The Journal of Wildlife Management 79(8):1361–1368; 2015; DOI: 10.1002/jwmg.

943

Research Article

Occupancy Modeling of Bird Point Counts:

Implications of Mobile Animals
DANIEL B. HAYES,1 Department of Fisheries and Wildlife, Michigan State University, 480 Wilson Road, Michigan State University,
East Lansing, MI 48824-1222, USA
MICHAEL J. MONFILS, Michigan Natural Features Inventory, Michigan State University Extension, P.O. Box 13036, Lansing,
MI 48901-3036, USA

ABSTRACT Occupancy modeling has been applied to a wide variety of taxa and sampling methods,
including bird point counts. A critical assumption of basic occupancy models is that sites are occupied
throughout the duration of the study, which is unlikely to be true for typical bird point-count studies. As
such, we evaluated the implications of mobile animals on parameter estimates. We simulated the movement
and detection of individual birds using an individual-based simulation model. We fit the basic occupancy
model to data that represented a range of animal mobility, and determined the bias relative to known
parameters used in the simulation. Occupancy depends on the size of the site selected, with smaller sites
leading to lower occupancy for a given area and number of individuals present. At low animal density,
occupancy scales approximately linearly with the area of sites, but at very high density, occupancy asymptotes
at 1.0 across all site sizes. Even small amounts of movement lead to bias in estimates of occupancy and
detectability, and the typical size of bird home ranges can lead to highly biased parameters. Moreover,
variation in home range size over time or across habitats can lead to varying degrees of bias. Because of the
potential for large bias in occupancy estimates, and their sensitivity to behaviors of birds (e.g., home range
size), we recommend against applying current occupancy models to bird point-count data. Ó 2015 The
Wildlife Society.

KEY WORDS birds, closure assumption, detectability, home range size, individual-based models, movement
behavior, occupancy modeling, point count.

Occupancy modeling (e.g., MacKenzie et al. 2002,2006) has
been widely applied in studies of the demography of many
animal groups, including birds (e.g., Frey et al. 2011, Miller
et al. 2011, Warren et al. 2013), mammals (Kalies et al.
2012), fish (Falke et al. 2012), amphibians (MacKenzie et al.
2002), and reptiles (Kery et al. 2009). This modeling is
intended to address shortcomings of approaches that assume
complete detectability, an assumption that is rarely upheld in
natural populations. As MacKenzie et al. (2006) articulate,
estimates of occupancy provide important insights into
various aspects of animal ecology, including habitat selection
and metapopulation dynamics. Because of the power and
flexibility of these models, their application has become
standard practice for many wildlife studies.
Assumptions of the basic occupancy model (MacKenzie
et al. 2006) are 1) occupancy status at each site does not
change over the survey season (closure); 2) the probability of
occupancy is constant across sites, or is a function of
covariates; 3) the probability of detection is constant across
Received: 27 February 2015; Accepted: 1 July 2015
Published: 13 August 2015
1
E-mail: hayesdan@msu.edu
sites and surveys, or is a function of covariates; 4) detection of
species and detection histories at each location are
independent; and 5) a species is not falsely detected when
absent. Of these assumptions, the closure assumption seems
most likely to be violated for bird point-count samples and is
the focus of this investigation. Although methods have been
proposed to accommodate lack of closure (e.g., Farnsworth
et al. 2002, Rota et al. 2009, Chandler et al. 2011, Amundson
et al. 2014), these methods require additional data (such as
distance measurements, or identification of individual
animals) or changes to sampling methods. Such an approach
(sometimes termed a robust design; e.g., Kery et al. 2009,
Rota et al. 2009), although useful, still requires closure at
some temporal or spatial scale.
In our research applying these models to point counts of
marsh birds (Monfils et al. 2014), we have become concerned
that the assumptions of the basic occupancy model (i.e.,
MacKenzie et al. 2006) are often violated, particularly the
assumption that the occupancy status of a site does not
change. Some authors have addressed this concern,
concluding that animal movement in and out of a site
does not seriously bias results (e.g., MacKenzie et al. 2004,
Kendall and White 2009), whereas other authors have
concluded that such movement is an issue that requires

