A variety of structural parameters were measured from wolf choruses recorded in the Superior National Forest, Minnesota, USA. Mean duration of 60s did not vary with pack size or composition. Packs replied to simulated howling after an average of 40s, often interrupting the stimulus howls. Choruses began with simply-structured howls, which became increasingly modulated as the chorus progressed. Little difference in mean fundamental frequency or other howl parameters was found among the choruses from packs of various sizes and compositions. In particular, choruses produced by single adult pairs did not differ from those of larger packs accompanied by pups. The lack of relationship between chorus parameters and pack size or composition indicates there is little useful information concerning a pack's size to be found in its chorus howling.
The observation that chorus howling by adult pairs is often perceived as that of larger groups with pups suggests that chorus structure has evolved to exaggerate the apparent size of the pack, especially those newly-established or otherwise reduced in number. If so, wolf howling choruses may represent a mammalian example of the Beau Geste effect, made particularly viable because of the relative immunity of the signal to probing.
Harrington, F.H. (1989). Chorus howling by wolves: acoustic structure, pack size and the Beau Geste effect. Bioacoustics 2(2): 117-136