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Variation in the Bout Structure of Northern Mockingbird (Mimus Polyglottus) Singing

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Variation in the Bout Structure of Northern Mockingbird (Mimus Polyglottus) Singing Bird Behavior, Vol. 13, pp. 93–98, 2000 1056-1383/00 $20.00 + .00 Printed in the USA. All rights reserved Copyright © 2000 Cognizant Comm. Corp. Variation in the Bout Structure of Northern Mockingbird (Mimus polyglottus) Singing NICHOLAS S. THOMPSON,* EMILY ABBEY,* JESSICA WAPNER,* CHERYL LOGAN,† PETER G. MERRITT,‡ AND ALBERT POOTH§ *Departments of Biology and Psychology, Clark University, Worcester, MA †Department of Psychology, University of North Carolina, Greensboro, NC ‡Treasure Coast Regional Planning Council, Stuart, FL §Department of Biology, University of Miami, Miami, FL The highly variable singing of the northern mockingbird (Mimus polyglottus) is distinguishable from that of other sympatric mimids by its organization into bouts: the bird’s tendency to repeat an element several times before proceeding to another. To determine the degree to which this bout structure is a common feature of mockingbird song, 10 samples of singing from widely different populations and circumstances were examined including examples from coastal and central New England, central North Carolina, and the Florida peninsula and examples of day and night and spring and summer song. Measures included note parameters (peak frequency, internote interval, and note duration) and bout parameters (songs per bout and mean values of note parameters). All samples were found to be organized in bouts, but the degree of differentiation of the bouts (i.e., the degree to which the boundaries between bouts were emphasized by the contrast between their songs) varied both within and between samples. Bout differentiation was not maxi- mized: songs of bouts sung in close temporal proximity were more similar than average, and the perfor- mance overall seemed to consist of runs of high and low values of note and/or bout parameters. Whether these variations in the bout structure reflect changes in the state of the singer or in his circumstances or serve to enhance the overall effectiveness of his performance remains to be determined. Mimidae Mockingbird Communication Song variation Song structure Among the primary functions of birds’ singing species, the male’s singing is practically invariant. are the communication of the species and individual Performances vary only subtly across the individuals identity of the singer. If singing is to identify sing- of the species, and each singer repeats the same sound ers both as species members and as individuals, then or, at most, sings one or two variants. The species- at least one property of the performance must be specific properties of such a performance are obvi- consistent across the members of the species, and at ous. In other species, the individual males sing hun- least one property must be variable from species dreds, perhaps thousands, of different sounds, and member to species member but consistent within a performances vary strikingly from individual to indi- species member (Marler, 1960). vidual. For species with such highly variable perfor- Birds differ strikingly in the degree of variation in mances, identification of the species-specific proper- their singing (Catchpole & Slater, 1995). In many ties of the performance may be less straightforward. 93 94 THOMPSON ET AL. One taxonomic group particularly known for the about 100 ms in duration separated in time from variety of its singing is the mimic thrushes. Within neighboring notes by silence. Notes are gathered into this group, three species breed sympatrically and units of repetition called songs. Songs can be classi- during overlapping time periods in the eastern United fied into readily recognized discrete types called States: the gray catbird (Dumetella carolinensis), the song types. During singing, the mockingbird imme- brown thrasher (Toxostoma rufum), and the north- diately repeats songs of one type several times be- ern mockingbird (Mimus polyglottus). All three are fore switching to songs of a different type. These repertoire-singing species with an extraordinary va- sequences of immediate repetition of song types are riety of song elements. Both the catbird and the called “bouts.” Mockingbirds may also produce mockingbird are known to sing hundreds of repeat- songs in temporal clusters called “groups.” Group able vocal elements, and the brown thrasher is boundaries do not necessarily correspond to bout thought to sing thousands (Boughey & Thompson, boundaries (Figs. 1 and 2). 