Animal Communication (Web Topic 14.1) Models of Environmental Signaling

Animal Communication: Web Topic 14.
CHAPTER 14
Chapter Outline
Summary
Web Topics
Topic 14.1
14. Environmental
EnvironmentalSignals
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LITERATURE CITED
Introduction
Nearly all of the interactions between individuals discussed in Chapter 14 can be
modeled with evolutionary game theory. It is instructive both for understanding the
specific interactions and for getting a broader feel for game theoretical modeling to
deconstruct the basic logic and outcomes of some of these models. We do not
cover all the models cited in Chapter 14: those applicable to the evolution of crypsis,
aposematism, and mimicry have been nicely reviewed elsewhere (Ruxton et al.
2004; Mappes et al. 2005; Skelhorn et al.
2010). However, the examples below
provide a broad sampling of ESS approaches. They are presented in the order
encountered in Chapter 14, and we classify the types of format used according to
the schema outlined in Web Topic 10.5.
Predator notification (Bergstrom and Lachmann

2001)
The Question: When does it pay for a prey animal to notify a predator that it has
been detected, and when does it pay for the predator to attend to such notification
signals?
Background: If predators were to attend to such signals, what would keep a prey
from giving the signal whether it has spotted a predator or not? In that case,
predators would do better to ignore the signal and then there would be no selection
for the prey to give it. There is clearly a conflict of interest between the two parties
and some honesty guarantee will be required for communication to
be favored. In
this model the proposed cost that insures honesty is the
risk that giving the signal
will reveal the presence of the prey to a predator that had not yet detected it.
Game Format:
The game is obviously asymmetric (prey versus predator).
The model is based on discrete alternative strategies (prey: signal or dont;
predator: chase prey or dont).
It is a single-shot model analyzed in extensive form with successive branch
points: nature decides whether a predator is near a prey or not, the prey then
decides whether to signal or not, and the predator (if present) then decides
whether to chase or not.
Payoffs at the terminus of each tree branch are fixed (a contest).
Special Assumptions: Only a single prey is considered; this excludes alarm
signaling to conspecifics and relative vulnerabilities of multiple prey as factors. The
model also excludes signals that indicate prey unsuitability for reasons other than
predator
detection (e.g., aposematism or relative escape abilities).
Synopsis:
http://sites.sinauer.com/animalcommunication2e/chapter14.01.html[10/16/2014 6:01:40 PM]
Animal Communication: Web Topic 14.1
Prey are exposed to stimuli that may or may not be generated by a nearby
predator. They then use Bayesian updating to combine prior probabilities of
predator presence with the conditional probabilities of
perceiving these stimuli
to generate an updated probability that a predator is present.
Prey can also estimate the risk of capture given the current context and likely
predator. The product of this risk and the updated probability that a predator
is present provides an overall measure of predation risk. Different stimuli and
different occasions result in different overall risks.
Emitting a predator detection signal costs a prey by increasing its
risk of
detection by other predators. Prey should only emit the signal if these signal
costs are less than the appropriately scaled overall risk of capture. For a fixed
average cost, this defines a threshold stimulus level below which no signals
should be emitted and above which they should.
The predators payoff depends on the probability of capturing a prey minus
appropriately scaled costs of a chase.
The analysis sought equilibrium (ESS) strategies in which prey only
signal if
stimuli exceed the threshold, and predators avoid chasing prey that so signal.
ESSs only exist if prey that estimate high overall risk (implying a predator is
almost surely present) can use this information to reduce their risk of capture.
That is, detecting a predator early in its hunt helps a prey to escape a
subsequent chase. A related requirement is that prey that are exposed to
stimuli below the threshold do not signal. If met, these conditions generate an
ESS in which emission of a predator detection signal is relatively honest and
thus valuable to both parties.
Two specific contexts were considered: in one case, predators always chased
prey that did not signal. In the other, predators did not pursue prey until at
least some signaled so that they could focus on the
ones that did not.
Conclusions:
The analysis for both contexts showed that predator detection signals are only likely
to be honest if there is a cost to senders (in this case, the risk of attracting other
predators). In addition, the benefit to prey of predator detection signals only accrue
if a prey is relatively sure that a predator is present, and if this knowledge improves
the preys chances of escaping if chased. This means that
animals that cannot
reduce their risks even if they detect a predator early (e.g., sessile or poorly mobile
species, animals far from refuges,
etc.) should not bother to give predator detection
signals. The model also predicts that the threshold for giving predator detection
signals should decrease when predators are more common, and the minimal cost
that guarantees sender honesty can be smaller if the costs of pursuit to

