Animal Behaviour Advantages No 1

ANIMAL
BEHAVIOUR:
ADVANTAGES AND
DISADVANTAGES
NO.1
Kevin Brewer
ISBN: 978-1-904542-36-0
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Kevin Brewer BSocSc, MSc

(http://kmbpsychology.jottit.com/)
An independent academic psychologist, based in England,

who has written extensively on different areas of
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Animal Behaviour: Advantages and Disadvantages No.1; Kevin Brewer; 2008 2

CONTENTS
Page Number
1. Advantages and Disadvantages of

Animal Migration 4

Ambush Hunting 19

Caching Behaviour 25

Territoriality 35

Delayed Breeding in Birds 47

1. ADVANTAGES AND DISADVANTAGES OF ANIMAL
MIGRATION
Migration refers to "regular movements between areas
uninhabited at different times of the year" (Cocker 1993)
or "long-distance travel, usually with a return, to
specific locations" (Grier and Burk 1997).
Migration is triggered by seasonal changes in
weather, air temperature or day length, or changing food
supply. For example, wildebeest move towards rain in the
dry season. Some animals move from one food source to
another, while others migrate to particular breeding
areas.
Most interest relates to bird migration, but other

animals also migrate through water or over land (table
1.1). It has been estimated that 5 000 million birds
migrate southwards across the Mediterranean area to
trans-Saharan Africa each year (Moreau 1972).
COMMON NAME SCIENTIFIC NAME WHERE
Caribou Rangifer tarandus North America
Wildebeest Connochaetes taurinus Africa
Saiga antelope Saiga tatarica (figure 1.1) Central Asia
Mongolian gazelle Procapra gutturosa Mongolia
White-tailed deer Odocoileus virginianus North America
Table 1.1 - Examples of migrating mammals.
(Source: US Federal Government)
Figure 1.1 - Saiga antelope.

There are advantages and disadvantages to migration
generally (table 1.2) as well as in relation to migration
for breeding or migration for food (table 1.3).
ADVANTAGES
1. Return to specialist site for breeding that does not need all year
round food supply, and often no (or few) predators.
2. Move to where food/prey available when not breeding, particularly

with young (ie: maximise feeding opportunity).
3. Stationary can mean increased predator risk.
4. Constant temperature conditions: escape bad weather and lower

temperatures (and greater risk of death), especially to give birth.
5. Able to have specialist breeding site (eg: no predators) and

another site for feeding.
6. Flexible strategy - some members of the species can migrate and

others not depending on where live.
7. Stationary animals risk exhausting food supply using it all year

round, particularly if competition from other species.
8. Opportunity for different members of the species to meet, and

greater breeding variety (eg: storks from western and eastern Europe;
figure 1.2).
9. Ideal when specialist food required because the earth's resources

are not evenly distributed. Most land mass and shallow seawater in
the northern hemisphere; eg: Arctic tern (Sterna paradisea) feed on
whitebait in shallow waters, and European swallow (Hirundo rustica)
on flying insects on land (Whitfield et al 1981).
10. Birds migrating at night usually safe from predators as few day-
time birds of prey adapt to night-time hunting.
DISADVANTAGES
1. Large amount of energy required to travel long distances.
2. Problems and risks of navigation.
3. Risk of forgetting sites or not being able to find again.
4. Leave home territory empty allowing for invaders, and then fights
on returning.
5. Risk at temporary stopovers from lack of local knowledge about

predators.
6. Vulnerable to weather changes or poor conditions in one year.
7. Many decisions required including optimal fuel load and optimal

time of departure.
8. Other risks like the change from salt to freshwater or vice versa
for some fish.

9. Evolutionary maladaptive behaviour in some cases; eg: green
turtles (Chelonia mydas)(figure 1.3) feed on eastern coast of South
America but breed on Ascension Island (south Atlantic); adequate
breeding sites nearer to feeding areas (Whitfield et al 1981).
10. Risks of night-time migration if animals normally active in day-

time (eg: bat predation of birds).
Table 1.2 - Advantages and disadvantages of migration.
(Source: Bamse)
Figure 1.2 - Migration routes of white stork.
(Source: US Fish and Wildlife Service)
Figure 1.3 - Green turtle.

Migration for breeding - 1, 5, 8
Migration for feeding - 2, 7, 9
Table 1.3 - Specific advantages in table 1.2 depending

upon type of migration.
DISTANCE
The length of migration varies from 1-2 metres per

day by zooplankton to thousands of miles by some birds.
For example, an Arctic tern, which was ringed, was found
to fly from Russia to the Antarctic (9000 miles)(Grier
and Burk 1992)(figure 1.4). Monarch butterflies fly up to
2000 miles between USA/Canada and Mexico.
(Source: Andreas Trepte)
Figure 1.4 - Migration routes of Arctic tern.
Studies of ringed populations of birds (figure 1.5)

found in different places is one way to study the
migration distances. For example, Black-browed
albatrosses (Diomedea melanophris) from the Falkland
Islands (south Atlantic) have been discovered in South
America mainly, but ringed birds from South Georgia
(south Atlantic) mainly found in southern Africa (quoted
in Cramp and Simmons 1977).

(Source: Julio Reis)
Figure 1.5 - The ringing process by "Cruzinha"

(national bird ringing authority) in Portugal.
There are cases from displacement experiments of

albatrosses migrating very long distances. Displacement
experiments involve taking birds to a point of release
that is different to their normal migration point, and to
see if they return to the original home. The birds are
ringed so records can be kept. For example, Laysan
albatrosses returned 5200km in 10-12 days from the west
coast of the USA to Midway Island (Pacific), or from the
Philippines to Midway Island (6500km taking one month)
(Alerstam 1993).
At the other extreme, the common toad (Bufo bufo)

moves a few hundred yards from vegetation to pond to
spawn.
VARIATIONS IN PATTERNS
Within some species certain individuals migrate and

others do not (eg: blackbirds found further north do).
There can also be differences between the sexes (eg:
chaffinch males more migratory than females).

Robins (Erithacus rubecula)(figure 1.6) are partially
migratory. This means that some "races" and some birds
within those "races" migrate, while others are sedentary.
The robins of Britain and Ireland that do migrate go
South South-West to Iberia (Spain/Portugal). This occurs
from September-October through to March-April (Roselaar
1988). Most migration occurs at night (Elphick 1995).
There are sex differences in migration among robins.

Males are more sedentary than females generally. Table
1.4 lists the estimates of the number of robins who were
sedentary in four different studies.
STUDY MALE FEMALE
Enniskillen (1926) 75 36
Devon (1965) 70 33
Cambridge (1984) 77 30
Oxford (1988) 69 3
(After Harper 1988)
Table 1.4 - Percentage of robins who are sedentary by sex

in four different studies.
(Source: Ramin Nakisa)
Figure 1.6 - Robin.
Whether the birds migrate or not depends on the

local conditions, like unpredictable winters. There are
three stages to the evolution of migratory behaviour in a

species (Gill 1995):
i) Partial migration - Some of the species migrates while

others do not;
ii) Division - Species separated into two clear

groupings: migratory and non-migratory;
iii) Natural selection leads to the elimination of one

group.
As a species, robins are probably between stages 1

and 2 above. Birds build up their body mass in late
summer to coincide with the "zugunruhe" behaviour
(migratory restlessness) (Biebach 1983).
Experiments with robins in steel cages with magnets

attached have found that they are influenced by magnetic
cues (Merkel and Wiltschko 1965 quoted in Gill 1995).
INSTINCT AND LEARNING
Though certain animals may have a "migratory

instinct", it is modified with learning in terms of how
often to migrate, and the direction travelled.
Perdeck (1958) took starlings, which usually

migrated from northern Europe in a south-west direction
to southern Europe, to Switzerland for release. Juvenile
birds flew south-west, but adults adjusted and flew to
their normal destination.
MIGRATION DECISIONS
Migration involves a number of decisions to produce

the optimal mating strategy:
1. Optimal fuel load
How much stored fat, protein and food stored in the

digestive tract before beginning. Too much slows the
animal down and reduces the ability to escape chasing
predators. Too little increases the risk of starvation
during the migration. A range equation gives "the
potential migration distance in relation to fuel load"
(Hedenstrom 2003a).
The ideal for birds should be short steps and small
fuel loads compared to large distances and large fuel
loads (Alerstam 2001).