Hayes and Monfils
Occupancy Modeling of Mobile Animals 1361

changes in sampling design or analytical methods (e.g., Rota
et al. 2009, Efford and Dawson 2012). We observe that the
basic occupancy model and assumptions continue to be
applied in many studies (e.g., Darrah and Krementz 2009,
Bolenbaugh et al. 2011, Valente et al. 2011), possibly because
of the conflicting advice provided in theoretical papers. The
goal of this paper is to examine the closure assumption of
occupancy models and where violations of this assumption
could lead to potentially erroneous results, with a focus on
applications to bird point counts. Our main objective was to
determine the performance of the standard occupancy model
and typical bird point-count data in light of the mobility of
birds as an example of mobile animals.
We focused on the common sampling design where N sites
are visited T times, assuming a constant probability of
occupancy (c) and constant detection probability (p) across
all sites and sampling visits. We note, however, that the
terminology associated with occupancy modeling can at
times be confusing, and for clarity, we reiterate the definition
of terms commonly used in occupancy modeling. The first
critical term is occupancy itself. At the scale of an individual
site, a site is occupied if it contains at least 1 or more
individuals of the focal species across the survey season,
consistent with assumption 1 above. For a broader region,
occupancy is often defined as the proportion of sites
containing 1 or more individuals, or the probability that a
site contains individual(s). If the closure assumption holds,
this definition also leads to the concept of proportion of area
occupied (PAO; MacKenzie and Royle 2005). For situations
where mobile animals lead to a violation of the closure
assumption, Chandler et al. (2011) use the term temporary
immigration or emigration to emphasize that movement
within an animal’s home range can lead to temporary
absences from an occupied site. MacKenzie and Royle (2005)
suggest that the term use should replace occupy, leading to
the concept of proportion of area used. When mobile animals
are considered, Efford and Dawson (2012) further distin-
guish asymptotic occupancy, which they define as the
accumulated area used over time, in contrast to instantaneous
occupancy, which is the proportion of sites used at a specific
point in time.
Within this definition of occupancy, the concept of what
constitutes a site is often flexibly defined across studies. Some
applications of the occupancy model focus on meta-
population dynamics, where a site is a patch large enough
to contain a population, and where the boundaries of the site
coincide with population boundaries. Clear examples of this
definition of a site are wetlands that contain breeding
subpopulations of amphibians (e.g., MacKenzie et al. 2002),
or islands containing discrete populations. However, in many
other cases, the size of a site is controlled by the investigator,
and is chosen for logistical concerns and not necessarily
related to the biology of the species. Efford and Dawson
(2012) use the phrase “arbitrary plots in continuous habitat”
to define this situation, which often occurs in studies of bird
populations. For example, a common definition of site for
bird point counts is a circle with a 50-m radius, where
detection depends on seeing or hearing an individual within
1362
that distance from the site’s center point (e.g., Rota et al.
2009, Monfils et al. 2014). For other taxa, such as small
mammals (e.g., deer mouse [Peromyscus maniculatus]) or
terrestrial amphibians (e.g., red-backed salamander [Pletho-
don cinereus]), sites are often square regions where traps are
set or the region searched (e.g., Otto and Roloff 2011). For
larger mammals, camera traps are commonly used to detect
the presence of target species (e.g., Clare et al. 2015).
Finally, we note that detection probability can be
interpreted in several ways. We simulated the situation
where individual animals have a probability of being detected
if present, and for simplicity, we treat this probability (p) as
being constant across all sites and sampling periods. In
contrast, the standard occupancy model is founded on a
definition of detection probability where p is the probability
that a species will be detected at a site if present, regardless of
the number of individuals present. Amundson et al. (2014)
further distinguish the probability of detection of individual
animals within a site as being a function of availability (pa)
and perceptibility (pd). Availability (pa) is the probability that
an animal is present at a site at the time of sampling, and
perceptibility (pd) is the probability that an animal will be
detected if present. We note, however, that the standard
occupancy model, where individual animals are not distin-
