Cory Hill, 2017. 

Musicology is the comparative study of music in all forms, while Bio-Musicology has been described by Fitch (2015) as the “Biological study of music in all forms”. The biological study of music focuses on the capacities (physiological, and psychological) which are possessed by human beings in order to perceive, and participate in behaviors which are considered musical. This notion of the study of musical capacities that are rooted in human biology are in stark contrast to the comparative studies of music from a cross cultural perspective (as is more conventionally studied). However, general musicology and bio-musicology are comparative, in that while seeking a comprehensive understanding of the nature and origins of musical capacities (from ontogenetic, phylogenetic, functional and mechanical levels of description, which I will later explain), the biological study of musical capacities extends to other species which share similar behavioral traits with humans. In this paper, I will outline and describe the benefits of approaching an understanding of human musicality from a multi-level evolutionary perspective as outlined by Fitch (2015), and how comparative studies of shared behavioral traits can better inform these levels of explanation. I will respond to the objection by Stephen Pinker that music should not be considered to be an adaptation (Pinker, 1997). I will conclude that the comparative approach to studying human musicality offers potential benefits and insights into pedagogical methods in exciting and creative ways which can greatly improve the learning of music students. 

While music itself is often thought of as a cultural phenomenon, the forms of which are highly diverse across cultures, there are still common elements across the varying forms of music. These common elements involve the use of variation in pitch and rhythm, to more abstract elements such as hierarchical organization of phrases and sections. These shared musical elements across cultures are stable and consistent, and are the result of the behavioral capacities of the humans which create the music. Behavioral capacities are supported by physiological and cognitive capacities, which are rooted (according to evolutionary theory) in our biological makeup and evolutionary history (Fitch, 2015). 

Fitch cites scientist Niko Tinbergen, who endorsed a multi-level form of explanation of behavioral traits (Tinbergen, 1963). The different levels of explanation are divided into two categories of kinds of questions that a multi-level explanatory model should seek to fulfill: proximate and ultimate. The proximate description category answers questions regarding why the behavioral trait exists within the individual, and its physiological processes (mechanistic), and how it develops through the lifespan (ontogenetic). The ultimate level of explanation regards the evolutionary history of the behavioral trait (phylogenetic) and how it supports survival (functional). Tinbergen supported the view that evolutionary biology should seek to pursue all levels of explanation in order to gain a more comprehensive look at the nature of biologically based behavioral traits. This would involve the biological aspects of musical behavior which consist in the behavioral and cognitive traits which enable music making. 

Answers on the questions about the origins of musical capacities in humans has been argued to be in vain by some. This is because unlike anatomy, cognitive and behavioral traits are not something that fossilize and for this reason there is no direct evidence with which we could trace such capacities as we could if we inquiring about the evolution of anatomy. However, it has been argued that behavioral traits can be traced to common origins in a similar way to anatomical traits (Honing et al., 2015). Shared traits are said to be either homologous or analogous. Homologously shared traits are traits shared between species because of a common ancestor, and analogously shared traits are said to have evolved between species independently of a common ancestor. 

Homologous traits allow us to trace the characteristics of long lost common ancestors, and it has been argued that if closely related species exhibit a similar behavioral trait, this behavioral trait reflects a common mechanism which is inherited from their common ancestor. This means that we can theoretically trace and date the origins of musical capacities and make further inferences about their emergence based on the environment of that ancestor. Analogous traits help evolutionary theory in a different way by providing multiple data points of behavioral traits for use in testing evolutionary hypotheses. A trait which can be found in many species which share a common ancestor only serve as one point of data with which to reference a hypothesis as to that’s traits origins. In contrast, a trait which has evolved independently many times in many species serves as multiple data points or contexts in which to test an evolutionary hypothesis as to the functional role the trait plays and the environmental factors which lead to the emergence of the trait (Honing et al., 2015). 

