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COMPARING HUMANS and BIRDS by Daniel C. Mann a Dissertation STABILIZING FORCES IN ACOUSTIC CULTURAL EVOLUTION: COMPARING HUMANS AND BIRDS by Daniel C. Mann A dissertation submitted to the Graduate Faculty in Linguistics in partial fulfillment of the requirement for the degree of Doctor of Philosophy, The City University of New York 2019 2019 DANIEL C. MANN All rights reserved ii This manuscript has been read and accepted for the Graduate Faculty in Linguistics in satisfaction of the dissertation requirement for the degree of Doctor of Philosophy. JULIETTE BLEVINS Date Chair of the Examining Committee GITA MARTOHARDJONO Date Executive Officer MARISA HOESCHELE DAVID C. LAHTI MICHAEL I. MANDEL Supervisory Committee THE CITY UNIVERSITY OF NEW YORK iii Abstract STABILIZING FORCES IN ACOUSTIC CULTURAL EVOLUTION: COMPARING HUMANS AND BIRDS By Daniel C. Mann Advisor: Professor Juliette Blevins Learned acoustic communication systems, like birdsong and spoken human language, can be described from two seemingly contradictory perspectives. On one hand, learned acoustic communication systems can be remarkably consistent. Substantive and descriptive generalizations can be made which hold for a majority of populations within a species. On the other hand, learned acoustic communication systems are often highly variable. The degree of variation is often so great that few, if any, substantive generalizations hold for all populations in a species. Within my dissertation, I explore the interplay of variation and uniformity in three vocal learning species: budgerigars (Melopsittacus undulatus), house finches (Haemorhous mexicanus), and humans (Homo sapiens). Budgerigars are well-known for their versatile mimicry skills, house finch song organization is uniform across populations, and human language has been described as the prime example of variability by some while others see only subtle variations of largely uniform system. For each of these species, I address several questions related to variability and uniformity: What is the typical range of variation? What are the limits of variation? How are those two issues related? And what mechanisms underlie variability and uniformity? iv In chapter 3, I investigate a potential domain of uniformity in budgerigar warble: the segment. Segments, units divided by acoustic transitions rather than silence, have been largely ignored in non-human animal communication. I find that budgerigars can achieve a high degree of complexity and variability by combining and arranging these small, more stereotyped units. Furthermore, I find that budgerigar segment organization is not only consistent across independent budgerigar populations but is consistent with patterns found in human language. In chapter 4, I investigate variability in house finch song. I present data showing that house finches learn sound patterns which are absent in wild house finch populations. These data suggest that cross-population variation in house finch song is narrower than what is permitted by the house finch song learning program. Finally, in chapter 5, I focus on human language, the most well-described communication system. Here, I research a sound pattern that is absent in the majority of known languages. I find that the rare pattern has independently developed at least six times. In every case, the historical pathway which led to the rare pattern was the same. The historical development in these six linguistic lineages suggests that the overall rarity of the sound pattern is the result of acoustic similarity. These data illuminate the evolutionary forces that give rise to, and limit, variation. The results of this dissertation have wide-ranging implications, from necessary revisions of linguistic theories, to understanding epigenetic interactions, to the application of evolutionary theory to complex behavior. While these projects within the dissertation are all different, evidence from all three projects support the following claims: (i) cross-population commonality is not evidence for what a species is able to learn; (ii) peripheral mechanisms have a strong influence in limiting cross-population variability; and (iii) high degrees of variation can emerge from uniform traits. v Acknowledgments First, I would like to thank my supervisor, Professor Juliette Blevins. Her knowledge, guidance, and patience have been invaluable throughout my PhD. I’m immensely grateful for her support when I wanted to take a not-so-typical route through a linguistics PhD. My committee has also been amazing, and I could not have done this without them. Professor David Lahti has an infectious passion for science which helped convince me to pursue a comparative approach and ultimately won me over to the bird side. Dr. Marisa Hoeschele is a fount of knowledge and has been a source of inspiration. She