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American Society of Mammalogists

Assessment of Abundance of San Joaquin Kit Foxes by Spotlight Surveys Author(s): Katherine Ralls and L. Lee Eberhardt Source: Journal of Mammalogy, Vol. 78, No. 1 (Feb., 1997), pp. 65-73 Published by: American Society of Mammalogists Stable URL: http://www.jstor.org/stable/1382639

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ASSESSMENT

OF ABUNDANCE

OF SAN JOAQUIN KIT FOXES

BY SPOTLIGHT SURVEYS

KATHERINE RALLS AND L.

LEE EBERHARDT

National Zoological Park, SmithsonianInstitution, Washington,D.C. 20008 (KR)

  • 2528 West Klamath,Kennewick, WA99336 (LLE)

Biologists of the California Department of Fish and Game have conducted quarterly spot-

light surveys of San Joaquin kit foxes,

Vulpes macrotis mutica, and potential species of

prey along seven 48-km routes since 1970. The annual reproductive cycle of the foxes was reflected by a seasonal cycle in the counts. Counts of foxes during surveys in June were correlated with total precipitation during the previous rainfall season, but not with concur- rent counts of prey. Number of foxes seen/0.8-km interval was more highly correlated with estimated sighting distance based on physical features, such as hills and plains, than with current vegetation density or type. Mean number of foxes counted per survey per route ranged from two to 20. Counts of foxes increased over time on one route, decreased over time on another, and showed a curvilinear trend on two routes. The average number of foxes seen over all routes did not show any long-term trend. However, small samples,

missing data, and lack of replication limited statistical analysis and interpretation of surveys. Statistical tests on simulated data similar to the data collected during the surveys indicated that larger samples, which could be obtained by replication, would be needed to detect

changes in the population of foxes with any efficiency (statistical power). An expanded spotlight-survey program of kit foxes, with additional routes and replicated surveys, could

be used to monitor population trends throughout the range of this endangered subspecies.

Key words: Vulpes macrotis mutica, kit fox, spotlighting, surveys

The San Joaquin kit fox, Vulpes macrotis

mutica (Mercure et

al.,

1993), is listed as of California and as

threatened by the state

endangered at the federal level. To monitor the abundance of this fox, the California

Department of Fish and Game initiated a series of spotlight surveys along rural roads

in 1970. Biologists have conducted quarter- ly surveys for kit foxes and potential prey

species (leporids, kangaroo rats) along these routes from 1970 to date.

Spotlighting

has

been

many species of wildlife

used

to

census

(Barnes and Tap-

per, 1985; Progulske and Duerre, 1964) in- cluding red foxes, Vulpes vulpes (Stahl, 1990; Stahl and Migot, 1990; Weber et al.,

1991). The annual reproductive cycle of red

foxes is reflected by a corresponding annual increase and decrease in the number of fox-

es counted during seasonal spotlight sur-

veys

(Weber et al., 1991). Because repro-

duction of kit foxes is also strongly season-

al, with young born in February (O'Farrell, 1987), we expected to observe a similar

seasonal cycle in the number of foxes

counted during surveys by personnel of the California Department of Fish and Game. The San Joaquin Valley is a desert hab- itat with annual precipitation occurring pri- marily as winter rains. Variations in annual

rainfall strongly influence the growth and

reproductive

success

of

annual

plants,

which

in turn affect densities

of rodents.

The density of nocturnal rodents, the pri- mary prey of San Joaquin kit foxes in many

areas (Morrell, 1972; White et al., 1996), correlates positively with the amount of

precipitation during the previous rainfall season (K. Allred, in litt.). Drought reduces reproductive success of kit foxes due to de-

Journal of Mammalogy, 78(1):65-73,

1997

65

66

JOURNALOF MAMMALOGY Vol. 78, No. 1

creased availability

of prey (Egoscue,

1975;

White

and Ralls,

1993).

Therefore,

we pre-

dicted

that

the

number

of

foxes

counted

during surveys

would

be

correlated

with

both counts of prey and precipitation

during

the previous

rainfall

season.

 

We examine

the survey

data of

the Cal-

ifornia Department

of Fish

and

Game

for

an annual cycle of abundance

of

kit foxes,

correlations

among counts of foxes,

counts

of prey, and annual precipitation,

effects

of

variations

in physical

features

and vegeta-

tion along

the route on the number

of foxes

seen,

and

trends

in

the

number

of

foxes

seen

over

time.

We also

consider

the effec-

tiveness

of

the

survey

design

for detecting

trends

in population

size

of

kit foxes.