guished and detection is recorded as a binary response for a
site, does not distinguish these components.
METHODS
We constructed an individual-based model to simulate the
main biological processes involved in occupancy modeling.
We created a 2,500-m  2,500-m region, containing 2,500
sites, each 50 m  50 m in size. We used square sites in the
simulation because circular sites do not provide complete
coverage of a region (i.e., circular sites do not form a regular
tessellation). Within this region, we randomly and indepen-
dently distributed a total of M animals by generating a
random uniform number U(0, 2,500) for both an x and y
position (Xc,Yc) that we treated as the centroid of the
animal’s home range, which we then converted to a site
location.
To ensure our estimation procedure replicated results from
previous studies, we included scenarios where animals were
immobile (i.e., home range is a point in space) as well as
having larger home ranges. In initial simulations, animals
were not allowed to move from the centroid of their home
range, thereby meeting the assumptions of the basic
occupancy model. In simulations where animals were mobile,
we treated the probability of detecting an individual animal if
present (p) as a constant across the entire study region and
across all survey periods. When detection probability was less
than 1.0 (i.e., incomplete detectability), the detection of
individual animals occurred at random, and we simulated
detection by generating a U(0, 1) random variate, with
detection occurring when this random variate was less than
the detectability (p) for that simulation scenario. We
aggregated observations of all animals occurring within a
site within a survey period, and developed a detection history
for each site based on whether no (coded as 0), or 1 or more
The Journal of Wildlife Management
79(8)
animals were detected (coded as 1) at the site. This Table 1. Relationship between home range multiplier (h), radius of a circle
encompassing 95% of simulated points, and area of the circle. P(central) is
formulation follows the conceptual model presented in Royle the proportion of detections that would take place within the site in which
and Nichols (2003). the circle’s centroid is located. We chose steps for h to represent 95% home
For simulations with mobile animals, we treated the range circles that are 10%, 50%, 100%, 2, 5, and 10 of the area of the
location generated above as the centroid or focal point of the 50  50-m site.
animal’s home range. For each sampling period, we h Radius (m) Area (m2) P (central)
simulated an animal’s position within its home range by 3.645 8.92 250 0.943
the following function: 8.152 19.95 1,250 0.874
11.527 28.21 2,500 0.824
ðXt ; Yt Þ ¼ ðXc þ Nð0; 1Þ  h; Yc þ Nð0; 1Þ  hÞ ð1Þ 16.302 39.90 5,000 0.758
25.775 63.08 12,500 0.631
36.451 89.21 25,000 0.503
where, (Xc, Yc) is the fixed centroid of the animal’s home
range, t is an index for survey period (e.g., whether it is the
first, second, or later visit to the site), and h is a multiplier
used to scale the size of an animal’s home range. This
implementation resulted in observations within the home provided us with a complete census of all possible sites,
range that were distributed as a bivariate normal variate allowing a direct evaluation of the bias and consistency of the
(Fig. 1). A consequence of this implementation is that home estimators (as defined by Cochran 1977) without the
ranges were circular in shape. We calculated the radius of a potential confounding effects of imprecision that would
circle enclosing 95% of generated points for 6 different occur with a subsample of sites. The topic of the precision of
scaling multipliers (h; Table 1). We then discretized the x,y estimates as a function of number of sites sampled, number of
positions of individual observations at time t to associate with visits, and detectability has been well addressed in the
a specific site, and summarized the data to produce a literature (e.g., Mackenzie et al. 2002), and as such was not a
detection history for each site (Fig. 2). focus of our simulations. Bias in occupancy and detection
Following data generation, we applied the basic occupancy probability was evaluated as a function of number of animals
model to estimate occupancy and detection probability across present, size of home range, number of repeat visits to each
the region simulated. We obtained estimates of occupancy site, and detectability. We evaluated 4 scenarios: 1) immobile
and detection probability by maximizing the likelihood animals with complete detectability; 2) immobile animals
function (Mackenzie et al. 2002, 2006): with incomplete detectability; 3) mobile animals with
complete detectability; and 4) mobile animals with incom-
Lðc;pÞ ¼ ðcn: Ppnt
t ð1  pt Þ
n:nt
Þ  ðcPð1  pt Þ þ ð1  cÞÞN n plete detectability.
ð2Þ

