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Peromyscus boylii

• INTRODUCTORY
• WILDLIFE DISTRIBUTION AND OCCURRENCE
• BIOLOGICAL DATA AND HABITAT REQUIREMENTS
• FIRE EFFECTS AND USE
• REFERENCES

INTRODUCTORY

AUTHORSHIP AND CITATION:
Luensmann, Peggy S. 2005. Peromyscus boylii. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: www.fs.usda.gov/database/feis/animals/mammal/pebo/all.html [].

FEIS ABBREVIATION:
PEBO

SYNONYMS:
Peromyscus boylei Baird [54]

COMMON NAMES:
brush deermouse
brush mouse

TAXONOMY:
The scientific name of brush deermouse is Peromyscus boylii Baird [7,115]. It is a member of the mouse, rat, and vole family, Muridae [9]. Subspecies in the United States include:

P. b. subsp. rowleyi (Allen) [46]
P. b. subsp. utahensis Durrant [46]

In the Timing Of Major Life History Events section of this review, some reproductive information specific to the brush deermouse was not available. In these cases, information on the entire genus is presented.  Additionally, in the Fire Effects And Use section, observations of mice from the genus Peromyscus are presented, although the mice were not identified to species in those publications.

ORDER:
Rodentia

CLASS:
Mammal

FEDERAL LEGAL STATUS:
None [109]

OTHER STATUS:
Information on state- and province-level protection status of animals in the United States and Canada is available at NatureServe, although recent changes in status may not be included.

WILDLIFE DISTRIBUTION AND OCCURRENCE

• GENERAL DISTRIBUTION
• ECOSYSTEMS
• STATES/PROVINCES
• BLM PHYSIOGRAPHIC REGIONS
• KUCHLER PLANT ASSOCIATIONS
• SAF COVER TYPES
• SRM (RANGELAND) COVER TYPES
• PLANT COMMUNITIES

AZ: Overstories where brush deermice are found are characterized by desert willow (Chilopsis linearis), Arizona sycamore (Platanus wrightii), and shrub live oak (Quercus turbinella) [16,32]. Shrub associates include evergreen sumac (Rhus virens), skunkbush sumac (R. trilobata), roundleaf snowberry (Symphoricarpos rotundifolius), New Mexico locust (Robinia neomexicana), common hoptree (Ptelea trifoliata), fendlerbush (Fendlera rupicola), Carruth's sagewort (Artemisia carruthii), catclaw acacia (Acacia greggii), broom snakeweed (Gutierrezia sarothrae), red barberry (Mahonia haematocarpa), netleaf hackberry (Celtis reticulata), and wait-a-minute (Mimosa aculeaticarpa var. biuncifera) [21,32,68]. Herbaceous and succulent species in brush deermouse habitats include Wheeler sotol (Dasylirion wheeleri), sacahuista (Nolina microcarpa), Palmer agave (Agave palmeri), goldenrod (Solidago spp.), lupine (Lupinus spp.), prickly-pear (Opuntia spp.), and ocotillo (Fouquieria splendens) [21,32,68]. Grasses found in brush deermouse habitat include Lehmann lovegrass (Eragrostis lehmanniana), bottlebrush squirreltail (Elymus elymoides), and other annual and perennial grasses [16,32,68].

NM: Habitats are characterized by rabbitbrush (Chrysothamnus spp.), Apache plume (Fallugia paradoxa), fourwing saltbrush, common hoptree and crispleaf buckwheat (Eriogonum corymbosum) in addition to a pinyon-juniper-mountain-mahogany (Pinus-Juniperus-Cercocarpus spp.) overstory [98].

UT: Rocky areas are dominated by cliffrose (Purshia spp.) and buffaloberry (Shepherdia spp.) on the slopes, and barberry (Mahonia spp.) and oak (Quercus spp.) at cliff bottoms [2].

TX: Plant associates include Pinchot juniper (J. pinchotii), algerita (M. trifoliolata), clapweed (Ephedra antisyphilitica), featherplume (Dalea formosa), and prickly-pear [38].