 

MATERIALSAND METHODS

The locations of the seven original 48-km sur-

vey routes range from Kern and San Luis Obis-

po counties in the south to Fresno and Merced

counties in the north (Fig.

1). All routes were

on public roads, some of which were unpaved.

An eighth route, Soda Lake, was added in 1989.

Due to increasingly dangerous driving condi-

tions, the Blackwells Comer route was replaced

by a new route (Allensworth, not shown in Fig.

1) in the same general area in 1990. One survey

was scheduled quarterly along each route.

Whenever possible, surveys were conducted

during March, June, September, and December.

Occasionally, however, surveys were conducted

in other months (e.g., the winter survey for one

year was delayed until January of the following

year) or missed because sections of the routes

which it was seen were recorded. Total numbers

of black-tailed jackrabbits (Lepus californicus),

desert cottontails (Sylvilagus audubonii), and

kangaroo rats (Dipodomys) also were recorded.

The California Department of Fish and Game

provided us with the survey data as a dbase file

that included the number of foxes, but not the

number of potential prey, counted during each

survey. We also obtained the original data sheets

for the Elkhorn route from which we tallied the

numbers of each potential prey species seen dur-

ing each survey. Thus, we were able to look for

seasonal cycles of abundance and trends in the

number of foxes counted over time on all survey

routes and were able to examine relationships

between numbers of foxes and numbers of po-

tential prey for the Elkhorn route.

We were interested particularly in the Elkhorn

route because it runs through the Carrizo Plain

Natural Area, where one of us (KR) has con-

ducted a radiotelemetry study of kit foxes (Ralls

and White, 1995; White and Ralls, 1993; White

et al., 1994). We obtained annual precipitation

data for this route from weather recording sta-

tions in the area. To study the possible effects

of terrain and vegetation on the number of foxes

seen, we drove the Elkhorn route in 1992 during

the day and categorized viewing distance based

on terrain, such as hills

and plains (which re-

mained constant over the years), density of veg-

etation (which might have changed over the

years), and the predominant vegetation type by

0.8-km intervals. The effect of terrain on view-

ing distance was scored in three categories: <75

m visibility, 75-250 m visibility, and 250 m (the

limit of the light beam). Density of vegetation

was scored in one of five classes ranging from

impenetrable with few or no openings (class 1,

were impassable.

 

poorest visibility) to sparse, short, or no vege-

Two observers, one sitting on each side of a

tation (class 5, highest visibility). The predomi-

vehicle

that enabled their eyes

to

be

m

nant vegetation was scored as either: 1) peren-

above the level of the road, conducted each -1.5 sur-

nial shrubs (e.g., Ephedra, Allenrolfea, or Atri-

vey by driving the designated route at ca.

16-

  • 24 km/h. Each observer scanned one side of the

route with a spotlight. Strength of spotlights in-

creased over the years but this had no discern-

able effect on the number of foxes seen (J. Lid-

berg, pers. comm.).

Spotlights

of

at

least

850,000 candle power currently are used. When

an animal was detected by the light reflected

from its eyes, the driver stopped the vehicle, and

the observers identified the animal. The species

of each predator and the time and mileage at

plex), (2) tall or bushy annual plants (e.g., Amar-

anthus, Brassica), 3) short annual plants or

fallow (e.g., Amsinckia, old grain fields), or 4)

non-native grassland and perennial shrubs.

We used data from summer surveys,

when

young foxes emerge from dens at night but have

not yet dispersed from their natal home ranges,

to examine relationships among counts of foxes,

counts of prey, and precipitation. Because den-

sities of rodents on the Elkhorn Plain are cor-

related with precipitation during the previous

February 1997

RALLS AND EBERHARDT-SURVEYS OF KIT FOXES

67

80 120 40 0 ... Kilometers 0 25 50 75 S - Miles - \ N----
80
120
40
0
...
Kilometers
0
25
50
75
S
-
Miles
-
\
N----
>
F
Tf
FeOrtigalita
1
Blackw
,
CornerI 's
' ..
ElkhornPlain
L
.
.
I
N
B
s
o
n
.-,
...
.
Belridge-McKittrick
,
Soda
Lake
.
I
Lake
SSoda

FIG. 1.-Map

showing the locations of the seven original survey routes, and the Soda Lake route

added in 1989, in relation to the probable range of the San Joaquin kit fox (shaded area). The dashed

line encloses all historic and current records of this fox. However, ca. 80% of this area is either

urbanized or intensely cultivated and has extremely low densities of foxes. Much of the remaining

undeveloped land has unsuitable or marginal habitat for kit foxes. Furthermore, there have been no

recent surveys for this fox in several parts of its range, such as the eastern side of the San Joaquin

Valley along Highway 99.

