Behavioral Ecology

doi:10.1093/beheco/arq032
Advance Access publication 15 March 2010

Determinants of false alarms in staging flocks of

semipalmated sandpipers
Guy Beauchamp
Faculty of Veterinary Medicine, University of Montreal, PO Box 5000, St-Hyacinthe, Quebec, Canada
J2S 7C6

nimals often interrupt their activities and flee to cover

upon detecting potential threats. Such threats, however,
often prove harmless and much time is thus spent fleeing at
the expense of fitness-enhancing activities such as feeding.
False alarms are a common feature in many species of birds
and mammals and account for a surprisingly large proportion
of alarms in some species (Hoogland 1981; Cresswell et al.
2000; Blumstein et al. 2004; Kahlert 2006). It is thought that
a high level of responsiveness to potential threats is a low-cost
strategy that reduces the likelihood of not responding
appropriately when a real attack does occur, the better-safethan-sorry principle.
For a single individual, the appropriate level of responsiveness should depend on the costs and benefits of responding or
not responding to potential threats whether they prove real or
harmless. The costs of responding include the energetic costs
imposed by fleeing and the foregone returns from any curtailed activity. Not responding upon detection of a potential
threat allows foragers to avoid these costs but at the risk of having to deal with a real predator. Not responding is also more
dangerous if threat detection is hampered. This can occur
when detection conditions are not ideal, for instance at low
light levels (Lima 1988), or in the presence of visual obstruction (Guillemain et al. 2001), or if harmless and real threats
share many features, making discrimination more difficult.
In a group, individuals can rely on their own detection as
described above but can also react to the responses of companions (Pulliam 1973). When relying on detection by companions, it is not always obvious to decide whether departures are
caused by a real alarm or are due to nonthreatening factors,
such as satiation for instance (Lima 1995). However, given

Address correspondence to G. Beauchamp. E-mail: guy.beauchamp

@umontreal.ca.
Received 10 September 2009; revised 17 December 2009; accepted
12 February 2010.

The Author 2010. Published by Oxford University Press on behalf of
the International Society for Behavioral Ecology. All rights reserved.
For permissions, please e-mail: journals.permissions@oxfordjournals.org

that multiple, simultaneous departures are more likely to represent independent responses to a real threat, individuals in
groups that have not detected the threat by themselves could
rely on these multiple departures to take appropriate action
(Lima 1994; Proctor et al. 2001; Beauchamp and Ruxton
2007b). Although the risk of not detecting a potential threat
is certainly lower in a group because many more eyes are
scanning the surroundings, false alarms can still occur in
a group given that the first few detectors are still relying on
their own potentially biased assessment of threats and nondetectors rely on this potentially biased information. In fact,
given that there are more potential detectors in a group, one
would predict that false alarms should actually increase with
group size because the odds of misclassifying a threat, at the
level of the group, increase with the number of independent
assessments of a potential threat (Treisman 1975). Not
responding in a group is also costly because individuals that
are left behind following the departure of companions may be
more at risk of attack by predators (Quinn and Cresswell 2005;
Beauchamp and Ruxton 2007a; Sirot and Touzalin 2009).
Surprisingly, few studies have examined the determinants of
false alarms in animal groups. In a study of foraging redshanks
(Tringa totanus), individuals were more likely to take flight in
alarm on cloudy days and later in the overwintering season
(Hilton et al. 1999). Clouds decrease light levels thus increasing the odds of misclassifying threats. Proneness to alarm later
in the season was interpreted as a low-risk strategy in birds that
are less likely to face starvation. In the same species, the ratio
of false alarms to real attacks decreased with the number of
real attacks and on rainy days as both are expected to increase
perceived predation risk. The effect of group size was not
investigated in this system.
One difficulty with an analysis of false alarms in foraging animals is that the foregone returns from interrupting foraging
are not negligible. Therefore, factors affecting the value of foraging are likely to influence the choice to respond or not to
potential threats making it more difficult to compare false

