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Kevin Brewer

ISBN: 978-1-904542-32-2
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Kevin Brewer BSocSc, MSc

An independent academic psychologist, based in England,

who has written extensively on different areas of
psychology with an emphasis on the critical stance
towards traditional ideas.

Orsett Psychological Services,

PO Box 179,
RM16 3EW

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 2
Page Number



2.1. Natural Selection 7

2.2. Sexual Selection 9


3.1. Safety From Predators 10

3.2. Food 13
3.3. Mate Access 18
3.4. Communal Care 19
3.5. Social Transmission of Information 21
3.6. Examples of Other Benefits 23


4.1. Increased Competition 25

4.2. Increase Risk of Infection 30
4.3. Exploitation by Other Animals 33
4.4. More Obvious to Predators 36
4.5. Risk of Inbreeding 36
4.6. Risk to Young 37


Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 3
(Source: US Federal Government)

Figure 1 - Herd of Caribou (Rangifer tarandus).

Different species of animals live in different ways
varying from solitary to massive groups. Some groups are
ordered and structured, and others are aggregations with
no order (individuals at the same place at the same time;
eg: flocks of seed-eating birds). Some groups are
permanent and others are temporary (eg: just for

Groups vary in size from two individuals to millions

(if not more) at the largest (eg: insects in super-
colonies). There are many different terms used for
groups, and mean and maximum size of groups (Reiczigel et
al 2008) (table 1).

Slater and Halliday (1978) distinguished seven types

of group situation (table 2).

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 4
Bonded pair.
Family group; eg red fox (Vulpes vulpes) mean 3, maximum 5.
Sibling group.
Harem - one male with females and offspring.
Matriline - dominant female, daughters and young.

Covey; eg quail.
Pack; eg: African wild dog (Lycaon pictus) mean 10, maximum 25).
Pod; eg killer whale (Orca orca) mean 5, maximum 15.
Troop; eg baboons.


Shoal/school - fish.
Flock - birds; eg American white pelican (Pelecanus erythrorhynchos)
mean 1600, maximum 15 000.
Herd; eg African buffalo (Syncerus caffer) mean 300, maximum 1500.
Aggregation - no order or structure.

Eusocial group eg ants
Super-organism - combined colonies

Table 1 - Examples of terms used to describe different

sized groups.

1. Solitary (eg sloth).

2. Aggregation for single activity - eg: meet in one place to mate,

but no group structure or organisation (eg: seals).

3. Aggregation for most activity - group without order or structure

(eg some grazing mammals.

4. Aggregation for some activity/stable group for others - eg:

aggregate at feeding sites, but organised group for rearing offspring
(eg: some birds).

5. Aggregation for some activity/solitary for some/stable group for

others (eg: bats).

6. Stable group - clear hierarchy and organisation to group (eg: pack

of wolves).

7. Solitary within stable group - solitary individuals join stable

group for a period (eg: dominant male takes over pride of lionesses).

Table 2 - Seven types of group structure according to

Slater and Halliday (1978).

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 5
How the species comes to live - alone or in a group,
and the group size - depends upon the evolutionary
advantages and disadvantages of doing so (table 3). Put
at its simplest, evolution is about staying alive
(feeding and not being eaten) and mating (passing genes
into the next generation). Thus group living must be seen
in relation to these behaviours.


1. Safety from predators.

2. Food; eg: co-operative hunting.
3. Mate access.
4. Communal care.
5. Social transmission of information.
6. Other benefits like thermoregulation.


1. Increased competition.
2. Increased risk of infection.
3. Exploitation by other animals.
4. More obvious to predators.
5. Risk of inbreeding.
6. Risk to young; eg: misdirected parental care.

Table 3 - Summary of the advantages and disadvantages of

group living for animals.

Group living and group size can vary within a Family

of animals, like African antelope (Daly and Wilson 1983;
Reiczigel et al 2008)(table 4). Body size determines the
food needed and where the animal can live. The biggest
animals live on grasslands in the largest groups, and the
smaller animals in the forest.


Kirk's dikdik (Madoqua kirki) 2 3

(small animal, 3-60kg, living in forest)

Mountain reedbuck (Redunca fulvorufula) 3 6

Impala (Aepyceros melampus) 7 80

African buffalo (Syncerus caffer) 300 1500

(large animal, 300-900kg, living on grasslands)

Table 4 - Group sizes of different African antelope.

There will always be a trade-off between the costs

and benefits of group living (Alexander 1974). This does
mean that some groups are dynamic and changing, known as

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 6
"fission-fusion" groups (Couzin 2006), depending upon
food availability or period in breeding cycle.

Popa-Lisseanu et al (2008) found both fission-fusion

groups and stable societies among giant noctule bats
(Nyctalus lasiopterus) studied in a Spanish park. The
composition of a single tree roost varied daily, but the
overall roosting groups were stable for over fourteen
years of the observations. The authors believed that
switching trees was a means of maintaining social bonds
in the whole of the colony. There are other advantages,
and disadvantages of sub-groups and small group switching
for animals in larger groups (table 5).


1. Flexible response to food availability.

2. Maintain social bonds in large group.
3. Anti-parasite strategy.


1. Conflict between sub-groups within the larger group.

2. Less defence against predators.
3. Less opportunity for benefits of thermoregulation.

Table 5 - Advantages and disadvantages of forming sub-

groups and small group switching within the larger group.

Evolution is the cornerstone of understanding non-
human behaviour. It is based around two central concepts,
proposed by Charles Darwin (1859, 1871): natural
selection and sexual selection.


This is the idea of the survival of animals within a

species with particular traits that give them an
advantage compared to others. This behaviour is
"adapted", and is well suited to the environment that the
animal lives in. These "fit" animals will survive and
leave more offspring, which means the spread of "adaptive
traits" in that species.
For example, running faster is an adaptive trait for
prey being chased by fast predators (figure 2).

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 7

Majority - animal A: runs slow**; few offspring in subsequent


2 offspring → 1 survive = 2 offspring → 1 survive = 2 offspring

Minority - animal B: runs fast*: many offspring in subsequent


1 offspring → 2 survive = 4 offspring → 4 survive = 8 offspring

KEY: * adaptive trait = run fast; ** non-adaptive = run slow; each

animal has 2 offspring

Figure 2 - Example of natural selection for adaptive


More formally, natural selection depends on three

principles (Dowling 1994):

i) Principle of diversity - there are a large number

of variant forms of the same species (known as members of
the population).

ii) Principle of interaction - these variant forms

interact with the environment to see which "fit"; eg:
animals that breathe air will not "fit" a permanent
underwater environment.

iii) Principle of differential amplification - the

variants that "fit" will spread at the expense of those
who don't "fit"; ie: more offspring.

In terms of leaving offspring, animals will have

evolved different strategies in relation to fecundity and
viability. The first term relates to the number of
fertilised eggs, and viability is the fertilised egg's
chances of surviving (table 6).


FISH high low Many eggs laid but

few survive

MAMMAL low high Few or single eggs

fertilised but most

Table 6 - Examples of fecundity and viability.