68

JOURNALOF MAMMALOGY Vol.78, No. 1

rainfallseason (K. Allred, in litt.), we

compared

summer counts of foxes and concurrentcounts

of prey to total precipitationduring the previous

rainfallseason

(e.g.,

June 1990 counts with rain-

fall from 1 July 1988 to 30 June 1989). We com-

puted Pearson product-moment correlationco- efficients among these variables and used Bon-

ferroni-adjustedprobabilities to reflect the num-

ber of

correlations being tested (Wilkinson,

1990).

To model a possible annual cycle

of abun-

dance on the Elkhorn route, numbers of foxes

counted at or near June of each

year were as- sumed to be the actual abundanceat that time,

and reduced by a constant rate of

change per

month (0.884) until about the following April.

The rate of change was

approximatedby a least-

NJune

St for the

squares fit using the model N, =

months of June to May (months 0-11), follow-

ing the approach of Eberhardt (1987). Here, Nt denotes the population at montht (the following

September,December, or March) and

s denotes

a range of change due to deathsand the dispersal

of young foxes from the area.

We examined trends in populations of foxes

over time for

each route by summingup the sea-

sonal counts to get a yearly total. Based on pop-

ulation simulations conducted by Eberhardt (1992), we transformedthe counts by natural

logarithms and

tested for linearity and curvilin-

would be ex-

earity. A log-linear relationship

pected

if the population were increasing or de-

creasing at a relatively constantrate. The

test for

curvilinearitycompared deviationsfrom a fitted

second-degreepolynomial to those from the lin-

ear

regression(Snedecor and Cochran, 1967).

To examine effectiveness

of

samples of

surveys

counts of the size typically obtainedin

to detect a population trend, we constructeda

simple

stochasticmodel based on the annual cy-

cle. We simulateddata for threeinitial count lev-

els

(high, intermediate,and low) for 5, 7, and

producing 1,000 simulationsfor each

highest-level

simulated counts were

10 years,

case. The

obtainedon the ElkhornPlain routeand the low- est were those obtainedon the Panoche and Or-

tigalita

routes. The model simulateda 10%/year

population.

We

applied

the

log-

decline in the

linear regression test of significance to these simulateddata, and recordedthe numberof re- sults that were significantat the 5% level.

RESULTS

There were numerous cases of missing

data, usually because

one

or more

of

the

quarterly surveys was not conducted. More

surveys were conducted during the dry sea- son (summer and autumn, n = 1,239) than

during the wet season (winter and spring, n = 883), when routes sometimes became im-

passible. Plots of the quarterly counts (Fig. 2) showed the expected annual cycle of abun- dance of foxes, with the highest counts most often occurring during June. Counts

of foxes during summer surveys on the Elk- horn route were correlated with total rain-

fall during the previous rainfall season (r = 0.74, P < 0.01, n = 21), but not with num-

bers of cottontails, jackrabbits, or kangaroo rats. The only counts of prey species that correlated with total rainfall were those for cottontails (r = 0.66, P < 0.05, n = 21). Total number of foxes seen per 0.8-km in- terval on the Elkhorn route was more high-

ly

correlated

with

0.001,

n

=

73, Fig.

terrain

(r2

=

0.24,

P

<

3), than with vegetation

density (r2 = 0.08, n.s.) or vegetation type

(r2

=

0.04,

n.s.).

Mean number of foxes (? 1 SE) counted

per survey

ranged from two

to

20,

with

considerable

variation among routes (Ba-

kersfield,

2.45

?

0.26,

n

=

73;

Belridge-

 

McKittrick, 3.42

?

0.34,

n

=

81; Black-

wells

Corner,

11.12

?

1.38,

n

=

41;

Elk-

horn, 20.34 +

1.42, n = 85; Ortigalita, 2.00

?

0.27,

n

=

55; Panoche, 2.40

?

0.31,

n

=

58;

Soda

Lake,

10.79

 

?

1.57,

n

=

14;

Taft-Fellows, 4.78 ?

0.39,

n

=

81).

 

Counts

of

foxes

 

on the Belridge-Mc-

Kittrick route increased over time while those on the Blackwells Corner route de-

creased (Table 1). A significant curvilin-

ear trend over time was evident

on

the

Elkhorn and Taft-Fellows routes (Table

1). A plot of the yearly means suggested no long-term trend in the number of foxes

seen along the combined survey routes. The trend in the number of foxes seen on the Soda Lake route often differed from

that seen

on

the

Elkhorn

route

(i.e.,

a

higher count on one route was accompa-

nied

by

a lower

count on the other),

al-

though these

two

routes

run down

the

February 1997

80

0

w 60

RALLS AND EBERHARDT-SURVEYS OF KIT FOXES

v

0

S40

20

69

00

0

12

80N

70

Go

24

0'"I

36

I."1*'

48

60

72

MONTH

84

I

"

96

V 0.

