Zigzag Survey Designs in

Line Transect Sampling

Samantha STRINDBERG and Stephen T. BUCKLAND
Zigzag survey lines are frequently used in shipboard and aerial line transect surveys
of animal populations. Analysis proceeds assuming that coverage probability through the
survey region is uniform. We show that the two types of zigzag design that are in wide use
do not generally have this property, and explore the degree of bias in abundance estimates
that can be anticipated. We construct a zigzag design for convex survey regions that has
even coverage probability with respect to distance along a design axis. We also provide
Horvitz-Thompson estimators that allow coverage probability to vary by location through
the survey region.

Key Words: Automated design algorithms; Coverage probability; Sighting surveys; Survey
design; Systematic designs; Zigzag designs.

1. INTRODUCTION
Shipboard and aerial line transect surveys are widely used for estimating the abundance
of marine mammals (e.g., Borchers et al. 1998; Buckland et al. 2001, pp. 288–290) and
seabirds (e.g., Clarke et al. 2003; Buckland et al. 2001, p. 290), and to a lesser extent,
fish (e.g., Foote and Stefansson 1993; O’Connell and Carlile 1993; Chen and Cowling
2001). Aircraft are also widely used for surveys of terrestrial mammals (White, Bartmann,
Carpenter, and Garrott 1989; Pojar, Dowden, and Gill 1995; van Hensbergen, Berry, and
Juritz 1996; Trenkel, Buckland, McLean, and Elston 1997; Pople et al. 1998; Wiig and
Derocher 1999). A design consists of a series of survey lines, along which the ship or
aircraft travels, and observers record animals sighted, together with the distances of those
observations from the survey line. A full description of the approach, including detailed
discussion of survey design and analysis, was given by Buckland et al. (2001).
Frequently in such surveys, the survey design comprises a series of parallel lines, either

Samantha Strindberg is Associate Conservation Scientist and Biometrician, Living Landscapes Program, Inter-
national Conservation Programs, Wildlife Conservation Society, 2300 Southern Boulevard, Bronx, NY 10460
(E-mail: sstrindberg@wcs.org). Stephen T. Buckland is Professor of Statistics and Director, Centre for Research
into Ecological and Environmental Modelling, The Observatory, Buchanan Gardens, St. Andrews KY16 9LZ,
Scotland (E-mail: steve@mcs.st-and.ac.uk).

2004
c American Statistical Association and the International Biometric Society
Journal of Agricultural, Biological, and Environmental Statistics, Volume 9, Number 4, Pages 443–461
DOI: 10.1198/108571104X15601

443
444 S. STRINDBERG AND S. T. BUCKLAND

Figure 1. A line transect survey comprises a series of lines. Illustrated here within a simple rectangular survey
region are examples of three commonly used designs: randomly spaced parallel lines (top); systematically spaced
parallel lines (middle); and a continuous zigzag sampler.

randomly spaced or systematically spaced with a random start (Figure 1). The disadvantage
of such a design is that the ship or aircraft must travel from one line to the next “off-
effort,” that is, without searching for the animals of interest. If searching is carried out on
these legs, the random design is compromised, as there tends to be greater effort along
the boundaries of the survey region where animal density may be atypical, so that design-
based estimation of abundance is biased. If searching is not carried out on these legs,
resources are not used efficiently, especially in the case of shipboard surveys, for which
ship time is expensive. Successive survey lines may be tens or even hundreds of kilometers
apart, so that the loss of search effort is unacceptable on cost grounds. As a consequence,
continuous zigzag designs are often preferred. These have the property that survey effort is
continuous. Typically, a design comprises a single zigzag survey line, although for analysis
purposes, successive legs of the zigzag are often assumed to be independent, just as the
 ZIGZAG SURVEY DESIGNS IN LINE TRANSECT SAMPLING 445

separate lines of a systematic parallel-line design are usually assumed to be independent

(Figure 1). Thus, variance estimation proceeds as if the legs of the zigzag are random lines,
independently placed within the survey region (Buckland et al. 2001, p. 238). Although
variance estimation may be biased, systematic designs are efficient because systematic
zigzags and systematic parallel lines are more evenly distributed through the survey region
than are randomly spaced parallel lines (Figure 1). In a shipboard survey of a large region,
lines tend to be long (perhaps several hundred kilometers) and resources typically allow few
lines per stratum. Thus, design efficiency is an important advantage of systematic sampling.
(If lines are shorter, then daylight hours are wasted in traveling from one line to the next for
parallel-line designs.)
We refer to the surveyed strip comprising the survey line together with its associated
width 2w (w either side of the line) as a sampler. We define coverage probability π(u)
at a point (x, y), denoted by u ≡ u(x, y), to be the probability that the sampler covers
the point u, given a rule for randomizing the location of the sampler. Generally, zigzag
survey designs do not provide uniform coverage probability throughout the survey region,
yet the standard analysis methods assume that they do. This can lead to substantial bias in
abundance estimates if animal density varies appreciably through the study area (Strindberg
2001).
We address how abundance estimation should be modified to take account of uneven
coverage probability, using a Horvitz-Thompson estimator and consider the properties of
commonly used zigzag designs, demonstrating their potential for bias in estimates of animal
abundance. We also construct a zigzag design within a convex-shaped survey region (termed
“convex survey region” for brevity) that has even coverage probability with respect to
distance along a design axis, and show how to modify the method for nonconvex survey
regions.
Annual surveys of Antarctic minke whales are conducted under the auspices of the
International Whaling Commission. Due to the cost of operating ships in the Antarctic, and
because of the size of the area, practical considerations dictate that zigzag designs be used.
We illustrate the design algorithms of this article, using these surveys. The example also
shows the importance of using an automated design algorithm that requires no subjective
decisions; the ice-edge, which delineates the southern boundary of the survey region, can
move by as much as 100 kilometers per day, so that the design must be developed as the
survey progresses.