predators
are higher.
Related Models:
Getty (2002) expanded on this model by noting that the prey face a signal
detection problem in which each seeks to minimize costly false alarms; at the
same time, the predator faces an optimal diet problem. Getty computed the
relevant red line that the prey should adopt along the predator stimulus axis
given a signal detection analysis. He also considers what happens when
predators have alternative
prey to pursue.
This model only considered a single prey animal. When multiple prey
are
present, a variety of consequences follow. Those relating to relative
vulnerabilities are outlined in the models presented below. However, the
simplest change is that instead of giving predator notification signals
independently, a group of prey does so in concert. This usually takes the form
of group predator inspection behavior. A relevant model for this tactic can be
found in Dugatkin and Godin (1992).
A number of other models consider predator notification signals for
other
reasons. Readers may want to compare the above model to those in which
prey signal to predators honest indications of their physical condition,

stamina, palatability, or agility (Nur and Hasson 1984; Hasson 1991; VegaRedondo and Hasson 1993; Maynard Smith and Harper 2003; Ruxton et al.
2004; Searcy and Nowicki 2005).
Vigilance behaviors (Sirot and Touzalin 2009)

The Question: What is the ESS level of vigilance in a
foraging group if vigilant
individuals are more likely to survive a predator attack than foraging individuals?
Background: Prey that are vigilant when a predator appears or attacks can often
escape sooner and more effectively than prey that are foraging. This is because
vigilant prey spot a predator first, know where it is, and thus know which direction
would move them away from it. Foragers may only flee after vigilant animals have
fled or
given alarms, and then have no idea where the predator is. Given these
differences, the risk to foragers increases as more and more members of their
group become vigilant; the risk to vigilant animals decreases as more and more
group members forage. The best strategy for escaping predation is always to be
vigilant; however, the animal will then die of
starvation. Is there an average level of
vigilance which if adopted by all in the group optimizes the tradeoff between
predation and starvation
risks?
Game Format:
The strategy set is the fraction of time an individual spends being vigilant
instead of foraging. It is thus continuous.
The game is a scramble because the payoff of adopting a given strategy
depends on the mix of strategies adopted by a defined circle of
neighbors.
Although relative location in a group may affect optimal strategy, all individuals
in the same neighborhood begin the game with otherwise similar properties.
The game is thus symmetric.
Special Assumptions: The game is spatially explicit;
payoffs depend on
abundances of different strategies adopted by local neighbors of a focal animal, not
by the population as a whole. The game is played on cellular automoton grids such
that each animal (at a node) has a fixed number of surrounding neighbors with
which it shares predator detection ranges, alarms, and attack vulnerabilities. No
coordination of vigilance based on mutualism, reciprocity, kin selection, or other

cooperative economics is included in this game. Each
animal just does what is best
for itself (although noting what neighbors are doing is a key part of that process).
Synopsis:
Prey animals need to acquire new energy through foraging and avoid being
killed by predators. Vigilance is assumed to be incompatible with foraging and
vice versa. There is thus a tradeoff in how to allocate actions.
The payoff currency in this model is the amount of energy reserves attained at
the end of the game. This depends on how fast energy can be obtained when
foraging and lost when not foraging, and the fraction of time that was spent
foraging instead of being vigilant. The potential final reserves are then
discounted by the probability of having survived
all predator attacks.
Mortality risks due to a predators attack are varied along two axes. The first
axis is the ratio between vigilant and foraging animal vulnerabilities: at one
extreme, vigilant animals are never killed if there is at least one foraging
animal in an attacked group; at
the other extreme, vigilant and foraging group
members have identical risks. The second axis is related to who, if anyone,
first spots an approaching predator: risk is assumed to be least for an animal
spotting
and responding to an approaching predator, somewhat higher for
animals who do not first spot the predator but are secondarily alerted by
another members predator detection and response, and highest if nobody in
the group spots the approaching predator.

