Estimating Wildlife Populations Based on Incomplete Area Surveys

Author(s): John R. Skalski

Source: Wildlife Society Bulletin (1973-2006), Vol. 22, No. 2 (Summer, 1994), pp. 192-203
Published by: Wiley on behalf of the Wildlife Society
Stable URL: http://www.jstor.org/stable/3783246
Accessed: 19-09-2016 14:08 UTC

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Society Bulletin (1973-2006)

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Wildl. Soc. Bull. 22:192-203, 1994

ESTIMATING WILDLIFE POPULATIONS BASED ON

INCOMPLETE AREA SURVEYS

JOHN R. SKALSKI, Center for Quantitative Science, School of Fisheries, University of Wash-
ington, Seattle, WA 98195

Key words: density estimation, line-transect, mark-recapture, random sampling, sampling with pro
ability proportional to size, stratified random sampling, survey sampling

The application of mark-recapture or line are particularly suited for surveying big game
transect methods has been traditionally re- (White et al. 1989, Bothma et al. 1990, Drum-
stricted to surveying animal abundance at a mer et al. 1990) and bird populations (Gates
single locale. Often this single locale is but a et al. 1968, Ratti et al. 1983, Brennan and Block
small component of a species range or area of 1986). Burnham et al. (1980) provide guide-
interest. However, management decisions of- lines for design and analysis of line transect
ten are of regional scope, making the result of surveys. Precision of line transect surveys de-
a site-specific survey of little practical conse- pends on the number of animals seen and the
quence. My purpose is to present a quantitative spatial variability in abundance among repli-
framework for making sound statistical infer- cate lines within a site. Ramsey and Scott (1979)
ences to the wildlife populations in the region. modified visual survey techniques using vari-
To do this, both the sampling variability of the able circular plots to estimate density of forest
survey techniques and the heterogeneity of avifauna. Alternatively, mark-recapture tech-
wildlife numbers across the landscape must be niques have been adapted to estimate animal
accounted for. My approach combines finite densities (MacLulich 1951, Smith et al. 1971,
sampling theory (see Hansen et al. [1953], Swift and Steinhorst 1976, Otis et al. 1978).
Cochran [1977], Jessen [1978], for a review) Among these mark-recapture methods, the
with density estimation techniques. These nested grid technique of Otis et al. (1978:67-
methods are not new, but rather a compilation 74) based on the methods of MacLulich (1951)
of sampling techniques suitable for wildlife is logistically the easiest to implement. More
surveys. The statistical inferences sought are recently, the trapping web design of Anderson
with regard to total abundance (NT) or mean et al. (1983) has been proposed for estimating
density (15) of animals within a specified geo- the density of small mammals. However, eval-
graphic area. uations of density estimators by Wilson and
In order to make inferences to specific areas, Anderson (1985a,b) suggest continued use of
survey techniques must have well-defined geo- the nested grid design over that of the trapping
graphic boundaries. However, abundance es- web.
timates using mark-recapture techniques (see General inability to conduct mark-recap-
Seber [1982] for review) typically survey wild- ture or line transect studies over the entire
life over an unspecified area around a trap grid region of statistical inference necessitates the
making area assessments impossible. For this area be subsampled. Consequently, a regional
reason, only density estimation techniques will survey transforms into a 2-stage sampling pro-
be considered. Abundance estimates (N) are gram. The first stage is the drawing of a prob-
computed by multiplying the density estimatesabilistic sample of the spatial area. Any sam-
(D) by plot size (a). pling scheme that yields a probabilistic sample
Two general approaches have been devised can be employed (see Cochran [1977] for re-
for estimating animal densities. Line transect view). In this paper, I consider only simple
methods (Eberhardt 1978, Burnham et al. 1980) random sampling (SRS), sampling with prob-

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 AREA SURVEYS OF WILDLIFE * Skalski 193

ability proportional to size (SPPS), and strati- 1 '32

fied sampling schemes. The second stage is the k-6

estimation of animal density at the study plots

selected in the first stage. Hence, overall esti-
mates of animal density or abundance in the
region will be composed of 2 sources of error,
the spatial variation in animal density and the Fig. 1. Schematic of a simple random sample for k
= 6 and K = 32 plots in a region of size A where all
measurement error in estimating animal den- plots are of equal area, a = A/K.
sity at the study plots. Survey designs presented
below quantify these sources of error in the
estimation of sampling variance and confi- jacent (Fig. 1), capture data from animals
dence interval construction. However, classicalcaught at >1 trap grid were first segregated
finite sampling methods assume that the ob- to only those captures that occurred at a par-
servations, animal abundance or density in this ticular site. Segregation of the capture data by
case, are measured without error (Cochran site was necessary to assure stochastic inde-
1977:380-383). Hence, modifications of finite pendence of the site-specific density estimates.
sampling methods are presented that account Results of the surveys are presented in Table 1.
for the error structure of wildlife surveys. Sur-
veys are illustrated using mark-recapture SURVEY DESIGNS
methods. Extensions to line transect and other
Two objectives of regional surveys of anima
survey methods are straightforward.
resources will be considered. One objective is
to estimate total wildlife abundance (NT) de-
EXAMPLE fined as

