STATUS OF THE RED-TAILED CHIPMUNK (TAMIAS RUFICAUDUS) IN BRITISH COLUMBIA

by D. Nagorsen

Illustration © Michael Hames

Victoria, B.C.

Wildlife Working Report No. WR. 115 December 2004

Wildlife Working Reports frequently contain preliminary data, so conclusions based on these may be subject to change. Working Reports receive little review. They may be cited in publications, but their manuscript status should be noted. Copies may be obtained on-line from http://www.env.gov.bc.ca/wld/ or from the Ministry of Environment, Ecosystems Branch, P.O. Box 9338 Stn. Prov. Govt., Victoria, BC V8W 9M1. © Province of British Columbia ISBN 978-0-7726-6288-0 Date: May 6, 2010 British Columbia. Ministry of Environment. Status of the Red-tailed Chipmunk (Tamias ruficaudus) in British Columbia Citation Nagorsen, D.W. 2004. Status of the Red-tailed Chipmunk (Tamias ruficaudus) in British Columbia. B.C. Ministry of Environment, Victoria, BC. Wildlife Working Report No. WR 115.

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DISCLAIMER

In cases where a Wildlife Working Report or Bulletin is also a species’ status report, it may contain a status recommendation from the author. The Province, in consultation with experts, will determine the official conservation status and consider official legal designation. The data contained in the status report will be considered during those processes. NOTE: The content of this report was completed in March of 2004 and reflects the state of our knowledge of this species at that time. It has not been updated prior to publication April 2010. Information on current status of native species in British Columbia can be found through B.C. Species and Ecosystems Explorer http://www.env.gov.bc.ca/atrisk/toolintro.html

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ACKNOWLEDGMENTS This status report was funded by the (then) British Columbia Ministry of Water, Air and Land Protection. Mark Fraker, who initiated the Kootenay chipmunk research project in 1996, collected some of the specimens and habitat data that provide a basis for this report. Nick Panter assisted with field work in the Kootenay chipmunk research project and prepared the voucher specimens and genital bone preparations used for identification. Ian Parfitt created the range map and provided a digital file of the map. Dave Grieve provided information on coalfields in the Kootenay region of British Columbia. I thank Laura Friis, John Krebs, and Ted Antifeau for their support. The following museums provided the author with their museum records and loans of Red-tailed Chipmunk specimens from British Columbia: Canadian Museum of Nature, Ottawa; Royal British Columbia Museum, Victoria; Royal Ontario Museum, Toronto; Cowan Vertebrate Museum, University of British Columbia, Vancouver.

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SUMMARY The Red-tailed Chipmunk (Tamias ruficaudus) is confined to a small geographic area in the Rocky and Columbia Mountains of Montana, Idaho, Washington, Alberta, and British Columbia. Two subspecies are recognized: T. r. ruficaudus and T. r. simulans. They differ in mitochondrial DNA, male and female genital bone morphology, pelage colour, and skull morphology. Although they appear to be incipient species, mitochondrial DNA introgression in central Idaho suggests that reproductive isolation among these two forms is not complete. In British Columbia, T. r. ruficaudus and T. r. simulans are allopatric, separated by about 180 km. T. r. ruficaudus inhabits a narrow elevational zone (1785–1950 m) in the extreme southern Rocky Mountains of British Columbia. T. r. simulans occupies a broad elevational range (560–1829 m) in the southern Selkirk Mountains of British Columbia where its range is delimited by the Columbia and Kootenay Rivers. In the Rocky Mountains, the Red-tailed Chipmunk is associated with subalpine forest of Englemann spruce (Picea englemannii) and subalpine fir (Abies lasiocarpa). In the Selkirk Mountains, the species is associated with forests of western hemlock (Tsuga heterophylla), Englemann spruce, and western redcedar (Thuja plicata). Although the Red-tailed Chipmunk is an arboreal chipmunk that constructs tree nests, it inhabits forests of various ages from mature to recent clearcuts. It is often most common in disturbed habitats such as clearings, roadsides, and forest edges with abundant shrubs and woody debris on the ground. The narrow elevational zone occupied in the Rocky Mountains reflects competition effects with the Yellow-pine Chipmunk (Tamias amoenus) and Least Chipmunk (Tamias minimus). The three species are contiguously allopatric along an elevational gradient. In the Selkirk Mountains where T. ruficaudus inhabits a wide range of elevations, T. amoenus appears to be rare and sparsely distributed. The Red-tailed Chipmunk is a facultative hibernator. Rather than accumulating heavy fat reserves, it depends on seeds stored in its burrow for winter survival. Breeding begins shortly after animals emerge from hibernation in spring. Females produce only one litter in the breeding season; only 11–15% of yearling females will breed. There are few threats to this species. Although forest harvesting is occurring in much of the range of the Red-tailed Chipmunk, the species readily exploits early successional stages associated with logging and forest fires. In the Rocky Mountains of Canada where much of the range is in protected areas, fire suppression could affect habitat quality. The distribution of T. r. ruficaudus is outside the range of any coalfields and potential open pit coal mines in the Rocky Mountains of British Columbia. This species is not listed by the International Union for Conservation of Nature and Natural Resources (IUCN) or the Committee on the Status of Endangered Wildlife in Canada (COSEWIC). The two subspecies, T. r. simulans and T. r. ruficaudus, represent nationally significant populations and warrant separate rankings. They probably diverged before the late Pleistocene glaciation. Their allopatric distributions in Canada are the result of postglacial dispersion patterns and competition effects from other chipmunk species.

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The British Columbia Conservation Data Centre currently ranks T. r. ruficaudus as S2 because of its limited range. Although it has a limited distributional range in British Columbia (extent of occurrence is 150 km2 and includes five known occurrences), much of its Canadian range occurs within protected areas (Waterton Lakes National Park, Akamina-Kishinena Provincial Park). Except for the possible effects of fire suppression, no threats are known. There is no evidence of habitat or population declines. Populations of T. r. ruficaudus in British Columbia are connected to populations in Montana and Alberta. A COSEWIC assessment of this subspecies will have to include data from the Alberta population. T. r. simulans is ranked as S3S4 by the British Columbia Conservation Data Centre. Its distributional range in British Columbia is restricted to a small area in the southern Selkirk Mountains (extent of occurrence is about 4070 km2 and includes about 15 occurrences). However, within this small area, it inhabits a broad elevational range and a variety of habitats, including early successional stages. No threats are known and there is no evidence of population or habitat declines. Priorities for conserving and managing this species include inventories in the mountains west of the Flathead River valley and south of Crowsnest Pass to determine the limits of the range of T. r. ruficaudus. Areas on the north side of the Kootenay River near Nelson and the west side of the Columbia River near Trail are needed to verify the limits of T. r. simulans. A high priority for management of T. r. ruficaudus is a study on the role of fire in maintaining habitat.

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TABLE OF CONTENTS

ACKNOWLEDGMENTS .............................................................................................................. iv SUMMARY .................................................................................................................................... v TABLE OF CONTENTS .............................................................................................................. vii 1  SPECIES INFORMATION ..................................................................................................... 1 

1.1  Name and Classification................................................................................................... 1 1.2  Description ....................................................................................................................... 2 1.3   Nationally Significant Populations ................................................................................... 2 

2  BIOLOGY ................................................................................................................................ 2 2.1  General ............................................................................................................................. 2 2.2  Reproduction .................................................................................................................... 3 2.3  Survival ............................................................................................................................ 3 2.4  Physiology ........................................................................................................................ 3 2.5  Movements/Dispersal ....................................................................................................... 3 2.6  Nutrition and Interspecific Interactions ............................................................................ 3 2.7  Behaviour and Adaptability ............................................................................................. 4 

3  HABITAT ................................................................................................................................ 5 3.1  Habitat Requirements ....................................................................................................... 5 3.2  Trends in Habitat Quality and Quantity ........................................................................... 8 3.3  Protection/Ownership ....................................................................................................... 8 

4  DISTRIBUTION ...................................................................................................................... 8 4.1  Global/ North American ................................................................................................... 8 4.2  British Columbia .............................................................................................................. 9 

