Language Evolution and Computation Bibliography

Using a simple model of signaling of fighting ability, I demonstrate that; (1) conventional, cost-free, signals of fighting ability can be an ESS, (2) signals with significant costs can be used at ESS as long as they are used to indicate weakness rather than strength, (3) that if a set of signals is used to indicate a set of fighting abilities through their costs, they must decrease in cost for stronger signalers. The reason for this is that individuals of higher fighting ability have less to gain by avoiding escalated contests, and are thus more sensitive to signal costs. These results are of particular relevance to badges of status and other simultaneous signals used to settle contests over minor resources

This thesis investigates communication between animals with conflicting interests. Particular attention is paid to conventional signalling, in which signals are not inherently costly and information is inferred by convention. This type of signalling is emphasised for two reasons: firstly it is communication in its purest sense, and secondly it seems to more accurately reflect the properties of many biological signals. The costs which maintain the evolutionary stability of communication are of great interest because of the apparent benefit to be gained through the use of misleading signals, such as bluffs. I argue that these stabilising costs emerge from the manner in which receivers respond to signals, rather than being inherent to the signals themselves.

The theoretical papers in this thesis begin with the most basic signalling game and proceed towards a more general understanding of conventional signalling. I begin by investigating the importance of signal cost in the simplest possible model of communication, the Action-Response game. I demonstrate that the signals used do not have to be costly to be reliable, even when the signaller and receiver are in a state of conflict. I then consider the effect of adding costs to signals in a game in which reliable conventional signalling already exists, and demonstrate that the costly signals will be used by the weaker, not stronger, signallers. This demonstrates a stabilising mechanism fundamentally different from that of the handicap hypothesis, with its stabilisation through signal cost. Finally, I identify the conditions other than cost which are necessary for conventional signalling to be evolutionarily stable. These conditions relate to the information which both signaller and receiver must gain over the course of an interaction. Most models used to investigate signalling cannot account for behaviour seen in more complicated biological interactions because they are too simple to produce results other than that of the handicap prediction.

The other work included in this thesis addresses issues raised by the models. I review the literature on threat display use by birds, and present evidence that these displays are conventional signals. The stability of conventional signalling rests upon the existence of some common interest within a larger conflict between signaller and receiver. I present a clear example of communication attributable to common interest between fighting opponents. Cichlids of the species Nannacara anomala use a distinct colour signal, the Medial Line display, to coordinate another agonistic behaviour, tail-beating. It appears that both individuals benefit from the clearer assessment of relative fighting ability that this coordination affords. These N. anomala colour displays are quite conspicuous. It has been assumed that when common interest exists, signals will be very subtle, whereas when signaller and receiver are in conflict, signals will be exaggerated and conspicuous. Using an evolving neural-network model, I demonstrate that selection for exaggerated signals may exist even when the signaller and receiver have complete common interests.

I present a simple game, the Basic Action-Response game, which allows investigation of the claim that signals must be costly to be reliable. The Basic Action-Response game is the simplest communication game possible, by investigating its parameters we are able to define clearly ``conflict'', ``handicap'', ``communication'' and other relevant concepts. I explore the conditions on the magnitude of the stabilizing cost and handicap that must hold in order to maintain the evolutionary stability of signalling. It will be demonstrated that stable communication need not make use of costly signals at ESS, not even ``on average'', and that ``negative handicaps'' can be stable as long as the stabilizing cost is large enough.