The basis of the area survey procedures will K K

be illustrated for Great Basin pocket mouse NT- aiDi Ni, (1)
1=1 i=1
(Perognathus parvus) populations using the
nested grid technique of Otis et al. (1978). The where
objective of the survey was to estimate mean
Ni = animal abundance on the ith study plot
density (5) and total pocket mouse abundance
(i = 1, ... IK),
(NT) on a 32-ha region. This contrived example
D, = animal density on the ith study plot (i
represents the results of trapping studies on six
= 1, . . .,K),
1-ha trapping grids established in big sage-
brush (Artemisia tridentata )-bluebunch
wheatgrass (Agropyron spicatum) habitat in Table 1. Density estimates (D) and measurement er-
rors for 6 hypothetical small mammal surveys on 1-ha
southeastern Washington. The 6 sites represent plots randomly selected in a 32-ha region (see Fig. 1)
a simple random sample of the 32 one-ha sites using the nested grid technique of Otis et al. (1978:67-
within the region (Fig. 1). Small mammals at 74).

each site were captured using a 13- x 13-trap Site N = D 1 Var(N, I N.)
grid with Sherman traps (6.5 x 16.5 X 5 cm)
1 63.8 135.7
spaced 7.7 m apart. Trapping was conducted 2 47.1 83.9
for 10 consecutive days and multiple mark- 3 28.7 50.1
4 52.3 94.6
recapture methods were used to survey pop-
5 19.3 43.4
ulations. Density was estimated independently 6 38.4 67.6
at each site using the most appropriate mark-
recapture model. When 2 trap grids were ad-
N = 41.6 V

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194 Wildl. Soc. Bull. 22(2) 1994

K = total number of plots into whichSimple the Random Sampling

region (A) has been subdivided, and
Theory.-When the region being surveyed
a, = area of the ith study plot (i = 1,.is .homogeneous
. , K). with regard to animal densities
or
In the case of the pocket mouse example, K = little is known about the spatial structure of
the
32 and a, = I ha for all i = 1, . . , 32. The population, simple random sampling (SRS)
may
other objective is to estimate mean density (D) be useful in sampling the spatial domain.
in the region, defined as In order to obtain a random sample of the area,
k of K plots are randomly selected with equal
K K
probability (i.e., P = k/K). The plots need not
z
D _1_ _ t=1 NT (2)
aiD,
be of equal size. 2Ni
D K KT 2 An unbiased estimator of total abundance
2; ai 2; ai (equation 1 given that D, are unbiased) can be
i=1 i=1 written as

Although the animal populations to be sur- Kk K k

NT=k aiD -=kNi. (4)
veyed will be quantified using density esti-
mation techniques, the analyses need to be
The estimator of total abund
based on plot-specific estimates of abundance
on multiplying the estimate of mean abun-
if plot sizes vary and unbiased regional esti-
dance by the number of possible study plots
mates are sought. Animal abundance (Ni) at a
in the region (K). An estimator of the variance
study plot will be estimated from an estimate
for total abundance (NT) based on SRS can be
of population density (Di) using the relation-
written (see Appendix A) as
ship
Var(NT) (5)
Ni = aiDi. (3)

It is important to note that the converse of

estimating density from an
=K2[(estimate
K) + Var(Ms of absolute
I Ni){\
abundance (i.e., Di = Ni/ai), known as the
naive density estimator (White et al. 1982:120),
where
results in positively biased estimates of density.
The reason for the bias is that thek effective
area of trapping is dependent on zhome (NiN)2 range
size and is greater than the immediate area s2 i=1
around a trap grid. where~~~~~~~~~r
i(k-i1)
In subsequent discussions, I assume that Di is k

unbiased with variance estimate Var(D' Di),

also unbiased. The abundance estimator (NA) N=i=1 and
will then have a variance of k

