Friday, April 17, 2020

Liberty Valance and the Pug-nosed Tree Frog

In the classic John Ford western, The Man Who Shot Liberty Valance, there is a traditional western shootout between an eastern lawyer, Rance Stoddard (James Stewart) and the vicious outlaw, Liberty Valance (Lee Marvin). Valance’s first shot injures Stoddard and just as Valance is about to finish him off, Stoddard manages to squeeze off a shot and Valance falls, dead. Unknown to the characters and the audience, at that point in the film, Valance was actually killed by Tom Doniphon (John Wayne), who shot a fraction of a second after Stoddard. Had the situation been real, a witness would have heard a single shot and thought that it was Stoddard’s gun that killed Valance.

This sonic illusion, called the precedence effect illusion (PEI),
occurs when two identical sounds, separated in space, are emitted close together in time. A witness perceives them as coming from the direction of the first emitted sound. This effect enables us to locate the true source of a sound and not be fooled by sonic reflections that bounce off other objects in the environment. Without the PEI we would be unable to identify the true source of a sound.

Animals such as frogs, birds, crickets, or katydids, use auditory signals (calls) to establish territories and attract mates. In most frogs neighboring males alternate their calls, giving an approaching female a clear choice between competing males. But a few frog species are known to call almost synchronously, potentially creating a PEI. It works like this: male A starts to call. A few thousandths of a second later, nearby male B calls. A female in the vicinity will perceive a single call coming in the direction of frog A, due to the PEI, and hop or swim toward male A.

Why did frog B not wait a bit before starting to call? If he delayed calling, he would stand out and have a better chance of attracting a nearby female. If mating preference was the only factor at work here then synchronous calling should never have evolved.

In Central America there are bats that use their keen sense of hearing to prey on calling male frogs. In addition, there are midges, mosquito-like flies, that hear with their antennae. They can locate a calling frog and, like mosquitoes, take a blood meal from it. In the act of feeding these midges can also transmit diseases to the frogs. Could such predators and parasites make synchronous calling advantageous? If they are subject to the PEI in the same way as humans they would be attracted to the frog that began calling first. Suddenly, synchronous calling doesn’t seem like such a bad choice when there are predators about.

Pug-nosed Tree Frog
(Wikimedia by Dario/CC0)
In Central America, found from Costa Rica through Panama, the Pug-nosed Tree Frog, Smilisca sila, is a synchronous caller and also preyed upon by the bats and midges. A research group, working at the Smithsonian Tropical Research Institute in Panama, recorded the calls of individual Pug-nosed frogs, creating a sound library. From this library they could use software to create an artificial chorus of frogs, broad cast from a pair of speakers separated by three to four feet. In this way they could randomly assign the calls from the sound library to each speaker and randomly determine which caller would be the leader (the first to initiate calling) and which the follower. The follower lagged the leader by 23 thousandths of a second, a value based on actual Pug-nosed choruses.

Captured bats were tested in a flight cage with a video recorder. Bats were judged attracted to a speaker is they approached it to a distance of less than 20 centimeters. To determine the attraction of midges traps were placed around the speakers and the number of midges collected at the leader and follower speakers were counted.

About 70% of the bat attacks were made on the leader speaker, 30% on the follower speaker. The response of midges was not as distinct, ~60% were trapped in the leader speaker, ~40% in the follower speaker. The difference, in both cases, was statistically significant.

So it appears that in Pug-nosed Tree Frogs it’s a matter of who croaks first!

Reference:
Legett, H.D., Hemingway, C.T., and Bernal, X.E. (2019). Prey Exploits the Auditory Illusions of Eavesdropping Predators. The American Naturalist doi: 10.1086/707719.