Aeroelastic flutter of feathers, flight and the evolution of non-vocal communication in birds.

Tonal, non-vocal sounds are widespread in both ordinary bird flight and communication displays. We hypothesized these sounds are attributable to an aerodynamic mechanism intrinsic to flight feathers: aeroelastic flutter. Individual wing and tail feathers from 35 taxa (from 13 families) that produce tonal flight sounds were tested in a wind tunnel. In the wind tunnel, all of these feathers could flutter and generate tonal sound, suggesting that the capacity to flutter is intrinsic to flight feathers. This result implies that the aerodynamic mechanism of aeroelastic flutter is potentially widespread in flight of birds. However, the sounds these feathers produced in the wind tunnel replicated the actual flight sounds of only 15 of the 35 taxa. Of the 20 negative results, we hypothesize that 10 are false negatives, as the acoustic form of the flight sound suggests flutter is a likely acoustic mechanism. For the 10 other taxa, we propose our negative wind tunnel results are correct, and these species do not make sounds via flutter. These sounds appear to constitute one or more mechanism(s) we call 'wing whirring', the physical acoustics of which remain unknown. Our results document that the production of non-vocal communication sounds by aeroelastic flutter of flight feathers is widespread in birds. Across all birds, most evolutionary origins of wing- and tail-generated communication sounds are attributable to three mechanisms: flutter, percussion and wing whirring. Other mechanisms of sound production, such as turbulence-induced whooshes, have evolved into communication sounds only rarely, despite their intrinsic ubiquity in ordinary flight.

Clark CJ ，Prum RO 《-》

Aeroelastic flutter produces hummingbird feather songs.

Clark CJ ，Elias DO ，Prum RO 《-》

Hummingbird feather sounds are produced by aeroelastic flutter, not vortex-induced vibration.

Clark CJ ，Elias DO ，Prum RO 《-》

Smithornis broadbills produce loud wing song by aeroelastic flutter of medial primary wing feathers.

Broadbills in the genus Smithornis produce a loud brreeeeet during a distinctive flight display. It has been posited that this klaxon-like sound is generated non-vocally with the outer wing feathers (P9, P10), but no scientific studies have previously addressed this hypothesis. Although most birds that make non-vocal communication sounds have feathers with a shape distinctively modified for sound production, Smithornis broadbills do not. We investigated whether this song is produced vocally or with the wings in rufous-sided broadbill (S. rufolateralis) and African broad bill (S. capensis). In support of the wing song hypothesis, synchronized high-speed video and sound recordings of displays demonstrated that sound pulses were produced during the downstroke, subtle gaps sometimes appeared between the outer primary feathers P6-P10, and wing tip speed reached 16 m s(-1) Tests of a spread wing in a wind tunnel demonstrated that at a specific orientation, P6 and P7 flutter and produce sound. Wind tunnel tests on individual feathers P5-P10 from a male of each species revealed that while all of these feathers can produce sound via aeroelastic flutter, P6 and P7 produce the loudest sounds, which are similar in frequency to the wing song, at airspeeds achievable by the wing tip during display flight. Consistent with the wind tunnel experiments, field manipulations of P6, P7 and P8 changed the timbre of the wing song, and reduced its tonality, demonstrating that P6 and P7 are together the sound source, and not P9 or P10. The resultant wing song appears to have functionally replaced vocal song.

Clark CJ ，Kirschel AN ，Hadjioannou L ，Prum RO ... -  《-》

Ways that Animal Wings Produce Sound.

There are at least eight ways that wings potentially produce sound. Five mechanisms are aerodynamic sounds, created by airflow, and three are structural sound created by interactions of solid surfaces. Animal flight is low Mach (M), meaning all animals move at <30% of the speed of sound. Thus in aerodynamic mechanisms the effects of air compressibility can be ignored, except in mechanism #1. Mechanism #1 is trapped air, in which air approaches or exceeds Mach 1 as it escapes a constriction. This mechanism is hypothetical but likely. #2 is Gutin sound, the aerodynamic reaction to lift and drag. This mechanism is ubiquitous in flight, and generates low frequency sound such as the humming of hummingbirds or insect wing tones. #3 is turbulence-generated atonal whooshing sounds, which are also widespread in animal flight. #4 are whistles, tonal sounds generated by geometry-induced flow feedback. This mechanism is hypothetical. #5 is aeroelastic flutter, sound generated by elasticity-induced feedback that is usually but not always tonal. This is widespread in birds (feathers are predisposed to flutter) but apparently not bats or insects. Mechanism #6 is rubbing sound (including stridulation), created when bird feathers or insect wings slide past each other. Atonal rubbing sounds are widespread in bird flight and insects; tonal stridulation is widespread in insects. #7 is percussion, created when two stiff elements collide and vibrate, and is present in some birds and insects. Mechanism #8 are tymbals and other bistable conformations. These are stiff elements that snap back and forth between two conformations, producing impulsive, atonal sound. Tymbals are widespread in insects but not birds or bats; insect cuticle appears predisposed to form tymbals. There are few examples of bat wing sounds: are bats intrinsically quiet, or just under-studied? These mechanisms, especially Gutin sound, whooshes, and rubbing (#2, #3, and #6) are prominent cues in ordinary flight of all flying animals, and are the "acoustic substrate" available to be converted from an adventitious sound (cue) into a communication signal. For instance, wing sounds have many times evolved into signals that are incorporated into courtship displays. Conversely, these are the sounds selected to be suppressed if quiet flight is selected for. The physical mechanisms that underlie animal sounds provide context for understanding the ways in which signals and cues may evolve.

Clark CJ 《-》