2. HORVITZ-THOMPSON ESTIMATOR
If a survey design that gives uneven coverage probability is used, then a Horvitz-
Thompson estimator (Horvitz and Thompson 1952) can be applied to obtain an unbiased
estimate of abundance. Cooke (1985) was the first to suggest this approach for line transect
sampling. When coverage probability is uneven, it is usually unknown. However, given
automated algorithms for generating zigzag and other samplers, we can readily estimate
coverage probability at any chosen point by simulation. To do this, we repeatedly generate
446 S. STRINDBERG AND S. T. BUCKLAND

realizations of a given design, and observe the proportion of realizations for which the
generated sampler covers the chosen point. For estimating animal abundance, we must obtain
this proportion for all points at which animals were detected. These coverage probabilities
are estimated, whereas the Horvitz-Thompson estimator assumes that they are known. This
presents no practical difficulty, however, as we can estimate them to any desired precision,
simply by increasing the number of realizations of the sampler that we generate. In practice,
except for strip transect surveys during which all objects within w of the line are counted,
the effective half-width of search µ must also be estimated, and its precision is determined
by the survey data, so that variance estimation should take this into account (this is discussed
in the following).
In conventional line transect sampling of a population when animals do not occur in
groups or “clusters,” animal abundance N is estimated by

An
N̂ = , (2.1)
2µ̂L
where A is the surface area of the survey region, n is the number of animals detected, µ̂
is the estimated effective strip half-width, and L is the total length of line (Buckland et al.
2001). This may be obtained as a Horvitz-Thompson-like estimator as follows.
If an animal i is located outside the sampled strips, it cannot be sampled. If it is inside
the strip, and we do not condition on its distance from the line, the probability that it is
detected is estimated by µ̂/w, where w is the half-width of the strip (Buckland et al. 2001,
pp. 37–38). Let πc denote the mean coverage probability, where πc ≈ 2w×L/A for small w.
(The exact value includes an additional term in w2 that depends on the angle of the sampler
to the design axis and on the shape of the survey region. Modified designs for which the
sampler does not overlap the region boundary that satisfy this result exactly are possible,
but abundance estimates are then biased if animal density is not typical near the boundary.)
Assuming even coverage probability, the probability that a sampled strip includes a given
animal is πc . Hence the estimated inclusion probability p̂i of animal i is the product of these
two quantities, and
n

n

n


1 1 1 An
N̂ = = = = . (2.2)
p̂i (µ̂/w)πc (µ̂/w)(2wL/A) 2µ̂L
i=1 i=1 i=1

Note that this is not a strict Horvitz-Thompson estimator since the inclusion probability
must be estimated when µ is unknown (Borchers et al. 1998).
We now allow the probability that the sampled strip includes a given animal to vary by
location. Denote the coverage probability at the location of detected animal i by πi . Then
n

n

n
1 1 w
1
N̂ = = = . (2.3)
p̂i (µ̂/w)πi µ̂ πi
i=1 i=1 i=1

If πi = πc for all i (even coverage probability), this gives the same result as before. This
equation may be further generalized by allowing animals within the strip to have their own
 ZIGZAG SURVEY DESIGNS IN LINE TRANSECT SAMPLING 447

individual effective strip half-widths µi :

n

n

n


1 1 1
N̂ = = =w . (2.4)
p̂i (µ̂i /w)πi µ̂i πi
i=1 i=1 i=1

The µi may be modeled as a function of their distance from the line and of other covariates
(Marques and Buckland 2003).
If animals occur in clusters, and the ith detected cluster is of size si , then the above
formulas yield estimates of cluster abundance, and animal abundance may be obtained by
n n
replacing i=1 1/p̂i with i=1 si /p̂i in each of the above expressions.
Variances for the various estimators may be estimated by bootstrapping transects
(Borchers et al. 1998; Marques and Buckland 2003). For zigzag survey designs these are
usually taken to be the separate legs of the zigzag sampler. This tends to be more robust than
the analytic variance for the Horvitz-Thompson estimator, because it does not require the
assumption that different detections on the same transect are independent, and uncertainty
due to estimating µ is readily incorporated into the bootstrap variance. The simple percentile
method is generally adequate for estimating confidence limits (Buckland et al. 2001, p. 82).
If analytic variances are preferred, Huggins (1989) showed how to include a component
into the variance of the Horvitz-Thompson estimator when the inclusion probabilities are
estimated. However, in the context of zigzag samplers, as for standard systematic sampling
methods, some joint inclusion probabilities are zero if a single sampler is used. It is generally
impractical to conduct surveys using several zigzag samplers, randomly located over the
survey region, so that the strategy of bootstrapping sampler legs has strong practical appeal.

3. PROPERTIES OF ZIGZAG SURVEY DESIGNS

In this and the section that follows, we assume that the survey region is convex. Methods
for nonconvex regions are considered in Section 5.

