TY - JOUR
AU1 - Liu, Wan-Chun
AU2 - Kroodsma, Donald E.
AB - Abstract Songbirds typically have small-to-large repertoires of different song types used in complex patterns over a day or season, but what remains poorly understood are patterns of song use by songbirds that have a single, simple song. Here, on the basis of extensive field observations, we reveal how a male Chipping Sparrow (Spizella passerina) varies its simple, repetitive song to create a dynamic singing performance. During the day, and beginning shortly after males arrive on their territories, males typically sing long songs at a relatively slow rate from the tops of trees near the center of their territories. About two weeks after males have arrived, they begin to sing well before sunrise; during this dawn chorus, songs are brief and delivered rapidly, typically while males face each other on the ground at territorial boundaries. Each male sings this way during the dawn chorus throughout the breeding season, except when his female is fertile. By contrast, daytime singing essentially stops after a male pairs. The dawn chorus appears to mediate social interactions among territorial neighbors, whereas daytime songs function in long-distance territory advertisement, particularly for female attraction. Comportamiento de Canto al Amanecer y Durante el Día en Spizella passerina Resumen Las aves canoras típicamente tienen repertorios pequeños a grandes de distintos tipos de canto empleados en patrones complejos a lo largo del día o de la estación, pero aún se conoce muy poco sobre los patrones de uso de los cantos por parte de aves canoras que tienen un solo canto simple. Con base en observaciones de campo intensivas, en este estudio revelamos cómo un macho de Spizella passerina varía su canto simple y repetitivo para crear un despliegue de canto dinámico. Comenzando poco después de que arriban a sus territorios, durante el día los machos típicamente emiten cantos de larga duración a una tasa relativamente lenta desde la parte alta de árboles ubicados cerca del centro de sus territorios. Aproximadamente dos semanas después de su llegada, los machos comienzan a cantar bastante antes del amanecer. Durante este coro del amanecer, los cantos son breves y emitidos rápidamente, típicamente mientras los machos se enfrentan entre sí en el suelo cerca de los límites de los territorios. Cada macho canta de este modo durante el coro del amanecer a lo largo de la época reproductiva, excepto cuando su pareja está fértil. En contraste, el comportamiento de cantar durante el día esencialmente se suspende luego de que el macho adquiere una pareja. El coro del amanecer parece mediar en las interacciones sociales entre vecinos territoriales, mientras que los cantos emitidos durante el día funcionan como anuncios territoriales a grandes distancias, particulamente para atraer a las hembras. Thirty minutes to an hour before sunrise, many animals engage in intense vocalizing, and the intensity of the dawn chorus then gradually subsides as sunrise approaches (Staicer et al. 1996). Among most songbird species, dawn singing is more variable, complex, or elaborate than daytime singing (Spector 1992). This song complexity or variability during the dawn chorus differs greatly among species, and is perhaps associated with, or constrained by, the song repertoire size of each species. A Song Sparrow (Melospiza melodia), for example, has a repertoire of eight or so different songs; the rate of switching among song types is much higher at dawn than later in the day, and the frequency of matching song types with neighbors changes over the season (Beecher et al. 1996). By contrast, among several genera of paruline warblers, males use two distinct categories of song, one primarily at dawn and the other during the day (Spector 1992), and these distinct diel singing behaviors seem to have different functions (Nelson and Croner 1991, Spector 1992, Staicer et al. 1996). It remains unclear how these patterns of repertoire use evolved. Perhaps by better understanding patterns of song use in a songbird in which males have but a single, simple song, we might better understand the forces that have selected for patterns of complexity in species that have larger repertoires. The Chipping Sparrow (Spizella passerina) has one of the simplest songs among songbirds (Fig. 1). Each male has only one song type, and each song consists of one type of syllable repeated many times (Marler and Issac 1960, Borror 1961, Middleton 1998). Here, we describe how a male uses his song at different times of day and under different social circumstances, revealing distinct dawn and daytime singing behaviors that change over the season. Fig. 1. Open in new tabDownload slide Chipping Sparrows gradually change their singing behavior from dawn to daytime. (A) Male 1 sang most intensely at 0430 hours, delivering about one song each second, but song duration and the intersong interval gradually increased until he was singing a much longer song about every six seconds by 0600 hours. (B) For some males, however, only the intersong interval changed appreciably, song duration remaining essentially unchanged. Methods During April–August of 1996-1999, we studied Chipping Sparrows at several locations in western Massachusetts: Quabbin Cemetery, Ware; Wildwood Cemetery, Wildwood Elementary School, Amherst; and the campuses of the University of Massachusetts, Hampshire College, and Amherst College, Amherst. At these locations, 125 territorial male Chipping Sparrows were mist netted and color-banded. When color bands could not be seen before sunrise, individual males were identified