thesis-presentation_vers1-4_final_english-version
TRANSCRIPT
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Enhedens navn
Welcome!
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Institute of Biology, Section for Ecology and Evolution
DAILY PATTERNS OF GROOMING BEHAVIOUR
IN CHACMA BABOONS (PAPIO URSINUS)
Internal supervisor: Associate professor, DSc. Torben DabelsteenExternal supervisors: Dr. Guy Cowlishaw
PhD stud. Alecia Carter & Harry Marshall Institute of Zoology, Zoological Society of London
A test of the biological market theory
Tsaobis Baboon Project - Namibia
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TSAOBIS BABOON PROJECT
• Guy Cowlishaw
• Ecology & behaviour – Desert-adapted baboon population
• Since ~ year 2000 • Central Namibia – Tsaobis Leopard Park
• 2 PhD’s + 6 volunteers
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PROGRAMME
Introduction
Methods
Results
Discussion
Conclusion & perspectives
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Introduction
• Care for/cleaning skin, fur, feathers etc.
• Behaviour in many animals
• Especially among primates
• Auto- vs. allo-grooming
• FunctionHygienicStress reducingSocial
• Cost Time for other activities Vigilance
WHAT IS GROOMING?
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Introduction
WHY GROOM OTHERS?
Altruism between relatives • Kin selection theory (Hamilton, 1964)• Factor of relatedness, r• Hamilton’s rule: rB > C
Cooperation between non-relatives• Reciprocal altruism (Trivers, 1971)• Three conditions:1) B (receiver) > C (donor)2) High probability for future encounters3) Receiver and donor remember each other -> behaviour reciprocated
Does the same behaviour need to be reciprocated?
Not necessarily!
?
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Introduction
THE BIOLOGICAL MARKET-THEORY
• Primates-> social complex group-structure
• Many social interactions
• Seyfarth (1977), Nöe & Hammerstein (1994), Henzi & Barrett (1999)
• Grooming as currency1) Can be reciprocated for itself2) Can buy other ”services”
• Grooming can be exchanged for:1) Access to infants2) Support during aggression3) Tolerance during feeding
• The market is dynamic- Supply and demand- The value of the service/commodity- Individual characteristics
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Introduction
Rank diff.
Chacma baboons
• Females stay in natal group -> matrilines
• Adult males emigrate
• Strong linear dominance hierarchy
• Males fight for their rank
• Females inherit their rank from their mother
Alpha-maleAdult malesMatriline 1 ( -female, daughters/sons)αMatriline 2 (…)Matriline 3 (…)…Lowest ranking female
Socially complex group-structure
Relatedness
Social bond
Sex, age, rank, sociality
Sex, age, rank, sociality
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Introduction
• Grooming buys tolerance -> Rank diff. btw. groom pairs greatest in the morning
• Family and ’friends’ exhibit tolerance -> groom more in the evening
• Tolerance’s value matches payment -> Greater effort in the morning when rank diff. is great
• Feeding competition raises the value of the tolerance
• Lowest ranking + social + older individuals negotiate more tolerance
It is not random, who grooms who at what time of day
PURPOSE, HYPOTHESES & PREDICTIONS
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Methods
STUDY AREA• Swakop river• Two habitat types
Semi-desert
Riparian woodland
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Methods
THE BABOONS
L (n=24) J (n=36)
• Two groups
• Habituated to observers
• Individually identifiable
L sleeping cliffsJ sleeping cliffsJ & L sleeping cliffsWater holesCamps
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Methods
DATA COLLECTION
• May – November 2009
• Follow groups from dawn to dusk
• Training period
• 1 h. focal observations
- Behavioural state
- Dominance interactions
• 10 min. focal observations
- Events: Approach/retreat
- 5 m. distance
• Ad libitum dom. interactions
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Methods
DATA SET
1 groom bout = groom activity btw. same pair; pause of max 10 seconds
• Individual + groom partner -> repeated observations
• Several groom bouts in the same observation
-> Data not independent
• Many predictors
• Interaction-effects
General linear mixed model (fixed + random effects)
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Methods
MODELS
4 model categories
Model 1: Rank difference
Model 2: Relatedness
Model 3: Social bond
Model 4: Groom effort per bout
Groom pair of
1) all types of individuals
2) only adult females
Choice ofpartner
?
?