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 8

The best strategy for passing the genes into the

next generation will vary between the male and female of
the species. The male is able to produce many sperm, and
so can theoretically have as many offspring as mates
But the female is restricted, in most species, by
giving birth to the offspring. Thus she has more invested
in its survival (table 7).

EXAMPLE - Male mates with ten females, who have one offspring
each in the breeding season


MALE 10 fathered; can Find many female mates; ie:

afford some not indiscriminate; little concern
to survive for post-natal care

FEMALE Each female has Female invests time and effort

one offspring in survival, but must exercise
and thus survival choosiness about male; ie: only
important mate with male who has "best

Table 7 - Sexual selection and strategies for males and


Different species behave in different ways depending

upon their environments, but generally the example in
table is the common strategy of sexual selection. "Female
choosiness" has led to the evolution of males who
compete, in some way, to show the female that their genes
are best for mating. This competition involves fights,
"shows of quality" (eg: ornaments like a peacock's tail),
or the collection of scare resources to give to the
female ("resource-holding power"; RHP).

The ideas of evolution from Charles Darwin are based

upon the survival of the individual. But Dawkins (1976),
more recently, has suggested that it is the survival of
the genes that matter. For example, a mother who
sacrifices herself for her three offspring will guarantee
three copies of half of her genes survive. This has an
evolutionary advantage over the survival of the mother at
the expenses of her offspring. This has led to the focus
on "inclusive fitness" (the survival of the individual
and their biological relatives).

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 9
3.1. Safety from Predators

Individual animals are vulnerable to predators, but

groups give individual animals greater protection. Groups
aid defence from predators in a number of ways:

i) Increased vigilance

Larger groups of animals detect predators sooner

than smaller ones.
More pairs of eyes to spot approaching predators,
and greater possibility of alarm calls (the "many-eyes"
hypothesis or collective detection; Lima 1990). For
example, the Arabian babbler (Turdoides squamiceps)
produced approximately 50% more calls per minute at the
sight of a snake when in a group than when the bird is
alone (Ostreiher 2003).

An individual animal can spend more time eating (or

even sleeping) and less time watching for predators than
when alone. This advantage is especially important for
animals that have to feed for many hours per day in order
to gain sufficient calories (eg: grazers).

So all members of the group are alerted to a threat

as soon as one member detects it. Lima (1995) tested this
idea with mixed flocks of dark-eyed juncas (Junco
hyemalis) and American tree sparrows (Spizella
arborea)(figure 3).
Lima scared individual birds in the flocks (targets)
by rolling a rubber ball towards them. This produced an
alert response - stop feeding, adopt upright position,
and move towards cover. Only 40% of the ball rolls
produced an alert response in non-target birds. In other
words, one bird noticing a threat more often did not
cause the whole flock to respond. But when the ball
rolling frightened a number of birds simultaneously the
flock responded. It seems that multiple detections are
needed to alarm the whole flock in these birds.

Increased vigilance within a group is formalised in

"sentinel behaviour": "one member from a cooperative
group standing guard in a prominent position while the
rest of the group forages in comparative safety" (Wright
et al 2001).
But there are times when the "sentinel" uses their
position to scrounge or steal food from others rather
than doing their duty (Hellier 2002).

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 10
(Source: US Fish and Wildlife Service)

Figure 3 - American tree sparrow (Spizella arborea).

ii) Reduced risk of capture

Individual animals have less risk of capture by

predators because there are so many prey to choose from.
For example, hundreds of wildebeest crossing a crocodile-
infested river during migration. So many prey overwhelms
the predators, and only a limited number of individual
animals get caught.

iii) Confuse predators

Predators can be confused in different ways, varying

from inability to visually distinguish individual animals
from the group to prey escaping in different directions
when attacked.
While certain types of lemur combine their calls to
make them louder, making it sound like a much larger
animal, and thus scare predator away.

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 11
iv) Increased risk of injury for predators

Groups pose the risk of injury for predators in ways

like stampede threat on attack. For example, eagles find
it hard to attack flamingos in massive groups (figure 4)
because of the risk of injury to wings from hitting other
birds during an attack on one bird.

(Source: Duncan Wright)

Figure 4 - Flamingos (Phoeniconaias minor) at Lake Nakuru

National Park.

v) Easier to fight back if large number of animals

A group of animals can respond to a predator with

mobbing behaviour (eg: birds; Curio 1978). This is where
a large number of animals gather around a lone predator
and harass it. It has a number advantages and
disadvantages as a predator defence (table 8).

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 12

1. Deters predators.

2. Alerts others to predator's presence, and thereby removes the

threat of ambush.

3. Assessment of risk of predator ie their motivation to hunt and

level of danger. Prey can adjust their behaviour accordingly.

4. Can signal quality to potential mates.

5. Teaches young about predators by observing adults.


1. Risk of injury or mortality by approaching predator.

2. Use of energy.

3. Lost opportunity for other behaviours, like foraging and mating.

4. Risk that conspecifics may take advantage at this time to steal

food or mates.

5. Mobbing of non-dangerous animals is a waste of energy (eg:

meerkats; Graw and Manser 2007).

Table 8 - Advantages and disadvantages of mobbing


3.2. Food

Living in a group gives various benefits in relation

to food.

i) Co-operative Hunting/Group Foraging/Social Predation

Animals working together to hunt can tackle prey

larger than themselves, and combat the group defence of
herding (table 9).

1. Conserve energy.
2. Capture larger or dangerous prey.
3. Use different skills.
4. Increase food intake when food scarce.
5. Protect kill from scavengers.

Table 9 - Advantages of group foraging.

Lions (Panthera leo) (figure 5) use co-operative

hunting strategies.
A meat diet of a carnivore provides more energy than
a vegetarian one, but requires a greater expenditure of
energy to catch the meal. Hunting together helps in
Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 13
resolving this dilemma.
Certain lions run around potential victims and chase
them back to where other lions lay in wait (Stander
1992). Lions are strong but cannot run for long, and may
easily be outrun by lighter, faster prey (Riley 1995).
The chase is often limited to 100 metres, though they can
achieve a maximum speed of 30 mph (Server 1998).
Working together also allows them to bring down prey
that weighs much more than a single hunter could catch.
Possibly up to twelve times heavier than a single animal
could capture (Schaller 1972). Group foraging allows the
capture of prey that are dangerous to the single
predator. For example, one kick from a giraffe could kill
an adult lion.

(Source: Miroslav Duchacek)

Figure 5 - Lions (Panthera leo).

Hunting in groups also conserves energy for

individual animals. It allows the combination of
different skills: certain lions may be better at
perceiving prey, and others at chasing (Bailey 1995).
Schaller (1972) noted that lions on the Serengeti
Plains are not that successful in their hunts. When
hunting alone, only 15% of the time were the lions
successful compared to 30% for group hunting. Thus there

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 14
is a lot of energy expended when food is not gained.

