Small Mammal Populations and Ecology 

in the Kings River Sustainable Forest 

Ecosystems Project Area1 

William F. Laudenslayer, Jr.

2

 and Roberta J. Fargo3 

Abstract 

Small mammals are important components of woodlands and forests. Since 1992, we have 

been studying several aspects of small mammal ecology in oak woodlands in western 

foothills of the southern Sierra Nevada. Assemblages of small, nocturnal mammal species are 

dominated by the brush mouse (Peromyscus boylii), California mouse (P. californicus), and 

dusky-footed woodrat (Neotoma fuscipes). Brush mice and California mice were found in 

highest numbers on sites with the densest vegetative cover. Populations of both species 

generally appeared to decline over the 4 years of study. Dusky-footed woodrats in our study 

sites were arboreal and often used specific travel paths to move about their home ranges. 

These behaviors made them difficult to capture with standard trapping grids, and we 

found that capture success was increased markedly by placing traps near their houses, 

either on the ground or in trees. Woodrats on our sites constructed several kinds of 

houses—some of which were difficult to detect—and house use was difficult to detect. 

We did not find a close correspondence between the numbers of woodrats and the 

numbers of their houses. 

Small mammals, principally rodents, are important residents of a variety of 

habitats in the Kings River Sustainable Forest Ecosystems Project Area. They 

affect the composition and structure of their environment through consumption 

and distribution of foliage, plant seeds, and fungal spores. They also serve as 

food for larger predators, including species of concern, such as the northern 

goshawk (Accipiter gentilis), California spotted owl (Strix occidentalis occidentalis), 

marten (Martes americana), and fisher (Martes pennanti). Although existing 

literature includes some reports of small mammal assemblages in California’s 

oak woodlands, information on species composition is lacking, even in relation 

to gross vegetation structure and particularly in relation to predator activity. 

The initial objective of this study was to learn more about small mammalspotted owl relations where the owls breed in riparian hardwood forests of the 

oak woodland zone of the western Sierra Nevada foothills. Critical methodical 

questions surfaced during the course of the work, however, some of which 

required answers before we could move forward effectively with studies of the 

mammal-owl relations. 

Repeat live trapping on grids is commonly used to detect the presence and 

estimate the abundance of small mammals. If traps or trapping locations are 

biased, however, comparisons between sampling locations and species are 

probably unreliable. Different types of traps may bias trap success toward some 

species and against others (Laudenslayer and Fargo 1997, O’Farrell and others 

1994, Willy 1985). The placement of traps within or outside of particular 

1 An abbreviated version of this 

paper was presented at the 

Symposium on the Kings River 

Sustainable Forest Ecosystems 

Project: Progress and Current 

Status, January 26, 1998, Clovis, 

California. 

2 	Research Wildlife Biologist, 

Forestry Sciences Laboratory, 

Pacific Southwest Research 

Station, USDA Forest Service, 

2081 E. Sierra Ave., Fresno, CA 

93710. 

3 	 11240 Mendenhall Loop Rd., 

Juneau, AK 99801. 

USDA Forest Service Gen. Tech. Rep. PSW-GTR-183. 2002. 133 
Laudenslayer, Fargo Small Mammal Populations and Ecology in the Kings River Sustainable Forest Ecosystems Project Area 

microhabitats, and their proximity to travel paths, may also bias trap success for 

some species. Reliable data on arboreal and fossorial species can be especially 

difficult to obtain because of uncertainties associated with identifying 

appropriate microhabitats and travel paths for setting traps. 

Typically, dusky-footed woodrats (Neotoma fuscipes) (hereafter, woodrats) 

build stick houses in which to nest. These houses are conspicuous and often are 

used as an index of woodrat presence and abundance (Hamm 1995, Sakai and 

Noon 1993, Vogl 1967). Some studies have suggested, however, that woodrat 

numbers are poorly correlated with numbers of houses (Fargo and Laudenslayer 

1999, Hamm 1995, Humphrey 1988). 

Woodrats are often arboreal, but little is known about their activity patterns 

on the ground and in the trees. Studies of the ecology of nocturnal small 

mammals, such as the woodrat, have used various methods, such as 

mark-recapture, radio telemetry, and laboratory simulations. Although these 

approaches provide important ecological information, data on behavior and 

home-range use are often incomplete. For example, mark-recapture and 

telemetry data usually provide two-dimensional spatial information on animals. 

