The evolution of animal communication systems : questions of function examined through simulation
Simulated evolution is used as a tool for investigating the selective pressures that have influenced the design of animal signalling systems. The biological literature on communication is first reviewed: central concepts such as the handicap principle and the view of signalling as manipulation are discussed. The equation of “biological function” with “adaptive value” is then defended, along with a workable definition of communication. Evolutionary simulation models are advocated as a way of testing the coherence of a given theory. Contra some ALife enthusiasts, simulations are not alternate worlds worthy of independent study; in fact they fit naturally into a Quinean picture of scientific knowledge as a web of modifiable propositions. Existing simulation work on the evolution of communication is reviewed: much of it consists of simple proofs of concept that fail to make connections with existing theory. A particular model (MacLennan & Burghardt, 1994) of the evolution of referential communication in a co-operative context is replicated and critiqued in detail. Evolutionary simulations are then presented that cover a range of ecological scenarios; the first is a general model of food- and alarm-calling. In such situations signallers and receivers can have common or conflicting interests; the model allows us to test the idea that a conflict of interests will lead to an arms race of ever more costly signals, whereas common interests will result in signals that are as cheap as possible. The second model is concerned with communication during aggressive interactions. Many animals use signals to settle contests, thus avoiding the costs associated with fighting. Conventional game-theoretic results suggest that the signalling of aggression or of strength will not be evolutionarily stable unless it is physically unfakeable, but some recent models imply that cost-free, arbitrary signals can be reliable indicators of both intent and ability. The simulation, which features continuous-time perception of the opponent’s strategy, is an attempt to settle the question. The third model deals with sexual signalling, i.e., elaborate displays that are designed to persuade members of the opposite sex to mate. The results clarify the question of whether such displays are the pointless result of runaway sexual selection, or whether they function as honest and costly indicators of genetic quality. The models predict the evolution of reliable communication in a surprisingly narrow range of circumstances; a serious gap remains between these predictions and the ethological data. Future directions for simulation work are discussed.