2. Number and duration of stopovers (optimal stopover
duration)
Each stopover risks local predators and taking

longer to reach destination against the need for fuel.
Taking longer may mean missing the best breeding sites or
opportunities.
Hedenstrom (2003b) calculated the proportion of time
spent on travelling against refuelling as 1:3 in running
migrates. In other words, three times as long feeding as
travelling.
3. Speed of movement
Moving faster uses more energy but arrives quicker.
4. Chosen or enforced detours
Whether to go directly from A to B or via another

place (eg: feeding site)(chosen detour). While enforced
detours are barriers for land migrating animals, like
rivers or mountains, and poor weather conditions for
birds (eg: blown off course).
5. Optimal time of departure
Before the weather changes negatively or food supply

exhausted here, and arrive when food supply best at
destination. Birds are known to leave stopover sites when
winds are favourable (Akesson and Hendestrom 2002).
KNOWING WHEN TO MIGRATE
Migration is about moving to the right place at the

right time. Birds seem to have circannual (annual)
biological clocks which trigger the migratory
restlessness (Zugunruhe) and put on weight beforehand
(Elphick 1995). The biological clock, probably
linked to the pineal gland in the brain and connected to
the visual system, responds to changes in light and
darkness with the seasons.
But there is also an endogenous biological clock
which works irrelevant of changes in the environment.
Gwinner (1986) kept wild birds in special cages for over
three years. The environment was controlled so that there
was always 12 hours of daylight and 12 hours of darkness.
Thus the removal of external cues to migration. But the
birds showed the migratory restlessness at the
appropriate time of the year still.
However, the Wandering albatross (Diomedea exulans)

breeds biennial, and so migrates to the breeding site
every other year; even less frequently in some cases. How
does the biological clock work in this situation?
MECHANISMS USED TO NAVIGATE
One area of major interest in relation to migration

is how the animals know where to go (migratory
orientation systems). Birds, which travel vast distances,
are of particular interest, and the cues they are using.
Table 1.5 shows the different cues that animals use to
navigate.
1. Sun compass - movement of sun; angle of sun; polarised light

(pattern of light based on sun's position and reflection on water).
2. Geomagnetic compass - sensitivity to magnetic North and the

earth's magnetic field.
3. Star compass or position of moon.
4. Other visual cues - patterns of waves; cloud patterns; landmarks.
5. Smell.
6. Sound.
7. Electric.
Table 1.5 - Different cues used by animals to navigate.
The use of a geomagnetic compass by birds is hotly

debated. The geomagnetic declination is the difference
between the North Pole and "magnetic North", and it
varies based on global position (Thorup et al 2006).
The development of satellite-based radiotelemetry

has meant that birds fitted with small transmitters can
be tracked during their migration in free-flying studies.
For example, Thorup et al (2006) tracked the route taken
by 25 peregrine falcons (Falco peregrinus)(figure 1.7) in
the autumn movement between North and South America, and
seven honey buzzards (Pernis apivorus) and thirteen
ospreys (Pandion haliaetus) between Europe and Africa.
The researchers wanted to see if the birds followed
a geographical compass (direct route from home to
destination) or a geomagnetic compass (curved path). For
example, in North America in autumn, birds following a
fixed magnetic course will curve anti-clockwise (south to
east) when migrating south-east in western North America
and clockwise (south to west) when flying south-west in
the east.

Figure 1.7 - Peregrine falcon eating on US Fish and

Wildlife Service boat.
The flight direction for all three species were

closer to a predicted constant geographical compass. This
suggested that the birds were flying directly towards
their destination (aided by celestial cues like stars
during night flight) and not using magnetic cues.
However, it is quite probable that birds use a
selection of orientation cues to establish the direction
of migration. For example, geomagnetic cues become
important when there is cloud cover. Thorup et al (2006)
were also pragmatic in admitting that different species
of birds may use different cues.
Other researchers use different methods to study

this same behaviour. Cue-conflict experiments study
orientation cues in laboratory-based surroundings. A
typical example (Akesson et al 2002) involved 36 juvenile
white-crowned sparrows (Zonotrichia leucophrys gambellii)
captured in northern Canada. They usually migrate south-
east to southern USA for the winter. The birds were kept
in cages surrounded by magnetic coils that shifted the
magnetic field by ninety degrees.
If the birds use celestial cues to migrate at night,
they will be unaffected by the shift in magnetic field,
whereas if magnetic cues are used they will be. The top
of the cages had Emlen funnels (Emlen and Emlen 1966).
These are funnels through which the birds can see the
sky, but it is too small for them to fly through. The
direction the bird is trying to fly is recorded on Tipp-
Ex paper which lined the funnel.
The sparrows showed that they were influenced by the
shifted magnetic field, and so must have been using
magnetic cues. However, released birds were able to
recalibrate quickly using celestial cues.
Free-flying studies have advantages and

disadvantages in relation to laboratory studies which
artificially change the magnetic fields and orientations
in cages (table 1.6).
FREE-FLYING STUDIES
Advantages
1. Studying birds in their natural habitat.
2. Study over larger distances than laboratory studies.
3. Aided by development of modern technology which overcomes the

problem of observers missing behaviour.
4. Little interference of bird's behaviour by researchers.
5. Gives fuller picture of behaviour than in short experiments.
Disadvantages
1. Birds fitted with tracking devices which may change their

behaviour; eg: radio-transmitters produce very weak magnetic field.
2. Expensive, particularly if bird dies or migration studied not

typical (like extreme weather).
3. Tend not to have control group.
4. Lack of control of variables.
5. Not possible to establish causality.
LABORATORY EXPERIMENTS
Advantages
1. Control over birds and variables studied.
2. Possible to establish cause and effect.
3. Can have multiple conditions each changing a different variable as

well as a control group.
4. Birds watched (or video-recorded) all the time and so unlikely to

miss changes in behaviour.
5. Replication possible.

Disadvantages
1. Artificial conditions.
2. Whether birds affected by capture and captivity.
3. Whether birds bred in captivity are typical of the species.
4. Behaviour measured for limited period only.
5. Narrowness of independent and dependent variables.
Table 1.6 - Advantages and disadvantages of free-flying

and laboratory studies of migration orientation.
RISK OF PREDATORS
Animals (particularly birds) are especially

vulnerable to predators at temporary stopovers either
from lack of knowledge of local predators or
preoccupation with large food intake.
Young birds on their first migration or experienced
birds affected by poor weather conditions stop at sites
never visited before and encounter new types of
predators. There is limited time during such stopovers to
gain local information about the risk because of the
demands of "refuelling" (Cimprich et al 2005).
Does this mean that birds change their behaviour

during migration? Are they willing to accept greater
risks from predators during temporary stopovers because
of the need for food? Research by Cimprich et al (2005)
suggested that this is not necessarily the case and
predator avoidance remains just as important during
migration as at home.
Cimprich et al (2005) used observation and

experiment to investigate the behaviour of blue-grey
gnatcatchers (Polioptila caerulea)(figure 1.8) and
American redstarts (Septophaga ruticilla) on temporary
stopovers at Bon Secour National Wildlife Refuge on the
coast of Alabama, USA.
During the observation part of the research, the
behaviour of these birds in response to the appearance of
a flying predator (sharp-shinned hawk; Accipter striatus)
was measured. In the experiment part, the researchers
simulated a gliding hawk using a balsa wood model.