Var(Ni I Ni) = aVar (Di I Di) _ Var(Ni - N1)

and estimated variance Var(Ni I N,) = il
k
Var(N, I NJ) = aVar(Di I Di)
Variance estimate (5) is composed of 2 com-
that also is unbiased. ponents: the first term in the brackets char-

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 AREA SURVEYS OF WILDLIFE * Skalski 195

acterizes the sampling error associated with

selecting k of K plots in the region and the tIa/2,k-1 =1/V - 1/0
second term characterizes the average mea- \Var()4
surement error associated with the survey tech-
leading to (1 - a)100% confidence limits
niques. The quantity (1 - k/K) is called the
finite population correction. As the number of 1
plots selected for survey (k) approaches the
total number of plots (K) in the region, the (1/0) ? tI-a/2,kI1 r/b74
first term of equation (5) vanishes and the vari-
ance of NT reduces to the sum of the mea- for 0 = NT or D. The transformation is rec-
surement errors at all plots. ommended when k ' 5.
Mean density (2) is estimated from equation Example.-Surveys of the six 1-ha plots (Ta-
(4) as ble 1) yield an estimate of total Great Basin
k
pocket mouse abundance (4) of

N
AA
= AT=Klk A (6) NT = 32(41.6) = 1,331.2
on the 32-ha site. The estimated variance (5)
with variance estimator
of total Great Basin pocket mouse abundance
Var(NT) is calculated to be
Var(D) = 2 (7)
Var(NT)
The estimates of mean density (D) and total
abundance (NT) can be assumed to be approx-
imately normally distributed for moderate
sampling effort within and between plots. Fol- = 2{_- 6(262.304) +7~7
lowing recommendations of Cochran (1977:
27), a (1 - a)100% confidence interval (CI)
[( 62)X 32
estimate of total abundance can be computed = 38,907.904.
using variance formula (5) as
Using these results, an approximate 95% CI
NT ? t1 a/2,k-1\/ai estimate of total Great Basin pocket mouse
abundance then would be
Similarly, a (1 - a)100% CI estimate of mean
density is computed using variance formula CI(1,331.2 - 2.571V3
(7) as
< NT < 1,331.2 + 2.571V3W8) =
D ? t1_a/2,k-1 )' CI(824.1 < NT < 1,838.3) = 0.95.

where tlIa/2k - is a critical value and follows Similarly, an estimate of mean density of Great
a Student's t-distribution satisfying the rela- Basin pocket mice on the 32-ha site is
tionship P( I t I < t a1/2,kI) 1 - a. For small 1,331.2
sample sizes, the estimators NT and D will tend D - - =41.6/ha
to be right-hand skewed. Both Ricker (1958) 32
and Cormack (1968) recommended using an with estimated variance
inverse transformation (e.g., 1/D) to eliminate
skewness when it occurs, in which case, CI _ : - 38,907.904 - 38.0.
Var(D I D) =-380
estimation should be based on the expression (32)2

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196 Wildl. Soc. Bull. 22(2) 1994

It then follows that an approximate 95% CL
estimate of mean pocket mouse density is
CI(41.6 - 2.571VX86
< D < 41.6 + 2.571V'_i87d) = 0.95
(i.e., Pi = k/K), estimator (8) reverts to equa-
tion (4). When sample size is predetermined
(i.e., k fixed) the Horwitz-Thompson estimator
(8) has the variance (Yates and Grundy 1953)
Var(NT)

CI(25.8 < D < 57.4) = 0.95.

K KN N
The precision of the pocket mouse survey as = 2 (PiPj -Pij) - )
i=l joi Pi Pi
measured by theA half-width of a 95% CI is
?38% of NT or D. In the sample size section, K Var(N, I Ni) (9)
I will investigate how alternative levels of sam-
pling (i.e., k of K) might be expected to influ-
ence this level of precision. where Pi; = probability the ith and jth
are both in the sample of k plots selected
Sampling Proportional to Size among the K total plots. Simple formula
the calculations of Pi and Pi1 are not ava
Theory.-Inspection of variance formula (5)
for all choices of k of K but must be com
for SRS indicates that the precision of regional
on a case-by-case basis and can be extrem
surveys is a function of the variability in abun-
tedious for even slightly large samples.
dance (N1, i =1 . . . K) between plots. When
Typically, when plot sizes vary and stu
plots are of equal size, S2 simply measures the
costs are fixed but dependent on the are
spatial heterogeneity in abundance across the
veyed, the number of plots chosen (k) be
landscape. However, with unequal plot sizes,
a random variable. In this case, the variance
SN is inflated by the variation in plot size. As
is based on the Horwitz-Thompson version of
the variation in sizes of plots increases, S2 can
the variance that is estimated by
be expected to increase. Hence, SRS in the
presence of unequal plot sizes can be impre- k (I - Pi)N?
cise. Basing calculations on the Di does not i=l i