3.1 COVERAGE PROBABILITY

A zigzag survey line is defined with respect to a design axis G that, without loss of
generality, starts at the origin and runs along the x-axis of a Cartesian coordinate system.
Let the angle of the line to the design axis G be either θ(x) or 180◦ − θ(x) at distance x,
where θ(x) may vary in the range [0, 90◦ ). Conventional zigzag designs keep θ(x) constant
within a leg, but we do not impose that constraint here.
To use standard distance sampling methodology, we would like a zigzag sampler to
provide even coverage probability, so that coverage probability at point u is π(u) = πc , ∀u ∈
R. To date, survey designs based on zigzag samplers have been ad hoc, with no check
that coverage probability is at least approximately uniform (e.g. Buckland, Cattanach, and
Gunnlaugsson 1992; Barlow 1994; Branch and Butterworth 2001; Hammond et al. 2002;
Hedley and Buckland 2004).
448 S. STRINDBERG AND S. T. BUCKLAND

Figure 2. The sampler s intersected by an infinitesimal strip perpendicular to the design axis G of width δx, height
H(x) and area (to first order) H(x)δx. The height hs (x) of s is determined by its width 2w and angle θ(x), where
hs (x) = 2w / cosθ(x). The filled parallelogram of area hs (x)δx indicates the covered proportion of the strip. The
h (x)δx h (x)
marginal coverage probability πx (x) is therefore given by πx (x) = s = s .
H(x)δx H(x)

Let hs (x) denote the height of the zigzag sampler s at location x along the design axis
G. Figure 2 shows that hs (x) is given by
2w
hs (x) = . (3.1)
cosθ(x)
The coverage probability π(u) achieved by a zigzag sampler typically varies spatially. We
consider how it varies along the design axis (the horizontal component πx (x)), integrated
over y. Similarly, we use the term “vertical component” to denote coverage probability at
point y, πy (y), after integration over x. Let H(x) denote the height of the survey region at
x. For all types of zigzag design, the horizontal component is given by
hs (x)
πx (x) = . (3.2)
H(x)
This is illustrated in Figure 2.
The vertical component πy (y) does not have an equivalent analytic form. Its value
depends on the type of zigzag sampler, the survey region shape and the total survey effort
L. To ensure that coverage probability will be even throughout the region, a solution is to
select a random orientation for G, provided we select a zigzag sampler such that πx (x)
as given by Equation (3.2) is constant. The algorithms for generating each of the zigzag
samplers described in this article are readily automated, so it is easy to estimate the combined
vertical and horizontal coverage probability as a function of location by simulation. If
this were considered to vary too much from a constant probability, the Horvitz-Thompson
 ZIGZAG SURVEY DESIGNS IN LINE TRANSECT SAMPLING 449

estimator, which does not require the assumption of an even coverage probability, can be
used as described in Section 2.

3.2 GENERAL CONSTRAINTS FOR THE ZIGZAG SAMPLER

Consider a survey region R and a zigzag sampler s that forms an angle of either θ(x)
or 180◦ − θ(x) with design axis G, where sampler angle θ(x) ∈ [0, 90◦ ). In general, θ(x)
varies with distance x along G. Suppose the boundary of the survey region is defined by a
set of straight edges E = {e1 , e2 , . . . , eK }, and let φ = {φ1 , φ2 , . . . , φK } denote the set
of angles formed by each of the ek with G, where k = 1, . . . , K. Let EU denote the set of
“upper” edges that can be intersected by s as it moves in the direction of increasing y. Let
EL denote the set of “lower” edges that can be intersected by s as it moves in the direction
of decreasing y.
Depending on whether an edge ek belongs to the set EU or EL and the angle φk of
ek , there is a corresponding allowable range for the zigzag angle θ(x) that prevents the
sampler s from “bouncing outside” the survey region R. Figure 3 shows four categories of
survey region edge along with the corresponding allowable sampler angle range. Thus, if
we consider a survey region R with edge angles φ for a selected design axis orientation, the
smallest possible sampler angle (corresponding to the least effort L), which ensures that s
remains within R, can be calculated by considering the edge that has the greatest absolute
slope with respect to G (any edge ek perpendicular to G, i.e., φk = 90◦ , is not considered
as the sampler is not required to “bounce off” such edges). Let φ denote the angle that
such an edge forms with G. Then the smallest possible allowable sampler angle is θ = φ
if 0 ≤ φ < 90◦ or θ = 180◦ − φ if 90◦ < φ ≤ 180◦ . (Note that the boundary is taken to
be part of the survey region and the sampler is allowed to coincide with the boundary). The
minimum amount of effort, that is, sampler length L, required to avoid bouncing outside R
can be estimated by applying the formula:
(xmax − xmin )
L= , (3.3)
cosθ
where xmin and xmax define the minimum and maximum values of x within the survey region
R. When the ends of R are not lines perpendicular to G the formula shown in (3.3) is an
approximation and will overestimate the amount of effort required.

4. ZIGZAG SAMPLER ALGORITHMS

4.1 THE EQUAL-ANGLE ZIGZAG SAMPLER

For the equal-angle zigzag sampler, θ(x) = θ for all x. The design may be implemented
by selecting a random (uniformly distributed) point along the design axis, and a sampler
orientation with respect to the design axis of θ or 180◦ − θ, with equal probability. The sam-
pler is extended from the selected point in both directions until it meets the upper and lower
450 S. STRINDBERG AND S. T. BUCKLAND