The species which is to be comparatively investigated depends on the trait. For example, if we are investigating vocal learning, our non-human primate relatives are not a candidate for comparison, bust more distantly related species such as parrots, cetaceans and songbirds are. When studying the making of percussive sounds, we can investigate near primate relatives such as gorillas and chimpanzees, but also woodpeckers and kangaroo rats. Frogs and insects can be investigated with respect to our shared behavioral traits with them in chorusing and turn taking in order to gain better functional understanding of the adaptive purpose of this behavior (Fitch, 2015). 

Up to this point, I have established that despite the fact that human music varys greatly from culture to culture, the underlying capacities which support the creation of music are far more stable and rooted in our biology. To better understand and explain human musical capacities, it is beneficial to take a comparative approach across species which share similar behavioral traits. Similar behavioral traits between closely related species point to a common underlying mechanism and a common ancestor, while similar behavioral traits in distantly related species give clues as to the adaptive function of the trait and underlying mechanism. I will soon explain why I think that an understanding of the selective pressures which give rise to the behavioral capacities of a species has great relevance to pedagogical methods. Before I do, I will provide a practical example of how this approach can inform our understanding of one mechanism of musicality, and also consider an objection regarding whether or not music can truly be described as an adaptation. 

Song is thought to be one of the few traits of human musicality that can be found in music across cultures. Song involves vocalizations of varying pitch and rhythm which are learned and shared among cultures, and newcomers to that culture. They evolve and change overtime through cultural transmission. The underlying capacity for song behavior is complex vocal learning, which is the mechanism by which animals learn to imitate heard sounds and create novel sounds by assimilating previously learned vocalizations (Mercado et al., 2014). Many species are said to have evolved this vocal learning ability independently and exhibit behaviors which resemble human song. However, our close non-human primate relatives do not possess complex vocal learning abilities. Because it has evolved many times independently, comparative studies of this mechanism across species can better inform us of the neural correlates of the trait (by means of subtraction, similar to that of fMRI imaging), and also to better construct a functional story about the mechanisms adaptive value (Fitch, 2015). 

To illustrate this point more clearly, Colbert-White et al., (2014), published an article regarding the prerequisites for speech, titled “Where apes and songbirds are left behind”. In this article they describe a comparative analysis of humans, parrots, apes and songbirds on multiple levels, including basic sociality, hemispheric asymmetry, complex sociality, vocal learning and heightened respiratory control. “Basic sociality” refers to whether or not the species has regular interaction with conspecifics, are able to recognize them individually and demonstrate parental care. “Hemispheric asymmetry for communication” refers to neural asymmetry of the brain hemispheres of regions related to communication. “Complex sociality” refers to all features of basic sociality but also incorporates hierarchical relationships among members of the individuals group. “Heightened respiratory control” reflects the physiological constraints on speech production which are clearly necessary to vocal imitation and also vocal learning. Between humans, parrots, apes and songbirds, the only two species which share all traits in common are parrots and humans. Apes do not possess adequate physiological mechanisms for vocal production, and songbirds do not exhibit complex sociality. Species which exhibit high degrees of sociality, and interact with conspecifics frequently, must be able to do so by virtue of cognitive mechanisms which allow them to keep track of this information. Because of the adaptive benefits of humans and parrots to share enhanced social relationships (partly due to environmental pressures, and also to longer lifespans), cognitive mechanisms took shape which allowed the emergence of novel means by which we make use of our physiology and cognition to further enhance these relationships through speech imitation. Similar environmental demands lead to similar behaviors and common underlying mechanisms. 

The study of bio-musicology takes an evolutionary perspective on the investigation of musical capacities across species. The assumption is that musical capacities are the result of selective pressures and are adaptive in nature. However, not everyone would agree that music can be considered to serve any adaptive value. In his book “How the Mind Works”, Steven Pinker objects to the idea of music providing any biologically adaptive value, saying “ What benefit could there be to diverting time and energy into the making of plinking noises, or to feeling sad when no one has died… [regarding] biological cause and effect, music is useless” (Pinker, 1997). Pinker goes on to list some of the proposed adaptive benefits of music, such as strengthening social bonds, coordinating action, enhancing ritual, and emotional regulation, yet objects that this does not answer the mystery as to why it is that these things are so. He concludes by saying that unlike basic faculties such as language, vision and social reasoning, if music were to disappear from the face of the earth the species would remain unaffected. If music does not have an adaptive value, then it is very difficult to fit it into any kind of evolutionary explanation, and it is not beneficial to investigate it across species. 