(Source: Albuttlee)
Figure 1.8 - Blue-grey gnatcatcher.
The appearance of a hawk (real or model) caused the

birds to move deeper into the shrubs (ie: out of sight),
and to reduce movement (birds of prey attracted by
movement). When there was no risk of hawks, most of the
birds were on the edge of the shrubs moving rapidly from
perch to perch. With a high risk of hawks, the birds
moved over 40cms into the shrubs, and changed perch less
than once per minute. The experiment produced similar
data (table 1.7). The birds responded to predators in
their normal way (hiding and/or freezing) despite the
negative impact upon amount of food gained.
BEFORE AFTER
Median depth into shrubs (cms) 8 18

Mean movement rate (perch changes/min) 30 20
(After Cimprich et al 2005)
Table 1.7 - Blue-grey gnatcatchers' behaviour in response

to model hawk.
As well as the risk of predation at stopovers, there

is also that risk during the movement of migration. For
example, Ibanez et al (2001) reported evidence that
greater noctule bats (Nyctalus lasiopterus) were
capturing nocturnal migrating birds in flight.
Based on bat dropping analysis from two regions of

Spain, the researchers found bird remains in the
migration periods, March-May and August-November, but not
during the rest of the year.
REFERENCES
Akesson, S & Hedenstrom, A (2000) Wind selectivity

of migratory flight departures in birds Behavioural
Ecology and Sociobiology 47, 140-144
Akesson, S et al (2002) Avian orientation: Effects

of cue-conflict experiments with young migratory
songbirds in the high Arctic Animal Behaviour 64, 469-
475
Alerstam, T (1993) Bird Migration Cambridge:

Cambridge University Press
Alerstam, T (2001) Detours in bird migration Journal

of Theoretical Biology 209, 319-331
Biebuch, H (1983) Genetic determination of partial

migration in European robin (Erithacus rubecula) Auk
100, 601-606
Cimprich, D.A et al (2005) Passerine migrants

respond to variation in predation risk during stopover
Animal Behaviour 69, 1173-1179
Cocker, J (1993) The migratory and navigational

behaviour of birds Psychology Teaching December, 2-12
Cramp, S & Simmons, K (1977) Handbook of the Birds

of Europe, Middle East and North Africa: Volume 1 -
Ostrich to Ducks Oxford: Oxford University
Elphick, J (1995) (ed) Atlas of Bird Migration

London: Collins
Emlen, S.T & Emlen, J.T (1966) A technique for

recording migratory orientation of captive birds Auk 83,
361-367
Gill, F (1995) Ornithology (2nd ed) New York: WH

Freeman
Grier, J.W & Burk, T (1992) Biology of Animal

Behaviour Dubuque, IO: W.C.Brown
Gwinner, E (1986) Circadian rhythm in the control of

avian rhythms Advances in the Study of Behaviour 16,
191-228

Harper, D (1988) Erithacus rubecula Robin: Field
characteristics. In Cramp, S (ed) Handbook of the Birds
of Europe, the Middle East and North Africa Vol V
Oxford: Oxford University Press
Hedenstrom, A (2003a) Optimal migration strategies

in animals that run: A range equation and its
consequences Animal Behaviour 66, 631-636
Hedenstrom, A (2003b) Scaling migration speed in

animals that run, swim and fly Journal of Zoology 259,
155-160
Ibanez, C et al (2001) Bat predation on nocturnally

migrating birds Proceedings of the National Academy of
Sciences, USA 98, 17, 9700-9702
Moreau, R.E (1972) The Palaearctic-African Bird

Migration Systems London: Academic Press
Perdeck, A.C (1958) Two types of orientation in

migrating starlings, Sturmus vulgaris L., and
chaffinches, Fringella coelbs L., as revealed by
displacement experiments Ardea 46, 1-37
Roselaar, C (1988) Erithacus rubecula Robin:

Breeding. In Cramp, S (ed) Handbook of the Birds of
Europe, the Middle East and North Africa Vol V Oxford:
Oxford University Press
Thorup, K et al (2006) Do migratory flight paths of

raptors follow constant geographical or geomagnetic
courses? Animal Behaviour 72, 875-880
Whitfield, P et al (1981) The Rhythms of Life

London: Marshall
Useful Websites
* MIGRATE (http://www.migrate.ou.edu/) open access

abstracts on:
white-crowned sparrows and migration

http://www.migrate.ou.edu/pubwiki/index.php/White-
crowned_Sparrow_%28Zonotrichia_leucophrys%29_Papers
American redstarts and migration

http://www.migrate.ou.edu/pubwiki/index.php/American_Re
dstart_%28Setophaga_Ruticilla%29_Papers
* BBC project "World on the Move"

http://www.bbc.co.uk/radio4/worldonthemove/

2. ADVANTAGES AND DISADVANTAGES OF AMBUSH
HUNTING FOR ANIMALS
Predators seek prey in different ways. The most
obvious is the chase. But some predators take a
completely different approach, and use "sit-and-wait"
foraging or ambush hunting. This means that the predator
picks a spot and waits, sometimes for a long period, for
the prey to come to them. There are ambush hunters
throughout the animal kingdom (table 2.1).
Many snakes
Many spiders and some other insects
Crocodiles and alligators (partly)
Some lizards and reptiles (eg: tuatura; Sphenodon

punctatus; picture:
http://www.bbc.co.uk/nature/wildfacts/factfiles/3052.shtm
l)
Few birds (eg: shrike; Laniidae; picture:

http://www.montereybay.com/creagrus/shrikes.html)
In sea (eg: monkfish; Lophius upsicephalus. Pacific angel

shark; Squatina californica. Anglerfish;
Lophiiformes)
Table 2.1 - Some examples of ambush hunters.
There are advantages and disadvantages of ambush

hunting (table 2.2), particularly in comparison with
chasing prey.
ADVANTAGES
1. Uses little energy, and much less than chases.
2. Works where prey have predictable habits, like using the same
routes.
3. Good way to get prey within reach of weapons - eg: teeth

(crocodile), poison (snake).
4. Avoids the risk of prey fighting back (and of injury) if ambush is

swift and effective.
5. Best where prey moves around or are geographically dispersed.
6. Catch prey when off-guard or not expecting attack - eg: spider's

web and flying insects.
7. Most effective when predator has no natural enemies.
8. Best when animal does not need to feed too often.

9. It is a strategy than can be used as well as others like chasing
the prey.
10. Effective strategy for well-camouflaged animals.
11. Disguises the number of predators competing for food, and risk of
fights.
DISADVANTAGES
1. Risk that prey will not come or not right type of prey, and thus
starvation.
2. If limited number of prey come, predator must have high success

rate or risk of starvation.
3. Risk of becoming prey themselves if stationary for long periods.
4. Not effective if spotted by prey; thus importance of camouflage.
5. Not effective if prey learns to avoid ambush sites, particularly

if predator always uses same sites.
6. Waste of limited resources if ambush unsuccessful - eg: snake's

poison.
7. If prey are unpredictable means must be alert at all times which

limits opportunity for sleep.
8. Risky strategy for animals with high calorie needs.
9. Depends on prey being mobile.
10. Too many prey can distract predator for individual kill.
11. Lower capture rates than active hunters of same species (eg:
lizards; Anderson and Karasov 1981).
Table 2.2 - Advantages and disadvantages of ambush

hunting for animals.
The best ambush hunters will be camouflaged and

attack very quickly. Crypsis is camouflage where the
animal shows visual resemblance to some part of the
environment, like crab spiders (Thomisidae) that assume
the same colour as flowers which deceives landing insects
(Thery and Casas 2002).
Pacific angel sharks (Squatina californica) hides on
the sea floor partly covered in soft substrata as well as
its crypsis. The duration of an attack was measured at
30-100 milliseconds (Fouts and Nelson 1999).
Within the same group of animals active and ambush

hunting can evolve. Cooper and Whiting (2000) collected
data on skinks (lizards) in southern Africa (table 2.3).

ACTIVE HUNTERS
More energy used

Higher capture rate
Greater stamina and greater average speed (eg: 0.03 m/s)
Search in wider area per unit time
Higher proportion of time spent moving (PTM)(eg: 0.49) and higher
number of movements per minute (MPM)(eg: 1.6)
More vulnerable to ambush predators
Examples: Mabuya sulcata (figure 2.1), Mabuya variegata
AMBUSH HUNTERS
Less energy used

Lower capture rate
Greater sprint speed (eg: 0.2 m/s vs 0.008 m/s) but lower average
speed (eg: 0.003 m/s)
Limited search area
Lower PTM (eg: 0.03) and lower MPM (eg: 0.3)
More vulnerable to active hunters
Examples: Mabuya acutilabris, Mabuya spilogaster
Table 2.3 - Comparison between active and ambush hunters

among skinks.
(Source: Hans Hillewaert)
Figure 2.1 - Mabuya sulcata.

