Social systems: Socio-ecology and social organization Matrilinear

General behaviour and social
behaviour
Social systems
Natural history
Communication and social behaviour
Dominance
Predator avoidance
Foraging
Time budgets
Infant care
Abnormal behaviour
Practical exercises
o
o
o
o
o
o
o
o
o
o
Social systems:
Observed differences in group size
o
Terrestrial primates larger
groups
o
Frugivores larger groups
than folivores
o
Diurnal species larger
groups than nocturnal
Making observations of primate behaviour
Socio-ecology and social
organization
Anti-predatory advantages
Resource defence advantages
Different anti-predatory strategies
solitary
harem
monogamous
Group size depends
on how females are
distributed
o
Food choice
Predation risk
Males go where
females are
o
chimp
multi male multi female
Matrilinear subgroups
Dispersal
o
o
o
Clumped resources
Females philopatric
o
o
o
males disperse
Several subgroups
per group
Macaques and
baboons
Not hamadryas
clan
Minimizes risk of inbreeding
Usually, only one gender
disperses
o
Baboons, macaques, langurs –
males
Chimpanzees and spider
monkeys – females
Gorillas and orangutans – both
genders
Stressing process
Leaving social security
Exposed to harassment
Exposed to predation
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Consequences in the lab
o
The dispersing
gender has
difficulty in
tolerating
individuals of the
same gender
So how do you know?
o
Look in the literature
o
Look at your monkeys
By simply looking at the animals, you can find
out what kind of social system they have in
nature
Potential aggression
and injury
Big males – small females
o
Field studies
Large male canines
Competition
between males
o
Large testicles
Competition
between males
Female preferences
o
o
o
Sperm competition
Indirect competition
between males
Three traits in
males selected for
increased fitness in
partner
Indicators of good
genes
Sensational
(non-adaptive)
(territory defence,
paternal care)
2
Morphological adaptation and
social organization
o
Morphological adaptation and
social organization
harem
o
size dimorphism
small testes
canines - weapons
concealed
ovulation
multi-male
infanticide
female choice
Morphological adaptation and
social organization
o
monogamy
o
arboreal
± no size dimorphism
no ”weapons”
small testes
concealed ovulation
long tails, some with
grasping function
hands and feet grasping
function
hanging or climbing
locomotion
terrestrial
males big responsibility
(not gibbons - but
siamangs)
often very territorial
Behavioural adaptations and habitat
shorter tails
no pronounced grasping
function of hands and feet
often locomote on all fours
Behavioural adaptations and habitat
o
Marmosets
(tropical mountain
/ valley forests)
o
consortships - female
choice
Morphological adaptations and
habitat
o
sperm competition relative testes size
large
no pronounced size
dimorphism
signalled ovulation
Gouge holes in gum
trees
Patas monkey (savannah/desert)
Fastest monkey on earth: 55 km/h
3
Behavioural adaptations and habitat
o
Guenons
(treetops)
Behavioural adaptations and habitat
o
Visual
communication
(head bobbing)
Mona monkey
Diana monkey
Vervet monkeys
(riverine
woodland)
Alarm calls
Different predators –
different calls!
Spot-nosed guenon
Communication in primates
o
Vocalizations
Communication in primates
o
Alarm calls
Species recognition
Vocalizations
Territorial calls
Long-distance calls of
low frequency
Carry farthest in early
mornings
Communication in primates
Facial expressions (larger primates)
Attracting mates
Keeping track of group
members
Pygmy marmosets trill in
sequence, each awaiting
their turn
Agonistic calls
Aggressive and
submissive
o
Food calls
Communicates affective
state
Social factors, too
Recognizing individual
voices
Communication in primates
o
Pilo-erection (smaller primates)
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Communication in primates
o
Scent-marking
Female reproductive conditions
Communication in primates
o
Physical
Inhibition of ovulation in subordinate
callithrichids
Territorial scent marking
Identity
Age
Sex
Urine washing
Grooming
Play
Testicle grabbing
Baboons, muriquis
Squirrel monkeys and owl monkeys on
vegetation
Capuchins on themselves
Females advertising sexual receptivity
Ritualized stink-fighting
Ring-tailed lemur males during
breeding season
Why do animals play?
o
Probably not to
practice fighting!
Why do animals play?
o
Other areas in brain
activated
Other behaviours shown,
different sequences of
behaviour
Role reversal – animals
both win and loose play
fights
Self handicap – bigger
animals ”give up” and
loose
Nerve growth in
cerebellum?
Byers & Walker: ”Refining the motor training hypothesis for the evolution
of play.” American naturalist, 146: 25-40. 1995.
Why do animals play?
o
Teaches young to
deal with novelty
and surprise?