where t is the number of visits to a site, N is the number of

sites surveyed, and n. is the number of sites where at least 1
detection occurred. Because our focus was the bias of
estimators, we simulated the situation where data from all
sites in the region for all time periods were collected. This

Figure 2. Example of 10 detection points for an animal with a home range

Figure 1. Example of 500 detection points for an animal with a home range
multiplier (h) of 50 across a 50  50-m grid.
multiplier (h) of 50 across an area broken into 50  50-m sites, with the 9
sites surrounding the centroid indicated by letters. The plus symbol denotes
the centroid of the animal’s home range, and the numbered points indicate
the location of the animal on each of 10 independent visits to the sample
area. Potential detections histories for the 9 sites surrounding the centroid
are: A (0000000000), B (0000000010), C (1000000000), D (0100000000),
E (0000101100), F (0001000001), G (0000000000), H (0010010000), and I
(0000000000).

Hayes and Monfils
Occupancy Modeling of Mobile Animals 1363

RESULTS
Scenario 1: Immobile Animals With Complete
Detectability
As expected, occupancy within a region increased asymptot-
ically with increasing number of animals present (Fig. 3)
when animals do not exhibit temporary emigration or
immigration and where they are detected with a probability
of 1.0. Clearly, if only 1 animal is present, then only 1 site can
be occupied. With N sites, this yields a probability of
instantaneous occupancy for any single site of 1/N. When
more than 1 animal is present, then each site may have 0, 1, 2,
up to M animals present. The probability that a specific
number of animals is present at a given site, assuming
animals are randomly and independently distributed can be
readily computed from the binomial distribution:
k!
PðxÞ ¼ px qkx
x!ðkxÞ!
where k ¼ number of trials (i.e., number of animals present
¼ M); p ¼ 1/N; q ¼ 1  (1/N), and x ¼ number of animals
present at a site. Note that in the traditional formulation of
the binomial distribution used here, p refers to the
probability of success, and not detection probability referred
to elsewhere in the paper. As an example, if 50 animals are
present within a region composed of 1,000 sites, the
probability that a site will have exactly 1 animal is:
50!
Pðx ¼ 1Þ ¼ 0:0011 0:999ð501Þ ¼ 0:0476
1!ð50  1Þ!
Using the binomial distribution as above, we can compute
the proportion of sites that are unoccupied as:
k!
Pð0Þ ¼ p0 qðk0Þ ¼ qk
0!ðk  0Þ!
And the proportion of the sites that are occupied (i.e., x  1)
by its complement:
Pðx  0Þ ¼ 1  qk
Figure 3. Comparison of simulated occupancy (diamonds) and theoretical
occupancy predicted by the binomial distribution (solid line) as a function of
animal abundance for a hypothetical region composed of 2,500 sites.
Animals were immobile in this simulation and were assumed to be 100%
detectable; we conducted 100 simulations for each level of abundance.
1364
Comparison of results predicted by the binomial distribution
and the simulation produced equivalent results (Fig. 3).
Because the probability of success (p) in the binomial
depends on both the number of animals in a region, and the
number of sites that the region is broken into, these results
have 2 important implications. Occupancy for randomly and
independently distributed individuals is inherently a density-
dependent process, with a nonlinear relationship between
density and occupancy. Occupancy depends critically on the
size of the site selected, with smaller sites leading to lower
occupancy for a given region and number of individuals
present. Further, the relationship between occupancy and
site size varies with the number of animals present. At low
animal density, occupancy scales approximately linearly with
the area of sites, with a doubling of site area halving
occupancy. At very high density, occupancy asymptotes at 1.0
in all cases (Fig. 4).
ð3Þ
Scenario 2: Immobile Animals With Incomplete
Detectability
Consistent with prior research, our results confirm that when
the assumption of immobility is met, the methods outlined in
MacKenzie et al. (2002, 2006) produce unbiased estimates of
occupancy and detectability across a range of population sizes
and levels of detectability. Results of this scenario confirm
correct implementation of the maximum likelihood estima-
tion procedure (Fig. 5).
Scenario 3: Mobile Animals With Complete
ð4Þ Detectability
Estimated occupancy was unbiased when home range
size ¼ 0 (as shown in scenario 2), but bias increased rapidly
as a function of the home range size (Fig. 6). As in the above
scenarios, occupancy was a function of the number of animals
present, and for a relatively small number of animals
ð5Þ (M ¼ 100–300; occupancy for immobile animals
¼ 0.04–0.11), increases in occupancy with home range size
ð6Þ
Figure 4. Occupancy predicted by binomial distribution as a function of
animal abundance and size of site for a region of 2,500  2,500 m. A site size
of 50 m results in 2,500 total sites. The number of sites for a site size of 70,
100, 141, and 204 m is approximately 1,250, 625, 312, and 150, respectively.
The Journal of Wildlife Management
79(8)
Figure 5. Estimates of occupancy and site-level detectability for simulations
with no movement, 3 visits per site, and across a range of modeled
detectability (p) and number of animals present in the region. Points
represent mean of 1,000 simulations; confidence intervals are within symbol.