5  POPULATION SIZE AND TRENDS ................................................................................... 11 6  LIMITING FACTORS AND THREATS .............................................................................. 11 7  SPECIAL SIGNIFICANCE OF THE SPECIES.................................................................... 12 8  EXISTING PROTECTION OR OTHER STATUS ............................................................... 12 9  RECOMMENDATIONS AND MANAGEMENT OPTIONS .............................................. 13 

9.1  Inventory ........................................................................................................................ 13 9.2  Habitat Protection and Acquisition ................................................................................ 13 

10  EVALUATION .................................................................................................................. 13 10.1  Provincial Ranking ......................................................................................................... 13 

10.1.1 T. r. simulans (see Appendix 3) ................................................................................. 13 10.1.2 T. r. ruficaudus (see Appendix 2) ............................................................................... 12

10.2  COSEWIC Ranking ....................................................................................................... 15 11  AUTHORITIES CONSULTED ......................................................................................... 16 12  LITERATURE CITED ....................................................................................................... 16 

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LIST OF FIGURES Figure 1. Logged area, Church Creek. T. r. simulans inhabited second- growth

and edge habitats ...................................................................................................................... 6 Figure. 2. Clearcut area, Giveout Creek. T. r. simulans inhabited edge habitats and

second-growth forest ................................................................................................................ 6 Figure 3. Wall Lake, Akamina-Kishinena Provincial Park. T. r. ruficaudus

inhabited forest bordering the lake ........................................................................................... 7 Figure 4. Middle Kootenay Pass. T. r. ruficaudus inhabited the subalpine

forest and lower portions of the old burn ................................................................................. 7 Figure 5. Distribution of the Red-tailed Chipmunk (Tamias ruficaudus) in

North America .......................................................................................................................... 9 Figure 6. Distribution of the Red-tailed Chipmunk (Tamias ruficaudus) in

British Columbia .................................................................................................................... 10 

LIST OF APPENDICES Appendix 1. Summary of all known occurrences (museum specimens, literature records)

of the Red-tailed Chipmunk (Tamias ruficaudus) in British Columbia.. ............................... 19 Appendix 2. Technical Summary - Tamias ruficaudus ruficaudus ............................................... 21 Appendix 3. Technical Summary - Tamias ruficaudus simulans .................................................. 23

1 SPECIES INFORMATION

1.1 Name and Classification The Red-tailed Chipmunk (Tamias ruficaudus [Howell 1920]) is a member of the rodent family Sciuridae. In the early literature, this species was classified as Eutamias ruficaudus; however, in the recent literature, all North American chipmunk species are classified in the genus Tamias (Wilson and Reeder 1993). Nevertheless, generic classification of the chipmunks is unresolved. Recent data from ectoparasites (Jameson 1999) and mitochondrial DNA (mtDNA) (Piaggio and Spicer 2001) suggest that western North American chipmunks are sufficiently distinct to be separated into the genus Neotamias. Based on molecular data, Piaggio and Spicer (2001) concluded that T. ruficaudus was a member of the minimus species group, which also includes the Least Chipmunk (Tamias minimus), Panamint Chipmunk (Tamias panamintinus), and Long-eared Chipmunk (Tamias quadrimaculatus). Howell (1929) recognized two subspecies of T. ruficaudus: T. r. simulans (Selkirk Mountains subspecies) and T. r. ruficaudus (Rocky Mountains subspecies) largely on the basis of pelage. Both occur in British Columbia. Based on their distinct male genital bones (i.e., bacula), White (1953) speculated that T. r. simulans and T. r. ruficaudus were distinct species. In a study of geographic variation among populations from Washington, Idaho, and Montana, Patterson and Heaney (1987) demonstrated that the two taxa were differentiated in male genital bone (i.e., baculum) morphology but overlapped in cranial morphology. They suggested that the two taxa were differentiated at the species level, but they recommended that detailed studies of possible hybridization in contact zones were needed. Nagorsen et al. (2002) demonstrated that in western Canada, the two subspecies are strongly divergent in male and female genital bone morphology, skull morphology, and pelage colour. A study of mtDNA by Good and Sullivan (2001) revealed distinct western and eastern clades of T. ruficaudus that are concordant with the distribution of the T. r. ruficaudus and T. r. simulans morphological forms. Sequence divergence among the two clades was about 4.7%. However, because Good and Sullivan (2001) found evidence for mtDNA introgression in the Clearwater Drainage of Idaho, they concluded that reproductive isolation was not complete and recommended that the two taxa be treated as subspecies. Interestingly, Good et al. (2003) found evidence for recent mtDNA introgression between the Yellow-Pine Chipmunk (Tamias amoenus) and T. r. ruficaudus in the Rocky Mountains and Rocky Mountain Trench of British Columbia. Specimens of T. amoenus (identified from genital bone morphology, pelage, and skull morphology) taken at seven locations were carrying the mtDNA haplotypes of T. r. ruficaudus: Harvey Creek, Flathead River valley, Kishinena Creek, Middle Kootenay Pass, Sage Creek, Todhunter Creek. Some of these locations (e.g., Todhunter Creek) are far outside the known range of T. r. ruficaudus in British Columbia. These results demonstrate that introgression must be considered when identifying chipmunks from mtDNA markers. An alternate common name for T. ruficaudus is the Rufous-tailed Chipmunk. A common name used for T. r. simulans is the Coeur D’ Alene Chipmunk (Anderson 1946). A more appropriate common name would be the Red-tailed Chipmunk–Columbia Mountains subspecies. Anderson (1946) used

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the species common name for T. r. ruficaudus. An appropriate common name would be the Red-tailed Chipmunk–Rocky Mountains subspecies.

1.2 Description

The Red-tailed Chipmunk is a large chipmunk with the typical body striping pattern of five dark stripes alternating with four light stripes. Pelage colour varies strikingly between the two subspecies. The Rocky Mountain subspecies (T. r. ruficaudus) has bright reddish pelage on the underside of its tail; the sides, shoulders, and back of the head are washed with rufous or deep orange. The fur on the dorsal surface of the hind feet is rufous or cinnamon coloured. The Selkirk Mountains subspecies (T. r. simulans) is paler with tan or yellow fur on the underside of the tail; the sides, shoulders, and back of the head are washed with tan or yellow. The fur on the dorsal surface of the hind feet is tawny. In both forms, the winter pelage tends to be paler. The dental formula is 1/1 incisors, 0/0 canines, 2/1 premolars, and 3/3 molars. The Red-tailed Chipmunk converges in pelage colour with some populations of the Yellow-pine Chipmunk. In British Columbia, convergence between the two species is most pronounced in the Columbia Mountains where T. r. simulans and T. a. luteiventris cannot be reliably identified from pelage colour (Nagorsen et al. 2002). However, the male and female genital bones of Tamias ruficaudus, the largest and most robust of the four chipmunk species found in British Columbia, are diagnostic (see Nagorsen 2002; Nagorsen et al. 2002). Mean representative body measurements (range in parentheses) for adult specimens of T. r. ruficaudus from British Columbia and Alberta are as follows: total length = 222 mm (207–235), tail vertebrae = 95 mm (85–102), hind foot = 34 mm (32–36), ear = 17 mm (14–19), and body mass = 65.3 g (53.5–78.7). Mean representative body measurements (range in parentheses) for adult specimens of T. r. simulans from British Columbia are as follows: total length = 225 mm (216–237), tail vertebrae = 102 mm (93–115), hind foot = 33 mm (30–35), ear = 16 mm (13–19), and body mass = 54.6 g (44.2–64.6).

1.3 Nationally Significant Populations

T. r. ruficaudus and T. r. simulans are two strongly divergent subspecies that differ in male and female genital bone morphology and mtDNA (Good and Sullivan 2001). They appear to represent two distinct phylogeographic lineages that diverged before Late Pleistocene climatic events (Good and Sullivan 2001). In Canada, where the two subspecies are allopatric separated by about 180 km, they also differ in cranial morphology, pelage colour, and ecology (Nagorsen et al. 2002). They warrant treatment as distinct evolutionary units for conservation and management.