Figure 3. A survey region edge ek belongs to either the set of “upper” or the set of “lower” edges (EU or EL ) that
can be intersected by the zigzag sampler s as it moves in the direction of increasing or decreasing y, respectively.
Consider a coordinate system whose origin O can be located at any point along ek . The sampler angle (either
θ(x) or 180◦ − θ(x)) may vary with distance x along the design axis G. The restrictions on θ(x), which ensure that
s remains within the boundary of R as it intercepts ek , are determined by its edge set and the angle φk (indicated
by the dotted line in the diagrams) of ek . As s approaches O from the left (as shown in diagrams 1 and 4) or from
the right (diagrams 2 and 3) θ(x) must be such that s passes through the portion of the quadrant indicated by the
vertical lines fill and the quadrant indicated by the diamond fill, which is completely accessible (i.e., s can take
on any orientation in that quadrant and still remain within the boundary of R).

region boundary. At each point of intersection of the sampler with the boundary (termed
a “waypoint”), the sampler “bounces off” the boundary. This is achieved by reflecting the
sampler in a line perpendicular to G passing through the intersection of the sampler with
the boundary. This procedure is continued until the sampler covers the entire x-value range
of G. Designs based on an equal-angle zigzag sampler are denoted by SEAZ .
An equal-angle zigzag sampler is generated using a specified value for either θ or L. We
may obtain θ from L or vice versa from the relationship shown in Equation (3.3). As noted
earlier the value obtained will either be exact or a conservative approximation depending
on whether or not the ends of R are lines perpendicular to G.
As θ is constant, Equation (3.1) yields
2w
hs (x) = =h
cosθ
say, from which Equation (3.2) gives

h
πx (x) = , ∀x ∈ R.
H(x)
 ZIGZAG SURVEY DESIGNS IN LINE TRANSECT SAMPLING 451

Figure 4. The construction of an equal-spaced zigzag sampler (solid line) using the intersection points of a
systematically spaced set of parallel lines (dashed lines) with the boundary of the survey region R. The sample
end segments are constructed using the intersection of the dotted and dot-dashed line.

Thus, the coverage probability is a function of distance along the design axis and is inversely
proportional to the height of the survey region.

4.2 THE EQUAL-SPACED ZIGZAG SAMPLER

To construct the equal-spaced zigzag sampler, a set of equally spaced parallel lines
is constructed perpendicular to, and randomly positioned along, the design axis G and
spanning its full length. Alternate intersections of these lines with the region boundary
form the waypoints of the zigzag sampler, as illustrated in Figure 4. Thus, if the parallel
lines pass north to south, the zigzag is formed by linking the north intersection of one line
with the boundary to the south intersection of the next line, and so on. This fails to define
the sections of sampler between xmin and the first parallel line, and between the last line
and xmax . We chose to define these sections as shown in Figure 4. We denote designs based
on an equal-spaced zigzag sampler by SESZ .
An equal-spaced zigzag sampler thus has waypoints equally spaced along the design
axis. For a specified spacing c, it is possible to estimate the expected total sampler length
for a realization of the design. The actual sampler length varies about this value, depending
on the shape of the survey region and the particular realization of the design. Similarly,
for a fixed total zigzag length L, an estimate of the spacing required to generate that total
line length can be calculated. To estimate L given c or vice versa the following formula is
applied:

 2
(xmax − xmin ) A
L= + c2 .
c xmax − xmin

The formula relies on the knowledge that the average height of the survey region R is given
by the surface area A of R divided by its width (xmax − xmin ) with respect to G. Thus L can
be approximated by multiplying the length of the hypotenuse of the right-angled triangle,
whose other edges include this average height and the spacing c, by the number of spacings
452 S. STRINDBERG AND S. T. BUCKLAND

that fit between xmin and xmax . Again, this formula will approximate values for L or c unless
the ends of R are lines perpendicular to G.
The equal-spaced zigzag sampler can be partitioned into K segments. For each sampler
segment sk the sampler angle θ(x) remains constant. That is, θ(x) = θk for every x ∈ sk ,
where k = 1, . . . , K. Thus, for each sampler segment sk , from Equation (3.1), the height
hk of sk is given by
2w
hs (x) = hk = ∀x ∈ sk , k = 1, . . . , K.
cos(θk )
Equation (3.2) yields
hk
πx (x) = πk = ∀x ∈ sk , k = 1, . . . , K.
H(x)
As for the equal-angle zigzag sampler, the coverage probability varies with distance
along the design axis. It is inversely proportional to the height of the survey region within
segment k. However, the angle of the sampler, and hence the constant of proportionality
hk , differs between sampler segments. This allows a closer approximation to even coverage
probability, with a smaller value for hk in shorter segments.

4.3 THE ADJUSTED-ANGLE ZIGZAG SAMPLER

The equal-angle zigzag and, to a lesser degree, the equal-spaced zigzag do not achieve
an even coverage probability because the surface area of a sampler subsection falling within
any strip of R is not proportional to the surface area of that strip. We can construct a zigzag
sampler that does have this property. From Figure 2, we simply need to ensure that sampler
height hs (x) is proportional to survey region height H(x). Then πx (x) = πc and from
Equations (3.1) and (3.2) together with the approximation for mean coverage probability
this gives
2w 2w × L
= .
cosθ(x) × H(x) A

Thus, the angle θ(x) must be continually adjusted in proportion to the survey region height
H(x). At any position along G, the angle of the sampler is given by
 