was approximately parallel (Fig. 6). Bias in occupancy

estimates occurred even for small amounts of movement; for
home ranges that were 10% of the area of the sampled site,
estimated occupancy was biased high by 22–24%. Home
range sizes that were equal to the site area resulted in
estimated occupancy 81–90% greater than that for immobile
animals.
Concurrent with increases in estimated occupancy,
estimated detectability decreased with increasing home
range size (Fig. 6). As expected, estimated detectability
was 100% when home range size was 0 across all values of M
but declined precipitously with increasing home range size.
Similar decreases in estimated detectability occurred across
all values of M (Fig. 6). As with estimated occupancy,
estimates of detectability were biased even for small amounts
of movement; detectability was only about 0.82 for a home
range size 10% of the site area. Estimated occupancy and
detectab6ility responded in opposite directions to movement,
and as such, were inversely related (Fig. 6). This relationship
was nonlinear, and depended on the number of animals
present (Fig. 6).
Scenario 4: Mobile Animals With Incomplete
Detectability
Results of these simulations demonstrated that differing
levels of detectability had little effect on bias in occupancy as
a function of home range size (Fig. 7), at least for a modest
population size of 300 animals. Estimated detection
probability for a home range size of 0 was unbiased (as in
Figure 6. Estimates of occupancy and site-level detectability for simulations
including movement but with complete detectability (P ¼ 1.0). Simulations
were conducted with 3 visits per site across a range of number of animals
present in the region (M). In the top panel, the true occupancy used in the
simulation was 0.039 for M ¼ 100, 0.077 for M ¼ 200, and 0.113 for
M ¼ 300. In the center panel, the true detectability was 1.0, and is
represented by a dashed line. Points represent mean of 1,000 simulations;
confidence intervals are within symbol.
scenario 2) but was increasingly biased low as home range
size increased (Fig. 7), with convergent estimates as home
range size reached 10 times the site size. Estimated
occupancy and estimated detectability were negatively
related in a nonlinear fashion (Fig. 7) that varied by the
level of detectability used for data generation. The number of
repeat visits to each site had less effect on estimates of
occupancy and detectability than home range size; however,
estimated occupancy was consistently higher as the number
of visits increased to 10 (Fig. 8).