mitigate this problem. k k

An alternative to SRS in the presence of + 2 2 (P -ij PPj)N
unequal plot sizes is to sample with probability i=1 j>i 0ij j

proportional to size (SPPS) of the plots. When

sampling without replacement, total abun- + 2 Var(Ni I Ni) (10)
k 10%- A ~ ~ (1

dance (NT) can be estimated using the Hor- i= Pi

witz-Thompson estimator (Horwitz and In the case where k is in
Thompson 1952) where the variance based on equation (9) will tend
k to be much smaller than that of the Horwitz-
Ni
NT= P Ni (8) Thompson variance (see Appendix B, equation
[18]).
and where Pi = probability the ith plot
Mean is in
density is estimated following SPPS
the sample of k plots selected from among
using the
estimator (6) with variance estimator (7).
K total plots. The Horwitz-Thompson esti- The variance of the Horwitz-Thompson esti-
mator of total abundance is based on escalating
mator (10) typically will be smaller than the
plot specific abundance estimates (N,) by the variance of the SRS estimator (5) when survey
probabilities of selection (Pi). When the prob-plots are of unequal size.
abilities of plot selection are equal for all plots Confidence interval estimators for total

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 AREA SURVEYS OF WILDLIFE * Skalski 197

abundance (NT) or mean density (15) using the

A C
results of SPPS are based on the standard nor-
mal distribution. A (1- a)100% CI estimate N=40 N =100
of total abundance is calculated using variance
a=1 a=2
estimate (10) and

NT ? Z-ia/2\/r), (11) B

or for mean density N=60

D ? Zi-a/2 Va7), (12) a=

where D

P( IZI < ZI a/2) 1 - C. N = 440

For example, for a 95% CI, ZO.975 = 1.96. a =4

Example.-A simple example can illustrate
the relative improvement in precision of SPPS
over SRS when plots are of unequal size. Con-
Fig. 2. Schematic of an hypothetical area survey of
sider the contrived example of a region sub- plots of unequal size (K = 4) with a total abundance
divided into K = 4 plots of unequal size (Fig. of NT = 640.
2) and a sample of size k = 2 is drawn. Esti-
mates of total abundance (NT) computed by
2 24 4 2
SRS and SPPS can be compared (Table 2). To _-2 + --4 = 0.41667,
8 6 8 4
simplify this comparison, measurement error
will be ignored and only the spatial variance
considered. The probabilities of joint selection where P(A I *) = probability th
of any 2 plots (i.e., k = 2 of K = 4) for SPPS selected before any other plot. Th
are computed as follows: P(B I A) denotes the probability th
selected after plot A has already b
P(AB) =P(A I *)P(B IA) + P(B I *)P(A I B) The joint probability of selecting
and B [i.e., P(AB)] is then compute
+ -- = 0.03571, the Law of Total Probability (Felle
87 87 7
P(AC) =P(A I *)P(C IA) + P(C I *)P(A I joint These C) probabilities can be use
1 2 2 1
trate that estimator (8) is unbiased
=I-
8 7- +
8 -
6 = 0.07738,
timate the sampling variance (i.e.,
tion [10]). The inclusion probabilit
P(AD) = P(A I *)P(D I A) + P(D | *)P(A | D
plot (i.e., Pi in equation [8]) are th
1 4 14 1 as follows:
=.- + --- = 0.19643,
8 7 8 4
P(A) = P(AB) + P(AC) + P(AD)
P(BC) =P(B I *)P(C I B) + P(C I *)P(B I C)
= 0.03571 + 0.07738 + 0.19643
1 2 2 1
_ 1 + - = 0.07738, = 0.30952,
8 7 8 6
P(B) = P(AB) + P(BC) + P(BD)
P(BD) = P(B I *)P(D I B) + P(D I *)P(B I D) = 0.03571 + 0.07738 + 0.19643
1 4 4 1 = 0.30952,
=-- + 4 = 0.19643, P(C) = P(AC) + P(BC) + P(CD)
P(CD) = P(C I *)P(D I C) + P(D I *)P(C I D) = 0.07738 + 0.07738 + 0.41667

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198 Wildl. Soc. Bull. 22(2) 1994

the total abundance NT = 640. However, the

standard error for SPPS of SE(NT) = 150.5 is
<40% of the standard error for SRS of
SE(NT) = 412.6 (Table 2). This reduction in
SE(NT) directly translates to a similar reduc-
tion in the width of a CI estimate of regional
abundance.