−1 A A
θ(x) = cos , ∀x ∈ Rs ⊂ R, ≤ 1, ∀x, (4.1)
LH(x) LH(x)
where Rs denotes the sampled subregion of R.
Note that even coverage probability for the above continuous sampler is achievable
only if H(x) ≥ (A/L) for all x ∈ R. As for other zigzag samplers, the procedure relies on
being able to “bounce off” a survey region boundary at an angle θ(x) to G. This is possible
only when the absolute value of the slope of the boundary of R is smaller than the absolute
value of the slope of the sampler. (Note that boundary sections at an angle of 90◦ , that is,
with an infinite slope, need not be considered as the sampler terminates when it reaches such
 ZIGZAG SURVEY DESIGNS IN LINE TRANSECT SAMPLING 453

sections.) This condition always holds as θ(x) → 90◦ , which occurs as L → ∞. For smaller
values of θ(x), subregions may occur in which a continuous sampler with constant coverage
probability along the design axis is not achievable. When H(x) = (A/L), θ(x) = 0◦ , and
at smaller values of H(x), even a sampler that is parallel to the design axis over-samples
an infinitesimal strip at x. For most surveys, bias from this source is likely to be minimal,
but if it is considered to be a significant problem, a solution is to reduce search effort along
this section of transect, for example, by traveling “off-effort” for short segments.
In the special case of a rectangular survey region of height H and width W , whose top
and bottom edge runs parallel to the design axis, the sampler angle is given by
   
−1 H ×W −1 W
θ(x) = cos = cos (L ≥ W ),
L×H L
independent of x. In this case, all three zigzag samplers considered here are equivalent.
To implement the adjusted-angle zigzag sampler, first select a starting point at random
within the survey region, and determine the corresponding angle θ(x) from Equation (4.1).
Select from θ(x) and 180◦ −θ(x) with equal probability to determine the angle of the sampler
to the design axis G. Extend the sampler from the selected point in both directions along
G, while continually adjusting the angle according to Equation (4.1). When it intersects the
upper or lower boundary of R, it is reflected in a similar manner to the equal-angle zigzag
is, except that the sampler angle is determined by the height of R at that x-coordinate. This
procedure is continued until the sampler covers the entire x-value range of G. We denote
designs based on an adjusted-angle zigzag sampler by SAAZ .

4.4 COMPARISON OF ZIGZAG SAMPLERS

As noted previously, general practice is to estimate animal abundance assuming that
the zigzag sampler design attains even coverage probability πc . Under that assumption, and
given knowledge of animal density D(x, y) within the area covered by the sampler, we
could obtain a Horvitz-Thompson estimator (Horvitz and Thompson 1952) of abundance
as
 xmax  ys (x)+hs (x)
1
N̂HT = D(x, y)dydx,
πc xmin ys (x)

where xmin and xmax define the full range of x within the survey region, and ys (x) is the
lower y-extreme of the sampler at distance x along the design axis. This equation can
equivalently be written as

1
N̂HT = D(u)z(u)du, (4.2)
πc R
where z(u) is an indicator function, equal to one if u ∈ Rs , and zero otherwise.
These results are not useful for analysis of real survey data, because D(x, y) (equiv-
alently D(u)) cannot be observed. However, they allow us to evaluate bias arising from
assuming that the equal-angle and equal-spaced zigzag samplers provide even coverage
454 S. STRINDBERG AND S. T. BUCKLAND

Figure 5. A trapezoidal survey region R, whose design axis G and edge coincide with the x-axis. Within R three
types of zigzag sampler are illustrated: equal-angle (dotted line), equal-spaced (dashed line), and adjusted-angle
(solid line).

probabilities, given an assumed form for D(u), without the need to simulate. Strindberg
(2001) evaluated the bias in N̂HT for three cases: D(u) = αx + β, ∀u ∈ R where α, β
are positive constants; D(u) = αx + βy + γxy + δ, ∀u ∈ R, where α, β, γ, δ are positive
constants; and D(u) = α(x)y 2 + β(x)y + γ(x), ∀u ∈ R where

4(Dmax (x) − Dmin (x))

α(x) = ,
(H(x))2

β(x) = −α(x)H(x),

γ(x) = Dmax (x),

and

Dmin (x), Dmax (x),

are the minimum and maximum density with respect to y for a given x. The functional
form of Dmin (x) and Dmax (x) is such that Dmax (x) > Dmin (x) > 0, ∀x ∈ R. The survey
region was taken to be a trapezium with vertices at (0, 0), (0, 100), (120, 20), and (120, 0),
respectively (Figure 5).
Each of the three zigzag samplers was constructed with the same total length L. To
compare them visually, we started all three from the origin in Figure 5, although in practice,
 ZIGZAG SURVEY DESIGNS IN LINE TRANSECT SAMPLING 455

Figure 6. Coverage probability πx (x) plotted against x for the three zigzag samplers in Figure 5. Also shown is
the height of the trapezium (not to scale) as a function of distance along the design axis, which indicates that the
equal-angle zigzag (dotted line) has too low coverage probability where the survey region is wide, and too high
where it is narrow. For the equal-spaced zigzag (dashed line), the coverage probability changes for each change
in sampler angle. The coverage probability for the adjusted-angle zigzag (solid line) remains constant, because
the angle varies as a smooth function of the trapezium height.