1976; Kroodsma & Parker, 1977). Furthermore, all Although the bout structure of mockingbird song three species incorporate the vocal elements of other is thought to be a species-identifying property, it is species in their performances. not an invariant one. Bout length varies in a mock- The extraordinary variability of mimic thrush song ingbird performance, and its frequency distribution elements raises questions concerning auditory dis- overlaps with the bout length distributions of the crimination among the three species. In the field, sympatric mimids (Boughey & Thompson, 1976, the three species are traditionally distinguished by 1981) (Fig. 2). In the field, the birds also appear to the number of repetitions of song elements: catbirds vary the salience of the bout structure by varying tend to sing each element only once and brown the distinctiveness of the songs that make up neigh- thrashers once or twice, whereas mockingbirds typi- boring bouts and by varying the duration of pauses cally repeat each song element several times before that occur within and between bouts. going on to the next (Mathews, 1904). Every mock- The purpose of this investigation was to examine ingbird performance has at least three levels of or- variations in the bout structure of northern mock- ganization called notes, songs, and bouts (see ingbird song performances taken from a range of Derrickson & Breitwisch, 1992; Howard, 1974; circumstances and locations to see if patterns of Moody, Kedoux, & Thompson, 1994; Wildenthal, variation therein were consistent with a species iden- 1965; for additional information and song terminol- tification role or if they suggested other communi- ogy). A note is a continuous utterance of sound of cative functions. Figure 1. A typical sequence of mockingbird song with multiple repetition of songs. VARIATION IN MOCKINGBIRD SINGING 95 Figure 2. A sequence of mockingbird song with very short bouts. In this unusual se- quence of mostly one-song bouts, the singer is linking temporal groups by using some part of each previous group in the next. Methods Macintosh Quadra 700 running Sound Designer II (Digidesign, 1992) and converted into spectrograms Ten samples of mockingbird song were obtained in Canary 1.1 or 1.2 (Bioacoustics Research Pro- that were designed to represent widely different lo- gram, 1995). The breeding season samples from cations and recording circumstances (Table 1). The North Carolina were digitized at 22,050 Hz; all other samples were obtained from Massachusetts, North samples were digitized at 44,100 Hz. After being Carolina, and Florida. Some were recorded in the digitized, each sample was separated into blocks of day, some in the night; some were recorded in the approximately 20 s. breeding season, some outside the breeding season. Once converted into spectrograms, samples were With one briefer exception, all samples were ap- measured, note by note, by student technicians. Four proximately 3 min in duration and contained 700– song features were measured for each note using 1100 notes. All samples were digitized on a Canary’s 1.x’s automatic logging feature: frequency Table 1. Sources of the Samples of Mockingbird Song Used in this Study and the Characteristics of the Samples Samplea Region Season Time of Day Mate Status (if known) Sample Length NC-1 Central North Carolina breeding day mated 232 s NC-2 Central North Carolina breeding day unmated 190 s NC-3 Central North Carolina nonbreeding day 239 s MA-1 Central Massachusetts breeding day 162 s MA-2 Coastal Massachusetts breeding night 376 s FL-2a Florida breeding night unmated 206 s FL-2b Florida breeding day unmated 45 s FL-2c Florida breeding night unmated 198 s FL-1a Florida breeding day 277 s FL-1b Florida nonbreeding day 196 s aArabic numerals (NC-1, NC-2, etc.) denote different individuals; a, b, c indicate different recordings from the same individual. 96 THOMPSON ET AL. at peak of amplitude (peak frequency) in Hz, dura- song of the next bout, and (4) the first song of the tion in ms, interval in ms, and peak amplitude (Ca- bout five bouts later in the sound record. The tetrads nary 1.1 and 1.2) (Peak amplitude measurements are provided the basis for three sets of sound similarity valid only for comparisons within sample.) The be- correlations differentiated by the distance apart in ginnings and endings of notes were determined by the sound record: correlations between songs within enlarging spectrograms and playing them at 1/8 the the same bout, between songs from neighboring normal speed. The reliability of each technician in bouts, and between songs five bouts apart in the per- determining where a note began and ended was formance. achieved by using an explicit measurement proto- col and was confirmed by correlating his or her Results measurements of the notes of a standard selection done by all previous technicians. The 10 samples exhibited the variability and volu- After samples had been measured, the place of bility characteristic of northern mockingbird sing- each note was identified in the bout/song/note struc- ing. All singers added new note and song types as ture. Song boundaries were determined by the be- their performances progressed and bouts of old types ginning and ending of a repeated sequence of notes, were seldom repeated. bout boundaries in a change in the sequence of songs that were repeated. These distinctions were straight- Variation Between Samples forward in most cases. Difficult cases were notes Every variable we examined varied significantly not obviously elements of songs. Single notes that between samples (one-way ANOVA for sample, were repeated were treated as a bout of one-note df =9, p< 0.0001).
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