: U.S. Department of Agriculture, 

Forest Service; U.S. Department of the Interior, Bureau of Land 

Management, U.S. Department of the Interior, Fish and Wildlife Service, 

and U.S. Department of the Interior, National Park Service. 427 p.

The Interagency Scientific Committee reviewed the literature and concluded that tree voles were positively associated with stand age based on 

pitfall-trapping data and may reach highest densities in old-growth forests. 

They conceded that the data were too limited to suggest trends in abundance based on geography or elevation. In table J1, they erred by including 

the tree vole as a prey of spotted owls in Washington, where tree voles do 

not occur.

414.   Thomas, J.W.; Raphael, M.G.; Anthony, R.G.; Forsman, E.D.; 

Gunderson, A.G.; Holthausen, R.S.; Marcot, B.G.; Reeves, G.H.; 

Sedell, J.R.; Solis, D.M. 1993. Viability assessments and management considerations for species associated with late-successional and 

old-growth forests of the Pacific Northwest: the report of the Scientific 
125

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Analysis Team. Portland, OR: U.S. Department of Agriculture, Forest 

Service. 530 p.

The Scientific Analysis Team included the red tree vole in the list of species that they evaluated. They assessed species viability under five alternative planning scenarios and discussed mitigation options to ensure species 

viability.

415.  Thomas, J.W.; Raphael, M.G. [and others]. 1993. Forest ecosystem 

management: an ecological, economic, and social assessment. Report of 

the Forest Ecosystem Management Assessment Team. Portland, OR: U.S. 

Department of Agriculture, Forest Service and U.S. Department of the 

Interior, Bureau of Land Management. [Irregular pagination].

The Forest Ecosystem Management Assessment Team used an expert 

panel process to estimate the outcomes of various management options on 

tree voles (pages IV-171–174). They concluded that, “...habitat for red tree 

voles will be sufficient and well distributed under Options 1, 3, 4, and 5, 

but lower under options 7, 8, and 9.” On page IV-176, they say that “Little 

is known about the red tree vole. Studies are needed to better understand 

its basic ecology, including its habitat associations and dispersal capabilities. Further genetic research is needed to determine whether Phenacomys 

pomo and P. longicaudus are distinct species.” In Appendix Table IV-C-1, 

they estimated the area of the range of the red tree vole in Oregon and 

California and estimated how much of the range was in areas protected 

from harvest.

416.  Thomas, J.W.; Ruggiero, L.F.; Mannan, R.W.; Schoen, J.W.; Lancia, 

R.A. 1988. Management and conservation of old-growth forests in the 

United States. Wildlife Society Bulletin. 16: 252–262.

The authors listed the red tree vole as a species that uses old-growth forest 

disproportionately to its occurrence.

417.   Thompson, J.L.; Diller, L.V. 2002. Relative abundance, nest site characteristics, and nest dynamics of Sonoma tree voles on managed timberlands 

in coastal northwest California. Northwestern Naturalist. 83: 91–100.

Thompson and Diller used line transects to estimate density of Sonoma 

tree vole nests in six stand-age classes (10 to 19, 20 to 29, 30 to 39, 40 to 

49, 50 to 59, >60 years). The density of occupied or recently occupied nests 

increased with increasing stand age. Median persistence time of nests was 

28.6 months (95 percent CI = 25.8 to 34.8 months). They found a few nests 

in stands as young as 10 to 16 years old, but suggested that tree voles
126

GENERAL TECHNICAL REPORT PNW-GTR-909

usually do not start to occupy young stands until trees are about 20 years 

old. The authors located one terrestrial nest in a 38-year-old stand below a 

small Douglas-fir.

418.  Todd, A. 1891. An Oregon mouse. West American Scientist. 7: 240–242.

In this paper, Aurelius Todd described the events surrounding his discovery of the type specimen of Phenacomys longicaudus, which was collected 

by L.J. Cornelius near Coos Bay, Oregon, in 1890 (USNM 19071/25883). 

Todd said that the type specimen was not the first tree vole he had seen. He 

captured another tree vole in the headwaters of Elk Creek, about 7 miles 

east of Yoncalla, Douglas County, Oregon, in 1886, but the specimen was 

lost in transit on its way to the National Museum. 

Todd suggested that tree voles were almost exclusively arboreal. He said 

that, “The nest is a novelty in itself, being about the size of a robin’s nest, 

and built after the usual manner of mice in shape, but almost exclusively 

of the leaves of the tree in which it lives, which are split into threads from 

end to end, forming very slender filaments, seldom broken, and each leaf 

is frequently split twice or more, making from two to four threads of each 

leaf. These threads are soft, dry and apparently warm, and they show 

much ingenuity in the general make-up of the whole nest. Rarely has a few 

unsplit leaves, moss and twigs on the outside of the nest been found.” The 

split leaves that Todd referred to were obviously discarded resin ducts from 

the needles of Douglas-fir.

419.   Toweill, D.E. 1986. Resource partitioning by bobcats and coyotes in 

coniferous forest. Corvallis, OR: Oregon State University. 155 p. Ph.D. 

dissertation. 

Toweill found remains of white-footed voles in 2 of 484 bobcat (Felis [= 

Lynx] rufus) scats and 3 of 844 coyote (Canis latrans) scats in the Cascade 

Range of Oregon. He also found remains of heather voles in three coyote 

scats. He did not say how he was able to distinguish remains of whitefooted voles from red tree voles. 

420.  True, F.W. 1890. Description of a new species of mouse, Phenacomys 

longicaudus, from Oregon. Proceedings National Museum. XIII, No. 826: 

303–304.

True described the type specimen of Phenacomys longicaudus, which was 

collected by L.J. Cornelius near Coos Bay, Oregon, and sent to the U.S. 

National Museum by Aurelius Todd. The specimen (USNM 19071/25883) 

included a dried skin and crushed skull. True described the bright rusty 
127

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

brown color of the vole and said that the molars were rooted and resembled 

those of the heather vole. He referred to the new species as a “mouse” but 

did not give it a common name.

421.   U.S. Department of Agriculture, Forest Service. 1996. Survey and manage inventory report: Mount Hood National Forest 1995. Sandy, OR. 105 p.

This report described the results of red tree vole surveys on the Mount 

Hood National Forest from 1995 through 1996. In 1995, surveyors located 

eight trees with tree vole nests in the Bull Run watershed and one nest in 

the South Fork of Eagle Creek. Not all trees were climbed so it is not clear 

how the observers confirmed the species that built the nests. In 1996, three 

red tree vole nests were confirmed by tree climbers in Eagle Creek, and 22 

more were classified as potential red tree vole nests that were not climbed.

422.  U.S. Department of Agriculture, Forest Service. 2007. Notice of availability (NOA) Record of Decision (ROD) for the final supplement to the 

2004 supplemental environmental impact statement to remove or modify 

the survey and manage mitigation measure standards and guidelines. 

Federal Register. 72: 41288–41289. 

In this announcement in the Federal Register, the U.S. Forest Service said 

that the Northwest Forest Plan had made it difficult for the Forest Service 

and Bureau of Land Management to meet goals for timber harvest, hazardous fuels treatment, and ecosystem restoration. They proposed to solve this 

problem by removing the tree vole from the “survey and manage” requirements in the Northwest Forest Plan. As an alternative, they proposed to 

change the tree vole to a “special status species,” which does not require 

specific surveys or protections except in limited areas.

423.  U.S. Department of Agriculture, Forest Service; U.S. Department of 

the Interior, Bureau of Land Management. 1994a. Final supplemental 

impact statement on management of habitat for late-successional oldgrowth forest related species within the range of the northern spotted owl, 

Volume 1. Portland, OR. [Irregular pagination].

The agencies concluded that the “Survey and Manage” standards and 

guidelines would help in providing a well-distributed population of red tree 

voles. In table 3 and 4-30 (page 3 and 4-184), they estimated the likelihood that tree voles would remain well-distributed within their historical 

range under each of the different land management alternatives in the draft 

supplemental environmental impact statement. Under alternative 9 (the 

selected alternative), the red tree vole was given a 73 percent likelihood 
128

GENERAL TECHNICAL REPORT PNW-GTR-909

of remaining well distributed, a 25 percent likelihood of becoming locally 

restricted, a 2 percent likelihood of becoming restricted to refugia, and a 

0 percent likelihood of extirpation. Under alternative 9, the Sonoma tree 

vole was given a 78 percent likelihood of remaining well distributed, a 23 

percent likelihood of becoming locally restricted, and a 0 percent likelihood of becoming restricted to refugia or going extinct.

424.  U.S. Department of Agriculture, Forest Service; U.S. Department of 

the Interior, Bureau of Land Management. 1994b. Final supplemental 

impact statement on management of habitat for late-successional oldgrowth forest related species within the range of the northern spotted owl, 

Volume 2: appendices. Portland, OR. [Irregular pagination].

In table B11-1 (page B-160), the red tree vole was the only mammal 

included on the list of “survey and manage” species. It was designated as 

deserving surveys prior to ground-disturbing activities.

425.  U.S. Department of Agriculture, Forest Service; U.S. Department 

of the Interior, Bureau of Land Management. 1994c. Final supplemental impact statement on management of habitat for late-successional 

old-growth forest related species within the range of the northern spotted 

owl: Appendix J2–results of additional species analysis. Portland, OR. 

[Irregular pagination].

The Species Analysis Team reviewed the panel notes from Thomas et al. 

(1993) and did not find support for further mitigation to meet the screening criteria for the Sonoma tree vole under alternative 9. The red tree vole 

failed to pass the screens because of its association with old forest, limited 

dispersal capabilities, and general lack of data on distribution, habitat 

requirements and population status. But the main concern for the red tree 

vole was that forest fragmentation and spatial isolation of old forests would 

prevent gene flow and demographic movement between metapopulations. The Species Analysis Team concluded that past management had 

fragmented red tree vole habitat and that timber harvesting practices had 

reduced tree vole numbers. The team concluded that more information was 

needed on dispersal and abundance of tree voles, especially in young forests. They also discussed the need for connectivity of old forest in reserves 

to facilitate movement.
129

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

426.  U.S. Department of Agriculture, Forest Service; U.S. Department of 

the Interior, Bureau of Land Management. 1994d. Record of Decision 

for amendments to Forest Service and Bureau of Land Management planning documents within the range of the northern spotted owl. Portland, 

OR. 74 p.

This record of decision was based on alternative 9 from the final environmental impact statement. The standards and guidelines established the 

Survey and Manage Program that required surveys for red tree voles prior 

to ground-disturbing activities and management of known sites.

427.   U.S. Department of Agriculture, Forest Service; U.S. Department 

of the Interior, Bureau of Land Management. 1994e. Standards and 

guidelines for management of habitat for late-successional and old-growth 

forest related species within the range of the northern spotted owl: attachment A to the Record of Decision for amendments to Forest Service and 

Bureau of Land Management planning documents within the range of the 

northern spotted owl. Portland, OR. [Irregular pagination].

The Forest Service and Bureau of Land Management concluded that they 

should conduct surveys to determine if red tree voles were present prior to 

conducting ground-disturbing activities. This decision added the tree vole 

to the so-called list of “survey and manage species.” The tree vole was the 

only mammal on the survey and manage list, which also included many 

plants, invertebrates, amphibians, and birds.

428.  U.S. Department of Agriculture, Forest Service; U.S. Department of 

the Interior, Bureau of Land Management. 1996. Draft survey protocol 

and management recommendations for the red tree vole Arborimus longicaudus in Oregon. Version 1.0. June 1996. Portland, OR. 44 p.

In this first attempt to develop a survey protocol for red tree voles on 

federal lands, the Tree Vole Working Group reviewed literature and unpublished reports to summarize what was known regarding the taxonomy, 

distribution, biology, and habitat of the red tree vole. They concluded that 

red tree vole habitat consisted primarily of moist Douglas-fir forests and 

that the network of old forest reserves established in the Northwest Forest 

Plan would support a well-distributed population of tree voles.
130

GENERAL TECHNICAL REPORT PNW-GTR-909

429.   U.S. Department of Agriculture, Forest Service; U.S. Department of 

the Interior, Bureau of Land Management. 1999. Draft supplemental 

environmental impact statement for amendment to the survey and manage, protection buffer, and other mitigation measures standards and guidelines. Portland, OR. 492 p.

This supplemental environmental impact statement listed the life-history 

traits and habitat requirements that led to the listing of the red tree vole 

as a “survey and manage” species in the Northwest Forest Plan. It also 

defined the interim survey protocol for red tree vole nests and summarized 

surveys conducted to date. The anonymous authors reported that few 

or no tree voles were found in the north Cascades and northern Coast 

Ranges of Oregon. They cited pitfall trapping studies and one nest survey 

(Meiselman and George 1996) in support of the conclusion that tree voles 

were significantly more abundant in old forest than in young forests. They 

mentioned but did not specifically cite a distance sampling study (Biswell 

unpublished data) in the central Coast Ranges where tree vole nests were 

more abundant in old forest than young forest (0.83 vs. 0.18 nests per hectare). They also reported that surveyors found 0.07 tree vole nest trees per 

hectare in predisturbance surveys in northern Josephine County, Oregon.

430.  U.S. Department of Agriculture, Forest Service; U.S. Department 

of the Interior, Bureau of Land Management. [2000a]. Management 

recommendations for the Oregon red tree vole, Arborimus longicaudus, 

Version 2.0. Portland, OR. 25 p.

This literature review included information on reproduction, ecology, 

nests, distribution, habitat, threats, information needs, and management 

recommendations of red tree voles. The anonymous authors used the term 

“active” to describe tree vole nests that were either occupied or recently 

occupied. The term “active” tree vole nest from different versions of the 

tree vole survey protocol (Biswell et al. [2000], 2002; Huff et al. 2012) was 

defined as nests containing green resin ducts, green fecal pellets, and/or 

green cuttings. 

431.   U.S. Department of Agriculture, Forest Service; U.S. Department of 

the Interior, Bureau of Land Management. 2000b. Final supplemental 

environmental impact statement for amendment to the survey & manage, 

protection buffer, and other mitigation measures standards and guidelines 

- Volume 1–chapters 1–4. U.S. Department of Agriculture, Forest Service, 

Regional Ecosystem Office, Portland, OR. XLVIII + 516 p.
131

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

The agencies described and compared four management alternatives for 

“survey and manage” species, including the red tree vole. They concluded 

that all four alternatives would provide sufficient habitat but would differ 

in the degree of certainty regarding persistence of tree voles.

432.  U.S. Department of Agriculture, Forest Service; U.S. Department of 

the Interior, Bureau of Land Management. 2000c. Final supplemental 

environmental impact statement for amendment to the survey & manage, 

protection buffer, and other mitigation measures standards and guidelines 

- Volume 2 - appendices. U.S. Department of Agriculture, Forest Service, 

Regional Ecosystem Office, Portland, OR. 346 p.

This document included background material for the analyses that were 

presented in the companion volume 1, including a long section that 

included responses to reviewer questions.

433.  U.S. Department of Agriculture, Forest Service; U.S. Department of 

the Interior, Bureau of Land Management. 2000d. Final supplemental 

environmental impact statement for amendment to the survey & manage, 

protection buffer, and other mitigation measures standards and guidelines. 

Portland, OR. 40 p.

The agencies concluded that the red tree vole would have enough habitat to 

maintain stable populations under all of the proposed alternatives except 

for alternative 2.