Two models are considered. The first (static) model looked for the ESS
average fraction of time that each individual in the group should be
vigilant
given its position in the group, its initial energy reserves, and the current
probability of predator attacks. The second (dynamic) model acknowledges
that different animals may elect to be vigilant at different times, thus making
the number of vigilant neighbors a stochastic variable. The second model
allows a focal animal to track these unpredictable variations and adjust its
own vigilance accordingly.
The static model sought an overall ESS in which each group member adopted
an average fraction of time spent vigilant that was optimal given their position
in the group and the fractions adopted by other members in the group. At this
mix of fractions, nobody benefited by adopting a different average fraction.
These equilibria could not be identified analytically, but were identified for
each set of initial conditions using asymptotic iterations. Of major interest was
how ESS mixtures of fractions changed depending upon initial reserves at the
start of the game, group size, and different relative vulnerabilities for foraging
and vigilant group members.
The second model focused on a fixed number of relevant neighbors (12) and
a fixed value for initial reserves, and played the cellular automaton game on a
torus to eliminate world boundary effects. The optimal (ESS) fraction of time
vigilant was computed using the equations
for the first model and plotted
against successively greater fractions of the group being vigilant. Equilibria
occurred whenever the ESS fraction for an individual equaled the fraction of
the group that was currently vigilant (using a method called rational reaction
sets
[Simaan and Cruz 1973]). Equilibria were stable if a slightly higher
fraction of vigilant neighbors benefitted from a lower optimal (ESS) vigilance,

and a slightly lower fraction benefitted from a higher optimal vigilance. Any
drift away from a stable equilibrium would thus trigger individual adjustments
back to the equilibrium point.
Even if all group members adopted the same optimal fraction of time
vigilant,
they might or might not synchronize their vigilance with that
of neighbors.
Simulations of the dynamic model were used to measure levels of spatial and
temporal autocorrelation in vigilance for different conditions.
Conclusions:
The static model predicted greater vigilance fractions for animals located on
the outer edge of a group (since they had fewer neighbors to share the risk),
for all neighbors given greater initial energy reserves
(since they did not need
to feed as much), when foragers were at higher
risk than vigilant animals
(since nobody wanted to be the straggler if attacked), and in contexts where
predator attacks were more common.
The dynamic model made different predictions depending upon: a) the
relative
vulnerability of foragers and vigilant animals, and b) the differences in risk
when at least one neighbor spotted the predator versus when nobody did:
Vigilant and foraging animals suffer similar capture probabilities, but
detection of an approaching predator significantly reduces everyones
risk. Here, an individuals optimal vigilance decreased to an asymptote
as the number of vigilant neighbors increased. An intermediate level of
vigilance was the stable optimum. Instantaneous snap shots of such
populations showed little spatial or temporal synchrony in being
vigilant. An individual animals timing of activity was little affected by
that of its neighbors.
Foraging animals suffer higher risks of being killed than vigilant animals,
and detection of an approaching predator significantly
reduces risk.
High levels of vigilance were optimal when few neighbors were vigilant
since this increased chances of the few vigilant
animals spotting the
predator and alerting all. High levels of vigilance were also favored
when most group members were vigilant since being one of the few
foraging animals greatly increased risk. When intermediate numbers of
neighbors were vigilant, there were enough eyes and ears at work to
lower ones own vigilance, and enough other foragers around to reduce
chances of being the only suitable prey if attacked. Stable equilibria
occurred at a somewhat less than intermediate fraction of time being
vigilant, and another when everyone was vigilant. Instantaneous
snapshots of the population showed dense patches of synchronously
vigilant animals (since nobody wanted to be the
sole vulnerable
forager) but no similar contagion of foraging.
Foraging animals suffer higher risks than vigilant animals of
being killed,
but early detection of a predator has little effect on anyones risk. The
only relevant factor here was the greater risk of being killed while
foraging when most of the neighbors were being vigilant. A stable ESS
occurred at a low average vigilance fraction. These conditions
generated contagious patches of synchronous vigilance as in the prior
case, but also contagious patches of foraging.
See Figure 14.13 in the text for graphic comparisons of the three conditions.
Related Models:
This model built on the results of a number of prior publications on vigilance in
groups of animals including Pulliam et al. 1982; Lima 1987; McNamara and
Houston 1992; Lima 1995b, a; Lima and Zollner 1996; Bednekoff and Lima
1998b, a; Bahr and Bekoff 1999; Hilton et al. 1999; Lima and Bednekoff 1999;
Beauchamp 2007; Pays et al. 2007a; Pays et al. 2007b. Readers may want to
consult these earlier treatments after having
worked through this one.
There are several parallel models examining the degree to which animals
should coordinate vigilance by observing which neighbors are vigilant
(Rodriguez-Girones and Vasquez 2002; Fernandez-Juricic et al. 2004;
Jackson and Ruxton 2006).
Sirot and Pays (2011) recently published a paper using similar logic to identify
the optimal amount of time a solitary forager should devote to scanning for
predators.
Sentinel behaviors (Bednekoff 1997)

