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 WestKlamath,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
threatened by the state of California and as
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 used to census
many species of wildlife (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-
Journal of Mammalogy, 78(1):65-73, 1997
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-
65
66 JOURNALOF MAMMALOGY
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
were impassable.
Two observers, one sitting on each side of a
vehicle that enabled their eyes to be m nant vegetation was scored as either: 1) peren-
above the level of the road, conducted each-1.5sur-
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
Vol. 78, No. 1
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,
poorest visibility) to sparse, short, or no vege-
tation (class 5, highest visibility). The predomi-
nial shrubs (e.g., Ephedra, Allenrolfea, or Atri-
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
February1997 RALLS AND EBERHARDT-SURVEYS OF KIT FOXES 67

40 80 120
0...

Kilometers

0 25 50 75

S -
- ..Miles

\ N----

> F

Tf

FeOrtigalita
1

, Blackw CornerI
's
'..

Plain
Elkhorn

. .
I

N B s o n
.-, ...
.

Belridge-McKittrick
...,

Soda Lake
. I
SSoda Lake

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 JOURNAL
OFMAMMALOGY
rainfallseason (K. Allred, in litt.), we compared
summercounts of foxes and concurrentcounts
of prey to total precipitationduringthe previous
rainfallseason (e.g., June 1990 counts with rain-
fall from 1 July 1988 to 30 June 1989). We com-
puted Pearson product-momentcorrelationco-
efficients among these variablesand used Bon-
ferroni-adjustedprobabilitiesto reflectthe num-
ber of correlations being tested (Wilkinson,
1990).
To model a possible annual cycle of abun-
dance on the Elkhornroute, numbersof 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-
squaresfit using the model N, = NJune St for the
months of June to May (months0-11), follow-
ing the approachof Eberhardt(1987). Here, Nt
denotes the populationat montht (the following
September,December,or March)and s denotes
a rangeof changedue to deathsand the dispersal
of young foxes from the area.
We examined trends in populationsof foxes
over time for each routeby summingup the sea-
sonal counts to get a yearly total. Based on pop-
ulation simulations conducted by Eberhardt
(1992), we transformedthe counts by natural
logarithmsand tested for linearityand curvilin-
earity. A log-linear relationshipwould be ex-
pected if the populationwere increasingor de-
creasingat a relativelyconstantrate.The test for
curvilinearitycompareddeviationsfrom a fitted
second-degreepolynomialto those from the lin-
ear regression(Snedecorand Cochran,1967).
To examine effectiveness of samples of
counts of the size typically obtainedin surveys
to detect a populationtrend, we constructeda
simple stochasticmodel based on the annualcy-
cle. We simulateddatafor threeinitialcount lev-
els (high, intermediate,and low) for 5, 7, and
10 years, producing 1,000 simulationsfor each
case. The highest-level simulated counts were
obtainedon the ElkhornPlain routeand the low-
est were those obtainedon the Panocheand Or-
tigalitaroutes.The model simulateda 10%/year
decline in the population.We applied the log-
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
Vol.78, No. 1
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 terrain (r2 = 0.24, P <
0.001, n = 73, Fig. 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
February1997 RALLS AND EBERHARDT-SURVEYS OF KIT FOXES 69

80 0 v 0

w 60

S40

20

V q a -
00 I 0.
" ..
0 12 24 0'"II."1*'
36 48 60 72 84 96 108 120 132 144
MONTH
80N
Go
70
VU-

60-
50 .
U-so4
w 40
w 30
S
2 .•II
z20

-
00

144 156 168 180 192 204 216 228 240 252 264 276 288
MONTH
FIG.2.-Number of foxes seen on the ElkhornPlain routeduringtwo successive 12-yearperiods.
The initial point for each plot is a summercount. Curves are fitted by least squaresto representan
annualcycle 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 all cases when only 5 years of
data were collected, even at the highest
70 JOURNAL
OFMAMMALOGY Vol.78, No. 1

3
5•

40
(),
-J
j

20u
>. 310
z z
I-
w
M i m
z
0 0

0 10 20 30 40 50 60 70 80

FROMSTARTOF ROUTE
HALF-MILES
FIG.3.-Mean numberof foxes sighted/0.8-kminterval (line with points; left-handscale) versus
category of viewing distancebased on terrain(line withoutpoints;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.
-
Log-linear
Route n relationship Curvilinearity
Bakersfield 17 0.26 0.34
Belridge-McKittrick 20 5.64 (P < 0.05) 0.03
Blackwells Corner 8 25.10 (P < 0.005) 1.48
Elkhorn 23 2.60 13.37 (P < 0.005)
Ortigalita 11 0.74 0.04
Panoche 13 2.50 (P < 0.10) 0.08
Taft-Fellows 21 0.09 5.23 (P < 0.05)
February1997 RALLS AND EBERHARDT-SURVEYS OF KIT FOXES 71

