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A formal investigation of Cultural Selection Theory: acoustic adaptation in bird song

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Abstract

The greatest challenge for Cultural Selection Theory lies is the paucity of evidence for structural mechanisms in cultural systems that are sufficient for adaptation by natural selection. In part, clarification is required with respect to the interaction between cultural systems and their purported selective environments. Edmonds et al. have argued that Cultural Selection Theory requires simple, conclusive, unambiguous case studies in order to meet this challenge. To that end, this paper examines the songs of the Rufous-collared Sparrow, which seem to exhibit cultural adaptations minimizing signal degradation relative to local environments. Specifically, the more forested the habitat, the more the tail end of the song resembles a whistle rather than a trill; yet, variation in song is uncorrelated with genetic variation. This paper explores the mechanisms responsible for these putative acoustic adaptations through a series of computer simulations. The main point of this research is not to test this model, but to demonstrate that models of this type have the resources to meet the in-principle objections that have been raised against Cultural Selection Theory. This research lends much-needed empirical support to Cultural Selection Theory by clarifying the nature of the interaction between culture and environment. It also contributes to evolutionary theory by clarifying the scope and limits of adaptation by natural selection.

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Notes

  1. For a compelling defense of memetics, see Gers (2008).

  2. A version of this simulation (Rand and Wilensky 2007) can be run online at: ccl.northwestern.edu/netlogo/models/ElFarol.

  3. See Anderson (2009) for an enlightening discussion of the relationship between strategic thinking and Cultural Selection Theory.

  4. It would be more precise to use the terms open habitat and closed habitat instead of field and forest, respectively. This is because the key factor is how much vegetation exists in the region. So, for example, a marsh might have the same acoustic properties as a field, as would the top of a body of water: all of these are open habitats in the acoustically relevant sense. Furthermore, the distinction between open and closed habitats is more precisely conceived as lying on a continuum. So, for example, the leafy treetops of a mature forest would be a very acoustically closed environment, while the floor of that mature forest might be slightly more open, a sapling forest might be more open still, and a clearing still more open than this. Moreover, the observed correlation in bird song type matches this gradient in habitat type very closely such that the more open the habitat, the more the song resembles a trill rather than a whistle. Thus, when I use the terms field and forest, it is in a slightly stylized sense that is intended to make this paper more readable.

  5. Degradation refers to distortions of the signal’s frequency and amplitude modulations caused by environmental obstacles. It is one of two kinds of signal loss that occurs as a sound is transmitted. The other is attenuation, which is the opposite of amplification.

  6. Interestingly, Brown and Handford learned [first by computer simulation (1996) and then by field experiment (2004)], that a further refinement of the concept of acoustic degradation is required—a distinction between average and consistent transmission fidelity. Whistles incur less average degradation than trills in both forests and fields; however, trills transmit more consistently than whistles in fields. This leads them to conclude that consistency and average transmission fidelity might produce competing selection pressures that together produce the observed correlation between song and habitat type. To avoid charges of Panglossianism, however, it would be necessary to test this new hypothesis further; for example, by verifying the relative effect of average versus consistent transmission fidelity on the song imitation of sparrows.

  7. This description presumes that trills and whistles are distinct, rather than appearing on a continuum. This simplifying assumption will be discussed later in the paper.

  8. By Gould and Lewontin’s description of Panglossianism, it would appear that this fallacy is frequently committed in evolutionary biology. Panglossianism is most accurately portrayed as occurring on a spectrum, where most of the transgressions are mild in degree. Adaptationist reasoning is an essential methodological device in evolutionary biology. Gould and Lewontin’s Panglossianism paper is most compellingly construed as a warning against a certain potential for inappropriate adaptationist reasoning which can occur when researchers relax their vigilance: an ideal limit of a lurking tendency all evolutionists need to avoid.

  9. Alexander and Skyrms refer to this imitation-based dynamic as imitate-the-best.

  10. In standard Evolutionary Game Theory, the probability a player has of playing against a particular strategy is given by the proportion of the population playing that strategy. This proportion changes over successive rounds of the game by the replicator dynamics: the proportion of the population playing a given strategy in one generation is multiplied by a fitness factor, which is the ratio of the average payoff of that strategy to the average payoff of the population.

  11. According to Nash’s (1950) definition, distributive justice holds when two criteria are met: all the players have equal outcomes and there are no alternatives under which all players are better off. For instance, in the Prisoner’s Dilemma, distributive justice holds only when both players cooperate.

  12. My simulations use the program NetLogo (Wilensky 1999), which can be downloaded for free at: http://ccl.northwestern.edu/netlogo.

  13. Technically, the key interaction between players occurs only when new neighbours are introduced into a neighbourhood, but analytically there is no difference between this and a fixed-player game.

  14. Currently territories on the far edge of the game-space share borders with those on the opposite edge, similar to the design of old arcade games: for example, if you took Pac-Man out the door on the right edge of the board, he would reappear through the door on the left edge.

  15. Of course, some human communications show characteristics of acoustic adaptation similar to those of the Rufous-collared Sparrow; for instance, yodels and other vocal signals intended for transmission of information over long distances. It is challenging to make a case that these human calls are the result of cultural selection for several reasons. Humans can update their communication strategies throughout the course of their lives which makes it difficult to identify population frequencies of their cultural units. Further, any cultural evolutionary analysis would have to accommodate the clear potential that human strategic reasoning contributed significantly to the development of such acoustically effective calls. For these reasons, the identification and measurement of selection pressures is much more challenging in human versus bird cultural systems.

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Acknowledgments

Special thanks go to the following people for their help in developing the ideas in this paper: Wayne Myrvold, Bill Wimsatt, Bill Harper, Bob Batterman, Scott McDougall-Shackleford, Matt Gers, Paul Handford, Chris Hajzler, Andrew Fenton, Dale Jamieson, Laura Franklin Hall, Bill Ruddick, Greg Bognar, Chris Schlottmann, Sandra Larsen, Kim Sterelny, and two anonymous reviewers. I am also grateful for feedback received from participants in the conference “Biological Explanations of Behavior: Philosophical Perspectives” which was organized in Hannover, Germany, by Katherine Plaisance and Thomas Reydon, with special thanks to Colin Allen, David Sloan Wilson, Omri Tal, and Armin Shulz. Of course, the author maintains full responsibility for any errors or omissions that may remain in this paper.

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  1. Department of Philosophy, Loyola University Chicago, 1032 West Sheridan Road, Chicago, IL, 60660, USA

    G. K. D. Crozier

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Crozier, G.K.D. A formal investigation of Cultural Selection Theory: acoustic adaptation in bird song. Biol Philos 25, 781–801 (2010). https://doi.org/10.1007/s10539-010-9194-6

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