OK: Rocky, brush-covered cliffs and slopes are characterized by netleaf hackberry, western soapberry (Sapindus saponaria var. drummondii), common hoptree, skunkbush sumac, American plum (Prunus americana), and oak [40]. Grassy areas include grama (Bouteloua spp.) and threeawn (Aristida spp.).

BIOLOGICAL DATA AND HABITAT REQUIREMENTS

• TIMING OF MAJOR LIFE HISTORY EVENTS
• PREFERRED HABITAT
• COVER REQUIREMENTS
• FOOD HABITS
• PREDATORS
• MANAGEMENT CONSIDERATIONS

Brown [19] reported that brush deermice reach sexual maturity at 12-19 weeks. However, Clark [23] reported that female brush deermice may reach sexual maturity in approximately 5-9 weeks. In Arizona, females born in late May or June were found pregnant during August of the same year [48]. In western Texas, Arizona, southern Utah, and Mexico, breeding appears to occur nearly year-round [25,38,48,66,92]. However, in northern California, Jameson [55] found that breeding in brush deermice peaked twice each year, once in late spring and again in late summer. This result may be related to food availability [55,56].

Female brush deermice carry 1-6 embryos each [11,17,48,55,66,100] with an average of 3-4 embryos [11,17,48,55]. Lactating females may become pregnant [48], although lactation may slow the development of embryos in Peromyscus species [102]. Bradley and Schmidly [17] reported that the gestation period of the brush deermouse is around 23 days. Females may have multiple litters per year [17,25]. The amount of time between litters is likely similar to other Peromyscus species and may be 25-31 days [100]. Young are weaned at 3-4 weeks of age [17].

Brush deermice have a promiscuous mating system [57,85]. Four out of 7 litters in a California study were fathered by several males [57]. Males and females did not share nests and mating pairs did not remain together for long periods of time [57].

Few Peromyscus spp. mice live longer than 6 months in the wild, but individuals may live up to 4-5 years in captivity [80].

PREFERRED HABITAT:
Vegetation in brush deermouse habitats may vary from location to location, but brush deermice are consistently captured in areas with medium to high densities of shrubs and tree cover under 16 feet (5 m) in height [2,25,31,36,48,49,73,75,82,103,111,114]. In California, mature chaparral (cover ≥50%) appears to provide more suitable habitat for brush deermice than young, open chaparral (cover <50%) [82]. Similarly, in Arizona, Duran [31] captured brush deermice most frequently in shrub live oak and birchleaf mountain-mahogany (Cercocarpus betuloides) understory habitats with 45% to 50% plant cover. Fewer brush deermice were captured in habitats with less plant cover [31]. Holbrook [50] observed that after vegetation crowns were removed in a manzanita (Arctostaphylos spp.)-oak shrubland, brush deermice avoided the newly-opened space. In another study, brush deermice were strongly restricted to habitats in which gaps between rocks or ceanothus (Ceanothus spp.) on the site were less than 4 feet (1.2 m) [81].

In addition to shrub density, the height of cover appears to influence brush deermouse distribution within a site. An average understory height of 5-6.5 feet (1.5-2.0 m) was preferred by brush deermice over lower understory cover [97]. In another study, brush deermouse presence was positively correlated with microhabitats of shrub cover up to 10 feet (3 m) tall, logs over 3 inches (7.5 cm) in diameter, and understory trees 10-33 feet (3-10 m) in height, but negatively correlated with grass-forb microhabitats [16].