The Question: Why and when should an animal in a foraging group act as a
sentinel?
Background: It is not initially obvious why any animal in a group would give up
foraging and take on sentinel duty to the benefit of other group members. One
answer is that, as suggested by the previous model, being a sentinel is often safer
than being a forager. Also following the logic of the previous model, being a sentinel
all the time would then be the safest anti-predator strategy, but the consequence
would be death by starvation. Is there an optimal mix of foraging and sentinel duty
and how might this mix vary depending upon the way in which sentinels and
foragers do or do not alert each other when a predator is spotted?
Game Format:
The strategy set is discrete: individuals can either act as a sentinel or forage.
The game is a dynamic one: it consists of many successive bouts, during
each of which each player decides to act as a forager or a sentinel for that
bout. The goal is to identify the optimal choreography
or policy to follow over
a long series of bouts.
All group members start out with similar resources so the game is symmetric.
The payoffs are discrete: either an animal survived the series of bouts or it did
not. It could die during any bout as a result of either predation or starvation.
Because the outcome of any bout depends not only on a focal players actions
in that bout but also on the fractions of the foraging group that have adopted
forager and sentinel roles, the game is
a scramble.
Special Assumptions: Reciprocity and kin considerations are excluded from this
model. By-product mutualism (in which doing what is best for oneself incidentally
helps others) does come into play.
Synopsis:
The model assumes that sentinels are less likely to be killed during a predator
attack than are foragers. The ratio of relative vulnerabilities is a critical
variable in the model.
ESS policies for a given set of conditions were derived using both forward and
backward induction methods (see Web Topic 10.5 and Mangel and Clark
(1988) for background).
ESS policies were independently derived for different patterns of information
sharing. Three cases were considered: a) no sharing:
detection of a predator
by either a sentinel or forager never alerts other group members, either
because the detector fails to give an alarm,
or because its flight is not
detectable by others; b) sentinel sharing: detection of a predator by sentinels
but not foragers alerts other group members; and c) full sharing: detection of
a predator by either sentinels or foragers alerts other group members. Being
alerted reduces predation risk.
The median and distribution of fractions of the group acting as sentinels were
computed for each ESS policy derived.
The initial model assumed a group size of five animals, four-fold lower
vulnerability to attacks for sentinels than for foragers, and 90% probability that
foragers would fail to detect approaching predators on their own. These
values were later varied in a sensitivity analysis to determine the robustness
of the initial results.
Conclusions:
No information sharing: Here, predation risk for either sentinels or foragers
increased as more of the individuals in a group became sentinels. This is
because the risk to remaining foragers increases as they become more rare
(same outcome as vigilance model of Sirot and Touzalin [2009] above), and
once foragers are sufficiently rare, being a sentinel becomes just as risky and
it no longer pays to be
a sentinel. As a consequence, the threshold in energy
reserve that was required before a forager switched to being a sentinel
decreased as the number of sentinels in the group increased. The distribution
in number of sentinels over time was highly variable with a mode at 0
sentinels. In short, sentinel behavior is unlikely to pay, even if sentinels have
lower vulnerabilities than foragers, when there is no information sharing.

