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　　Vocalization in Frogs

The Túngara frog is a small terrestrial vertebrate that is found in Central America. Túngara frogs breed in small pools， and breeding groups range from a single male to choruses of several hundred males. The advertisement call of a male Túngara frog is a strange noise， a whine that starts at a frequency of 900 hertz and sweeps downward to 400 hertz in about 400 milliseconds. The whine may be produced by itself， or it may be followed by one or several chucks or clucking sounds. When a male Túngara frog is calling alone in a pond， it usually gives only the whine portion of the call， but as additional males join a chorus， more and more of the frogs produce calls that include chucks. Scientists noted that male Túngara frogs calling in a breeding pond added chucks to their calls when they heard the recorded calls of other males played back. That observation suggested that it was the presence of other calling males that incited frogs to make their calls more complex by adding chucks to the end of the whine.

What advantage would a male frog in a chorus gain from using a whine-chuck call instead of a whine？

Perhaps the complex call is more attractive to female frogs than the simple call. Michael Ryan and Stanley Rand tested that hypothesis by placing female Túngara frogs in a test arena with a speaker at each side. One speaker broadcast a pre-recorded whine call， and the second speaker broadcast a whine-chuck. When female frogs were released individually in the center of the arena， fourteen of the fifteen frogs tested moved toward the speaker broadcasting the whine-chuck call.

If female frogs are attracted to whine-chuck calls in preference to whine calls， why do male frogs give whine-chuck calls only when other males are present？ Why not always give the most attractive call possible？ One possibility is that whine-chuck calls require more energy than whines， and males save energy by only using whine-chucks when competition with other males makes the energy expenditure necessary. However， measurements of the energy expenditure of calling male Túngara frogs showed that energy cost was not related to the number of chucks. Another possibility is that male frogs giving whine- chuck calls are more vulnerable to predators than frogs giving only whine calls. Túngara frogs in breeding choruses are preyed upon by a species of frog-eating bats， Trachops cirrhosus， and it was demonstrated that the bats locate the frogs by homing on their vocalizations.

In a series of playback experiments， Michael Ryan and Merlin Tuttle placed pairs of speakers in the forest and broadcast vocalizations of Túngara frogs. One speaker played a recording of a whine and the other a recording of a whine-chuck. The bats responded as if the speakers were frogs： they flew toward the speakers and even landed on them. In five experiments at different sites， the bats approached speakers broadcasting whine-chuck calls twice as frequently as those playing simple whines （168 approaches versus 81）. Thus， female frogs are not alone in finding whine-chuck calls more attractive than simple whines. An important predator of frogs also responds more strongly to the complex calls.

Ryan and his colleagues measured the rates of predation in Túngara frog choruses of different sizes. Large choruses of frogs did not attract more bats than small choruses， and consequently the risk of predation for

an individual frog was less in a large chorus than in a small one. Predation was an astonishing 19 percent of the frogs per night in the smallest chorus and a substantial 1.5 percent per night even in the largest chorus. When a male frog shifts from a simple whine to a whine-chuck call， it increases its chances of attracting a female， but it simultaneously increases its risk of attracting a predator. In small choruses， the competition from other males for females is relatively small， and the risk of predation is relatively large.

Under these conditions it is apparently advantageous for a male Túngara frog to give simple whines. However， as chorus size increases， competition with other males also increases while the risk of predation falls. In that situation， the advantage of giving a complex call apparently outweighs the risks.