The strategy of ambush or active hunter also depends
upon the risk to the predator from their predators (eg:
jumping spiders:
http://www.bgu.ac.il/life/Faculty/Bouskila/when.html).
AMBUSH SITE SELECTION
The choice of ambush site is very important for

"sit-and-wait" foragers. Successful ambush sites must
have certain characteristics (Shine and Sun 2002):
i) Site visited frequently and/or in large numbers by

prey;
ii) Possible for predator to hide - ie: opportunity for

camouflage/evade detection;
iii) Predator can detect prey immediately on arrival.

This is important if the prey only visit the ambush site
briefly;
iv) Allows for prey capture. The prey needs to come

within the striking distance of the predator.
These characteristics may vary in importance

depending upon the availability of prey. In the case of
scarcity, success in finding prey is paramount. Whereas
if the prey are abundant but hard to catch, ambush
factors related to capture become most important.
Sine and Sun (2002) investigated the site choice of

one type of pit-viper (Gloydius shedaoensis) in ambushing
migrating birds that land on branches. The study took
place on Shedao island in north-eastern China, which lies
on major migratory bird routes, in May 2000. The snakes
only feed on twice-yearly migrating birds, which can be a
risky strategy (table 2.4).
ADVANTAGES
1. High availability of prey twice a year.
2. Birds tired and less wary of predators.
3. Birds not familiar with area and with predators.
DISADVANTAGES
1. Risk of no food if migration patterns change (eg: storm blows

birds off course).
2. Birds may learn to be wary of snakes' behaviour and change their

behaviour.

3. Snakes shut down in between migrations and this involves risks.
Table 2.4 - Advantages and disadvantages of pit-viper

strategy of waiting for migrating birds.
One hundred and forty-nine snakes were observed to

discover the choice of ambush site on a tree. One hundred
and twenty-seven trees were chosen, and the snakes showed
preferences for their ambush sites as follows:
To perch in trees on the edge (rather than in the

middle) of thickets or in isolated trees;
Branches facing outward;
Branches lower to the ground. Nearly two-thirds of the

snakes were found on branches 0.5 metres above the
ground, and less than 10% 1.5 metres or more above the
ground. The majority of the prey observed (564 bird
perching events) preferred outward facing branches, and
branches less than 1 metre above the ground. This
follows criteria (i) above for a successful ambush
site;
Diameter and angle of branch that allowed snake

stability to strike (criteria iv above);
Cool backgrounds which allowed a contrast for the

snakes to use their heat-sensitive facial pits to spot
birds (criteria iii above). Outward facing branches
have this.
The branches chosen also allowed camouflage as the

snake's body colour and scalation were a visual match to
the tree's appearance (criteria ii above). The
researchers admitted that "we often failed to see pit-
vipers in ambush poses until we had approached them far
more closely than we desired".
BEING SPOTTED BY PREY
Ambush hunting is only successful if the prey does

not spot the ambusher. Weislo and Schatz (2003) looked at
the evasive action taken by female sweat bees
(Lasioglossum umbripenne) when detecting ants (Ectatomma
ruidum) waiting in ambush near their nest in Panama. The
bees nest in mounds in the soil, and the ants, that are
twice as big, wait for returning bees and lunge at them.
The ants cannot enter the nest because they are too big.
The researchers placed different stimuli at the nest
entrance:

A dead ant (either with or without its natural smell)
A black cardboard rectangle (the size of an ant)
A black cardboard square (the size of an ant)
No stimulus
Live ant
When there was no stimulus by the nest, the majority

of bees (85.5%) flew directly into the nest ("approach").
When the bees detected an ambusher they changed their
behaviour ("approach and withdraw"). This included re-
approaching the nest from the opposite side (54% of times
when live ant present), zigzagging flights, or landing
and walking into the nest.
Sometimes the ant was still able to capture the bee,
and, on other occasions, the ant was distracted by other
returning bees. "Flexibility in nest entering-exiting
behaviour of L.umbripenne, in the face of flexible
hunting strategies by E.ruidum.. illustrates the dynamics
of a complex predator-prey relationship.." (p187).
The bees did not change their behaviour in the
presence of the cardboard models which suggested that
they visually detect ants.
REFERENCES
Anderson, R.A & Karasov, W.H (1981) Contrasts in

energy intake and expenditure in sit-and-wait and widely
foraging lizards Oecologia 49, 67-72
Cooper, W.E & Whiting, M.J (2000) Ambush and active

foraging modes both occur in the Scincid Genus Mabuya
Copeia 1, 112-118
Fouts, W.R & Nelson, D.R (1999) Prey capture by the

Pacific angel shark, Squatina californica: Visually
mediated strikes and ambush-site characteristics Copeia
2, 304-312
Shine, R & Sun, L-X (2002) Arboreal ambush site

selection by pit-vipers, Gloydius shedaoensis Animal
Behaviour 63, 565-576
Thery, M & Casas, J (2002) Predator and prey views

of spider camouflage Nature 10/1, p133
Weislo, W.T & Schatz, B (2003) Predator recognition

and evasive behaviour by sweat bees, Lasioglossum
umbripenne (Hymenoptera: Halictidae) in response to
predation by ants, Ectatomma ruidum (Hymenoptera:
Formicidae) Behaviour Ecology and Sociobiology 53, 182-
189

3. ADVANTAGES AND DISADVANTAGES OF CACHING
BEHAVIOUR
Some animals eat their food as soon as they find or
capture it. Others will store (cache or hoard) it for
later. Caching occurs in some species of birds (eg:
western scrubjays) and mammals (eg: squirrels). This is
particularly useful when food supplies are low or
unpredictable at certain times of the year, like winter
(table 3.1).
There are factors that influence whether a food item

is cached or eaten immediately (Dally et al 2006):
Perishability;
Handling time (ie: time taken to eat item);
Presence of other animals who eat same food (ie:

potential pilferers from same or different species) and
risk of kleptoparasitism (food stealing);
Observer identity - whether observer is attentive

(threat)(eg: social species like rooks) or inattentive
(not pilfering threat)(eg: asocial species like Clarks
nutcrackers);
Position in social hierarchy;
Food value (ie: energy gained).
ADVANTAGES
1. Means food available during winter.
2. Means food available during shortages or when unpredictable.
3. Better for animals that active throughout the year rather than
those which build up fat stores and hibernate.
4. Not dependent on living near large food sources or waiting for

binge periods (eg: migration).
5. Useful when too much food to carry home or eat at one time.
6. Quicker to store food than consume it. This is important when

there is a high risk of predation, or living with food competitors as
in groups.
7. Immediate consumption of food limited by amount of fat that animal

can store or else food item wasted.
8. Animals can cache in one place (larder) or in multiple sites

(scatter caching).

9. Allows storage for when greater demand (eg: offspring born).
10. Part of the evolution of sophisticated behaviours like planning.
DISADVANTAGES
1. Problems of storing food, like degradation.
2. Risk of theft of cache (pilferage) from own or other species.
3. Risk of forgetting where cache hidden. Needs animal to have well

developed memory to work if scatter caching used.
4. Risk of predation while retrieving cache.
5. Cache may be inaccessible when needed (eg: snow covers ground

where food buried).
6. Requires the ability to plan.
7. Immediate consumption of food stops competitors from getting it.
8. Problems of defending larder.
9. Energy wasted with pilfering avoidance strategies (eg: moving

further away to cache or pretending to cache).
10. Not good strategy for food with low energy content (ie: expend
more energy caching and retrieving than gaining energy), unless
absolute food shortage.
Table 3.1 - Advantages and disadvantages of caching food.
FINDING THE CACHE
Grey squirrels may have left food in over 10 000

different locations, which they recover with a
combination of visual landmarks, scent marking and memory
(Cline 2005).
They are using scatter caching. There is evidence of
them refreshing their memory of cache sites by checking
them and moving around caches ("Life of Mammals:
Chisellers" 2002; BBC Television).
Kamil and Jones (1997) showed that Clarks

nutcrackers (Nucifraga columbiana)(figure 3.1) use a
"cognitive map" to find their cache. The experiment used
a sandy room with two landmarks which were changed in
position each time. The experimenters always buried the
seeds half way between the two landmarks. The birds were
allowed into the room four times per day for 50-60 days.
Within fifteen days they could accurately find the seeds
each time. They had learnt the geometrical relationship
between the buried seeds and the two landmarks, which is
the characteristic of a cognitive map.