their location within the survey region would be random. The curve of the adjusted-angle
zigzag becomes especially apparent in Figure 5 as the height of the survey region becomes
small. Also apparent is the much better approximation of the equal-spaced zigzag sampler to
the adjusted-angle sampler than that of the equal-angle sampler. This is reflected in Figure 6,
which shows that the equal-spaced sampler provides less variable coverage probability than
does the equal-angle sampler.
Strindberg’s (2001) evaluations of bias conform to expectation, with the equal-spaced
sampler (SESZ ) yielding much lower bias under the assumption of even coverage probability
than the equal-angle sampler (SEAZ ). Using Equation (4.2), the model D(u) = αx + β with
α equal to .02, .2, or 2 and β equal to .05, .5, or 5 gave percentage biases in N̂HT ranging
from 4.5% to 28.6% (median = 27.1%) for SEAZ and from 1.0% to 6.1% (median = 5.8%)
for SESZ . The model D(u) = αx + βy + γxy + δ with α equal to .002, .02, or .2, β equal
to .003 or .03, γ equal to .001 or .01, and δ equal to .005 gave percentage biases in N̂HT
ranging from 1.1% to 26.0% (median = 10.5%) for SEAZ and from .2% to 5.6% (median
= 2.2%) for SESZ . The model D(u) = α(x)y 2 + β(x)y + γ(x) with Dmin (x) = αd x + βd ,
Dmax (x) = γd Dmin (x), ∀x ∈ R and αd equal to .0001, .001, .01, .1, 1, or 10, βd equal
to .3 or 3 and γd equal to 2.5, gave percentage biases in N̂HT ranging from .05% to 28.6%
(median = 17.4%) for SEAZ and from .01% to 5.8% (median = 3.6%) for SESZ .
456 S. STRINDBERG AND S. T. BUCKLAND

These simulations give a clear message. If conventional analyses that assume even
coverage probability are used, then equal-spaced zigzag samplers give substantially lower
bias than equal-angled samplers. Bias from this source can be eliminated entirely by using
the adjusted-angle zigzag sampler, at the cost of increased complexity of design.

5. SURVEYS OF MINKE WHALES IN THE ANTARCTIC

Annual surveys of minke whales are conducted in the Antarctic under the auspices of the
International Whaling Commission. The Antarctic is divided into six different management
areas, and in most years, a single management area is surveyed. These surveys provide a
good illustration of the value of automated design algorithms. Whale density is highest at
the ice-edge, so sampling effort is concentrated within the ice-edge stratum. The ice-edge is
very irregular, and can move by up to 100 kilometers in a single day, so it is not possible to
lay down the design in advance. Instead, an algorithm must be defined, to allow the transects
to be determined as the survey progresses.
The zigzag samplers were defined for convex survey regions. For nonconvex regions,
depending on the characteristics of the region, the amount of available effort and the orien-
tation of the design axis, it may be possible to generate the zigzag samplers. If, however,
some parts of the region are inaccessible to the sampler, the properties explored here do
not hold. Minke whale surveys within the ice-edge strata can provide an extreme depar-
ture from convexity. To ensure that the entire survey region is accessible and that coverage
probabilities are nonzero throughout the region, one of three approaches can be employed.
These are illustrated in Figure 7 and are as follows:

1. When departure is mild, the problem may be overcome by designing the survey
within a convex hull around the survey region. Sections of sampler that fall outside the
survey region are then not sampled. It is advantageous to use a convex hull rather than a
minimum bounding rectangle of the survey region for survey design purposes, as the relative
sampler discontinuity is less and a more even coverage probability πx (x) can be achieved
by using the former. However, Figure 7(a) shows that this solution is far from satisfactory
for this particular ice-edge stratum.

2. Consider dividing the stratum into subregions, each of which is convex. This strategy
can be successful, but again, for the ice-edge stratum of Figure 7, far too many subregions
would be required. Instead, define convex hulls around almost convex subregions of the
original survey region, giving a reasonable design as shown in Figure 7(b). In this case,
three subregions were defined, and a convex hull was established around each one. The
solution is not ideal as inefficiency arises from sampler discontinuity and the need for
relocation between subregions, but an advantage is that the orientation of the design axis
can be separately defined for each subregion.

3. The last approach attempts to generate the zigzag samplers within the nonconvex
survey region itself. In the example shown in Figure 7(c), this approach minimizes the sam-
 ZIGZAG SURVEY DESIGNS IN LINE TRANSECT SAMPLING 457

Figure 7. The survey stratum WN and a more complex ice-edge stratum WS in the Antarctic. (a) A single realization
of a zigzag sampler is shown in the convex hull of WS. The sampler is discontinuous (because sections outside the
survey region are not covered) and thus inefficient. (b) WS has been divided into three almost convex sub-regions,
and a zigzag sampler placed within the convex hull of each, reducing the sampler discontinuity. (c) A zigzag
sampler is generated in the nonconvex WS giving the least sampler discontinuity in this instance. Within WN three
types of zigzag sampler are illustrated: equal-angle (dotted line), equal-spaced (dashed line) and adjusted-angle
(solid line).
458 S. STRINDBERG AND S. T. BUCKLAND

Table 1. Estimates of Pod Abundance for the WN Stratum of the 1987–1988 International Decade of
Cetacean Research survey of Antarctic Minke Whales

Assumed zigzag sampler Estimate 95% confidence interval

Constant coverage probability 4260 (510, 17040)
Equal-angle 3740 (430, 14790)

pler discontinuity. It may, however, not be a viable solution for certain types of nonconvexity,
especially when attempting to generate the adjusted-angle zigzag sampler.