108

q

120

..

60-

50

U- so4

.

w 40

w 30

2

z20

VU-

.•II

00 -

144

156

168

180

192

204

216

228

MONTH

240

252

264

a

-

132

144

S

276

288

FIG.2.-Number of foxes seen on the ElkhornPlain route during two successive

12-yearperiods.

The initial point for each plot is a summercount. Curves are fitted by least squares to represent an

annual cycle of abundance (s = 0.88, see text).

western and eastern sides, respectively, of

the same valley and are only separated by 6-8 km.

We used simulated data to examine the effectiveness of the surveys for detecting

population trends over time. Although the simulated populations decreased at 10%/

year, a significant change was detected in

<40%

of

data were

all cases

when only

5 years

of

collected,

even

at the highest

70

JOURNALOF MAMMALOGY

Vol.78, No. 1

5•

40

>.

3 10

z

w

M

0

3 (), -J j 20u z I- i m z 0 0 10 20 30 40
3
(),
-J
j
20u
z
I-
i
m
z
0
0
10
20
30
40
50
60
70
80

HALF-MILESFROM START OF ROUTE

FIG.3.-Mean

numberof foxes sighted/0.8-km interval (line with points; left-hand scale) versus

category of viewing distance based on terrain (line without points; right-handscale) on the Elkhorn

Plain route.

population level (Fig. 4). Results improved if 7 years of data were collected but statis-

tical power became satisfactory only with 10 years of data, and then only for the two

higher population levels.

DIscusSION

The number of foxes

counted during a

spotlight survey is influenced by both the

number of foxes present and the ease with which they can be seen. The mean number of foxes seen per survey varied substantial-

ly across the survey routes, ranging from >20 on the Elkhorn to two on the Ortigalita route. Estimated sighting distance account- ed for only ca. 25% of the variation in the number of foxes seen at various points along the Elkhorn route. Thus, it seems

TABLE1.-F-values for two tests for trend in size of populations offoxes over time along individual

survey routes; n is the number of years

in which at least three surveys were conducted on each

route. P-values are provided only for tests significant at P

0.10.

-

Route

Bakersfield

Belridge-McKittrick

Blackwells Corner

Elkhorn

Ortigalita

Panoche

Taft-Fellows

n

17

20

8

23

11

13

21

Log-linear

relationship

0.26

5.64 (P

25.10 (P

2.60

<

<

0.74

2.50 (P

<

0.09

0.05)

0.005)

0.10)

Curvilinearity

0.34

0.03

1.48

13.37 (P

< 0.005)

0.04

0.08

5.23 (P

<

0.05)

February 1997

1.0

RALLS AND EBERHARDT-SURVEYS OF KIT FOXES

71

HIGHESTPOPULATION LEVEL

0

0.8

 

w

0.6

 

INTERMEDIATE

 

o

0.4

 

POPULATIONLEVEL

c

0.2

 
  • 0 ,o

LOWESTPOPULATION

LEVEL

 

0.0

 
 

5

6

7

8

9

10

 

NUMBEROF YEARSDATA COLLECTED

FIG.4.-Results

of statisticaltests on simulated survey data with a 10% decrease in size of pop-

ulation

per

year. The left-hand scale (power) indicates how often the test reported a statistically

significantchange. The test was applied to data generated for 5, 7, and 10 years.

 

likely that differences in the mean number of foxes seen on the various routes reflect true abundance of foxes to some extent. Further research to relate mean counts of foxes on the various routes to mean sighting distance and estimates of density of foxes along the routes would facilitate interpre- tation of survey results. As predicted, the highest counts of foxes

tended to occur during summer surveys, in-

dicating that a single survey often is suffi-

cient to detect reproduction of foxes.

Be-

cause kit foxes

live

in pairs and can pro-

duce three to five young annually that dis- perse in the autumn (O'Farrell, 1987), the summer population can be several times that of other seasons. We predicted that the number of foxes seen during June surveys would be positively correlated with both counts of prey and the amount of precipi- tation during the previous rainfall season. Summer counts of foxes on the Elkhorn route were related to precipitation, but not counts of prey. Because density of kit foxes

is related to density of prey and rainfall in- fluences density of foxes via its effects on

density of prey (White and Rails, 1993), these results suggest that spotlight surveys are not a sensitive method for assessing the abundance of prey of kit foxes. Habitat destruction is the greatest threat to the survival of the San Joaquin kit fox

(Williams, 1992). However, most of the sur- vey routes are in relatively remote areas, and counts of foxes declined over time only on the Blackwells Corner route. This de- cline was likely due to the conversion of natural habitat to cultivated agricultural

lands along much of the route (G. Presley,

pers. comm.).