2 BIOLOGY

2.1 General Most of the information on the general biology of the Red-tailed Chipmunk is based on Mirza Beg's (Beg 1969) dissertation research on T. r. ruficaudus in Montana. Little ecological research has been done on either subspecies in Canada. Nagorsen et al. (2002) analyzed taxonomy and distribution. Panian (1996) evaluated habitat use by T. r. simulans in the Selkirk Mountains of British Columbia, but his study is seriously flawed because the identification of most of his chipmunk captures was not verified from genital bone morphology, and some of his captures may be T. amoenus (see Nagorsen et al. 2002; Good et al. 2003).

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2.2 Reproduction Reproduction in British Columbian populations has not been studied. In Montana, Beg (1971a) found that mating occurred from 28 April to 24 May, shortly after animals emerged from hibernation. The timing of the breeding season was correlated with elevation; high-elevation populations showed a 7 to 10-day lag in their breeding schedules. Of the males captured from 7 April to 30 May, 88.7% had scrotal testes (Beg 1971a). Non-breeding males were all yearlings. By late May, the testes begin to regress in size. The gestation period in the Red-tailed Chipmunk is assumed to be about 31 days (Best 1993). Beg (1971a) estimated that in his study area in Montana, parturition occurred from 26 May to 12 June. Mean litter size, based on embryo counts and placental scars, for Beg's (1971a) two study sites was 4.85. Females more than two-years old evidently produce larger litters than younger animals. Beg (1971a) estimated pregnancy rates (% of females pregnant) at 50.0–73.3%, which are lower than most other chipmunks. The pregnancy rate varies from year to year among sites and with sexual maturity. Only 11.1–15.0% of the yearling females in Beg's (1971a) study sites bred. Females produce only one litter each year.

2.3 Survival According to Beg (1971b), mortality is high in the first year; the survival rate is only 25%. Survivorship is higher after the first year. From recapture data, Beg (1971b) found that several Red-tailed Chipmunks survived three to five years, and one individual lived eight years. About 6–11% of the individuals survived to five years or older. Causes of mortality have not been documented. Potential predators in British Columbia include the Northern Goshawk (Accipter gentilis), Ermine (Mustela erminea), Coyote (Canis latrans), Long-tailed Weasel (Mustela frenata), American Marten (Martes americana), and Red Fox (Vulpes vulpes).

2.4 Physiology Physiology of the Red-tailed Chipmunk has not been studied, but this species is presumably a facultative hibernator, entering torpor in winter. As is typical of all chipmunks, the Red-tailed Chipmunk does not accumulate fat reserves in autumn; it relies instead on seeds hoarded in its burrow for winter survival (Beg 1972). It remains inactive in its underground burrow from mid November until late March. In Montana, Beg (1969) observed Red-tailed Chipmunks above ground as late as 4 December and as early as 29 March. Males appear to emerge in the spring earlier than females.

2.5 Movements/Dispersal No radio-telemetry studies have been done on this rodent. The only data on movements are based on recapture data from Beg (1970). He estimated that the maximum distances moved averaged 117 m (adult males), 81 m (adult females), and 71 m (juveniles). The longest movements observed were 458 m (adult males), 259 m (adult females), and 290 m (juveniles). The longest movements, which were usually taken by males, occurred in June and July.

2.6 Nutrition and Interspecific Interactions The diet includes green plant material, fruits, seeds, and invertebrates, but seeds are the most important food item. From an analysis of cheek pouch contents, Beg (1969) identified 51 different

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types of seeds. He also documented, from direct observation, the use of flowers, leaves, fruits, and seeds of about 30 plant types. The diet varies seasonally in response to plant availability. In spring, most of the Red-tailed Chipmunk’s foraging activity is on the forest floor where it moves quickly through the ground vegetation feeding on flowers and leaves. Beg (1969) estimated that Red-tailed Chipmunks typically moved 46–60 m on the ground within 3 to 4 minutes while foraging. Common dandelion (Taraxacum officinale) and heart-leaved arnica (Arnica cordifolia) flower heads are favourite food items at this time of year. In late summer and particularly autumn, the diet shifts to fruits and seeds, with much of the foraging occurring above ground in shrubs or trees. Seeds of bull thistle (Cirsium vulgare), ponderosa pine (Pinus ponderosa), common snowberry (Symphoricarpos albus), saskatoon (Amelanchier alnifolia), and mallow ninebark (Physocarpus malvaceus) are staple foods, and they form the bulk of seeds cached for winter stores. Although he made no quantitative estimates, Beg (1969) observed that immense quantities of seeds were carried in the cheek pouches in autumn. Lockner (1972) concluded that the Red-tailed Chipmunk was a “natural hoarder” that demonstrated both scatter hoarder and larder hoarder behaviours. Given its diverse diet and ability to respond to seasonal availability, food is not limiting for the Red-tailed Chipmunk. The most likely competitor for food resources is T. amoenus. In Montana, Beg (1969) found no significant differences in diet or foraging habits between the two species. Although their feeding niches overlap, competition between the two species in British Columbia is limited. In the Rocky Mountains, the two species are contiguously allopatric along an elevational gradient, co-occurring only in limited contact areas; in the Selkirk Mountains, where T. ruficaudus is widespread, T. amoenus appears to be rare—it is known from only a few locations (Nagorsen et al. 2002; Good et al. 2003).

2.7 Behaviour and Adaptability The social biology of the Red-tailed Chipmunk is unknown, but the species is presumably solitary. No estimates of home range area exist for this species. Using distances between captures in live traps, Beg (1970) found that adult males occupied larger home ranges than females and juveniles. Breeding and non-breeding adult females showed no differences in their home range size. After emerging from their maternal burrow, juveniles occupy small home ranges; however, by August, they move similar distances to adults. The Red-tailed Chipmunk is the most arboreal of the four chipmunk species found in British Columbia. Dens are located either in underground burrows or above ground in trees. Underground burrows are used mostly in winter and in spring, whereas tree nests are used in summer (Broadbrooks 1974). The young may be born in underground burrows or in tree nests. Two underground burrows examined by Broadbrooks (1974) were about 90 cm long and 28 cm deep, and had a nest of dried grass and lichen at one end. Three tree nests found in eastern Washington (Broadbrooks 1974) were all in Englemann spruce (Picea englemannii) trees that were 12–24 m in height. The nests, constructed from dead grass, were situated near the trunks, in dense growths of branches and small twigs located under witches brooms, about 5.8–18.3 m above ground. The tree nests were about 36 cm in diameter. Orr (1943) observed cavities in a dead fir tree 5.4 m high that supported six individual Red-tailed Chipmunks. Rust (1946) found a nest (17.8 x 25.4 cm) made of

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dried grass located in the forks of a bush; it contained four small, undeveloped young that were only about one-third grown with unopened eyes. Despite its limited range in Canada, the Red-tailed Chipmunk is a generalist, scansorial rodent capable of exploiting a wide range of habitats, including those disturbed by fire or forest harvesting. This species also exploits man-made habitats and often becomes habituated to campgrounds and picnic areas in parks.