This is often incomplete, however, because many small mammals, including 

woodrats, are at least partially arboreal and thus use a third dimension. 

Since 1993, we have been studying small mammal assemblages in three 

2.25-ha oak woodland sites in eastern Fresno County, California. Efforts to date 

have examined rodent responses to five general questions: Are standard trapping 

grids for small mammals effective for estimating presence and abundance of 

nocturnal small mammals in these oak woodland habitats? Do the abundances of 

small mammal species vary in relation to habitat structure and through time? Do 

woodrats construct their visually obvious stick houses under all habitat 

structures examined? Can surveys of stick houses suffice to estimate relative 

abundance of woodrats? How do woodrats use their three-dimensional 

environment? 

Study Sites 

Our study sites are adjacent to Pine Flat Reservoir on the Kings River, Fresno 

County, California. All sites are within 2 km of the Kings River, and elevations 

range from 300-450 m. We chose three 2.25-ha study sites—Pine Flat, Camp 

4-1/2, and Secata. Although all sites were classified as oak woodland, they 

differed markedly in the structure and composition of their vegetation (table 1). 

Table 1—Percent of plant species 1-4 m and >4 m high at Pine Flat, Secata, and Camp 4-1/2 study sites (frequency is 

based on the species of tree or shrub providing the highest percentage of canopy cover over each survey point) 

Pine Flat Secata Camp 4-1/2 

Species 1–4 m >4 m 1–4 m >4 m 1–4 m >4 m 

_______________________________________________________________________________ 

Blue oak 4.0 16.7 50.0 68.4 4.3 21.7 

California ash 4.0 4.2 — — 13.0 4.3 

California buckeye — — 5.6 5.3 17.4 13.0 

California redbud — — — — 4.3 — 

Foothill pine — 20.8 — — — — 

Interior live oak 40.0 54.2 11.1 21.1 39.1 60.9 

Manzanita 16.0 4.2 22.2 5.3 — — 

Poison oak 8.0 — — — — — 

Redberry 8.0 — — — 4.3 — 

Wedgeleaf ceanothus 20.0 — 11.1 — 17.4 — 

134 USDA Forest Service Gen. Tech. Rep. PSW-GTR-183. 2002. 
Small Mammal Populations and Ecology in the Kings River Sustainable Forest Ecosystems Project Area Laundenslayer, Fargo 

The Pine Flat site, with 80 percent canopy cover, is characterized by a dense, 

shrubby cover dominated by interior live oak (Quercus wislizenii), wedgeleaf 

ceanothus (Ceanothus cuneatus), and poison oak (Toxicodendron diversiloba), with 

some manzanita (Arctostaphylos sp.), blue oak (Q. douglasii), and foothill pine 

(Pinus sabiniana). The Camp 4-1/2 study site, with 65 percent canopy cover, is 

dominated by interior live oak, blue oak, and California buckeye (Aesculus 

californica) and has a moderately open understory of poison oak and ceanothus. 

The Secata study site is relatively open—45 percent canopy cover—with patches 

of large trees dominated by blue oak and lesser amounts of interior live oak, 

California buckeye, and manzanita. 

The largest oaks in bole and canopy diameter are at Secata, followed by 

Camp 4-1/2, and then Pine Flat. The tallest blue oaks are at Secata and the tallest 

interior live oaks are at Camp 4-1/2. Shrubs also vary in size across the sites with 

the largest poison oaks at Camp 4-1/2 and largest manzanitas at Secata (tables 2, 

3, and 4). 

Table 2—Mean bole diameters (cm) of trees and shrubs at Pine Flat, Secata, and Camp 4-1/2 study sites 

Species Pine Flat Secata Camp 4-1/2 

Blue oak 13.4 34.0 18.8 

California ash 2.7 1.2 10.2 

Foothill pine 28.0 — — 

Interior live oak 8.3 20.6 15.0 

Manzanita 5.5 11.7 — 

Poison oak 1.1 1.0 3.5 

Wedgeleaf ceanothus 3.6 5.5 5.4 

Table 3—Mean crown diameters (m) of trees and shrubs at Pine Flat, Secata, and Camp 4-1/2 study sites 