Brush deermice are also commonly captured at locations with a high proportion of rock cover and/or slash piles in habitats characterized by chaparral-mountain shrub, oak/shrub, oak-juniper-pinyon pine, juniper-pinyon pine, and oak-pine communities, as well as riparian habitats [1,2,25,27,30,36,38,40,41,48,49,50,51,66,73,103,110,111,114]. The brush deermouse in Texas has been found in all major habitats present (desert, grassland, riparian, and montane) although it is typically associated with rock outcrops within these habitats [25]. Riparian sites with abundant brush deermouse populations had high shrub cover, high frequency of debris piles with low grass, litter, and tree cover [32]. In a Mexico study, a canyon was dominated by exposed rock, grasses, pines, hardwoods, and brush [30]. In western Texas, brush deermice favored fallen logs and brush piles [26]. Modi [71] discovered that brush deermice were common in riparian zones dominated by pecan (Carya illinoensis), American sycamore (Platanus occidentalis) and live oak (Quercus virginiana), in an oak community with a partially open canopy and dense understory, and in a pine forest with little understory and scattered boulders. In New Mexico, brush deermouse populations were significantly higher (P<0.05) on sites that were bulldozed or thinned (98 and 115 captures, respectively) than untreated or bulldozed and burned sites [95]. Populations were lowest on sites that had not been treated (45 captures). Sites that had increased slash from bulldozing and burning had more brush deermice (57 captures) than the untreated sites, but the difference was not significant (P>0.05). No influence of canopy cover on brush deermice was observed in the study [95].

Besides high tree, shrub, and rock densities, brush deermice appear to prefer locations with low grass cover [32,81]. At the same time, grasses are often present in the understory indicating that grasses do not exclude brush deermice [25,40]. Brush deermice utilized grazed and ungrazed pastures and ceanothus plots [81], but they were concentrated around rocky outcrops and vegetation continuous with the rock outcrops. No brush deermice were captured in the grasslands more than 20 feet (6 m) from rocks, shrubs, or trees [81]. Litter depth also appears negatively correlated to brush deermouse presence [97,111]. For instance, brush deermice in Arizona were captured in litter depths of only 0.9 inches (2.4 cm) [111].

Brush deermice also utilize fire-affected habitats. In one study, brush deermice were captured in burned and unburned chaparral as well as burned and unburned pine-oak forest. The highest number of captures were recorded in unburned forest while the lowest captures occurred in the unburned chaparral [73]. These results are somewhat inconsistent with other observations which show the brush deermouse favoring dense chaparral habitat. Small mammal capture data in the study were collected from 14 months to 3 years after fire [73]. The time frame of sampling after fire may influence the perceived response of the brush deermouse to postfire habitats.

Elevation, in addition to habitat characteristics, may play a role in habitat suitability in some areas. For instance, in the northern Sierra Nevada of California, brush deermice were captured in brush habitats at 3,500-5,000 feet (1,000-1,500 m), but not at 6,500 feet (2,000 m) [53]. Aspect may influence the distribution of brush deermice on a site as well. For example, in New Mexico, 51% of all brush deermice captured were taken on south-facing slopes, 24% on west-facing slopes, with 13% and 12% of mice captured on east- and north-facing slopes, respectively [114]. The south-facing canyon slopes may provide more cover for brush deermice due to higher numbers of shrubs [114].

Site characteristics of brush deermouse habitat:

Although brush deermice are found on a variety of slopes, including flat mesas and gradual slopes, they seem to prefer locations with very steep slopes, such as hillsides, mountain sides, and canyons (including some slopes with >45% gradient) over more gradual slopes in the same areas [1,30,40,41,73,97,101,111]. Findley [35] reported that brush deermice were captured on hillsides in an oak/sacahuista (Nolina spp.) community. In another study, brush deermice were common in canyon bottoms, on hillsides, and in arroyos (water channels in arid regions) characterized by oak woodlands [36]. Brush deermice have also been captured along the sides of brush covered canyons and burned slopes of an oak/brush association [114].

Density and Home Range
Kalcounis-Ruppell [57] discovered brush deermouse population densities were 40-72 mice per hectare in coastal California. Similarly, Kalcounis-Ruppell and Millar [58] determined that densities of brush deermice in coastal California were 42-89 mice per hectare. Brush deermouse densities appear greatly influenced by weather. Densities were 17-20 per acre one year, but were reduced to 6 mice per acre following a severe winter [41]. At another study site, brush deermice increased from 4 individuals per acre to 12 per acre after a mild winter [41]. Populations in a canyon in Mexico were estimated at a mean of 10.8 brush deermice per acre or 6.0 males and 4.8 females per acre [30]. The range was 9.0-14.0 brush deermice per acre [30].