3 HABITAT

3.1 Habitat Requirements In Canada, the two subspecies of the Red-tailed Chipmunk demonstrate major habitat differences. In the Selkirk Mountains of southern British Columbia, T. r. simulans inhabits a broad elevational range from 549 m in the floodplain of the Creston valley to 1829 m on the Salmo-Creston Summit (Appendix 1). Known occurrences are in the Interior Cedar–Hemlock (ICHmw, ICHdw) and Englemann Spruce–Subalpine Fir (ESSFwc) biogeoclimatic zones (Nagorsen et al. 2002). Habitats used include forests of western hemlock (Tsuga heterophylla), Englemann spruce, and western redcedar (Thuja plicata) of various ages from mature to recent clearcuts. T. r. simulans seems to be most common in disturbed habitats, such as clearings, roadsides, and forest edges with abundant shrubs and woody debris on the ground (Figures 1, 2). T. r. simulans occupies similar habitats in Washington State (Johnson and Cassidy 1997). In the Rocky Mountains of British Columbia, T. r. ruficaudus is restricted to a narrow elevational range from about 1785 to 1950 m (Appendix 1) where it is associated with subalpine coniferous forests. The few known occurrences are all within the Englemann Spruce–Subalpine Fir (ESSFdk) biogeoclimatic zone (Nagorsen et al. 2002). In Akamina-Kishinena Provincial Park, this chipmunk was found in a mature Englemann spruce–subalpine fir (Abies lasiocarpa) forest with an understory of thimbleberry (Rubus parviflorus) and black gooseberry (Ribes lacustre) (Figure 3). In Middle Kootenay Pass, it inhabited mature Englemann spruce–subalpine fir forests and disturbed open areas with abundant woody debris created by a recent forest fire (Figure 4). In the southern Rocky Mountains of Alberta, T. r. ruficaudus occupies a similar narrow elevational range from 1586 to 2135 m (Soper 1973). Habitat data for Alberta are based largely on data from Waterton Lakes National Park. According to Nielsen (1973), T. r. ruficaudus is associated with mixed-wood forest, spruce–fir associations, and krummholz. Wallis and Wershler (1997) reported that T. r. ruficaudus prefers open Englemann spruce–subalpine fir, especially with bear-grass (Xerophyllum tenax); other habitats used include stunted subalpine fir–larch (Larix sp.) and open subalpine fir–larch–whitebark pine (Pinus albicaulis). Both subspecies readily exploit habitats recently disturbed by forest harvesting or fire (Ramirez and Hornocker 1981). Scrivener and Smith (1984) found Red-tailed Chipmunks in forests of various ages, but they were most abundant in mid successional stages (11–79 years old) that follow timber harvesting. One of Scrivener's and Smith's early successional study areas (1–10 years old) also supported a high abundance of Red-tailed Chipmunks. Clearcuts that were lightly burned were recolonized by Red-tailed Chipmunks one year following the burn; heavily burned sites where the logging slash was destroyed, were not recolonized until three years after the burn (Halvorson 1982).

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Figure 1. Logged area, Church Creek. T. r. simulans inhabited second- growth and edge habitats. (Photograph by M. Fraker)

Figure. 2. Clearcut area, Giveout Creek. T. r. simulans inhabited edge habitats and second-growth forest. (Photograph by M. Fraker)

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Figure 3. Wall Lake, Akamina-Kishinena Provincial Park. T. r. ruficaudus inhabited forest bordering the lake. (Photograph by M. Fraker)

Figure 4. Middle Kootenay Pass. T. r. ruficaudus inhabited the subalpine forest and lower portions of the old burn. (Photograph by D.W. Nagorsen)

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The narrow subalpine elevational zone inhabited by T. r. ruficaudus in the Rocky Mountains of Canada reflects competitive exclusion with T. amoenus and the Least Chipmunk (T. minimus). Beg (1969) concluded that the entire fundamental niche of T. ruficaudus was a subset the T. amoenus fundamental niche, and that species excludes T. ruficaudus from low elevation forests. The absence of T. ruficaudus above treeline can be attributed to the presence of the smaller T. minimus, a chipmunk that is better adapted to extreme alpine conditions (Sheppard 1965). Hoffmann (1974) described a similar zonation pattern for the three species in Glacier National Park, Montana. Although these chipmunks are largely allopatric along an elevational gradient, there are narrow zones where they overlap in their distributions. At Middle Kootenay Pass, for example, Nagorsen et al. (2002) found all three chipmunk species coexisting in an elevational transect spanning a few hundred metres elevation. In the southern Selkirk Mountains of British Columbia where T. minimus is absent and T. amoenus is rare, T. ruficaudus inhabits lowland valleys, mid-elevation, and subalpine forests. It probably also occurs in alpine areas. Although T. ruficaudus and T. amoenus are generally segregated in their distributions along an elevational gradient, this simple pattern is not evident in the Selkirk Mountains. The only verified (from genital bone morphology and mtDNA) occurrence of T. amoenus in the southern Selkirk Mountains of British Columbia is near Topaz Creek at 900 m elevation (Nagorsen et al. 2002; Good et al. 2003). Historical museum records (identification unconfirmed) also exist for Fruitvale in the Beaver River valley, Nelson, and the west side of the Creston valley (Nagorsen, unpublished data). These locations are within the elevational range of T. r. simulans. In the Selkirk Mountains, these two chipmunks may exploit different microhabitats. Beg (1969) described subtle habitat differences between T. r. ruficaudus and T. amoenus in Montana that were related to topography, aspect, and slope.

3.2 Trends in Habitat Quality and Quantity Although Red-tailed Chipmunk habitat has been modified by forest harvesting, road construction, and small-scale farming, this species has adapted to various successional stages. There is no evidence of declines in the quality or quantity of habitat in British Columbia.

3.3 Protection/Ownership There are no quantitative data on land tenure classification for the Red-tailed Chipmunk’s range in British Columbia. Except for three provincial parks totalling 27 878 ha, and small private farmlands on the west side of the Creston valley, most of the range of T. r. simulans is on Crown land. In contrast, two of the five known occurrences of T. r. ruficaudus in British Columbia are in Akamina-Kishinena Provincial Park (10 291 ha). The Middle Kootenay Pass area is Crown land.

4 DISTRIBUTION

4.1 Global/ North American The Red-tailed Chipmunk has a small distributional range that is confined to northern Montana and Idaho, northeastern Washington, southeastern British Columbia, and southwestern Alberta (Figure 5). About 10% of this species' range occurs in Canada.

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4.2 British Columbia Although most published maps (see Figure 5) show the Red-tailed Chipmunk continuously distributed throughout southern British Columbia, Nagorsen et al. (2002) demonstrated that T. r. simulans and T. r. ruficaudus are disjunct in British Columbia and are separated by 180 km and major physiographic barriers such as the Rocky Mountain Trench and Purcell Mountains (Figure 6).

T. r. simulans is restricted to the southern Selkirk Mountains where its range is delimited by the Kootenay and Columbia Rivers. The eastern limits of its range are delimited by Kootenay Lake and the Creston valley. Anderson (1934) and Munro (1950) concluded that T. r. simulans did not occur on the east side of the Creston valley, a distributional pattern that was confirmed by Nagorsen et al. (2002). Historical museum specimens from the Purcell Mountains that were identified as T. r. simulans are actually misidentified T. amoenus. Maillard (1932) collected a series of chipmunks from the Creston area on the east side of the Kootenay River that he identified as T. r. simulans, but these specimens appear to be T. amoenus (Nagorsen, unpublished data). Cowan and Guiguet (1965) showed the range of T. r. simulans extending into the Purcell Mountains and the Rocky Mountain Trench, presumably on the basis of Crowe's (1943) T. r. simulans specimens from Invermere. Nagorsen et al. (2002) determined that Crowe's specimens were actually T. amoenus. T. r. simulans has not been found west of the Columbia River or on the north side of the Kootenay River and West

Figure 5. Distribution of the Red-tailed Chipmunk (Tamias ruficaudus) in North America. 1 = T. r. ruficaudus, 2 = T. r. simulans. (Taken from Hall 1981)

9

Arm of Kootenay Lake in British Columbia (Figure 6). The area of occupancy of T. r. simulans in British Columbia is unknown; its extent of occurrence is about 4070 km2. This represents about 10–15% of this subspecies' global range.

In British Columbia, T. r. ruficaudus is restricted to a small area in the southern Rocky Mountains where it ranges from the Akamina Pass–Wall Lake area north to Middle Kootenay Pass (Figure 6). On the Alberta side of the Rocky Mountains where it is mostly confined to Waterton Lakes National Park, the northernmost record is from the headwaters of the Castle River east of Middle Kootenay Pass (Bennett 1999). The precise distributional area of T. r. ruficaudus in Canada is not clear. This chipmunk could inhabit subalpine areas in the mountains west of the Flathead River in British Columbia. Similarly, there are no physiographic barriers in the Middle Kootenay Pass–Castle River area that would delimit the northern limits of its range. Nagorsen et al. (2002) hypothesized that the range may extend as far north as Crowsnest Pass, a wide, low-elevation pass (1360 m) that could be a dispersal barrier to the Red-tailed Chipmunk. The area of occupancy of T. r. ruficaudus in British Columbia is unknown; its known extent of occurrence is about 150 km2. This represents about 5–10% of this subspecies’ global range.