Species Pine Flat Secata Camp 4-1/2 

Blue oak 2.9 5.0 3.5 

California ash 1.2 1.0 3.2 

Foothill pine 4.5 — — 

Interior live oak 3.4 4.3 3.3 

Manzanita 2.4 2.9 — 

Poison oak 1.1 1.0 1.6 

Wedgeleaf ceanothus 2.0 1.7 1.7 

Table 4—Mean heights (m) of trees and shrubs at Pine Flat, Secata, and Camp 4-1/2 study sites 

Species Pine Flat Secata Camp 4-1/2 

Blue oak 5.1 9.9 7.8 

California ash 1.8 1.0 4.3 

Foothill pine 10.0 — — 

Interior live oak 4.8 5.6 6.3 

Manzanita 2.7 3.0 — 

Poison oak 1.4 1.0 2.6 

Wedgeleaf ceanothus 2.0 2.0 2.0 

USDA Forest Service Gen. Tech. Rep. PSW-GTR-183. 2002. 135 
Laudenslayer, Fargo 

4	 Mention of trade names or 

products is for information 

only and does not imply 

endorsement by the U.S. 

Department of Agriculture. 

Small Mammal Populations and Ecology in the Kings River Sustainable Forest Ecosystems Project Area 

Methods 

Effectiveness of Standard Trapping Grids 

Initially, we randomly placed a 7 x 7 trap-station grid with 15-m spacing in each 

study site. We positioned one Sherman XLK (7.7 x 9.5 x 30.5 cm) and one 

Tomahawk 201 (12.7 x 12.7 x 40.6 cm)4 at each station in 1993. Because attempts to 

capture woodrats on the grid were essentially unsuccessful, we also set 

Tomahawk traps at woodrat houses, both on the ground and in trees. Traps set at 

houses on the ground were positioned along woodrat travel paths, if evident. 

Because this approach was successful, in 1994 and 1995 we set only one Sherman 

XLK at each grid station, in addition to Tomahawks at the houses. In 1995, we 

also used Shermans at houses on the ground and in trees for a short period to 

confirm that the woodrats were not avoiding this type of trap. In 1996, we 

concentrated the trapping adjacent to woodrat houses, based on questions that 

had arisen during the previous years of trapping. Nocturnal trapping was done 

in the spring (April-June) and in the late summer/early fall (SeptemberNovember), and traps were checked in the early morning to reduce incidental 

mortality from long exposures to the weather. 

Woodrat Stick Houses 

Each study site was thoroughly surveyed for woodrat houses. In addition to 

houses found during initial establishment of study sites, newly built houses 

found over the course of the study were added to the inventory. Occupancy of 

individual houses was determined by observations at the houses and by the 

presence or absence of woodrat “sign.” Direct observations of woodrats often 

indicated which houses were being used. For other houses, the presence of fresh 

vegetative cuttings, recently deposited fecal matter, and travel paths and house 

entrances clear of debris and spider webs demonstrated their current use. All 

houses found were classified as typical, intermediate, or atypical, based on stick 

content and nest location. 

We did vegetation surveys in the spring of 1998 to compare house types with 

vegetation structure and composition on the sites. House information collected 

included house type, size, substrate, and cover. Site information included tree 

and shrub characteristics such as species, bole diameter, height, and percent 

cover; and ground cover attributes such as rock outcrops, logs, vegetation, and 

woody material. 

Woodrat Behavior and Movements 

To study the behavior and track the movements of woodrats, we set 10 

Tomahawk traps at randomly selected, active woodrat houses each night of 

tracking. The first two adult or subadult woodrats captured were generally 

selected as the subject animals for the night. Each was equipped with a 

light-emitting diode (LED) package and dusted with fluorescent powder before 

release. Different colors of LEDs and fluorescent powder permitted 

discrimination of activity patterns between the two woodrats being observed 

(Fargo and Laudenslayer 1997). 

Using audio cassettes, two observers (one per woodrat) verbally recorded 

the movements of the tagged woodrats throughout each night of observation, 

noting the time and duration of each detection of an animal, microhabitat at the 

animal’s location (for example, tree canopy), characterization of the animal’s 

location as a “travel route” or “destination,” position of the animal (height above 

the ground and the azimuth and distance from its house), and the animal’s 

activity (for example, walking, tail and foot tapping). 