Animals engaged in cooperative hunting benefit from

using less energy to capture the prey, and gain from the
calories eaten after the kill. Creel (1997) calculated
the overall benefit of Net Kilojoules (NetkJ) in African
wild dogs (Lycaon pictus)(figure 6) living in the Selous
Game Reserve, Tanzania. This figure is the calories
gained from eating after energy expended in hunting is
From observing 905 hunts, Creel calculated that pack
sizes of 10-11 adults produced a NetkJ of 30 000 per day
for each adult dog, and 80 000 for groups of twenty. The
figure for solitary hunting dogs was difficult to
calculate because the packs attack larger prey (eg:
wildebeest, impala) that a single dog could not. But
small pack sizes (less than four adults) averaged under
20 000 Net kJ per dog.
Furthermore, larger packs can make simultaneous
kills per hunt. Packs of 10-12 adults could kill a mean
number of 1.5 animals per hunt compared to only one for
very small packs.

(Source: Bart Sawnson)

Figure 6 - Wild dogs (Lycaon pictus) in Kruger National

Park, South Africa.

But, in lions, individuals in groups do not

automatically gain more calories than hunting alone.
Lions hunting in pairs had 10% more food intake per day
per individual than loners, but individuals in a group of
six adults had 50% less intake per individual than lone
hunters (Caraco and Wolf 1975).
The relationship between food intake and group size
is also affected by the availability of prey. Packer et
al (1990), after many hours of observation, noted that
group foraging was an advantage when prey was scarce. The
average food intake per adult female was eight kilograms
per day when hunting alone in times of scarcity, but over

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 15
11 kgs when in groups of five or six adults.
In times of abundance, there was no advantage -
average intake 10 kgs per day when alone compared to
eight when in group of four.

Co-operative hunting allows for specialist skills by

an individual animal. This can be seen in chimpanzees
hunting agile red colobus monkeys (Procolobus badius).
Individual chimpanzees take different roles like
"drivers" (scaring the monkey out of hiding), "blockers"
(limiting their escape routes), and "ambushers" (making
the kill)(Boesch and Boesch-Achermann 2000).

Gilby et al (2008) collected data on over six

hundred hunts by chimpanzees between 1996-2003 at
Kanyawara chimpanzee community in Uganda. They found that
two males seemed to initiate many of the group hunts
("impact hunters") rather than a group-initiated activity
as in lions. This may be because a small amount of meat
is a nutritious extra to the chimpanzees' usual plant-
based diet, whereas for lions meat is their main source
of food.

There are other examples of teamwork like crocodiles

who need others to hold large prey (eg: zebra) while
they break off part to eat. Small animals, like jackals,
can harass seal pup mothers one jackal at a time until
she is exhausted and the pup can be stolen, or one animal
distracts while the other steals the pup.

ii) Food sharing

Those animals with food can share with those animals

not successful in the hunt.
Wilkinson (1984) observed blood sharing by vampire
bats (Desmodus rotundus)(figure 7) in Costa Rica. One
hundred and ten regurgitations of blood were witnessed
and most were for genetic relatives. But between one-
quarter and one-third were non-genetic relatives. The
evolutionary benefit of sharing with non-genetic
relatives was a system of reciprocity that existed in a
cave. In other words, one bat gives tonight and can
benefit from receiving another night.

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 16
(Source: Unknown)

Figure 7 - Vampire bat (Desmodus rotundus).

iii) Help in finding food

One member of the group can communicate to others

where the food source is situated. Von Frisch (1967)
noticed that individual honey bees (Apis millifera)
(figure 8), returning to the hive, performed a "dance"
(particular movements) to communicate to the others where
flowers were to be found (direction and distance).
Subsequent research showed that sound and olfactory
communication was also used (Kirchner and Towne 1994).

iv) Defending the food

A groups of animals are better able to defend a kill

from conspecifics or scavengers while it is consumed, or
defend a territory containing food.

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 17
(Source: Bjorn Appel)

Figure 8 - Honeybees (Apis millifera).

3.3. Mate Access

Group living means that mates are readily available.

Some species may group together just for breeding as well
as permanent groups. While other groups are a male and
his females (harem)(eg: gorillas).
There are advantages and disadvantages of group
living here compared to solitary animals looking for a
mate (table 10).

Darling (1938) noted a faster reproduction in

crowded areas. This is the "Fraser Darling effect" - the
stimulation of reproduction by the presence of other
members of the species not the mating pair. It may be due
to social facilitation where behaviour is generally
influenced by the presence of others as opposed to alone.

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 18

1. Little energy expended in finding mate.

2. Availability of choice and variety of genes, including the best


3. Little risk of non-breeding season through failure to find mate.

4. Opportunity to mate with more than one partner.

5. Females mating with many males guarantees multiple support for

raising offspring.

6. In permanent groups, opportunity to accurately assess the quality

of other animals over a period of time.

7. Best for species with brief mating period, like one night a year
(scramble competition polygyny; Alcock 2005).

8. Allows "mate-sampling" (short-term liaisons) before finding life-

long monogamous partner (eg: barnacle geese; Constable 2002).


1. Risk of conflict and injury, particularly male-male competition.

2. Males have no guarantee that female will not mate with someone
else, unless they "mate guard" ("paternity certainty" hypothesis;
Ridley 1978).

3. If temporary group, males often leave after mating and females

left to raise offspring.

4. If permanent group, males may help raise offspring that are not
genetically related because of female extra-pair copulation (EPC).

5. Many permanent groups have dominance hierarchies which means that

subordinate animals limited in their mating opportunities.

6. Females in harems restricted and controlled by dominant males.

7. Risk of inbreeding in large groups.

8. Evolutionary costs of group living for males; eg: larger body size
for competition; strategies to guarantee paternity.

Table 10 - Advantages and disadvantages of group living

for mating.

3.4. Communal Care

There are four types of communal care (CC) according

to Gittelman (1985):

 Nuclear family with reproductive pair and offspring

from previous seasons; eg: beaver;
 Matriarchy with reproductive female only; eg: little
brown bat;

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 19
 Harem; eg: Northern elephant seal;
 Multi-male/multi-female group containing both related
and unrelated individuals; eg: lions.

CC has both advantages and disadvantages (table 11).


1. Predator defence.
2. Communal suckling.
3. Acquisition of food easier.


1. Attract predators.
2. Mix-up of litters.
3. Disease spreads quickly.
4. Infanticide possible.

Table 11 - Advantages and disadvantages of communal care.

The young are vulnerable to predators even more than

adults as well as needing large amounts of food. Group
living helps in these two problems.

a) Protection of the young

A group can defend the young by physical protection

(eg forming a circle around them in the presence of
predators), or by having members to look after them
("auntying") while the parents forage.
Some animals, like wildebeest, have "nursery herds"
formed by cows and their calves to aid defence of the
young (Clamp and Russell 1998).

b) Feeding the young

Groups give more "pairs of hands" to acquire food

for the young. In some cases, like lionesses, "communal
suckling" occurs where all the litter is fed by a
lactating female. Usually all the young are genetic
relatives in some way. Bertram (1978) calculated the
average coefficient of genetic relatedness between
members of the same pride to be 0.15 (15%). Because
females remain in the pride (and males leave), future
generations will still be genetically related to the
adult females, but in smaller amounts.
Lionesses in a pride show synchronous breeding which
means that they all benefit from co-operative cub rearing
(Packer et al 2001).