A study utilizing radiotelemetry and trapping data in Arizona estimated mean home range size for male brush deermice at 0.72-1.6 acres (0.29-0.64 hectare), and 0.32-0.79 acre (0.13-0.32 hectare) for females [86]. In a another study at the same location, home ranges for male brush deermice ranged from means of 1.2-1.5 acres (0.47-0.62 hectare) and for females, means of 0.64-1.2 acres (0.26-0.49 hectare) [85]. Mean home range for male and female brush deermice in Arizona, based on radiotelemetry, was 0.30 acre (0.12 hectare) [42].

COVER REQUIREMENTS:
No significant differences in habitat use between genders were observed in Arizona [43]. Typical brush deermouse habitat in southern Arizona was characterized by 74% tree cover, 60% leaf cover, 21% shrub cover, and 16% rock cover. Additionally, 67% of all brush deermice relocated by radiotelemetry in the study were located in a riparian zone with the other observations occurring in uplands and an intermittent stream channel. Brush deermice used sites with significantly (P<0.05) more rock cover (19% to 22%) during winter and spring than at other times of the year. Also in spring, plots with the most brush deermice also had significantly (P<0.05) more shrub cover (21% vs.14%) and succulents (9% vs. 2%) than random plots [43].

In a New Mexico study, 27% of brush deermice captured were taken around rocky areas, 10% under Gambel oak (Q. gambelii), 9% under juniper, and 7% each around woodpiles and sacahuista [114]. Less frequently, brush deermice were taken under or around pinyon pine, gray oak (Q. grisea), apache plume, rabbitbrush, mountain-mahogany, white fir (Abies concolor), Douglas-fir (Pseudotsuga menziesii), locust (Robinia spp.), prickly-pear, cholla (Opuntia spp.), chokecherry (Prunus spp.), bricklebush (Brickellia spp.), and grape (Vitis spp.) [114].

In southern California, brush deermice were captured on leaf mold in an oak hardwood association of coast live oak, white alder (Alnus rhombifolia), whiteflower currant, and sumac (Rhus spp.) [110]. They were also trapped under logs and dense vegetation and on wet seepage slopes next to a creek [110]. In coastal California, brush deermice were captured primarily under coast live oak, false-willow (Baccharis douglasii), California buckeye, and California bay [58].

Fallen logs and rock outcrops provide nest sites for the brush deermouse [17]. They may also construct nests in tree hollows and underground burrows [113]. Brush deermice living in mine shafts or caves build nests similar to those of goldfinches [48]. Davis [26] noted that the brush deermouse nest is a "globular structure" and is constructed primarily of dried grasses within natural cavities. Brush deermouse nest sites in California were found on sites with a high density coast live oak (Q. agrifolia) overstory and an open understory with low vegetation and ground cover [57].

FOOD HABITS:
Brush deermice are semiarboreal and can be found foraging in shrubs and trees for leaves and fruits [35,49,51,56]. Females were captured more often than males foraging in canyon live oaks (Q. chrysolepis) (P<0.05) [69]. The individuals with the longest tails appear to spend more time climbing than those with shorter tails [35].

