Graphical AbstractHighlights d Imitation, not social factors, is thought to drive zebra finch song learning d We tested whether non-vocal visual feedback from females affects song learning d Song learning was facilitated by contingent feedback; yoked controls did not learn d Socially guided vocal learning is a crucial mechanism of song development In BriefThe zebra finch is the most common model of human speech development but, unlike humans, is thought to learn only via imitation. Carouso-Peck and Goldstein show that song learning is affected by non-vocal, visual feedback from females. Young males given feedback contingent on their immature songs developed higher-quality songs than yoked controls.

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The capacity to learn novel vocalizations has evolved convergently in a wide range of species. Courtship songs of male birds or whales are often treated as prototypical examples, implying a sexually selected context for the evolution of this ability. However, functions of learned vocalizations in different species are far more diverse than courtship, spanning a range of socio-positive contexts from individual identification, social cohesion, or advertising pair bonds, as well as agonistic contexts such as territorial defence, deceptive alarm calling or luring prey. Here, we survey the diverse usages and proposed functions of learned novel signals, to build a framework for considering the evolution of vocal learning capacities that extends beyond sexual selection. For each function that can be identified for learned signals, we provide examples of species using unlearned signals to accomplish the same goals. We use such comparisons to generate hypotheses concerning when vocal learning is adaptive, given a particular suite of socio-ecological traits. Finally, we identify areas of uncertainty where improved understanding would allow us to better test these hypotheses. Considering the broad range of potential functions of vocal learning will yield a richer appreciation of its evolution than a narrow focus on a few prototypical species. This article is part of the theme issue ‘Vocal learning in animals and humans’.

Human infants are altricial, born relatively helpless and dependent on parental care for an extended period of time. This protracted time to maturity is typically regarded as a necessary epiphenomenon of evolving and developing large brains. We argue that extended altriciality is itself adaptive, as a prolonged necessity for parental care allows extensive social learning to take place. Human adults possess a suite of complex social skills, such as language, empathy, morality, and theory of mind. Rather than requiring hardwired, innate knowledge of social abilities, evolution has outsourced the necessary information to parents. Critical information for species-typical development, such as species recognition, may originate from adults rather than from genes, aided by underlying perceptual biases for attending to social stimuli and capacities for statistical learning of social actions. We draw on extensive comparative findings to illustrate that, across species, altriciality functions as an adaptation for social learning from caregivers. Expected final online publication date for the Annual Review of Developmental Psychology, Volume 2 is December 15, 2020. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

Socially guided vocal learning, the ability to use contingent reactions from social partners to guide immature vocalizations to more mature forms, is thought to be a rare ability known to be used only by humans, marmosets and two unrelated songbird species (brown-headed cowbirds and zebra finches). However, this learning strategy has never been investigated in the vast majority of species that are known to modify their vocalizations over development. We propose a novel, preliminary evolutionary modelling approach that uses ecological, reproductive and developmental traits to predict which species may incorporate social influences as part of their vocal learning system. We demonstrate our model using data from 28 passerines. We found three highly predictive traits: temporal overlap between sensory (memorization) and sensorimotor (practice) phases of song learning, song used for mate attraction, and social gregariousness outside the breeding season. Species with these traits were distributed throughout the clade, suggesting that a trait-based approach may yield new insights into the evolution of learning strategies that cannot be gleaned from phylogenetic relatedness alone. Our model suggests several previously uninvestigated and unexpected species as likely socially guided vocal learners and offers new insight into the evolution and development of vocal learning. This article is part of the theme issue ‘Vocal learning in animals and humans’.