Figure 6. Distribution of the Red-tailed Chipmunk (Tamias ruficaudus) in British Columbia. Taken from Nagorsen et al. (2002); map by Ian Parfitt, Columbia Fish and Wildlife Compensation Program.

10

5 POPULATION SIZE AND TRENDS The only population estimates are Beg's (1971b) live capture data for Montana. He estimated densities of about 4–13 animals per hectare in his study grid. Year-to-year variation over the three years of his study was minor, but the Red-tailed Chipmunk demonstrated pronounced seasonal variation. Populations were small in May but increased in June because of immigration. Peak populations occurred in July and early August with the emergence of young-of-the-year from their maternal burrows. In response to mortality and emigration, populations declined in August and September. Population density is habitat dependent and population numbers evidently change rapidly with habitat disturbances such as fire and forest harvesting (Halvorson 1982; Scrivener and Smith 1984). The total population size and the number of mature individuals of the Red-tailed Chipmunk cannot be determined. No data exist on population trends. Its area of occupancy is unknown, and no habitat suitability models have been developed. The Canadian populations of T. r. simulans and T. r. ruficaudus are linked to adjacent populations in the United States. The map in Johnson and Cassidy (1997) suggests that T. r. simulans is found in counties adjacent to the British Columbia border. No physiographic barriers exist in Washington State that would impede cross border movements. T. r. ruficaudus occupies a narrow elevational zone in the southern Rocky Mountains of Canada, but populations in Alberta and British Columbia are connected to adjacent populations Montana. Although populations inhabiting the east and west sides of the Canadian Rocky Mountains are separated by alpine populations of the Least Chipmunk, passes such as Akamina Pass, Sage Pass, South Kootenay Pass, and Middle Kootenay Pass provide corridors for dispersal and gene-flow among British Columbia and Alberta populations of T. r. ruficaudus.

6 LIMITING FACTORS AND THREATS Threats and limiting factors are few. The narrow elevational zonation of T. r. ruficaudus in the Rocky Mountains appears to reflect interspecific competition with T. amoenus and T. minimus rather than habitat limitations. Forest harvesting is occurring within the range of both subspecies in British Columbia, but any disturbance from logging is short term because this species readily exploits early successional stages. Bennett (1999) identified two possible threats to this species in Alberta: logging in the Castle River area and fire suppression in Waterton Lakes National Park. Although Bennett (1999) argued that T. r. ruficaudus preferred mature forest, this seems inconsistent with the literature (Halvorson 1982; Scrivner and Smith 1984). Jeff Good (pers. comm.) captured most of his Red-tailed Chipmunks in northern Montana in roadsides, second-growth forest, and ~10-year old clearcuts with no substantial cover. Nagorsen et al. (2002) reported similar habitats for their capture sites of T. r. simulans in British Columbia. Any negative effects from forest harvesting are short term. Although this chipmunk is arboreal and uses tree dens, it survives in early disturbed habitats where it presumably excavates underground burrows. With most of the known Canadian occurrences of T. r. ruficaudus confined to two protected areas (Waterton Lakes National Park and Akamina-Kishinena Provincial Park), fire suppression is of concern. Bennett (1999) suggested that fire suppression could reduce the understory, forest openings, and edges. Alternatively, an intense fire may destroy coarse woody

11

debris, an essential component of chipmunk habitat. Another potential threat in the southern Rocky Mountains of British Columbia is open pit coal mining associated with coalfields. However, according to the map of coalfields in the Southern Rocky Mountain Management Plan (British Columbia Ministry of Sustainable Resource Management 2003), the known range of T. r. ruficaudus does not coincide with any coalfields. The nearest coalfields are on the west side of the Flathead River valley.

7 SPECIAL SIGNIFICANCE OF THE SPECIES The Red-tailed Chipmunk is a somewhat obscure, little known, small mammal in Canada. With its limited range in the Rocky and Columbia Mountains, this chipmunk is a regional endemic and a model for understanding evolutionary processes and historical biogeography in the western Cordillera. A recent phylogeographic study by Good and Sullivan (2001) demonstrated that the patterns of genetic divergence in this species reflect complex multiple isolation events in refugia that were associated with past glaciation events. The curious allopatric distribution of the two subspecies in Canada reflects postglacial dispersal patterns and competition effects from other chipmunks.

8 EXISTING PROTECTION OR OTHER STATUS The Red-tailed Chipmunk is not listed in the International Union for Conservation of Nature and Natural Resources (IUCN) Red Book, and it has not been assessed by the Committee on the Status of Endangered Wildlife in Canada (COSEWIC). Globally, the species is listed as G5. In Idaho and Montana, it is listed as S4 and S5. These jurisdictions do not give separate designations for the two subspecies. The Washington designation of S2? applies to T. r. simulans, the only subspecies found in the state. In Canada, the Red-tailed Chipmunk is listed nationally as N3N4 (vulnerable to secure). The Alberta designation, which applies to the subspecies T. r. ruficaudus, is S2 (imperilled). The British Columbia status is S2S3 (imperilled to vulnerable), and the species is on British Columbia's Blue List (Cannings et al. 1999). British Columbia has also listed the two subspecies, with T. r. ruficaudus S2 (imperilled) and T. r. simulans S3S4 (vulnerable to secure). In British Columbia, the Red-tailed Chipmunk is protected from killing under the provincial Wildlife Act. However, its habitat is protected only in provincial parks and ecological reserves. This rodent is not listed as an Identified Wildlife Species by the province and, therefore, has no protection under the revised British Columbia Forest Practices Code. Three provincial parks are within the distributional range of T. r. simulans: Stagleap Provincial Park (1133 ha), West Arm Provincial Park (25 319 ha), and Champion Lakes Provincial Park (1426 ha). Much of the range of T. r. ruficaudus in British Columbia is in Akamina-Kishinena Provincial Park (10 291 ha). This park is contiguous with Waterton Lakes National Park in Alberta.

12

9 RECOMMENDATIONS AND MANAGEMENT OPTIONS

9.1 Inventory Despite the recent field surveys by Nagorsen et al. (2002), more inventory work needs to be done. Existing records strongly suggest that T. r. simulans is absent from the Purcell Mountains. Nevertheless, no chipmunk inventories have been done on the moist west slope of the Purcell Mountains above Kootenay Lake. Similarly, more surveys are required to verify the absence of this subspecies from areas north of the Kootenay River and west of the Columbia River. T. r. ruficaudus is known from only five general locations in British Columbia. No surveys have been done in the subalpine areas in the Crowsnest Pass area to establish the northern limits of its range. Potential subalpine habitat exists in some of the mountains on the west side of the Flathead River valley, an area that has not been studied.

9.2 Habitat Protection and Acquisition Much of the Canadian range of T. r. ruficaudus is in large, contiguous, protected areas. A high priority for research and management is a study on the role of fire in maintaining Red-tailed Chipmunk habitat in the southern Rocky Mountains. Acquiring more protected area is not critical for managing T. r. simulans, a subspecies that seems to be adaptable to habitat disturbances. More research should be done in the Selkirk Mountains to determine the distribution of T. amoenus and its effects on the distribution and abundance of T. r. simulans.

10 EVALUATION

10.1 Provincial Ranking The two subspecies T. r. simulans and T. r. ruficaudus represent nationally significant populations and warrant separate rankings.

10.1.1 T. r. simulans (see Appendix 3) This taxon was ranked as S2 (Red List) by Cannings et al. (1999), but is currently listed as S3S4 by the British Columbia Conservation Data Centre, largely on the basis of its small distributional area. Compared to T. r. ruficaudus, T. r. simulans occupies a broader elevational range and more habitats, including the floodplain of the Creston valley, mid elevation forests (mature and logged), and subalpine habitat in Stagleap Provincial Park. Its population is contiguous with those in Washington and Idaho; therefore, there is considerable potential for rescue effect. No threats are known. Number of Subpopulations: None. Number of Occurrences: Using the recommended separation distance criteria of 1 km, the number of known occurrences for the British Columbian range is estimated at about 15 (Appendix 1). Because this includes both historical and recent records, the number of occurrences with good viability is unknown. Population Size: The current population size and number of mature individuals in British Columbia are unknown.