Figure 3.1 - Clarks nutcracker.
The hippocampus is the area of the brain linked to

memory, and particularly spatial memory. It can expand in
some species (eg: black-capped chickadees) in the autumn
when caching for the winter (Smulders et al 1995). While
rats with damage to that area could not remember the
location of a platform in the Morris selection task (or
open-field water maze procedure)(Jarrard 1995).
The Morris selection task (Morris 1981, 1984)
involves a pool of water with a hidden platform.
Initially, the rats swim around until they hit the
platform by chance and then climb on. Subsequently, when

put in the pool, they swim straight to the platform
showing evidence of spatial memory.
CACHING AND SOPHISTICATED BEHAVIOUR
Caching is associated with sophisticated behaviour

by the animals that use the strategy.
Adaptability
Squirrels cache nuts. In North America, they can

distinguish two types of oak nuts - those that germinate
soon which are eaten immediately, and those that
germinate later which are cached. When there is a
shortage of the latter, the squirrels collect oak nuts
that germinate soon and remove the germinating part of
the seed before caching ("Life of Mammals: Chisellers"
2002; BBC Television).
Foresight and Planning
Western American crows (Corvus brachyryhchus

hesperis) were observed to eat bigger walnuts immediately
and bury smaller ones 1-2kms away. This meant high energy
now and the loss of smaller nuts not such a problem
(Cristol 2001).
Western scrubjays (Aphelocoma californica)(figure
3.2) were found to recover perishable items first from
their cache. This could be evidence of an internal
representation of the future (Clayton et al 2001).
These birds have shown planning in a recent
experiment (Raby et al 2007). The scrubjays were put in
special cages divided into sections in the evening and in
the morning. In one section there were given food in the
morning ("breakfast room") and in another not ("no-
breakfast room"). When given pine nuts in the evening,
the birds buried three times as many in the sand of the
"no-breakfast room" as they did not know which section
they would be placed in the morning.
Using the same cage set-up, the birds were given

different foods (peanuts or dried dog food) in different
sections of the cage in the morning. When in the cage in
the evening with food, they buried the opposite food to
where given in the morning. So they buried dried dog food
in the cage section where fed peanuts in the morning and
vice versa.

(Source: US National Park Service)
Figure 3.2 - Western scrubjay.
Crows also show evidence of planning. They cache

clams during low tide because not available during high
tide ("Bird Brains" 1999; National Geographic Channel).
PILFERAGE RISK
There is always a trade-off between the benefits of

hoarding food and the cost of loss from pilferage.
Cache pilfering has been seen in a number of species
who cache food (Dally et al 2006)(table 3.2).
Stealing from other species (heterospecifics)

Grey jays (Perisoreus canadensis) and Stellar's jays (Cyanocitta
stelleri)
Great tits (Parus major) from marsh tits (Poecile palustris)
Eurasian jackdaws (Corvus monedula) from rooks (Corvus frugilegus)
Stealing from own species (conspecifics)

Grey squirrels (Sciurus carolinensis)
Pinyon jays (Gymnorhinus cyanocaphalus)
Mexican jays (Aphelocoma ultramarina)
Clarks nutcrackers (Nucifraga columbiana)
Western scrubjays (Aphelocoma californica)
Common ravens (Corvus corax)
Table 3.2 - Animals known to do cache pilferage.

Cache pilfering may not be such a problem if the
pilferers are genetically related to the storer (indirect
fitness)(Smulders 1998), or if the storers are also
pilferers themselves and make up for own losses
("reciprocal pilferage")(Vander Wall and Jenkins 2003).
For example, North American red squirrels
(Tamiasciurus hudsonicus) lost 25% of their stored cones,
but pilfered 26% of those they ate (Gerhardt 2005). In
terms of indirect fitness, it is the benefits to the
genes rather than to the individual. So theft of food by
a brother, for example, makes evolutionary sense.
When there is risk of pilfering, the cachers can use

cache protection behaviours (Dally et al 2006):
i) Eat food immediately;
ii) Cache more to offset predicted loss (eg: eastern

chipmunks; Tamias striatus). But hard to predict loss and
extra effort needed;
iii) Cache in many different sites making it

unprofitable for pilferers to find. But needs cacher to
have good memory;
iv) Cache less or not at all in presence of

observers.
v) Cached behind a large obstacle when observers

present (eg: ravens; Bugnyar and Kotrschal 2002);
vi) Delay caching until alone (ie: "out of view" of

competitors);
vii) Defend cache items ("larder-hoarders" keep all

cache in one place). But risks from fighting;
viii) Cache in sites difficult to find for

competitors (ie: "hard-to-see" sites);
ix) Move caches after observers gone (recaching);
x) Misinformation - pretend to cache or appear not

to have food item.
If observers go, then cacher has waiting risks (eg:

predation, loss of food), but a benefit when alone.
Different strategies are needed if food competitors are
constantly present (table 3.3).

CONSTANT PRESENCE COMPETITOR PRESENT BOTH
OF FOOD COMPETITOR SOMETIMES
ii; iii; vii; viii iv; v; vi; ix i; v; x

(Numbers refer to cache protection strategies listed above)
Table 3.3 - Cache problem strategies depending on whether

food competitors are constantly observing or not.
Animals that cache have been shown to adapt their

behaviour in response to cache pilferage in different
ways (Dally et al 2006):
Change food type cached;

Change from scatter to larder hoarding;
Eating caches immediately.
Western scrubjays also show evidence of the ability

to imagine another individual's behaviour. Birds that had
stolen food from other caches were more careful in hiding
their own cache from observers (Emery and Clayton 2001).
One way to combat cache pilferage is through

"behavioural deception". This is "the use of false
signals to modify the behaviour of a receiver in a way
that benefits a sender, at some cost to the receiver"
(Semple and McComb 1996).
The use of behavioural deception has been seen in
grey squirrels (figure 3.3). This includes covering
additional empty cache sites ("deceptive caches") if
other squirrels were present (Steele et al 2008).
Squirrels were observed at three different sites in
north-eastern USA. Deceptive caching was observed in
around 10% of caches, and was more likely when at least
three other squirrels were present and in close proximity
(average of six metres away). Overall, pilferage
avoidance behaviours were evident in around one-third of
cachings.
Leaver et al (2007) observed the pilferage avoidance

strategies of eastern grey squirrels at their university
campus (University of Exeter, England). The number of
caches made in the presence or absence of another
squirrel was not significantly different (mean 3.2 vs
4.2), nor in the presence or absence of birds (eg:
carrion crows, Corvus corone)(mean 3.3 vs 4.4). But the
cachers were more likely to face away from other
squirrels when burying food (52% of caches), yet not when
only birds present. Also caches were spaced further away
(average three metres apart) because "travelling further
between successive caches may serve to distract the
attention of an observer, which may descrease the

likelihood of that observer engaging in area-localised
search" (p26).
(Source: MONGO)
Figure 3.3 - Grey squirrel.
The strategies used by cachers and pilferers is an

example of an "evolutionary arms race" (Bugnyar and
Kotraschal 2002). This is where the development of a
strategy on one side leads to the development of
competing strategy on the other. For example, the
evolution of the ability to learn where cache sites from
observing cachers is countered by caching "out of view".
REFERENCES
Bugnyar, T & Kotrschal, K (2002) Observational

learning and the raiding of food caches in ravens, Corvux
corax: Is it "tactical" deception? Animal Behaviour 64,
185-195
Clayton, N et al (2001) Elements of episodic-like

memory in animals Philosophical Transactions of the Royal
Society B 356, 1483-1491
Cline, P (2005) Animal Cameo: Grey squirrel -

