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The Vocal Behavior of the American Crow, Corvus Brachyrhynchos

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The Vocal Behavior of the American Crow, Corvus Brachyrhynchos THE VOCAL BEHAVIOR OF THE AMERICAN CROW, CORVUS BRACHYRHYNCHOS THESIS Presented in Partial Fulfillment of the Requirements for the Degree Master of Sciences in the Graduate School of The Ohio State University By Robin Tarter, B.S. ***** The Ohio State University 2008 Masters Examination Committee Approved by Dr. Douglas Nelson, Advisor Dr. Mitch Masters _________________________________ Dr. Jill Soha Advisor Evolution, Ecology and Organismal Biology Graduate Program ABSTRACT The objective of this study was to provide an overview of the vocal behavior of the American crow, Corvus brachyrhynchos, and to thereby address questions about the evolutionary significance of crow behavior. I recorded the calls of 71 birds of known sex and age in a family context. Sorting calls by their acoustic characteristics and behavioral contexts, I identified and hypothesized functions for 7 adult and 2 juvenile call types, and in several cases found preferential use of a call type by birds of a particular sex or breeding status. My findings enrich our understanding of crow social behavior. I found that helpers and breeders played different roles in foraging and in protecting family territories from other crows and from predators. My findings may also be useful for human management of crow populations, particularly dispersal attempts using playbacks of crows’ own vocalizations. ii ACKNOWLEDGEMENTS I would like to thank Dr. Kevin McGowan of Cornell, Dr. Anne Clark of Binghamton University, and Binghamton graduate student Rebecca Heiss for allowing me to work with their study animals. McGowan, Clark and Heiss shared their data with me, along with huge amounts of information and insight about crow behavior. None of my work would have been possible without their help. I would also like to thank Dr. Douglas Nelson, Dr. Mitch Masters, Dr. Larry Phelan and Dr. Jill Soha of The Ohio State University for their help at all the stages of my research. Finally, I owe a great deal of thanks to my husband, Anton Mates for all of his help – from accompanying me in the field, to programming help, to editing and revising my thesis. iii VITA May 29,1982 ......................................Born – Walnut Creek, California 2000 ....................................................B.S. Integrative Biology, The University of California at Berkeley 2004 – present.....................................Graduate Teaching and Research Associate, The Ohio State University FIELDS OF STUDY Major Field: Evolution, Ecology and Organismal Biology iv TABLE OF CONTENTS Page Abstract……………………………...…………………................................…….....……ii Acknowledgements……………………………………...……................................….....iii Vita…………….…………………................................…...……………………….…….iv List of Figures…………………………………………...............................………....…..vi Introduction………………………………………………...............................…….....…..1 Methods………………………………………….................................………….......……3 Results………………………………………………………......................................…....9 Discussion……………………………………………...............................….......………29 Bibliography……………………………...……………..….......................................…..37 v LIST OF FIGURES Figure Page 1. Spectrograms of short, rough, fading, and scream calls...............................................11 2. Spectrograms of nasal, double short, and double rough calls.......................................12 3. Spectrograms of coo and rattle calls.............................................................................18 4. Fraction of adults that gave each call type by sex and breeding status.........................22 5. Call types shown separately by sex and breeding status...............................................23 6. Spectrograms of begging and babble calls....................................................................25 7. Fraction of juveniles of each sex that gave each call type............................................28 vi INTRODUCTION The American Crow, Corvus brachyrhincos, is a highly intelligent songbird, with an exceptionally large brain relative to its body size (Lefebvre et al. 2002). American crows display advanced problem solving skills and exceptional memories. They mature slowly and disperse late compared with other passerines (Caffrey 1992). They live in extended families, generally composed of juveniles, young from previous years, and one pair of breeding adults. Individuals may disperse as early as 2 months post fledging or stay in their natal group for 7 years or longer and different populations display either male or female biased dispersal (Marzluff & Angell 2005 and Caffrey 1992 respectively). It is unknown whether non-dispersing individuals (referred to as helpers) gain fitness benefits from this cooperative breeding behavior; so far no studies have succeeded in finding reproductive benefits of cooperative breeding for either helpers or breeders (Caffrey 2002). There are several reasons to study American crows other than their ubiquity, intelligence and unusual social system. American crows have great economic impact on many industries, including agriculture and civic planning. California pistachio growers, for instance, suffer an average damage of over a hundred dollars per hectare annually, and growers of nuts and fruits lose 3-4% of their yield to crows (Delwiche et al. 2007). 1 Crow communal roosts inconvenience humans across the country. In the winter, crows form giant groups of as many as two million individuals (McGowan 2005), often within city limits due to the light, warmth and lack of predators in urban habitats, making noise and leaving waste where they gather. There has been considerable interest in using crows’ own calls to disperse these roosts (Delwiche et al. 2007, Frings and Frings 1957), and the relatively low auditory frequency cutoff (5.6 kHz) found in the closely related hooded crow suggests that these species are adapted to long-range communication, which would make them good targets for acoustic dispersal (Jensen and Klokker 2006). The questions of how cooperative breeding behavior is maintained, and how humans can best disperse unwanted congregations of crows can both be partially resolved by examining the vocal behavior of individually tagged and human-habituated wild crows. The fact that my study population was almost entirely comprised of tagged, human-habituated animals sets it apart from all other studies of crow acoustics except one (Yorzinski et al 2006). Untagged crows are not reliably identifiable as individuals by human observers, and since at a distance there are no known indicators of their age or sex, observations of untagged individuals usually cannot be classified accordingly. With this problem removed, I was better able to ascertain contexts for the call types that I observed, and in many cases describe the responses they evoked in other known individuals. The differences I observed between the vocalizations of breeders and helpers sheds light on the benefits of cooperative breeding in this species. Moreover, I hypothesize functions related to group movement and foraging for certain call types, raising the possibility of exploiting those calls to relocate nuisance birds. 2 METHODS My study took place in Cayuga Heights, New York (42°N, 76°W) from the beginning of June to the end of August, 2006. At the beginning of my study, the young had just fledged and at the end they were beginning to become more active and some ventured away from their families. Beginning in 1989, the crow population in this area has been monitored and nestlings have been marked with unique combinations of leg bands and wing tags to facilitate individual recognition (Yorzinski 2006). The majority of the individuals in my study had had blood samples taken in the nest, which were used to determine their sex. The first phase of my research was a general observation of the animals in the study area. I spent 2 weeks following the members of the Ithaca crow project learning about the habits of the animals in their study. After this period of observation, I choose 10 families to focus on for the rest of my study. My selection was based on several factors. First, I wanted to use families with multiple tagged individuals so that I could identify the animals I was recording. I also wanted families that were habituated enough to human activity on their territories that they would behave normally while I recorded them. It was also important for me to include individuals of varying age and reproductive status in my study to allow me to examine the ontogeny of crow acoustic 3 behavior and to compare the behavior of helpers and breeders. All of the families I chose had 2 breeders, from 0 to 5 adult nonbreeders (helpers), and from 2 to 5 juveniles (first year birds). Ages ranged from less than 1 to 13 years. I wanted to have as much variation in the composition of my study families as possible in order to examine the possible effects that different composition might have on behavior. Finally, because I decided to make all of my recordings from inside my car, it was necessary to choose families that frequented car accessible locations. I chose to do all of my focal watches from my car because I found that, when I approached the crows on foot, the only vocalizations I could record were alarm-associated. Parr (1997) made the same observation. I was able not only to tell individuals apart and compare their behaviors with respect to their sexes and breeding conditions, but also to get closer
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