Neotoma cinerea



INTRODUCTORY


  Photo courtesy of R.B. Forbes, Mammal Images Library of the American Society of Mammalogists

AUTHORSHIP AND CITATION:
Ulev, Elena 2007. Neotoma cinerea. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ [].

FEIS ABBREVIATION:
NECI

COMMON NAMES:
bushy-tailed woodrat
bushy-tailed packrat
packrat
trade rat
woodrat

TAXONOMY:
The currently accepted scientific name for the bushy-tailed woodrat is Neotoma cinerea (Ord) [15,16,63,126]. It is a member of the family Crecetidae [126]. The 13 currently recognized subspecies are listed below [63,64,126]:

Neotoma cinerea acraia (Elliot)
Neotoma cinerea alticola Hooper
Neotoma cinerea arizonae Merriam
Neotoma cinerea cinerea (Ord)
Neotoma cinerea cinnamomea J. A. Allen
Neotoma cinerea drummondii (Richardson)
Neotoma cinerea fusca True
Neotoma cinerea lucida Goldman
Neotoma cinerea macrodon Kelson
Neotoma cinerea occidentalis Baird
Neotoma cinerea orolestes Merriam
Neotoma cinerea pulla Hooper
Neotoma cinerea rupicola J. A. Allen

SYNONYMS:
None

ORDER:
Rodentia

CLASS:
Mammal

FEDERAL LEGAL STATUS:
No special status

OTHER STATUS:
None


ANIMAL DISTRIBUTION AND OCCURRENCE

SPECIES: Neotoma cinerea
GENERAL DISTRIBUTION:
Bushy-tailed woodrats occur from the Yukon Territory and Northwest Territories south to Arizona and New Mexico and from California east to the Badlands of the upper Missouri River drainage area of South Dakota and Nebraska [29,73,111].

The specific ranges of the 13 subspecies are listed below [63]:

Neotoma c. acraia: east-central California to southeastern Idaho, Utah, and northern Arizona

Neotoma c. alticola: southern Washington, Oregon, southern Idaho, northeastern Caliifornia, and northern Nevada

Neotoma c. arizonae: Utah, Colorado, New Mexico, and Arizona

Neotoma c. cinerea: southeastern British Columbia, southern Alberta, and southwestern Saskatchewan south to central Idaho, western Wyoming, Montana, and western North Dakota

Neotoma c. cinnomomea: southwestern Wyoming, northeastern Utah, and northwestern Colorado

Neotoma c. drummondii: Yukon and Northwest Territories south to eastern British Columbia and western Alberta

Neotoma c. fusca: the Coast Ranges of Washington and Oregon

Neotoma c. lucida: southeastern California and southwestern Nevada

Neotoma c. macrodon: east-central Utah

Neotoma c. occidentalis: from Yukon Territory south to Washington, the Cascade Range of Oregon, and northern Idaho

Neotoma c. orolestes: southeastern Montana and southwestern South Dakota to northern New Mexico

Neotoma c. pulla: southwestern Oregon to northern California

Neotoma c. rupicola: southeastern Montana and North Dakota to western Nebraska and northeastern Colorado [63]

The following lists are speculative and are based on the habitat characteristics and species composition of communities bushy-tailed woodrats are known to occupy. There is not conclusive evidence that bushy-tailed woodrats occur in all the habitat types listed, and some community types, especially those used rarely, may have been omitted. See Preferred Habitat for more detail.

ECOSYSTEMS [57]:
FRES19 Aspen-birch
FRES20 Douglas-fir
FRES21 Ponderosa pine
FRES22 Western white pine
FRES23 Fir-spruce
FRES24 Hemlock-Sitka spruce
FRES25 Larch
FRES26 Lodgepole pine 
FRES27 Redwood 
FRES28 Western hardwoods 
FRES29 Sagebrush 
FRES30 Desert shrub 
FRES34 Chaparral-mountain shrub 
FRES35 Pinyon-juniper 
FRES36 Mountain grasslands 
FRES37 Mountain meadows 
FRES39 Prairie 
FRES41 Wet grasslands 
FRES44 Alpine  

STATES/PROVINCES: (key to state/province abbreviations)
UNITED STATES

AZ CA CO ID MT NE NV NM ND OR
SD UT WA WY

CANADA
AB BC NT SK YK

BLM PHYSIOGRAPHIC REGIONS [19]:
1 Northern Pacific Border
2 Cascade Mountains
3 Southern Pacific Border
4 Sierra Mountains
5 Columbia Plateau
6 Upper Basin and Range
7 Lower Basin and Range
8 Northern Rocky Mountains
9 Middle Rocky Mountains
10 Wyoming Basin
11 Southern Rocky Mountains
12 Colorado Plateau
13 Rocky Mountain Piedmont
14 Great Plains
15 Black Hills Uplift
16 Upper Missouri Basin and Broken Lands