Stratified Sampling

If the region to be surveyed has the notice-

Stratum I Stratum 2 able differences in habitat and expected den-
K, 20 K, 10 sities of animals, then the precision of the sur-
k=5 k,=3 vey may benefit from stratifying the area before
Am-A, Ama-A, sampling (Fig. 3). Stratification can reduce the
overall variance of a regional survey by elim-
Fig. 3. Schematic of a hypothetical st
sample of study inating
plots the between-strata
in a variability
region in den- o
where all plots sities. Following stratification,
within a stratum the survey pro- ar
= AJ/Kj. ceeds as a series of independent SRS or SPPS
of plots within strata. The resulting estimates
of mean density (D) or total abundance (NT)
= 0.57143, are then sums of the within-strata survey re-
P(D) = P(AD) + P(BD) + P(CD) sults.
= 0.19643 + 0.19643 + 0.41667 I define total abundance for a stratified sur-
= 0.80953, vey design as
L L
also based on the Law of Total Probability.
These inclusion probabilities are subsequently NT=: NTJ= ADj, (13)
w=1 j=h
used in estimating total abundance (8). Both
SRS and SPPS provide unbiased estimates of where

Table 2. Characterization of all possible samples of size k = 2 of K = 4

Fig. 2.

Simple random sampling Sampling with probability proportional to size

Sample drawn Probability of draw N Probability of draw 9Ti

A B 0.1667 200 0.0357 323.1
A C 0.1667 280 0.0774 304.2
A D 0.1667 960 0.1964 672.8
B C 0.1667 320 0.0774 368.8
C D 0.1667 1000 0.1964 737.4
B D 0.1667 1080 0.4167 718.5
E(NT)a = 640 E(NT) = 640

SE(NT) = 412.6 SE(NT) = 150.5

aExpected value of total abundance estimate E(N9) = z P(N) NT.

' SE(N ) = V) -N1)2.

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 AREA SURVEYS OF WILDLIFE * Skalski 199

L = number of strata in the region,

of 2 specifications, the maximum size of the
relative error in estimation that will be toler-
NT} = total abundance in the jth stratum
(j= 1,...,L), ated (e), and the desired probability (1 - a) a
sample survey will attain at most that level of
Aj = area of the jth stratum
error. In other words, equation (14) specifies
(j = 1,. . ., L), and
that the relative error of the estimate of mean
DJ = mean density in the jth stratumdensity (D) or total abundance (NT) should be
(j = 1, .. . ,L).
< E, (1 - a)100% of the time. For example,
It then follows that an unbiased estimator of one might seek a precision of ?20% of NT,
total regional abundance for a stratified sur- 90% of the time (i.e., e = 0.20, a = 0.10).
vey is
L
Simple Random Sampling
NT = N T
j=1 Assuming the survey estimates (i.e., equa-
tions [4] or [6]) are normally distributed, the
with estimated variance
required number of plots to sample (k) in an
L
SRS of the region to satisfy precision expression
Var(NT) = Var(NT)4 (14) is
j=1

The estimator of total abundance using strat-

K(SNj + Var(N1 I N1))
ified sampling is simply the sum of the within-
k = _& 2 , (15)
strata estimates of total abundance (NT) Be-
K(N) +S2
cause sampling is conducted independently
t 1-a12,k-1 N
between strata, the overall variance for a strat-
ified survey is the sum of the within-stratumwhere S2 = variance in animal abun-
variance estimates. Mean density (D) based on dance among the K plots,
stratified sampling is estimated analogous to
K
equation (6) with associated variance estimator
(NI N2
(7). i=1

Confidence interval estimates should be (K-1)

based on a standard normal distribution and
equations (11) or (12) for total abundance and Equation (15) must be iteratively solved w
mean density, respectively. successive entries into a t-table to find va
of t that satisfy the relationship where P(
< tl-a/2,k-) = 1 - a. It is important to
SAMPLE SIZE CALCULATIONS
that for some arbitrarily small (, a sample
of k = K will be insufficient to yield the de
The sampling effort required for a regional
survey will depend on the precision sought. level of precision. In such case, the average
Precision of a regional survey can be defined asmeasurement error of the survey techniques
[i.e., Var(Ni I N1)] must be reduced by increas-
ing the within-plot sampling effort.
P(D < <E)
Example.-The Great Basin pocket mouse
study now can be reexamined to determine
| NT NT < 1-(a. (14) the level of spatial sampling needed to achieve
a prescribed level of precision. Suppose a level
Precision as defined by equation (14) consists of precision defined by