It would appear that what is at the root of Pinker’s objection is that music is not a fundamental trait which has evolved to benefit us in the same way as the evolution of our sensory modalities or less complex behaviors which support social cohesion. For Pinker, music is not fundamental enough to be considered to offer any significant value or require any functional story. Those who would read his objection could conclude that the studies of bio-musicology are not envisioning music in the right context. However, I would remind them that the approach of bio-musicology is not to search for the evolutionary origins of music per se, but rather to take a more bottom up approach by considering the capacities that allow for musical behavior in humans and extending this analysis across species. A comparative analysis of the behavioral and psychological capacities which underlie human music making across species can still be said to provide adaptive explanations because these same capacities aid survival for other contexts. For example, in “Imitating sounds, a cognitive approach to understanding vocal imitation”, Mercado et al., make a hypothesis that the vocal imitation abilities of dolphins have to do with cognitively representing the heard vocalizations of their conspecific and subtracting the difference between the sound as it was heard in the environment and the copied sound which is emitted, the difference reflecting the distance of the conspecific (Mercado et al., 2014). If a dolphin is doing this with more than one conspecifics at a time, this is also reflective of attentional and working memory capacities which are supporting the behavior which ultimately is adaptive for social cohesion. Vocal imitation is part of vocal learning, and vocal learning supports the human capacity for song. Therefore, even if Pinker is correct that music in the cultural sense does not provide any biologically adaptive value, it is still supported by mechanisms which are adaptive. These are the mechanisms which are investigated by bio-musicology, and rightly so. 

In light of all of this, I would like to end by saying a few words about how bio-musicology can benefit music education. Phylogenetic and functional descriptions provided by bio-musicology on relevant musical capacities can greatly inform and complement our understanding of the ontogeny of musical abilities. If we can understand what caused the independent emergence of musical traits across species, we could better construct pedagogical exercises which facilitate the development of these capacities in order to cultivate expert musical skill in the performance of human music. For example, if vocal imitation and vocal learning are said to be the product of basic and complex sociality in conjunction with adequate physiological mechanisms, then we can conclude that there is more to learning to sing than simply listening to music and performing vocal exercises. The enrichment of psychological faculties is every bit as relevant as the physiological development in building an expert musician. Exercises should be given which allow the student to maintain perceptual control over the contents of selective attention, working memory and the processes which involve consolidating those contents into long term memory. These are the processes which originally supported complex and basic sociality and are part of the functional story of the emergence of the mechanisms which support human song. 

Colbert-White, E., Corballis, M. C., & Fragaszy, D. M. (2014). Where apes and songbirds are left behind: A comparative assessment of the requisites for speech. Comparative Cognition & Behavior Reviews, 9, 1-28. 
Fitch WT. 2015 Four principles of bio-musicology. Phil. Trans. R. Soc. B 370: 20140091. 
Honing H, ten Cate C, Pertez I, Trehub SE. 2015 Without it no music: cognition, biology and evolution of musicality. Phil. Trans. R. Soc. B 370: 20140088. 
Mercado,Eduardo, I.,II, Mantell, J. T., & Pfordresher, P. Q. (2014). Imitating sounds: A cognitive approach to understanding vocal imitation. Comparative Cognition & Behavior Reviews, 9, 1-57. 
Pinker, S. (1997). How the mind works W W Norton & Co, New York, NY. 
Tinbergen N. 1963 On aims and methods of ethology. Z. Tierpsychol. 20, 410-433.

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