KUCHLER [79] PLANT ASSOCIATIONS:
K001 Spruce-cedar-hemlock forest
K002 Cedar-hemlock-Douglas-fir forest
K003 Silver fir-Douglas-fir forest
K004 Fir-hemlock forest
K005 Mixed conifer forest
K006 Redwood forest
K007 Red fir forest
K008 Lodgepole pine-subalpine forest
K010 Ponderosa shrub forest
K011 Western ponderosa forest
K012 Douglas-fir forest
K013 Cedar-hemlock-pine forest
K014 Grand fir-Douglas-fir forest
K015 Western spruce-fir forest
K016 Eastern ponderosa forest
K017 Black Hills pine forest
K018 Pine-Douglas-fir forest
K019 Arizona pine forest
K020 Spruce-fir-Douglas-fir forest
K021 Southwestern spruce-fir forest
K022 Great Basin pine forest
K023 Juniper-pinyon woodland
K024 Juniper steppe woodland
K025 Alder-ash forest
K026 Oregon oakwoods
K028 Mosaic of K002 and K026
K029 California mixed evergreen forest
K030 California oakwoods
K033 Chaparral
K034 Montane chaparral
K037 Mountain-mahogany-oak scrub
K038 Great Basin sagebrush
K040 Saltbush-greasewood
K050 Fescue-wheatgrass
K051 Wheatgrass-bluegrass
K052 Alpine meadows and barren
K055 Sagebrush steppe
K056 Wheatgrass-needlegrass shrubsteppe
K063 Foothills prairie

SAF COVER TYPES [48]:
12 Black spruce
16 Aspen
63 Cottonwood
107 White spruce
201 White spruce
204 Black spruce
205 Mountain hemlock
206 Engelmann spruce-subalpine fir
207 Red fir
208 Whitebark pine
209 Bristlecone pine
210 Interior Douglas-fir
211 White fir
212 Western larch
213 Grand fir
215 Western white pine
216 Blue spruce
217 Aspen
218 Lodgepole pine
220 Rocky Mountain juniper
221 Red alder
222 Black cottonwood-willow
223 Sitka spruce
224 Western hemlock
225 Western hemlock-Sitka spruce
226 Coastal true fir-hemlock
227 Western redcedar-western hemlock
228 Western redcedar
229 Pacific Douglas-fir
230 Douglas-fir-western hemlock
231 Port-Orford-cedar
232 Redwood
234 Douglas-fir-tanoak-Pacific madrone
235 Cottonwood-willow
237 Interior ponderosa pine
238 Western juniper
239 Pinyon-juniper
243 Sierra Nevada mixed conifer
244 Pacific ponderosa pine-Douglas-fir
245 Pacific ponderosa pine
246 California black oak
247 Jeffrey pine
248 Knobcone pine
253 Black spruce-white spruce

SRM (RANGELAND) COVER TYPES [108]:
101 Bluebunch wheatgrass
102 Idaho fescue
107 Western juniper/big sagebrush/bluebunch wheatgrass
108 Alpine Idaho fescue
109 Ponderosa pine shrubland
110 Ponderosa pine-grassland
202 Coast live oak woodland
203 Riparian woodland
207 Scrub oak mixed chaparral
208 Ceanothus mixed chaparral
209 Montane shrubland
210 Bitterbrush
213 Alpine grassland
216 Montane meadows
302 Bluebunch wheatgrass-Sandberg bluegrass
304 Idaho fescue-bluebunch wheatgrass
314 Big sagebrush-bluebunch wheatgrass
315 Big sagebrush-Idaho fescue
401 Basin big sagebrush
402 Mountain big sagebrush
403 Wyoming big sagebrush
406 Low sagebrush
408 Other sagebrush types
409 Tall forb
410 Alpine rangeland
411 Aspen woodland
412 Juniper-pinyon woodland
415 Curlleaf mountain-mahogany
422 Riparian
501 Saltbush-greasewood
503 Arizona chaparral
504 Juniper-pinyon pine woodland
612 Sagebrush-grass
614 Crested wheatgrass

PLANT COMMUNITIES:
In addition to the plant communities listed above, occurrence of the bushy-tailed woodrat was noted in the following habitats:

Bushy-tailed woodrats were found inhabiting rock outcrops surrounded by big sagebrush (Artemisia tridentata)/cheatgrass (Bromus tectorum), big sagebrush/Sandberg bluegrass (Poa secunda), and big sagebrush/needle-and-thread (Stipa comata) community types at the Hanford Site in Washington [58].

In the Slim Buttes area of northwestern South Dakota, bushy-tailed woodrats occur in green ash (Fraxinus pennsylvanica)/chokecherry (Prunus virginianus) habitat [109].