by their unique songs (songs were recorded and sonograms were later compared to songs in a catalogue of songs for the population). Songs were recorded on a Marantz PMD 222 taperecorder with a Dan Gibson parabola and Sennheiser ME 62 microphone. We used a Kay Elemetrics 5500 DSP real-time spectrum analyzer to analyze the songs (bandwidth: 2–10 kHz). The dawn chorus occurred 30–60 min before sunrise (as listed in a meteorological table) and ended in one of two ways, depending on the mating status of the focal male. Because paired males rarely sing after sunrise, we defined the end of their dawn chorus as the time they stopped singing from the ground. Unpaired males sing from the ground during the dawn chorus (as do paired males), but they sing from elevated perches during the day, so we defined the end of their dawn chorus as the time they stopped singing from the ground and flew to a tree where they resumed singing. Daytime song, by definition, occurred after the dawn chorus ended and throughout the day. We recorded daytime song as late as 1100 hours EST. To compare dawn and daytime songs of unpaired males from late April to mid-May, we chose 15 males from the four study areas for which we had the best information (i.e., quantity and quality of recordings). For dawn song, we chose ≥20 songs from each male within the first 15 min of the dawn bout (every 5th to 10th song was chosen during each dawn bout, depending on a male's song rate). For daytime song, we chose 20–36 songs from each male recorded. For each male, several song features were analyzed: dawn bout length (total duration of dawn singing, in minutes), singing rate (number of songs per minute), song duration, and the time interval between songs. To compare dawn and daytime singing behaviors, we used 22 males (6 in 1997, 7 in 1998, and 9 in 1999). Each male was part of a “neighborhood” of at least two territorial males. During each dawn chorus, we recorded two to four neighboring males, and each male's song was recorded for 2–5 min before we recorded another male. During each recording, we spoke into the microphone and described the songperch height, singing location and direction (singing direction was determined by whether a male sings facing toward or away from its closest neighbor's territory; each male has one or two immediate neighbors, with no others in proximity), and associated behavior of each male. After sunrise, we recorded the daytime singing behavior of each of these males for ≥10 min on any given day, for 53 days in 1997, 32 days in 1998, and 66 days in 1999. To describe how these dawn and daytime singing patterns change with breeding stage, we followed seven territorial males from two study areas and recorded their daily singing behaviors in 1996 (35 days of recordings from three males) and 1997 (37 days of recordings from four males). Each male's dawn songs were recorded every one to three days. During the day (0600–0900 hours), each male's singing behavior was recorded for half an hour to an hour every one to three days. After sunrise, we confirmed the breeding stage as nest building, egg laying, incubation, or nestling. When comparing dawn and daytime songs and behaviors, we used a two-tailed, Wilcoxon matched-pairs signed-ranks test, unless otherwise stated. For all analyses, sample sizes are always the number of males used in a test (the mean of multiple measures from a given male was used in the test). Results Song features.— Dawn songs of unpaired males were especially brief, typically less than half the duration of daytime songs (1.2 ± 0.44 s vs. 2.52 ± 0.56 s, t = 6, P < 0.001, n = 15 males, 20–30 songs per male). Even longer songs were sung occasionally by paired males during the day, especially during incubation (8.51 ± 3.76 s, based on 26 recordings from 14 males; compared with daytime song of unpaired males given above, n1 = 15, n2 = 14, P < 0.01). Dawn songs were brief because they contained fewer syllables (Fig. 1A), not because the syllable duration changed (dawn vs. daytime syllable duration, t = 47, P = 0.16, n = 15 males, 30–40 syllables per male). The singing rate for these unpaired males was much higher at dawn than during the day (21.6 ± 7.19 songs min−1 vs. 6.43 ± 1.07 songs min−1; t = 7, P < 0.001, n = 15 males; Fig. 2A). The dawn singing rate was higher not only because of the briefer songs at dawn (see above) but also because the time interval between two songs was much briefer at dawn than during the day (1.39 ± 0.62 s vs. 6.78 ± 1.03 s, t = 7, P < 0.001, n = males, 20–30 songs from each). These data characterize average differences between dawn and daytime singing; but during the dawn chorus, song duration and singing rate changed gradually (Figs. 1A and 2A). After sunrise, however, males sang longer songs with little variation during most of the day (see below). Fig. 2. Open in new tabDownload slide Dawn chorus and daytime singing differ in both singing rate and song-perch height. (A) Males sang with the greatest frequency at dawn, but they gradually reduced song rate toward sunrise. (B) At dawn, males typically stood on the ground, but after sunrise they moved to higher singing perches. Each data point represents the mean (± SE) of song rate (A) or song-perch height (B) from eight territorial males, and each male's song rate or perch height is the average of four days' recordings, one hour each day, during mid-May 1997. Dawn singing patterns also differed among individuals. Most males shortened both song duration and intersong interval at dawn. Some males, however, shortened only their intersong interval, whereas the song duration remained the same as daytime songs (Fig. 1B). Owing largely to the high variation of song duration and intersong interval, the dawn singing rate was highly variable among individuals (21.6 ± 7.2 songs min−1, coefficient of variation (CV) = 33.3%, n = 15 males). During the day, the variation in song rate among unpaired individual males was much smaller (6.4 ± 1.1 songs min−1, CV = 16.3%, F = 3.21, df = 14 and 14, P < 0.05, n = 15 males; Sokal and Braumann 1980). (Daytime song rate of the paired males is not included here, because these birds rarely sang during the day.) Song-perch height.