There is another benefit to group living here known

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 20
as "co-operative breeding". This is where individuals
(usually kin) aid the parents in raising the offspring.
It is estimated to occur in 8% of bird species and 3% of
mammals (Canestrari et al 2008). The kin members who help
tend not to breed themselves that season, but gain an
evolutionary advantage because some of their genes are
surviving in the form of nephews/nieces or grandchildren.
This is known as an indirect fitness benefit (Hamilton
Generally there is a positive correlation between
group size (ie: number of adult helpers) and number of
young who survive to adulthood. This has been found in a
number of bird species.

Canestrari et al (2008) showed this effect in

carrion crows (Corvus corone corone) in Northern Spain.
They collected data from ninety-nine bird territories
between 1995-2004. Reproductive success (ie: surviving
offspring) is related to success in egg hatching, and
survival of nestlings to age of fledging (ie: leaving
nest). The first is dependent upon stopping predators
eating the eggs, and the latter is influenced by that and
availability of food (particularly in first ten days of
Two parents produced an average of 0.5 fledglings
per breeding period compared to 0.8 for groups of four
adults. This was a significant difference.

3.5. Social Transmission of Information

Individuals living in groups can learn from other

animals, most notably through observation learning. For
example, macaque monkeys in Koshima, Japan were observed
to copy each other in washing a sweet potato in sea water
before eating (Imanishi 1957).
Or in terms of survival, learning from others which
new food is safe. Galef and Wigmore (1983) offered pairs
of rats (Rattus norvegicus) two new foods. One rat ate
their choice first ("demonstrator") and the other
observed. The observer mostly chose the same as the
demonstrator rat.

Franks and Richardson (2006) found evidence of

teaching of route from the nest to food by ants. One ant
("teacher") travels with another ant ("pupil"), known as
"tandem running", and the "pupils" learnt the route four
times quicker than when learning alone.

Bats changing tree roosts daily within a large

colony can transfer information about food sources and
colony members (Popa-Lisseanu et al 2008).

Chimpanzees (Pan troglodyte)(figure 9)have been the

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 21
most studied animal in terms of the social transmission
of information.

(Source: Thomas Lersch)

Figure 9 - Chimpanzee (Pan troglodyte).

Jane Goodall (1986) is probably best known for her

painstaking observations of chimpanzees in their habitat.
This research and that of others as part of seven long-
term projects have together produced 151 years of
chimpanzee observation (Whiten et al 1999).
From these projects has come observations of
differences in the behaviour repertoires of chimpanzees
that has been called cultural variation (Whiten et al
Whiten et al (1999) identified thirty-nine different
behaviour patterns between the seven sites in Africa:
Bossu (Guinea), Budongo (Uganda), Gombe (Tanzania),

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 22
Kibale (Uganda), Mahale (2 separate projects) (Tanzania),
and Tai Forest (Ivory Coast).
The differences in behaviours between the sites were
classed as culture where it appeared that the behaviour
had been learnt by chimpanzees at that particular site.
For example, eight behaviour patterns were seen as unique
to Tai Forest chimpanzees and three to chimpanzees at
Gombe. Table 12 gives some examples of behaviours.
"Each local chimpanzee community has a unique array
of specific traditions, representing a 'package' that can
be described as its local culture.." (Whiten 2005 p53).


Cracking nuts Hit with wood (hammer) against wood

(anvil), or stone hammer on wood anvil,
or stone hammer on stone anvil

Fishing for insects Put stick into termite mound and pull out
with insects on, wipe stick with hand and
place insects in mouth, or put stick
directly into mouth

Grooming Pick off parasite and squash in hand, or

squash on skin

Gaining attention Knock knuckles loudly, or slap branches,

or bend branches nosily

Table 12 - Examples of behaviour differences between


3.6. Examples of Other Benefits

i) Specialisation

Among eusocial insects, different shapes and sizes

have evolved to perform specialist tasks in the colony.
In the Asian Marauder ant (Pheidologeton diversus), for
example, "minor" workers are one five-hundredth of the
weight of "majors" (soldiers)(Randerson 2003).

ii) Thermoregulation

Huddling together at night for warmth; eg: Emperor

penguins (figure 10)(Sparks 1969).

iii) Control population level

In situations of high population density, only

stronger animals survive and are able to mate.

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 23
(Source: US Antartic Program Photo Library)

Figure 10 - Emperor penguins (Aptenodytes forsteri).

iv) Save energy on movement

Individual fish in massive schools use less energy

as they move compared to alone.

v) Weight gain

Fritzsche et al (2000) took the usually solitary

golden hamster (Mesocricetus auratus) and kept them in
female pairs for five weeks. Each animal had increased
their body weight by 25% compared to less than 5%
increase for the solitary animals.

vi) Synchronization of circadian rhythms

Groups of fruit flies, degus, birds, fish, bats and

beavers, but not rats and hamsters, all show a
synchronization of circadian rhythms (bodily rhythms over
24 hours) ie sleeping and eating at the same time
(Davidson and Menaker 2003).

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 24
4.1. Increased Competition

The presence of many animals means that there will

be more competition for food and mates, and the
consequent risk of fights and injury.

i) Competition for food

Groups find it less easy to hunt by surprise or

ambush compared to individuals. Thus hunting often means
chasing, and this is not the most successful method in
relation to energy costs (eg: 15% of chases successful
for lone lions; Schaller 1972).
The presence of large numbers can overwhelm food
sources (eg: eating all the grass). The increased size of
the population affects red deer, for example, directly
through lack of food, and indirectly through calf
survival and growth. Males tend to move area more in
situations of high population density. This may be
because they need more food to achieve the large increase
in body weight ready for the rut.
There is also the pressure to share food with other
members of the group.

ii) Competition for mates

Competition for mates in the group can mean that

some individuals may not find one, or there is guarantee
against cuckoldry, particularly for males, when a mate is
The competition between males for mates occurs
before mating and after. Competition between males can be
both direct and indirect.
Direct competition involves males confronting each
other and fighting, as with red deer (Cervus elaphus)
(figure 11). This leads to the evolution of larger body
size in the male compared to the female of the species.
This is known as body size dimorphism (Harvey and
Bradbury 1991), and is found most commonly in monogamous
or harem situations.
However, in situations where many males are living
with many females, a more indirect type of male
competition evolves. Here it is not body size that
matters because the males rarely confront each other. It
is the ability to produce a lot of sperm quickly. This is
known as sperm competition Parker 1970), and leads to the
evolution of larger testes relative to body size (eg: in

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 25
(Source: Oliver Deme)

Figure 11 - Red deer (Cervus elaphus).

