The Pause That Enmeshes

by Michael Greenfield with Roger Martin

In most species of noisemaking animals, females prefer the sexual calls of some males over others. Usually, the preference is based on the energy of the call. Females often prefer loud calls or calls delivered at a high rate. But females also may exhibit preferences that have nothing to do with loudness or frequency.

For example, among many species of frogs and acoustic insects, females seem to prefer the male who makes the first of two or more calls that occur closely in time. In psychophysics, this is known as a "precedence effect," and variations of it also occur among animals that signal each other visually, rather than acoustically, such as fireflies and fiddler crabs.

A precedence effect influences human hearing, too -- although, as far as we know, not in the context of mate choice! That is, humans pay less attention to a second sound in a series when the second comes too close on the heels of the first.

It's currently unclear why animals would have evolved to give preferential notice to the first of two signals they receive from potential mates or other sources.

Precedence effects in hearing have had an effect on the evolution of calling behavior in animals. That is, evolution has favored the survival of males that do not call immediately after the onset of a neighbor's call. The males that wait increase their production of leading calls and presumably get more attention from females.

Now in situations where many animals are crowded together, there's a danger: A male's calling rate might drop sharply. Therefore, we hypothesize that the males learn to attend selectively to those around them -- just as humans, say, learn to focus on one person or a small group of individuals at a cocktail party and tune out others.

Túngara frogs are among the more common amphibians in Central America and northern South America. Female túngara frogs display a strong preference for leading male calls, and males are inhibited from calling following a neighbor. However, playback experiments conducted in the acoustic laboratories at the Smithsonian Tropical Research Institute in Panama demonstrated that males apply this inhibitory mechanism in relation only to a subset of nearby signaling neighbors. This research will be published in a future issue of Ethology under the title "Frogs Have Rules: Selective Attention Algorithms Regulate Chorusing in Physalaemus Pustulosus (Leptodactylidae)."

Selective attention is regulated by a combination of several rules. If those rules were contained in an instruction manual passed out to túngara frogs at birth, here's how they'd read:

In determining when to make your courting call

  • Pay attention to (allow yourself to be inhibited by) the loudest (usually nearest) neighbor and any additional ones whose calls are only 6 to 8 decibels weaker than the loudest.
  • Pay attention only to two neighbors when the calls are all weak or the second one is much weaker than the first.
  • Pay attention to three neighbors when the calls are loud or all neighbors are approximately equal in loudness.

These rules suggest that frog chorusing is a complex social feature that ultimately evolved in the context of sexual competition. Getting there first, or avoiding being drowned out by coming in second a hair after whoever does get there first, seems to be important to a male túngara frog in search of a mate.

 

Croak first:

Male Tungara frogs that do tend to get the girl.