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Methods
PREDICTORS & STATISTICS
• Time of day, where the groom bout occurred
• Rank difference
• Relatedness
• Social bond
• Feeding competition
• Initiator’s rank, age & sociality
• Interactions – all variables with time of day
• Two statistical tests
1) Likelihood ratio test
2) Markov chain Monte Carlo test
Model with only the important factors
Only if not the response-variable
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Results
HOW MUCH TIME IS SPENT GROOMING?
• 1780 hours focal observations (1 t. focal obs.)
• 1844 groom bouts
• Juvenile og adult females spend most time (14%) – sub adult males least (3%)
Forage
TravelRest
Allo-groomingDrinkPlayAuto-grooming
All individuals (60)
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Results
DO PAIRS HAVE A HIGHER RANK DIFF. IN THE MORNING?
YES!
• Estimate±SE: -0.008 ±0.004, P<0.01
• Not for adult female pairs
• Relatedness correlates positively (P<0.001)
Effect of other factors?• Feeding competition: No
• Rank, age og sociality: No
Time of day
Rank difference
Model 1: RANK DIFFERENCE
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Results
DO RELATIVES GROOM MORE LATER IN THE DAY?
NO!
• BUT … more in the morning
• Estimate±SE: -0.008 ±0.004, P=0.09
• Also among adult female pairs
Effect of other factors?• Feeding competition: No
• Rank, age og sociality: No
Relatedness
Time of day
Model 2: RELATEDNESS
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Results
DO ’FRIENDS’ GROOM MORE LATER IN THE DAY?
NO!
Effect of other factors?• Feeding competition: No
• Age og sociality: No
BUT…• Adult females: Daily pattern
affected by initiator’s rank, P<0.05
Only when the groom bout’s
initiator is the dominant
Model 3: SOCIAL BOND
Late
Early High rank
Low rank
Strength of social bond
Time of day Rank
(dom
.)
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Results
GROOM PARTNER CHOICE
• The prediction about rank difference was correct!
• Mornings: Pairs with high rank diff. og relatedness
• No effect of feeding competition,
rank, sociality og age
BUT
• Within adult female pairs - effect of rank
-> when the dominant initiator’s rank is low
-> grooms with subordinate ’friends’ in the morning
?
?Model 1-3: RANK DIFFERENCE, RELATEDNESS & SOCIAL BOND
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Results
IS THERE A GREATER GROOM EFFORT EARLY IN THE DAY?
NO!
• But… greater later in the day
• Estimate±SE: 0.07 ±0.03, P<0.01
• Higher rank, less effort
Effect of other factors• Feeding competition: No
BUTThe daily pattern depends on:
• Relatedness, when initiator is subord.Time of day
Initiator’s groom effort
18 min.
2.5 min.
55 sec.
3 sec.
Model 4: GROOM EFFECT
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Results
GROOM EFFORT, WHEN INITIATOR IS SUBORDINATE
Depends on relatedness (P<0.05)
• Close relatives -> greatest effort in the morning
• Non-relatives
-> least effort in the morning
Influence of other factors?• Large rank difference (P<0.001)
• Higher rank (P<0.05)
• Older (P<0.05)
Greater effort
Late
Early Closely related
Not related
Groom
effort (subord.)
Time of day Relat
edne
ss
Model 4: GROOM EFFORT
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Results
GROOM EFFORT
• Generally smaller groom effort earlier in the day
• No extra effort in the morning with high rank difference
• Higher rank, smaller effort
• BUT - when initiator is the subordinate
-> greater effort when rank is higher
• No effect of feeding competition
• When the subordinate grooms a close relative
-> greater effort in the morning
Model 4: GROOM EFFORT
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Discussion
WHY GREATER RANK DIFFERENCE IN THE MORNING?
• In accordance with the biological market theory
• More beneficial to negotiate tolerance earlier
• Why not negotiate tolerance the whole day?
1) A subordinate also has parasites!
2) Has costs
3) Risk of aggression
The choice of who & when is important!
Model 1: RANK DIFFERENCE
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Discussion
WHY NO DAILY VARIATION IN RANK DIFF. IN AD. FEMALES?
They do not choose each other strategically during the day• They do not negotiate tolerance with each other
• High competition for adult females
The value is high – it pays all day long• Tolerance is returned over several days
Model 1: RANK DIFFERENCE
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Discussion
Tolerance is also negotiated between relatives?• More beneficial in the morning
• Lower risk of aggression
• Better ’rate’
Greater benefits of getting groomed in the morning?• Higher stress levels?
• More ecto-parasites?