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False alarms occur when animals flee abruptly upon detection of a threat that subsequently proved harmless. False alarms are
common in many species of birds and mammals and account for a surprisingly high proportion of all alarms. False alarms are
expected to be more frequent in larger groups, where the odds of misclassifying threats are higher, and under environmental
conditions where detection of threats is compromised, such as low light levels. In addition, false alarms should be less frequent
when the energetic cost of fleeing increases. I examined these hypotheses in roosting flocks of staging semipalmated sandpipers
(Calidris pusilla) over 2 years. False alarms increased with group size but the effect of group size was confounded by the fact that
more attacks by falcons (Falco spp.) were directed at larger roosts. False alarms were more frequent at low light levels and later
during staging. As individuals double their body mass during staging, the energetic cost of fleeing must greatly increase thus
contributing to decreased responsiveness. A simple reduction in responsiveness caused by repeated exposures to harmless signals
would also produce a temporal decrease in responsiveness but this hypothesis cannot account for the effect of group size and
light level. Study of the determinants of false alarms provides an opportunity to examine adjustments in behavior in relation to
changes in perceived predation risk. Key words: false alarm, group size, predation risk, roosting, semipalmated sandpiper,
stopover ecology. [Behav Ecol 21:584587 (2010)]

Beauchamp

False alarms in staging flocks

585

observation duration in the model and forcing the parameter

estimate to be equal to one, which is what the offset does, it is
therefore possible to compare the number of false alarms even
when focal observations differ in duration. The independent
variables included year (2008 vs. 2009), stopover phenology
(early vs. late), log10 of roost size, the occurrence of rain, wind
velocity, air temperature, light intensity, and number of real
alarms. The final model was obtained through sequential removal of nonsignificant variables. Using information about
roost size at this and other sites in the Bay, I classified focal
observations as occurring early in staging (up to the 7th of
August, after peak arrival) or late. The proportion of birds
that have doubled their mass since arrival is low early in staging and higher later (Hicklin 1987).
The prevalence of attacks by avian predators was examined
in relation to the previous independent variables in a logistic
regression model. I used the prevalence of attacks, that is
whether or not a roost was attacked at all during the focal observation on a given day, rather than the actual number of
attacks since repeated attacks on a given day at the same roost
may not be independent events. I used cumulative time spent
in the roost as an offset.
I compared the number of false alarms at the same roost in
the first and in the last 10 min of roosting for all focal observations that lasted more than 20 min. I divided the number of
false alarms by the cumulative time spent roosting in
each period to calculate false alarm rate. I compared early
and late false alarm rates for matched data using a paired t-test.

MATERIALS AND METHODS

Data collection
The study was conducted from 28 July to 13 August in 2008 and
2009 at Daniels flats, New Brunswick, Canada (lat 45.73 N,
long 64.65 W). Daniels flats are located at the northern
end of Chignecto Bay in the upper Bay of Fundy. Tides, averaging 11.5 m in height, expose mudflats twice daily in the area.
Using a 20 3 60 scope, I monitored birds at a diurnal high-tide
roost located on a pebbly beach approximately 100-m away.
From my vantage point, I could easily count the number of
roosting birds and detect avian predators because I had an unobstructed field of view around the roost.
At the beginning of each focal observation, I counted the
number of birds either directly, when the roost was small, or
using local landmarks to determine the approximate size of
the roost and then multiplying the area by 100 birds per m2
(Mawhinney et al. 1993). I also noted time of day, air temperature (C), wind velocity (km/h), and light intensity (Lux)
obtained from handheld devices. I also noted the occurrence
of rain. Roosting birds frequently left the roost in alarm and
returned some time later at the same location. For each
alarm, I recorded the time when birds departed and returned
to the roost and assessed the cause of each alarm. Alarms were
characterized as real when an attack by an avian predator
(mostly Falco peregrinus and Falco columbarius but occasionally
Circus cyaneus) took place. When I could not detect the presence of a predator following an alarm, I classified the alarm as
false. I could not determine the exact cause of false alarms in
roosting flocks.
A focal observation ended when roost size changed due to
the arrival or departure of companions or when the birds
started feeding. A new focal observation started on the same
day if the newly formed flock subsequently reconvened to roost
at the same site. Changes in roost size in subsequent focal observations on any given day were substantial with up to 10-fold
changes in size. There were from 1 to 4 focal observations each
day but most days provided only one focal observation.