The evening following observations, woodrats were recaptured to remove 

the LED units, and their fluorescent powder trails were tracked. Information 

136 USDA Forest Service Gen. Tech. Rep. PSW-GTR-183. 2002. 
Small Mammal Populations and Ecology in the Kings River Sustainable Forest Ecosystems Project Area Laundenslayer, Fargo 

collected included length of the powder trail within each microhabitat, 

characterization of sections of the powder trail as “travel route” or “destination,” 

position of the powder trail (height above the ground and the azimuth and 

distance from the woodrat’s house), and canopy cover over each powder trail. 

Vegetation surveys will be done to compare each woodrat’s use of the 

microhabitat in its range with what is available. For each woodrat observed, 

vegetation information within a specified area around each house will be 

collected as described above for woodrat houses. 

Results 

Small Mammal Fauna 

Nine species of small mammals were captured at the study sites, with most 

being active mainly at night (table 5). The brush mouse, California mouse, and 

dusky-footed woodrat were the most abundant mammal species captured. Six 

other species were captured but in low numbers. Despite their occurrence in 

similar habitats elsewhere in California, we have yet to capture any piñon mice 

(Peromyscus truei). 

Effectiveness of Standard Trapping Grids 

Both Sherman and Tomahawk traps successfully captured woodrats, but few 

were captured on trapping grids. Instead, most were captured at woodrat 

houses, either on the ground or in nearby trees (table 6). Both species of mice 

were captured more successfully in Sherman than in Tomahawk traps. This was 

especially true for brush mice, regardless of trap placement (table 6), because 

nearly all of them could escape through the Tomahawk trap’s mesh and the 

space around the door. Although results are inconclusive because of changes in 

trapping methods, capture rates for both mice may be better at traps associated 

with trees and woodrat houses (table 6). 

Abundance of Small Mammals 

Woodrat numbers varied markedly over the 4 years of the study, but the annual 

variability was not consistent among the study sites (table 7). Woodrats were 

most abundant at Pine Flat, which generally had the most immatures and the 

highest female to male ratio. This site also had the greatest canopy cover and 

Table 5—Small mammal species captured in oak woodland study sites between 1993 and 1996 

Common name Scientific name Activity period 

Brush mouse Peromyscus boylii Nocturnal 

California ground squirrel Spermophilus beecheyi Diurnal 

California mouse Peromyscus californicus Diurnal 

California pocket mouse Chaetodipus californicus Nocturnal 

California vole Microtus californicus Nocturnal 

Deer mouse Peromyscus maniculatus Nocturnal 

Dusky-footed woodrat Neotoma fuscipes Nocturnal 

Merriam’s chipmunk Tamias merriami Diurnal 

Shrew Sorex sp. Nocturnal 

Western harvest mouse Reithrodontomys megalotis Nocturnal 

USDA Forest Service Gen. Tech. Rep. PSW-GTR-183. 2002. 137 
Laudenslayer, Fargo Small Mammal Populations and Ecology in the Kings River Sustainable Forest Ecosystems Project Area 

Table 6—Small mammal capture success [pct success = 100 x (number captured/number of trap nights)] compared by trap type and trapping 

location (numbers of trap nights: on grid—Shermans = 4,525, Tomahawks = 2,183; at houses on ground—Shermans = 52, Tomahawks = 2,106; at 

houses in trees—Shermans = 10, Tomahawks = 1,877) 

On grid1 At houses on ground2 At houses in trees3 

Animal 

Sherman4 Tomahawk5 Sherman Tomahawk Sherman Tomahawk 

Brush mouse 17.46 1.05 26.92 0.09 70.00 0.11 

California mouse 2.34 0.78 9.62 0.57 10.00 0.59 

Dusky-footed woodrat 0.49 1.65 23.08 8.74 10.00 12.63 

1 Traps set in grid on the ground and in trees.


2 Traps set at woodrat houses on the ground.


3 Traps set at woodrat houses in adjacent trees.


4 Sherman XLK trap.


5 Tomahawk 201 trap.


Table 7—Numbers of individuals captured on trapping grids from 1993 to1996 

Pine Flat Secata Camp 4-1/2 

Animal 1993 1994 1995 1996 1993 1994 1995 1996 1993 1994 1995 1996 

Brush mouse 55 59 54 15 47 8 15 6 104 37 50 24 

California mouse 21 14 13 5 2 0 0 0 19 6 4 

Dusky-footed woodrat 24 34 19 25 10 13 12 18 9 25 16 20 

densest understory vegetation. Captures at Secata, which had the least canopy 

cover and sparsest understory vegetation, were approximately half the number 

at Pine Flat in 1993 and 1994, but increases at Secata and decreases at Pine Flat 

narrowed the difference in 1995 and 1996 (table 7). 