At Bass Lake, California, 1 bushy-tailed woodrat was found in a forest dominated by incense-cedar (Calocedrus decurrens), white fir (Abies concolor), California black oak (Quercus kelloggii), ponderosa pine (Pinus ponderosa), sugar pine (Pinus lambertiana), and white alder (Alnus rhombifolia). In Lake Tahoe, California, bushy-tailed woodrats were found inhabiting a building surrounded by Jeffrey pine (Pinus jeffreyi), sugar pine, lodgepole pine (Pinus contorta), incense-cedar, white fir, western juniper (Juniperus occidentalis), California black oak, and chaparral. Chaparral consisted of greenleaf manzanita(Arctostaphylos patula), huckleberry oak (Quercus vaccinifolia), whitethorn ceanothus (Ceanothus cordulatus), bush chinquapin (Castanopsis semperivirens), bitter cherry (Prunus emarginata), big sagebrush, and other species [112].


BIOLOGICAL DATA AND HABITAT REQUIREMENTS

SPECIES: Neotoma cinerea

 

  Photo courtesy of Jessica Allewalt

TIMING OF MAJOR LIFE HISTORY EVENTS:
Mating: Female bushy-tailed woodrats are seasonally polyoestrous [16,47,51]. The breeding period occurs from January to August [16,34,85] and peaks between March and June [16]. In southwestern Oregon, 55% (n not given) of female bushy-tailed woodrats were found lactating until fall, indicating a longer breeding period [29].

Social Organization: Bushy-tailed woodrats occur in small family groups, which are often widely separated, and distribution is often patchy [29,47]. Males are territorial and aggressively defend dens and foraging areas [16,29,47]. Bushy-tailed woodrats are considered polygynous by some authors [16,29,47]; however, this is not based on observed matings, but on the size and relative overlap of male and female ranges [34].

Gestation period and litter size: Gestation of bushy-tailed woodrats is 27 to 32 days [16,47]. On the Oregon Coast, young are born in March [85]. Litter size ranges from 1 to 6 [47,85,114], and averages 3.5 [16]. In the Sierra Nevada, mean litter size was 2.5 (n=11) [47]. Number of litters is typically 1 per year, especially in northern parts of the bushy-tailed woodrat's range [16]. In southern parts of the bushy-tailed woodrat's range, 2 litters may be produced [85].

Development: Young bushy-tailed woodrats are altricial [16,85], and the male parent does not help raise offspring [47]. Young are weaned between 26 and 30 days of age [16], and dispersal from the nest occurs at 2 months of age [114]. Bushy-tailed woodrats reach sexual maturity at 11 months [16], and breeding begins during spring and summer, ~1 year following birth [47].

Dispersal: Female bushy-tailed woodrats are generally philopatric and breed on the same rock outcrop over successive years. Male yearlings are more mobile than female yearlings and may disperse greater distances in search of suitable habitat and den sites [47]. Dispersal distances of 1.4 to 2.0 miles (2.2-3.2 km) have been recorded [29]. Once a territory is established, both sexes are relatively sedentary. The separation of family groups may attract predators and lead to extinction of the colony via emigration, predation, and random demographic processes such as death and low birth rates. Therefore, dispersal of offspring is important in the recolonization of vacated habitats [29,47].

Habits: Bushy-tailed woodrats are primarily nocturnal and are most active during the half hour after sunset and at dawn, year-round [16,29,47,51,85,114,118].

Survival: Litters produced early in the season have a greater chance of survival than litters produced later in the season [47]. Bushy-tailed woodrats may live 3 to 4 years [47,114].

Annual survival rates for bushy-tailed woodrats in dry forest cover types in the Cascade Range [82]

  Year

Cover type

1997-1998 1998-1999 1999-2000
Open ponderosa pine 0.14 0.13 0.05
Young mixed conifer
(grand fir (Abies grandis), Douglas-fir (Pseudotsuga menziesii), and ponderosa pine)
0.25 0.17 0.05
Mature mixed conifer (same species as above) 0.08 0.21 0.20

PREFERRED HABITAT:
Bushy-tailed woodrats are the most boreal species within the Neotoma genus [47,51]. They inhabit areas from sea level to 14,110 feet (4,300 m) and are most often associated with montane habitats [47,61,114]. Bushy-tailed woodrats are arboreal rodents [13,21,26,29,30,47], associating primarily with ponderosa pine, Douglas-fir, spruce (Abies spp.), and quaking aspen (Populus tremuloides) forests [47,51,85]. Other common habitat types inhabited by the bushy-tailed woodrat include pinyon (Pinus spp.)-juniper (Juniperusspp.) woodlands [5,22,44,50,56,67,78,93,110,125] and big sagebrush habitat [5,45,49,58,72,89,125].

Occurrence of bushy-tailed woodrats is closely tied to the geology of a landscape [47]. Shelter and topography are important determinants of habitat suitability [47,51]. The availability of rock shelters may be a more important resource for the bushy-tailed woodrat than the associated plant community [34,47,51]. Preferred habitat varies depending on the subspecies of bushy-tailed woodrat; however, most subspecies are associated with rocky environments such as talus slopes [16,51,116], rockslides [47,114], boulder fields [34], rock outcrops [16,34,51,85,116], cliffs [51,116], and caves [16,51,61,106,114,116].