Posture
Balance
Coordination
Play in the laboratory?
o
Complex environment stimulates play
o
Novelty
Brain development
”vaccination” against stress
Stimulates the SEEKING-system
Curiosity, interest, expectation
Hypothalamus: dopamine
These neurons active while searching for food and shut off
when food is localized and eaten
Risky – Neophobia!
Spinka et al.: Mammalian Play: Training for the Unexpected.
Quarterly Review of Biology, 76 (2): 141-168. 2001.
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Social behaviour
o
Agonistic behaviour
o
How to assess dominance
o
threats and aggression
dominance displays
submission
redirected aggression
Usually, subordinate is more risktaking
Dominant has priority of access to resources
o
Aggressive behaviour
Affiliative behaviour
Exploration
o
play
sex / mounting
grooming
Shown by both dominant and subordinate
Submissive behaviour
Always shown by subordinate
Displacements / supplants
Bared-teeth displays
removing parasites
ritualization social function
Ranking order within matriline
Competition and rank
(macaques, baboons)
rank
o
o
Reduces cost of
aggression
Rank correlates:
Size, age, sex,
hormones, health
age
A daughter has the rank immediately below
her mother and above her older sisters
Ranking order in Callithrichids
Consequences in the lab
(marmosets, tamarins)
rank
o
Creating groups in captivity
age
Breeding female highest ranking individual. Older offspring
outrank younger offspring. Very young offspring have high
privileges, which they loose before adolescence
Structure cages appropriately
Visual barriers
Different heights available for sitting
x
y
Check for compatibility
Avoid unnecessary regroupings
Non-contact familiarization
α
β
αβ
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Group size and complexity
Compatibility
o
o
o
o
o
Clear dominance
No monopoly
No depression
Grooming relation
Reconciliation
Keep an eye out in
bigger groups!
Consequences in the lab –
Keeping track of others…
o
Socially interested – play a lot
o
Chimpanzee play in triads 25% of
play; baboons 3-5%
o
o
Adequate social grouping
o
Animals have knowledge of
the relationship between
other individuals
o
ex play-back experiment
baboon
rank in wrong context
vervet monkey
mother-offspring
o
o
Predator avoidance
o
o
o
o
o
o
Sentry behaviour
Mobbing behaviour
Vertical flight
Cryptic slow
climbing behaviour
Freezing; branch
dropping
Polyspecific
associations
No single housing
No crowding
Species-specific grouping patterns
Macaques and baboons: female relatives
Chimpanzees and spider monkeys: brothers
Compatibility checks
Visual barriers
Training opportunities
Predator avoidance in lab?
Saddleback
tamarin
Emperor
tamarin
o
o
o
Vertical dimension
Visual barriers
Habituation / training to participate
in procedures
Several species together
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Foraging
o
The lion.
o
Foraging
o
Opportunistic feeder
Search for prey
Rapid energy expenditure
(outcome unpredictable)
Banquetteer
Possessive behaviour
o
o
o
o
Two feeding bouts
early morning and late afternoon
Predictable food supply
Continous feeding
No food competition
Feeding habits matter
- influences on home range size
Foraging strategies among primates
o
Clumped food
Food competition
Mental maps of home range
Extractive foods
Lion-tailed macaques ”peel” foods lacking protective
covering
The gazelle.
Diurnal frugivore primates.
Frugivores
Insectivores
Folivores
Gumivores
Omnivores
Diana monkey
Folivores have smaller home ranges than insectivores and frugivores
15ha
70ha
Black and white colobus
Western red colobus
Generalist feeder
Specialist feeder
(all kinds of leaves)
Do primates share?
o
o
usually mother – infant
chimpanzee + capuchin
begging and celebration
passive tolerance - active generosity
selective and strategic generosity
punishing cheaters
(shoots, flowers, fruits)
Foraging vs silver platter servings
o
”meals” – for whose convenience?
o
Approach and ingest
Conditioned expectations of food availability
Feeding patterns vary
Foraging (visual and auditory senses)
Contact – searching or sentinel behaviour
SEEKING system activated
Acquisition – hunting or harvesting
Processing (olfactory and gustatory senses)
Preparing – reducing food to ingestible portions
Ingesting – masticating and swallowing
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Consequences in the lab
o
o
Most laboratory primate species
don’t share
Distribute food with care
Avoid monopolization
Visual barriers
Substrate – scatter feeding
Allow for foraging (looking for food)
Animals willing to work for access to food
Sympatric primates
o
Niche divergence
Canopy level
Degree of structural
support needed
Diet
Contrafree loading
Sympatric primates;
different diets
Sympatric primates;
different habitat use
Golden-handed
tamarin
Squirrel monkey
Putty-nosed monkey:
Mostly leaves
Sedentary prey
Crowned guenon:
More fruit and
insects
mobile prey
Moustached
monkey:
Intermediate
Dietary overlap is greater when food is abundant
Smaller species use thinner
branches, leap more, and use
forest understory and liane forest
Larger species use thicker branches,
climb more and use canopy more
Consequences in the lab
o
Feeding habits
Frugivores
Folivores
Active, search for food, eat buffé (moving)
Infants in mammals
o
Two mutually exclusive strategies:
o
Altricial species
More passive, eat banquet (sitting down)
o
Habitat use
Thick or thin branches preferred?