Plots of the mean yearly

number of foxes seen on all routes (includ-

ing the Blackwells

Corner route) did not

show any long-term trend, suggesting that populations of foxes are persisting where natural habitat remains.

How might the design of the surveys be improved? Due to the topography and large number of private landowners in the areas

  • 72 JOURNAL OF MAMMALOGY

Vol. 78, No.

I

to be surveyed, it would be impossible

to

conduct conventional line-transect surveys.

There are few specific guidelines for the de- sign of spotlight surveys using existing

roads,

although

Norton-Griffiths

(1975)

recommends using a number of repeated counts during a short interval to estimate

sampling error. Small samples (i.e., the low

number of

foxes

seen

per

survey

along

most routes), missing data (i.e., one or more

of the quarterly surveys was not conduct- ed), and lack of replication limited our anal-

ysis and interpretation of survey data. Sta- tistical tests on simulated data similar to that collected during the surveys indicated that larger samples would be needed to de- tect changes in populations of kit foxes with any efficiency (statistical power) using spotlight surveys. These larger samples could be collected by repeated runs of in- dividual routes during the same month. However, it is impossible to determine the number of replicates required to obtain ad- equate statistical power to detect population trends without some measure of the vari- ability of surveys.

Stahl and Migot (1990)

estimated vari-

ability of spotlight surveys for red foxes in France by conducting a series of 28 trials consisting of three replicate surveys during each trial. They found that coefficients of variation ranged from 0 to 180% and in- creased with estimates of abundance of fox- es based on the same counts. However, es-

tool. An expanded spotlight-survey pro-

gram of kit foxes with some form of

rep-

licated surveys could be a cost-effective

way of monitoring

population

trends

throughout the range of the subspecies.

ACKNOWLEDGMENTS

We thankR. Schlorff, J. Lidberg, G. Gersten- berg, G. Presley, and W. Asserton, III for the survey data, and personnel at the San Luis Obis- po County Engineering Office and D. Williams for data on rainfall. The California Department of Fish and Game and the United States Fish and Wildlife Service provided financial support. C. Vanderbilt-Whitescored physical features

and vegetation on the Elkhorn route, and P. Kel- ly and S. Philips assisted with Fig. 1. B. Cypher

and P. J.

White made helpful comments on the

manuscript.

LITERATURE CITED

BARNES, R. E W., ANDS. C. TAPPER.1985. A method

for counting hares by spotlight. Journal of Zoology

(London), 206:273-276.

EBERHARDT, L.

L.

1987.

Population projections from

simple models. The Journal of Applied Ecology, 24:

103-118.

menting

?n1992.

the

An analysis of procedures for imple-

dynamic

response

method.

Marine

Mammal Science, 8:201-212.

EGOscuE,

fox

in

H. J.

1975.

Population dynamics of the kit

western Utah. Bulletin of the Southern Cali-

fornia Academy of Sciences, 74:122-127.

MERCURE, A.,

K.

RALLS, K.

P. KNOEPFLI, AND R.

K.

WAYNE. 1993. Genetic subdivisions among small

canids: mitochondrial DNA differentiation of swift,

kit, and Arctic foxes. Evolution, 47:313-1328.

MORRELL, S. H.

1972. Life history of the San Joaquin

timates of variability are likely to be species

 

kit fox. California Fish and Game, 58:162-174.

 

M.

NORTON-GRIFFITHS,

1975.

Counting animals. Af-

and site specific, so their results cannot be

 

rican Wildlife Leadership Foundation, Nairobi, Ke-

extended to kit foxes in California. The

 

nya, 110 pp.

   

Carrizo Plain Natural Area seems the logi-

 

O'FARRELL, T. P. 1987.

Kit fox.

 

Pp. 423-431, in Wild

cal place to estimate variability of surveys;

 

furbearer management and conservation in North

 

America.

(M. Novak,

J.

A.

Baker, M. E. Obbard,

two survey routes (Soda Lake and Elkhorn)

 

and B. Malloch, eds.). Ontario Ministry of Natural

run through this area and counts of foxes

Resources, Ottawa, Ontario, 1150 pp.

PROGULSKE, D.

R., ANDD. C. DUERRE.1964. Factors

on the Elkhorn tend to be higher than those

 

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Submitted 5 December 1994. Accepted 30 May 1996.

Associate Editor was James G. Hallett.

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