Hayes and Monfils
Occupancy Modeling of Mobile Animals 1365


Figure 7. Estimates of occupancy and site-level detectability for simulations
with varying degrees of movement, expressed as home range size as a ratio to
the site area. Simulations were conducted with M ¼ 300 animals, and 3 visits
per site, across a range of detectability. In the top panel, the true occupancy
used in the simulation was 0.113 and is represented by a dashed line. In the
center panel, the true detectability (p) used is indicated in the legend. Points
represent mean of 1,000 simulations; confidence intervals are within symbol.
DISCUSSION
In bird point-count studies, the size of sites are generally
chosen by the investigator, but do not typically correspond to
the boundaries of individual animal’s home ranges,
constituting what Efford and Dawson (2012) term arbitrary
plots in continuous habitat. Researchers typically select site
sizes and sampling designs that address logistical and
funding constraints and ensure independence of sampling
units. Our results show that violations to the closure
assumption caused by temporary emigration introduce
substantial bias to estimates of occupancy and detectability
and that this bias is not reduced by increasing the number of
site visits. Bias in estimated occupancy and detectability
occurred for home ranges that are a small fraction (e.g., 10%
1366
Figure 8. Estimates of occupancy and site-level detectability for simulations
with varying numbers of site visits (T). We conducted simulations with
M ¼ 300 animals, across a range of home range sizes and with detectability
(p) ¼ 0.5. The true value of occupancy (0.113) and detectability (0.5) are
represented by dashed lines in both panels. Points represent mean of 1,000
simulations; confidence intervals are within symbol.
or 250 m2 [0.03 ha]) of the plot size used in our simulations
(50  50 m), which is likely to be much smaller than the
home range of a typical songbird or marsh bird. For example,
the ovenbird (Seiurus aurocapillus) is a readily detectable
species with estimated breeding territory sizes ranging from
0.20 ha to 1.80 ha in the Midwest and Ontario (Hann 1937,
Stenger 1958, Wenny 1989), which is double the typical plot
size of 0.78 ha (50-m circular plot). Although the amount of
bias depends on a number of factors, such as number of visits
and actual detectability, a home range size that is twice the
area of the plot can lead to estimates of occupancy that
are more than twice the true level. For marsh birds, which
were the impetus for this work, the home range size is likely
to be much larger. For example, Pickens and King (2013)
report that the home range of king rail (Rallus elegans), as
determined by radio telemetry, was between 0.80 ha and
32.8 ha. Moreover, they found that although home range size
did not vary by sex or year, it was negatively correlated with
the amount of open water. As such, estimates of detection
probability and occupancy could be confounded with habitat
features through the effect on home range size instead of
actual changes in the detectability of individual animals when
located within a plot.
An additional issue that arises is that the home range size of
birds typically varies seasonally and across habitats, potentially
leading to systematic bias in estimates of detectability and
occupancy across time or location. As such, the application of
The Journal of Wildlife Management
79(8)
occupancy models to bird point-count data has the potential to
lead to seriously biased results. The degree of bias will depend
on specific details of the biology of the species under
consideration and site size, and should be carefully considered
by the investigator. The examples above, however, suggest that
50-m-radius plots that are commonly used are small relative to
the home range of many common bird species. In other
applications, where the site boundaries coincide with natural
boundaries that relate to the biology of the organism (e.g.,
wetland anuran surveys), the assumptions of occupancy models
are more likely to hold, leading to reasonable estimates of
detectability and occupancy.
Our results are in contrast to some previous authors but
concur with others. MacKenzie et al. (2006) claim that “we
would thus expect the occupancy estimator to be unbiased if
species randomly moved in and out of a sampling unit”.
Likewise, Kendall and White (2009) suggest that mobile
species can mitigate some problems of subsampling locations
within a site. Consistent with Rota et al. (2009), we found
that the implication of assuming immobility, when animals
are in fact mobile, is that estimates of occupancy are biased
high and estimates of detectability are biased low, with the
degree of bias depending on the size of the animal’s range
relative to the size of the site used in the study.
We offer several reasons for the discrepancy between our
simulation results and previous analyses or simulations
concluding that violations to the closure assumption by
random movement results in minimal bias. Conceptualizing
the problem in an individual-based framework highlights
some of the differences; only 1 site can contain the core area
or focal point of a bird’s home range (as seen in Fig. 2), and all
other sites that are used have lower probabilities of use.
Because of this, all occupied sites do not have equal number
of detections, even when detection probability is 100%. As
such, simulations using an assumed level of occupancy across
a region, and simply perturbing that level of occupancy
downward to account for random periods when a bird
is absent misses a critical point, namely that mobile animals
create a situation where the occupancy status of adjacent
sites are not independent of one another. Similar to other
authors (e.g., Ramsey et al. 2015), we used a bivariate normal
distribution to represent animal movement within their
home range. Other statistical distributions are also plausible,
and would likely influence the degree to which estimates are
biased. The critical element, however, is that any movement
within the home range leads to some probability that animals
will not be present at a site when it assumed to be present,
leading to bias in detection probability.
Another way in which our simulation differs is how