Acorns are commonly eaten by brush deermice wherever they are available [10,17,26,54,57]. Arthropods and cutworms (Protorthodes rufula) are also eaten throughout the year [31,41,54,56,98]. A variety of fruits and seeds from Douglas-fir, ponderosa pine (Pinus ponderosa), pinyon, California buckeye, manzanita (A. patula and A. viscida), silktassel (Garrya spp.), oneseed juniper (Juniperus monosperma), hackberries (Celtis spp.), New Mexico groundsel (Senecio neomexicanus var. neomexicanus), trailing fleabane (Erigeron flagellaris), annual sunflower (Helianthus annuus), broom snakeweed, common dandelion (Taraxacum officinale), western yarrow (Achillea millefolium), white sweetclover (Melilotus albus), threenerve goldenrod (Solidago velutina), prickly-pear, desert wheatgrass (Agropyron desertorum), Kentucky bluegrass (Poa pratensis), and blue grama (Bouteloua gracilis) are eaten throughout the year when available [17,26,31,41,54,56,57,98]. Other plant parts, such as leaves, stems, flowers, pollen cones and new sprouts are typically eaten in lower quantities than other foods [31,41,54,56,98]. Fungi are typically consumed when other foods are scarce [54,56]. Infrequently, stomach contents of brush deermice contained pieces of mammals, birds, and fence lizards (Sceloporus spp.) [54].

Brush deermice have been observed caching pinyon pine seeds [78,79]. This observation suggests that the brush deermouse may play a role in seed dispersal for some plant species.

PREDATORS:
Predators of the brush deermouse include birds of prey such as the northern goshawk (Accipiter gentilis) and spotted owl (Strix occidentalis) [60,63,84,100,108]. Potential mammalian predators of the brush deermouse may include the coyote (Canis latrans), common gray fox (Urocyon cinereoargenteus), red fox (Vulpes vulpes), swift fox (Vulpes velox), ringtail (Bassariscus astutus), American marten (Martes americana), fisher (Martes pennanti), ermine (Mustela erminea), long-tailed weasel (Mustela frenata), striped skunk (Mephitis mephitis), hooded skunk (Mephitis macroura), white-backed hog-nosed skunk (Conepatus leuconotus), northern raccoon (Procyon lotor), bobcat (Lynx rufus), and ocelot (Leopardus pardalis) [38,63,100,112]. Snakes also prey on brush deermice [100,119].

MANAGEMENT CONSIDERATIONS:
The activities of humans since the arrival of Europeans have created more brushlands than previously existed, thus creating favorable habitat for the brush deermouse [10]. Brush deermouse populations appear positively correlated to the amount of downed woody debris present on a site [95]. In New Mexico, bulldozed and thinned sites had 2.5-3 times more slash (10% to 13% slash cover) than untreated sites (3% slash cover). These same sites had at least twice as many brush deermice captured as the untreated site. A site that was bulldozed, piled, and burned (6% slash cover) also had a higher brush deermouse population than the untreated site, but lower than the bulldozed or thinned only sites [95]. Similarly, a habitat assessment in California suggests that the removal of downed woody debris and the reduction of tree and shrub canopy cover to 40% during fuels treatments would have a negative impact on the brush deermouse [15].

Brush deermice are primary carriers of Sin Nombre virus, which causes hantavirus pulmonary syndrome in humans [42]. Because their presence in and around buildings has been documented [42], reducing habitat suitability by the removal of vegetation and wood piles around man-made structures may reduce the transmission of the disease to humans.

FIRE EFFECTS AND USE

• DIRECT FIRE EFFECTS ON ANIMALS
• HABITAT-RELATED FIRE EFFECTS
• FIRE USE

Observations of small mammals before, during, and after fire suggest that most small mammals are able to escape fire by burrowing into the soil [99]. However, rodents may be susceptible to the heat effects of fire when soil temperatures are high. For instance, soil temperatures reaching or exceeding 145 °F (63 °C) appear to almost always be lethal to small mammals [52]. Rodents buried up to 6 inches (15 cm) died from lethal temperatures resulting from fire. Temperatures were especially high under fallen logs and near burning stumps. Rodents also died in rock crevices and under ceanothus when the temperature reached at least 140 °F (60 °C). Similarly, rats buried 1.5-5 inches (4-13 cm) below the soil surface in an open sunny site outside the burned area died when temperatures from solar radiation alone reached at least 145 °F (60 °C). However, a rat that was buried 7 inches (18 cm) under the surface survived where temperatures only reached 138 °F (59 °C) [52]. No data were found on brush deermouse survival in burrows during fire.