Sciurus carolinenesis Feedback September, p5
Cristol, D (2001) American crows cache less

preferred walnuts Animal Behaviour 62, 331-336
Dally, J.M et al (2006) The behaviour and evolution

of cache protection and pilferage Animal Behaviour 72,
13-23
Emery, N & Clayton, N (2001) Effects of experience

and social context on prospective caching strategies by
scrubjays Nature 414, 443-446
Gerhardt, F (2005) Food pilfering in larder-hoarding

red squirrels (Tamiasciurus hudsonicus) Journal of
Mammalogy 86, 108-114
Jarrard, L.E (1995) What does the hippocampus really

do? Behavioural Brain Research 71, 1-10
Kamil, A.C & Jones, J.E (1997) The seed-storing

corvid Clark's nutcracker learns geometric relations
among landmarks Nature 390, 276-279
Leaver, L et al (2007) Audience effects on food

caching in grey squirrels (Sciurus carolinensis):
Evidence for pilferage avoidance strategies Animal
Cognition 10, 23-27
Morris, R.G.M (1981) Spatial localization does not

require the presence of local cues Learning and
Motivation 12, 239-261
Morris, R.G.M (1984) Developments of a water-maze

procedure for studying spatial learning in the rat
Journal of Neuroscience Methods 11, 47-60 (pdf at
http://pages.towson.edu/bdevan/Morris%20articles/Morris,%
201984.pdf)
Raby, C.R et al (2007) Planning for the future by

western scrubjays Nature 445, 919-925
Semple, S & McComb, K (1996) Behavioural deception

Trends in Ecology and Evolution 11, 34-37
Smulders, T.V (1998) A game theoretical model of the

evolution of food hoarding: Applications to the Paridae
American Naturalist 151, 356-366
Smulders, T.V et al (1995) Seasonal variation in

hippocampal volume in a food-storing bird, the black-
capped chickadee Journal of Neurobiology 27, 1, 15-25
Steele, M.A et al (2008) Cache protection strategies

of a scatter-hoarding rodent: Do tree squirrels engage in
behavioural deception? Animal Behaviour 75, February,
705-714
Vander Wall, S.B & Jenkins, S.H (2003) Reciprocal

pilferage and the evolution of food-hoarding behaviour
Behavioural Ecology 14, 656-667

4. ADVANTAGES AND DISADVANTAGES OF
TERRITORIALITY
Some animals have a fixed territory (territoriality)
while others are mobile. Territoriality is defined as
"the defence of a fixed physical space with the purpose
of excluding individuals that are not members of the
social group" (Darden and Dabelsteen 2008 p905).
Importantly, it is the area defended against individuals
of the same species (Gese 2001).
The territory may be defended permanently (all year
round) or seasonally, and the defended area can include
nesting sites, food supplies, and/or sexual partners.
Territory is defended in different ways including
aggressive physical contact (direct defence) and signals
of borders (eg: scent markings)(indirect defence).
In some cases, animals may have more than one

territory: for example, oystercatchers have two
territories - inland where they nest, and on beach where
they feed (Sparks 1969).
Territoriality has a number of advantages and

disadvantages (table 4.1).
ADVANTAGES
1. Males able to hold on to resources show their evolutionary fitness

and are attractive to females.
2. Exclusive access to food, particularly at times of shortage.
3. Exclusive area for breeding and raising young.
4. Space for sexual display and courtship.
5. Spacing of animals avoids competition.
6. Reduces aggression.
7. Local knowledge of predators and resources.
8. Exclusive place to retreat and shelter.
9. Dispersion of nests reduces predation.
10. Higher survival rates.
11. Benefits in growth (eg: convict cichlids in larger territory grew

quicker; Kim et al 2004).
DISADVANTAGES
1. Cost of defending territory including risk of physical contact,

and displays of strength.

2. Need to be vigilant for intruders.
3. Defending territory is time that could be feeding or mating.
4. Vocal or visual communication of territory ownership makes the

individual vulnerable to predation.
5. Difficult for smaller animals to hold territory; ie: more likely

to be attacked than larger animals (eg: smaller European common toads
received 240 attacks compared to 35 for larger toads; Davies and
Halliday 1979).
6. Difficult to move if resources exhausted.
7. Importance of territory size. If too large, then hard to maintain

control. If too small, not enough resources for effort of defending.
8. Higher risk of predation if territory within predator's territory.
9. Easy for predators to find.
10. Ever present threat of take-over as surplus of animals without

territory.
11. Extra vigilance required at certain times of the year (eg:

breeding season).
Table 4.1 - Advantages and disadvantages of

territoriality.
There are three main reasons for territoriality

which varies between species (table 4.2).
REASON FOR TERRITORY NUMBER OF SPECIES

(from 40; Wilson 1975)
1. Control food supply 14
2. Retreat; shelter; nest 8
3. Access to females; space

for sexual display; courtship 18
Table 4.2 - Main reasons for territories among different

species.
Maher and Lott (2000) assessed the variables that

influenced whether an animal is territorial or not (table
4.3).
But how each of the variables influences

territoriality can be unclear with contradictory results
from studies. For example, twelve studies reported that
territoriality declined as food quantity increased and
four studies that increased food produced increased
territoriality, while a few studies found no relationship
(Maher and Lott 2000). Only some of the studies were
experiments.

FOOD NON-FOOD OTHER
CHARACTERISTICS RESOURCES
- quantity - distribution - population density

- predictability - quantity - habitat features
- distribution - predictability - mating
- quality - quality - space
- renewal rate - refuges/spawning/
- type home sites
- density - predation pressure
- accessibility - host nests (for
brood parasites)
- energy availability
Table 4.3 - Factors influencing whether an animal is

territorial or not.
Maher and Lott felt that the relationship between

many variables and territoriality is an inverted U-shape;
eg: "very abundant food and very scarce food would not be
defended, but intermediate levels would be defended"
(p18).
Not only individuals but groups also defend

territory. Gese (2001) observed 112 instances of
territory defence in Yellowstone National Park, Wyoming,
USA by coyote (Canis latrans)(figure 4.1) packs. A number
of patterns were seen:
The main coyotes involved in territory defence were

alpha males (77% of chases observed) or alpha females
(59%);
The majority (three-quarters) of the intruders were

evicted without physical contact (ie: chasing was
enough). Physical contact did not involve serious
injury;
The average chase was 2.87 minutes and up to 1km until

the territory boundary;
Territory defence increased during the breeding season

(January-February), the gestation season (March), and
when the pups emerged from the den (June);
Territorial coyotes had advantages over transient

animals, but the nature of the advantages varied with
position in the pack social hierarchy. Alpha coyotes
had the greatest advantage in terms of breeding, while
beta animals rarely did and transients not at all. All
territory members gained in feeding terms compared to
transients - greater capture rate of small mammals (2.3
per hour vs 2.0) and greater capture success (alphas
38.2% vs 32.3% for transients).

(Source: marya (emdot))
Figure 4.1 - Coyote.
Usually residents are protective of their territory

from outsiders. But Davis and Houston (1981) noted two
unrelated pied wagtails (Motacilla alba)(figure
4.2)(owner and "satellite") defending winter feeding
territory together because they could achieve a one-third
higher feeding rate than alone. Lone birds spotted 60% of
intruders on their territory while two birds spotted 85%.
This is an example of mutualism where co-operation
between two individuals benefits both more than
competition between them (Hinde 1982).
Territorial defence is costly. For example, Gill and

Wolf (1975) estimated the cost of territory defence for
individual golden-winged sunbirds (Nectarinia reichenowi)
was three times more per unit of time of energy compared
to feeding for the equivalent time (1000 kCals foraging
vs 3000 kCals chasing intruders away). But these birds in
East Africa that collect nectar from flowers in their
territory saved 780 calories by having a territory rather

than mobile foraging.
(Source: sannse)
Figure 4.2 - Pied wagtail.
COMMUNICATING TERRITORY OWNERSHIP AND DISPUTES
One way to communicate territory ownership is

through vocal signals, like barking. Darden and
Dabelsteen (2008) investigated such behaviour among the
male swift fox (Vulpes velox)(figure 4.3) observed in
north-east Colorado, USA. The study included an acoustic
playback experiment (table 4.4) which involved playing
the call of a strange male fox to measure the response of
resident male foxes.
The researchers found that resident males responded
with barking to the recording of a strange male with
greater intensity and much quicker when the playback
occurred in the core area of the territory compared to
the edge (average 226 seconds to respond for inner area
vs 471 seconds for outer area).