434.  U.S. Department of Agriculture, Forest Service; U.S. Department of 

the Interior, Bureau of Land Management. 2001. Record of Decision 

and standards and guidelines for amendment to the survey and manage, 

protection buffer, and other mitigation measures standards and guidelines. 

Portland, OR. 86 p.

This record of decision amended the “survey and manage” guidelines by 

increasing or decreasing levels of protection afforded different species. It 

included a recommendation that the agencies conduct strategic surveys of 

tree voles to learn more about their distribution and abundance. An annual 

species review process was established to evaluate new information on 

whether species should be added or removed from the list of survey and 

manage species.

435.  U.S. Department of Agriculture, Forest Service; U.S. Department of 

the Interior, Bureau of Land Management. 2003a. Draft supplemental 

environmental impact statement to remove or modify the survey and manage mitigation measure standards and guidelines. Portland, OR. 334 p.
132

GENERAL TECHNICAL REPORT PNW-GTR-909

In this 2003 draft supplement to the 2000 final supplemental environmental impact statement for the northern spotted owl, the agencies presented 

three alternatives that would either remove or modify the “survey and 

manage” standards and guidelines that were adopted as part of the Northwest Forest Plan. The stated reasons for doing the analysis were that (1) it 

was costing too much money and time to survey and protect species like 

tree voles, and (2) the survey and manage standards and guidelines were 

making it difficult to meet the resource management goals and objectives 

set forth in the Northwest Forest Plan. Of the three alternatives considered, 

the preferred alternative was to get rid of the survey and manage standards 

and guidelines. If adopted this would mean that the agencies would no longer be required to survey for tree voles in most areas proposed for harvest.

436.  U.S. Department of Agriculture, Forest Service; U.S. Department of 

the Interior, Bureau of Land Management. 2003b. Process for determination of red tree vole sites as non-high priority sites within the “pilot” 

area. Portland, OR. 15 p.

This report described the process that field units were supposed to use to 

evaluate whether a site could be classified as a “non-high-priority site” 

for surveys of tree voles in central Coast Ranges and central Cascades 

of Oregon. Determination of non-high-priority sites was to be conducted 

at two scales: the watershed scale and survey polygon scale. The four 

criteria for determining non-high-priority sites were (1) the percentage of 

federal lands within a watershed, (2) habitat quality, (3) survey effort, and 

(4) number of “active” tree vole nests located. For watersheds with high 

amounts of reserves and red tree vole habitat, these criteria could be used 

to programmatically release areas from “survey and manage” measures for 

the red tree vole.

437.   U.S. Department of Agriculture, Forest Service; U.S. Department of 

the Interior, Bureau of Land Management. 2004a. Final supplemental environmental impact statement to remove or modify the survey and 

manage mitigation measure standards and guidelines, Volume 1. U.S. 

Department of Agriculture, Forest Service, Regional Ecosystem Office, 

Portland, OR. 450 + 332 p.

In this followup to the 2003 draft supplemental environmental impact 

statement, the agencies announced that their preferred alternative was to 

remove the “survey and manage” standards and guidelines from the Northwest Forest Plan. Under this alternative the tree vole would be classified 
133

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

as a Special Status Species on federal forest lands in the northern Coast 

Ranges. The stated reasons for retaining the tree vole as a Special Status 

Species in the northern Coast Ranges were low population numbers and 

lack of federal lands. 

438.  U.S. Department of Agriculture, Forest Service; U.S. Department of 

the Interior, Bureau of Land Management. 2004b. Final supplemental 

environmental impact statement to remove or modify the survey and manage mitigation measure standards and guidelines, Volume 2-Appendices. 

Portland, OR. 414 p.

This document included background material for the analyses that were 

presented in Volume 1 of the 2004 final supplemental environmental 

impact statement (FSEIS). There is a large section in which the agencies 

responded to questions raised by reviewers during preparation of the 

FSEIS, including specific questions regarding tree voles on pages 243–246. 

439.   U.S. Department of Agriculture, Forest Service; U.S. Department of 

the Interior, Bureau of Land Management. 2004c. Record of Decision 

to remove or modify the survey and manage mitigation measure standards 

and guidelines in Forest Service and Bureau of Land Management planning documents within the range of the northern spotted owl. Portland, 

OR. 44 p.

In this March 2004 Record of Decision the agencies announced their 

decision to remove the “survey and manage” mitigation measure from the 

Northwest Forest Plan. This decision would allow the agencies to conduct 

timber sales without protecting tree vole nest sites or conducting surveys to 

determine if tree voles were present.

440.  U.S. Department of Agriculture, Forest Service; U.S. Department of 

the Interior, Bureau of Land Management. 2004d. To remove or modify the survey and manage mitigation measure standards and guidelines. 

Federal Register. 69: 3316−3317.

This notice informed the public that the U.S. Forest Service and Bureau 

of Land Management had completed a final supplemental environmental 

impact statement in which they proposed to remove the “survey and 

manage” mitigation measure from the Northwest Forest Plan. Instead, 

they proposed to manage species like tree voles using, “...other elements 

of the Northwest Forest Plan and their existing Special Status Species 

Programs....”
134

GENERAL TECHNICAL REPORT PNW-GTR-909

441.   U.S. Department of Agriculture, Forest Service; U.S. Department 

of the Interior, Bureau of Land Management. 2005. Bureau of Land 

Management Districts and National Forests within the range of the northern spotted owl; western Oregon and Washington, and northwestern 

California; removal of survey and manage mitigation measure standards 

and guidelines. Federal Register. 70: 73483.

In this notice, the U.S. Forest Service and Bureau of Land Management 

announced that they would prepare a supplemental environmental impact 

statement (SEIS) to respond to a court order in the case of Northwest 

Ecosystem Alliance v. Rey, dated August 1, 2005. In this case, Judge 

Pechman ruled that the 2004 SEIS was deficient for several reasons. In this 

notice, the agencies stated that they would address these deficiencies in a 

new SEIS.

442.  U.S. Department of Agriculture, Forest Service; U.S. Department of 

the Interior, Bureau of Land Management. 2006. Draft supplement to 

the 2004 final supplemental environmental impact statement to remove or 

modify the survey and manage mitigation measure standards and guidelines. Portland, OR. XIV + 140 p. + appendix.

This draft supplemental environmental impact statement (SEIS) was 

prepared in response to the August 1, 2005, court order in the case of 

Northwest Ecosystem Alliance v. Rey. It was not a stand-alone document. 

It specifically addressed deficiencies in the 2004 final environmental 

impact statement that were identified by the court. It included sentences or 

sections that replaced or expanded on sections in the 2004 final SEIS. The 

section on red tree voles (pages 87–92) briefly summarized new research. 

The agencies continued to argue that they could drop the “survey and manage” standards and guidelines, but they acknowledged that the proposed 

action might not provide for stable populations of some species within the 

area of the Northwest Forest Plan. In the case of the red tree vole (pages 

87–92), they acknowledged that the preferred alternative would not provide 

enough habitat to support stable populations in some parts of the range of 

the species.

443.  U.S. Department of Agriculture, Forest Service; U.S. Department of 

the Interior, Bureau of Land Management. 2007a. Supplemental to the 

July 2006 draft supplemental for the 2004 FSEIS to remove or modify 

the survey and manage mitigation measure standards and guidelines. 

Portland, OR. VII + 66 p + appendix.
135

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

In this supplement to the supplement, the U.S. Forest Service and Bureau 

of Land Management added a no-action alternative that attempted to 

address the 2006 U.S. 9th Circuit Court ruling that they violated federal 

law in their attempt to remove or modify “survey and manage” mitigation 

for the red tree vole.

444.  U.S. Department of Agriculture, Forest Service; U.S. Department of 

the Interior, Bureau of Land Management. 2007b. Final supplement 

to the 2004 supplemental environmental impact statement to remove or 

modify the survey and manage mitigation measure standards and guidelines, Vol. 1, chapters and text. Portland, OR. XXV + 414 p.

In yet another final supplemental environmental impact statement, the U.S. 

Forest Service and Bureau of Land Management stuck to their guns and 

selected a preferred option that removed the “survey and manage” mitigation measure from the Northwest Forest Plan. The expected effects of the 

various options on tree voles were discussed on pages 285–297.

445.  U.S. Department of Agriculture, Forest Service; U.S. Department of 

the Interior, Bureau of Land Management. 2001. Survey and manage—

fiscal year 2001 annual status report, Northwest Forest Plan. 10 p.

In this annual report, the agencies reported that they found tree vole nests 

in 1,239 predisturbance surveys. They also announced plans to start a 

new survey in which a contractor would conduct tree vole surveys in 400 

randomly selected plots starting in 2002.

446.  U.S. Department of Agriculture, Forest Service; U.S. Department of the 

Interior, Bureau of Land Management. 2003. Survey and manage—fiscal 

year 2003 annual status report, Northwest Forest Plan. 14 p.

Little new information on tree voles was presented in this report, but the 

authors did mention that a radio-tracking study of red tree voles was being 

conducted.

447.   U.S. Department of the Interior, Bureau of Land Management. 2007. 

Record of Decision to remove the survey and manage mitigation measure 

standards and guidelines from Bureau of Land Management land and 

resource management plans within the range of the northern spotted owl. 

Portland, OR. 44 p.

In this record of decision, the Under Secretary of Land and Minerals 

Management for the U.S. Department of the Interior (C. Stephen Alfred) 

announced his decision to remove the “survey and manage” mitigation 
136

GENERAL TECHNICAL REPORT PNW-GTR-909

measure standards and guidelines as they applied to Bureau of Land Management lands in the Northwest Forest Plan. He specifically addressed the 

red tree vole on pages 22–23 where he said that the Bureau of Land Managment would discontinue predisturbance surveys for tree voles in all areas.

448.  U.S. Department of the Interior, Fish and Wildlife Service. 2004. 

Species of concern in Oregon. http://www.fws.gov/oregonfwo/Species/

Documents/SOC-Nov05.pdf. (March 2009).

The red tree vole and white-footed vole were listed as species of concern in 

Oregon. However, the U.S. Fish and Wildlife Service also concluded that 

more information was needed to determine the status of both species. 

449.  U.S. Department of the Interior, Fish and Wildlife Service. 2008. 

Endangered and threatened wildlife and plants: 90-day finding of a petition to list a distinct population segment of the dusky tree vole (Arborimus 

longicaudus silvicola) as endangered or threatened. Federal Register. 73: 

63919–63926.

The U.S. Fish and Wildlife Service announced that they had received 

a petition requesting that they list the dusky tree vole as endangered or 

threatened. They concluded that the petition had merit and announced 

that they would start the evaluation process to determine if listing was 

warranted.

450.  U.S. Department of the Interior, Fish and Wildlife Service. 2011. 

Endangered and threatened wildlife and plants: 12-month finding of a 

petition to list a distinct population segment of the red tree vole as endangered or threatened. Federal Register. 76: 63720–63762.

After 3 years of review, the U.S. Fish and Wildlife Service concluded that 

listing of the tree vole in the northern Coast Ranges of Oregon was warranted, but they declined to begin the listing process because other species 

had priority.

451.   Verts, B.J. 1976. Red tree vole range maps. In: Allan, W.G.S.; Patton, 

C.P.; Plank, R.D. Atlas of Oregon. Eugene, OR: University of Oregon: 

154–157.

Verts compiled the maps for the mammal section in this atlas, including 

the red tree vole. These maps are now out of date.

452.  Verts, B.J.; Carraway, L.N. 1984. Keys to the mammals of Oregon. 

Corvallis, OR: Oregon State University Book Stores. 178 p.

This document included a morphological key and range maps for Oregon 

mammals, including the red tree vole and white-footed vole. 
137

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

453.  Verts, B.J.; Carraway, L.N. 1995. White-footed vole: Phenacomys albipes. Mammalian Species. 494: 1–5.

This species account described the taxonomy, distribution, habitat, food, 

behavior, and reproduction of the white-footed vole.

454.  Verts, B.J.; Carraway, L.N. 1998. Land mammals of Oregon. Berkeley, 

CA: University of California Press. XVI + 668 p.

In this incredibly detailed book, the authors provided a wealth of information on all land mammals in Oregon, including the red tree vole and 

white-footed vole. They presented data on morphological measurements, 

distribution, habitat, geographic variation, reproduction, ontogeny, mortality, behavior, and taxonomy. They limited their analyses of distribution 

primarily to data from museum specimens.

455.  Voge, M. 1955. A list of cestode parasites from California mammals. 

American Midland Naturalist. 54: 413–417.

Voge listed a cestode parasite (Andrya spp.) collected from a Sonoma tree 

vole 0.8 km east of Stewarts Point, Sonoma County, California.

456.  Voge, M.; Bern, H.A. 1949. Cecal villi in the red tree mouse, 

Phenacomys longicaudus. Anatomical Record. 104: 477–485. 

Voge and Bern found that the red tree vole, white-footed vole, and heather 

vole all had large numbers of hair-like villi in the cecum, whereas voles 

in the genus Microtus, Clethrionomys [= Myodes], Lagurus, Lemmus and 

Ondatra did not have such villi. They suggested that these villi could 

slow the passage of food through the cecum, but did not say whether they 

thought these villi could increase the nutrient absorption of the organ. 

They suggested that the structure of the tree vole cecum might be an 

adaptation to the specialized diet.

457.   Voge, M.; Bern, H.A. 1952. Cecal villi in a pika (Ochotona princeps). 

Journal of Mammalogy. 55: 104–105.

Voge and Bern compared the cecal villi of the American pika (Ochotona 

princeps) and Phenacomys spp. The cecal villi of the pika were shorter, 

wider, and flatter. Unlike the red tree vole, the epithelium of the cecal villi 

in the pika did not show intense alkaline phosphatase activity, suggesting 

that villi in the cecum did not serve the same function in both species.

458.  Voth, E.H.; Maser, C.; Johnson, M.L. 1983. Food habits of Arborimus 

albipes, the white-footed vole, in Oregon. Northwest Science. 57: 1–7.
138

GENERAL TECHNICAL REPORT PNW-GTR-909

The authors examined the stomach contents and fecal pellets of 21 whitefooted voles captured in snap-traps in riparian habitats in the Oregon Coast 

Ranges. Based on the percentage of occurrence, the diet consisted entirely 

of leaves from vascular plants, including 41 percent red alder, 9 percent 

red willow (Salix lasiandra), 15 percent shrubs, and 23 percent forbs. They 

found no seeds, fruits, fungi, or invertebrates in the diet. They concluded 

that the white-footed vole was a browser with a niche that extended from 

the ground into the forest canopy.

459.  Vrieze, J.M. 1979. Unpublished field notes. On file with: Humboldt State 

University, Vertebrate Museum, Arcata, CA.

In his field notes for 8 to 10 July 1979, Vrieze described a field trip in 

which he, Brian McNab, and McNab’s two sons collected three adult 

Sonoma tree voles east of Arcata, Humboldt County, California. We think 

McNab (1992) may have used these tree voles in his analyses of energetics 

and then discarded the specimens, as we did not find the specimens in any 

museums.

460.  Vrieze, J.M. 1980. Spatial patterning of red tree mouse, Arborimus longicaudus, nests. Arcata, CA: Humboldt State University. 37 p. M.A. thesis.