In addition to lethal temperatures, small mammals may perish from exposure to flames. Deer mice (P. maniculatus), pinyon mice (P. truei), Townsend's chipmunks (Tamias townsendii), Trowbridge's shrews (Sorex trowbridgii), and dusky-footed woodrats (Neotoma fuscipes) were seen fleeing from burning slash piles in a prescribed fire in California [106]. Woodrats in the fire waited until their cover was fully engulfed in flames before escaping. Some woodrats caught on fire from burning pitch that dropped onto them while they were hiding. Such individuals appeared to run in a panicked manner, starting small fires along the way. Woodrats not on fire tended to run from one patch of cover to the next in a less erratic fashion. Several woodrats perished either from catching on fire or by refusing to leave their homes [106]. Although observations of mouse (Peromyscus spp.) behavior during the fire was limited, it is possible that they may behave in a similar manner to the woodrats.

Immediately after the fire, approximately one third of the original number of mice captured on the site were recaptured [106]. Nearly all were collected from the edge of the burn. Individuals captured within the burn were collected from islands which did not catch fire. Thus, all the mice that were not able to find cover outside the fire appear to have either perished or fled the area entirely [106]. Fire effects on the brush deermouse may be comparable because of the biological similarities between brush deermice and other Peromyscus species.

HABITAT-RELATED FIRE EFFECTS:
During the 2.5 weeks after a prescribed fire in California, 13 mice (P. maniculatus or P. truei) were captured [106], or approximately one third the original number of captures before the fire. The deep ash covering the burned area prevented the movement of the mice. Several days after the fire, heavy rain hardened the surface of the ash. Within 5 days, 51 mice were captured within the burned area [106]. Thus, fire-altered habitat appears to restrict the movement of mice until postfire conditions change.

Brush deermice in New Mexico preferred sites with significantly lower tree cover (P<0.01), lower conifer density and basal area (P<0.001) and less litter (P<0.001) than random sites [111]. This suggests that fire may benefit brush deermice by keeping tree cover and litter depth low. However, they also preferred sites with significantly higher shrub and oak sapling and seedling densities (P<0.001) [111]. This result suggests that fires removing low shrub and tree cover would be detrimental to the brush deermouse. In an Arizona study, small mammals were sampled in burned and unburned chaparral and pine-oak forest from 14 months to 3 years after fire [73]. The wildfire burned more than 90% of the vegetation that was present before the fire. More than twice as many brush deermice were captured in the unburned forest habitat than any other habitat sampled. The lowest number of captures came from the unburned chaparral. The burned habitats may have recovered sufficiently 3 years after fire, allowing brush deermice to reinvade the area [73].

Fire may alter the habitat structure and/or the community composition, which may influence brush deermouse populations [67]. Brush deermice are able to recolonize areas that were burned previously. For instance, in a chaparral habitat of southern California, brush deermice recolonized sites 3 years and 4 months after wildfire or prescribed fire [116]. Presumably, the fires severely altered the chaparral habitat so that it was unsuitable to brush deermice for more than 3 years. Furthermore, brush deermice occupied fire-caused secondary forest growth in dry forest habitats of southern Mexico [88]. Additionally, brush deermice were captured in a former mixed conifer woodland dominated by white fir, Douglas-fir, incense-cedar (Calocedrus decurrens), sugar pine (Pinus lambertiana), and ponderosa pine that had succeeded to a California black oak (Q. kelloggii)-manzanita-ceanothus-Fremont silktassel (Garrya fremontii) association 19-38 years after fire [53].

Fire regimes: The following table provides fire return intervals for plant communities and ecosystems where the brush deermouse is important. Find fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find Fire Regimes".

Frequent fires would likely be necessary to keep brush deermouse populations low. This goal would be valuable when the spread of hantavirus to humans was a concern. On the other hand, if increasing brush deermouse populations as a food source for predators such as the spotted owl were necessary, then less frequent fires may be desirable.

Peromyscus boylii: REFERENCES

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