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Population Trends: No data are available on population trends. Population density fluctuates seasonally. There is no evidence of declines in suitable habitat. Extent of Occurrence: The total extent of occurrence is about 4070 km2. Area of Occupancy: The area of occupancy is unknown. Threats: There are no obvious threats to this subspecies. Environmental Specificity: Low-a generalist with all key requirements common. Number of Protected Occurrences: One known occurrence (Kootenay Pass) is in Stagleap Provincial Park. T. r. simulans probably occurs in West Arm and Champion Lakes Provincial Parks, but this has not been confirmed. Intrinsic Vulnerability: Not intrinsically vulnerable. 10.1.2 T. r. ruficaudus (see Appendix 2) This subspecies is ranked as S2 (Red List) in British Columbia because of its limited range and few known locations (Cannings et al. 1999). Similarly, it is ranked as S2 by the Alberta Natural Heritage Information Centre and is on that province’s Sensitive List (see Bennett 1999). In British Columbia, T. r. ruficaudus has a small range and is limited to a narrow elevational zone. Nonetheless, much of its distributional range occurs within the boundaries of Akamina-Kishinena Provincial Park. Moreover, because the British Columbia populations are connected with populations in Alberta and adjacent areas of Montana, there is potential for a rescue effect. Although extensive logging is occurring within this subspecies’ elevational range in the Flathead River valley of British Columbia, the subspecies inhabits early and later successional stages. Fire suppression in national and provincial parks is the only potential impact on this subspecies. The known range of this subspecies in British Columbia is outside the range of known coalfields. Number of Subpopulations: None. Number of Occurrences: Using the recommended separation distance criteria of 1 km, the number of known occurrences for the British Columbian range is estimated at about five (Appendix 1). The number of occurrences with good viability is unknown. Population Size: The current population size and number of mature individuals in British Columbia are unknown.

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Population Trends: No data are available on population trends. Population density fluctuates seasonally. There is no evidence of declines in suitable habitat. Extent of Occurrence: The extent of occurrence is about 150 km2. Area of Occupancy: The area of occupancy is unknown. Threats: The only potential threat to this subspecies is habitat loss from fire suppression in Akamina-Kishinena Provincial Park. Scope: low, < 20–60% of the area affected Severity: low Imminence: high, threat is operational Environmental Specificity: Low-a generalist with all key requirements common. Number of Protected Occurrences: Of the five known occurrences, two occur within Akamina-Kishinena Provincial Park, and one borders Waterton Lakes National Park. Intrinsic Vulnerability: Not intrinsically vulnerable.

10.2 COSEWIC Ranking The Red-tailed Chipmunk has not been assessed by COSEWIC. Because the entire Canadian range of T. r. simulans occurs within British Columbia, data in this report could be used to make a COSEWIC assessment. However, any COSEWIC assessment of T. r. ruficaudus must include data from the Alberta population. The technical summary for T. r. ruficaudus (Appendix 2) in this report is based only on the British Columbian population. For a national COSEWIC status assessment, categories such as extent of occurrence and number of extant locations must include the entire Canadian range. Although an Alberta provincial status report has been written for T. r. ruficaudus (Bennett 1999), the report is not in the COSEWIC format and lacks essential data for a COSEWIC designation.

Because the population size and number of mature individuals of either subspecies are unknown, the only COSEWIC criterion based on the IUCN Red List categories that can be applied to them is criterion B. Although the extent of occurrence of these subspecies is less than 20 000 km2, T. r. simulans and T. r. ruficaudus do not meet two of the three conditions, and therefore do not qualify as Endangered or Threatened. Despite its provincial listing as S3S4, T. r. simulans is probably Not at Risk. T. r. ruficaudus may qualify for the COSEWIC designation of Special Concern because of its few known occurrences and small range that is restricted to a narrow elevational range.

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11 AUTHORITIES CONSULTED Antifeau, Ted. Rare and Endangered Species Biologist, B.C. Ministry of Water, Air and Land

Protection, 401-333 Victoria Street, Nelson, BC V1L 4K3. Tel: (250) 354-6163. Email: Ted.Antifeau@gems5.gov.bc.ca

Good, Jeff. Graduate student, Department of Ecology and Evolutionary Biology, University of Arizona, Biosciences West, Tucson, AZ 85721. Email: jgood@email.arizona.edu

Grieve, Dave. Mineral Land Planner, B.C. Ministry of Sustainable Resources Management, 205 Industrial Road G, Cranbrook, BC V1C 7G5. Tel: (250) 489-8514. Email: Dave.Grieve@gems6.gov.bc.ca

Guy, Stewart. Project Manager, Wildlife Conservation Planning, B.C. Ministry of Water, Land and Air Protection. PO Box 9338 Stn Prov Govt, Victoria, BC V8W 9M1. Tel: (250) 387-0060. Email: Stewart.Guy@gems9.gov.bc.ca.

Fraker, Mark. Consultant, TerraMar Environmental Research Ltd., Sidney, BC V8L 1M8. Tel: (250) 656-3972. Email: maf@terramar.bc.ca

Krebs, John. Wildlife Biologist, Columbia Basin Fish and Wildlife Compensation Program, 401-333 Victoria Street, Nelson, BC V1L 4K3. Tel: (250) 352-6874. Email: John.Krebs@bchydro.bc.ca

Ramsay, Leah. B.C. Ministry of Sustainable Resources Management, Conservation Data Centre, PO Box 9344 Stn Prov Govt, Victoria, BC V8W 9M1. Tel: (250) 387-9524. Email: Leah.Ramsay@gems4.gov.bc.ca

12 LITERATURE CITED Anderson, R.M. 1934. Review of the "Birds and mammals from the Kootenay valley,

southeastern British Columbia". Can. Field-Nat. 48:21–24. Anderson, R.M. 1946. Catalogue of Canadian recent mammals. Nat. Mus. Can. Bull. 102:1–238. Beg, M.A. 1969. Habitats, food habits, and population dynamics of the red-tailed chipmunk,

Eutamias ruficaudus. Thesis, Univ. Montana, Missoula. 153pp. Beg, M.A. 1970. Movement patterns in the red-tailed chipmunk, Eutamias ruficaudus. Pakistan J.

Zool. 2:199–206. Beg, M.A. 1971a. Reproductive cycle and reproduction in red-tailed chipmunks, Eutamias

ruficaudus. Pakistan J. Zool. 3:1–13. Beg, M.A. 1971b. Population dynamics of the red-tailed chipmunk, (Eutamias ruficaudus), in

western Montana. Pakistan J. Zool. 3:133–145. Beg, M.A. 1972. Seasonal changes in body weight of the red-tailed chipmunk, Eutamias

ruficaudus. Pakistan J. Zool. 4:13–16. Bennett, R. 1999. Status of the red-tailed chipmunk (Tamias ruficaudus) in Alberta. Alberta

Environment. Prot., Fish. Wildl. Div., Alberta Conserv. Assoc., Edmonton, Alta. Report No. 19. 15pp.

Best, T.L. 1993. Tamias ruficaudus. Mamm. Species 452:1–7. British Columbia Ministry of Sustainable Resource Management. 2003. Southern Rocky

Mountain Management Plan. Public Review Draft, February 14, 2003. Victoria, B.C. 111pp.+ maps.

Broadbrooks, H.E. 1974. Tree nests of chipmunks with comments on associated behavior and ecology. J. Mammal. 55:630–639.

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Cannings, S.G., L.R. Ramsay, D.F. Fraser, and M.A. Fraker. 1999. Rare amphibians, reptiles, and mammals of British Columbia. B.C. Minist. Environ., Lands and Parks, Wildl. Branch and Resour. Inventory Branch, Victoria, B.C. 198pp.

Cowan, M., and C.J. Guiguet. 1965. The mammals of British Columbia. B. C. Prov. Mus., Victoria, BC. 414pp.

Crowe, P.E. 1943. Notes on some mammals of the southern Canadian Rocky Mountains. Am. Mus. Nat. Hist. Bull. 80:391–410.