(Source: en:user:cburnett)
Figure 4.3 - Swift fox.
ADVANTAGES
1. Researchers can control and test exactly what interested in

studying.
2. Includes control group (eg: sound of neutral animal like a cow) to

compare with the experimental group.
3. Able to establish cause and effect.
4. Standardised procedure and recordings.
DISADVANTAGES
1. Problems of accurately recording calls.
2. Involves interference by the researchers into the life of the

animal.
3. Artificial - eg: no associated smell when using speaker, so

resident may realise not real intruder.
4. Time-consuming to collect recordings and expensive in terms of

equipment.
Table 4.4 - Advantages and disadvantages of acoustic playback

experiments.
In the case of frogs, if another male approaches the

territory, a "vocalisation battle" occurs. If the
intruder does not withdraw, the two males will wrestle

until one is forced onto his back (Emlen 1968). Wells
(1981) reported observing a fight of over two hours in
Trinidad. Residents tend to do better in fights, and win
over 90% of contests (Stewart and Rand 1991).
After disputes between neighbours are resolved, the
residents only respond aggressively to strangers. This is
known as the "dear-enemy truce-relationship" (Jaeger
1981), and saves energy and reduces the risk of endless
fights.
ROBINS
Generally the robin (Erithacus rubecula) is a

solitary bird, and is strongly territorial (except during
severe winters; Lack 1965). Migrant robins have been
found to be territorial even during a stop-over in
migration. Nelson (1907) observed territorial behaviour
on a boat at sea where a group of robins had rested.
Both sexes have their own individual territories

outside of the breeding season. There are two phases to
the territorial behaviour in the year. The individual
territory for most of the year, and the joining together
of two birds to breed.
a) Individual territory
The average size in English studies varies between

0.27-0.73 hectares (approximately 450-2000m²) per bird
(Harper 1988). There is no evidence of sex differences in
the size of the territory, nor it being related to food
density.
Sedentary (non-migratory) birds defend the same
territory for their whole life, while migrants return to
the same sites each year.
b) The breeding territory
Two birds will join together during the breeding

season, usually by the female moving to a male's
territory (78% of cases observed). In 10% of
observations, the male goes to the female's wintering
territory, and 9% of times adjacent territories are
fused together.
Occasionally, the birds will pair together and then
move to a new site: only 3% of 92 pairs studied in
Cambridge did this (Harper 1988). The average size of the
breeding territory is 0.55-1.44 hectares (approximately
900-2500m²) (Harper 1988).
Unpaired males can be evicted from their territories
at this time.

Aggression
Peek (1972) noted three levels of defence in birds:
i) Vocal communication as a long-range warning;

ii) Visual displays when visual contact made;
iii) Chased and attacked if other levels failed to repel
intruders.
Because robins are strongly territorial, there is a

high level of antagonistic encounters. Most occur when
territories are being established or when there are
boundary changes (at the beginning of the breeding
season). Usually the display of the red breast is enough.
The back of male European robins is camouflaged, but
the puffed-up red chest is a warning signal to other
males. Lack (1943) has found the red breast is a "sign
stimulus" 1 for territorial aggression against intruders.
Robins will attack just red feathers, such is the
strength of the stimulus. The resident bird flies at the
intruder with a high-pitched call. If the intruder does
not leave, the resident then perches on a branch and
displays its red breast. Occasionally this "show of
strength" does not work, and a fight ensues.
But 13% of 1067 encounters observed end in fights
(Harper 1988). These fights can be fatal: in a Cambridge
study, 10% of 98 males and 3% of 86 females died in this
way (Harper 1988).
Once the territories have been established, the

residents are able to distinguish the calls of their
neighbours from strangers occupying territories further
away. In this situation, the direct neighbour is not seen
as a threat. This is known as the "dear enemy phenomenon"
(Wilson 1975). Recognition of the neighbour avoids
wasteful conflicts, and makes clear the boundary edges of
the territory.
There are a number of possible explanations for the
"dear enemy phenomenon":
Habituation to the neighbour's song - each time it is

heard, it produces less of a territorial aggression
response until there is no response at all to the song;
Mere exposure effect - similar to above;
Dialect convergence - the similarity in the neighbours'
1
"Sign stimulus" is a trigger for instinctive behaviour (Hinde 1982). For example, the red of the
breast triggers the territorial aggression response. The trigger must have certain specific
characteristics. Lack (1943) found no territorial aggression response to models of robins without the
red breast.

songs reduces the territorial aggression response.
The "dear enemy phenomenon" is dependent on the
resident recognising their neighbours' calls from
strangers' calls. Hardouin et al (2001) found that male
little owls (Athene noctua)(figure 4.4) did not respond
to neighbours' hoots played back from the usual location,
but only if played from an unusual location. Based in
west France, this study used the "four-stimuli playback
discrimination paradigm". This involves recordings of two
sets of calls (neighbour and stranger) played from two
different places (usual and unusual locations).
(Source: Artur Mikolajewski)

Figure 4.4 - Little owls.
Where the playback came from the usual location of
neighbours, there was a clear difference in response by
residents towards neighbours and strangers. Stranger
calls elicited more hoots per minute (13 vs 9 for
neighbours), and for longer duration (average 200 seconds
vs 125 seconds). But when the playback came from an
unusual location, there was no difference in response by
residents towards neighbours and strangers.
Residents learn to discriminate familiar calls of
neighbours when they originate from the expected
direction. The two characteristics of sound and direction
is what the resident becomes habituated towards (Bee and
Gerhardt 2001).
Residents Always Win
Research has found that residents in a territory

tend to defeat the challengers in most cases. Maynard
Smith and Parker (1976) suggest possible explanations:
i) Value asymmetries - greater investment and local

knowledge about predators and resources gives the
resident advantages to win the contest;
ii) Resource-holding power (RHP) variations - those

in residence have superior abilities (eg size) that
allows them to defeat intruders;
iii) "Owners always win" convention - the individual

in possession of the territory fights harder than the
intruder.
This is an example of a "bourgeois strategy"; ie: a

strategy that cannot be bettered in a population. This
idea is the part of the application of game theory to
animal behaviour (Maynard Smith 1982).
Tobias (1997) tested the first explanation with a

series of "removal-replacement experiments" with 75
robins. This involves removing the residents from their
territories for a period of time, and then returning
them, to see if they can win the territory back from the
"new residents". Tobias removed 37 males and 13 females
from their winter territories, and 19 males and six
females from the spring territories. The length of the
removal varied from 1-14 days.
Two observations came from the research:
For winter territories, the longer the resident was

away the less likely they were to win back the
territory. Less than five days away, all won back their
territory, but if away for more than 10 days, no

returnees won;
With the spring territories, there was an immediate
loss no matter how short the period away. This is
because territory is crucial for males at this time of
the year. Males without territory have little chance of
breeding.
REFERENCES
Bee, M.A & Gerhardt, H.C (2001) Neighbour-stranger

discrimination by territorial male bullfrogs, Rana
catesbeiana: II. Perceptual basis Animal Behaviour 62,
1141-1150
Darden, S.K & Dabelsteen, T (2008) Acoustic

territorial signalling in a small, socially monogamous
canid Animal Behaviour 75, 905-912
Davies, N.B & Halliday, T.R (1979) Competitive mate

searching in male common toads, Bufo bufo Animal
Behaviour 27, 1253-1267
Davies, N.B & Houston, A.I (1981) Owners and

satellites: The economics of territory defence in the
pied wagtail, Motacilla alba Journal of Animal Ecology
50, 1, 157-180
Emlen, S.T (1968) Territoriality in the bullfrog,

Rana catesbeiana Copeia 2, 240-243
Gese, E.M (2001) Territorial defence by coyotes

