(Arborimus Longicaudus), Sonoma Tree Vole (A. Pomo), and White-Footed Vole (A
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United States Department of Agriculture Annotated Bibliography of the Red Tree Vole (Arborimus longicaudus), Sonoma Tree Vole (A. pomo), and White-Footed Vole (A. albipes) Forest Pacific Northwest General Technical Report August Service Research Station PNW-GTR-909 2016 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 pro- grams). 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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: [email protected]. 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that we could find on tree voles (Arborimus longicaudus, A. pomo) and white-footed voles (A. albipes), including many unpublished sources in museum archives, court proceedings, and agency documents.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. 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 stud- ies 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 geo- graphic 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 bib- liography, 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 1 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 white- footed 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 longi- caudus 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 differ- ences 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 subspe- cies (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 2 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 regard- ing 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.