Sentinel sharing and full sharing: These two cases had very similar outcomes.
Predation risk when no sentinels were present decreased dramatically to a
very low value when one sentinel was present; adding additional sentinels
further reduced risk for either foragers or sentinels but at a much decelerated
rate. As a result, the threshold energy reserves required before an animal
switched from foraging to being a sentinel rose dramatically once one other
group member became a sentinel and increased only slightly when higher
fractions acted as sentinels. Observed distributions of numbers of sentinels in

multiple simulations showed a significantly narrow peak at one sentinel per
group. Sentinel behavior can thus be favored as long as
information is shared
in some way.
Alternation: When information is shared, a threshold ratio between sentinel
and forager vulnerabilities can be determined at which a well-fed animal could
do equivalently well as a sentinel or forager. This sets the scene for
alternation in which different individuals take successive turns in the sentinel

role (a common observation in natural groups). Given the other parameters
examined in these models, this threshold was always met if the conditions
favoring being a sentinel at all were met.
Robustness of results: The optimal fraction of the group that should act as
sentinels was independent of group size. Reduced mean
food intake when
foraging or greater unpredictability of food intake reduced the ESS fraction of
the group serving as sentinels. Variation in
other model parameters such as
predator attack rates, failure of foragers to detect approaching predators, and
initial energy reserves had only minor effects on predictions.
See Figure 14.14 in the text for graphic representation of some of these
results.
Related Models:
This author developed the same model further, including the special, but
common, case of pairs of animals in Bednekoff (2001).
The literature citing or building on this paper overlaps extensively with those
papers cited for the prior game model (Sirot and Tourazin 2009).
A large number of field studies have since examined the assumptions
and
predictions of this model. Citations can be found in the main text.
Some authors have extended the information sharing aspect of sentinel
behavior by noting that many sentinels emit repeated calls while on duty. The
evidence that this enhances forager efficiency versus
has some alternative
function remains conflicting (Bednekoff et al. 2008; Hollen et al. 2008; Ellis
2009).
False alarm call rates (Beauchamp and Ruxton 2007)

The Question: Why and how often should members of a foraging group attend to
false alarms?
Background: It is well known that foraging groups of
some species experience
frequent false predator alarms. Some false alarms are simply due to mistaken
stimulus interpretation, but others may be intentional manipulations by some
members of a group. Given the relative risks of predator attack and starvation, what
is the optimal level of false alarms that can be tolerated by a foraging group, and are
there other modulators of responses to false alarms (in particular, how

many group
members have taken flight) that alarm responders should consider?
Game Format:
The strategy set was discrete: individuals could either forage or be vigilant.
Individuals did not begin the game identically, but because their parameters
were drawn from the same random distributions, the game was stochastically
symmetric.
Payoffs were 0 if an individual starved or was killed by a predator, and equal
to its accumulated energy reserves if a survivor. Because both factors could
vary depending on what others in the group did, this game was a scramble.
Like the prior example, the game was dynamic: within a generation, each
individuals lifetime was divided into many successive bouts within each of
which it could choose to be vigilant or forage.
Special Assumptions: As with prior examples, kin selection and reciprocity were
not allowed in the models.
Synopsis:
As in prior examples, individuals were assigned to groups. Different group
sizes were examined but held constant for a given analysis.

