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BoundedRationality_TheAdaptiveToolbox.pdf

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242 Kevin N. Laland<br />

example, bats that are unsuccessful in locating food alone follow previously<br />

successful bats to feeding sites (Wilkinson 1992). Starlings can use the foraging<br />

success of other birds to assess patch quality, and exploit this information in their<br />

judgements as to whether to stay or switch patches (Templeton and Giraldeau<br />

1996). For redwing blackbirds, the social learning of a food preference is a£<br />

fected by whether the demonstrator bird becomes sick or remains well (Mason<br />

1988). At this stage it is not clear which, if any, of these heuristics provides the<br />

best description of the rules that animals employ when they learn from others.<br />

INNOVATION AND DIFFUSION<br />

When a novel behavior pattern spreads through an animal population, typically<br />

one individual will be its inventor or discoverer. There have been precious few<br />

empirical investigations of who these innovators are. However, evidence is beginning<br />

to emerge that innovators are sometimes individuals for whom the current<br />

prevailing risk-averse foraging strategies are unproductive, and who are<br />

driven to search for alternatives, perhaps incurring some risk. Such animals are<br />

often small, young, low-ranking, or poor competitors. Primate studies appear to<br />

indicate that innovators are frequently low in rank or poor competitors on the<br />

outskirts of the social group (Kummer and Goodall 1985). For instance, Sigg<br />

(1980) found that peripheral female hamadryas baboons were significantly<br />

better at learning novel tasks than central females. A recent review found that of<br />

38 independent cases of innovation reported in the primate literature, 25 were by<br />

low-ranking individuals (S.M. Reader, pers. comm.).<br />

Kummer and Goodall (1985) found no shortage of innovation in primate<br />

populations, but these innovative behaviors rarely spread, in spite of their apparent<br />

utility. It seems that often individuals do not adopt a better alternative, even<br />

when other members of the population exhibit such behavior. This is perhaps because<br />

on many occasions when ecological and technical innovations occur, the<br />

innovator is alone, or at least freed from social distractions (Kummer and<br />

Goodall 1985). If the twin observations, that it is poorer competitors that inna<br />

vate, and that poorer competitors find themselves on the outskirts of aggregations,<br />

are common to many species, much innovation will tend to occur<br />

unobserved by others, decreasing the chance that the novel behavior pattern will<br />

spread. This may be one explanation for both the slow diffusion of new behaviors<br />

in the wild (Kummer and Goodall 1985), and the rarity of stable animal traditions<br />

(Boyd and Richerson 1988).<br />

Empirical research has established a number of factors that influence the stability<br />

of animal traditions, including the number of individuals in a social group,<br />

the rate of change of group composition, and the amount of time available to forage<br />

(Galef and Allen 1995). Social transmission may be more stable when it reinforces<br />

a prior preference (i.e., for a more palatable diet) than when it conflicts<br />

with one (Laland and Plotkin 1991). Whether such pre-established biases reflect

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