Due to the occasional scarcity of rocky habitats in the Pacific Northwest, hollow trees, logs, dwarf mistletoe (Arceuthobium spp.) brooms, and coarse woody debris are used for denning, foraging, and shelter [21,26,34,35,66,77,82,85,94,96,124]. In the Columbia River Basin, bushy-tailed woodrats use grand fir and white fir snags most often for den sites [77,94]. Tree cavities created by the pileated woodpecker (Dryocopus pileatus) are used by bushy-tailed woodrats in grand fir, western larch, and ponderosa pine trees [21,24,25,26].

Bushy-tailed woodrats may also associate with streamside habitats [29,34,47,82,87]. In southwestern Oregon, streamside forests provide the most suitable habitat for bushy-tailed woodrat [29]. In wet Douglas-fir and western hemlock (Tsuga heterophylla) forests in western Washington, bushy-tailed woodrats are found primarily in rocky, streamside areas [82]. Riparian quaking aspen woodlands in rangeland communities are commonly used by bushy-tailed woodrats in northeastern Nevada [87].

Human-made buildings [40,51,85,112,114] and mine tunnels are sometimes used by bushy-tailed woodrat [51].

Stand- and landscape-level habitat: Bushy-tailed woodrats may be found in early through climax stages of succession [4,29,34,82,125].

In the Pacific Northwest, one of the preferred landscapes for the bushy-tailed woodrat is closed pole-sapling stands, which are somewhat ephemeral as they succeed to small saw-log stands. Extensive areas of Douglas-fir old-growth (>200 years) may be most suitable for maintaining a viable population of bushy-tailed woodrats. Contiguous areas of suitable habitat or corridors connecting suitable habitat are also important [29].

In western Oregon and western Washington, bushy-tailed woodrats preferred mature (>80 years), unmanaged upland Douglas-fir-western hemlock streamside forests more than young (35 to 80 years), "managed" streamside forest. Streams in this habitat were narrow, deeply cut, and associated with forested boulder fields. The adjacency of streams was not sufficient to ensure occupancy by bushy-tailed woodrats.

Percent frequency of occurrence of bushy-tailed woodrats in Douglas-fir/western hemlock streamside forests [34]
 

Stream present

Stream not present

  Stands (n) % occupied Stands (n) % occupied
Mature, unmanaged forest 27 78 34 21
Young, managed forest 12 33 46 17

In the Pacific Northwest, Douglas-fir forests can be "sources" or "sinks" for the bushy-tailed woodrat, depending on the stand condition, presence of streams, and occurrence of predators such as the northern spotted owl (Strix occidentalis caurina) [29]. "Sources" are defined as an environment capable of sustaining a viable population. "Sinks" are defined as an environment capable of maintaining a population for a short time, but supporting low reproduction or periods of unsuitability for occupancy.

Habitat ratings for stand conditions in Douglas-fir forests in the Pacific Northwest [29]
Closed sapling-pole
(20 years old; average DBH=1 to 21 inches (3-53 cm); canopy cover >60%)
Source and sink
Small sawtimber
(60 years old, DBH and canopy cover not given)
Sink
Large sawtimber
(100 years old; average DBH=21 inches (53 cm); canopy cover <100%; decay and decadence lacking)
Sink
Old-growth
(Stand 700 years old, at least 2 tree layers, decay in living trees, snags, fallen trees)
Source and sink

Information on landscape suitability for the bushy-tailed woodrat is sparse outside the Pacific Northwest. In Douglas-fir and mixed-conifer forests in southwestern Oregon and northern California, bushy-tailed woodrats were found in young and mature forests containing diverse understory vegetation [86]. Bushy-tailed woodrats were transients or ephemeral residents within the quaking aspen seral stage of a forest dominated by Engelmann spruce (Picea engelmannii) in the Wasatch Mountains in Utah [4]. In Montana, bushy-tailed woodrats are found in old-growth western larch (Larix occidentalis)/Douglas-fir forests [65]. In alluvial valleys of the Green and Yampa rivers in northwestern Colorado and northeastern Utah, bushy-tailed woodrat presence was examined in 3 habitats: high floodplain, low floodplain, and upland. Bushy-tailed woodrats were found in high floodplain and upland habitats, but the numbers in upland habitats were small. In high floodplain habitat, flooding occurred at longer intervals than in low floodplain areas, which flooded every other spring. High floodplain habitat contained the most mature cottonwood (Populus fremontii) and woody debris of the 3 habitats studied. In upland habitat, where flooding never occurred, dominant vegetation included sagebrush (Artemisia spp.), greasewood (Sarcobatus spp.), and saltbush (Atriplex spp.) [49].

Home range and density: Bushy-tailed woodrats have a larger home range than any other Neotoma species. This may due to the requirement of existing geologic structures for den sites, such as fissures in rocky outcrops. Denning and nesting sites are usually limiting resources in bushy-tailed woodrat habitat. This may lead to concentrations of bushy-tailed woodrats in certain areas [47]. Importance of a suitable den site may outweigh the costs of maintaining a large home range [47].

