Bulla and colleagues quantified male song complexity (number of element types in 50-element sequences) for 4,940 passerine species from 18,065 citizen-science recordings (Xeno-canto, Macaulay Library), validating their metric against published repertoire metrics (r = 0.53β0.77) and song-control brain region HVC (r = 0.58) .
The headline descriptive result is a latitudinal gradient: the most complex songs occur in temperate open-habitat assemblages with relatively drab males (Palearctic, Saharo-Arabian, parts of Nearctic), while tropical species sing simpler songs . Phylogenetic signal is strong (Pagel's Ξ» = 0.74 all passerines, 0.71 Oscines, 0.68 Suboscines), and song-learning Oscines exceed innate-song Suboscines by 0.87 SD (95% CI 0.42β1.32) .
Three traditional hypotheses fared poorly. Six sexual-selection proxies (size dimorphism, plumage dimorphism, polygyny, territoriality, social-bond stability, extra-pair paternity) showed no strong effects . The species-recognition hypothesis also lacked support. Notably, song complexity was negatively correlated with brain size (N = 679 species) β a pattern the authors attribute to a confound: migrants have smaller brains yet more complex songs, and adjusting for migration eliminates the brain-size association . This directly contradicts the widely accepted intelligence-signaling view and aligns with a companion suboscine analysis finding that sexual selection does not increase complexity there either .
What survives: in Oscines, complexity is higher in open habitats (consistent across latitudes), migratory species, and habitat generalists β and these ecological strategies largely account for the latitudinal pattern. The songβplumage trade-off is context-dependent, appearing only in temperate forests .
Strengths: unprecedented taxonomic coverage, multiple sexual-selection proxies tested simultaneously, sensitivity analyses (PGLS, clade-specific models, heteroscedasticity), open R code and data, and a sharp falsification of a popular cognitive-signaling narrative with an explicit confound analysis.
Limitations: (1) ~17% of passerines are missing, likely non-randomly (rarer, tropical species β potential sampling bias); (2) only male songs were analyzed, ignoring female song and non-song vocalizations; (3) the complexity metric (element types per 50 elements) ignores repertoire size, sequencing rules, and performance quality β different metrics could yield different gradients; (4) recordings vary in quality and identification reliability across citizen-science archives; (5) the design is observational, so 'acoustic adaptation' and 'migration' remain correlational accounts, and the authors themselves note assemblage patterns may reflect lineage distribution rather than adaptation. Confidence in the descriptive gradient is high; confidence in the mechanistic interpretation (ecology over sexual selection) is moderate until experimental or within-species tests corroborate it. Interestingly, single-population work in blue tits shows some song dimensions do predict fitness (clutch size), suggesting sexual selection may act on qualities this global metric cannot capture β a key counterpoint the cross-species design cannot exclude.
What would change the conclusion: a positive song complexityβbrain size relationship after rigorous migration correction across more species, or within-species evidence that females prefer complex songs independent of habitat and migration ecology.
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