Illustration of the Horvitz-Thompson estimators of Section 2 is not straightforward,

because automated algorithms for zigzag samplers have not been available until now. How-
ever, we use the “west north” (WN) stratum of the 1987–1988 surveys for illustration, as
the realized design is a close approximation to an equal-angle zigzag design. Within this
stratum of area 148,820 nautical miles squared, there were seven legs of effort, totaling
1,500 nautical miles. Half of each leg was covered in “passing mode” and half in “closing
mode.” In “closing mode,” the survey vessel ceases searching when a pod (school) of whales
is detected, and closes with it, to confirm pod size. Because this procedure can create some
bias in the estimated encounter rate, “passing mode” is also used, in which pod sizes are
not confirmed, but encounter rate is likely to be less biased. Full details of the analysis of
the 1987–1988 survey were given by Borchers and Haw (unpublished). Within the WN
stratum, 44 minke whale pods were detected within 1.5 nautical miles of the trackline, of
which 33 were detected while in passing mode.
We do not attempt a full reanalysis of the data here. We use the estimated effective strip
half-widths from Borchers and Haw (unpublished): .555 nautical miles with coefficient
of variation 40.3% for passing mode, and .418 nautical miles with coefficient of variation
25.2% for closing mode. These were found using a truncation distance of 1.5 nautical miles,
so that the survey strips are three nautical miles wide. For each pod detected, the probability
that it would have fallen within a survey strip for a single random design of the equal-angle
zigzag sampler was estimated from 10,000 realizations of the design. Conventional analysis
assumes that coverage probability is the same everywhere, in which case each pod has the
same probability of 1,500 × 3/148, 820 = .0302.
We show the resulting estimates of numbers of minke whale pods in Table 1. Assum-
ing constant coverage probability, the estimated pod abundance N̂CCP was obtained using
Equation (2.3) for each of closing mode and passing mode, and summing the two estimates
(with µ̂i equal to .555 for pods detected while in passing mode, and .418 for closing mode).
The estimated pod abundance N̂EAZ was obtained similarly, using the coverage probabilities
corresponding to each sighting achieved by an equal-angle zigzag design. Bootstrapped con-
fidence intervals were obtained using the percentile method. Parametric bootstrap replicates
of the effective strip half-width were generated, assuming a log-normal distribution, and
nonparametric resampling of effort legs was conducted to quantify variability in encounter
rate.
The equal-angle sampler’s coverage probabilities varied between .0207 and .0557,
 ZIGZAG SURVEY DESIGNS IN LINE TRANSECT SAMPLING 459

with an abundance estimate that is 520 pods lower than for the conventional analysis. This
suggests substantial bias occurs when the properties of the zigzag samplers are ignored,
although the high variability in these estimates makes it difficult to draw strong conclusions.

6. DISCUSSION
For some survey regions, when the available survey effort is limited, it will be impossible
to avoid a boundary and sampler orientation combination that forces the zigzag sampler
out of R. This problem could be resolved by letting the sampler run along the boundary
to minimize the discontinuity, while maintaining the same total line length, although this
leads to over-sampling along the edge. The effect on coverage probability could be assessed
by simulation. Depending on the shape of the survey region, it may be possible to avoid
the problem by suitable choice of orientation of the design axis. Failing this, a pragmatic
option is to approximate the shape of the survey region in a way that avoids the problem.
For populations known to exhibit a density gradient, the zigzag sampler orientation
should usually be chosen so that each leg as far as possible cuts across all density levels.
This minimizes variation in encounter rate among lines (Buckland et al. 2001, pp. 238–239),
so that better precision in the estimated abundance is achieved. For zigzag designs, this is
accomplished by placing the design axis approximately parallel to the density contours.
In our example, minke whales have highest density along the ice edge, so that the design
axis is chosen to be roughly parallel to the ice edge. If nothing is known about density
gradients within the population, and assuming that the sampling unit is taken to be a single
leg of the zigzag sampler, then a sampler orientation that maximizes the number of legs
is advantageous. This increases the sample size for variance estimation and yields a more
precise estimate of variance.
Zigzag samplers have an associated width, so overlap at the boundaries of the survey
region occurs between successive legs of the sampler. This overlap is not a problem provided
detected animals that fall within the area of overlap are recorded for one of the legs and
that the effort in the area of overlap contributes to total effort only once. Another problem
is that part of the sampler may fall outside the survey region at the boundary. This can lead
to a coverage probability that is smaller than expected. Usually, the sampler width is very
small compared to the sampler length, so that overlap and other edge effects are negligible.
Many ship and aerial surveys span large geographic regions, so that the loss of survey
effort in transiting from one line to another of a parallel-line design can be substantial. In
these circumstances, zigzag surveys should be considered. In the case of shipboard surveys,
if lines of a parallel-line design are say 20 kilometers apart, it may take an hour to traverse
from one line to the next. Given total line length, line separation and number of lines,
a straightforward calculation reveals the percentage of potential search effort lost during
such traverses. A zigzag design eliminates almost all of this loss. Aircraft travel faster, and
a break between lines can be helpful, giving observers the chance of a short rest, which
improves their concentration while on-effort. The operation costs per hour are also typically
much less than for a ship. Hence a greater loss in potential survey time might be deemed
460 S. STRINDBERG AND S. T. BUCKLAND

acceptable. However, if loss of search effort is of the order of 10% or more relative to a
zigzag design, we suspect that most survey planners would opt for a zigzag design.
We recommend that equal-angle zigzag samplers not be used. In practice, equal-spaced
zigzag samplers are likely to prove satisfactory in most surveys, typically with low bias,
even if constant coverage probability is assumed. Increasingly, surveys are conducted with
sophisticated navigation systems, and in these circumstances, the adjusted-angle zigzag
sampler may be practical, and offer a worthwhile improvement over equal-spaced zigzag
samplers when the shape of the survey region is irregular.

ACKNOWLEDGMENTS
We thank the two referees for their thorough and constructive reviews. The research for this article was
completed within the School of Mathematics and Statistics at the University of St Andrews, Scotland.

[Received September 2002. Revised February 2004.]