Sperm competition occurs also in the size of the

penis and the number of sperm in an ejaculation. The
presence of other males in the vicinity as in multi-
male/multi-female groups leads to larger number of sperm
in each ejaculation. This requires larger testes to carry
them (Baker 1996). Table 13 shows some data for primates.
Gorilla who live in harems have an average testes
size of 10g relative to body size of 250kg. Testes are
0.02% of body weight. Male chimpanzees, who live in
multi-male/multi-female groups, have testes' size of
0.03% of body weight - 60g to 50kg body weight (Short




(adult male divided by adult 1.1 1.7
1.3 female weight; the higher the male
number, the larger the male competition
relative to the female) direct


(a measure of testes size after male
body size affects have been competition
removed; the higher the number, indirect
the larger the testes relative
to body size)
(* Harcourt et al 1981)
(After Clutton-Brock and Harvey 1984)

Table 13 - A comparison of body size dimorphism, relative

testes size and mating system among primates.
Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 26
Pre-copulation competition in red deer

The stags are competing throughout the rutting

period for access to females. They are competing in
asymmetric contests (Maynard Smith and Parker 1976), in
that individual animals differ in strength or fighting
How to signal that ability to other males? The most
obvious way is by winning the fight, but this is costly
in terms of the risk of injury, and the loss of strength
and using up energy.
Also during a fight, the females are left unattended
which could allow the opportunity for subordinate males
to mate with them. Dawkins and Krebs (1978) call this the
"sneaky fucker" strategy.

Because of the risks involved in fighting,

"ritualised fighting" or pre-fighting assessments have
evolved. These are contests that show which male is
stronger without actual fighting.
But weaker animals may be able to bluff in such
situations, and signal strength that they do not really
have. Thus the "ritualised fighting" must be such that it
is a genuine signal of ability.
Only a small number of approaches (20%) end in
fights. Where the stags did fight, they usually had equal
roaring rates (Clutton-Brock and Albon 1979). The stags
may take up to 30 minutes to assess each other.
This pre-fight assessment is an "Evolutionary Stable
Strategy" (ESS) in game theory (Maynard Smith 1982. It is
the best strategy based on costs and benefits of fighting
(table 14).
Game theory is a mathematical theory applied to
decision-making. An "Evolutionary Stable Strategy" (ESS)
is a strategy that if most members of the population
adopted cannot be bettered; ie: the best individual
strategy depends on the population strategy (Dawkins


eg: injury

BENEFITS Pre-fight assessment Always fight

FROM and "ritualised
FIGHTING fighting" evolves
eg: access
to females

NO BENEFITS No fighting Co-existence

Table 14 - Costs and benefits and strategies of fighting.

This trade-off between fighting and not fighting

seems a good strategy. However, the benefits are very

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 27
high; ie: the value of the resource is great. Thus the
stags are willing to risk injury in fighting because of
the limited availability of female each year. Mating
opportunities are so few that this makes the benefits of
winning so much greater. This can be seen in that the
peak time for fights, injuries and conceptions is mid-
October on Rhum (Clutton-Brock et al 1982) (table 15).


FIGHTING eg: mating eg: females only available
often in year for few weeks each year

Pre-fight assessment Pre-fight assessment

very important; little important, but males more
actual fighting willing to fight if necessary

Table 15 - Benefits of fighting

Stages in "ritualised fighting"

a) Roaring contest

The first stage of the contest between two males

involves the ability to roar. As the contest develops,
the stags increase their speed of roaring until one of
them cannot keep up. In experiments with tape recordings,
stags compete until the tape becomes too fast, and then
they withdraw.
This signal of strength cannot be bluffed because of
the great energy demands of roaring. Zahavi (1977) argues
that animals can signal their dominance by the degree of
cost it is willing to incur. Also the maximum and mean
roaring rates have been found to correlate with fighting
ability (Goodenough et al 1993).

b) Parallel walking

If a contest cannot be decided by roaring, then the

next stage is for the animals to walk side by side
allowing them to assess each other's body size.
Both males face in the same direction, and perform a
slow walk with antlers held high. Parallel walks often
occur between evenly matched males, and those who hold
harems (Alvarez 1993).

c) Fighting

This involves the interlocking of antlers and


Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 28
Post-copulation competition among males

After the male has mated with a particular female,

it is still important to stop other males from mating
with her. There are strategies used by the male and his
competitors (Wilson 1975) (table 16).



- Mating plugs and - Sperm displacement (remove

repellents first male's sperm)

- Prolonged copulation - Induce abortion and then


- Male remains attached - Infanticide of first male's

to female offspring and then

- Guarding

- Departure of mated pair

from rival males

Table 16 - Post-copulation competition strategies by


Within a group, behaviour takes place in the

presence of others. This can change behaviour, and is
known as the "audience effect" ("the effect of an
observing animal on the behaviour of another individual";
Plath et al 2008 p22). For example, in Siamese fighting
fish (Betta splendons)(Dzieweczynski et al 2006), male-
male aggression intensity can vary in the presence of an
audience of females or males, or no audience.

Plath et al (2008) investigated the effect of an

audience of a male on the mate choice of another male in
a species of fish, Atlantic molly (Poecilia mexicana)
found in freshwater in Central America and Mexico. The
experiment was conducted in a tank with different
compartments. In the middle area was the male ("focal
male"), and at each end a different female ("stimulus
This allowed the researchers to record the time
spent with each female (mate choice) as the end of the
tank chosen by the male. The "audience male" was in a
separate compartment in the middle of the tank, and so
could see all around, and be seen by the focal male. The
mate choice for either a large or a small female was
tested alone and then in the presence of the audience.
When alone the focal male spent an average of 394
seconds near the larger female and 110 seconds near the
smaller one. Forty-one males were tested individually.
Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 29
There was a clear preference for the larger female.

The presence of an audience male produced a

significant change in behaviour in the focal male. They
spent less time with the larger female and more time with
the smaller female, such that the average time spent with
both females was similar (approximately 160 seconds
each). In other words, with an audience, the male tried
to hide their preference for a particular female.

Why did this happen? One possibility was to avoid a

costly fight over a female. More likely, according to the
authors, an evolutionary response to sperm competition.
Females may mate with more than one male, and this is
risky for the first male. Assuming that a male competitor
will copy their choice, the first male appears to choose
the smaller female.

iii) Conflict of interest in decision making

Group decisions can produce a conflict of interests

for individual animals in that group. There is a need for
a "consensus decision" which will not cause the group to
split. For example, a flock of birds deciding whether to
leave a foraging area to find another, or finding a new
nest site in eusocial insects (Conradt and Roper 2005).
In the case of honey bees (Apis mellifera), the
"scouts" explore the local area for a suitable nesting
site. They communicate their choice to the hive, and
their consensus is accepted by the remaining 95% of bees
(Seeley and Visscher 2004). The majority have no say in
the decision and accept it. This produces a speedy
decision, which may be traded off against accuracy.
In many situations the group decision benefits the
individual, but, in others, the individual sacrifices
their preference to remain with the group. In very large
groups the individual animal simply follows those closest
to them.

4.2. Increased Risk of Infection

Animals living in close proximity are at higher risk

from the spread of disease than solitary animals. In
cliff swallows, for example, nestlings in massive
colonies (over 5000 birds) had five times more swallow
bugs on their bodies than in small colonies (less than
100 birds). Bugs reduced survival by 50%(Brown and Brown
Group living also leaves animals vulnerable to
parasite infection. Parasites survive by living within
host animals and following a particular life cycle, which
can involve "encouraging" one host to be eaten by a

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 30
predator, and thus the parasite moves on. For example, a
parasite may reduce the host's ability to move and thus
make them more vulnerable to capture.

