USDA Forest Service RMRS-RN-41. 2009. 1

Introduction

Salamanders play important roles in forested ecosystems. They are long-lived, have low mobility, and 

are sensitive to environmental changes, thus they are 

good indicators of ecosystem health (Davic and Welsh 

2004; Russell and others 1999; Welsh and Droege 

2001). Terrestrial salamanders (Plethodontidae), in 

particular, require moist conditions and are vulnerable to land management activities, such as prescribed 

burns and logging, that alter forest floor environments 

(Welsh and Droege 2001). The study of terrestrial 

salamanders is important to understanding the impact 

of these activities and making informed management 

decisions.

Fire can alter populations both through direct mortality and through indirect effects via habitat alteration. 

Information on salamander response to fire is limited 

and primarily restricted to the southeastern United 

States (Pilliod and others 2003). Although little information is available, direct mortality is thought to be 

fairly low (Renken 2006; Russell and others 1999). 

Individuals may find refuge underground because 

the majority of many salamander populations are 

subterranean (Bailey and others 2004; Petranka and 

Murray 2001; Taub 1961). On the other hand, high 

susceptibility to fire has been suggested because terrestrial salamanders have small home ranges, move 

slowly, and are probably limited in their ability to disperse (Kleeberger and Werner 1982; Staub and others 

1995). Studies of prescribed fire’s effects in the southeastern United States found no changes in relative 

abundance of terrestrial salamanders (Ford and others 

1999; Keyser and others 2004; Kilpatrick and others 

2004; Moseley and others 2003), whereas salamander numbers were lower in forests where prescribed 

fire and thinning were applied in Maryland (McLeod 

and Gates 1998). Studies of prescribed fire in western 

North America are more limited and have generally 

been conducted in coastal climates that host different salamander communities and natural fire regimes 

compared to interior forests. No changes in relative 

abundance of slender salamanders (Batrachoseps sp.) 

were found for prescribed fire in coastal California, 

(Vreeland and Tietje 2002). However, in coastal 

Oregon, some salamander species declined from sites 

that were burned and clearcut (Cole and others 1997). 

Bagne, Karen E.; Purcell, Kathryn L. 2009.Response of two 

terrestrial salamander species to spring burning in the Sierra 

Nevada, California. Res. Note RMRS-RN-41. Fort Collins, CO: 

U.S. Department of Agriculture, Forest Service, Rocky Mountain 

Research Station. 10 p.

Authors

Karen E. Bagne is a wildlife biologist with the USDA Forest Service, 

Rocky Mountain Research Station in Albuquerque, New 

Mexico. She completed this research in partnership with the 

Pacific Southwest Research Station, Sierra Nevada Research 

Center in Fresno, CA, while working on her Ph.D. in Biology at 

the University of California, Riverside.

Kathryn L. Purcell is a research wildlife biologist with the Pacific 

Southwest Research Station’s Sierra Nevada Research Center 

in Fresno, CA. She received her Ph.D. from the University of 

Nevada, Reno in Ecology, Evolution, and Conservation Biology.

United States

Department

of Agriculture

Forest Service

Rocky Mountain

Research Station

Research Note 

RMRS-RN-41

September 2009

Response of Two Terrestrial Salamander Species 

to Spring Burning in the Sierra Nevada, California

Karen E. Bagne and Kathryn L. Purcell

Abstract—Terrestrial salamanders may be vulnerable to prescribed fire applications due to their moist, 

permeable skin and limited mobility. We present data collected on terrestrial salamander populations in a 

ponderosa pine-dominated forest in the Sierra Nevada where fire was applied in the spring. Two species, 

Sierra ensatina (Ensatina eschscholtzi platensis) and gregarious slender salamander (Batrachoseps 

gregarius), were captured under coverboards. Capture rates of ensatinas declined within the first 2 years 

after burning, but postfire captures were similar to or greater than capture rates on unburned plots. Capture 

rates of slender salamanders were more variable, but high capture rates on burned plots suggest they 

persist following fire. We captured fewer small ensatinas within 2 years of burning, but sizes of slender 

salamanders pre and post burning were similar. Salamanders were captured in both closed and open canopy 

forests, and presence under individual coverboards was associated with deeper litter and greater canopy 

closure. Coverboards may be avoided for a year or more by gregarious slender salamanders, and capture 

rates were highest during winter and early spring. Though sample sizes were small and conclusions should 

be made with caution, results indicated no strong adverse effects from spring burning. Suitable habitat may 

have been maintained by the patchy burn pattern characteristic of spring burns.
2 USDA Forest Service RMRS-RN-41. 2009.