Good, J.M., and J. Sullivan. 2001. Phylogeography of the red-tailed chipmunk (Tamias ruficaudus), a northern Rocky Mountain endemic. Molecular Ecol. 10:2683–2695.

Good, J.M., J. Demboski, D.W. Nagorsen, and J. Sullivan. 2003. Phylogeography and introgressive hybridization: chipmunks (genus Tamias) in the northern Rocky Mountains. Evolution 57:1900–1916.

Hall, E.R. 1981. The mammals of North America. Vol. 1. John Wiley and Sons, New York, N.Y. 600+90pp.

Halvorson, C.H. 1982. Rodent occurrence, habitat disturbance, and seed fall in a larch-fir forest. Ecology 63:423–433.

Hoffmann, R.S. 1974. Terrestrial vertebrates. Pp. 475–568 in J.D. Ives and G.B. Roger, eds. Arctic and alpine environments. Methuen and Company Limited, London. 999pp.

Howell, A.B. 1920. Description of a new chipmunk from Glacier National Park, Montana. Proc. Biol. Soc. Wash. 33:91–92.

Howell, A.B. 1929. Revision of the American chipmunks. North Am. Fauna 52:1–157. Jameson, E.W.J. 1999. Host-ectoparasite relationships among North American chipmunks. Acta

Theriologica 44:225–231. Johnson, R.E., and K.M. Cassidy. 1997. Terrestrial mammals of Washington State: location data

and predicted distributions. Pg. 304 in K.M. Cassidy, C.E. Grue, M.R. Smith, K.M. Dvornich, eds. Washington Sate Gap Analysis-Final Report. Wash. Coop. Fish Wildl. Res. Unit, Univ. Washington, Seattle, Wash.

Lockner, F.R. 1972. Experimental study of food hoarding in the red-tailed chipmunk, Eutamias ruficaudus. Zeitschrift fur Tierpsychologie 31:410–418.

Maillard, J. 1932. Birds and mammals from the Kootenay valley; southeastern British Columbia. Proc. Calif. Acad. Sci. 20:269–290.

Munro, J.A. 1950. The birds and mammals of the Creston region, British Columbia. B.C. Prov. Mus., Occas. Pap. 8:1–89.

Nagorsen, D.W. 2002. An identification manual to the small mammals of British Columbia. B.C. Minist. Sustainable Resour. Manag., B.C. Minist. Water, Land, and Air Prot., and R.B.C. Mus., Victoria, B.C 153pp.

Nagorsen, D.W., N. Panter, and M.A. Fraker. 2002. Chipmunks of the Kootenay region, British Columbia: distribution, identification, taxonomy, conservation status. Columbia Basin Fish and Wildlife Compensation Program, Nelson, B.C. 142+viii pp. [http://www.cbfishwildlife.org/reports/chipmunk_pdfs.php]

Nielsen, P.L. 1973. Mammals of Waterton Lakes National Park. Can. Wildl. Serv., Edmonton, Alta. 176pp.

Orr, R.T. 1943. Mammals of the Clearwater Mountains, Idaho. Proc. Calif. Acad. Sci. 23:511–536.

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Panian, M.A. 1996. Chipmunks in the Kootenays: red-tailed chipmunk subspecies simulans. Columbia Basin Fish and Wildlife Compensation Program, Nelson, B.C. 69pp. [http:/srmwww.gov.bc.ca/kor/wld/reports/zipfiles/chip.zip]

Patterson, B.D., and L.R. Heaney. 1987. Preliminary analysis of geographic variation in red-tailed chipmunks Eutamias ruficaudus. J. Mammal. 68:782–791.

Piaggio, A.J., and G.S. Spicer. 2001. Molecular phylogeny of chipmunks inferred from mitochondrial cytochrome b and cytochrome oxidase II gene sequences. Molecular Phylogenetics and Evolution 20:335–350.

Ramirez, P.J., and M. Hornocker. 1981. Small mammal populations in different aged clearcuts in northwestern Montana. J. Mammal. 62:400–403.

Rust, H.J. 1946. Mammals of northern Idaho. J. Mammal. 27:308–327. Scrivner, J.H., and H.D. Smith. 1984. Relative abundance of small mammals in four successional

stages of spruce-fir in Idaho. Northwest Sci. 58:171–176. Sheppard, D.H. 1965. Ecology of the chipmunks Eutamias amoenus luteiventris (Allen) and E.

minimus oreocetes Merriam, with particular reference to competition. Ph.D. Thesis, Univ. Saskatchewan, Saskatoon, Sask. 229pp.

Soper, J.D. 1973. The mammals of Waterton Lakes National Park Alberta. Can. Wildl. Serv. Rep. Ser. 23:1–55.

Wallis, C., and C. Wershler. 1997. Waterton Lakes National Park ecological land classification (wildlife component) 1995–96. Unpublished report. Canadian Heritage, Calgary. 195pp.

White, J.A. 1953. The baculum in the chipmunks of western North America. Univ. Kansas Publ., Mus. Nat. Hist. 5:611–631.

Wilson, D.E., and D.M. Reeder. 1993. Mammal species of the world. A taxonomic and geographic reference. Smithsonian Inst. Press, Washington, D.C. 1206pp.

Appendix 1. Summary of all known occurrences (museum specimens, literature records) of the Red-tailed Chipmunk (Tamias ruficaudus) in British Columbia. Because of identification problems in the Selkirk Mountains with the morphologically similar Yellow-pine Chipmunk (Tamias amoenus), occurrences of T. r. simulans are limited to specimens identified from genital bone morphology and mtDNA (see Good and Sullivan 2001; Nagorsen et al. 2002; Good et al. 2003). Source1 Location Zone Northing Easting Elevation

(m) T. r. ruficaudus CMN 16016 Akamina-Kishinena Prov Park ;

Akamina Pass; W of Alberta 11 5434000.00 715000.00 1798

RBCM 3571 Akamina-Kishinena Prov Park ; Akamina Pass; W of Alberta

11 5435000.00 713200.00 1798

UBC 1627 Akamina-Kishinena Prov Park ; Akamina Pass

11 5434000.00 715000.00 1798

UBC 1628 Akamina-Kishinena Prov Park ; Akamina Pass

11 5434000.00 715000.00 1798

UBC 1629 Akamina-Kishinena Prov Park ; Akamina Pass

11 5434000.00 715000.00 1798

RBCM 19683 Akamina-Kishinena Prov Park ; Wall Lake; 30 m from lake

11 5432200.00 713660.00 1785

RBCM 19875 Middle Kootenay Pass; Rainy Ridge; Site 004

11 5458944.00 689045.00 1950

RBCM 19880 Middle Kootenay Pass; Site 003 11 5458987.00 688667.00 1850 RBCM 19885 Middlepass; Site 005 11 5458628.00 689020.00 1880 RBCM 19884 Middlepass Creek; Site 005 11 5458723.00 689101.00 1880 RBCM 19887 Middlepass Creek; Site 005 11 5458434.00 688795.00 1850 RBCM 19906 Middlepass Creek; Upper; Site 003 11 5458701.00 688446.00 1785 RBCM 19907 Middlepass Creek; Upper; Site 003 11 5458656.00 688217.00 1795 RBCM 19914 Middlepass Creek; Upper; Site 003 11 5458519.00 688562.00 1810 RBCM 19915 Middlepass Creek; Upper; Site 003 11 5458554.00 689046.00 1825 RBCM 19916 Middlepass Creek; Upper; Site 003 11 5458879.00 688246.00 1795 RBCM 19917 Middlepass Creek; Upper; Site 003 11 5458883.00 688355.00 1795 RBCM 19918 Middlepass Creek; Upper On Road;

Site 003 11 5458827.00 688345.00 1845

RBCM 19919 Middlepass Creek; Upper; Site 003 11 5458806.00 688273.00 1845 Cowan and Guiguet (1965)