Vigilant animals never erred in detecting an approaching predator or by fleeing
or alarm signaling to innocuous stimuli (i.e., they never gave false alarms).
Foragers however, could err in both ways. They could
thus generate false
alarms.
Each individual was haploid and carried three genes on a single
chromosome: one specified the probability that the individual would be
vigilant at any given time, one specified the probability that the individual
would flee if it heard one group members alarm or saw one group member
fleeing, and the third gene
specified the probability that the individual would
flee when it saw two or more group members fleeing. A model run began by
assigning random
values to each individuals genes.
Unlike the prior example, evolution was then allowed to proceed through many
successive generations. At the end of each generation, its members were
ranked according to individual payoffs. The next generation
was created by:
a) replicating the top half of the ranks in the ending generation, and b)
replacing the lower half of the prior generation by random choices of
individuals from the top half of the rankings. Mutations were then added to
each gene at a low rate.
After a sufficient number of generations, each of the three gene loci tended to
converge on a stable distribution with a distinct mode and median. The
combination of the three asymptotic medians for a given set of initial
conditions defined an ESS. This type of multi-generational analysis is called
genetic algorithm modeling (Vrugt and Robinson 2007).
ESS mixes were identified for a variety of different ambient conditions.
Sensitivity analyses were then run to evaluate the robustness of these
outcomes.
Conclusions:
The median value of the vigilance gene decreased as group size was
increased, and also when the rate at which foragers misclassified stimuli was
increased.
The median value of the gene controlling the probability of fleeing
after two
group members fled was always higher than that for the probability of fleeing
after only one members flight. The ESS thus favored reliance on more than
one alarmed individual before fleeing.
The median value for both genes controlling the probability of flight decreased
as group size increased, but the reduction was much smaller for the gene
controlling flight after multiple alarms. Animals tend to rely on multiple flights
or alarms at about the same rate regardless of group size or rates of forager
misclassification of stimuli.
The average fraction of false alarms at the ESS was 20% for a group
of six
but as high as 55% for a group of 20 individuals. Larger groups thus tolerated
surprisingly high rates of false alarms.
Related Models:
Lima (1994) and Proctor et al. (2001) consider similar questions, but include a
number of constraints that are relaxed in this model.
Pollard (2011) models the degree to which identifying which
group members
produce the alerts might reduce false alarms, Bell et al.
(2009) review
refinement of alarm responses when animals integrate multiple sources of
information such as recent surveillance outcomes, and Thompson and Hare
(2010) show that where successive alarms are emitted can be used to predict
an approaching predators trajectory.
Flower marking by bees (Stout and Goulson 2002)

Question: Why and when should foraging bees avoid nectar collection at a flower
marked by an earlier forager?
Background: Bees visiting flowers remove most of the
available nectar per visit. The
flower then begins replenishing the nectar at a species-specific rate. The bee will
obtain the largest subsequent nectar load if it postpones a return visit until the
depleted
flower has sufficiently replenished its nectar. Many bee species mark
flowers with complex pheromone footprints whose components volatilize at