Vrieze examined the horizontal and vertical distribution of tree vole nests 

in young forests (≤50 years old) east of Arcata, California. He found that 

the horizontal distribution of nests did not significantly deviate from 

random. On the vertical axis, tree voles appeared to build their nests near 

the lowest branches with green needles. He concluded that differences in 

nest height between study areas were due to differences in the heights of 

the lowest green branches in the various study areas. Vrieze climbed to all 

nests to document activity and species of cuttings on nests. He classified 

tree vole nests as “active” if they had fresh conifer cuttings, fecal pellets, 

and resin ducts, and he assumed that “active” nests were occupied by one 

or more tree vole. This was the first reference we are aware of that classified tree vole nests as “active” or “inactive.” Unfortunately, this has led to 

a great deal of confusion because all subsequent groups that have used the 

term “active” to refer to tree vole nests have used the term to include any 

nest with evidence of recent use, regardless of whether the nest is actually 

occupied. As a result, none of the surveys conducted using the “active” or 

“inactive” classification data can be used to estimate the actual density of 

tree voles.
139

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

461.  Waldien, D.L. 2005. Population and behavioral responses of small mammals to silvicultural and down wood treatments in the Oregon Coast 

Range. Corvallis, OR: Oregon State University. 114 p. Ph.D. dissertation. 

Waldien used Sherman live-traps (52,800 trap-nights) and Tomahawk 201 

live-traps (10,800 trap-nights) to sample small mammals in Oregon State 

University’s McDonald Forest near Corvallis, Benton County, Oregon. He 

did not catch any red tree voles or white-footed voles.

462.  Walker, A. 1928. Some Oregon Phenacomys. Journal of Mammalogy.

4: 254–255.

Walker said that the only prior collections of dusky tree voles were the type 

of specimen collected near Tillamook in 1916 and a female and four young 

collected near Corvallis by Wight (1925). He described an interesting case 

in which loggers near Tillamook used dynamite to remove a hollow western 

hemlock snag, only to find four or five dazed tree voles staggering about in 

the hole left by the stump. Peter Walker captured one of the injured voles and 

gave it to his brother Alex. When Alex visited the site the next day, he found 

another mangled tree vole in the hole. He suspected that the voles had been 

nesting inside the base of the snag and were left in the hole when their nest 

was blasted apart. Fragments of a nest were found in the depression where 

the snag had stood. The site was on a hillside covered by a forest of large 

Douglas-fir and western hemlock. The trees in the immediate vicinity of the 

nest were exclusively hemlock and had just been felled and were still lying 

on the ground. The two voles that were collected were an adult male and 

female. This episode is puzzling because adult tree voles are rarely found 

together in the same nest, and it was unclear if there were multiple nests or if 

the nest was located in the stump of the snag (i.e., a ground nest), or fell from 

a location higher up in the snag.

463.  Walker, A. 1930. Notes on the forest Phenacomys. Journal of 

Mammalogy. 11: 233–235.

Walker reported that a logger brought him a live dusky tree vole that was 

captured on Cape Meares, Tillamook County, Oregon, in 1929. The vole 

was captured in the branches of a Sitka spruce that had just been felled. 

The logger saw another vole at the same time, but did not catch it. Walker 

kept the vole alive for several months and described its behavior, including climbing and dangling by one foot. The vole preferred to eat western 

hemlock and would ignore spruce and Douglas-fir if hemlock was available. Walker stated that fresh young tips of limbs were preferred.
140

GENERAL TECHNICAL REPORT PNW-GTR-909

Walker also described a trip that he and his brother Peter made to Cape 

Lookout, Tillamook County, Oregon, in 1929. On that trip they found four 

tree vole nests in young Sitka spruce trees and captured one adult female 

and two young, all of which died soon after capture. They also found a nest 

in a western hemlock that had fresh nest material, but was unoccupied. All 

nests in Sitka spruce were 12 to 15 m above ground and had spruce cuttings on top. The nest in the western hemlock was 4.5 m up in a bushy tree 

and had hemlock cuttings on top.

464.  Waltari, E.; Hijmans, R.J.; Peterson, A.T.; Nyári, Á.S.; Perkins, S.L.; 

Guralnick, R. 2007. Locating Pleistocene refugia: comparing phylogeographic and ecological niche model predictions. PLoS ONE. 2: 1–11.

The authors used ecological niche models to explore spatially explicit 

hypotheses regarding past geographic patterns of distribution. They 

predicted three phylogeographic refugia for the red tree vole during the

last glacial maximum: Oregon Cascades, Oregon Coast Ranges, and 

northwestern California Coast Ranges.

465.  Ward, J.P., Jr. 1990. Spotted owl reproduction, diet and prey abundance 

in northwest California. Arcata, CA: Humboldt State University. 70 p. 

M.S. thesis. 

James (Pat) Ward identified 495 prey items from nine northern spotted 

owl territories near Dinsmore, Humboldt County, California. Of the 495 

prey items, 8.9 percent were tree voles. Ward also used Sherman traps 

and Tomahawk live-traps to sample prey (30,980 trap-nights) but did not 

capture any tree voles, even though many traps were placed on tree trunks.

466.  Ward, J.P., Jr.; Gutiérrez, R.J.; Noon, B.R. 1998. Habitat selection by 

northern spotted owls: the consequences of prey selection and distribution. 

Condor. 100: 79–92.

The authors reported that Sonoma tree voles were important prey items of 

northern spotted owls in northwestern California but could not be effectively sampled with Sherman live-traps that were placed on the ground.

467.  Waters, J.R.; Zabel, C.J.; McKelvey, K.S.; Welsh, H.H., Jr. 2001.

Vegetation patterns and abundances of amphibians and small mammals in 

a northwestern California watershed. Northwest Science. 75: 37–52.

From 1995 through 1996, the authors used pitfall traps to sample small 

mammals and amphibians near the border of Humboldt and Trinity Counties in northwestern California. They reported capturing Sonoma tree voles 

at greater than 7 of the 42 streamside sites they sampled, but they did not 
141

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

report the actual number of tree voles captured or what they did with the 

specimens.

468.  Watson, H.C. 1932. Unpublished field notes. On file with: University of 

California, Berkeley, Museum of Vertebrate Zoology.

Watson said that he collected an adult female and juvenile tree vole at the 

Reilley Ranch in the South Fork Mountains, Trinity County, California, in 

1932. He was working with Chester Lamb (1932).

469.  Webb, M.G. 1981–1982. Unpublished field notes. On file with: Humboldt 

State University, Vertebrate Museum, Arcata, CA.

Webb’s field notes for 1981−1982 described trips in which he, Bill Gannon, 

Doug Kelt, Tim Lawlor, and Doug Kain collected Sonoma tree voles east 

of Arcata, California. We found eight tree vole specimens in museums that 

were collected by Webb, but his field notes only listed four.

470.  Weng, C.; Jackson, S.T. 2000. Species differentiation of North American 

spruce (Picea) based on morphological and anatomical characteristics of 

needles. Canadian Journal of Botany. 78: 1367–1383.

Weng and Jackson described the structure and placement of resin ducts 

in Sitka spruce needles, which are an important food source for tree voles 

in the northern Coast Ranges of Oregon. They found that Sitka spruce 

needles had discontinuous resin duct canals, which the authors referred 

to as resin duct sacs. Sitka spruce had an average of 4.9 mm of resin duct 

sacs per needle. In figure 5, they showed a cross section of a Sitka spruce 

needle that had a single resin duct sac on one side of the needle.

471.  Whitaker, J.O., Jr.; Maser, C. 1979. Parasitic mites of voles of the genera Arborimus and Lagurus in Oregon. Northwest Science. 53: 224–227.

Whitaker and Maser found that 15 of 17 red tree voles examined had mites, 

including at least 5 different species. All but 1 of 33 white-footed voles had 

mites, including 7 different species. The authors concluded that the damp 

arboreal nests of tree voles and subterranean nests of white-footed voles 

were ideal habitats for mites.

472.  Whitaker, J.O., Jr.; Walters, B.L.; Castor, L.K.; Ritzi, C.M.; Wilson, 

N. 2007. Host and distribution list of mites (Acari), parasitic and phoretic, in the hair or on the skin of North American wild mammals north of 

Mexico: records since 1974. Lincoln, NE: Faculty Publications from the 

Harold W. Manter Laboratory of Parasitology. http://digitalcommons.unl.

edu/parasitologyfacpubs/1/. (August 2014). 
142

GENERAL TECHNICAL REPORT PNW-GTR-909

The red tree vole was host to Radfordia arborimus and Haemogamasus 

reidi. The white-footed vole was host to Eulaelaps stabularis, H .ambulans, H. occidentalis, and Macrocheles praedafimetorum. Both the red tree 

vole and white-footed vole were hosts to Glycyphagus hypudaei, Androlaelaps fahrenholzi, and Quasilistrophorus microticolus.

473.  Whitaker, J.O., Jr.; Wilson, N. 1974. Host and distribution lists of mites 

(Acari), parasitic and phoretic, in the hair of wild mammals of North 

America, north of Mexico. American Midland Naturalist. 91: 1–67.

Whitaker and Wilson reported that the white-footed vole was host to 

the mite Haemogamasus reidi and the red tree vole was host to the mite 

Androlaelaps fahrenholzi.

474.  Wiens, J.D. 2012. Competitive interactions and resource partitioning 

between northern spotted owls and barred owls. Corvallis, OR: Oregon 

State University. 141 p. Ph.D. dissertation. 

On his study area in the Oregon Coast Ranges, David Wiens identified 179 

tree voles in pellets of spotted owls and 155 tree voles in pellets of barred 

owls. Tree voles accounted for 14.7 and 4.6 percent of prey items identified 

in the pellets of spotted and barred owls, respectively. 

475.  Wiens, J.D.; Anthony, R.G.; Forsman, E.D. 2014. Competitive interactions and resource partitioning between northern spotted owls and barred 

owls in western Oregon. Wildlife Monographs. 185: 1–50.

This was the published version of the information in Wiens (2012).

476.  Wight, H.M. 1925. Notes on the tree mouse, Phenacomys silvicola.

Journal of Mammalogy. 6: 282–283.

On a zoology field trip led by Howard Wight in 1924, a student climbed a 

tree and found a nest that contained four juvenile tree voles near Corvallis, Oregon. The nest was about 14 m up inside the broken top of a 60-cm 

Douglas-fir. The young were removed and three snap-traps were placed on 

top of limbs radiating out from the nest. The next morning the adult female 

was found caught by a leg and hanging from one of the traps. Though the 

young were fed a mixture of cow and goat milk, they all died within a 

week and were preserved in alcohol. The female was preserved as a study 

skin. Her stomach contained fir needles, bark and lichen, all in a finely 

pulverized state. 

477.   Wight, H.M. 1974. Nongame wildlife and forest management. In: Black, 

H.C., ed. Wildlife and forest management in the Pacific Northwest. 

Corvallis, OR: Oregon State University, School of Forestry: 27–38.
143

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

In this report, Howard Wight listed the red tree vole as a snag-dwelling 

mammal. He got it right, because tree voles do occasionally nest in snags 

that are in contact with the limbs of live trees (Swingle 2005: 29).

478.  Williams, D.F. 1986. Mammalian species of special concern in 

California. Sacramento, CA: California Department of Fish and Game, 

Habitat Conservation Planning Branch. 100 p.

The California Department of Fish and Game listed the red tree vole as 

a species of concern owing to habitat loss. In this report, Williams gave 

a species account that included information on distribution, population 

status, and habitat. He suggested that selective logging may be less detrimental to tree voles than clearcutting. Williams stated that red tree voles 

have “a spotty pattern of dispersion,” citing Taylor (1915) as his source. 

Williams said that little was known about white-footed voles, especially 

distribution and habitat.

479.   Wilson, T.M.; Forsman, E.D. 2013. Thinning effects on spotted owl 

prey and other forest-dwelling mammals. In: Anderson, P.D.; Ronnenberg, 

K.L., eds. Density management for the 21st century: west side story. 

Gen. Tech. Rep. PNW-GTR-880. Portland, OR: U.S. Department of 

Agriculture, Forest Service, Pacific Northwest Research Station: 79–90.

Wilson and Forsman suggested that opening up the forest canopy by thinning in young stands was detrimental to tree voles because it eliminated 

arboreal pathways between trees and reduced the availability of trees that 

had good structure for nest support.

480.  Witmer, G.W.; deCalesta, D.S. 1986. Resource use by unexploited sympatric bobcats and coyotes in Oregon. Canadian Journal of Zoology. 64: 

2333–2338.

Witmer and deCalesta examined seasonal variation in the frequency of 

occurrence of prey in scats of bobcats and coyotes on Elliott State Forest 

in the Oregon Coast Ranges. Arborimus voles occurred in 0 to 4 percent of 

coyote scats and 0 to 8 percent of bobcat scats. The authors did not attempt 

to differentiate between remains of red tree voles and white-footed voles.

481.  Wooster, T.W. 1994. Red tree vole (Arborimus longicaudus) observations 

in Humboldt, Mendocino, and Sonoma Counties, California 1991–1994. 

23 p. Unpublished report. On file with: California Department of Fish 

and Game, Region 3, Sacramento, CA. (Available upon request from the 

compilers).
144

GENERAL TECHNICAL REPORT PNW-GTR-909

Wooster found 224 Sonoma tree vole nests from 1991 through 1994, during 

surveys of areas slated for harvest in northwestern California. He provided 

no analysis of the data, but made the following assertions: (1) When one 

nest is found in a stand, many others would be found; (2) there was no 

pattern of nest tree selection by size or age; (3) nests were placed in trees 

that received more light; (4) tree voles girdled trees; and (5) tree voles were 

associated with valley bottoms. He also questioned the assertion that tree 

voles are weak dispersers because he found one case where tree voles were 

able to reinvade forests growing on an area that had been harvested. He 

concluded that Sonoma tree voles were quite common in previously logged 

areas and that the California Forest Practices Rules provided sufficient 

protection of tree vole habitat.

482.  Wooster, T.W. 1996. Red tree vole (Arborimus longicaudus) observations 

in Humboldt, Mendocino, and Sonoma Counties, California 1991–1995. 

29 p. Unpublished report. On file with: California Department of Fish and 

Game, Region 3, Sacramento, CA. 

In this wide-ranging discussion regarding habitat and behavior of tree 

voles in California, Wooster concluded by saying that his surveys indicated 

that tree voles were very adaptable and were thriving in numerous younggrowth stands. It is difficult to interpret his results because he provided 

no quantitative comparisons between young and old forests or between 

thinned and unthinned stands. But this uncertainty does suggest the need 

for long-term studies of tree vole persistence and population density in 

managed forests.

483.  Wooster, T.; Town, P. 1998. California red tree voles within a coastal 

second-growth forest. 11 p. Unpublished manuscript. On file with: USDA 

Forest Service, Pacific Northwest Research Station, 3200 Jefferson Way, 

Corvallis, OR 97331. (Available upon request from the compilers)

Wooster handed out this unpublished manuscript at a meeting of tree vole 

biologists at Jackson Demonstration State Forest, California on 3 March 

2005. The paper described a survey of tree voles at one site in Mendocino 

County, California. Wooster and Town suggested that forest capacity to 

hold water could be more important than forest age in providing suitable 

habitat for tree voles.

484.  Wooster, T.; Town, P. 2002. Newly discovered food and habitat use by 

California red tree voles. California Fish and Game. 88: 181–185.
145

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Wooster and Town reported that they found four nests in which tree voles 

were living and feeding on Bishop pine in Mendocino County, California. 

They stated that tree voles that fed on Bishop pine mechanically removed 

the resin ducts in the edges of the needle, just as they do with Douglas-fir. 