Seppala et al (2008) studied the parasite,

Diplostomum spathaceum (trematoda eye fluke) in rainbow
trout (Oncorhynchus mykiss)(figure 12). This parasite
matures in bird intestines after being swallowed from an
infected fish. It is to the parasite's advantage that
their fish host is eaten by a bird predator. In fish, the
parasite lives in the eye lenses and reduces vision.
The researchers observed small shoals in
experimental tanks. Parasite-infected fish formed smaller
shoals than non-infected ones, and the shoals were less
compact when a predator threat existed. The distance to
nearest neighbour (DNN) 7 cm for infected fish and 3 cm
for non-infected fish. A predator threat was simulated by
moving a shape over the tank as if a bird flying over.
Group defences work in fish by large shoals tightly
bunching together.

By affecting the trout's vision, the parasite

reduces the host's ability to group, and thereby makes it
more vulnerable to predation. The eye fluke reduced the
trout's ability to shoal as group defence in a number of

 Trout less able to see food, increased starvation, and

less likely to remain in shoal. Individual fish more
vulnerable to predation;

 Trout unable to join and keep up with group, including

keeping distance to nearest neighbour;

 Less likely to see approach of predator.

But there is a risk for the parasite that the host

prey may be eaten by another fish (different type of
predator) which is not suitable for the parasite's

Davies et al (1991) found higher rates of mosquito

bites (and risk of malaria) in larger groups of primates
in South America. Those primates (eg: Bald-headed uakari;
Cacajao calvus)(figure 13) living in the largest groups
(around fifty individuals) had a 50% infection rate for
malaria compared to 5% for White-faced saki (Pithecia
pithecia)(figure 14) who lived in groups of three.
Mosquitoes detect animals odours (like carbon dioxide)
which will be higher in larger sleeping groups.

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 31
(Source: US Department of Agriculture)

Figure 12 - Rainbow trout (Oncorhynchus mykiss).

(Source: Evgenia Kononova)

Figure 13 - Male uakari (Cacajao calvus).

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 32
Eusocial insects may, in fact, benefit from improved
immunity from disease by group living. Traniello et al
(2002) found a "social transfer" of infection resistance
to fungus in dampwood termites (Zootermopsis
angusticolis). The immunity of non-immunized termites
improved in the presence of immunized nestmates, through,
for example, grooming behaviour.

4.3. Exploitation by Other Animals

Animals in groups are at risk of exploitation by

other members of the group. This is particularly so in
groups with dominance hierarchies.

i) Exploitation by dominant animals

Dominance hierarchies developed in groups as a way

of maintaining order and avoiding costly confrontations
too often. The dominant animals benefit from more (and
better) food, and access to mates. For subordinate
animals, life in a group can be hard.
Among meerkats in South Africa, where only one
dominant female and one dominant male breed each season,
the subordinate animals are "forced" to babysit, feed
pups, and guard the burrow (Clutton-Brock et al 2004).

The position in the hierarchy influences sexual

activity through sexual contraception in some species. In
naked mole rat (Heterocephalus glaber)(figure 15)
colonies, the queen (dominant female) only breeds and
urinary chemico-signals "switch off" the release of
hormones and ovulation in subordinate females (such that
90% never breed); thus ignored by males (Faulkes and
Bennett 2001). Social stress may also result from the
queen's bullying of females.

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 33
(Source: User: Cacophony)

Figure 14 - White-faced saki (Pithecia pithecia).

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 34
(Source: Ltshears)

Figure 15 - Naked mole rat (Heterocephalus glaber).

ii) Exploitation by conspecifics

Even in groups where there are not dominance

hierarchies, animals can be exploited by group members.
Food stealing (kleptoparasitism) is one example. The
animal who does not join the hunt saves energy and gains
from the food.
This is the risk of "free riders" or "cheaters" -
animals gaining without doing their share of the work.
Because of this risk, animals in a group would be
expected to watch other members to make sure they are
doing their "fair share". This has been observed among
humans (Fehr and Renninger 2004), but not, for example,
in birds like dark-eyed juncas and American tree sparrow.

Lima (1995) wanted to see if these birds would

monitor the vigilance of groupmates in the case of
"collective detection". Food deprived birds were added to
the flock. These birds would be concerned to eat and not
"do their turn" at watching for predators. The rest of
the flock did not change their vigilance behaviour
suggesting that they were not monitoring the groupmates'

Deception can take place between group members.

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 35
Observation at the Okavango lion project suggests that
the females have little bond to the males. Kat (2000)
reported the case of attempted deception by "Vouvray"
(lioness). She had found a carcass killed by a leopard,
and tried to call her cubs from the pride to eat. But two
adult males followed, so "Vouvray" took the cubs to the
water-hole away from the carcass. The males followed. She
then tried to sneak off with her cubs to the food, but it
failed and the males found the food.

4.4. More Obvious to Predations

Large groups are easy to spot for predators (ie:

more conspicuous), and there is less chance that such a
large group can hide.
It seems obvious that small groups are at more risk
in some species, but so are groups that are too large.
Thus there is an optimal size for groups in different

McGuire et al (2002) investigated the optimal group

size for prairie voles (Microtus ochrogaster)(figure 16)
based on a population living in Illinois, USA. The groups
were studied over seven years.
The ideal size of a group was three adults and
offspring. Larger groups were more likely to have
disappeared during the study, mainly due to predation by
weasels. For example, adults in groups of eighteen adults
had an average survival of less than fifty days compared
to 150 days for adults in the optimal group size.

(Source: National Park Service, USA)

Figure 16 - Prairie vole (Microtus ochrogaster).

4.5. Risk of Inbreeding

Mating is paramount, yet, at the same time, it is

important not to mate with those who are too genetically
related because of the higher risk of recessive traits
appearing. Recessive genes require both copies (ie: one
from each biological parent) to appear, and can transmit
genetic flaws.

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 36
Dispersion of the offspring at puberty is one
mechanism that animals can use to avoid incest. For
example, adolescent lions are driven out of their birth
pride and wander looking for other prides.
While the "Westermarck hypothesis" (Westermarck
1891) originally suggested that biology turns off sexual
arousal to close genetic relatives.

Research with humans by Wedekind et al (1995) has

highlighted another mechanism: females prefer the smell
of males who are different in terms of the immune system,
or specifically the major histocompatibility complex
(MHC) region of the genome, as measured by antigens in
body fluids. Such combination of different immune systems
help the offspring to fight parasites.
In their experiments in Switzerland, the researchers
asked women to smell the sweat on clothes of men
genetically related and unrelated to them. The women
preferred the smell of those unrelated, except when
taking oral contraceptives, where the opposite was true.

In eusocial insects, like bees, all individuals in

the colony are closely related (eg: sisters have three-
quarters of genes in common instead of usual half)
because the colony queen is the only female who mates
usually. It could mean that a genetic flaw or
vulnerability to infection becomes common in the colony.