Insights into salamander response to prescribed fire 

may be available from wildfire studies, but differences in fire season and severity are likely to alter 

response. Following a New Mexico wildfire, no effect 

on salamander presence was observed, but there was 

a shift to smaller size classes in a terrestrial salamander (Cummer and Painter 2007).

Prescribed fire is commonly used in coniferous 

forests of the western United States where decades 

of fire suppression, climatic changes, and other types 

of disturbance have altered forest structure and natural fire regimes (Pyne 1984). Fire regime changes in 

California’s Sierra Nevada have generally increased 

tree and shrub densities, as well as lengthened fire return intervals and increased fire severities (McKelvey 

and others 1996). Historically, fires on the west slope 

of the Sierra Nevada occurred late in the growing 

season from mid-summer to early fall (Caprio and 

Swetnam 1995), but high fuel levels and air quality 

limitations have led managers to prescribe fires during periods of high moisture, including in the spring. 

On one hand, these periods coincide with terrestrial 

salamander emergence from subterranean habitats, 

creating a higher risk of direct mortality (Pilliod 

and others 2003). On the other hand, because these 

fires are generally less severe and burn incompletely, 

habitat alteration is small compared to contemporary 

wildfires.

Many salamander species in the Sierra Nevada 

are endemic and little is known about their population status (Jennings 1996) much less fire’s affect on 

the species’ populations or habitats. Because many 

studies lack replication and pre-fire data (Russell 

and others 1999), we collected data on salamanders 

before and after prescribed burns on multiple plots. 

We present data collected on terrestrial salamander 

populations up to 6 years following burning in the 

Sierra Nevada, California, where prescribed fire was 

applied in the spring. We also measured individual 

size because differences in prey size or probability 

of surface activity can lead to size-dependent fire 

response (Cummer and Painter 2007). For instance, 

small salamanders have a larger surface area to volume ratio and may be more vulnerable to disturbance 

(Hairston 1987). Additionally, a greater proportion of 

small (young) individuals may indicate that individuals are using suboptimal habitats (Welsh and others 

2008). We also collected data on physical characteristics of coverboard arrays to investigate their influence 

on capture rates.

Methods

The study took place from 2001 to 2004 in the 

Sierra National Forest, Fresno County, California, at 

elevations ranging from 1,000 to 1,400 m, approximately 65 km east of the city of Fresno (37°02’N, 

119°15’W). The forests of the study area were dominated by ponderosa pine (Pinus ponderosa) but 

also contained canyon live oak (Quercus chrysolepis), black oak (Quercus kelloggii), incense cedar 

(Calocedrus decurrens), sugar pine (Pinus lambertiana), and white fir (Abies concolor). Ground cover 

consisted primarily of a thick layer of pine needles 

interspersed by mountain misery (Chamaebatia foliolosa), a common perennial ground-cover species. 

Forested areas were intermixed with granitic outcrops 

and shrub fields dominated by whiteleaf manzanita 

(Arctostaphylos viscida). Precipitation fell primarily during the winter in the form of rain or snow. 

Cumulative precipitation from January through May 

at the Pine Flat Dam (National Weather Service 

Cooperative Station 46896, approximately 24 km 

from the study area) was 30.7 cm, 17.8 cm, 24.6 cm, 

and 19.1 cm from 2001 to 2004, respectively. The 

average precipitation for the preceding 10 years was 

41.9 cm.

The study area consisted of six plots, four of which 

the U.S. Forest Service burned during the study for its 

fuel management programs. Prior to 1997, the most 

recent fire in the area was in 1941. Two plots were 

burned in April of 2002. Two plots that were burned in 

April 1997 and May 1998, respectively, were burned 

again in June 2003. Thus, there were two types of 

plots that were burned once: those burned in 2002 

and those burned in 1997/1998. In 2003, the plots 

burned in 1997/1998 were burned again (table 1). The 

remaining two plots remained unburned. Three plots 

(one in each treatment type) were on the Rush Creek 

drainage and three were approximately 4 km away on 

the Big Creek drainage, both of which flow into Pine 

Flat Reservoir on the Kings River.