Taylor (1915) also observed a Bishop pine cutting in a tree vole nest along 

with grand fir cuttings but did not speculate that tree voles were feeding 

on pine. Stagnaro (2008) also reported evidence of tree voles feeding on 

Bishop pine.

485.  Wright, J.; Wright, D. 1967. Oregon’s unique inhabitant. Oregon 

Holidays. 6: 6.

In this article intended for a lay audience, the authors described the interesting little mouse that lived in the trees, complete with illustrations.

486.  Yensen, E. 1998. Arborimus albipes (Merriam 1901) white-footed vole. 

In: Hafner, D.J.; Yensen, E.; Kirkland, G.L., Jr., eds. Status survey and 

conservation action plan North American rodents. Gland, Switzerland: 

The World Conservation Union: 86.

In this short species account, Yensen suggested that the rarity of whitefooted vole specimens could be due to the difficulty of capturing them. 

The World Conservation Union Red List Category lists the white-footed 

vole as data deficient because so little is known about its distribution and 

abundance.

487.  Zabel, C.J.; McKelvey, K.; Ward, J.P., Jr. 1995. Influence of primary 

prey on home-range size and habitat-use patterns of northern spotted owls 

(Strix occidentalis caurina). Canadian Journal of Zoology. 73: 433–439.

The authors reported that red tree voles made up 36 percent of prey items 

identified in 84 pellets collected from northern spotted owls near the 

Chetco River, Curry County, Oregon. At another study area near Mad 

River, Humboldt County, California, Sonoma tree voles comprised 12 

percent of 581 prey items in spotted owl pellets. No tree voles were located 

in 76 spotted owl pellets collected in the Klamath Mountains, 48 km west 

of Happy Camp, California.

488.  Zahler, P. 1996. Rediscovery of the woolly flying squirrel (Eupetaurus 

cinereus). Journal of Mammalogy. 77: 54–57.

Zahler described the woolly flying squirrel (Eupetaurus cinereus), a species native to northern Pakistan, that appeared to feed primarily on conifer 
146

GENERAL TECHNICAL REPORT PNW-GTR-909

needles. Other than tree voles, this is the only other mammal that we are 

aware of that feeds heavily on needles of conifers.

489.  Zahler, P.; Khan, M. 2003. Evidence for dietary specialization on pine 

needles by the wooly flying squirrel (Eupetaurus cinereus). Journal of 

Mammalogy. 84: 480–486.

Zahler and Khan analyzed fecal samples from four woolly flying squirrels and found that 92 to 100 percent of the fecal material was from pine 

(Pinus spp.) needles. They concluded that the woolly flying squirrel was 

highly specialized for feeding on pine needles. They suggested that free 

water may be a limiting factor for woolly flying squirrels in the arid region 

of northern Pakistan because they assumed that wooly flying squirrels 

required lots of water to counteract and purge the volatile secondary 

compounds in pine needles.

490.  Zentner, P.L. 1977. The nest of Phenacomys longicaudus in northwestern California. Sacramento, CA: California State University. 59 p. M.A. 

thesis.

Zentner reported that he located 53 Sonoma tree vole nests in northwestern 

California from 1973 to 1977. He reported that nests in small (<30 cm dbh) 

Douglas-fir trees were usually located in the upper third of the live crown, 

whereas nests in medium Douglas-firs (30 to 60 cm dbh) were usually 

located in the top half of the crown, and nests in larger Douglas-fir trees 

(>60 dbh) were located throughout the live crown. Zentner described the 

shape and size of nests and concluded that large trees had larger nests. He 

stated that twigs and cuttings in vole nests were 8 to 30 cm long and that 

nest cores consisted of shredded Douglas-fir bark. He also reported that 

14 percent of the inner nest core was the outer edges of fir needles. He 

described the inner chamber as being located in the center of the nest and 

lined with “finely-chewed material.” He said that he documented tree voles 

removing snow from their nests, but he never said how he documented 

this. Zentner said that if he climbed nest trees, the voles would abandon 

their nests for 2 to 7 days or longer, and that just walking to within sight of 

a nest usually caused desertion for 1 to 3 days. We think he could not have 
147

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

documented any of this without dissecting nests to see if they were

actually occupied.

The most perplexing part of this thesis is that Zentner claimed to have 

found tree vole nests much farther inland in northern California than any 

previous researchers. Based on these records, he put forth the hypothesis 

that tree voles in northern California had been forced to expand their range 

eastward in response to logging of their habitat near the coast. Given the 

improbability of a mass exodus of tree voles from logged areas into a 

previously unoccupied region, and the negative results of recent surveys for 

tree voles in the inland regions where Zentner claimed to have found tree 

vole nests, we think a much more likely scenario is that Zentner mistook 

nests of other species for tree vole nests, and tree voles did not occur there 

in the first place. But this needs further investigation. 
148

GENERAL TECHNICAL REPORT PNW-GTR-909

Appendix 1—Number of tree voles (Arborimus longicaudus, A. pomo) or whitefooted voles (A. albipes) captured during trapping studies conducted in western 

Oregon and northwestern California (continued) 

Citationa

Trapnights Trap typeb Tree volecWhite-footed 

volec

8—Anthony et al. (1987) 10,800 ST 0 9 (0.083)

11—Aubry et al. (1991) NRd PF >1 NRd

44—Borrecco (1973) 63,360e LT 1 (0.002) 0

53—Bury and Corn (1987) 38,800 PF 17 (0.044) 0

82—Cole et al. (1998) 50,256 PF, LT 1 (0.002) 1 (0.002)

87—Corn and Bury (1986) 38,800 PF 18 (0.046) 0

88—Corn and Bury (1988) NRd PF 18 0

89—Corn and Bury (1991) 108,864 PF 13 (0.012) 0

90—Corn et al. (1988) NRd PF 17 0

102—Dizney et al. (1985) 65,600 PF, LT 0 0

103—Doyle (1985) 92,904 LT 0 1 (0.001)

104—Doyle (1990) 40,152 LT 0 0

108—Fisher (1942) NRd ST 1 0

110—Fontaine (2007) 8,900 LT 1 (0.011) 0

141—Gilbert and Allwine (1991) 165,310e PF, ST 9 (0.005) 0

145—Gillesberg and Carey (1991) 2,310e LT 0 0

146—Gitzen et al. (2007) 256,896 PF 1 (<0.001) 1 (<0.001)

148—Gomez (1992) 100,800 PF 23 (0.023) 59 (0.059)

149—Gomez and Anthony (1998) 100,800 PF 23 (0.023) 59 (0.059)

150—Gomez et al. (1997) 7,290 PF 4 (0.055) 4 (0.055)

178—Hayes et al. (1996) 14,400 LT 0 1 (0.007)

182—Hooven and Black (1976) 15,750e LT 3 (0.019) 0

208—Jewett (1915) NRd ST 0 1

233—Johnston (2006) 17,600 LT 0 0

257—Maguire (2002) 86,400e PT >1 >1

260—Manning (2002) 23,760e LT 0 0

261—Manning and Edge (2004) 2,400e LT 0 0

262—Manning and Edge (2008) 23,760e LT 0 0

263—Manning and Maguire (1999) 21,168 PT 1 (0.005) 0

264—Manning et al. (2003) NRd PF 0 13

268—Martin (1998) 127,900 PT 42 (0.033) 85 (0.066)

270—Martin and McComb (2002) 127,900 PF 42 (0.033) 85 (0.066)

288—McComb et al. (1993a) 28,800 PF <10 0

289—McComb et al. (1993b) 31,200 PF, ST 2 (0.006) 11 (0.035)

350—Ralph et al. (1991) 141,120 PT, ST, LT 22 (0.016) 0

352—Raphael (1988) 899,431 PF 19 (0.002) 0

366—Rosenberg (2003) 36,000e LT NRd NRd
149

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Appendix 1—Number of tree voles (Arborimus longicaudus, A. pomo) or whitefooted voles (A. albipes) captured during trapping studies conducted in western 

Oregon and northwestern California (continued) 

Citationa

Trapnights Trap typeb Tree volecWhite-footed 

volec

368—Russell (1942) 16e, f ST 8 (50) 0

375—Sherrell (1969) NRd ST 0 1

376—Sherrell (1970) 5,585e ST 0 1 (0.018)

398—Sturges (1955) NRd PF, ST 0 2

399—Suzuki (1992) 19,300 PF, ST 0 14 (0.073)

400—Suzuki (2000) 139,600 PF >1 >1

401—Suzuki and Hayes (2003) 139,600 PF 9 (0.006) 432 

(0.0310)

407—Swingle et al. (2004) 184f LT 3 (1.6) 0

412—Tevis (1956) 3,400e ST 0 0

461—Waldien (2005) 52,800 LT 0 0

467—Waters et al. (2001) NRd PF >1 0

476—Wight (1925) 1e,f ST 1(100) 0

a

 Bibliography number precedes citation.

b

 PF = pitfall trap, ST = snap trap, LT = live-trap. 

c

 Captures per 100 trap-nights are in parentheses and were estimated from the data if not reported in 

the original manuscript. 

d

 NR = not reported.

e

 Trap-nights were estimated based on data presented in the methods section.

f

 Traps were placed on top of nests.
150

GENERAL TECHNICAL REPORT PNW-GTR-909

Subject Index by Bibliography Number

Bolding and underlining indicate species referred to in each reference, as follows:

Bold numbers without underlining = tree voles

Bold numbers with underlining = both tree voles and white-footed voles

Underlined, but not bolded = white-footed voles

Not bolded or underlined = references that do not specifically mention either species.

Arborimus longicaudus silvicola ............7, 15, 27, 43, 48, 51, 67, 68, 72, 80, 132, 140, 164, 165, 166, 176, 185, 186, 193, 

201, 202, 211, 217, 224, 227, 244, 247, 255, 256, 275, 276, 277, 278, 280, 283, 284, 306, 307, 309, 310, 316, 320, 324, 

325, 333, 347, 429, 430, 435, 437, 444, 449, 454, 462, 463, 476, 485

Behavior

Activity patterns ................................................................................................................................................131, 402, 405

Arboreal ..............................................................................................................................59, 264, 279, 410, 453, 454, 458

Climbing/movements ........................14, 15, 28, 80, 85, 131, 186, 217, 277, 284, 319, 320, 331, 334, 374, 408, 410, 454

Female aggression towards male .................................................................................................................80, 170, 218, 224

Foraging ...................................................................................................................................131

Gnawing....................................................................................................................................80

Solitary behavior...............29, 80, 130, 131, 176, 186, 195, 209, 277, 279, 341, 380, 402, 408, 410, 430, 450, 454, 485

Temperament .................................................................................15, 28, 80, 170, 186, 209, 217, 277, 283, 408, 454, 463

Vocalizations ......................................................................................................................................80, 170, 217, 408, 410

Captive voles .................15, 28, 29, 48, 63, 79, 80, 98, 122, 126, 129, 161, 169, 170, 175, 176, 177, 180, 186, 214, 216, 217,

218, 224, 225, 227, 231, 246, 277, 278, 331, 410, 450, 463, 476

Colony/colonial ....................................................................................................... 2, 29, 48, 61, 62, 80, 93, 186, 258, 293, 410

Commensals/inquilines

Amphibians ...........................................................28, 29, 96, 128, 131, 170, 213, 217, 231, 277, 278, 343, 387, 402, 408

Invertebrates .....................................................................4, 13, 28, 80, 128, 170, 186, 217, 231, 277, 408, 471, 472, 473

Conifer needle anatomy and chemistry

Anatomy ................................................... ............................................................................................9, 204, 241, 361, 470

Chemistry ............................................................ ........................................................................................97, 231, 241, 450

Resin ducts ......................................................... ....................................................................9, 29, 167, 176, 241, 361, 470

Court cases .........................3, 21, 71, 91, 92, 183, 228, 280, 318, 336, 337, 339, 340, 431, 434, 435, 438, 439, 383, 441,

442, 443, 444, 447

Current vegetation survey (CVS)/Forest inventory and analysis (FIA) plots .............70, 105, 266, 323, 362, 437, 442, 444, 450

Diseases and injuries .................................................................................................................................................80, 121, 170

Distribution .....................................................2, 7, 8, 19, 29, 33, 34, 44, 49, 50, 53, 56, 58, 59, 63, 65, 66, 67, 68, 77, 80, 82, 

83, 87, 88, 89, 90, 93, 94, 95, 102, 103, 108, 110, 116, 130, 139, 141, 145, 146, 148, 149, 150, 151, 157, 158, 161, 164, 

165, 166, 176, 182, 186, 193, 195, 208, 221, 231, 240, 249, 250, 254, 257, 260, 263, 264, 268, 269, 277, 280, 282, 

283, 284, 288, 289, 304, 307, 309, 313, 319, 324, 325, 341, 342, 345, 350, 352, 368, 375, 376, 380, 384, 394, 398, 399, 

400, 401, 407, 410, 428, 429, 430, 442, 443, 444, 449, 450, 451, 452, 453, 454, 476, 478, 481, 486, 490
151

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Elevation ..................................2, 33, 34, 49, 59, 72, 87, 116, 130, 158, 176, 193, 217, 249, 263, 273, 277, 280, 282, 284, 381,

430, 443, 449, 450, 454, 468, 478

Field notes/lab notes/specimen catalogs ................13, 25, 28, 45, 57, 60, 96, 98, 108, 128, 130, 138, 170, 171, 175, 180, 192,

198, 207, 214, 216, 217, 234, 246, 249, 258, 273, 274, 278, 283, 290, 302, 314, 335, 351, 368, 373, 393, 396, 408, 459, 

468, 469

Fire effects ..................................................72, 110, 125, 130, 133, 195, 217, 231, 265, 277, 280, 431, 442, 444, 447, 449, 450

Food and water consumption

Alnus rubra ..................................................................................................................49, 83, 127, 161, 162, 453, 458, 486

Conifer needles ...............................................................................................13, 14, 15, 28, 29, 34, 50, 62, 63, 65, 72, 80, 

93, 94, 129, 130, 131, 139, 169, 172, 175, 176, 186, 193, 195, 217, 227, 231, 241, 247, 255, 277, 278, 279, 280, 281, 

283, 284, 313, 319, 341, 380, 384, 390, 408, 428, 429, 430, 431, 449, 450, 454, 456, 460, 463, 482, 484

Cuttings harvested per day ...............................................................................................................................................131

Cutting size ..............................................................................................................................131, 231, 279, 283, 284, 410

Eating needles whole ................................................................................................................. 29, 277, 279, 283, 313, 454

Energetics .........................................................................................................................................................231, 291, 292

Exotic foods in captivity ..........................................................15, 28, 29, 80, 167, 170, 186, 200, 217, 277, 410, 449, 463

Lichens .......................................................................................................................................80, 101, 371, 449, 454, 476

Food consumption rate ................................................................................................................. 15, 80, 139, 186, 231, 380

Food preference ...................15, 80, 129, 130, 134, 167, 175, 186, 217, 227, 241, 255, 264, 277, 279, 281, 449, 450, 463

Forbs/shrubs......................49, 59, 83, 95, 161, 162, 172, 187, 250, 264, 277, 279, 282, 283, 341, 453, 454, 458, 478, 486

Mycophagy .......................................................................................................................................................................264

Resin duct removal .....................................2, 28, 29, 80, 167, 169, 170, 176, 186, 209, 217, 231, 277, 279, 283, 284, 450

Roots.................................................................................................................................................................187, 453, 454

Twig bark and cambium ............................2, 80, 93, 94, 130, 131, 132, 133, 134, 139, 159, 167, 176, 186, 195, 209, 217,

231, 241, 277, 278, 279, 281, 283, 284, 319, 449, 454, 476, 484

Water consumption ..........................2, 15, 28, 29, 33, 34, 50, 58, 63, 64, 65, 66, 72, 80, 83, 122, 130, 139, 145, 169, 170,

186, 241, 277, 279, 280, 283, 293, 294, 334, 380, 410, 428, 430, 449, 450, 483

Fog ....................33, 34, 63, 64, 65, 122, 169, 186, 193, 195, 231, 236, 239, 277, 279, 293, 341, 380, 410, 449, 450, 478, 483

Fossil history/evolution.......................1, 22, 26, 41, 72, 73, 74, 75, 84, 106, 160, 176, 221, 224, 231, 242, 244, 271, 272, 277,

279, 283, 304, 355, 356, 357, 358, 360, 369, 370, 395, 450, 453, 454, 464

Genetics and taxonomy...........................................................1, 2, 27, 34, 39, 40, 54, 58, 59, 63, 66, 67, 68, 72, 73, 74, 75, 83, 

84, 99, 135, 139, 140, 160, 161, 164, 176, 184, 185, 186, 189, 195, 217, 220, 221, 222, 224, 229, 231, 235, 242, 243,244, 

255, 256, 277, 279, 284, 295, 296, 303, 304, 305, 306, 307, 308, 309, 310, 311, 315, 316, 320, 355, 356, 357, 363, 380, 

410, 428, 430, 442, 444, 447, 449, 450, 453, 454, 478

Habitat

Barriers .........................................................................................................62, 63, 130, 186, 277, 280, 328, 341, 449, 450

Canopy cover/canopy layering ........................................... 20, 62, 66, 72, 80, 136, 294, 328, 388, 389, 402, 421, 479, 483

Corridors....................................................................................................................................................322, 378, 385, 450
152

GENERAL TECHNICAL REPORT PNW-GTR-909

Crown interconnectivity ........................................................................................ 64, 72, 80, 259, 319, 388, 389, 402, 479

Deciduous forests .....................................................................................................................................148, 149, 162, 458

Forest age

Old forest ..............................................................................................................................................................................

