In most vocalizing vertebrates, lighter animals tend to produce acoustic signals of higher frequency than heavier animals. Two hypotheses propose to explain this negative relationship in vespertilionid bats: (i) mass-signal frequency allometry and (ii) emitter-limited (maximum gape) signal directionality. The first hypothesis, that lighter bats with smaller larynges are constrained to calls with higher frequencies, is supported at the species level. The second hypothesis proposes that in open space, small bats use higher frequencies to achieve narrow sonar beams, as beam directionality increases with both emitter size (maximum gape) and signal frequency. This hypothesis is supported within a comparative context but remains untested beyond a few species. We analyzed gape, body mass, and echolocation data under a phylogenetic comparative framework to test these hypotheses, and considered forearm length as both a proxy for wing design and an alternative measure of bat size. Controlling for mass, we found no support for the directionality hypothesis. Body mass and relative forearm length were negatively related to open space echolocation call peak frequency, reflecting species-specific size differences, but also the influence of wing design and preferred foraging habitat on size-independent species-specific differences in echolocation call design.
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