ADVANCED

1. ADVANCED LEC 10 University of Rio Grande Donald P. Althoff, Ph.D. ORNITHOLOGY Sound Production & Song Reference Chapter 8

2. Sound Production There are four basic ways birds can produce sound • ___________– through the syrinx • _____ clipping (think Royal Albatross) and Pecking(think woodpeckers) using the bill/beak • __________(think woodcock with wing feathers and manakin with wing and tail feathers) http://vimeo.com/40657519 • _______________ (think prairie chickens and sage grouse)

3. Feathers Sound Production • Manakins http://vimeo.com/40657519 red-capped manakin http://vimeo.com/40652880 club-winged manakin • Woodcock—3 outer primaries narrower…produce “whistling” sound in flight • Prairie Chickens & Sharp-tailed Grouse: as part of breeding displays “wag” tail that produces some sound (video clip)

4. Air Sac Sound Production…with a little help from the syrinx • Prairie chickens–anterior end of _________modified as a vocal sac/resonatingchamber • Air is forced into the sac directly from the trachea via the pharynx • As sac inflates, sound waves from syrinx strike the tense walls acting as resonators

5. Vocalizations • Involve the syrinx • Two broad distinctions: A) _____–associated with _______territorial males B) _____–not associated with breeding i) distress/alarm calls ii) flight calls iii) begging calls iv) feeding calls v) nests calls vi) flock calls • Technically, there is ___real dichotomy between songs and calls in their _______________________________ _____________________________________________

6. ___________– visual display of the frequency of a sound distributed in relation to time Example of two “songs” (top and bottom) of buntings. _________are the principal basic units. ______are 1 or more syllables. Gill Fig 8-2 p217

7. Range of Vocalizations • From short clicks of swifts…. • …to quavering whistles of the tropical, partridge-like tinamoushttp://www.youtube.com/watch?v=llBEEeBe-P0 • …to the long melodies of wrens http://www.youtube.com/watch?v=Kbl-mWkSId8 • …to the annoying mockingbird http://www.youtube.com/watch?v=NNNX3f3_svo • Range from deep infrasounds to high pitches inaudible • Range from almost no vocalizations (mute swans, turkey vultures) to almost unlimited “vocabularies” (mynas, parrots, mockingbirds)

8. Modulations within bird songs/calls are ______among animal sounds • Modulation—in the form of frequency (pitch) and amplitude (energy) • Example—little modulation: white-throated sparrow http://www.youtube.com/watch?v=dtZkV_hiiKY • Example—considerable modulation: song sparrowhttp://www.youtube.com/watch?v=wdaE7eaayKM

9. Fundamental Dichotomy • _____________vs. _____________songs/calls • Whistled songs consist of nearly pure (lacking harmonics) sinusoidal waveforms. Sinusoidal waveforms exist when with higher pitch, there are more frequent oscillations of sound waves. • Harmonic songs have overtones with frequencies that are multiples of fundamental frequency—the number of harmonics and their relative amplitude determines general tonal quality of the notes. One dominant harmonic has more energy or greater amplitude than do the others in the spectrum.

10. Whistled vs. Harmonic Songs/Calls • Whistled Blackpoll warbler http://www.allaboutbirds.org/guide/blackpoll_warbler/sounds Spruce Grouse http://www.allaboutbirds.org/guide/spruce_grouse/sounds • Haromonic Black-capped chickadee http://www.allaboutbirds.org/guide/black-capped_chickadee/sounds

11. Detailed Analysis of the Black-capped Chickadee’s dee-dee-dee-dee-dee scold call • Sonogram of one “dee” phrase • The number of “dees” _________in relation to the threat of predation • Relative amplitude of the tones of the harmonic series Gill Fig 8-4, p219

12. More on the Black-capped Chickadee’s “information-loaded” call sharing abilities Number of “dee” notes per call Gill Box 8-2, p223

13. _______________ of sound affects the ease with with the listener—predator or neighbor—can locate its source • Calls used to locate and/or attract one another consist of ____________ with broad frequency ranges…thereby enriching the information about __________________ • In contrast, alarm calls are _________ (i.e., narrow frequency range), __________________ ____________that conceal the sender’s whereabouts.

14. Eurasian BlackbirdLocate/attract vs. Alarm Call Call when mobbing an ______ Call to alert that a _______is flying over Sonograms(Gill Fig 8-5, p220)

15. Physical structure alsodetermines the distance sound will travel…and how much distortion it will sustain before reaching its listener • Interference, absorption, and scattering of sound waves affected by a) __________ b) __________ c) __________ • ____________ sounds (like calls of grouse, cuckoos, doves, and large owls) are most effective for ______ __________ communication

16. Physical structure…con’t • Forest-dwelling birds tend to produce simple sounds • vs. more complex, frequency modulated songs/calls are produced by birds in open habitat • Examples: rose-breasted grosbeak (forest) http://www.allaboutbirds.org/guide/rose-breasted_grosbeak/sounds vs. grasshopper sparrow (prairie) http://www.allaboutbirds.org/guide/grasshopper_sparrow/sounds

17. Other considerations… • Birds adjust singing behavior to their surroundings…aka singing environment. a) hummingbirds sing ______in the presence of increased background noise (like a creek) b) great tits now sing at ________ frequencies brought on by auto traffic • Cassowaries—large solitary flightless birds that live in dense rainforests—use low-frequency sounds (as do elephants and some whales) to communicate over long-distance (remember low frequency sound carries further in a “dense” environment). Achieved by ‘doubling width of neck’ (aka inflated)

18. Sound Production by the Syrinx • Syrinx operates with “nearly 100%” efficiency to create loud, complex sounds. • It is a _________…that is unique to birds (note: that makes three unique features to birds….feathers, pecten, and now the syrinx) • It produces sound ________vocal cords • It is the made of _______ tissues in some species, ________tissues in some species, or both tracheal and bronchial tissues. • Sound is produced by the ____________ of the air column as air passes through the syringeal passageway.

19. Juncture of the trachea and the two bronchi Gill Fig 8-6, p224 Simple (left) and complex (right) syringeal musculature Gill Fig 8-7, p225

20. 1) last free cartilaginous ring 2) tympanum 3) first group of syringeal rings 4) pessulus (cartilage) 5) membranatympaniformislateralis 6) membranatympaniformismedialis 7) second group of syringeal rings 8) main bronchus 9) bronchial cartilage

21. Sound Production by the Syrinx…con’t • The airways become constricted by the internal and external __________(see enlarged handout, bottom schematic) • There is a glass-clear membrane—the internal typmanium membrane, which with the surrounding interclavicular air sac gets pushed into the bronchial air space. • __________of the labia (internal and external)—not the tympaniform membranes—determine the basic characteristics of the sounds produced by the syrinx. • _________________ control syrinx action during vocalization. If the bird species lacks these muscles it can only grunt or hiss.

22. Sound Production by the Syrinx…con’t • Most nonpasserine birds have just ___pairs of “extrinsic muscles” either side of the trachea above the syrinx • Most passerine birds have as many as ___pairs of “extrinsic muscles”… • Thus, “__________” have more complex musculature but not necessarily more complex vocalizations vs. those with just 2 pairs. 1 2 3 4 5 5 6 X

23. Sound Production miscellaneous… • Some species, like swans, cranes, some curassows, and guineafowl that produce loud, rich resonating trumpet-like calls have usually long trachea that are ________in the body cavity or in the bony sternum itself.