2, 6, 11, 12, 15, 32, 33, 34, 36, 50, 62, 63, 64, 65, 66, 72, 76, 80, 87, 88, 89, 90, 93, 94, 105, 128, 129, 130, 131, 132, 

133, 136, 141, 145, 148, 149, 150, 152, 172, 176, 183, 195, 196, 199, 217, 231, 236, 237, 279, 281, 283, 289, 293, 294, 

297, 328, 341, 352, 380, 402, 413, 415, 416, 425, 429, 430, 431, 433, 435, 442, 443, 444, 449, 450, 454, 479, 481

Young forest .......................................... 6, 11, 15, 32, 33, 34, 36, 44, 50, 62, 63, 65, 66, 72, 76, 80, 87, 105, 120, 125, 128, 129, 

130, 131, 132, 133, 155, 171, 172, 176, 185, 186, 193, 195, 196, 206, 211, 217, 231, 236, 273, 277, 279, 280, 281, 283, 

288, 293, 294, 341, 352, 380, 402, 415, 417, 425, 430, 431, 435, 442, 444, 449, 450, 454, 462, 481, 482, 483

Forest composition

Abies concolor ....................................................................................................................................................... .263, 430

Abies grandis .................... 2, 15, 29, 33, 34, 58, 60, 61, 94, 128, 130, 131, 139, 145, 167, 170, 176, 196, 217, 224, 236,

250, 277, 283, 284, 314, 334, 380, 408, 410, 411, 430, 449, 450, 454, 478, 484

Alnus rubra .................................................................................59, 83, 127, 250, 264, 277, 279, 283, 341, 454, 458, 478

Corylus cornuta ......................................................................................................................................................162, 264

Picea sitchensis ....................2, 12, 15, 33, 34, 43, 58, 80, 72, 93, 129, 130, 134, 139, 176, 201, 209, 217, 224, 231, 236,

247, 250, 277, 279, 280, 283, 284, 319, 380, 410, 411, 430, 449, 450, 463, 478, 484, 485

Pinus muricata .................................................................................................................. 61, 130, 170, 334, 408, 410, 484

Pseudotsuga menziesii ...................................... 2, 15, 28, 29, 33, 34, 43, 58, 72, 76, 80, 93, 94, 128, 129, 130, 131, 132,

139, 145, 152, 167, 170, 172, 176, 193, 196, 201, 209, 211, 217, 224, 231, 236, 237, 247, 250, 277, 279, 280, 283, 

284, 293, 294, 314, 319, 334, 341, 380, 384, 388, 389, 408, 410, 411, 413, 418, 420, 430, 449, 450, 454, 460, 478, 

484, 485, 490

Sequoia sempervirens ...........................................28, 29, 76, 138, 139, 170, 196, 234, 236, 277, 283, 319, 334, 380, 484

Tsuga heterophylla ....................... 2, 12, 15, 33, 34, 43, 58, 72, 80, 94, 129, 130, 132, 134, 176, 201, 217, 224, 231, 236,

247, 277, 279, 280, 283, 284, 380, 430, 449, 450, 454, 462, 463, 478, 484, 485

Habitat associations ..................................................2, 6, 11, 12, 32, 33, 34, 36, 58, 62, 63, 64, 65, 66, 87, 89, 90, 93, 94, 105,

130, 141, 148, 149, 152, 155, 176, 194, 195, 196, 199, 231, 236, 237, 251, 264, 266, 268, 270, 280, 283, 288, 289, 293, 294, 

297, 380, 413, 415, 416, 417, 425, 429, 431, 433, 435, 442, 443, 449, 450, 454, 479, 483

Habitat connectivity ............................................................................................... 194, 265, 415, 425, 431, 435, 442, 443, 444

Habitat threats ................... 2, 11, 12, 28, 33, 34, 58, 59, 62, 63, 65, 66, 72, 80, 88, 89, 90, 95, 125, 130, 132, 152, 155, 161, 

176, 193, 195, 202, 217, 231, 251, 252, 265, 273, 277, 280, 281, 283, 324, 325, 341, 352, 377, 380, 413, 414, 425, 428, 429, 

430, 431, 444, 449, 450, 478

Poison oak (Toxicodendron diversilobum) in nest trees....................................................................................170, 212, 217, 227

Riparian forest.......................... 8, 59, 83, 95, 148, 149, 161, 178, 185, 186, 187, 208, 240, 250, 264, 277, 279, 281, 282, 283,

284, 313, 341, 384, 398, 399, 452, 453, 454, 458, 471, 486

Streams ................................................................................................................................................7, 15, 49, 51, 83, 158

Thinning effects ...........................................................................................20, 72, 125, 259, 388, 442, 444, 447, 450, 479
153

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Juvenile development

Activity outside nest ........................................................................................................................131, 277, 279, 430, 450

Anal perforation ............................................................................................................................................................... .80

Birth weight/juvenile growth rate. ......................................................... 2, 80, 109, 169, 170, 186, 214, 217, 248, 299, 300

Climbing behavior ...........................................................................................................131, 180, 186, 211, 279, 283, 463

Coprophagy ............................................................................................................................................. 169, 241, 277, 334

Dispersal ..................................................................................................................... 15, 34, 131, 279, 280, 283, 402, 450

Eyes open ................................................................................................................................................ 2, 15, 80, 169, 277

Infanticide ................................................................................................................................................................217, 277

Maternal care .....................................................................................................................................15, 28, 29, 50, 80, 169

Nursing ............................................................................................................................................................................... 170

Pelage development ................................................................................................................................................... 48, 80

Sexual maturity ......................................................................................................................................................... 80, 170

Solid food consumption .............................................................................................................................. 29, 80, 169, 277

Weaning .....................................................................................................................2, 50, 72, 80, 169, 221, 277, 320, 430

Management

Conservation status .............................58, 76, 83, 93, 94, 95, 155, 161, 202, 237, 238, 239, 266, 324, 325, 327, 328, 329,

333, 334, 341, 367, 414, 437, 438, 448, 449, 453, 454, 478, 484, 486

Distinct population segment (DPS) ............................................................................... 3, 72, 130, 215, 256, 265, 449, 450

Endangered Species Act (ESA) ............................................................................................. 3, 72, 215, 256, 265, 449, 450

Federal Land Policy and Management Act (FLPMA) .......................................................................... 3, 318, 443, 444, 447

Federal tree vole database

GeoBob (Geographic Biotic Observations) ............................................................... 130, 195, 266, 442, 443, 444, 447

Interagency species management system (ISMS) ........................................... 34, 72, 432, 435, 438, 442, 444, 445, 446

NRIS (Natural Resource Information System....................................................................................................130, 195

Forest Ecosystem Management Assessment Team (FEMAT) ....................72, 266, 280, 415, 425, 430, 431, 432, 434, 435,

438, 443, 447

Habitat conservation plan .................................................................................................. 76, 196, 197, 266, 323, 332, 389

Interagency Scientific Committee (ISC) ..........................................................................................................................413

Management recommendations .............................58, 59, 76, 83, 152, 193, 195, 196, 199, 280, 341, 428, 429, 430, 431,

433, 434, 435, 437, 438, 439, 440, 442, 444, 447, 478

National Environmental Policy Act (NEPA) ....72, 266, 318, 336, 337, 339, 422, 431, 434, 438, 439, 442, 443, 444, 447

National Forest Management Act (NFMA) ............................................................ 193, 422, 426, 431, 434, 439, 442, 444

Non-high-priority sites/high-priority sites ..................................................... 3, 71, 195, 266, 340, 429, 436, 438, 443, 446

Northwest Forest Plan ..........................18, 21, 34, 71, 72, 91, 92, 105, 130, 195, 206, 237, 280, 312, 318, 323, 333, 336,

337, 339, 383, 389, 422, 423, 424, 426, 429, 430, 431, 432, 433, 434, 435, 437, 438, 439, 440, 441, 443, 444, 447, 

449, 450
154

GENERAL TECHNICAL REPORT PNW-GTR-909

Oregon state forests ..................................................................................................................72, 130, 133, 202, 328, 382

Pechman Exemption .................................................................................................................................................92, 338

Salvage logging ........................................................................................................72, 257, 266, 415, 423, 424, 426, 444

Scientific Analysis Team (SAT) ...............................................................................................414, 425, 426, 430, 431, 432

Sensitive species program ...............................................................................................................................432, 438, 439

Special status species........... 72, 215, 266, 333, 336, 337, 339, 422, 435, 437, 438, 439, 440, 441, 442, 444, 446, 447, 449

Species review process/annual species review ............................................... 318, 339, 432, 434, 437, 438, 443, 444, 447

Strategic surveys .......................................35, 105, 183, 206, 266, 312, 333, 431, 432, 434, 435, 438, 442, 443, 444, 445

Survey and Manage Program ..............................3, 18, 21, 33, 34, 35, 71, 72, 91, 92, 105, 130, 183, 195, 215, 237, 266, 

280, 312, 318, 323, 333, 336, 337, 339, 362, 383, 421, 422, 423, 424, 426, 427, 429, 430, 431, 432, 433, 434, 435, 437, 

438, 439, 440, 441, 442, 443, 444, 445, 446, 447, 449, 450

Morphology

Anatomy ........................................................................................................................................................... 99, 100, 221

Baculum .................................................................................................................. 100, 168, 176, 221, 229, 278, 453, 469

Bobtailed specimens............................................................................................................................................... 170, 217

Cecum/cecal villi ................................................................................................................ 2, 221, 229, 354, 453, 456, 457

Descriptions .................................................. 2, 7, 14, 15, 43, 51, 77, 80, 83, 208, 240, 247, 279, 282, 283, 304, 342, 392

Dwarf specimens ............................................................................................................................................. 170, 217, 277

Glands ..................................................................................................................... 176, 181, 221, 229, 346, 348, 349, 454

Isabellinism ..................................................................................................................................... 278, 304, 410, 449, 450

Mandible ............................................................................................................................................................................. 176

Manus ..................................................................................................................................... 176, 227, 231, 244, 391, 410

Melanism .......................................................................................................................... 72, 175, 176, 177, 402, 449, 450

Molt ..................................................................................................................................... 28, 80, 171, 277, 278, 410, 450

Myology .......................................................................................................................................... 227, 231, 242, 243, 244

Pelage .......................2, 7, 14, 15, 28, 34, 43, 48, 51, 63, 66, 72, 80, 83, 132, 139, 164, 165, 166, 170, 171, 175, 176, 185,

186, 250, 279, 283, 284, 285, 296, 304, 319, 320, 334, 342, 392, 394, 402, 410, 418, 420, 428, 430, 449, 450, 453, 

454, 485

Size/mass ............................................................................................................................2, 7, 15, 34, 43, 45, 51, 63, 66, 

72, 80, 83, 93, 94, 95, 164, 165, 166, 170, 171, 176, 186, 211, 224, 250, 278, 279, 282, 283, 284, 300, 303, 304, 310, 

314, 320, 345, 375, 392, 396, 402, 408, 418, 420, 428, 430, 449, 450, 454, 462, 463, 468, 469

Skull ....................... 80, 165, 166, 176, 184, 185, 186, 221, 224, 229, 231, 242, 243, 284, 296, 304, 347, 394, 410, 418,

420, 449, 452, 453, 454

Teeth ................. 2, 43, 74, 80, 160, 176, 184, 185, 186, 200, 217, 224, 229, 231, 242, 243, 271, 277, 303, 304, 319,

320, 342, 357, 358, 360, 394, 410, 418, 420, 449, 450, 452, 453, 454

Testes ......................................................................................170, 171, 217, 221, 229, 263, 278, 282, 284, 314, 375, 402

Vibrissae .........................................................................................................................................170, 214, 410, 418, 420
155

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Movements

Dispersal 2, 31, 34, 62, 64, 72, 87, 176, 193, 194, 328, 334, 402, 415, 425, 428, 430, 442, 443, 444, 447, 449, 450, 454, 

479, 481

Home range ................................34, 47, 50, 72, 87, 170, 172, 277, 341, 402, 404, 405, 410, 428, 430, 442, 444, 449, 450

Non-forest movements .........................................................................................31, 87, 402, 405, 428, 430, 449, 450, 454

Terrestrial movements ......... 28, 29, 31, 34, 63, 66, 72, 80, 159, 167, 176, 186, 195, 319, 350, 402, 405, 410, 415, 420,

428, 430, 431, 442, 450, 454, 478

Nests

Arboreal nests of white-footed voles.................................................................................................................. 127, 133, 195

Branch/bole structure .....................2, 12, 28, 30, 33, 50, 63, 65, 66, 72, 80, 128, 129, 132, 134, 137, 138, 144, 155, 183,

195, 196, 205, 217, 236, 247, 258, 273, 277, 283, 298, 402, 430, 450, 476, 479 

Composition and construction ............................... 2, 13, 14, 15, 28, 34, 45, 48, 58, 64, 65, 66, 72, 80, 105, 127, 128, 129, 

130, 131, 132, 133, 134, 138, 139, 144, 145, 152, 155, 167, 171, 175, 176, 186, 195, 200, 209, 211, 217, 231, 241, 

246, 247, 273, 277, 278, 279, 280, 281, 283, 284, 293, 313, 314, 319, 320, 334, 341, 343, 362, 371, 374, 380, 

