that learn vocalizations? A comparative approach also allows us to test
hypotheses of the ecological and behavioral factors that favor vocal learning. Are certain social structures conducive to vocal learning? Are individual signatures and group signatures influenced equally by vocal learning?
Is there support for the hypothesis that echolocation predisposes species to
learn social calls? Although we are close to the point where we can ask
these questions, we need to broaden even further the taxonomic groups we
choose to study. When designing new studies, careful consideration of the
phylogenetic relationship of a taxon to those for which we already have
data on vocal learning is warranted.
5.3.2. Cultural Evolution and Vocal Learning
The study of learned vocalizations is an especially pertinent type of social
learning that can inform both theoretical and empirical work on gene–
culture evolution. Nonvocal traits that are transmitted culturally can affect
the course of evolution for other traits (Feldman and Laland 1996). In the
nonhuman literature, examples have focused on cultural transmission of
foraging strategies, probably because foraging was thought to be under
strong selection to be optimal. This focus has yielded important advances
in theoretical work and some very interesting empirical results. Yet, the
ability to modify the evolution of other traits might be especially true for
behavioral traits that affect social organization or mating. Social interaction
is nearly always involved in cultural transmission, and traits are often transmitted between the members of social groups. Consequently, cultural traits
that themselves affect social organization can powerfully influence both
their own transmission and other aspects of a species’ biology. Culturally
transmitted traits that affect the likelihood of mating almost certainly alter
the fate of traits transmitted genetically. Many of the vocalizations reviewed
here are integral to the smooth functioning of social groups. In at least one
case, learned vocalizations actually facilitate another form of social learning—social foraging (greater spear-nosed bats—Boughman 1998;Wilkinson
and Boughman 1998).
Certainly, there is a rich history of this kind of work in the birdsong literature, including applications of the “meme” concept (Payne et al. 1988;
Payne and Westneat 1988; Trainer 1989; Ficken and Popp 1995). Much of
this work has focused on the fate of song variants, and fewer studies have
explored genetic consequences or tested gene–culture models (Gibbs 1990;
Grant and Grant 1996). We urge increased focus on the processes and consequences of culturally transmitted vocalizations outside of oscine song.
Both a fine-scale focus on individual species and large-scale consideration
of mechanisms and patterns across species are likely to yield plentiful
insights into how genes and culture coevolve. This work will require simultaneous study of vocal and genetic variation, both time-consuming occupations but likely to be well worth the effort.
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