(Canis latrans) in Yellowstone National Park, Wyoming:
Who, how, where, when and why Canadian Journal of Zoology
79, 980-987
Gill, F.B & Wolf, L.L (1975) Economics of feeding

territoriality in the golden-winged sunbird Ecology 56,
2, 33-45
Hardouin, L.A et al (2001) Neighbour-stranger

discrimination in the little owl, Athene noctua Animal
Harper, D (1988) Erithacus rubecula Robin: Social

pattern and behaviour. In Cramp, S (ed) Handbook of the
Birds of Europe, the Middle East and North Africa Vol V
Oxford: Oxford University Press
Hinde, R (1982) Ethology London: Fontana
Jaeger, R (1981) Dear enemy recognition and the

costs of aggression between salamanders American
Naturalist 117, 962-974

Kim, J.W et al (2004) Interactions between patch
size and predation risk affect competitive aggression and
size variation in juvenile convict cichlids Animal
Behaviour 68, 1181-1187
Lack, D (1943) The Life of the Robin London: H.F &

G Witherby
Lack, D (1965) The Life of the Robin London:

Penguin
Maher, C.R & Lott, D.F (2000) A review of ecological

determinants of territoriality within vertebrate species
American Midland Naturalist 143, 1-29
Maynard Smith, J (1982) Evolution and the Theory of

Games Cambridge: Cambridge University Press
Maynard Smith, J & Parker, G (1976) The logic of

asymmetric contests Animal Behaviour 24, 159-175
Nelson, T (1907) The Birds of Yorkshire London: A

Brown & Sons
Peek, F.W (1972) An experimental study of the

territorial function of vocal and visual displays in the
male red-winged blackbird (Agelaius phoeniceus) Animal
Sparks, J (1969) Bird Behaviour Harmondsworth:

Penguin
Stewart, M & Rand, A (1991) Vocalisations and the

defence of retreat sites by male and female frogs,
Eleutherodactylus coqui Copeia 1991, 1013-1024
Tobias, J (1997) Asymmetric territorial contests in

the European robin: The role of settlement costs Animal
Behaviour 54, 9-21
Wells, K (1981) Territorial behaviour of the frog,

Eleutherodactylus urichi in Trinidad Copeia 1981, 726-
728
Wilson, E (1975) Sociobiology Cambridge, MA:

Belknap Press

5. ADVANTAGES AND DISADVANTAGES OF DELAYED
BREEDING IN BIRDS
From an evolutionary viewpoint, it makes sense to
have as many offspring as possible over a bird's
lifetime. Yet there are some birds that have delayed
breeding. This means that breeding does not take place as
soon as the bird is physically able. For example, the
albatross first breeds at between 7-13yrs old.
There are a number of reasons for delayed breeding

(Gill 1995):
i) The life expectancy of the bird.
Birds that live longer have greater opportunities

for breeding over the lifetime, so can afford to wait.
For example the Wandering Albatross (Diomedea
exulans)(figure 5.1) produces one chick every two years
for up to 50 years (Gill 1995).
(Source: Mila Zimkova)
Figure 5.1 - Wandering albatross.
ii) The interval between generations (mean

generation time).
Successful breeding is not just about having many

offspring, but about them surviving into adulthood.
Spacing out the offspring allows more time to be spent on
their upbringing, and thus increases their survival.
iii) Costs of early reproduction can be severe.
Birds that breed young may be more likely to die

from lack of food or poor predator avoidance, and thus so
will the offspring die. Older birds may be better at
collecting food for the incubation/pregnancy period, and
finding nests that avoid predators.
Ainley and DeMaster (1980) studied the risk of

breeding among Adelie Penguins (Pygoscelis adeliae), who
show delayed breeding patterns. In one year, 39% of
breeding animals died compared to only 22% of non-
breeders. While 75% of 3-year-old females died at first
breeding compared to only 10% of 11-year-olds at first
breeding. There are survival advantages to delayed
breeding.
iv) Divorce more common among younger individuals of

monogamous species (eg: short-tailed Shearwater; Wooller
and Bradley 1996).
Table 5.1 compares delayed breeding to immediate

breeding in two birds.
ALBATROSS DUCK
Type of breeding delayed immediate
Survival before
breeding (per year)(%) 30 15
First reproduction
(age in years) 7-13 1
Fecundity (young
per year) 0.2 3
Adult mortality
(per year) (%) 5 50
(After Gill 1995)
Table 5.1 - Comparison of delayed and immediate breeders.
Table 5.2 lists the general advantages and

disadvantages of delayed breeding for birds.
ADVANTAGES

1. Older birds can learn about raising offspring from time helping
relatives raise offspring.
2. Older birds better skilled at collecting food, which important

when offspring born.
3. Less older birds die at first breeding.
4. Suitable for long-living birds that can produce offspring for many
years.
DISADVANTAGES
1. Bird could die before breeding and thus no offspring.
2. Not suitable for short-living birds.
3. Goes against basic principle of evolution which is to have as many

offspring as possible.
4. What does the bird do while waiting for age to breed?
Table 5.2 - Advantages and disadvantages of delayed

breeding.
REFERENCES
Ainley, D.G & DeMaster, D.P (1980) Survival and

mortality in a population of Adelie Penguins Ecology 61,
522-530
Gill, F.B (1995) Ornithology (2nd ed) New York: WH

Freeman
Woller, R & Bradley, S (1996) Monogamy in long-

lived seabird: The short-tailed Shearwater. In Black, J.M
(ed) Partnerships in Birds Oxford: Oxford University
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ANIMAL

1. All work is sourced to the original authors. The

This work is licensed under the Creative Commons

2. Details of the author are included so that the level

Kevin Brewer BSocSc, MSc

An independent academic psychologist, based in England,

Orsett Psychological Services,

Animal Behaviour: Advantages and Disadvantages No.1; Kevin Brewer; 2008 2

1. Advantages and Disadvantages of

2. Advantages and Disadvantages of

3. Advantages and Disadvantages of

4. Advantages and Disadvantages of

5. Advantages and Disadvantages of

Animal Behaviour: Advantages and Disadvantages No.1; Kevin Brewer; 2008 3

Most interest relates to bird migration, but other

Wildebeest Connochaetes taurinus Africa

Saiga antelope Saiga tatarica (figure 1.1) Central Asia

Mongolian gazelle Procapra gutturosa Mongolia

White-tailed deer Odocoileus virginianus North America

Table 1.1 - Examples of migrating mammals.

(Source: US Federal Government)

Figure 1.1 - Saiga antelope.

Animal Behaviour: Advantages and Disadvantages No.1; Kevin Brewer; 2008 4

2. Move to where food/prey available when not breeding, particularly

3. Stationary can mean increased predator risk.

4. Constant temperature conditions: escape bad weather and lower

5. Able to have specialist breeding site (eg: no predators) and

6. Flexible strategy - some members of the species can migrate and

7. Stationary animals risk exhausting food supply using it all year

8. Opportunity for different members of the species to meet, and

9. Ideal when specialist food required because the earth's resources

1. Large amount of energy required to travel long distances.

2. Problems and risks of navigation.

3. Risk of forgetting sites or not being able to find again.

5. Risk at temporary stopovers from lack of local knowledge about

6. Vulnerable to weather changes or poor conditions in one year.

7. Many decisions required including optimal fuel load and optimal

Animal Behaviour: Advantages and Disadvantages No.1; Kevin Brewer; 2008 5

10. Risks of night-time migration if animals normally active in day-

Table 1.2 - Advantages and disadvantages of migration.

Figure 1.2 - Migration routes of white stork.

(Source: US Fish and Wildlife Service)

Figure 1.3 - Green turtle.

Migration for feeding - 2, 7, 9

Table 1.3 - Specific advantages in table 1.2 depending

The length of migration varies from 1-2 metres per

(Source: Andreas Trepte)

Figure 1.4 - Migration routes of Arctic tern.

Studies of ringed populations of birds (figure 1.5)

Animal Behaviour: Advantages and Disadvantages No.1; Kevin Brewer; 2008 7

Figure 1.5 - The ringing process by "Cruzinha"

There are cases from displacement experiments of

At the other extreme, the common toad (Bufo bufo)

Within some species certain individuals migrate and

Animal Behaviour: Advantages and Disadvantages No.1; Kevin Brewer; 2008 8

There are sex differences in migration among robins.

STUDY MALE FEMALE

(After Harper 1988)

Table 1.4 - Percentage of robins who are sedentary by sex

(Source: Ramin Nakisa)

Figure 1.6 - Robin.

Whether the birds migrate or not depends on the

Animal Behaviour: Advantages and Disadvantages No.1; Kevin Brewer; 2008 9