384, 390, 402, 405, 408, 410, 421, 426, 430, 443, 449, 450, 454, 459, 468, 478, 481, 482, 483, 484, 490

Detectability ............... 29, 72, 80, 105, 129, 132, 139, 145, 167, 185, 186, 217, 231, 236, 277, 280, 317, 383, 402, 410,

450, 481, 483

Fidelity/desertion ......................................................................................131, 170, 231, 277, 280, 393, 410, 418, 454, 490

Height .....................2, 7, 14, 15, 28, 34, 48, 50, 51, 58, 60, 63, 65, 66, 80, 93, 94, 131, 139, 159, 170, 171, 175, 176, 195,

217, 231, 247, 273, 277, 281, 283, 284, 313, 319, 341, 342, 343, 374, 402, 408, 411, 421, 430, 478

Nest predation/destruction .................................................................................................80, 154, 170, 231, 277, 410, 490

Nest tree girdling .....................................................................................................................................................410, 481

Nest types

Branch whorls ..................14, 15, 29, 63, 72, 128, 170, 183, 195, 196, 211, 217, 231, 236, 277, 278, 279, 283, 402, 410,

421, 428, 463, 479

Broken tops ....................... 72, 80, 120, 128, 130, 132, 133, 134, 144, 145, 155, 175, 183, 186, 195, 217, 231, 277, 278,

283, 383, 402, 405, 430, 476

Cavity nests ...... 30, 63, 64, 65, 128, 129, 132, 134, 144, 145, 155, 183, 195, 196, 217, 231, 236, 277, 343, 402, 430, 462

Dwarf mistletoe (Arceuthobium spp.) brooms ............................................72, 134, 155, 170, 195, 280, 402, 428, 430, 479

Epicormic branches/palmate branches .....................................................................120, 132, 133, 402, 405, 428, 430, 479

Feeding platforms ................................................................................................................ 170, 231, 273, 454

Forked trunks ............................ 72, 120, 130, 133, 155, 171, 175, 183, 195, 196, 217, 231, 236, 277, 402, 405, 430, 479

Hollow limbs ..........................................................................................................................................144, 145, 277, 383

Sloughing bark ....................................................................................................................................................... 170, 402

Moss covered limbs ........................................................................................................................................... 64, 65, 134

Secondary leaders/multiple tops .................................................................... 120, 133, 195, 196, 217, 231, 236, 278, 402

Snags .............................................................................................................. 170, 217, 231, 273, 277, 377, 402, 462, 477

Structures built by other species ..................... 2, 15, 28, 29, 63, 66, 76, 80, 132, 167, 176, 186, 195, 200, 231, 273, 277,

279, 283, 284, 313, 314, 319, 334, 374, 380, 402, 410, 454
156

GENERAL TECHNICAL REPORT PNW-GTR-909

Nests next to bole .............................. 2, 14, 29, 48, 80, 144, 145, 170, 186, 192, 209, 211, 231, 236, 249, 277, 278, 279,

280, 283, 313, 319, 402, 410, 417, 421, 428, 454, 463, 479

Nests on limbs away from bole .................... 2, 29, 63, 80, 145, 170, 186, 196, 209, 217, 231, 236, 277, 278, 280, 283,

313, 319, 402, 410, 417, 421, 428, 454

Terrestrial nests ......................... 2, 7, 15, 56, 58, 63, 66, 87, 139, 145, 155, 186, 225, 231, 277, 279, 320, 341, 380, 417

Persistence/longevity ..................................................................................................................... 155, 277, 380, 417, 430

Placement ................. 2, 29, 50, 63, 66, 80, 128, 144, 145, 170, 176, 209, 217, 231, 236, 258, 273, 277, 279, 283, 284,

293, 313, 380, 402, 410, 454, 460, 483, 490

Relationship to bole ................................................................................................................ 80, 186, 196, 293, 294, 402

Slope aspect at nest site ........................................................................................................... 80, 105, 196, 277, 293, 294

Sexual differences in nest structure ............... 28, 29, 50, 139, 167, 170, 186, 217, 231, 258, 277, 280, 319, 402, 410, 450

Size ........................ 2, 13, 14, 28, 48, 51, 80, 138, 139, 159, 170, 171, 175, 176, 200, 211, 217, 231, 247, 273, 277, 280,

313, 319, 380, 402, 408, 410, 411, 421, 454

Spatial distribution of nests .................................................................................................... 2, 6, 80, 231, 280, 380, 410

Tree species used for nesting

Abies grandis ........................2, 15, 29, 77, 128, 131, 139, 170, 176, 186, 193, 196, 277, 281, 284, 402, 408, 409, 410,

449, 450, 454, 460, 478, 481, 482, 483

Acer macrophyllum ............................................................................................... 176, 217, 277, 402, 449, 450, 481, 482

Alnus rubra ................................................................................................................................................... 277, 481, 482

Arbutus menziesii ...................................................................................................................................... 2, 196, 314, 490

Castanopsis chrysophylla ............................................................................................................................. 402, 449, 450

Notholithocarpus densiflorus ................................................................................................ 155, 196, 417, 460, 481, 482

Picea sitchensis ... 2, 15, 43, 51, 72, 77, 120, 127, 133, 139, 175, 176, 186, 193, 209, 217, 277, 278, 279, 280, 281, 283, 

284, 319, 409, 449, 450, 454, 478, 485

Pinus muricata ..................................................................................................................................................................484

Pseudotsuga menziesii .........................................................................................2, 13, 14, 15, 28, 29, 43, 48, 51, 72, 77,

80, 96, 120, 128, 130, 131, 132, 133, 138, 139, 145, 155, 170, 171, 175, 176, 183, 186, 192, 193, 196, 200, 209, 210, 

211, 212, 217, 234, 236, 247, 249, 258, 259, 273, 277, 278, 279, 281, 283, 284, 293, 294, 313, 314, 319, 343, 368, 

374, 386, 393, 402, 407, 409, 417, 418, 420, 421, 428, 449, 450, 454, 459, 460, 468, 469, 476, 478, 481, 482, 483, 

485, 490

Quercus agrifolia .............................................................................................................................................................170

Quercus wislizenii ............................................................................................................................................................196

Sequoia sempervirens ................................................................2, 28, 29, 96, 170, 196, 217, 277, 319, 460, 481, 482, 483

Taxus brevifolia ........................................................................................................................................277, 402, 449, 450

Thuja plicata .....................................................................................................................................................................217

Tsuga heterophylla ....................2, 43, 51, 72, 120, 133, 175, 176, 193, 217, 277, 278, 279, 280, 281, 283, 284, 402, 449,

450, 454, 460, 462, 478, 485

Umbellularia californicus ..............................................................................................................................170, 196, 460
157

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Parasites

Cestodes/Nematodes ......................................................................................................2, 29, 80, 173, 174, 277, 354, 455

Fleas .....................................................................................................................2, 80, 176, 190, 191, 253, 277, 453, 490

Hemoflagellates ...............................................................................................................................................................80

Mange ..............................................................................................................................................................................29

Mites ..................................................................................2, 28, 29, 42, 80, 107, 163, 170, 176, 277, 278, 471, 472, 473

Ticks .....................................................................................................................................................2, 80, 170, 176, 277

Phenacomys intermedius ....... 27, 74, 77, 84, 88, 159, 173, 200, 201, 221, 223, 231, 242, 277, 295, 347, 357, 358, 370

Photographs/drawings ...........................................................................................................................61, 81, 372, 386, 390

Population parameters

Density estimates....................................................................................................................................................231, 277

Limiting factors ........................................................................................................................63, 64, 66, 72, 93, 231, 328

Longevity ........................................................................................................................................................80, 219, 231

Nest density/abundance..............................................28, 32, 34, 37, 72, 155, 196, 236, 237, 293, 380, 417, 429, 430, 436

Regional relative abundance ......................................................................111, 115, 116, 119, 130, 209, 413, 442, 444, 450

Sex ratios

Adults .....................................................................................................29, 80, 87, 129, 130, 186, 231, 277, 402, 409, 410

Juveniles ............................................................................................................................80, 130, 186, 231, 277, 402, 410

Survival rates ...................................................................................................................................402, 406, 442, 444, 450

Viability (persistence) ......................................................................414, 415, 423, 425, 430, 431, 435, 438, 443, 445, 450

Predators and predation

Birds......................2, 12, 23, 24, 34, 46, 50, 52, 65, 66, 72, 93, 94, 111, 112, 113, 114, 115, 116, 117, 118, 119, 121, 139,

153, 154, 155, 176, 186, 200, 231, 276, 277, 279, 281, 283, 284, 301, 341, 359, 366, 368, 380, 381, 402, 406, 410, 

413, 414, 417, 429, 430, 431, 433, 438, 444, 449, 450, 454, 465, 466, 474, 475, 487

Mammals ................5, 12, 15, 34, 72, 78, 80, 147, 176, 186, 200, 231, 277, 279, 283, 321, 379, 380, 402, 406, 410, 419,

430, 449, 450, 454, 480

Snakes......................................................................................................................................................402, 406, 449, 450

Escape behavior.......................10, 13, 14, 28, 63, 65, 66, 80, 121, 131, 132, 138, 170, 175, 176, 186, 206, 209, 211, 217,

246, 258, 273, 277, 278, 279, 283, 314, 319, 331, 334, 374, 410, 454, 469, 481, 482

Protein electrophoresis ..............................................................................................................74, 217, 220, 221, 449, 450

Resin ducts

Abies grandis ..........................................................................................................................................131, 195, 277, 410

Picea sitchensis .........................................................................................................................................80, 241, 320, 470

Pinus muricata ................................................................................................................................................390, 482, 484

Pseudotsuga menziesii .............9, 14, 15, 28, 29, 45, 56, 58, 64, 66, 72, 80, 105, 128, 131, 132, 133, 138, 144, 145, 152,

155, 167, 171, 175, 176, 186, 195, 200, 201, 209, 211, 217, 231, 241, 247, 255, 273, 277, 278, 279, 281, 283, 284, 293, 

313, 314, 319, 320, 334, 341, 362, 374, 384, 402, 421, 430, 443, 449, 450, 454, 478, 481, 482, 483, 484, 490
158

GENERAL TECHNICAL REPORT PNW-GTR-909

Tsuga heterophylla .....................................................................................72, 80, 133, 134, 195, 241, 255, 361, 380, 450

Recolonization/reintroduction .......................................................................................199, 231, 245, 277, 417, 431, 450

Reproduction

Age at first breeding .................................................................................................................................................80, 170

Breeding season ................ 15, 28, 29, 34, 50, 63, 65, 66, 72, 80, 83, 93, 94, 95, 132, 139, 186, 200, 201, 277, 279, 283,

284, 319, 320, 341, 384, 402, 410, 411, 428, 430, 449, 450, 454

Captive breeding .............................................................................................. 80, 169, 170, 186, 216, 217, 218, 224, 380

Corpora luteum .....................................................................................................................................169, 170, 231, 278

Cross fostering .................................................................................................................................................28, 217, 459

Delayed implantation ........................................................................................................................................... 2, 55, 169

Embryos ................................................................... 96, 165, 170, 217, 231, 277, 278, 282, 314, 368, 396, 454, 459, 469

Estrous .................................................................................................................................................2, 80, 169, 170, 320

Gestation period ................................... 2, 34, 50, 63, 65, 66, 72, 80, 93, 94, 169, 170, 201, 221, 299, 300, 320, 428, 430

Hybrids ................................................................................................................................2, 80, 217, 218, 224, 227, 344

Intromission ....................................................................................................................................................................170

Lactation .................................................................................................... 55, 96, 169, 170, 175, 217, 278, 402, 459, 469

Litter frequency.........................................................................................................................................................80, 131

Litter size ........................2, 7, 15, 47, 49, 50, 63, 65, 66, 80, 83, 93, 94, 95, 109, 131, 139, 159, 165, 166, 169, 170, 200,

201, 210, 221, 229, 246, 248, 277, 279, 283, 284, 299, 300, 319, 320, 341, 342, 377, 396, 410, 411, 428, 430, 449, 

450, 453, 454

Mating behavior ..................................................................................................................... 2, 28, 80, 131, 169, 170, 217

Nulliparous.........................................................................................................................................................................170

Ovaries ...................................................................................................................................................................170, 276

Overlapping litters in nest .............................................................. 131, 169, 170, 186, 231, 277, 279, 283, 284, 341, 450

Ovulation ...........................................................................................................................................................2, 169, 341

Placental scars ........................................................................................................................................................ 170, 231

Post-partum breeding ..............................................2, 48, 50, 65, 66, 72, 80, 131, 169, 176, 279, 283, 334, 449, 450, 454

Uterus ............................................................................................................................................................... 80, 170, 217

Vaginal cornification.................................................................................................................................................... 2, 169

Sampling methods

Arboreal live-trapping..................................................................................................................................................... 407

Cameras/video ............................................................................................................................................... 131, 286, 287

Distance sampling .................................................................................................................32, 36, 37, 133, 236, 237, 429

Dynamite ...................................................................................................................................................15, 231, 277, 462

Fixed area plots ...............................................................................................................................................277, 417, 483
159

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Individual tree sampling .........................................................................105, 130, 134, 142, 143, 144, 145, 183, 195, 362

Live-trapping ....................38, 44, 102, 103, 104, 110, 130, 145, 150, 176, 178, 182, 227, 233, 260, 261, 262, 282, 284,

350, 366, 453, 454, 461, 465, 466

Modeling ................ 20, 70, 94, 95, 105, 179, 193, 194, 236, 266, 328, 330, 388, 389, 421, 436, 442, 444, 445, 450, 464

Nest dissection in tree/dislodging nest from ground ................17, 170, 186, 231, 249, 273, 277, 278, 410, 469, 481, 482

Nest probing ..................................................................................... 39, 128, 130, 132, 133, 195, 402, 405, 406, 407, 450

Pitfall trapping ................11, 53, 62, 72, 82, 86, 87, 88, 89, 90, 93, 102, 129, 130, 132, 141, 145, 146, 148, 149, 150, 176,

193, 257, 263, 264, 268, 269, 270, 288, 289, 350, 351, 352, 380, 399, 400, 401, 413, 429, 450, 454, 467, 478, 486

Predisturbance surveys .............18, 21, 33, 34, 72, 92, 105, 130, 195, 266, 280, 312, 317, 318, 323, 339, 424, 427, 429,

430, 431, 432, 433, 434, 435, 437, 438, 442, 443, 444, 445, 446, 447, 449

Radio-telemetry ............................................................31, 72, 93, 402, 403, 404, 405, 415, 428, 442, 444, 446, 449, 450

Random double sampling ..........................................................................72, 105, 130, 312, 362, 437, 442, 443, 444, 447

Road kill ................................................................................................................................................................... 25, 394

Road surveys........................................................................................................................................................... 120, 130

Sawing off limbs .................................................................................................................................................... 217, 258

Shooting nests .................................................................................................................................. 28, 217, 258, 273, 277

Snap-trapping ...................... 8, 13, 14, 15, 78, 82, 108, 130, 141, 176, 184, 186, 187, 192, 208, 209, 212, 217, 234, 277,

282, 284, 289, 302, 350, 368, 375, 376, 399, 408, 412, 453, 454, 458, 476, 478

Transect surveys ...................6, 32, 33, 34, 35, 36, 37, 66, 72, 76, 93, 105, 130, 133, 145, 155, 195, 196, 236, 317, 362,

380, 417, 421

Tree climbing ......................................................................2, 6, 10, 13, 14, 15, 28, 29, 33, 34, 35, 39, 45, 60, 66, 76, 80,