REFERENCES

Barlow, J. (1994), “Abundance of Large Whales in California Coastal Waters: A Comparison of Ships Surveys in
1979/80 and in 1991,” Report of the International Whaling Commission, 44, 399–406.
Borchers, D. L., Buckland, S. T., Goedhart, P. W., Clarke, E. D., and Hedley, S. L. (1998), “Horvitz-Thompson
Estimators for Double-platform Line Transect Surveys,” Biometrics, 54, 1221–1237.
Borchers, D. L., and Haw, M. D. (unpublished), “Estimation of Minke Whale Abundance from the 1987/8
IWC/IDCR Antarctic Assessment Cruise in Area III,” Paper SC/41/SHMi4; available from the International
Whaling Commission.
Branch, T. A., and Butterworth, D. L. (2001), “Estimates of Abundance South of 60◦ S for Cetacean Species
Sighted Frequently on the 1978/79 to 1997/98 IWC/IDCR-SOWER Sighting Surveys,” Journal of Cetacean
Research and Management, 3, 251–270.
Buckland, S. T., Cattanach, K. L., and Gunnlaugsson, Th. (1992), “Fin Whale Abundance in the North Atlantic,
Estimated from Icelandic and Faroese NASS-87 and NASS-89 Data,” Report of the International Whaling
Commission, 42, 645–651.
Buckland, S. T., Anderson, D. R., Burnham, K. P., Laake, J. L., Borchers, D. L., and Thomas, L. (2001), An
Introduction to Distance Sampling, Oxford: Oxford University Press.
Chen, S. X., and Cowling, A. (2001), “Measurement Errors in Line Transect Surveys where Detectability Varies
with Distance and Size,” Biometrics, 57, 732–742.
Clarke, E. D., Spear, L. B., McCracken, M. L., Marques, F. F. C., Borchers, D. L., Buckland, S. T., and Ainley,
D. G. (2003), “Application of Generalized Additive Models to Estimate Size of Seabird Populations and
Temporal Trend from Survey Data Collected at Sea,” Journal of Applied Ecology, 40, 278–292.
Cooke, J. G. (1985), “Estimation of Abundance from Surveys,” unpublished manuscript.
Foote, K. G., and Stefansson, G. (1993), “Definition of the Problem of Estimating Fish Abundance Over an Area
from Acoustic Line Transect Measurements of Density,” ICES Journal of Marine Science, 50, 369–381.
Hammond, P. S., Berggren, P., Benke, H., Borchers, D. L., Collet, A., Heide-Jørgensen, M. P., Heimlich, S., Hiby,
A. R., Leopold, M. F., and Øien, N. (2002), “Abundance of Harbour Porpoise and Other Cetaceans in the
North Sea and Adjacent Waters,” Journal of Applied Ecology, 39, 361–376.
 ZIGZAG SURVEY DESIGNS IN LINE TRANSECT SAMPLING 461

Hedley, S. L., and Buckland, S. T. (2004), “Spatial Models for Line Transect Sampling,” Journal of Agricultural,
Biological, and Environmental Statistics, 9, 181–199.
Horvitz, D. G., and Thompson, D. J. (1952), “A Generalization of Sampling without Replacement from a Finite
Universe,” Journal of the American Statistical Association, 47, 663–685.
Huggins, R. M. (1989), “On the Statistical Analysis of Capture Experiments,” Biometrika, 76, 133–140.
Marques, F. F. C., and Buckland, S. T. (2003), “Incorporating Covariates into Standard Line Transect Analyses,”
Biometrics, 59, 924–935.
O’Connell, V. M., and Carlile, D. W. (1993), “Habitat-specific Density of Adult Yelloweye Rockfish Sebastes
ruberrimus in the Eastern Gulf of Alaska,” Fishery Bulletin, 91, 304–309.
Pojar, T. M., Bowden, D. C., and Gill, R. B. (1995), “Aerial Counting Experiments to Estimate Pronghorn Density
and Herd Structure,” Journal of Wildlife Management, 59, 117–128.
Pople, A. R., Cairns, S. C., Clancy, T. F., Grigg, G. C., Beard, L. A., and Southwell, C. J. (1998), “An Assessment
of the Accuracy of Kangaroo Surveys Using Fixed-Wing Aircraft,” Wildlife Research, 25, 315–326.
Strindberg, S. (2001), “Optimized Automated Survey Design in Wildlife Population Assessment,” PhD thesis,
University of St. Andrews.
Trenkel, V. M., Buckland, S. T., McLean, C., and Elston, D. A. (1997), “Evaluation of Aerial Line Transect
Methodology for Estimating Red Deer (Cervus elaphus) Abundance in Scotland,” Journal of Environmental
Management, 50, 39–50.
van Hensbergen, H. J., Berry, M. P. S., and Juritz, J. (1996), “Helicopter-based Line-transect Estimates of Some
Southern African Game Populations,” South African Journal of Wildlife Research, 26, 81–87.
White, G. C., Bartmann, R. M., Carpenter, L. H., and Garrott, R. A. (1989), “Evaluation of Aerial Line Transects
for Estimating Mule Deer Densities,” Journal of Wildlife Management, 53, 625–635.
Wiig, Ø., and Derocher, A. E. (1999), “Application of Aerial Survey Methods to Polar Bears in the Barents Sea,”
in Marine Mammal Survey and Assessment Methods, eds. G. W. Garner, S. C. Amstrup, J. L. Laake, B. F. J.
Manly, L. L. McDonald, and D. G. Robertson, Rotterdam: Balkema Press, pp. 27–36.