WHY DO RELATIVES GROOM MORE IN THE MORNING?
Model 2: RELATEDNESS
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?
WHO IS CHOSEN IN THE MORNING?
Close relativeHigh rank difference
Close relative
Model 1 & 2: RANK DIFFERENCE & RELTEDNESS
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Discussion
WHY NO DAILY PATTERN IN ’FRIENDSHIP’?
• Tolerance is not negotiated between ’friends’
• More and closer social bonds -> higher fitness
BUT!
Why does a low ranking adult femalestart to groom a subordinate ’girlfriend’more in the morning?
• Competition for females/risk too high
• Females trades with e.g. males in the morning
• Special strategy: Use ’girlfriends’ to get groomed?
Model 3: SOCIAL BOND
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Discussion
WHY DOES FEEDING COMPETITION NOT HAVE AN EFFECT?
• Feeding competition does not vary considerably
• Dispersal behaviour? Optimal exploitation of available food sources?
• Competition for partners -> Not an unrestricted choice!
• If tolerance is negotiated -> less aggression
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Discussion
WHY IS THE GROOM EFFORT LOWEST IN THE MORNING?
• Does not support the hypothesis
HOWEVER
• Highest ranking individuals need to groom least
• Higher rank difference greater effort for the subordinate
• At the bottom of the hierarchy less effort especially limited in time?
ALSO FOUND
• Both high relatedness and rank difference in the morning
Groom bouts shorter in the morning (found!)
Mornings could be a hectic time
Model 4: GROOM EFFORT
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Discussion
EFFECT OF RELATEDNESS ON GROOM EFFORT
• Generally lower effort in the morning
BUT…
• The daily pattern depends on relatedness, when the initiator is subordinate
WHY?
• Great effort strengthens a social bond important after a night?
• Tolerance negotiated between relatives Beneficial in the morning
• Help a close relative removing parasites + reduce stress
It pays for a subordinate to spend extra time in the morning grooming a close relative
Model 4: GROOM EFFORT
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Conclusion
WHAT DID I FIND?
Grooming in the morning
• High rank difference
• High relatedness
• Low groom effort
It is NOT random who grooms who, when and for how long
In accordance with the biological market theory! tolerance is negotiated with both dominants and relatives the baboons structure their choice in an optimal way during the day
New hypothesis - ’Groom-less-with-more’
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Perspectives
’GROOM-LESS-WITH-MORE’ IN THE MORNING?
• Optimises the benefit• Bet on more individuals -> lower risk of losing all effort
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Perspectives
FUTURE STUDIES
• MISSING LINK! Grooming in the morning Tolerance later?
• What type of pair trade with each other in the morning?
• Higher stress levels and more parasites in the morning?
• Higher groom partner shift rates in the morning?
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THANK YOU!GENERAL SUPERVISIONTorben Dabelsteen (DK)Guy Cowlishaw (UK)Alecia Carter (AU)Harry Marshall (UK)
STATISTICSRoger Mundry (GE)Thorsten Balsby (DK)Gösta Nachman (DK)Michael K. Borregaard (DK)Karina Banasik (DK)
DATA COLLECTIONAlecia Carter (AU)Harry Marshall (UK)Katherine Forsythe (AU)Rebecca Bodenham (UK)Will Symes (UK)Jenie Iles (NZ)Will Birkin (UK)Hannah Peck (UK)
INSTITUTIONSInstitute of Biology, University of CopenhagenInstitute of Zoology, Zoological Society of LondonDesert Research Foundation of NamibiaNamibian Ministry of Lands and ResettlementNamibian Ministry of Environment and Tourism
OTHERSSolveig Walløe HarpøthNana HeslerMikkel Bjelke Kristiansen
FINANCIAL SUPPORTCarlsberg’s Fond for Studenterrådet ved Københavns UniversitetDitlev Marcussen Buch og hustru Maren Buch, Baltzergaard StiftelseFriedrick Wilhelm Frank og hustru Angelina Frank’s MindelegatGreve A. Brockenhuus-Schacks Legat for den slesvigske ungdomGrosser Wilhem Rackwitzs LegatHotelejer Anders Månsson og hustru Hanne Månssons MindelegatKolding Gymnasiums VennerKøbenhavns Kommunes Legat for uddannelse mv.Købmand Jørgen Sørensen og hustru’s fondOticon Fonden Studiehjælpen Valdemar Shiøtts MindeThorkild og Thea Rosenvolds Legatfond