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 ... -  《-》

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 《-》

Bisonic Mechanical Wing Songs and Complex Kinematics in Aerial Displays of the Subtropical Doradito (Pseudocolopteryx acutipennis).

Loud mechanical sounds with a communication role are called sonations. Male Subtropical Doraditos (Pseudocolopteryx acutipennis) exhibit five conspicuously modified primaries suspected of sonating. Here we (1) describe feather modifications, (2) describe three different territorial/aggressive contexts for these sounds: one-perch aerial displays (1PADs), two-PADs, and Chukrut pursuits, (3) investigate the kinematics of the most common display (1PADs) and the physical mechanisms of sonation using synchronized high-speed video and audio, and (4) assess the roles of modified wing feathers in all contexts by experimental manipulation in four individuals. Primaries p3-p7 were modified in adult males but not in females: p3 was pointed with a reduced distal third to the outer vane; p4 and p5 were slim and falciform with pointed tips curved outwards; p6 was broad, massive, and subtly S-shaped, with a spatulate tip; and p7 was large with the distal third of the outer vane abruptly reduced, and the inner vane with a shallow concave sub-apical emargination. One-PADs consisted of perched short nasal introductory syllables accelerating until the bird performed a super-rapid circular flight of ∽15 cm in diameter from and to the same branch, during which two syringeal syllables and three mechanical syllables were given (chik… chik…. chik-chik frrrottt). The syllables were produced during rapid downstrokes by fluttering feathers and were bisonic, being conformed by two simultaneous main tonal, flat, narrow band sounds: a low-pitched note (f0 ∽1 kHz) and a high-pitched note (f0 ∽3.7 kHz). Primary p7 is the necessary and sufficient sound source of the low-pitched note (removal of p7 caused the sound to disappear) and p3 is the sound source of the high-pitched note, being necessary but perhaps not sufficient (removal of p3 caused the sound to disappear); the other modified feathers seem involved in different roles related to either producing the sonation (p4 and p5 interacting with p3) or allowing it (p6 raising dorsally letting p7 flutter freely; removal of p6 did not affect sound production). The specialized shape of p6 might be compromised to allow sonation of p7 without losing flight functionality. Sonations in Subtropical Doraditos occupy the position of the vocal flourish in the songs of other Pseudocolopteryx suggesting the evolutionary replacement of vocal by mechanical sounds. We propose that wing songs in flying birds may be constrained to occur in temporally broken patterns due to intrinsic features of flapped flight and structurally constrained by the demands of creating an airfoil.

Jordan EA ，Areta JI 《-》

Specialized primary feathers produce tonal sounds during flight in rock pigeons (Columba livia).

For centuries, naturalists have suggested that the tonal elements of pigeon wing sounds may be sonations (non-vocal acoustic signals) of alarm. However, spurious tonal sounds may be produced passively as a result of aeroelastic flutter in the flight feathers of almost all birds. Using mechanistic criteria emerging from recent work on sonations, we sought to: (1) identify characteristics of rock pigeon flight feathers that might be adapted for sound production rather than flight, and (2) provide evidence that this morphology is necessary for in vivo sound production and is sufficient to replicate in vivo sounds. Pigeons produce tonal sounds (700±50 Hz) during the latter two-thirds of each downstroke during take-off. These tones are produced when a small region of long, curved barbs on the inner vane of the outermost primary feather (P10) aeroelastically flutters. Tones were silenced in live birds when we experimentally increased the stiffness of this region to prevent flutter. Isolated P10 feathers were sufficient to reproduce in vivo sounds when spun at the peak angular velocity of downstroke (53.9-60.3 rad s(-1)), but did not produce tones at average downstroke velocity (31.8 rad s(-1)), whereas P9 and P1 feathers never produced tones. P10 feathers had significantly lower coefficients of resultant aerodynamic force (CR) when spun at peak angular velocity than at average angular velocity, revealing that production of tonal sounds incurs an aerodynamic cost. P9 and P1 feathers did not show this difference in CR These mechanistic results suggest that the tonal sounds produced by P10 feathers are not incidental and may function in communication.

Niese RL ，Tobalske BW 《-》

Sounds of Modified Flight Feathers Reliably Signal Danger in a Pigeon.

In his book on sexual selection, Darwin [1] devoted equal space to non-vocal and vocal communication in birds. Since then, vocal communication has become a model for studies of neurobiology, learning, communication, evolution, and conservation [2, 3]. In contrast, non-vocal "instrumental music," as Darwin called it, has only recently become subject to sustained inquiry [4, 5]. In particular, outstanding work reveals how feathers, often highly modified, produce distinctive sounds [6-9], and suggests that these sounds have evolved at least 70 times, in many orders [10]. It remains to be shown, however, that such sounds are signals used in communication. Here we show that crested pigeons (Ochyphaps lophotes) signal alarm with specially modified wing feathers. We used video and feather-removal experiments to demonstrate that the highly modified 8th primary wing feather (P8) produces a distinct note during each downstroke. The sound changes with wingbeat frequency, so that birds fleeing danger produce wing sounds with a higher tempo. Critically, a playback experiment revealed that only if P8 is present does the sound of escape flight signal danger. Our results therefore indicate, nearly 150 years after Darwin's book, that modified feathers can be used for non-vocal communication, and they reveal an intrinsically reliable alarm signal.

Murray TG ，Zeil J ，Magrath RD 《-》