Sage Pass 11 5446500.00 706800.00 -

RBCM 19920 Middlepass Creek; Upper; Site 003 11 5458595.00 688620.00 1820 T. r. simulans ROM 28444 Boundary Lake 11 5428000.00 508000.00 1311 RBCM 19656 Church Creek; Church Creek Road 11 5428449.00 466640.00 1086 RBCM 19666 Church Creek; Church Creek Road,

Between Km 9 & 10 11 5429000.00 466999.00 1086

RBCM 19667 Church Creek; Church Creek Road 11 5428349.00 467834.00 1086 RBCM 19668 Church Creek; Church Creek Road 11 5428349.00 467834.00 1086 Panian (1996)3 Clearwater Creek 11 5470000.00 488000.00 1460 CMN 10169 Creston; French's Farm 11 5437000.00 531000.00 549 ROM 28422 Creston; Kootenay Flats 11 5437000.00 531000.00 549

19

20

ROM 28453 Creston; West; 200 Ft Above Kootenay Flats

11 5437000.00 531000.00 549

ROM 28454 Creston; West; 200 Ft Above Kootenay Flats

11 5439000.00 531000.00 549

RBCM 19658 Giveout Creek; Giveout Ck Fs Rd Between Km 9 & 10

11 5479133.00 475690.00 1348

RBCM 19659 Giveout Creek; Giveout Ck Fs Rd Between Km 9 & 10

11 5479133.00 475690.00 1348

RBCM 19661 Giveout Creek; Giveout Ck Fs Rd Between Km 9 & 10

11 5479133.00 475690.00 1348

RBCM 19662 Giveout Creek; Giveout Creek Fs Road, Km 9.9

11 5428349.00 467834.00 1470

RBCM 20038 Giveout Creek; Forest Service Rd,14 Km From Hwy 3a; site 99-12

11 5478381.00 473995.00 1390

RBCM 19660 Giveout Creek; Giveout Ck Fs Rd Between Km 9 & 10

11 5479133.00 475690.00 1348

RBCM 19654 Gold Creek; Km 4.9 Gold Creek Road

11 5476042.00 479446.00 1470

RBCM 19655 Gold Creek; Km 4.9 Gold Creek Road

11 5476042.00 479446.00 1470

RBCM 20036 Gold Creek; Site 99-18 11 5143608.00 478430.00 1240 CMN 41264 Kootenay Pass, South; 30 Mi W

Creston 11 5434700.00 496400.00 1829

CMN 41266 Kootenay Pass, South; 30 Mi W Creston

11 5434700.00 496400.00 1829

CMN 41267 Kootenay Pass, South; 30 Mi W Creston

11 5434700.00 496400.00 1829

CMN 41269 Kootenay Pass, South; 30 Mi W Creston

11 5434700.00 496400.00 1829

CMN 41272 Kootenay Pass, South; 30 Mi W Creston

11 5434700.00 496400.00 1829

CMN 41274 Kootenay Pass, South; 30 Mi W Creston

11 5434700.00 496400.00 1829

CMN 41277 Kootenay Pass, South; 30 Mi W Creston

11 5434700.00 496400.00 1829

CMN 41282 Kootenay Pass, South; 30 Mi W Creston

11 5434700.00 496400.00 1829

CMN 41283 Kootenay Pass, South; 30 Mi W Creston

11 5434700.00 496400.00 1829

CMN 41286 Kootenay Pass, South; 30 Mi W Creston

11 5434700.00 496400.00 1829

Panian (1996)3 Porcupine Creek 11 5458000.00 490000.00 1696 CMN 1008 Salmon River (=Salmo River?);

South of 11 5430000.002 485000.002 -

Panian (1996)3 Toad Mountain 11 5473000.00 478000.00 1450 1 CMN=Canadian Museum of Nature, Ottawa; RBCM=Royal British Columbia Museum, Victoria; ROM=Royal Ontario Museum, Toronto; UBC=Cowan Vertebrate Museum, University of British Columbia, Vancouver 2 coordinates arbitrary 3 identified from radiographs taken on live animals that showed the male genital bone morphology

Appendix 2. Technical Summary - Tamias ruficaudus ruficaudus Red-tailed Chipmunk–Rocky Mountains subspecies Alberta, British Columbia Extent and Area Information • extent of occurrence (EO)(km²) 1501 • specify trend (decline, stable, increasing,

unknown) Stable

• are there extreme fluctuations in EO (> 1 order of magnitude)?

No

• area of occupancy (AO) (km²) Unknown • specify trend (decline, stable, increasing,

unknown) -

• are there extreme fluctuations in AO (> 1 order magnitude)?

-

• number of extant locations 52 • specify trend in # locations (decline, stable,

increasing, unknown) Stable

• are there extreme fluctuations in # locations (>1 order of magnitude)?

No

• habitat trend: specify declining, stable, increasing or unknown trend in area, extent or quality of habitat

Stable

Population Information • generation time (average age of parents in the population)

(indicate years, months, days, etc.) 2 years

• number of mature individuals (capable of reproduction) in the Canadian population (or, specify a range of plausible values)

Unknown

• total population trend: specify declining, stable, increasing or unknown trend in number of mature individuals

Stable

• if decline, % decline over the last/next 10 years or 3 generations, whichever is greater (or specify if for shorter time period)

-

• are there extreme fluctuations in number of mature individuals (> 1 order of magnitude)?

-

• is the total population severely fragmented (most individuals found within small and relatively isolated (geographically or otherwise) populations between which there is little exchange, i.e., < 1 successful migrant / year)?

No

• list each population and the number of mature individuals in each

-

• specify trend in number of populations (decline, stable, increasing, unknown)

-

• are there extreme fluctuations in number of populations (>1 order of magnitude)?

-

Threats (actual or imminent threats to populations or habitats)

21

-open pit coal mines -fire suppression in provincial national parks Rescue Effect (immigration from an outside source) Moderate • does species exist elsewhere (in Canada or outside)? Yes • status of the outside population(s)? Not at Risk • is immigration known or possible? Yes • would immigrants be adapted to survive here? Yes • is there sufficient habitat for immigrants here? Yes Quantitative Analysis - David Nagorsen, January 2003 1 British Columbia portion of range only 2 Number of element occurrences for British Columbia portion of range only

22

Appendix 3. Technical Summary - Tamias ruficaudus simulans Red-tailed Chipmunk–Columbia Mountains Subspecies British Columbia Extent and Area Information • extent of occurrence (EO)(km²) 4070 • specify trend (decline, stable, increasing,

unknown) Stable

• are there extreme fluctuations in EO (> 1 order of magnitude)?

No

• area of occupancy (AO) (km²) Unknown • specify trend (decline, stable, increasing,

unknown) -

• are there extreme fluctuations in AO (> 1 order magnitude)?

-

• number of extant locations 151 • specify trend in # locations (decline, stable,

increasing, unknown) Stable

• are there extreme fluctuations in # locations (>1 order of magnitude)?

No

• habitat trend: specify declining, stable, increasing or unknown trend in area, extent or quality of habitat

Stable

Population Information • generation time (average age of parents in the population)

(indicate years, months, days, etc.) 2 years

• number of mature individuals (capable of reproduction) in the Canadian population (or, specify a range of plausible values)

Unknown

• total population trend: specify declining, stable, increasing or unknown trend in number of mature individuals

Stable

• if decline, % decline over the last/next 10 years or 3 generations, whichever is greater (or specify if for shorter time period)

-

• are there extreme fluctuations in number of mature individuals (> 1 order of magnitude)?

-

• is the total population severely fragmented (most individuals found within small and relatively isolated (geographically or otherwise) populations between which there is little exchange, i.e., < 1 successful migrant / year)?

No

• list each population and the number of mature individuals in each

-

• specify trend in number of populations (decline, stable, increasing, unknown)

-

• are there extreme fluctuations in number of populations (>1 order of magnitude)?

-

Threats (actual or imminent threats to populations or habitats) -no threats known

23

24

Rescue Effect (immigration from an outside source) High • does species exist elsewhere (in Canada or outside)? Yes • status of the outside population(s)? Not at Risk • is immigration known or possible? Yes • would immigrants be adapted to survive here? Yes • is there sufficient habitat for immigrants here? Yes Quantitative Analysis David Nagorsen, January 2003 1 elements of occurrence, some historical