Bushy-tailed woodrats occur in small, widely separated family groups [29]. The minimum habitat size needed per family group has been estimated at 80 acres (32 ha) [29]. It is rare to find more than one family group on a talus slope covering 2 to 5 acres (0.8-2.0 ha). Banfield [16] reported an average population density of 1 bushy-tailed woodrat per 20 acres (8 ha) in Canada. Bushy-tailed woodrats may forage up to 450 feet (137 m) from the den; this constitutes a 15 acre (6 ha) home range [29]. In southwestern Oregon, Carey [29] recorded movements of up to 1,250 feet (381 m) within trapping grids that were only 1,300 feet (396 m) wide, which suggests a maximum home range of about 110 acres (45 ha). The average maximum distance moved per individual was 280 feet (85.3 m).

The size of the home range is positively correlated with the number of females in the group [47]. Male bushy-tailed woodrats have larger home ranges than females [29]. In the Kananaskis Valley in southwestern Alberta, average home range size for male bushy-tailed woodrats was 6.12 ha (n=23, range 1.6 to 11.2 ha), and for females was 3.56 ha (n=27, range 0.1 to 10.4 ha) [116].

In mixed-conifer (grand fir and Douglas-fir) habitat in southwestern Oregon, bushy-tailed woodrat density was greatest (1.08 individuals/ha 0.51 (SE)) in streamside closed-canopy habitat. In Douglas-fir forests, bushy-tailed woodrat density was greatest (0.59 individuals/ha 0.57 (SE)) in rocky sites. For more details about bushy-tailed woodrat density in different habitat types in Oregon, see Carey and others [32].

COVER REQUIREMENTS:
Den sites are primary limiting factors of bushy-tailed woodrat habitat [29,47,111]. Depending on the subspecies of bushy-tailed woodrat, dens may be located in openings between rocks, coarse woody debris (snags and logs), or dwarf mistletoe brooms [29]. In areas with heavy snowfall, rock shelters are the most important resource for the bushy-tailed woodrat [47]. In areas with mild climates and low availability of rock shelters, coarse woody debris, mistletoe brooms, and rock crevices along streams are utilized more often [21,26,35,66,77,82,85,94,96,124].

Bushy-tailed woodrat shelter consists of 2 parts, the den and the nest [29,47]. Dens refer to a shelter constructed from items such as sticks [47], leaves, grass, bark, feathers, paper, small stones [114] and human artifacts [16]. Dens are important for protection against predators and are almost never constructed in the open without a bush, rock crevice, or tree for support [47]. They are used as eating platforms, a platform for the nest, and a drying area for food [16]. Within the den are tunnels, chambers, and passages. One of the chambers may contain 2 or more nests, and other chambers are used for food storage and feeding. A lifetime may be spent using the same den [114].

Nests refer to a small cup-like structure usually located inside the den. Occasionally a nest may be found without a den [47]. On the Oregon coast and Coast Ranges, bushy-tailed woodrats may construct nests in trees up to 50 feet (15 m) above ground [29,85]. Nests are 6 to 8 inches (15-20 cm) in diameter [85] and are constructed of fibrous shredded vegetation [16,51,85].

FOOD HABITS:
Bushy-tailed woodrats are herbivores [4,67,114] with a broad, flexible diet [47,55,85]. Regional specialization on certain locally abundant food items may occur [47,51]. Due to their flexible diet, food availability is likely not a limiting factor in habitats occupied by bushy-tailed woodrats. Generally, food is a more abundant resource than shelter [47]. In general, bushy-tailed woodrats do not travel far from their den to forage [55,116]. In Gunnison County, Colorado, bushy-tailed woodrats satisfied their foraging needs within a 98 foot (30 m) radius of their dens, but would occasionally travel upto 500 feet (152 m) from their den for unknown reasons [55]. Food is dried on rocks before storage [16,47,51], and is stored extensively in middens located at the periphery of the den in late August and September [16,47,51]. As plant diversity increases in bushy-tailed woodrat habitat, diversity of midden contents increases [55]. Middens may be several feet deep [51] and have been radiocarbon dated to be thousands of years old [122].

Foods eaten include cones and needles of coniferous trees [85,114], berries, leaves [47], shrubs [16,72], and forbs [67]. Fungi are a major food item in lodgepole pine and grand fir habitats in northeast Oregon. Bushy-tailed woodrats are vectors of spore dissemination of hypogeous mycorrhizal fungi in conifer forests [86].

In Canada, bushy-tailed woodrats showed a preference for the leaves of quaking aspen, cherry (Prunus spp.), rose (Rosa spp.), snowberry (Symphoricarpos spp.), currants (Ribes spp.), elderberries (Sambucus spp.), and willows (Salix spp.). The twigs and needles of western white pine (Pinus monticola), Douglas-fir, Engelmann spruce (Picea engelmannii), subalpine fir (Abies lasiocarpa), and juniper (Juniperus spp.), and the seeds and/or fruits of Douglas-fir, anemone (Anemone spp.), fireweed (Epilobium angustifolium), gentian (Gentianella spp.), honeysuckle (Lonicera spp.), cinquefoil (Potentilla spp.), gooseberry (Ribes spp.), raspberry (Rubus spp.), and elderberry were also eaten [16].