4.6. Risk to the Young

There are risks to the young of being raised in a


i) Misdirected parental care

The young may suffer if the parent(s) fail to feed

and care for them because the care is misdirected to non-
genetic animals. This is also a risk for the parent(s).
Where many young are raised in proximity, it is
crucial for the mother to distinguish her own offspring.
Providing care for non-genetic offspring is an
evolutionary disadvantage, especially in mammals where
lactation is costly for the mother.
So the mother must be able to recognise her
offspring, and one mechanism is olfactory (smell).
Jesseau et al (2008) tested the ability of degu (Octodon
degus)(South American rodent) mothers to recognise the
smell of their offspring compared to sisters' offspring
(genetic relatives), co-nesting mothers' offspring
(familiar but genetically unrelated) and strangers'
offspring (unfamiliar and unrelated).

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 37
In the experiments performed, mothers could
discriminate the odours of their own pups (familiar own -
FO) from non-genetic related pups (familiar alien, FA,
and unfamiliar alien, UFA), but they could not
distinguish between familiar and unfamiliar non-genetic
related pups at two weeks old. However, with pups at six
weeks old, the mothers could not distinguish between the
odours of their own pups and co-nesting genetic unrelated
pups, but they could tell the difference between FA and
UFA. Importantly, degu pups are weaned by six weeks old,
so recognition of own offspring is not so crucial.
However, there is still a risk of social grooming or
uttering alarm calls to non-genetic related animals.
There is a risk of misdirected care against the
probability that the familiar alien young are distant
genetic relatives. Thus indirect fitness benefits to
nursing alien young.

But, even if mothers can recognise their offspring,

there are cases of mothers nursing alien young in the
group (eg lions; Pusey and Parker 1994).
What are the benefits in such situations?

 Defence of territory - females without territory have

limited opportunities to reproduce.

 Protection against infanticide - arriving males can

kill the cubs of the pride to bring the females back to
reproductive susceptibility. A group of lionesses can
protect the cubs from a single male.

 Cubs grow slowly and need baby-sitting while the mother

hunts. Death by disappearance from the nest is a risk.

ii) Risk to health of young

In larger groups where there are many animals (both

young and adults), the young may be injured or killed by,
for example, adults fighting which ends in the young's
The greatest risk to the young is death. Bourke
(2001) reported that in singly-mated queen wood ant
(Formica exsecta)(figure 17) colonies, worker ants killed
male offspring to maintain the balance of daughters to
sons (sex ratio). This is fraticide.

The young of some species are also at risk from

infanticide. In lions, for example, incoming males to the
pride will kill the cubs less than nine months old
already in the pride. Pusey and Parker (1992)
believed that up to a quarter of all cubs die this way.
The reason is sperm competition.

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 38
(Source: User: Cfp)

Figure 17 - Wood ants (Formica exsecta).

The infanticide causes the females to become

reproductively susceptible immediately, whereas normally
lionesses give birth every two years.
The benefits to the male are obvious from this
behaviour, particularly as the male will stay for around
two years.

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 39
Alcock, J (2005) Animal Behaviour: An Evolutionary Approach
(8th ed) Sunderland, MA: Sinauer Associates

Alexander, R.D (1974) The evolution of social behaviour Annual

Review of Ecology and Systematics 5, 325-383

Alvarez, F (1993) Risks of fighting in relation to age and

territory holding in fallow deer Canadian Journal of Zoology 71,

Baily, J (1995) Encyclopaedia of Animal Behaviour Oxford:


Baker, R (1996) Sperm Wars London: Fourth Estate

Bertram, B.C.R (1978) Kin selection in lions and evolution. In

Bateson, P & Hinde, R (eds) Growing Points in Ethology New York:
Cambridge University Press

Boesch, C & Boesch-Achermann, H (2000) The Chimpanzees of the

Tai Forest. Behavioural Ecology and Evolution Oxford: Oxford
University Press

Bourke, A.F.G (2001) Social insects and selfish genes Biologist

48, 5, 205-8

Brown, C.R & Brown, M.B (1986) Ectoparasitism as a cost of

coloniality in cliff swallows (Hirundo pyrrhonota) Ecology October,

Canestrari, D et al (2008) Reproductive success increases with

group size in cooperative carrion crows, Corvus corone corone Animal
Behaviour 75, 403-416

Caraco, T & Wolf, L (1975) Ecological determinants of group

sizes of foraging lions American Naturalist 109, 343-352

Clamp, A & Russell, J (1998) Comparative Psychology London:

Hodder & Stoughton

Clutton-Brock, T.H & Albon, S.D (1979) The roaring of red deer
and the evolution of honest advertisement Behaviour 69, 145-170

Clutton-Brock, T.H & Harvey, P.H (1984) Comparative approaches

to investigating adaptation. In Krebs, J.R & Davies, N.B (eds)
Behavioural Ecology: An Evolutionary Approach (2nd ed) Oxford:
Blackwell Science

Clutton-Brock, T.H et al (1982) Red Deer: Behaviour and Ecology

of Two Sexes Chicago: University of Chicago Press

Clutton-Brock, T.H et al (2004) Behaviour tactics of breeders

in cooperative meerkats Animal Behaviour 68, 1029-1040

Conradt, L & Roper, T.J (2005) Consensus decision making in

animals Trends in Ecology and Evolution August, 449-456

Constable, T (2002) Saucy geese BBC Wildlife April, p21

Couzin, I.D (2006) Behavioural ecology: Social organisation in

fission-fusion societies Current Biology 16, R169-R171

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 40
Creel, S (1997) Cooperative hunting and group size: Assumptions
and currencies Animal Behaviour 54, 1319-1324

Curio, E (1978) The adaptive significance of avian mobbing. I.

Teleonomic hypotheses and predictions Zeitschift fur Tierpsychologie
48, 175-183

Daly, M & Wilson, M (1983) Sex, Evolution and Behaviour (2nd

ed) Belmont, CA: Wadsworth

Darling, F (1938) Bird Flocks and the Avian Breeding Cycle

Cambridge: Cambridge University Press

Darwin, C (1859) The Origin of the Species London: Macmillan

Darwin, C (1871) The Descent of Man and Selection in Relation

to Sex London: John Murray

Davidson, A.J & Menaker, M (2003) Birds of a feather clock

together - sometimes: Social synchronization of circadian rhythms
Current Opinion in Neurobiology 13, 765-769

Davies, C.R et al (1991) Malaria infection rate of Amazonian

primates increases with body weight and group size Functional Ecology
5, 655-662

Dawkins, R (1976) The Selfish Gene Oxford: Oxford University


Dawkins, R (1989) The Selfish Gene (2nd ed) Oxford: Oxford

University Press

Dawkins, R & Krebs, J.R (1978) Animal signals: Information or

manipulation. In Krebs, J.R & Davies, N.B (eds) Behavioural Ecology:
An Evolutionary Approach Oxford: Blackwell Science

Dowling, H (1994) Horizon: The Man Who Made Up His Mind

London: BBC/Broadcasting Support Services

Dzieweczynski, T.L et al (2006) Effect of a dummy audience on

male-male interactions in Siamese fighting fish, Betta splendens
Behavioural Ecology 16, 1025-1030