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200 Wildl. Soc. Bull. 22(2) 1994

k; = number of plots surveyed in the jth

(I D i <0.20)
stratum (j = 1, . . ., L),
S2= variance in abundance among plots
= NT- NT < 0.20) = 0.95
(i.e., S2f) in the jth stratum

is sought. Before a sample size can be calcu-

lated, the variance components from the pre- and
vious study must be estimated (see Appendix
Var(N, I)j
C, equation [19]). The spatial variance in abun-
-average measurement error
dance (S2 ) is estimated to be
for surveys in the jth stratum

j= 262.304 - 79.217 = 183.087 (j= 1,...,L).

for Var(N, I N,) = 79.217 (Table 1). Assuming

similar survey effort at study plots in the fu- CONCLUSION

ture, the number of plots that must be surveyed

Classical methods of finite sampling (i.e., S
is calculated by the equation
SPPS) can be used to incorporate the result
32(183.087 + 79.217) wildlife surveys for the purpose of estima
I =
2
mean density (6) or total abundance (4). Den
32 (0.20)41.j8 + 183.087 sity estimation techniques must be used to
tO.975,k-1 /
vert area estimates of abundance at study p
A value of k = 13 (tO.975,12 = 2.179) satisfies theto regional estimates. Although the point e
equation, indicating the need to sample 13 of timates of mean density (D) or total abund
the 32 one-ha plots to be within ? 20% of D (NT) lead directly from sample survey met
or NT 95% of the time. ods, variance estimates must account for t
measurement error of the wildlife survey t
Other Sampling Designs niques. Variances of regional population esti-
mates are composed of 2 error sources, the
For the SPPS survey design presented, sam-
spatial variance in density across the landscape
ple size (k) is a random variable constrained and the average measurement error of the
by total fixed study costs (i.e., variance formula
wildlife surveys at the study plots. Should the
[11]). As such, sample size calculations are in-
entire region be surveyed, the resulting re-
appropriate for SPPS. For the case of stratified
gional estimates would still possess the mea-
random sampling (STRS) sample size refers to
surement error of the line-transect or mark-
the number of study plots/stratum. For pre-
recapture technique.
cision defined by expression (14), the required
Often project logisitics and study costs dic-
sample sizes for STRS are the values of k,(j tate that inferences to wildlife resources in a
1, . . . , L) that satisfy the equation
region be based on surveying a single "rep-
/ k.\ resentative plot." Nevertheless, researchers
need to appreciate that the measurement error
E2N2 L K K(1 - -) S + Var(N2 I
associated with that survey does not convey
1-_a/2-
Z2
j=1k the true degree of uncertainty ascribed to re-
where gional inferences. The measurement error at
the representative plot is only 1 of 2 error
Kj = number of plots in the jth stratum
sources comprising the variance of a regional
(i = 1,. . . I,), survey. The variance formula presented herein

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 AREA SURVEYS OF WILDLIFE * Skalski 201

provides the basis for weighting the relative SPPS typically will result in a smaller variance.
contributions of the spatial variation (i.e., S2N) However, when plots are of equal size, SPPS
and sampling variability of wildlife census and SRS are equivalent. Stratified sampling
techniques (i.e., Var(D, I Di)). Through proper
designs also are suggested to help reduce the
variance of area survey results. Appendices A
design and analysis of regional surveys, reliable
and B show variance formulae for finite sam-
inferences to wildlife resources are possible with
ascribed error rates. Furthermore, the human pling methods must be modified to account for
resources necessary to conduct useful wildlife error in animal densities measurements. Unless
surveys for management purposes can be ob- adjustments in the variance estimates are in-
jectively assessed using sample size calcula- corporated, classical finite sampling proce-
tions. dures will underestimate the actual variance
The formulae presented here appear com- in area surveys of wildlife. The consequence
plicated at first inspection. However, computer would be a false sense of precision and the
software such as CAPTURE (White et al. 1982) failure to appreciate the uncertainty of re-
is well suited to density estimation using mark-source management decisions.
recapture data. Similarly, DISTANCE (Laake
et al. 1993) conveniently estimates density us-
ing line-transect data. The remainder of the LITERATURE CITED
formulae use sample means and sample vari-
ANDERSON, D. R., K. P. BURNHAM, G. C. WHITE, A
ances from among replicate sites to estimate D. L. OTIS. 1983. Density estimation of small
regional totals and associated variance esti- mammal populations using a trapping web and
mates. Virtually any scientific hand-held cal- distance sampling methods. Ecology 64:674-680.
BOTHMA, J. D., M. J. S. PEEL, S. PETTIT, AND D. GROSSMAN.
culator can be used to calculate these summary 1990. Evaluating the accuracy of some commonly
values and tally them to produce the desired used game-counting methods. South Afr. J. Wildl.
Res. 20:26-32.
regional statistics.
BRENNAN, L. A., AND W. M. BLOCK. 1986. Line tran-
sect estimates of mountain quail density. J. Wildl.
SUMMARY Manage. 50:373-377.
BURNHAM, K. P., D. R. ANDERSON, AND J. L. LAAKE.