Surveys for terrestrial salamanders were conducted 

from 2001 to 2004 using coverboard arrays (DeGraaf 

and Yamasaki 1992). Boards were made of 30- by 30- 

by 2.5-cm plywood and left to weather outside at least 

two months before placement (Grant and others 1992; 

Monti and others 2000) at study locations in early 

2001. Boards were arranged on each of the 6 plots 

in 3 separate arrays of 18 boards each (324 boards 

total). The boards were placed 12 m apart based on 
USDA Forest Service RMRS-RN-41. 2009. 3

home range estimates (Staub and others 1995) to 

maximize unique individuals available for capture. 

Arrays were generally rectangular and near creeks or 

seeps. We placed two arrays on each plot in a closed 

canopy forest and one in an open forest dominated by 

whiteleaf manzanita because we did not know specific habitat associations. We made a shallow depression 

underneath each board to create a space for the salamanders. Litter was removed during initial placement 

so boards were in contact with the ground, but thereafter litter was allowed to accumulate on the boards 

(fig. 1). We recorded the general characteristics of 

each array before fire, including the slope and aspect 

of the overall terrain and distance to permanent water, 

which was measured from the center of each array using ArcMAP v.8.1 (Environmental Systems Research 

Institute, Inc., Redlands, California). Litter depth 

(mm) and canopy closure (%) were measured at each 

board. Canopy closure was calculated by averaging 

cover values obtained with a spherical densitometer 

from the four cardinal directions while standing at 

the edge of the coverboard and facing away from the 

center.

We conducted surveys of all boards on 6 plots on 

each of 13 days, November through July from 2001 to 

2004. We conducted surveys at least two weeks apart, 

usually the day after rainfall when the surface soil 

was moist and salamanders were expected to emerge 

(Fellers and Drost 1994; Marsh and Goicochea 2003). 

Surveys were not conducted during rainfall (Jaeger 

1980). Winter snow and dry conditions from June 

to October limited our survey opportunities. We removed boards prior to burning and replaced them 

immediately following fire on one plot in 2002, but 

found low fire intensities made it unnecessary. We 

replaced lost or damaged boards with boards of the 

same age that had been stored outside to weather 

equally.

We identified the species of each captured salamander and measured from the tip of the snout to the 

posterior tip of the vent (i.e., snout-vent length or SVL) 

(Petranka 1998). We marked individuals of Sierra ensatina with a visible implant elastomer (Northwest 

Marine Technologies) for individual identification 

(Donnelly and others 1994). Marks were placed on 

the belly close to the limbs in three places with two 

of four possible colors. Marking slender salamanders 

in the field proved to be difficult and was abandoned.

We calculated capture rates of salamanders as the 

number of salamanders per survey effort (captures 

per 100 coverboard survey days). Capture rates were 

used to evaluate survey timing, differences in array 

placement by habitat, and burn effects. We note that 

captures and comparisons are for the “visible population” and acknowledge that the majority of the 

population is subterranean (Taub 1961). Results were 

Table 1—Numbers of salamanders captured under coverboards for survey days from 2001 to 2004. 

Numbers are grouped by burn history with 2 replicate plots per treatment (control, burned once, 

burned twice) and 3 arrays of 18 boards each. Diagonal hatching indicates surveys done after 

plots were burned once and cross-hatching after plots were burned twice. Where no hatching 

appears, no fire had been recorded for greater than 50 years.

Year 2001 2002 2003 2004

Number of survey days 4 2 5 2

Sierra ensatina Plot

(Ensatina eschscholtzi platensis)

Control U1 1 0 6 2

U2 1 0 2 1

Burned 2002 B1 5 0 5 1

B2 3 0 5 1

Burned 1997/98 and 2003 P1 0 1 5 0

P2 0 1 9 2

Gregarious slender salamander

(Batrachoseps gregarius)

Control U1 0 0 0 0

U2 0 0 1 0

Burned April 2002 B1 0 1 14 1

B2 0 0 0 0

Burned April 1997 and June 2003 P1 0 1 8 1

Burned May 1998 and June 2003 P2 0 0 0 0
4 USDA Forest Service RMRS-RN-41. 2009.

not analyzed statistically because capture rates per 

treatment were low. Appropriate statistical analysis 

for the study design would require a time by treatment interaction, which would further reduce the 

available degrees of freedom. We present all data and 

discuss capture rates and mean salamander size related to burning qualitatively.

