Trust and deceit in the animal kingdom Liesbeth Sterck Animal Ecology, UU Ethology Research, BPRC Trust Nature in tooth and claw • Monkey business • Sly as a fox Cooperation in theory • Evolution on level individual • Altruism: benefit for other, costs for individual • Problem with cheaters Cooperation in theory • Mutualism: simultaneous exchange of benefits • Altruism: donor incurs cost; recipient benefits – Kin selection – Reciprocal altruism Kin selection (Hamilton 1964) • Altruism when: • C < r*B • Costs < relatedness * Benefits Kin selection (Hamilton 1964) • C < r*B r=1/2 r=1/4 r=1/8 Kin selection (Hamilton 1964) • C < r*B Animals can be nice to kin Cooperation with non-kin? http://www.youtube.com/watch?v=o2B1AJ3ZaUA Reciprocal altruism (Trivers 1971) • ‘If you scratch my back, I’ll scratch yours’ • Non-kin • Not simultaneous Prisoners Dilemma • Cooperate or cheat (defect) Reciprocal altruism (Trivers 1971) • • • • • ‘If you scratch my back, I’ll scratch yours’ Non-kin Not simultaneous Repeated encounters Individual recognition, e.g. in a group Humans • Humans are a group-living species – Family / kin – Friends / acquaintances / colleagues -- Warfare Groups are diverse • Size – From 2 - 100 - 10.000s individuals • Composition – Changing- fixed • Recognition group members – Anonymous –categories – individuals • Reproduction – One female (+ male) - everyone Cooperation in practice 1 • Why do primates live in groups? Group living: advantages • Warning against predators (van Schaik 1989) Group living: disadvantages • Competition (Wrangham 1980; Sterck et al. 1997} Females: – Food Males: – Matings Group living: advantages • Infanticide avoidance (Sugiyama 1965; 1966; Hrdy 1977; Sterck et al. 1997) Balance in costs and benefits Protection against predators and infanticide Competition Primate peculiarity: groups contain both females and males Cooperation in practice 2 • How do primates live in groups? How primates live in groups • How complex is living in a primate group • Do they take benefit others into account Methods • From individual to group: • Observations • Behavioural experiments Methods • From individual to group: • Observations • Behavioural experiments • Computer simulations How primates live in groups • How complex is living in a primate group – Dominance – Good relationships Bob (29) Relationships in group (Massen, Sterck & de Vos 2010) 1.6 Sitting together 1.4 1.2 20 29 Timon (20) 1 0.8 0.6 0.4 0.2 0 verwant 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18niet 19familie 20 21 22 23 24 25 26 27 28 30 31 32 33 34 35 Kin Dominance Friendship Good relationships Advantages male – female relationships Advantages Friendship and mating success Grooming by males (sec/hour) (Massen, Sterck et al. 2012) 120 * Mating wel paren 100 niet No paren mating 80 60 40 20 0 2006/07 2007/08 Mating season Friendship and paternity (Massen, Sterck et al. 2012) Proximity of male (rank) 6 * 5 4 3 2 1 0 otheralle male vader father males Friendship • Do they know their friends? Computer simulation of behaviour (Hemelrijk 1998, 2000; Evers, Sterck et al. 2011, 2012, 2014) Computer simulation of behaviour (Evers, Sterck et al. MS) Dominance Grooming= friendship Dominance Computer simulation of behaviour (Evers, Sterck et al. MS) 1.6 1.4 1.2 1 0.8 0.6 Dominance Grooming = friendship Dominance 0.4 0.2 0 niet verwant familie Primate group living • Compete and cooperate in group • Relationships – Kinship – Dominance – Friendship = ‘trust’ • Friendships take time to build and last long Primate cooperation in practice 2 • Do they take benefit others into account? – Other-regarding preferences – Inequity aversion • Yerkes 1930’s: http://www.youtube.com/watch?v=zrv91Pa3jgs Cooperation: pro-social behaviour 35 (Massen, vd Berg, Spruijt en Sterck 2010: PlosOne 5(3): e9734) • Do primates bestow other a favor? Cooperation: pro-social behaviour (Massen, vd Berg, Spruijt en Sterck 2010: PlosOne 5(3): e9734) • To kin • To high-ranking animals Bestow Neutral Withhold High rank 36 Low rank Inequity Aversion "Inequity exists for a person whenever his perceived job inputs and/or outcomes stand psychologically in an obverse relation to what he perceives are the inputs and/or outcomes of another" (Festinger 1957) 37 Inequity Aversion • Negative inequity aversion – React to getting less than other • Positive inequity aversion – React to getting more than other 38 Inequity Aversion (Brosnan & de Waal 2003) http://www.youtube.com/watch?v=KSryJXDpZo capuchin monkey refuses food 39 Inequity Aversion (IA): critisism • Newer data: no IA 40 The task for macaques 41 The Conditions 1. No effort (provisioning) Equity 2. No effort (provisioning) Inequity 3. Small effort Equity 4. Small effort Inequity 5. Large effort Equity 6. Large effort Inequity 7. Large effort Reward & effort inequity 42 O,35 kg 2, kg O,35 kg 2, kg Proportion of acceptance/performance of all equity conditions for all animals Friedman test: n = 19, 2 = 15.84, df = 2, p = 0.001 Post-hoc Wilcoxon ranks tests: Significant dropsigned in performance (for low value reward) with increasing workload Provisioning vs. 0,5 kg: n = 19, T+=62.5, pexact= 0.115 Provisioning vs. 2,3 kg: n = 19, T+= 152, pexact< 0.001 43 0,5kg vs. 2,3 kg: n = 19, T+= 117.5, pexact= 0.008 Proportion of acceptance/performance of dominant subjects Wilcoxon signed ranks tests: Disadvantageous Inequity in =small Provisioning: n = 12, aversion T+= 21, pexact 0.719effort test Small Effort: n = 12, T+= 34, pexact= 0.023 Large Effort (reward): n = 12, T+= 8.5, pexact= 0.211 Large Effort (reward & effort): n 44 = 12, T+= 15.5, pexact= 0.250 AIA: Proportion of acceptance/ performance of subordinate partners Wilcoxon signed Performance forranks low tests: value reward decreases when workload increases Provisioning: n = 9, T+= 3, p = 0.156 T += exact Small Effort: n = 9, 4, high pexact= 0.219reward At high workload, performance for value Large Effort (reward): n = 9, T+= 0, pexact= 0.008 significantly better then for low value reward Large Effort (reward & effort): n 45 = 9, T+= 4, pexact= 0.055 --> no advantageous inequity aversion Nature in tooth and claw? • Monkey business • Sly as a fox Care and friendship • C < r*B Conclusion • Deceit kept in limits: otherwise no cooperation • Competition and cooperation in group – Dominance and friendship • Trust and friendship crucial for cooperation Thank you Humans are smart and have large brains Social Intelligence Hypothesis • Primates have relatively large brains • Both cooperation and competition in group • Several hypotheses – Primates ‘smart’ through social complexity • Socially complex = socially intelligent Social intelligence • Evidence in favour: • Group size and brain size are correlated in primates, cetaceans and carnivores (Dunbar 1998)