Feb 26: Song Dialects: Birds Why Do Birds Sing? Why this is an ill-posed question • need to specify the correct level of analysis (for now, we are referring to the functional question of survival value) • even considering it as a functional question, it is TOO GENERAL Three more specific questions about bird song • Why do different species sing different songs? • Why do only males sing (usually)? • Are dialects (interpopulation variation within a species) adaptive? Feb 26: Bird Song Dialects: Digression A Functional Perspective on Communication What communication is--a broad definition: Production of signals by one individual (sender) that change the behavior of other animals (receivers) This broad definition is TOO broad • Includes signals that change the behavior of receivers to the benefit of the receiver but to the detriment of the sender (e.g., the rustling of a mouse that attracts a predator) • Natural selection would not favor traits in sender that work against the interests of the sender (there is no "function" of the mouse's rustling) A narrower definition: Production of signals by one individual that change the behavior of a receiver in a way that benefits the sender • This type of communication need not only occur among members of the same species • Also, the receiver might also benefit by having changed its behavior in response to the signal • Thus, a rattlesnake's rattle signals to potential predators that the snake is dangerous (this is good for the snake as well as for the would-be predator) Feb 26: Bird song dialects: Why do different species sing different songs? Why Do Different Species Sing Different Songs? H1: Songs diverge by chance (genetic drift) after speciation (perhaps associated with selection for consistency within species) Most likely hypothesis for most species H2: perhaps differences reflect adaptation to different local conditions (e.g., different acoustical environments Some evidence of this in Great Tits (in Europe)--woodland populations have lower frequency songs than grassland populations H3: Selection for divergence of traits involved in species recognition--to prevent hybridization No evidence for more pronounced song differences in indigo and lazuli buntings where their ranges overlap (in Nebraska) Feb 26: Bird song dialects--Why only males? Why Do Only Males Sing? H1: Tendency of males to sing (and females to not sing) is side-effect of sexual differentiation of reproductive behavior H2: Song involves costs (time, energy, exposure to predators); Perhaps only the males reap benefits of singing in most species Feb 26: Bird song dialects--Why only males? Why Do Only Males Sing? (cont’d) Hypothesis 2a: Males benefit by attracting females • Choosy females use song as indicator of male quality (we postpone for now the question of why females tend to be the choosy sex--see Chap. 12) Evidence: • In many species, males sing only when trying to attract mates • In many species, females clearly choose among males based on song--e.g., song complexity • In starlings, males with larger song repertoires are chosen earlier, and have young earlier Alcock Fig. 8.3 Feb 26: Bird song dialects--Why only males? Why Do Only Males Sing? (cont’d) Hypothesis 2b: Males benefit by more effectively repulsing rival males • Female is a resource for which males compete • Any trait that enhances competitiveness would be favored • Perhaps songs serve to warn other males of presence and quality of singer Evidence: • In some species, males sing most when mates are fertile, hence when male has most to lose by intrusion of rival male • In white-throated sparrows, removal of singing male results in intrusion by rival, but tape recording delays intrusion • In starlings, recordings of complex songs are more effective than recordings of simple songs; thus, in starlings, song complexity has survival value for both mate attraction and rival repulsion Alcock Fig. 8.4 Feb 26: Bird song dialects--Are dialects adaptive? Are Dialects Adaptive? H1: NO--they result from non-adaptive genetic or cultural drift? In indigo buntings, dialects do drift around over time--doesn’t mean they are non-adaptive H2: YES--perhaps they represent adaptation to local physical environment Scant evidence of this H3: YES--perhaps they represent adaptation to local social environment Feb 26: Bird song dialects--Are dialects adaptive? Are Dialects Adaptive? - cont’d H3a: Dialects are adaptive through process of female choice: perhaps females prefer to mate with male who sings like Dad (hence are adapted to same environment in which Dad was successful) Evidence: • In brown-headed cowbirds, females do prefer to mate with males singing their own dialect • However, ability of males to change dialect over lifetime (e.g. white-crowned sparrow) means that dialect doesn’t predict place of birth H3b: Dialects are adaptive through process of rival repulsion: perhaps males are more effective in repelling rivals if they can “talk the talk”--indicates familiarity with habitat, staying power • Song-matching in song sparrows Alcock Fig. 8.7 Feb 26: Song Dialects: Birds Why Do Birds Sing? Three more specific questions about bird song • Why do different species sing different songs? Apparently a combination of selection for species recognition and drift • Why do only males sing? Intersexual selection (mate attraction) & Intrasexual selection (rival repulsion) • Are dialects (interpopulation variation within a species) adaptive? Differences may be useful in indicating familiarity with local neighborhood
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