We modeled salamander presence under individual boards using a generalized linear model with a 

binomial distribution to investigate factors related to 

habitat and topography (PROC GENMOD in v.8.01, 

SAS Institute, Inc., Cary, North Carolina). We tested 

litter depth and average canopy closure in this model 

with a repeated measures statement identifying the 

array as the subject. Measures related to arrays (i.e., 

slope and distance to water) were also compared 

using generalized linear models with the repeated 

subject designated as the plot. Variables were considered to be related to salamander presence when 

P < 0.05 for the parameter estimate from the generalized estimating equation (GEE), which fits models to 

the correlated responses.

Results

Fires were generally of low intensity and left a 

patchwork of burned and unburned areas. Few of the 

boards left in place during burning were consumed. 

Of these nine burned arrays, three lost one board, one 

lost two, and one lost three.

We found 78 terrestrial salamanders of two species: 51 Sierra ensatina and 27 slender salamanders 

(table 1). Recapture rates of Sierra ensatina were 

low with nine recaptures of six individuals over four 

Figure 1—Coverboard lifted with adult Sierra ensatina (Ensatina eschscholtzi platensis) underneath (photo by Karen E. Bagne). 

Photo was scanned from 35 mm film and was adjusted for brightness and contrast.
USDA Forest Service RMRS-RN-41. 2009. 5

years. Slender salamanders were identified as gregarious slender salamanders (Batrachoseps gregarius) 

based on the location of the study area and physical characteristics of the species as described by 

Jockusch and others (1998) (fig. 2). A single adult 

California newt (Taricha torosa sierrae) was the 

only other salamander captured. The only other vertebrates we found were three Western fence lizards 

(Sceloporus occidentalis taylori) and one Northern 

alligator lizard (Elgaria coerulea palmeri).

Though we conducted surveys on various dates 

when soils were moist, captures in March (3.7 captures per 100 coverboard survey days) and April 

(2.5 captures per 100 coverboard survey days) were 

the highest (fig. 3). Capture rates were lower in May 

(1.2 captures per 100 coverboard survey days) and 

only two salamanders were captured on each of our 

survey days in July, November, and December. In addition, salamander presence under boards increased 

over time, but no gregarious slender salamanders 

were captured in 2001 (table 1).

Figure 2—Gregarious slender 

salamander (Batrachoseps 

gregarious) after capture under 

coverboard (photo by Karen E. 

Bagne). Photo was scanned from 

35 mm film and was adjusted for 

brightness and contrast.

Jan Feb Mar April May June July Aug Sept Oct Nov Dec

4

3.5

3

2.5

2

1.5

1

0.5

0

Capture rate

Month

Figure 3—Capture rates for 

all salamanders combined 

by month for 2001 to 

2004. Capture rate is 

number of captures per 

100 coverboard survey 

days. Months where no 

capture rate appears had 

no surveys rather than no 

salamanders.
6 USDA Forest Service RMRS-RN-41. 2009.

The percentage of canopy closure differed between 

the two sampled habitats, averaging 90.9% (SD = 7.1) 

for closed forest coverboards and 67.0% (SD = 24.9) 

for open forest coverboards. Gregarious slender salamander capture rates between the two habitats were 

similar, with 0.6 vs. 0.7 captures per 100 coverboard 

survey days for closed and open forest, respectively. 

More Sierra ensatinas were captured in closed forest arrays than open ones (1.5 vs. 0.6 captures per 

100 coverboard survey days). Salamander presence 

at arrays was not related to slope (deviance = 1.06, 

Z = -0.55, P = 0.58; table 2) or distance to water (deviance = 1.06, Z = 0.93, P = 0.35; table 2) though 

the maximum distance for any array was 225 m. 

Terrestrial salamanders were found under boards with 

greater litter depth (deviance = 0.99, Z = 2.16, P = 

0.03; table 2) and increased canopy closure (deviance 

= 0.96, Z = 3.12, P = 0.002; table 2). Litter depth and 

canopy closure were positively correlated (r2

 = 0.17).

