Potato and grain washing in macaquesPotato and grain washing in macaques –Imo Cream pilfering in blue titsCream pilfering in blue tits Gull hunting in captive orcasGull hunting in captive orcas Potential examples of animal culture

Inadvertent social information (ISI) use can produce differences among populationsInadvertent social information (ISI) use can produce differences among populations –E.g., mate choice copying Natural selection requiresNatural selection requires 1.Variation in a trait 2.Fitness implication 3.Transmission Not necessarily heritability!Not necessarily heritability! Social influence and culture

Recall…Recall… –V P = V E + V G –Does V E account for all environmental variance? V E = V DE + V AE + V AP + V ASV E = V DE + V AE + V AP + V AS –A = “additive”: variation is transmitted to offspring Fulfills transmission requirement of selectionFulfills transmission requirement of selection V DE direct influence of environmentV DE direct influence of environment V AE results from genotypic / environmental associationsV AE results from genotypic / environmental associations –E.g., Inheritance of a territory, influences of prior niche construction –Influence parent offspring regressions: lumped into V AG if not considered explicitly V E reconsidered I

V E = V DE + V AE + V AP + V ASV E = V DE + V AE + V AP + V AS V AP results from parental effectsV AP results from parental effects –E.g., the maternal environment, milk content, paternal care… –Late effects can be controlled by partial cross- fostering –Early influences are ~ impossible to control Generally end up (inappropriately) lumped with V AGGenerally end up (inappropriately) lumped with V AG V AS results from social influencesV AS results from social influences –E.g., ISI, teaching… –Additive effect can be very strong –Will be treated as V AG unless social influence is controlled V E reconsidered II

Why limit our interest to heritability (V AG /V P ) if selection and drift operate on components other than V AG ?Why limit our interest to heritability (V AG /V P ) if selection and drift operate on components other than V AG ? Transmittability = (V AG +V AE +V AP +V AS +V S*G ) / V PTransmittability = (V AG +V AE +V AP +V AS +V S*G ) / V P –Where V S*G is the genetic predisposition for social learning Measures the evolutionary potential of a population in terms of phenotypic change, whatever the mechanism of transmissionMeasures the evolutionary potential of a population in terms of phenotypic change, whatever the mechanism of transmission Transmittability

“Culture is the part (V AS ) of phenotypic variance that results from information transmitted across generations through social influences”“Culture is the part (V AS ) of phenotypic variance that results from information transmitted across generations through social influences” Cultural traits:Cultural traits: 1.Social learning 2.Transmitted across generations 3.Generalized across situations 4.Long term modification An evolutionary definition of culture

Learning: An adaptive change in behaviour through the effect of acquired informationLearning: An adaptive change in behaviour through the effect of acquired information –Includes imitation, copying, teaching, etc. –Strong role of ISI Social learning From Nordby et al Animal Behaviour 59:

Horizontal transmission alone does not result in tranmittabilityHorizontal transmission alone does not result in tranmittability Successive generations must interact to produce cultureSuccessive generations must interact to produce culture Transmission across generations Salmon returning to their natal streams is not a cultural traitSalmon returning to their natal streams is not a cultural trait –Culture is only part of non-genetic transmittability

Generalized female choiceGeneralized female choice Short lasting social influences will not tend to be transmittedShort lasting social influences will not tend to be transmitted Generalization and durability

Requires overlapping generationsRequires overlapping generations Some non-genetic transmittable information does not produce cultureSome non-genetic transmittable information does not produce culture –Parental effects –Territorial inheritance –Niche construction Cultural transmission vs. parental effectsCultural transmission vs. parental effects –Learning implies cognition –Cultural traits can be transmitted horizontally and obliquely Notes on culture

A meme is “that which is copied”A meme is “that which is copied” Individuals acquire memes (learn) and transmit them (culture)Individuals acquire memes (learn) and transmit them (culture) Mutate rapidly, non-randomlyMutate rapidly, non-randomly Population memetics Horizontal transmission homogenizes groupsHorizontal transmission homogenizes groups Increases differences between groupsIncreases differences between groups Can lead to group-level selectionCan lead to group-level selection The tax memeThe tax meme Dominance of cooperative memes can be enforced by moralistic aggressionDominance of cooperative memes can be enforced by moralistic aggression

1.Individual A modifies its behavior only in the presence of naïve observer B 2.A receives no immediate benefit or incurs a cost 3.A’s behavior results in B acquiring knowledge or skills more rapidly than it would have in the absence of A Teaching Older pup calls Younger pup calls

Geographic variation in bird songsGeographic variation in bird songs –True dialects –The four criteria LearningLearning Across generationsAcross generations Lasting modificationLasting modification Generalized (?)Generalized (?) –Arguably the strongest animal example of culture in non-humans –Effects on female choice, speciation? Bird songs and culture From Wilson et al The Condor 102: