Advertisement calls of closely related species often differ in quantitative features such as the repetition rate of signal units. These differences are important in species recognition. Current models of signal-receiver coevolution predict two possible patterns in the evolution of the mechanism used by receivers to recognize the call: (i) classical sexual selection models (Fisher process, good genes/indirect benefits, direct benefits models) predict that close relatives use qualitatively similar signal recognition mechanisms tuned to different values of a call parameter; and (ii) receiver bias models (hidden preference, pre-existing bias models) predict that if different signal recognition mechanisms are used by sibling species, evidence of an ancestral mechanism will persist in the derived species, and evidence of a pre-existing bias will be detectable in the ancestral species. We describe qualitatively different call recognition mechanisms in sibling species of treefrogs. Whereas Hyla chrysoscelis uses pulse rate to recognize male calls, Hyla versicolor uses absolute measurements of pulse duration and interval duration. We found no evidence of either hidden preferences or pre-existing biases. The results are compared with similar data from katydids (Tettigonia sp.). In both taxa, the data are not adequately explained by current models of signal-receiver coevolution.
Various mechanisms have been proposed as the neural basis for pulse-rate recognition in insects and anurans, including models employing high- and low-pass filters, autocorrelation, and neural resonance. We used the katydid Tettigonia cantans to test these models by measuring female responsiveness on a walking compensator to stimuli varying in temporal pattern. Each model predicts secondary responses to certain stimuli other than the standard conspecific pulse rate. Females responded strongly to stimuli with a pulse-rate equal to half the standard rate, but not to stimuli with double the standard rate. When every second pulse or interval was varied in length, females responded only when the resulting stimuli were rhythmic with respect to the period of the standard signal. These results provide evidence rejecting the use of either high-/low-pass filter networks or autocorrelation mechanisms. We suggest that rate recognition in this species relies on the resonant properties of neurons involved in signal recognition. According to this model, signals with a pulse rate equal to the resonant frequency of the neurons stimulate the female to respond. The results are discussed with regard to both neural and evolutionary implications of resonance as a mechanism for signal recognition.
SUMMARYAcoustic pattern recognition is important for bringing together males and females in many insect species. We used phonotaxis experiments on a walking compensator to study call recognition in the katydid Neoconocephalus affinis, a species with a doublepulsed call and an atypically slow pulse rate for the genus. Call recognition in this species is unusual because females require the presence of two alternating pulse amplitudes in the signal. A Fourier analysis of the stimulus-envelopes revealed that females respond only when both the first and second harmonics of the AM spectrum are of similar amplitude. The second harmonic is generated by the amplitude difference between the two pulses making up a pulse-pair. Females respond to double pulses that have been merged into a single pulse only if this amplitude modulation is preserved. Further experiments suggest that females use a resonance mechanism to recognize the pulse rate of the call, supporting a neural model of rate recognition in which periodic oscillations in membrane potential are used to filter the pulse rate of the signal. Our results illustrate how a reduction in pulse rate extends the opportunities for females to evaluate fine-scale temporal properties of calls, and provide further evidence for the importance of oscillatory membrane properties in temporal processing. The results are discussed with regard to evolutionary changes in call recognition mechanisms within the genus.
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