Captures varied considerably by plot. Gregarious 

slender salamanders were only present on plots in the 

Big Creek drainage and none were found in the Rush 

Creek drainage (table 1). Captures of Sierra ensatinas 

declined after burns in 2002 (table 3), but preburn 

capture rates for these two plots were higher than 

those of the unburned plots, and postburn capture 

rates were similar to the unburned plots. Postburn 

capture rates after 2002 burns were similar to those 

on plots 3 to 6 years postburn (table 3). No gregarious slender salamanders were captured on burned 

plots prior to burning, but postburn capture rates in 

2002 were greater than captures on the plot burned 

in 1997 or the unburned plot (table 3). We captured 

too few salamanders on the two surveys following the 

second burn in 2003 to evaluate the effects of burning 

twice (table 1). However, both Sierra ensatinas and 

gregarious slender salamanders were present after the 

second burn.

The mean size of captured salamanders also varied 

by plot. Results from plots burned in 2002 suggest the 

mean size of Sierra ensatinas increased after burning, 

indicating fewer small salamanders (table 4). Mean 

size of gregarious slender salamanders was similar on 

plots of all burn histories (table 4).

Discussion

Although our study design provided for pre and 

postburn data collection, low counts were problematic 

Table 3—Number and capture rate for salamanders found under coverboards for survey days 2001 to 2004. 

Capture rate is number of captures per 100 coverboard survey days. P1, burned April 1997, and P2, burned 

May 1998, were burned again in 2003, but post-2003 burn data are excluded here.

Sierra ensatina Gregarious slender salamander

(Ensatina eschscholtzi platensis) (Batrachoseps gregarius)

Burn status Plot Number Capture rate Number Capture rate

Unburned U1 9 1.28 0 0

U2 4 0.57 1 0.14

1-2 years postburn B1 pre 5 2.31 0 0

B1 post 6 1.23 16 3.29

B2 pre 3 1.39 0 0

B2 post 6 1.23 0 0

3-6 years postburn P1 6 1.01 9 1.52

P2 10 1.68 0 0

Table 2—Mean (SE) physical characteristics of boards where terrestrial 

salamanders were present (n = 268) or absent (n = 56) over the entire study 

period 2001 to 2004. P-values are from generalized linear models testing 

differences in salamander presence.

Variable Present Absent P

Slope (%) 17.9 (1.4) 15.0 (0.5) 0.58

Distance to water (m) 62.4 (8.5) 79.2 (4.0) 0.35

Litter depth (mm) 53.8 (4.5) 41.0 (2.0) 0.03

Average canopy closure (%) 89.6 (1.2) 81.6 (1.2) 0.002
USDA Forest Service RMRS-RN-41. 2009. 7

Table 4—Mean salamander snout-vent length (mm) (SVL) and standard errors (SE) by burn status and plot. Three- to 

six-year data from plots that were burned again in 2003 are excluded.

Sierra ensatina Gregarious slender salamander

(Ensatina eschscholtzi platensis) (Batrachoseps gregarius)

Burn status Plot Number Mean SVL ± SE Number Mean SVL ± SE

Unburned U1 4 46.3 ± 6.6 0 0

U2 9 53.2 ± 6.7 1 32.9

1-2 years postburn B1 pre 5 41.0 ± 8.4 0 0

B1 post 6 59.0 ± 5.5 15 31.2 ± 1.0

B2 pre 3 34.4 ± 3.8 0 0

B2 post 6 37.8 ± 5.1 0 0

3-6 years postburn P1 6 34.4 ± 5.9 8 32.3 ± 1.3

P2 10 51.5 ± 5.3 0 0

in that they prevented both evaluation of population 

changes over time and using recapture data to estimate 

detectability or population size. While we recognize 

the limitations of the data, we know of no other published studies on the effects of fire on salamanders 

from the Sierra Nevada. Thus, these data are a first 

step toward filling a gap in our understanding of prescribed fire effects on terrestrial salamanders and offer 

insights to aid future study design. To our knowledge, 

no information outside of taxonomic studies has been 

published on gregarious slender salamanders.

Both Sierra ensatinas and gregarious slender salamanders persisted after burn applications up to 6 years 

following fire. Despite initial concerns about vulnerability, salamander numbers were not lower on burned 

sites. Capture rates of Sierra ensatinas declined in the 

first two years after burning but were similar to unburned plots and plots 3 to 6 years postburn. Only 

one gregarious slender salamander was caught on unburned plots and none were captured in the preburn 

period, but high capture rates on burned plots indicate 

that these salamanders persisted following fire.