Both brush mice and California mice appeared to be generally more 

abundant at Camp 4-1/2 and Pine Flat than at Secata, and our capture rate 

tended to show a general decline from 1993 to 1996, although a decline in brush 

mice at Pine Flat was evident only in 1996 (table 7). 

Woodrat Stick Houses 

Typical and intermediate woodrat houses were found in high numbers at Pine 

Flat, where they were primarily found on rock outcrops or at the bases of shrubs 

or trees (fig. 1). Atypical houses were prevalent at Secata, where most were 

associated with tree cavities or rock outcrops. Camp 4-1/2 had nearly equal 

numbers of the three house types, with many in tree cavities. 

Individual woodrats used one to four houses at any given time, and this 

number often changed over time. 

Woodrat Behavior and Movements 

Results to date indicate that woodrats spent slightly more time in trees (45.27 

percent) than on the ground (40.01 percent) (table 8). While on the ground, 

138 USDA Forest Service Gen. Tech. Rep. PSW-GTR-183. 2002. 

2 
Small Mammal Populations and Ecology in the Kings River Sustainable Forest Ecosystems Project Area Laundenslayer, Fargo 

Figure 1—Numbers of typical, 

atypical, and intermediate woodrat 

houses at Pine Flat, Camp 4-1/2, and 

Secata study sites. 

Table 8—Amount of time woodrats spent in various microhabitats in the three study sites combined 

Microhabitat Time (seconds) Percent of time 

Downed log


Downed woody material


Ground (unknown substrate)


Grass/forb


Rock outcrop


Shrub canopy


Shrub limb


Tree Bole


Tree canopy


Tree limb


11 0.08 

336 2.44 

3,222 23.43 

184 1.34 

2,360 17.16 

314 2.28 

999 7.27 

1,142 8.31 

1,227 8.92 

3,955 28.76 

however, they spent much of their time on rock outcrops. Whether remaining 

stationary or moving about, they usually stayed under dense canopy cover. They 

tended to use established travel paths along limbs, on rocks, or on the ground. 

Travel paths on the ground were characteristically clear of debris and vegetation. 

Discussion 

Effectiveness of Standard Trapping Grids 

Our results indicate that capture success for different species of small mammals 

varies by trap type and trapping location. Effects of trap type on capture success 

were based, at least in part, on the ability of small mammals to escape from the 

traps and probably to some extent on selection of particular trap types by certain 

species and/or individuals. Similarly, Willy (1985) found that Tomahawk traps 

captured only woodrats, and at a higher rate than Shermans, whereas Shermans 

captured woodrats as well as other species. O’Farrell and others (1994) also 

found variation in capture success based on trap type, with wire mesh traps 

generally capturing more species and individuals of small mammals than 

Shermans in a variety of western habitats and geographic locations. 

Variation in our trapping success in relation to trap location appeared to be 

associated with microhabitat affinities and travel corridors. Similarly, Murray 

(1957) reported varying trap success between parallel trap lines depending on 

USDA Forest Service Gen. Tech. Rep. PSW-GTR-183. 2002. 139 
Laudenslayer, Fargo Small Mammal Populations and Ecology in the Kings River Sustainable Forest Ecosystems Project Area 

the structure and composition of vegetation, such that small differences in trap 

placement could seriously alter results. Differences in capture success could bias 

estimates of small mammal populations, thereby affecting decisions based on the 

assessments. It is important, therefore, to carefully evaluate methods to ensure 

that they are unbiased, or at least biased equally among the habitats and species 

being compared. 

Kirkland and Griffin (1974), Price (1978), Sakai and Noon (1993), and Tietje 

(1995) all found that patchiness of the animals’ habitat and their use of it 

influenced capture success. We captured woodrats more successfully in traps 

placed at their houses than on the trap grid, indicating that these animals, too, 

are not randomly distributed in their habitat. 