Area surveys of animal resources are typi- 1980. Estimation of density from line transect
sampling of biological populations. Wildl. Mono
cally based on a 2-stage nested sampling de- 72.202pp.
sign. The first stage is the selection of a subset COCHRAN, W. G. 1977. Sampling techniques. John
of possible study plots within the region of Wiley and Sons, New York, N.Y. 428pp.
CORMACK, R. M. 1968. The statistics of capture-re-
inference. The second stage then consists of capture methods. Oceanographic Mar. Biol. Annu.
surveying animal densities at the plots selected Rev. 6:455-506.
in the first stage. Consequently, both sampling DRUMMER, T. D., A. R. DEGANGE, L. L. PANK, AND L.
L. McDONALD. 1990. Adjusting for group size
error associated with subsampling the region influence in line transect sampling. J. Wildl. Man-
and measurement error associated with the age. 54:511-514.
survey method contribute to the variance of EBERHARDT, L. L. 1978. Transect methods for pop-
ulation studies. J. Wildl. Manage. 42:1-31.
estimates of total abundance and mean den- FELLER, W. 1968. An introduction to probability the-
sity. Sample size calculations need to take into ory and its applications. Vol. I. John Wiley and
account both of these error sources in the de- Sons, New York, N.Y. 509pp.
GATES, C. E., W. H. MARSHALL, AND D. P. OLSEN. 1968.
sign of area surveys. Line transect method of estimating grouse popu-
Use of sampling with probability propor- lation densities. Biometrics 24:135-145.
tional to size (SPPS) is recommended over sim- HANSEN, M. H., W. N. HURWITZ, AND W. G. MADOW.
1953. Sample survey methods and theory. Vol. I.
ple random sampling (SRS) when study plots John Wiley and Sons, New York, N.Y. 638pp.
are of unequal size. When plots vary in size, HORWITZ, D. G., AND D. J. THOMPSON. 1952. A gen-

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All use subject to http://about.jstor.org/terms
202 Wildl. Soc. Bull. 22(2) 1994

eralization of sampling without replacement from APPENDIX A

a finite universe. J. Am. Stat. Assoc. 47:663-685.
JESSEN, R. J. 1978. Statistical survey techniques. John The standard estimate of the variance of a sample
Wiley and Sons, New York, N.Y. 520pp. mean (x) from an SRS (Cochran 1977:26) is
LAAKE, J. L., S. T. BUCKLAND, D. R. ANDERSON, AND K.
P. BURNHAM. 1993. Distance user's guide, Ver-
sion 2.0. Colo. Coop. Fish. and Wildl. Res. Unit, (K) xi
Colo. State Univ., Fort Collins. 72pp. Var(f) = k (16)
MAcLULICH, D. A. 1951. A new technique of animal
census, with examples. J. Mammal. 32:318-328. where
OTIS, D. L., K. P. BURNHAM, G. C. WHITE, AND D. R.
k
ANDERSON. 1978. Statistical inference for capture
data from closed populations. Wildl. Monogr. 62. Dxi
135pp.
k
RAMSEY, F. L., AND J. M. SCOTT. 1979. Estimating
population densities from variable circular plot and
surveys. Pages 151-181 in R. M. Cormack, G. P.
Patil, and D. S. Robson, eds. Sampling biological
populations. Int. Coop. Publ. House, Fairland, Md.
z (Xi -x)2
2 l
RATTI, J. T., L. M. SMITH, J. W. HUPP, AND J. L. LAAKE.
1983. Line transect estimates of density and the
SX (k- 1)
winter mortality of gray partridge. J. Wildl. Man- However, when the xi are me
age. 47:1,088-1,096. that the expected value is
RICKER, W. E. 1958. Handbook of computations for
biological statistics of fish populations. Bull. Fish.
E(ix) = xi
Board Can. 119:1-300. with variance Var(i, I xi), variance formula (16) is no
SEBER, G. A. F. 1982. The estimation of animal abun- longer unbiased. The expected value of the variance
dance and related parameters. Charles Griffin, estimate (16) when the xi are measured with error (i.e.,
London, U.K. 654pp. ii) is
SMITH, M. H., R. BLESSING, J. G. CHELTON, J. B. GENTRY,
F. B. GOLLEY, AND J. T. MCGINNIS. 1971. De-
termining the density of small mammal popula- (1K- + Var(i, I xi))
tions using a grid and assessment lines. Acta Theri-
ologica 16:105-125. E[Var(f)] =k
Swwr, D. M., AND R. K. STEINHORST. 1976. A tech-
nique for estimating small mammal population although the true variance of the sample
densities using grid and assessment lines. Acta
Theriologica 1:471-480.
WHITE, G. C., D. R. ANDERSON, K. P. BURNHAM, AND
D. L. OTIS. 1982. Capture-recapture and re- Var(f) = k + Var(k[ x,)
moval methods for sampling closed populations.
LA-8787-NERP. Los Alamos Natl. Lab., Los Al- where
amos, N.M. 235pp. K