85, 96, 98, 105, 120, 123, 124, 129, 130, 131, 132, 133, 134, 138, 152, 170, 175, 183, 191, 192, 195, 196, 206, 209, 

211, 212, 217, 227, 246, 249, 277, 278, 283, 284, 293, 294, 313, 334, 362, 374, 402, 407, 408, 421, 437, 450, 469, 

476, 483

Tree felling ...........13, 14, 15, 16, 17, 62, 63, 66, 80, 93, 124, 129, 130, 142, 143, 144, 145, 186, 196, 217, 278, 418, 463
160

GENERAL TECHNICAL REPORT PNW-GTR-909

Author Index by Bibliography Number

Abramson, N.I. ..........................................................................................................1

Adam, M.D. ......................................................................................................2, 178

Adams, D.M. .........................................................................................................388

Ager, T.A. ..............................................................................................................357

Aiken, A. ...................................................................................................................3

Albert, D.L. ...........................................................................................................179

Alegria, J. ..............................................................................................................362

Alexander, C.P. ..........................................................................................................4

Alexander, L.F. ..........................................................................................................5

Allan, W.G.S. .......................................................................................................451

Allwine, R. ............................................................................................................141

Alton, C. ........................................................................................................238, 239

Anderson, P.D. ......................................................................................................479

Anderson, S. ............................................................................................................67

Andrews, L.S. ...................................................................................................6, 130

Anthony, H.E. ...........................................................................................................7

Anthony, R.G. ... .8, 12, 114, 115, 116, 149, 150, 178, 251, 289, 366, 380, 406, 414, 

475

Apple, M. ..................................................................................................................9

Arbogast, B.S. .........................................................................................................40

Ash, S. .....................................................................................................................10

Ashton, P.M.S. ......................................................................................................361

Aubry, K.B...................................................11, 12, 89, 141, 144, 193, 252, 350, 367

Audubon Society of Portland ..............................................................72

Bailey, V.O. ...........................................................................................13, 14, 15, 16

Bake, D.B. ...............................................................................................................17

Baker, D. .................................................................................................................18
161

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Baker, R.H. ..............................................................................................................19

Baker, R.J. .............................................................................................................235

Bannikova, A.A. ........................................................................................................1

Barbour, R.J. ...........................................................................................................20

Barnard, J. ...............................................................................................................21

Barnosky, A.D. ........................................................................................................22

Barrows, C.W. ...................................................................................................23, 24

Behle, W.H. .............................................................................................................25

Bell, C.J. ............................................................................................................22, 26

Bellinger, M.R. ................................................................................................27, 309

Benirschke, K. .......................................................................................................189

Benson, S.B. ......................................................................................................28, 29

Benton Forest Coalition ..........................................................................................71

Berlyn, G.P. ...........................................................................................................361

Bern, H.A. .....................................................................................................456, 457

Bigger, D. ................................................................................................................76

Bigley, R.E. .............................................................................................................93

Bingham, B.B. .........................................................................................................30

Biswell, B.L. ...................................................12, 31, 32, 33, 34, 35, 36, 37, 66, 130

Black, H.C. ......................................................................................38, 182, 377, 477

Blakesley, J.A. .........................................................................................................93

Blois, J.L. ....................................................................................................39, 40, 41

Blow, M. ............................................................................................................33, 34

Bochkov, A.V. .........................................................................................................42

Bolster, B.C. ............................................................................................................83

Bond, B.J. ..................................................................................................................9

Bond, M.L. ............................................................................................................199

Booth, E.P. ...............................................................................................................43
162

GENERAL TECHNICAL REPORT PNW-GTR-909

Borell, A.E. .............................................................................................................29

Borman, B.T. .........................................................................................................266

Borrecco, J.E. ..........................................................................................................44

Boscolo, M.E. .......................................................................................................204

Bradley, R.D. .........................................................................................................235

Breckel, R. ......................................................................................................33, 362

Brock, E.M. .............................................................................................................45

Brouwers, E.M. .....................................................................................................357

Brown, E.R. ...............................................................................................46, 47, 117

Brown, L.N. ............................................................................................................48

Brunet-Lecomte, P. ..................................................................................................73

Brylski, P. ..........................................................................................................49, 50

Burnett, K.M. ........................................................................................................322

Burt, W.H. ...............................................................................................................51

Burton, K. ................................................................................................................52

Bury, R.B. .................................................................................53, 86, 87, 88, 89, 90

Buzan, E.V. ..............................................................................................................54

Buzzio, O.L. ............................................................................................................55

Cahalane, V.H. ........................................................................................................56

California Academy of Sciences .............................................................................57

California Fish and Game .................................................................................58, 59

Cameron, I.R. ........................................................................................................361

Camp, C.L. ........................................................................................................60, 61

Carey, A.B. ....11, 24, 62, 63, 64, 65, 66, 89, 141, 142, 143, 144, 145, 252, 350, 367

Carleton, M.D. ..........................................................................................67, 68, 316

Carraway, L.N. ................................................................................69, 452, 453, 454

Carreno, N.B. ..........................................................................................................55

Carroll, C. ................................................................................................................70
163

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Carter, L.D. ...........................................................................................................357

Cascadia Wildlands ..........................................................................................71,172

Castor, L.K. ...........................................................................................................472

Castro-Vazquez, A. ..................................................................................................55

Center for Biological Diversity .........................................................................71, 72

Chaline, J. ..........................................................................................................73, 74

Chambers, C.L. ...............................................................................................82, 288

Chamblin, D. ...........................................................................................................37

Chapman, J.A. ...............................................................................................226, 344

Chavez, A.S. ............................................................................................................75

Chinnici, S.J. ...........................................................................................................76

Christensen, J.R. .....................................................................................................77

Christy, J. ......................................................................................................238, 239

Chrostowski, H. ....................................................................................................341

Clark, L.A. ............................................................................................................203

Clifton, P.L. .................................................................................................78, 79, 80

Clothier, C. ..............................................................................................................81

Cockrum, E.L. .......................................................................................................165

Cody, M.L. ..............................................................................................................93

Cole, E.C. ................................................................................................................82

Collins, P.W. ............................................................................................................83

Conroy, C.J. .....................................................................................................84, 269

Cook, J.A. .......................................................................................................84, 269

Coriell, T.R. .............................................................................................................85

Corn, P.S. ...................................................................................53, 86, 87, 88, 89, 90

Coughenour, J.C. ...............................................................................................91, 92

Courant, F. ...............................................................................................................73

Courtney, S.P. ..........................................................................................................93
164

GENERAL TECHNICAL REPORT PNW-GTR-909

Cox, J. ...................................................................................................................378

Crites, M.J. ..............................................................................................................11

Cromack, K., Jr. ....................................................................................................136

Csuti, B. ....................................................................................................94, 95, 330

Dalquest, W.W. ........................................................................................................96

Dash, J.A. ................................................................................................................97

Davis, R.J. .....................................................................................................130, 195

Davis, W.B. .............................................................................................................98

Dearden. L.C. ..................................................................................................99, 100

deCalesta, D.S. ......................................................................................................480

Denison, W.C. ...............................................................................................101, 136

Diller, L.V. .............................................................................................................417

Dinnie, C.E. ...........................................................................................................205

Dizney, L. ..............................................................................................................102

Dodd, N.L. ............................................................................................................380

Doyle, A.T. ............................................................................................103, 104, 294

Doyle, E. ...............................................................................................................238

Dumbacher, J.P. .................................................................................................70, 93

Dunk, J.R. .......................................................................................................70, 105

Dyrness, C.T. .........................................................................................................283

Edge, W.D. ............................................................................................261, 262, 330

Edwards, T. ...........................................................................................................362

Eisenberg, J.F. .......................................................................................................106

Engstrom, M.D. .....................................................................................................235

Fain, A. ..................................................................................................................107

Farrell, T.P. ................................................................................................................5

Feldhamer, G.A. ............................................................................................226, 344

Finley, L. .....................................................................................................33, 34, 35
165

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Fisher, H.I. ............................................................................................................108

Fleischer, R.C. .........................................................................................................93

Fleming, T.H. ........................................................................................................109

Fontaine, J.B. ........................................................................................................110

Ford, E.D. ......................................................................................................204, 205

Forsman, E.D. ................. 8, 27, 36, 37, 111, 112, 113, 114, 115, 116, 117, 118, 119,

120, 121, 122, 123, 124, 125, 126, 127, 128, 129, 130, 131, 132, 133, 134, 

241, 309, 310, 343, 403, 404, 405, 406, 407, 413, 414, 475, 479 

Foster, J.B. ............................................................................................................135

Franklin, A.B. ..........................................................................................................93

Franklin, J.F. ....................................................................93, 136, 137, 215, 281, 283 

Gaines, E.P. .......................................................................................94, 95, 238, 239

Gannon, W.T. ........................................................................................................138

Gardner, S.L. .........................................................................................................174

George, S.B. ..........................................................................................139, 140, 224

Ghiselin, J. ..............................................................................................................79

Gilbert, E.H. ..........................................................................................................382

Gilbert, F.F. ...........................................................................................................141

Gillesberg, A-M. ...........................................................................142, 143, 144, 145

Gitzen, R.A. ..........................................................................................................146

Golightly, R.T. .......................................................................................................147

Gomez, D.M. .........................................................................................148, 149, 150

Goodwin, G.G. ........................................................................................................78

Gordon, K.L. .........................................................................................................151

Gould, G.I. ............................................................................................................152

Grady, F. ................................................................................................................358

Graf, J-D. ................................................................................................................74

Graham, C.H. ........................................................................................................179

Graham, S.A. .........................................................................................132, 153, 154
166

GENERAL TECHNICAL REPORT PNW-GTR-909

Green Diamond Resource Company ....................................................................155

Green, G.A. ...............................................................................................................8

Greenway, G. ........................................................................................................156

Grinnell, J. .....................................................................................................157, 158

Grossenheider, R.P. .................................................................................................51

Grosvenor, M.B. ....................................................................................................159

Guilday, J.E. ..........................................................................................................160

Gunderson, A.G. ...................................................................................................414

Guralnick, R. .........................................................................................................464

Gutiérrez, R.J. ...........................................................................................24, 93, 466

Hadly, E.A. ........................................................................................................22, 41

Hafner, D.J. ...................................................................................................161, 486

Hagar, J.C. .............................................................................................................162

Hagele, T. ..............................................................................................................163

Hagestedt, R. .........................................................................................................194

Haig, S.M. .......................................................................................27, 309, 310, 311

Hall, E.R. ...............................................................................................164, 165, 166

Halpern, C.B. ........................................................................................................146

Hamilton, W.J., Jr. .........................................................................................169, 170

Hamilton, W.J., III. ........................................................................................167, 168

Hammer, E.W. .......................................................................................171, 225, 232

Hänfling, B. .............................................................................................................54

Harmon, M.E. .......................................................................................................388

Harris, J. ..................................................................................................................50

Harris, L.D. ...........................................................................................................172

Hart, B.S. ...............................................................................................................181

Hatch, N.R. ...................................................................................................131, 132

Haukisalmi, V. .......................................................................................................173
167

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Haverkost, T.R. .....................................................................................................174

Hawley, J.J.V.G. ....................................................................................................105

Hayes, J.P. .............................................................2, 12, 20, 175, 176, 177, 178, 401

Haynes, R.W. ........................................................................................................266

Henttonen, H. ........................................................................................................173

Hernandez, P.A. .....................................................................................................179

Higley, J.M. ...................................................................................................147, 379

Hijmans, R.J. .........................................................................................................464

Hinckley, T.M. .......................................................................................................204

Hoffmann, R.S. .....................................................................................................235

Holthausen, R.S. ...........................................................................................193, 414

Hooper, E.T. ..................................................................................................180, 181

Hooven, E.F. ....................................................................................................38, 182

Hopkins, D.S. ................................................................................................183, 333

Horn, K.M. ............................................................................................................117

Howell, A.B. .........................................................................184, 185, 186, 187, 188

Howell, K. .............................................................................................................239

Hsu, T.C. ...............................................................................................................189

Hubbard, C.A. ...............................................................................................190, 191

Huey, L.M. ............................................................................................................192

Huff, M.H. .................................................11, 89, 141, 144, 193, 194, 252, 350, 367

Huff, R.D. .....................................................................................................195, 323

Hughes, C. .............................................................................................................195

Humboldt Redwood Company .....................................................................196, 197

Humboldt State University Vertebrate Museum ...................................................198

Hunter, J.E. ............................................................................................................199

Huso, M.M.P. ....................................................................................................94, 95

Hutchinson, W.F. .....................................................................................................54
168

GENERAL TECHNICAL REPORT PNW-GTR-909

Ingles, L.G. ...................................................................................................200, 201

Institute for Natural Resources .............................................................................202

Irwin, L.L. .............................................................................................................203

Ishii, H. ..........................................................................................................204, 205

Jackson, S.T. ..........................................................................................................470

Jacobs, K.M. .........................................................................................................264

Jaros, G. ................................................................................................................206

Jass, C.N. .................................................................................................................26

Jenness, R. ...............................................................................................................97

Jewett, S.G. ...........................................................................207, 208, 209, 210, 211

Jobanek, G.A. ........................................................................................................212

Johnson, D.H. ........................................................................................194, 213, 214

Johnson, D.S. ..........................................................................................................70

Johnson, E. ..............................................................................................................76

Johnson, K.N. ................................................................................................215, 388

Johnson, M.L. ..............140, 216, 217, 218, 219, 220, 221, 222, 223, 224, 225, 226,

227, 228, 229, 230, 231, 232, 282, 354, 458

Johnson, S .....................................................................................................226, 344

Johnston S. ..............................................................................................................20

Johnston, A.N. .......................................................................................................233

Jollie, M. ...............................................................................................................234

Jones, C. ................................................................................................................235

Jones, C.A. ............................................................................................................235

Jones, J.K., Jr. ..........................................................................................................67

Jones, J.M. ...............................................................................................35, 236, 237

Jones, L.L.C. .........................................................................................................367

Jones, P.D. .............................................................................................................102

Juday, G. ................................................................................................................136
169

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Kagan, J. ........................................................................................................238, 239

Kaufman, B. ..........................................................................................................163

Kays, R.W. ............................................................................................................240

Kellogg, R. ............................................................................................................307

Kelsey, R.G. ..........................................................................................................241

Kelson, K.R. ..........................................................................................................166

Kenagy, G.J. ............................................................................................................75

Kennedy, R.S.H. ....................................................................................................389

Kenworthy, J. ........................................................................................................369

Kerbo, R. ...............................................................................................................369

Kerner, H.M. ...........................................................................................................30

Kesner, M.H. .................................................................................227, 242, 243, 244

Khan, M. ...............................................................................................................489

Kimerling, A.J. ..................................................................................................94, 95

Kirkland, G.L., Jr. .........................................................................................161, 486

Klompen, H. ..........................................................................................................163

Koepke Wise, L. ....................................................................................................238

Koffman, R. ...........................................................................................................245

Koford, M. ............................................................................................................246

Koninckx, A. ...........................................................................................................55

Kritzman, E.B. ......................................................................................................247

Krohne, D.T. ..........................................................................................................248

Krystufek, B. ...........................................................................................................54

Lamb, C.C. ............................................................................................................249

Lancia, R.A. ..........................................................................................................416

Landon, C.P. ..........................................................................................................343

Larrison, E.J. ...................................................................................................77, 250