Leaping or climbing locomotion?
Canopy, understory or ground?
White-faced saki
o
Many, immature infants (6-15)
Dog, cat, rat, mouse
Precocial species
Few, mature infants (1-2)
Horse, cow, sheep, elk, zebra
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Lactation and suckling
Are primates altricial or precocial?
o
o
Continuum of strategies:
o
Sparse suckling
Neither!
Few infants (1 – precocial trait)
Immature (altricial trait)
o
Frequent suckling
Parenting in primates
o
Unique life history
o
o
o
o
socially important
Attachment important
for social development
Infant and juvenile
phases long - learning!
Deprivation studies maladjusted individuals
o
H Harlow noted the
development of a
strong attachment to
pieces of cloth in
separated infants.
Built mother
surrogate including
head and face
Harry Harlow, 1959
o
o
o
o
o
Non-explorative
Stereotypies
Abnormal social
behaviour
Abnormal sexual
behaviour
”motherless
mothers”
Milk low in nutrition (more lactose)
Social behaviour – attachment
ex: primates
Surrogate mothers
Many suckling sessions
o
not altricial / precocial
Nutritious milk (more fat and protein)
Practical behaviour
ex: rabbits
Consequences in the lab
o
Early separations infant – mother
might influence scientific studies
Behaviour
Endocrinology
Neurology
Physiology
Harlow & Zimmerman: affectional responses in the infant monkey. Science, 130: 421-432
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Abnormal behaviours in primates
o
Abnormal behaviours in primates
Socially inadequate
background
o
Self-aggression
Withdrawal, depression
Hyper aggression
”juvenile” behaviour
Abnormal behaviours in primates
o
Hair loss - alopecia
Un-stimulating environment
Hyper activity
Aggression
Stereotypies
Abnormal behaviours in primates
o
Hair pulling and ingesting
Hair loss - alopecia
Hair pulling and ingesting
Difficult to treat
Dermatosis
Hair shedding due to (social) stress
Hair shedding due to endocrine changes
Adjustment problems in stressful
group-housing environment
in macaques:
Remove stressor – remove problem
95% subordinate victims
3% self-directed
associated with moderate /
intense crowding
Temporary phenomenon
Hair shedding associated with seasonal
changes (molt)
Chronic boredom
Temporary phenomenon
Abnormal behaviours in primates
o
Hair loss - alopecia
Partial alopecia caused
by hair pulling
Improved by:
Opportunities for exercise
Social interaction
Positive interactions with humans
Food-searching activities
Exploration of destructible objects
Abnormal behaviours in primates
o
Hair loss - alopecia
Total alopecia caused by
spontaneous shedding
If not temporary, then
improved by:
Moving the animal to species-adequate
cage-housing arrangements
Reduces frequency but does not
”cure” the condition
Reinhardt, 2006: Hair pulling: a review
Reinhardt, 2006: Hair pulling: a review
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Abnormal behaviours in primates
o
Aggression in primates
Inescapable stress
o
Learned helplessness
Then: aggression - ”dispersal
mechanism”
Lassitude,
Weakness
Lack of interest in
environment
Self-mutilation
Difficulty in learning
Weight loss
Appetite loss
Social and sexual deficits
Reversible!
o
inter-individual distance defined by aggression
anti-social behaviour
Now: aggression - ”cohesive
mechanism” in social groups
studies of reconciliation
former opponents mutually attracted
social behaviour
Reconciliation
o
o
o
o
o
TOM - Theory of Mind
increased frequency of affiliation between
former opponents after conflict. Restorative
mechanism.
cooperation and tolerance increased.
stress decreased
reconciliation = strategic
valuable relationship hypothesis
o
o
apes - chimpanzees o urangutan
children
older than 18 months
TOM - Theory of Mind
Self awareness
Attribute intention
to others
Tactical
deception
monkeys
inspection, making faces
o
o
react socially (threat - submission)
- explains the high level of cooperation in
primates.
o
Self recognition - mirror experiment
(Gallup)
Experimental approach: the mark
test.