RESULTS
Roosts formed at the same location each day shortly after high
tide. Undisturbed birds started to leave the roost about 2-h
later to start feeding. Overall, attacks on roosting birds, primarily by falcons, occurred every other day and up to 4 attacks
occurred on any given day. Median focal observation duration
was around 15 min in 2008 and 30 min in 2009 (Table 1).
Roost sizes varied from 25 to 15 000. Approximately one false
alarm took place for every minute spent roosting (range 04),
and there were generally 5 false alarms for every real alarm
(Table 1). All birds left the roost during an alarm. After a false
alarm, birds returned to the roost about 30-s later. Birds
often abandoned the roost for the day after an attack. Descriptive statistics for the independent variables are presented
in Table 1 for each year.
Table 1
Median (range) of the various independent variables measured in
the 2 study years at Daniels flats, New Brunswick, Canada

Statistical analysis
For each focal observation, I calculated the cumulative time
spent in the roost (excluding flight duration). I used a negative
binomial regression model to examine the determinants of the
highly rightly skewed number of false alarms during each focal
observation. The cumulative time spent in the roost during a focal observation was used as an offset to control for different
duration of stay by the birds at the roost. By including focal

Variable

2008

2009

Number of focal
observations

20

19

Number of falcon
attacks per minute
roosting

0 (00.37)

0.01 (01.52)

Ratio false to real

alarma

4.5:1

6.75:1

Log group size

Wind velocity (km/h)
Temperature (C)
Light level (Lux)
Focal observation
duration (s)

3.24 (1.534.18)
2.48 (1.404)
4.9 (018)
11.3 (2.918.5)
18.9 (16.522)
21.3 (12.722.8)
10 000 (400018 000) 20 000 (440020 000)
978 (2052140)
1710 (3756600)

Calculated only when an attack took place (n 7 in 2008 and n 10

in 2009).

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alarms across conditions. A further difficulty is that false alarms

in a foraging context may be used deceptively by individuals to
usurp resources from others or to increase access to food
(Mller 1988; Kahlert 2006; Wheeler 2009). Here, I propose
an analysis of false alarms in resting animals where the only
cost of responding to a potential threat is the energetic cost of
fleeing. Deceptive use of alarm signals is unlikely in animals
that are not competing for any resources but simply resting. I
examine false alarms in staging semipalmated sandpiper
(Calidris pusilla) flocks roosting on the shore at high tide. In
particular, I examined the hypothesis that false alarms should
be more frequent in larger groups and under environmental
conditions that increase perceived predation risk, such as low
light levels. Sandpipers double their body mass during fall
staging (Hicklin 1987). This large increase in body mass will
undoubtedly increase the energetic cost of flying and I predicted that false alarms should occur less often with length of
stay in the staging area.

Behavioral Ecology

586

Table 2
Final negative binomial regression model of the number of false
alarms in staging flocks of semipalmated sandpipers controlling for
focal observation duration
b (SE)

Variable
Year
Phenology
Group size

P value

Relative change in the

number of false alarms

23.59 (1.59) 0.02

97% lower in 2009 than 2008
22.09 (0.44) ,0.0001 88% lower later than earlier
0.80 (0.24) 0.0007 123% higher per unit increase
in log10 of roost size
0.08 (0.31)

0.79

Group size in
2009

1.52 (0.41)

0.0002 357% higher per unit increase

in log10 of roost size

Light level

22.08 (0.93)

0.03

9% higher per unit increase

in log10 of roost size

88% lower per unit increase

in Lux

In the final negative binomial regression model, controlling

for focal observation duration, the number of false alarms was
statistically significantly lower in 2009 than in 2008, increased
with group size, decreased with light intensity, and decreased
later during staging (Table 2). The effect of group size differed
between the 2 years of study and was absent in 2008 (Table 2).
No other interactions were detected (P . 0.34) indicating that
the effect of light intensity was similar in the 2 years of study
and that the effect of group size and light intensity was similar
early and late in staging. The number of false alarms was not
related to temperature (b [standard error {SE}] 20.11 [0.09],
P 0.19), wind velocity (b [SE] 0.02 [0.03], P 0.55), the
occurrence of rain (b [SE] 20.28 [0.39], P 0.48), and the
number of attacks (b [SE] 0.28 [0.17], P 0.11).
The odds of attacks increased by a factor of 3.1 for each unit
increase in log group size (b [SE] 1.15 [0.53], P 0.03). This
effect was independent of year and staging phenology.
None of the other independent variables reached statistical
significance.
The number of false alarms per min spent roosting did not
differ statistically between the first 10 min of the beach
(median [range]: 0.59 [02.5]) and the last 10 min (0.43
[02.9]; t 20.47, P 0.64, n 18).
DISCUSSION
Despite a relatively low frequency of attacks by avian predators, staging semipalmated sandpipers interrupted roosting
and flew in alarm frequently. Most of these alarms turned
out to be false in the sense that no real danger was actually
present. False alarms were more frequent in larger roosts,
at low light intensity, and earlier in staging. I discuss these
results in turn.
The positive effect of group size on false alarm rate appears
at first to confirm the hypothesis that in a large group where
individuals share information about potential threat detection,
the sudden departures of a few detectors is sufficient to trigger
the departure of all remaining birds. Given that at the level
of the group, the odds of misclassifying a signal increase with
group size, false alarms were thus expected to be more frequent in larger groups. However, the effect of group size only
occurred in one of the 2 study years even though the range of
group sizes was similar between years. The nonsignificant effect
of group size occurred in the year with the highest frequency of
false alarms suggesting that the effect of group size may have
been trumped that year by a lower threshold for responding to
potential threats. In other avian studies, false alarm rate

