Summary (Oscines): (2) Birds must practice (sub-, plastic, and ………………………………………… (3) crystallized song) Trimmed, cut, and frozen If not, song performance is greatly distorted (1)Bird must have a tutor during the sensitive period Dependent care Independent Juveniles Maturing Adult

Crude template Template matched to song heard Exact template Own species song heard MEMORIZATION PHASE Song matched to template Hears own song Song output Testosterone MOTOR PHASE Day length increases Gonad size increases AUDITORY TEMPLATE MODEL I III II Requires Tutor Requires hearing yourself This is a (brief) window of crystallization Forgetting or culling

Brown-headed cowbirds Brood parasites HOW DO THEY LEARN THEIR SONGS?

Action-based Learning Eastern and Southern sub-spp when transplanted chicks raised by the ‘other’ sub-spp they learn that sub-spp songs males respond to female wing-flicks by repeating songs more when females respond this way

Indirect pathway between HVC and RA – recursive loop HVA=Higher vocal center RA=Robust nucleus of the Arcopallium Anterior Vocal pathway (learning) Posterior Vocal pathway (production)

Many of the nuclei have neurons that recognize specific sounds in complex song

Summary: Some neurons are specific to individual syllables (higher firing rate) Gene induction higher for species-specific songs, and in some case novelty “ “ for so-called “sexy” songs (canaries) The feedback loop, singing behavior itself (feedback?) increase neuron production

Vocal fighting and vocal flirting Males sing differently to males and females

Two studies how that females prospect for males based on song Pied flycatcher Great reed warbler # females caught 10 5

So what are females looking for in song or singing that indicates a good male?

Specific song structures: - Sexy song in canaries (sensory bias or signals precise vocal coordination?) - Potent songs in cowbirds: males punish rivals singing potent songs, thus it indicates the # fights won and lost (like isolated males) - Swamp sparrows and limits to vocal performance

Song frequency (over time): - Song frequency is correlated to males nest defense and feeding the young (parental care) WHY? What does it indicate?

Songs per 5 min Number of birds paired first - In pied flycatchers suppl. feeding increases singing 2-fold) and results in faster pairing between males and females

Song repertoires: - Females show preferences for males with larger repertoires (song or syllable) Repertoire size Mean # displays

Song repertoires: Years on territory - Older birds (open-ended learners) sing more songs - Or song repertoire correlates with survivorship (close-ended learners) Song Sparrow

Repertoire size Mean # displays Lifetime rep success - Males with larger repertoires have higher LRS - And females prefer them - But it would nice to this in offspring…

Pairing date Mean # displays Repertoire size Genetic offspring survival Repertoire size

Male canaries affected with malaria lower repertoire size smaller HVC But also in response to nutritional stress in: song sparrows, canaries, and starlings

Song familiarity (local song structure): -Female song sparrows prefer the local dialect over ‘foreign’ dialect …this sets up a performance index…

Many notes; high accuracy Few notes; poor accuracy Intermediate

Vocal fighting and vocal flirting Males sing differently to males and females

Yasukawa’s experiment (1981) - Mute males via removing a portion of the hypoglossal nerve Krebs’ Experiment (1977) - Remove males - Monitor settlement

songs per min Neighbor stranger

- Muted males creates more territory intrusions - Experimental playbacks delays territorial settlement in the absence of males - There is a level of sophistication  males recognize neighbors from strangers and have different responses relative to their position in space with respect to territory intrusion Songs to advertise territory ownership and aggression to other males – 3 examples

Track 38 (Kroodsma) #1 Song (Type) Matching Counter singing Marsh Wrens

Repertoire Matching Beecher’s Studies on the song sparrow A N B E H J L K O P U V W X T S R Q G CI D F Bird 1 Bird 3 Bird 4 Bird 2

(1) Match song exactly Type Matching (2) Match with another shared song Repertoire Matching (3) Sing a unshared song

Type Match Repertoire Match Unshared - Neighbors tend to repertoire match more than expected by chance - Neighbors type-match early and repertoire match later in the season Song functions to communicate to territory ownership in very sophisticated ways