We are cautious in our conclusions, but feel there 

is enough evidence to conclude that low intensity 

spring burning did not significantly harm terrestrial 

salamanders up to 6 years postfire. These results are 

consistent with others who have found neutral responses to fire despite differences in the studied fire 

intensities (Ford and others 1999; Keyser and others 

2004; Kilpatrick and others 2004; Moseley and others 

2003; Vreeland and Tietje 2002).

It is possible that differences in capture probabilities related to fire were responsible for the patterns in 

capture rates we observed. Coverboards may be more 

attractive and have greater probability of capture following fire if natural cover is reduced, but capture 

rates were lower or unchanged on plots immediately 

following fire. Additionally, differences in coverboard capture rates following fire may not represent 

a change in numbers, but a change in surface activities. Temperature and surface moisture are known to 

affect surface activity and consequently capture rates 

(Petranka and Murray 2001). Our captures increased 

in years of higher precipitation, but we were able to 

reduce this bias by completing surveys on all plots 

within a single day when conditions were similar 

for burned and unburned plots. Differences in surface activity related to fire may explain why, on the 

plot where capture rates were most reduced following burning, we also found an increase in mean size. 

Smaller salamanders may be captured less often at 

the surface following fire if they are more sensitive to 

surface conditions and if fire reduces favorable conditions. Fire’s affect on surface activity needs further 

investigation.

Based on our findings, we can make a number of 

recommendations for future studies. Though we did 

not investigate alternative sampling methods, we 

felt the use of coverboards was generally successful. Surveys could be completed in one day by two 

people, the survey area was consistent in space and 

time, and habitat disturbance was minimal. Numbers 

of individuals captured were low overall (1.85 captures per 100 coverboard survey days), but capture 

rates were similar to those reported by Vreeland and 

Tietje (2002) from the California oak woodlands 

(1.74 captures per 100 coverboard survey days). 

Although spring and fall are common survey months, 

at this location capture rates were highest during late 

winter and early spring with few captures in the fall 

(fig. 3). Low precipitation during most of the study 

period may have contributed to low capture rates. 
8 USDA Forest Service RMRS-RN-41. 2009.

With these capture rates and the very low ensatina recapture rate, we concluded the number of sampling 

days and/or coverboards needs to be increased. We 

were limited by funding and winter access to our 

study area, which may be improved in other studies, 

but we were also limited by the restricted period of 

precipitation in a Mediterranean climate. We did not 

sample in January or February, but these may also be 

good sampling periods for this climate considering 

our high capture rates in March. Although replication 

of plots and arrays helped reduce bias related to plot 

differences, the strong differences between capture 

rates highlights the need for replication. Gregarious 

slender salamanders avoided coverboards for at least 

one year after installation, longer than for Sierra ensatinas or reported for other species (Monti and others 

2000). This should be taken into account when initially placing boards. Based on captures by habitat, 

captures in this region can be maximized by sampling 

only closed canopy forest, but because both salamander species were also captured in open forest mixed 

with whiteleaf manzanita, the suitability of this habitat should be noted for management.

Association of salamander presence with litter 

depth and persistence after spring fire indicates that 

spatial heterogeneity in burning may be an important 

factor in predicting effects. Fire intensities were low, 

resulting in incomplete consumption of the litter layer 

and logs as well as limited affect on canopy closure. 

As a result, suitable terrestrial salamander habitat was 

retained. Although high intensity fires would likely 

have a different impact, low intensity burns are typical of management burning on National Forest land in 

the Sierra Nevada where high fuel loads and air quality issues limit burning to moist times of year. Our 

findings are encouraging evidence for the persistence 

of terrestrial salamanders during spring management 

burning up to 6 years postfire. Finally, terrestrial 

salamanders are still threatened by climate change 

(Parra-Olea and others 2005), disease (Byrne and 

others 2008), and other forest management activities 

such as logging (Herbeck and Larsen 1999; Petranka 

and others 1993) and road building (deMaynadier and 

Hunter 2000), and deserve more active study.

Acknowledgments

Funding for this project was provided by the U.S. 

Forest Service, Pacific Southwest Research Station, 

Sierra Nevada Research Center, and Joint Fire 

Sciences Program. We would like to thank the field 

crew members who helped collect data and, in particular, G. Burton who generously donated his time. 

Anonymous reviewers supplied comments that improved the manuscript. Permission to handle and mark 

salamanders was provided through a scientific collecting permit issued by the California Department of Fish 

and Game (SC-004962) and an animal use agreement 

with the University of California, Riverside.

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