Consumption of woody vegetation by bushy-tailed woodrats may decrease competition for food with other forb-eating small mammals. In a pinyon-juniper habitat in the Piceance Basin in Colorado, 75% of the bushy-tailed woodrat diet was composed of woody plants, primarily antelope bitterbrush (Purshia tridentata). Forbs composed 15% of their diet, but the percentage of forbs in the diet may increase in years of heavy rainfall. For a list of foods eaten by bushy-tailed woodrat in pinyon-juniper habitat in Colorado, see Haufler and Nagy [67]. In basin big sagebrush (Artemisia tridentata ssp. tridentata) habitat in south-central Idaho, plains prickly-pear (Opuntia polyacantha) was the main food eaten by bushy-tailed woodrats [72].

PREDATORS:
Bushy-tailed woodrats add substantially to the prey base of birds and mammals [29,34]. Their presence and abundance may affect the fitness of spotted owls (Strix occidentalis) throughout their range [29,32,34,92,117,118,123], but their importance varies locally [32]. Bushy-tailed woodrats comprised >10% relative frequency and total biomass of the Mexican spotted owl's (Strix occidentalis lucida) diet in the northern portion of its range [123]. In the Olympic Peninsula, Coast Range, Umpqua River Valley, and Klamath Mountains of Oregon, bushy-tailed woodrats comprised 2% to 70% of prey biomass for the northern spotted owl [32].

Other predators of bushy-tailed woodrats include the great-horned owl (Bubo virginianus) [16,29,78], boreal owl (Aegolius funereus) [68], American marten (Martes americana) [16,23,29], fisher (Martes pennanti) [130], and bobcat (Lynx rufus) [29].

Incidental predators include the coyote (Canis latrans) [29,45], western spotted skunk (Spilogale putorius), long-tailed weasel (Mustela frenata) [29], red-tailed hawk (Buteo jamaicensis), northern goshawk (Accipiter gentilis) [47], rattlesnake (Viperidae), wolverine (Gulo gulo) and fox (Vulpes spp. and Urocyon spp.) [16].

Due to the large amount of time spent in dens, bushy-tailed woodrats are particularly susceptible to ectoparasites such as ticks, chiggers, lice, mites, fleas, cone-nosed bugs, and warble flies [51].

MANAGEMENT CONSIDERATIONS:
Due to their dependence on trees, arboreal rodents such as the bushy-tailed woodrat are sensitive to timber harvesting [71] and may experience population declines in intensively managed areas [13]. Huff and others [71] rated the vulnerability of arboreal mammal species to local extirpations resulting from the fragmentation or loss of old-growth Douglas-fir forests. The index ranged from "1-low vulnerability", to "10-high vulnerability". The bushy-tailed woodrat was rated a "4" [71].

Coarse woody debris is an important habitat element for the bushy-tailed woodrat, especially in the Pacific Northwest [21,26,34,35,39,82,85,94,96,124] (see Preferred Habitat). Specific information about the density and types of logs required to provide viable habitat for bushy-tailed woodrat populations is unavailable [24]; however, loss of logs due to timber management or fire should be mitigated due to their importance for cover [82]. According to Pearson [96], coarse woody debris management should focus on maintaining a diverse array of sizes and decay classes for various animal species. Suggestions for coarse woody debris management include the creation of debris piles from thinning slash, and management of streamside habitat to restore coarse woody debris and complex vegetation structure [34]. For detailed recommendations about how to manage trees and logs in the Interior Columbia River Basin, see Bull and others [26]. For information about snag management for wildlife use, see Wellersdick and Zalunardo [124] and Pearson [96].

Managing for habitat patchiness may be suitable for the bushy-tailed woodrat [26,34]. Carey and others [31,33] suggest variable-density thinning in wet Douglas-fir and western hemlock forests in western Washington, with the retention of large snags and woody debris on the scale of 0.5- to 1.2-acre (0.2 to 0.5 ha) patches.

Livestock grazing: Bushy-tailed woodrats may either be negatively or positively affected by grazing by domestic livestock in mixed-conifer and spruce (Picea spp.)-fir (Abies spp.) forests in Arizona and New Mexico, depending on the grazing regime. Seed production of shrubs and forbs could be increased by grazing; however, heavy livestock use may degrade woody cover. Bushy-tailed woodrats are a "priority species" in Arizona and New Mexico. This means that bushy-tailed woodrats should receive greater consideration than non-priority wildlife species during development of management strategies related to livestock grazing [131].