Several species of Peromyscus, including the piñon mouse and the California 

mouse, often construct their nests within woodrat houses (Cranford 1982; Merritt 

1974, 1978). Our findings, based on only a few trap nights, show that both brush 

and California mice are more successfully captured near woodrat houses. We 

did not, however, confirm cohabitation by searching woodrat houses for 

Peromyscus nests. 

Abundance of Small Mammals 

Although preliminary and without replication, our data suggest that these three 

study sites support different numbers of the three commonly captured rodent 

species. Secata, the most open site, supported the lowest numbers of all three 

species. Both Pine Flat and Camp 4-1/2, with denser vegetation, support higher 

numbers of all three species. 

Only a few California mice were captured at Secata, and in only 1 year, 

suggesting that California mice are not typically resident there, that they are 

present only in low numbers, or that our trapping methods are ineffective for 

capturing California mice at this site. 

Additional work is needed over a longer period to more clearly understand 

the relations between populations of these small mammals and the vegetation. 

Woodrat Stick Houses 

Woodrat houses have been described as conspicuous, conical piles of sticks with 

many passageways and compartments (Ashley and Bohnsack 1974, Cameron 

1971, English 1923, Gander 1929, Linsdale and Tevis 1951). Some observers have 

described “atypical” houses built in hollow limbs and rock crevices (Davis 1934, 

Gander 1929), but these are usually mentioned only briefly and apparently are 

considered to be uncommon. 

The presence of numerous atypical and cryptic woodrat houses in our study 

sites, however, suggests potentially large biases in estimates of woodrat 

populations if based on house counts. The Key Largo woodrat (Neotoma floridana 

smalli), for example, is generally known for its large stick houses, but it is also 

known to reside within rock crevices in some habitats, thereby escaping detection 

during house-count surveys (Barbour and Humphrey 1982). In spite of this 

problem, some observers have based population estimates on counts of houses 

(Hammer and Maser 1973, Sakai and Noon 1993, Vogl 1967), either assuming 

that each active house contained one woodrat (Vestal 1938, Vogl 1967) or 

applying some standard number of houses per woodrat across the entire study 

area. Not all locations may include cryptic houses, however, so investigators 

need to become familiar with all house types characteristic of their study site and 

to search accordingly for all houses before using house counts to estimate 

population size. 

Woodrats in our study sites also differed in the number of houses they used, 

and that number changed over time. English (1923) reported that dusky-footed 

140 USDA Forest Service Gen. Tech. Rep. PSW-GTR-183. 2002. 
Small Mammal Populations and Ecology in the Kings River Sustainable Forest Ecosystems Project Area 

woodrats near Corvallis, Oregon, were familiar with several houses in their 

home range, and Cranford (1977), in a study of radio-tagged woodrats, found 

that they resided within a house for an average of 34 days. When a home range 

contained only one house, it tended to be larger than average, and the woodrat 

resided in it for a longer period. Such variation in house use may also bias 

population estimates when based on one woodrat per house. In some seasons, 

more than one woodrat may reside in a house, especially during spring and 

summer when males reside with females for breeding (Donat 1933, English 1923) 

and when females are nursing young (Linsdale and Tevis 1956). Without further 

study, we consider it risky to use counts of woodrat houses for much more than 

determining woodrat presence (but not absence!). 

Woodrat Behavior and Movements 

Woodrats in our study sites appear to restrict their travel routes to limbs, rocks, 

and cleared paths that likely expedite their movements and reduce the noise 

produced. These travel paths are also generally beneath dense canopies. This 

behavior probably aids in reducing predation. As our research progresses and 

questions are answered about woodrat ecology (for example, habitat 

requirements), management decisions can be better informed regarding this 

species and others that depend upon it for their sustenance. 

Acknowledgments 

Annette Tenneboe, Tamra Sandoval, Jeff Schneiderman, and Ruth Elbert assisted 

with field work, including vegetation surveys and trapping of small mammals. 

Sandi Andreas advised us on database design, helped to convert data for analysis 

from the Data General minicomputer to the Macintosh for analysis, and 

converted data to IBM computer formats. David Grubbs and Michelle Selmon 

provided constructive suggestions on a draft of this paper. We gratefully 

acknowledge all of these contributions. 

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