different
rates (Goulson et al. 2000). A bee can then monitor the changing composition of a
footprint to determine how long a flower has had to replenish. Since different flower
species have different nectar replenishment rates, optimal return times will differ for
different species of flowers. Bumblebees have been shown to adjust return rates
according to flower species. One cost of the time mark system is that other bees,
even other species, detect and monitor the marks. A cheater bee can drain a
flower before the marking
bee would normally return. The cheater only gains a
partial load, but they at least get this nectar before any competitor does. The
resulting arms race could, in principle, undermine the entire marking strategy by
forcing competitors to make earlier and earlier visits to marked flowers. For a given
flower replenishment rate and density of competitors, is there an ESS that
maximizes individual rates of nectar acquisition but preserves the flower marking
strategy?
Game format:
The strategy set is the estimated time since a flower was last visited above
which a bee will collect the nectar and below which it will reject the flower for
now. The set is thus continuous.
The payoff to be maximized is the rate at which nectar is acquired per unit
time.
All bees have equal access to the same set of flowers and the game is thus
symmetric.
Because the nectar return for a bee adopting a given return time depends on
the return times adopted by other bees foraging in the same neighborhood,
the game is a scramble.
Special Assumptions: The authors further assumed that:
The time required to remove most of a flowers nectar (the handling time) was
independent of the amount of nectar being removed.
The bees always removed all the available nectar per visit.
The energetic costs of flight and handling were similar enough to use time
spent as a common currency.
The interacting bees were unrelated excluding kin cooperation from playing a
role.
Synopsis:
The average rate of nectar acquisition was computed based on the search
and handling times per flower, the relative fraction of flowers that were
currently acceptable (above threshold nectar replenishment), and the average
nectar load obtainable from those flowers.
It was assumed that there were enough bees and a finite number of flowers
such that all flowers in the neighborhood had been visited recently. New
flower production was thus assumed to be negligible over short time periods.
The fraction of flowers that were acceptable thus depended on the rate of
replenishment and return visit times used by the
average bee.
If all bees cooperated by observing the same return time, a pareto optimal
time could be computed that maximized individual forager intakes
for a given
flower species. No bee could do better without reducing the
intake of others.
Assuming most bees had adopted the pareto return time, the rate of nectar
accrual was then computed for a single mutant bee that visited flowers
randomly without regard to return time marks. Using
observed values for the
relevant parameters, the rate of nectar accrual
given random visits was
usually higher than that for bees respecting the pareto rate of return. This
meant that there was ample incentive for
some bees to cheat by visiting
flowers earlier than that proscribed by the pareto strategy.
An equilibrium occurs when bees using chemical marks to select or reject
flowers adopt a return time that yields a nectar accrual rate equal to that
enjoyed by bees that visit flowers randomly. If the bees relying on marking
were to use a longer time, cheaters would be favored and increase in
numbers; if cheaters were to use a shorter period, bees that relied on marks
would do better. The equilibrium is thus stable and
an ESS.
The study then compared the actual return times of wild bumblebees to the
pareto optimal and ESS predicted values.
Conclusions:
As expected, average return times to different flower species by wild
bumblebees were inversely related to the rate of nectar replenishment.
For several combinations of bumble species and flower species, particularly
flowers with rapid replenishment times, the pareto return time was
considerably longer than the ESS time. Where this was the case,
the average
return time exhibited by the wild bumblebees was much closer to the ESS
value than to the pareto value.
Where the pareto and ESS return times were very similar, the wild
bumblebees exhibited average return times very close to the shared value
of
both predictions.
Related Models:
No related models since this publication were found. However, there remains much
debate over whether the footprint marks of foraging bees are signals actively
deposited on flowers to allow replenishment time monitoring or instead are

inadvertent cues. Relevant citations are given in the text.
Literature cited
Bahr, D.B. and M. Bekoff. 1999. Predicting flock vigilance from simple passerine
interactions: modelling with cellular automata. Animal Behaviour 58: 831839.
Beauchamp, G. 2007. Vigilance in a selfish herd. Animal Behaviour 73: 449472.
Beauchamp, G. and G.D. Ruxton. 2007. False alarms and the evolution of
antipredator vigilance. Animal Behaviour 74: 11991206.
Bednekoff, P.A. 1997. Mutualism among safe, selfish sentinels: A dynamic game.
The American Naturalist 150: 373392.
Bednekoff, P.A. 2001. Coordination of safe, selfish sentinels based on mutual

benefits. Annales Zoologici Fennici 38: 514.
Bednekoff, P.A., R. Bowman and G.E. Woolfenden. 2008. Do conversational

gutturals help florida scrub-jays coordinate their sentinel behavior? Ethology 114:
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