, R. M. BARTMANN, L. H. CARPENTER, AND R. A.

z (xi - X)
GARROTT. 1989. Evaluation of aerial line tran-
S2 -i=
sects for estimating mule deer densities. J. Wildl.
x, (K-1)
Manage. 53:625-635.
WILSON, K. R., AND D. R. ANDERSON. 1985a. Evalu-and
ation of a density estimator based on a trapping K

web and distance sampling theory. Ecology 66: _ _ Var(i1 I xi)

1,185-1,194.
Var(ix I xi) = K
, AND . 1985b. Evaluation of a nested K
grid approach for estimating density. J. Wildl.
Manage. 49:675-678. under the case of measurement error. The bias in the
variance estimator (16) under measurement error is
YATES, F., AND P. M. GRUNDY. 1953. Selection without
replacement from within-strata with probability
proportional to size. J. R. Stat. Soc. (B) 15:253- Var(i, I x,) (17)
261. K

Received 16 September 1992. Hence, an unbiased variance estimator for a sample

Accepted 29 November 1993. mean under SRS with measurement error is equation
Associate Editor: Flather. (16) corrected by an estimate of the bias (17) leading to

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All use subject to http://about.jstor.org/terms
 AREA SURVEYS OF WILDLIFE * Skalski 203

+ 2 K K (P i
f-l j=l j ij
k
Var(i) = - ______ K
_ ,>i

K (1 - P,)Var(i, I x1)
where
k
i=1 Pi
_ Var(2I x,) resulting in a negative bias of
Var(i ) = i=X k K

bias = -: Var(i1 I xi).

such that
Consequently, an unbiased variance estimator can be
E[Var(xi I xi)] Var( I xi),
written as
and
k (PI - pk)2 k k (p j Pi )ij
k

i(j )2
Var(X) = + 2 2; "pip-PjP)ii
i=1 i J=1 j j t;
j>1
Sg2 = -
x~i (k-1 k Var(ix I xi)

i=1 P
APPENDIX B because

The standard Horwitz-Thompson estimator (Coch- E ;Var(i, I xi)) = Var(i, I xe).

ran 1977:261) of a total is
k
(= Pi ) =

i=l i
APPENDIX C
with unbiased variance estimator
Preliminary survey data can be used to estimate the
k I- P.) variance components used in equation (15) for sampl
Var(Xi) p xi size calculations. From a preliminary sample of n plots
i=l
the average sampling error can be estimated as
k kc (P i - PjP,)x'xj
+ 2; 2; (8P P )
i=l j=l i ij
j>i _ Var(N, I Nj)
for k random. When the observations are measured Var(N, I N,) = i='
n
with error [E(i I xi], the estimator and the spatial variance can be estimated from the
k Xi
relationship
i=1 i

has the variance S2- S2 Var(N, I N)j (19)

K (1 K
where s2 is calculated as
Var(X x1 +P
) 2
= (1's,
2xi2-
i Pl i pip fp ff i i1 1=1 i n
j>1
(Ni-N)2
K Var(i, l x,) 2 (=n-)

il Pi
However, the expected value of (18) when the xi are
measured with error is
K (1 _Fp)
E[Var(X)]= p2 xi2
i=

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