detection probability is conceptualized and represented. We
simulated the detection probability for individual animals
(i.e., the probability an individual will be detected if it is
present), as constant across space and time, similar to Royle
and Nichols (2003), and aggregated detections of multiple
individuals into a single measure. In contrast, the usual
parameterization of occupancy models is that detection
probability at the site level is fixed regardless of the number of
individuals present. When a population is at low density, and
Hayes and Monfils
Occupancy Modeling of Mobile Animals
the probability of having more than 1 animal present within a
site is low, the 2 conceptualizations differ little in practical
terms. At higher densities, however, variation in the number
of animals per site can result in heterogeneity in site-level
detection probability without any variation in the detectabil-
ity of individual animals (Royle and Nichols 2003).
Some authors have alluded to the issue of animal
movement by distinguishing between use and occupancy
(e.g., MacKenzie and Royle 2005, MacKenzie et al. 2006,
Kendall and White 2009). Our reading and understanding of
this distinction is unclear; MacKenzie and Royle (2005) state
that “the proportion of area ‘used’ by a species will often be
larger than the proportion of area where species physically
occurs.” We fail to see the difference between physical
occurrence and usage. From the perspective of the data
entering occupancy models, the detection–no detection input
used by the models simply does not make a distinction
between animals that are occupying a site or animals that are
using a site. As such, the basic occupancy model does not
allow one to separate availability from detectability, as
defined by Amundson et al. (2014). Further, the closure
assumption in the basic occupancy model effectively implies
that availability is 100% throughout the survey period, and
that all non-detections at sites where there is at least 1
detection are due to a lack of perceptibility.
Although not the initial focus of our investigation, we
found that the effects of site size are critical in our
understanding of occupancy, and relate closely to the
mobility of the organisms under study. We show that for
a given number of animals (M) in a region, occupancy varies
widely for different site sizes. For example, a 50-m radius
plot has an area of 7,584 m2 and a 70-m radius plot has an
area nearly twice as much at 15,394 m2, and the occupancy
for a given number of animals would also be nearly double for
the 70-m radius plot. This highlights the sensitivity of
occupancy estimates to even relatively small differences in
sampling design, or to misestimates of actual distance from
the plot center point. Issues of plot size also are important
considerations when a broader region is subsampled as
Kendall and White (2009) discuss.
One design consideration to minimize the influence of
animal mobility would be to enlarge the size of the sampled
sites, thereby reducing the ratio of the home range area to the
size of sites (e.g., Rota et al. 2009). Three negative
consequences of such a decision would be 1) occupancy
would inherently increase as shown in our discussion of the
effects of site size; 2) larger sites are more difficult to
completely assess (Kendall and White 2009), so detectability
would likely decline for many species as costs of conducting
the study would increase; and 3) larger sites are likely to
encompass greater habitat heterogeneity, and thus, impor-
tant questions regarding the influence of site characteristics
on occupancy or use rates would likely be obscured.
Unfortunately, we do not have direct advice for addressing
the issues we raise if the basic occupancy model and data
collection methods are used. Several authors have proposed
methods for addressing violations to the closure assumption.
For example, Rota et al. (2009) recommend subdividing the
1367
sampling time to provide a robust design; this method
assumes closure within a visit to a site, which is clearly
violated when birds are seen entering or leaving the site
during sampling. Other methods include measuring dis-
tances to individual birds (Amundson et al. 2014), or using a
removal sampling method where individual birds are counted
and density is estimated (e.g., Chandler et al. 2011). These
methods hold promise for improving estimates of occupancy
and detectability for mobile animals, but they require the
collection of auxiliary data, and as such, do not provide a
means for addressing the lack of closure for commonly
applied methods of conducting bird point counts where such
data are not collected. Although investigators formulating
new surveys should consider the use of these emerging
methods, analysts confronted with historical surveys without
such auxiliary data are left with little recourse. Although we
agree that multiple visits to individual sites are of value in
addressing imperfect detection, and estimating detectability
is of importance when interpreting study results, the
incorporation of more biologically realistic assumptions
into the models used is of greater importance.
MANAGEMENT IMPLICATIONS
We show that commonly used occupancy models address 1
untenable assumption (i.e., complete detectability) by
making another equally untenable assumption (i.e., immo-
bility) when applied to bird point-count data, leading to
serious bias in estimated detection probability and occupan-
cy. Moreover, the meaning and interpretation of occupancy
itself is unclear in situations where the home range of the
study organism is larger than the size of the sample plot. As
such, we recommend against the application of the basic
occupancy model to bird point-count surveys where the only
data available are detection histories for individual sites.
ACKNOWLEDGMENTS
We thank G. Roloff for his helpful review of a draft of this
manuscript. We thank the U.S. Fish and Wildlife Service,
Michigan Natural Features Inventory, Michigan State
University Extension, Michigan State University Depart-
ment of Fisheries and Wildlife, and the Michigan
Department of Natural Resources for funding and time to
perform this research.
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Associate Editor: Michael Morrison.
The Journal of Wildlife Management
79(8)