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Group size in
2008

was not related to group size (Lindstrom 1989; Cresswell

et al. 2000).
A difficulty in the interpretation of the results is that larger
roosts also attracted more attacks. A similar finding was observed for foraging sandpipers (Beauchamp 2008; Sprague
et al. 2008) and has also been documented in many other
species (e.g., Lindstrom 1989; Cresswell 1994; Guillemain
et al. 2007). Models dealing with false alarms assume that
attack risk is independent of group size (Lima 1994; Proctor
et al. 2001; Beauchamp and Ruxton 2007b). The results thus
suggest that this assumption is not always met. If larger groups
are more at risk of attack, then more false alarms should be
expected in those groups even when sharing information
about predation threats, which is also expected to increase
false alarms in large groups (Giraldeau et al. 2002), does
not take place. Therefore, future work is needed to tease apart
the effect of overall predation risk from the effect of group
size on false alarms especially given that group size does not
seem to have a consistent effect on false alarms in the species
examined thus far.
Low light levels are thought to increase the odds of misclassifying threats and therefore false alarms should be more frequent under these conditions. In staging semipalmated
sandpipers, false alarms did indeed increase at low light levels.
Given that low light levels often occur early and late in the day,
false alarms are thus probably most frequent at these times of
day. These results are compatible with the observation that foraging sandpipers were more likely to leave a foraging patch
abruptly early and late in the day, controlling for food density
(Beauchamp and Ruxton 2008). Increased skittishness is
probably not a response to increased attack rate at low light
levels given that there was no relationship between time of
day and frequency of attack by falcons on roosting birds
and more generally on foraging birds in the same area
(Beauchamp 2008).
As staging progresses, sandpipers double their body mass
over the 1014 days individuals spend in the Bay during fall
staging. I found that false alarms were less common later during staging. The increased energetic cost of fleeing in fatter
birds should increase the response threshold to potential
threats and thus lead to fewer false alarms. The prevalence
of attacks at the roost did not vary with staging phenology suggesting that predation risk did not vary to a large extent over
the course of staging and was not directly responsible for
a change in the number of false alarms. Increased skittishness
has been documented in 2 species after exposure to predator
attacks (Hilton et al. 1999; Martn et al. 2009) but in staging
sandpipers the number of false alarms was not related to the
actual number of attacks.
An alternative hypothesis can account for decreased responsiveness as staging progresses. Birds may learn to become less
responsive to potential threats with increased exposure to
harmless signals (Bouskila and Blumstein 1992; Edelaar and
Wright 2006). However, the learning hypothesis does not
make any prediction related to group size and to light intensity, 2 factors which have been found to influence the number
of false alarms in this species, and thus cannot explain all the
present results. I found no effect of learning within days in the
sense that false alarms did not vary with time spent roosting,
but it may be the case that learning occurs between rather
than within days. The fact that roosting birds are mainly adults
at this time of year (Hicklin 1987) also means that many of the
sandpipers present at the roosts have been exposed to the
potential threats in the Bay at least once before thus making
the learning hypothesis less relevant. A more direct test of the
hypothesis would be to examine changes in false alarm rate in
wintering birds as the season progresses. Given that overwintering birds need not accumulate fat until closer to migration

Beauchamp

False alarms in staging flocks

time, a reduction in false alarm rate between days would only

be compatible with the learning hypothesis.
In conclusion, staging semipalmated sandpipers adjust their
level of responsiveness to potential threats in a dynamic fashion
depending on the costs and benefits of responding or not
responding to environmental signals. Study of the determinants of false alarms thus provides an opportunity to examine
adjustments in behavior in relation to changes in perceived
predation risk.
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