TOM - Theory of Mind
o
Cognitive empathy consolation
compare ”emotional
contagion”
picture oneself in someone
elses situation without losing
distinction between self and
other
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Tool use
o
o
o
Chimpanzee cultures
Chimpanzee and
orangutan in nature
All apes in captivity
Superior to tools used
by other animals:
o
o
o
Different tools used in
specific order
Planning ahead
Using tool to make tool
Modifying tools
New situations – new
ways of using tools
Time budgets
Are time budgets useful?
o
Different populations = different time
budgets
o
Only a few behavioural categories
Clutton-Brock 1977 (unknown location)
Sussman 1993: Mauritius
Wheatley 1980: Borneo
24%
25%
31%
37%
42%
45%
8%
39%
36%
13%
o
Locomotion
Foraging
Grooming
Inactive
Ethogram
o
Catalog of animal’s entire
behavioural repertoire
o
o
Habituation / video
Only most common behaviours shown
Unusual behaviours may be as important!
Valid in a single population in a specific
environment over a short period of time
Ethogram
o
Where?
Baboons: 200 behaviours
In 100h: 90% can be obtained
2-3h: general idea (30 or so behaviours)
But good overview
Discrete categories of behaviour – as little overlap as possible
Thousands of hours of observation
Termite fishing in several cultures
Taï (Ivory Coast): nutcracking
Gombe (Tanzania): drinking from leaf
o
Cage use
3 dimensions
”Out of sight”
Who?
Affiliation
Aggression
Active – passive
recipient
o
What?
Social behaviours
Affiliative behaviours
Aggressive behaviours
Foraging
Locomotion
Inactivity
Other
Comfort behaviours
Abnormal behaviours
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Observing behaviour
Behavioural observations today!
o
Focal observations
time
o
o
Behavioural states
Feeding AND contact sit
o
o
Behavioural events
o
o
o
3 groups
Get info and data sheet
Watch animals, take data
Summarize data
o
How
We meet again in one hour!
Scan sampling – how and when
o
Good technique if observational conditions are
less than perfect
Cons
Behavioural synchrony among animals unknown
Time consuming to collect and analyse data
If too much happens, observer can’t keep up
All social behaviour, their durations, the senders and
receivers, distance relations of the participants,
neighbor relationships and temporal patterns of
behaviour
How
The animal can be followed in the field...
o
Complete record obtained (%, rate, duration,
events, states...)
Good if you don’t know which behaviours are interested
ahead of time
Focal observations – pros and cons
Pros
Continuous observation of 4 individual for 10
minutes each
All behaviours (states and events) noted
continuously
When useful?
o
Monitoring group for 40 minutes
Behavioural events noted as they occur
Focal observations – how and when
o
o
Instantaneous sampling on whole group
Behaviour states noted every 30s for 40 minutes (with
breaks)
”All occurrences”
Instantaneous behaviour (sneeze, vocalizations)
Practical exercise
o
Scan sampling
Continuous behaviour (climbing, grooming etc)
Sometimes overlapping
Continuous observation of 4 individual for 10 minutes
each
All behaviours (states and events) noted continuously
o
Instantaneous sampling on whole group
Behaviour states noted every 30s for 40 minutes
(with breaks)
When useful?
When studying group behaviour and synchrony
% of time in various activites
Categories should be easily and quickly
distinguished
More suited to non-social behaviour
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Scan sampling – pros and cons
o
Pros
”All occurrences” – how and when
o
Data on a large number of group
members obtained quickly
Not very taxing for the observer
o
How
o
Cons
When useful?
Scanning a whole group is not
instantaneous
Error in duration estimates (focal animal
sampling better)
Events not recorded
”All occurrences” – pros and cons
o
Pros
o
Cons
Quality of ethogram
o
Behaviours mutually exclusive
Clumped categories
Stopwatch
Use tape recorder
Observer fatigue
Observational conditions need to be excellent
Very few behaviours can be monitored
simultaneously
Behaviour need to be attention-attracting
Behaviours don’t occur too frequently to record
Agonistic encounters
Sexual behaviours
Sampling techniques - hints
o
Rate of occurrence obtained
Similar to ”focal animal sampling” on whole
group for the behaviours monitored
Behavioural synchrony can be monitored
Monitoring group for 40 minutes
Behavioural events noted as they occur
Experience
Focal animal sampling more tiring than ”all
occurrences” or scan sampling
Two observers
Only behavioural events, not states
Literature
o
Smuts & Cheney
o
Strier
o
Russon, Bard & Parker
o
de Waal
o
Goodall
Primate societies
Primate behavioural ecology
Reaching into thought: the minds of the great apes
Good natured
In the shadow of man
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