Other: When bushy-tailed woodrat density was compared between unchained and chained areas in a mature pinyon-juniper (Pinus edulis-Juniperus osteosperma) woodland in the Piceance Basin, Colorado, density was greatest (n=8 individuals) in unchained control areas. Each study area consisted of 3 trapping grids, which included 32 trap stations (4 x 8 feet (1 x 2m)) at intervals of 49 feet (15 m). In 1-year-old chained areas, bushy-tailed woodrat density was 5 per trapping grid. Bushy-tailed woodrats were absent in 8-year-old or 15-year old chained areas [93]. O'Meara and others [93] suggest interspersing blocks of chained and unchained vegetation to maintain pinyon-juniper woodland species such as the bushy-tailed woodrat.


FIRE EFFECTS AND USE

SPECIES: Neotoma cinerea
DIRECT FIRE EFFECTS ON ANIMALS:
Bushy-tailed woodrats can probably escape most fires by taking refuge in rocky habitats. A study in California showed that during a prescribed fire, woodrats (Neotoma spp.) evaded harm by escaping to rock outcrops [70]. Woodrats that do not escape to these refugia may be killed by fire [128]. Bushy-tailed woodrats residing in riparian areas are probably protected by their habitat, which may burn infrequently and in a patchy pattern.

HABITAT-RELATED FIRE EFFECTS:
Fire may influence bushy-tailed woodrat populations by altering habitat structure and/or plant community composition [83]. The rapidity of recolonization of a disturbed area by small mammals depends on the size and severity of the disturbance and the presence of refugia [74].

As of this writing, no information is available on the habitat-related fire effects on the bushy-tailed woodrat. Despite the lack of information, some generalizations based on their habitat requirements may be possible. Coarse woody debris and mistletoe brooms are important resources for the bushy-tailed woodrat, primarily in the Pacific Northwest [21,26,34,35,39,82,85,94,96,124] (see Preferred Habitat). Conflicts in coarse woody debris management have arisen between retaining logs for wildlife use and lowering fuel levels to decrease the risk of wildfire [24]. Coarse woody debris and mistletoe brooms close to the ground torch easily and would probably be consumed by low-severity prescribed fire [69]. In ponderosa pine and dry Douglas-fir habitats, low-severity, high-frequency fire regimes will likely reduce bushy-tailed woodrat populations unless fire prescriptions mitigate the loss of coarse woody debris and mistletoe brooms [82]. In 12 vegetation types in coastal and interior British Columbia, wildlife community structure was highly correlated with fire regimes. As fire size and intensity increased, the proportion of species using woody cavities decreased due to the loss of snags and woody debris. As the interval between fires increased and downed woody debris accumulated, proportions of species using downed wood increased [27].

Bushy-tailed woodrats are found in early successional through climax stages of succession [4,29,34,125], and their presence depends more on cover and food availability than on the seral stage [47,51]. In the spruce-fir zone of northern Utah, for example, bushy-tailed woodrats were found in meadows and quaking aspen stands. These communities are commonly the first successional stages after fire [103]. In the Sierra Nevada, bushy-tailed woodrats often inhabit early postfire successional stages of Jeffrey pine and lodgepole pine communities [20,76]. Some berry-producing shrubs that provide food for bushy-tailed woodrats, such as blackberries, raspberries, and gooseberries, often thrive after fire [127]. However, severe, stand-destroying fires that consume the organic layer can kill the roots of many berry-producing shrubs, reducing the potential for sprouting and delaying revegetation [97,119]. According to Lehmkuhl and others [82], unmanaged mature forests are source habitat for bushy-tailed woodrats compared to thinned and/or burned forest stands. Closed-sapling pole habitats are also source habitat for bushy-tailed woodrats (see Stand- and landscape-level habitat) [29].

In eastern Washington and Oregon, wildlife appears to be well-adapted to mixed-severity fire regimes in mixed-conifer forests [81]. A critical habitat feature affecting wildlife habitat in mixed-severity fire regimes is the mosaic of vegetation conditions that are created [2]. Mixed-severity fires in dry forest types create mosaics of varying burn intensity and a complex of open and closed-canopy forest structure containing woody debris, snags, and understory vegetation [3,81]. Woody debris may be abundant in mixed-severity vegetation types, but availability varies over time. Consumption of woody debris on the forest floor is offset by the creation of snags in patches burned at moderate and high severities [3]. Burning to create patchy fire severity may therefore benefit the bushy-tailed woodrat [82].

Snags with large cavities are important for many species of birds and mammals including the bushy-tailed woodrat [21]. A bushy-tailed woodrat nest was found in an old-growth log with a fire scar opening 3 feet (0.9 m) high and 2 feet (0.6 m) wide [29]. Old pileated woodpecker cavities used by bushy-tailed woodrats may be at risk of destruction by high-severity fire [82]; however, high-severity fire may also create snags [26].

To reduce slash from logging operations while maintaining coarse woody debris, broadcast burning may be useful. Because of the time of year that broadcast burning is conducted, large diameter materials are usually not consumed [98,99,101]. In addition to broadcast burning, slash <6 inches (15 cm) diameter can be piled and burned to reduce fine fuels while maintaining coarse woody debris [96]. Prescribed fire has less impact on small mammals in western larch habitat than mechanical scarification [104], which destroys small mammal habitat [84].