Laudenslayer, W.F., Jr. ..........................................................................................257
170

GENERAL TECHNICAL REPORT PNW-GTR-909

Lawlor, T.E. ...........................................................................................................251

Leach, R. .................................................................................................................37

Lebedev, V.S. .............................................................................................................1

Lee, D.C. ...............................................................................................................266

Leeming, J.P. ...........................................................................................................82

Lehmkuhl, J.F. .......................................................................................................252

Lesher, R. ..............................................................................................................312

Lewis, J.H. ............................................................................................................253

Lewis, M. ................................................................................................................37

Lewis, R.E. ............................................................................................................253

Lidicker, W.Z., Jr. ..................................................................................................228

Lint, J.B. ..........................................................................................................33, 413

Lisle, T.E. ..............................................................................................................257

Little, L. ................................................................................................................254

Livingston, S. ........................................................................................255, 256, 265

Lowe, R.W. ...........................................................................................................150

Lundsten, S. ............................................................................................................37

Luoma, D.L. ..........................................................................................................264

Madsen, S. ...............................................................................................................34

Maguire, C.C. ................................................................................146, 257, 263, 264

Mailliard, J. ...................................................................................................258, 259

Mannan, R.W. .......................................................................................................416

Manning, J.A. ........................................................................................260, 261, 262

Manning, T. ...........................................................................................146, 263, 264

Manriquez, A.C. ....................................................................................................204

Marcot, B.G. ...........................................................................12, 265, 266, 312, 414

Marincovich, L., Jr. ...............................................................................................357

Marks, S.B. ...........................................................................................................387
171

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Marshall, S.A. .......................................................................................................340

Martin, A.C. ..........................................................................................................267

Martin, J.R. ...........................................................................................................266

Martin, K.J. ...........................................................................................268, 269, 270

Martin, L.D. ..........................................................................................................271

Martin, R.A. ..........................................................................................................272

Marzluff, J.M. .........................................................................................................93

Maser, C. ......118, 136, 171, 229, 230, 231, 232, 253, 273, 274, 275, 276, 277, 278, 

279, 280, 281, 282, 283, 284, 297, 321, 458, 471

Master, L.L. ...........................................................................................................179

Mate, B.R. .............................................................................................................283

Matthews, S.M. .....................................................................................................379

Max, T. ..................................................................................................................362

Mayer, W.V. ...........................................................................................................285

Mazurek, M.J. ...............................................................................................286, 287

McComb, B.C. ......................................................................................................389

McComb, W.C. ...............................................................................82, 270, 288, 289

McDonald, M.A. ...................................................................................................132

McGarigal, K. .......................................................................................................289

McGrath, M.T. ......................................................................................................389

McGroddy, M.E. ...................................................................................................361

McGuire, J.L. ..........................................................................................................41

McKee, A. .............................................................................................................136

McKelvey, K.S. .............................................................................................467, 487

McLellan, J.E. .......................................................................................................290

McNab, B.K. .................................................................................................291, 292

Meiselman, N. ...............................................................................................293, 294

Mellen, T.K. ..........................................................................................................194
172

GENERAL TECHNICAL REPORT PNW-GTR-909

Mellen-McLean, K. ...............................................................................................195

Merriam, C.H. ...............................................................................................295, 296

Meslow, E.C. ......................................................................... 114, 115, 119, 297, 413

Michel, A.K. ..........................................................................................................298

Millar, J.S. .....................................................................................................299, 300

Miller, A.H. ...................................................................................................301, 302

Miller, G.S., Jr. ..............................................................303, 304, 305, 306, 307, 308 

Miller, M.P. ...........................................................................................309, 310, 311

Miller, S.A. ............................................................................................................310

Mires, G.W. ...................................................................................................117, 154

Molina, R. .....................................................................................................266, 312

Montgomery, G.G. ................................................................................................106

Montuire, S. ............................................................................................................73

Moore, D.L. ...........................................................................................................382

Mowdy, J.S. ...................................................................................................120, 133

Mullins, T.D. ...........................................................................................................27

Murie, O.J. ............................................................................................................313

Murray, M.A. ................................................................................................314, 315

Musser, G.G. .............................................................................................67, 68, 316

Nawa, R.K. ............................................................................................................317

Nelson, A.L. ..........................................................................................................267

Nelson, D.W. .........................................................................................................318

Nelson, E.W. .........................................................................................................319

Nelson, S.K. ..............................................................................................................8

Newton, M. .....................................................................................................82, 288

Noon, B.R. ....................................................................................................413, 466

Norse, E.A. ............................................................................................................353

Nowak, R.M. .........................................................................................................320
173

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Nussbaum, R.A. ....................................................................................................321

Nyári, Á.S. ............................................................................................................464

O’Neil, T.A. ......................................................................................94, 95, 194, 330

Olsen, K. ...............................................................................................................389

Olson, D.H. ...................................................................................................322, 323

Olterman, J.H. .......................................................................................324, 325, 326

Oregon Department of Fish and Wildlife ..............................................................327

Oregon Department of Forestry ............................................................................328

Oregon Natural Heritage Information Center .......................................................329

Oregon Wild ............................................................................................................72

Otto, I.A. ................................................................................................................121

Owings, D.H. ........................................................................................................331

Pabst, R.J. ..............................................................................................................389

Pacific Lumber Company .....................................................................................332

Paez, R.A. .............................................................................................................318

Palmer, C. ................................................................................................................18

Pampush, A. ..........................................................................................................333

Paradiso, J.L. .........................................................................................................320

Parmalee, P.W. ......................................................................................................160

Parmer, R. .............................................................................................................334

Paton, P.W.C. .........................................................................................................350

Patton, C.P. ............................................................................................................451

Patton, D.R. .....................................................................................................90, 352

Pearson, O.P. .........................................................................................................335

Pechman, M.J. ...............................................................................336, 337, 338, 339

Penland, T.F. ..........................................................................................................147

Perkins, S.L. ..........................................................................................................464

Perron, B. ..............................................................................................................340
174

GENERAL TECHNICAL REPORT PNW-GTR-909

Peterson, A.T. ................................................................................................174, 464

Pettinato, J.S. .........................................................................................................340

Phillips, C. .............................................................................................................341

Phillips, D. ................................................................................................................9

Piirto, D.D. ..............................................................................................................76

Pinto, C. ................................................................................................................377

Plank, R.D. ............................................................................................................451

Pratt, H.S. ..............................................................................................................342

Price, A.L. .............................................................................................120, 122, 133

Price, W.W. ....................................................................................................134, 343

Pugh, S.R. .............................................................................................................344 

Quay, W.B. ............................................................................345, 346, 347, 348, 349

Ralph, C.J. .............................................................................................................350

Raphael, M.G. .......................................................................351, 352, 353, 414, 415

Rausch, R.L. ..................................................................................................173, 354

Rauscher, R.J. ........................................................................................................109

Raymond, R.B. ......................................................................................................382

Reeder, D.M. ...................................................................................................68, 316

Reeder, W.................................................................................................................79

Reeves, G.H. .................................................................................................388, 414

Rehn, J.A.G. ..........................................................................................................308

Repenning, C.A. ............................................................................355, 356, 357, 358

Reynolds, R.T. .......................................................................................................359

Řičánková, V. ........................................................................................................363

Rice, D.W. .............................................................................................................235

Richards, R.L. .......................................................................................................360

Richardson, A.D. ...................................................................................................361

Rittenhouse, B. ......................................................................................................362
175

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Ritzi, C.M. .............................................................................................................472

Roberts, D.R. .........................................................................................170, 231, 232

Robovský, J. ..........................................................................................................363

Rock, D.C. .............................................................................................................203

Rock, S.L. .............................................................................................................203

Roest, A.I. .....................................................................................................364, 365

Ronnenerg, K.L. ....................................................................................................479

Rosenberg, D.K. ....................................................................................................366

Rosentreter, R. .......................................................................................................371

Ruedas, L.A. .........................................................................................................102

Ruff, S. ............................................................................................................65, 139

Ruggiero, L.F. ...................................................11, 89, 141, 144, 252, 350, 367, 416

Russell, W.C. .........................................................................................................368

Santucci, V.L. ........................................................................................................369

Sawyer, J.O., Jr. .......................................................................................................30

Schmalenberger, F. ..................................................................................................35

Schmelzer, S. ...........................................................................................................10

Schmidly, D.J. .......................................................................................................235

Schmidt, K. .............................................................................................................34

Schoen, J.W. ..........................................................................................................416

Sedell, J.R. ....................................................................................................136, 414

Semken, H.A., Jr. ..................................................................................................370

Severson, K.E. .................................................................................................90, 352

Sharnoff, S. ...........................................................................................................371

Shaughnessy, M.M. ...........................................................................................94, 95

Shea, P.J. ...............................................................................................................257

Shelton, A.C. .........................................................................................372, 373, 374

Sherrell, P.E. ..................................................................................................375, 376
176

GENERAL TECHNICAL REPORT PNW-GTR-909

Sierra Club ..............................................................................................................72

Sillett, S.C. ............................................................................................................387

Silovsky, G.D. .......................................................................................................377

Simberloff, D. ...............................................................................................378, 385

Slauson, K.M. .......................................................................................................379

Smith, W.P. ............................................................................................................380

Smoluk, A. ............................................................................................................381

Snow, M. ...................................................................................................................9

Snyder, K.U. ..........................................................................................................382

Sobb, N. ................................................................................................................383

Soeldner, A. ...............................................................................................................9

Solis, D.M. ............................................................................................................414

Sondenaa, A.C. ......................................................................................................384

Soulé, M.E. ...........................................................................................................385

Sovern, S.G. ..........................................................................................................407

Spade, A. ...............................................................................................................386

Spickler, J.C. .........................................................................................................387

Spies, T.A. .......................................................................................90, 137, 388, 389

Stagnaro, B. ...........................................................................................................390

Standiford, R.B. ......................................................................................................76

Steidl, R.J. .............................................................................................................330

Stein, B.R. .............................................................................................................391

Stephens, F. ...........................................................................................................392

Stone, W. ...............................................................................................................394

Stone, G.C. ............................................................................................................393

Storer, J.E. .............................................................................................................395

Storer, R.W. ...........................................................................................................396

Storm, R.M. ...........................................................................................................284
177

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Strong, W.D. ..........................................................................................................397

Stuart, J.D. ...............................................................................................................76

Sturges, F.W. .........................................................................................................398

Sullivan, T.J. ..........................................................................................................382

Suzuki, N. ..............................................................................................399, 400, 401

Swanson, F. ...........................................................................................................136

Swindle, K.A. ........................................................................................................366

Swingle, J.K. ..................37, 120, 123, 124, 125, 126, 127, 128, 129, 130, 131, 132,

133, 134, 241, 310, 402, 403, 404, 405, 406, 407 

Szaro, R.C. ......................................................................................................90, 352

Sztukowski, L. ........................................................................................................93

Tamarin, R.H. ........................................................................................................344

Taylor, C.A. ...........................................................................................................350

Taylor, W.P. ...................................................................................408, 409, 410, 411

Terres, J.K. ............................................................................................................411

Tesakov, A.S. .............................................................................................................1

Tevis, L., Jr. ...........................................................................................................412

Thomas, J.W. .................................................................................413, 414, 415, 416

Thompson, B.C. ....................................................................................................344

Thompson, D.R. ....................................................................................................318

Thompson, J.L. .....................................................................................................417

Tiekotter, K. ..............................................................................................................9

Tingey, D. ..................................................................................................................9

Todd, A. .................................................................................................................418

Tolle, T. ...................................................................................................................18

Toweill, D.E. .........................................................................................................419

Town, P. .........................................................................................................483, 484

True, F.W. ..............................................................................................................420
178

GENERAL TECHNICAL REPORT PNW-GTR-909

Tucker, G.F. .............................................................................................................20

U.S. Department of Agriculture, Forest Service ..........................421, 422, 423, 424,

425, 426, 427, 428, 429, 430, 431, 432, 433, 434, 435, 436, 437, 438, 439, 

440, 441, 442, 443, 444, 445, 446

U.S. Department of the Interior, Bureau of Land Management ...........423, 424, 425,

426, 427, 428, 429, 430, 431, 432, 433, 434, 435, 436, 437, 438, 439, 440, 

441, 442, 443, 444, 445, 446, 447

U.S. Department of the Interior, Fish and Wildlife Service ..................448, 449, 450 

Valentine, B.E. ......................................................................................................257

Van Gelder, R.G. .....................................................................................................78

Van Norman, K. ............................................................................................195, 323

Verner, J. .......................................................................................................297, 413

Verts, B.J. ............................................................5, 69, 325, 326, 451, 452, 453, 454

Viriot, L. ..................................................................................................................73

Voge, M. ................................................................................................455, 456, 457

Voth, E.H. ..............................................................................................................458

Vrieze, J.M. ...................................................................................................459, 460

Vrilakas, S......................................................................................................238, 239

Waldien, D.L. ........................................................................................................461

Walker, A. ......................................................................................................462, 463

Wallace, S.C. .........................................................................................................370

Waltari, E. .............................................................................................................464

Walters , B.L. ........................................................................................................472

Ward, J.P., Jr. .........................................................................................465, 466, 487

Waters, J.R. ...................................................................................................380, 467

Watson, H.C. .........................................................................................................468

Weatherspoon, C.P. ...............................................................................................257

Webb, M.G. ...........................................................................................................469
179

Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. Pomo)...

Weller, T.J. ...............................................................................................................76

Welsh, H.H., Jr. .............................................................................................387, 467

Weng, C. ................................................................................................................470

West, S.D. ........................................................................................................11, 146

Whitaker, J.O., Jr. ..................................................................107, 163, 471, 472, 473

Wiens, J.D. ....................................................................................................474, 475

Wight, H.M. ..........................................................................................119, 476, 477

Williams, D.F. .......................................................................................................478

Wilson, D.E. ..............................................................................65, 68, 139, 240, 316

Wilson, M.E. .........................................................................................................204

Wilson, N. .....................................................................................................472, 473

Wilson, T.M. ..........................................................................................................479

Winter, S. ...............................................................................................................298

Wise, L. .................................................................................................................239

Witmer, G.W. ....................................................................................................8, 480

Witt, J.W. .........................................................................................................66, 178

Wooster, T.W. ................................................................................481, 482, 483, 484

Wright, C.A. ..........................................................................................................222

Wright, D. .............................................................................................................485

Wright, J. ...............................................................................................................485

Yensen, E. .....................................................................................................161, 486

Young, J. ...................................................................................................................9

Zabel, C.J. ............................................... 12, 114, 115, 116, 237, 251, 380, 467, 487

Zahler, P. .......................................................................................................488, 489

Zentner, P.L. ..........................................................................................................490

Zielinski, W.J. .................................................................................70, 147, 286, 287

Zim, H.S. ...............................................................................................................267

Zrzavý, J. ...............................................................................................................363
Pacific Northwest Research Station

Web site http://www.fs.fed.us/pnw

Telephone (503) 808-2592

Publication requests (503) 808-2138

FAX (503) 808-2130

E-mail pnw_pnwpubs@fs.fed.us

Mailing address Publications Distribution

	 	 Pacific Northwest Research Station

 P.O. Box 3890

	 	 Portland, OR 97208-3890
U.S. Department of Agriculture

Pacific Northwest Research Station

1220 SW 3rd Ave., Suite 1400

P.O. Box 3890

Portland, OR 97208-3890

Official Business

Penalty for Private Use, $300