Fehr, E & Renninger, S-V (2004) The Samaritan Paradox

Scientific American Mind 1, 14-21

Faulkes, C.G & Bennett, N-C (2001) Family values: Group

dynamics and social control of reproduction in African mole-rats
Trends in Ecology and Evolution April, 184-190

Franks, N.R & Richardson, T (2006) Teaching in tandem-running

ants Nature 439, p153

Fritzsche, P et al (2000) Effects of social stress on behaviour

and corpus luteum in female golden hamsters (Mesocricetus auratus)
Physiology and Behaviour 68, 5, 625-630

Galef, B.G & Wigmore, S.W (1983) Transfer of information

concerning distant foods: A laboratory investigation of the
"information-centre" hypothesis Animal Behaviour 31, 748-758

Gilby, I.C et al (2008) Economic profitability of social

predation among wild chimpanzees: Individual variation promotes
cooperation Animal Behaviour 75, 351-360

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 41
Gittelman, J.L (1985) Functions of communal care. in Greenwood,
P.J et al (eds) Evolution Cambridge: Cambridge University Press

Goodall, J.S (1986) The Chimpanzees of Gombe: Patterns of

Behaviour Cambridge, MA: Harvard University Press

Goodenough, J et al (1993) Perspectives on Animal Behaviour

New York: John Wiley & Sons

Graw, B & Manser, M.B (2007) The functioning of mobbing in

cooperative meerkats Animal Behaviour 74, 507-517

Hamilton, W.D (1964) The genetic evolution of social behaviour

I, II Journal of Theoretical Biology 7, 1-52

Harcourt, A.H et al (1981) Testis weight, body weight and

breeding systems in primates Nature 293, 55-57

Harvey, P.H & Bradbury, J.W (1991) Sexual selection. In Krebs,

J.R & Davies, N.B (eds) Behavioural Ecology: An Evolutionary Approach
Oxford: Blackwell Science

Hellier, C (2002) Criminal crows BBC Wildlife May, p22

Imanishi, K (1957) Identification: A process of enculturation

in the subhuman society of Macaca fuscata Primates 1, 1-29

Jesseau, S.A et al (2008) Mother-offspring recognition in

communally nesting degus, Octodon degus Animal Behaviour 75, 573-582

Kat, P (2000) Prides: The Lions of Morengi Holland: Southern


Kirchner, W.H & Towne, W.F (1994) The sensory basis of the
honeybee's language Scientific American June, 52-59

Lima, S.L (1990) The influence of models on the interpretation

of vigilance. In Bekoff, M & Jamieson, D (eds) Interpretation and
Explanation in the Study of Animal Behaviour: Vol 2. Explanation,
Evolution and Adaptation Boulder, Cl: Westview Press

Lima, S.L (1995) Back to the basics of anti-predatory

vigilance: The group-size effect Animal Behaviour 49, 11-20

Manning, A & Stamp Dawkins, M (1998) An Introduction to Animal

Behaviour (5th ed) Cambridge: Cambridge University Press

Maynard Smith, J (1982) Evolution and Theory of Games

Cambridge: Cambridge University Press

Maynard Smith, J & Parker, G.A (1976) The logic of asymmetric

contests Animal Behaviour 24, 159-175

McGuire, B et al (2002) Fitness consequences of sociality in

prairie voles, Microtus ochrogaster: Influence of group size and
composition Animal Behaviour 64, 645-654

Ostreiher, R (2003) Is mobbing altruistic or selfish behaviour?

Animal Behaviour 66, 145-149

Packer, C et al (1990) Why lions for groups: Food is not enough

American Naturalist 136, 1-19

Packer, C et al (2001) Egalitarianism in female African lions

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 42
Science 27/9, 690-693

Parker, G (1970) Sperm competition and its evolutionary

consequences in insects Biology Review 45, 525-567

Plath, M et al (2008) Audience effect alters mating preferences

in a livebearing fish, the Atlantic molly, Poecilia mexicana Animal
Behaviour 75, 21-29

Popa-Lisseanu, A.G et al (2008) Highly structured fission-

fusion societies in an aerial-hawking, carnivorous bat Animal
Behaviour 75, 471-482

Pusey, A.E & Packer, C (1992) Infanticide in lions. In

Parmigiani, S et al (eds) Infanticide and Parental Care London:
Academic Press

Pusey, A.E & Packer, C (1994) Non-offspring nursing in social

carnivores: Minimizing the costs Behavioural Ecology 5, 4, 362-373

Randerson, J (2003) Together we are strong New Scientist 15/3,

Inside Science, 1-4

Reiczigel, J et al (2008) Measures of sociality: Two different

views of group size Animal Behaviour 75, 715-721

Ridley, M (1978) Parental care Animal Behaviour 26, 904-932

Ridley, M (1995) Animal Behaviour (2nd ed) Oxford: Blackwell

Schaller, G (1972) The Serengeti Lions Chicago: University of

Chicago Press

Seeley, T.D & Visscher, P.K (2004) Group decision making in

nest-site selection by honey bees Apidologie 35, 101-116

Seppala, O et al (2008) Shoaling behaviour of fish under

parasitism and predation risk Animal Behaviour 75, 145-150

Server, L (1998) Lions New York: Todtri

Short, R (1994) Why sex. In Short, R & Balaban, E (eds) The

Difference Between the Sexes Cambridge: Cambridge University Press

Slater, P.J.B & Halliday, T.R (1978) Behaviour and Evolution

Cambridge: Cambridge University Press

Sparks, J (1969) Bird Behaviour Harmondsworth: Penguin

Stander, P.E (1992) Co-operative hunting in lions: The role of

the individual Behavioural Ecology and Sociobiology 29, 445-454

Traniello, J.F.A et al (2002) The development of immunity in a

social insect: Evidence for the group facilitation of disease
resistance Proceedings of the National Academy of Sciences, USA 99,
10, 6828-6842

Von Frisch, K (1967) The Dance Language and Orientation of Bees

Cambridge, MA: Bellknap

Wedekind, C et al (1995) Major Histocompatibility Complex-

dependent mate preferences in humans Proceedings of the Royal Society
of London 260, 245-249

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 43
Westermarck, E (1891) History of Human Marriage London:

Whiten, A (2005) The second inheritance system of chimpanzees

and humans Nature 1/9, 52-55

Whiten, A et al (1999) Culture in chimpanzees Nature 17/6,


Wilkinson, G.S (1984) Reciprocal food sharing in the vampire

bat Nature 308, 181-184

Wilson, E.O (1975) Sociobiology: The New Synthesis Cambridge,

MA: Bellknap Press

Wright, J et al (2001) Cooperative sentinel behaviour in the

Arabian babbler Animal Behaviour 62, 973-979

Zahavi, A (1977) Reliability in communication systems and the

evolution of altruism. In Stonehouse, B & Perrins, C.M (eds)
Evolutionary Ecology London: Macmillan

Advantages and disadvantages of group living for animals; Kevin Brewer; 2008 44