The following table provides fire-return intervals for plant communities and ecosystems where bushy-tailed woodrats may occur. For further information, see the FEIS review of the dominant plant species listed below.

Community or Ecosystem Dominant Species Fire Return Interval Range (years)
silver fir-Douglas-fir Abies amabilis-Pseudotsuga menziesii var. menziesii >200
grand fir Abies grandis 35-200 [8]
California chaparral Adenostoma and/or Arctostaphylos spp. <35 to <100
sagebrush steppe Artemisia tridentata/Pseudoroegneria spicata 20-70 [95]
basin big sagebrush Artemisia tridentata var. tridentata 12-43 [102]
mountain big sagebrush Artemisia tridentata var. vaseyana 15-40 [10,28,90]
Wyoming big sagebrush Artemisia tridentata var. wyomingensis 10-70 (x=40) [120,129]
saltbush-greasewood Atriplex confertifolia-Sarcobatus vermiculatus <35 to <100 [95]
curlleaf mountain-mahogany* Cercocarpus ledifolius 13-1,000 [12,105]
California montane chaparral Ceanothus and/or Arctostaphylos spp. 50-100
California steppe Festuca-Danthonia spp. <35 [95]
green ash Fraxinus pennsylvanica <35 to >300 [46,121]
western juniper Juniperus occidentalis 20-70
Rocky Mountain juniper Juniperus scopulorum <35 [95]
western larch Larix occidentalis 25-350 [9,18,38]
Engelmann spruce-subalpine fir Picea engelmannii-Abies lasiocarpa 35 to >200 [8]
black spruce Picea mariana 35-200 [43]
blue spruce* Picea pungens 35-200 [8]
pinyon-juniper Pinus-Juniperus spp. <35 [95]
Rocky Mountain bristlecone pine P. aristata 9-55 [41,42]
whitebark pine* Pinus albicaulis 50-200 [1,6]
Rocky Mountain lodgepole pine* Pinus contorta var. latifolia 25-340 [17,18,115]
Sierra lodgepole pine* Pinus contorta var. murrayana 35-200 [8]
Colorado pinyon Pinus edulis 10-400+ [53,59,75,95]
Jeffrey pine Pinus jeffreyi 5-30
western white pine* Pinus monticola 50-200
Pacific ponderosa pine* Pinus ponderosa var. ponderosa 1-47 [8]
interior ponderosa pine* Pinus ponderosa var. scopulorum 2-30 [8,14,80]
Arizona pine Pinus ponderosa var. arizonica 2-15 [14,37,107]
aspen-birch Populus tremuloides-Betula papyrifera 35-200 [43,121]
quaking aspen (west of the Great Plains) Populus tremuloides 7-120 [8,62,88]
mountain grasslands Pseudoroegneria spicata 3-40 (x=10) [7,8]
Rocky Mountain Douglas-fir* Pseudotsuga menziesii var. glauca 25-100 [8,10,11]
coastal Douglas-fir* Pseudotsuga menziesii var. menziesii 40-240 [8,91,100]
Pacific coast mixed evergreen Pseudotsuga menziesii var. menziesii-Lithocarpus densiflorus-Arbutus menziesii <35-130 [8,36]
California oakwoods Quercus spp. <35 [8]
California black oak Quercus kelloggii 5-30 [95]
redwood Sequoia sempervirens 5-200 [8,52,113]
western redcedar-western hemlock Thuja plicata-Tsuga heterophylla >200
western hemlock-Sitka spruce Tsuga heterophylla-Picea sitchensis >200
mountain hemlock* Tsuga mertensiana 35 to >200 [8]
*fire return interval varies widely; trends in variation are noted in the species review

FIRE USE:
As of 2006, there was no research on the effects of wildfire or prescribed burning on the bushy-tailed woodrat. Data are needed to make detailed management recommendations. Based on the habitat requirements for the bushy-tailed woodrat, fire management generalizations may be possible. Bull [24] recommends that if maintaining a viable population density for a particular animal is a management goal, then the habitat requirements for a species and the ecological processes resulting from fires must be assessed. Mixed-severity fire regimes may be beneficial for the bushy-tailed woodrat due to the mosaic of landscapes that are created [3,81]. Habitat corridors may be important to connect suitable habitat [29].

High-severity wildfire and prescribed burns may have negative effects on the bushy-tailed woodrat due to the loss of coarse woody debris, mistletoe brooms, and food sources. Plants eaten by the bushy-tailed woodrat may not regenerate for several years after severe fire [97,119]. Low-severity fire may not have as great an impact on bushy-tailed woodrats as high-severity fire. Any type of fire prescription including an objective of maintaining bushy-tailed woodrat habitat should mitigate the loss of coarse woody debris, dwarf mistletoe brooms, and snags [82].


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