1.0
LEVEL
HIGHESTPOPULATION

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
OF YEARSDATA COLLECTED
NUMBER
FIG.4.-Results of statisticaltests on simulatedsurvey 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 generatedfor 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
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-
timates of variability are likely to be species
and site specific, so their results cannot be
extended to kit foxes in California. The
Carrizo Plain Natural Area seems the logi-
cal place to estimate variability of surveys;
two survey routes (Soda Lake and Elkhorn)
run through this area and counts of foxes
on the Elkhorn tend to be higher than those
on other routes.
The current survey program does not
cover the entire range of the San Joaquin
kit fox. The addition of more routes, partic-
ularly in the northeastern San Joaquin Val-
ley and the Salinas Valley, would increase
its effectiveness as a population-monitoring
Vol. 78, No. I
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 personnelat the San Luis Obis-
po County EngineeringOffice and D. Williams
for data on rainfall.The CaliforniaDepartment
of Fish and Game and the United States Fish
and Wildlife Service providedfinancialsupport.
C. Vanderbilt-Whitescored physical features
and vegetationon the Elkhornroute, and P. Kel-
ly and S. Philips assistedwith 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.
An analysis of procedures for imple-
?n1992.
menting the dynamic response method. Marine
Mammal Science, 8:201-212.
EGOscuE,H. J. 1975. Population dynamics of the kit
fox in 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
kit fox. California Fish and Game, 58:162-174.
NORTON-GRIFFITHS, M. 1975. Counting animals. Af-
rican Wildlife Leadership Foundation, Nairobi, Ke-
nya, 110 pp.
O'FARRELL, T. P. 1987. Kit fox. Pp. 423-431, in Wild
furbearer management and conservation in North
America. (M. Novak, J. A. Baker, M. E. Obbard,
and B. Malloch, eds.). Ontario Ministry of Natural
Resources, Ottawa, Ontario, 1150 pp.
PROGULSKE, D. R., ANDD. C. DUERRE.1964. Factors
influencing spotlighting counts of deer. The Journal
of Wildlife Management, 28:27-34.
RALLS,K., ANDP. J. WHITE.1995. Predation by larger
canids on San Joaquin kit fox. Journal of Mammal-
ogy, 276:723-729.
SNEDECOR, G. W., ANDW. G. COCHRAN.1967. Statis-
tical methods. Sixth ed. Iowa State University Press,
Ames, 593 pp.
STAHL, P. 1990. Suivi de l'abondance d'une popula-
tion de renards (Vulpes vulpes) par comptages noc-
February 1997 RALLS AND EBERHARDT-SURVEYS
turnes: evaluation de la m6thode. Gibier Faune Sau-
vage, 7:293-309.
STAHL, P., AND P. MIGOT. 1990. Variabilit6 et sensi-
bilit6 d'un indice d'abondance obtenu par comptages
nocturnes chez le renard (Vulpes vulpes). Gibier
Faune Sauvage, 7:311-323.
WEBER, J., S. AUBRY, N. LACHAT,J. MEIA, C. MERMOD,
ANDA. PARATTE.1991. Fluctuations and behavior
of foxes determined by nightlighting: preliminary
results. Acta Theriologica, 36:285-291.
WHITE, P. J., AND K. RALLS. 1993. Reproduction and
spacing patterns of kit foxes relative to changing
prey availability. The Journal of Wildlife Manage-
ment, 57:861-867.
WHITE, P. J., K. RALLS,AND R. A. GARROTT.1994.
Coyote-kit fox spatial interactions based on radio-
OF KIT FOXES 73
telemetry. Canadian Journal of Zoology, 72:1831-
1836.
WHITE,P. J., C. A. VANDERBILT WHITE,and K. RALLS.
1996. Functional and numerical responses of kit
foxes to a short-term decline in mammalian prey.
Journal of Mammalogy, 77:370-376.
WILLIAMS, D. E 1992. Introduction. Pp. ix-xi, in En-
dangered and threatened species of the San Joaquin
Valley, California (D. E Williams, S. Byrne, and T
A. Rado, eds.). California Energy Commission, Sac-
ramento, California, 588 pp.
L. 1990. SYSTAT: the system for statis-
WILKINSON,
tics. SYSTAT, Inc., Evanston, Illinois, 667 pp.
Submitted 5 December 1994. Accepted 30 May 1996.
Associate Editor was James G. Hallett.