believed in me when I didn’t believe in myself. Dr. Michael I. Mandel was generous with his time and his input was essential in helping me refine my chapters. I would also like to thank the Ernst Mach Grant committee and the Austrian Science Fund for funding parts of this dissertation, namely Chapter 3. I am also grateful to the Florence Bloch Dissertation Fellowship committee for providing the funds for writing this thesis. My PhD has been a journey full of self-doubt and self-discovery. This journey has taken this little Arkie from the Graduate Center to the University of Santa Cruz, then to Queens College, and finally to the University of Vienna. Along the way so many colleagues, collaborators, staff, and friends have provided insight, support, and guidance. I want to thank the members of the Lahti Lab, the Pinniped Cognition and Sensory Systems Laboratory, and the Budgerigar Lab. I would also like to thank May Ahmar, Dr. Arafat Angulo-Perkins, Sabrina Bettoni, Nishi Bissoondial, Alexandra Bohmann, Dr. Daniel Bowling, Professor Diane Bradley, Dr. Caroline Casey, Katherine Dawson, Sarah Deventer, Professor Bob Dooling, Adam Fishbein, Professor W. Tecumseh Fitch, Frances Geller, Professor Stefano Ghirlanda, Felix Haiduk, Riccardo Hofer, Professor John Locke, Nadja Kavik, Theresa Matzinger, Phillip Maier, Dr. Ignacio Montoya, the late Professor Paul C. Mundinger, Dr. Jinook Oh, Petra Pesak, Dr. Stephan Reber, Dr. Colleen Reichmuth, Dr. Andrew Richards, Dr. Cliodhna Quigley, Dr. vi Michelle Spierings, Lauren Spradlin, Gabriel Staubmann, Dr. Hsiao-Wei (Vicky) Tu, Laura Waddick, Bernhard Wagner (hi Bernhard!), and Mason Youngblood for thoughtful discussions and input during my PhD. I’d like to give a special thanks to Shahrzad Afroozeh for her extensive technical help and, more importantly, extended conversations about budgie drama and to Dr. Christen Madsen II for moral support from day one of my PhD and, of course, for the countless hours we spent at chain restaurants in NYC. Finally, I’d like to thank my family. To Maria and Alfred, Fabi, Andreas and Martina, Oma, and the Loimersdorf Großeltern: for taking care of Cass so that Raffi and I could have a break and for the words of encouragement. Now that the dissertation is complete I can focus more on learning German so that I can understand Andreas’s jokes. To Shelli, for the laughs and for being a source of strength. I might have the titles but I’m proud that my little sister is still the smart one. To my dad, Gary, for teaching me to question everything and to keep an open mind. Because of him I learned to love the written word. To my mom, Marcella, words can’t express how grateful I am for all of your support. You taught me the value of hard work and perseverance. You sacrificed so much to give us a better life. All my successes I owe to you. Last, but not least, to Cass and Raffi. My dog, Cass, tried his best to give emotional support when I was swearing at my computer, but, bless his heart, he’s a shit emotional support dog. Still, he was always there with loving eyes. He kept me grounded, made me exercise, and forced me to be calm (if only to not wake him up so he wouldn’t walk over and “help”). Raffi, you have been an amazing partner. You helped me keep perspective, you made me laugh, you kicked my ass when I needed it. I’m in awe of your brilliance. vii Table of Contents Chapter 1. Understanding limits on variation in human language and elsewhere ............................................................................................................................... 1-1 1.1. Introduction ..................................................................................................................... 1-1 1.1.1. Background and terminology .................................................................................... 1-4 1.1.2. Vocal learning ........................................................................................................... 1-5 1.1.3. Units of analysis ........................................................................................................ 1-8 1.1.4. Terminology ............................................................................................................ 1-11 1.2. Variation and uniformity in human language ............................................................... 1-15 1.2.1. Degree of variation in human language ................................................................... 1-15 1.2.2. Limits on variation in human language ................................................................... 1-17 1.2.3. Causes of uniformity and variation in human language .......................................... 1-20 1.3. Variation and uniformity in birdsong ..........................................................................
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