— Male Chipping Sparrows sang at dawn while standing on the ground or on low perches (0.9 ± 0.9 m, n = 15 males, 7–18 samples from each). After (or close to) sunrise and throughout the day, however, males sang from a higher perch, such as the top of a tree or building (8.5 ± 2.3 m, n = 15 males; dawn and daytime heights different: t = 7, P < 0.001, n = 15; see Fig. 2B). Singing location.— At dawn, males frequently came to the boundary of their territories and sang (distance to territory boundary: 12.4 ± 9.2 m, n = 15). Near sunrise, these males returned to the centers of territories and continued singing there. Throughout the day, males sang mostly near the territory center, and singing locations during the day were much farther from the nearest territory boundary (35.8 ± 17.6 m; dawn and daytime locations different: t = 23, P < 0.05, n = 15). The singing location at dawn was also farther from the nest site than daytime song locations (32.4 ± 12.3 m vs. 12.2 ± 8.0 m, t = 21.5, P < 0.05, n = 15 males). After nest failure, female Chipping Sparrows often shifted their nest site. Their mates, however, did not change their dawn singing locations after a female's nest site changed (11 of 12 males). Singing direction.— At dawn, a male typically faced the territory of his immediate neighbor while singing (an average of 92% of the time, singing males faced the neighboring territory; this is based on the averages of 8–23 samples from each of 10 males). During the day, however, the singing of males was less directional (only 71% of time spent facing the nearest neighbor, n = 10 males). Breeding stage.— At dawn, males sang during all breeding stages, except during the fertile period of their females (nest-building and mating period; Fig. 3). Of 29 territorial males recorded, 26 gave no or little dawn song on days when the territorial male and his female were copulating, days when the female would typically build the nest after sunrise. During these days, the territory owner and his immediate neighbors, more distant neighbors, or unknown intruders frequently chased and fought each other. Fig. 3. Open in new tabDownload slide Dawn and daytime singing patterns differ across the breeding stages (data collected from seven territorial males with 72 days of recording; on each day, each male's dawn and daytime singing was recorded for ≥30 min; see text for details). The dawn chorus occurred throughout the breeding season, except when a male's female was fertile (M,B = mating or copulations, nest-building); continuous daytime singing occurred only when a male was unpaired. U = unpaired, P = just paired, L = egg-laying period, I = incubation period, N = nestling period, F = fledgling period. During other breeding stages and regardless of pairing status, males consistently sang during the dawn chorus. The duration of the dawn bout for a male remained fairly constant throughout the breeding season (except during the fertile period), but the dawn bout was somewhat longer during the incubation period than during the nestling and fledgling periods (incubation vs. nestling period, t = 10, P < 0.05, n = 7 males; incubation vs. fledgling period, t = 7, P < 0.025, n = 7 males; corrected by Bonferroni test). Only unpaired males sang continuously throughout the day (at a fairly constant singing rate of 6.7 ± 0.9 songs min−1, n = 22 unpaired males). After pairing, however, they sang only occasionally (0.0 ± 0.1 songs min−1, n = 12 paired males; Fig. 3). Seasonal patterns.— After claiming a territory in early to mid-April, unpaired males started singing during the day. During this time, these males sang, at most, a few minutes of dawn songs, but most initially sang no dawn songs at all (Fig. 4). During early May, two to three weeks after the beginning of daytime singing, males began singing a dawn bout of 20–40 min. At this time, most male neighbors had arrived and females had begun arriving. During late July, males shorten their dawn bout length and stop dawn singing when they stop defending territories (late July to mid-August; Fig. 4). Fig. 4. Open in new tabDownload slide Seasonal patterns of duration of the dawn chorus. These data were acquired from 21 males in seven groups; 8 males in two groups were recorded every two to four days from arrival date in April to the end of the breeding season in early August 1996, and the other 13 males in five groups were recorded every two or three days from May through late July 1997. For each five-day period, we first determined the mean duration of the dawn bout for each male; here, we present the grand mean (± SE) for all 21 males. Frequency of close-range alternating and overlapping.— During the dawn chorus, neighboring males at territorial boundaries often engaged in intense countersinging duels. When <10 m apart, they often faced each other while standing on the ground, with one male beginning his song during the intersong interval of his neighbor. During these alternating bouts, it was often clear that the males adjusted their song duration or delivery rate to accommodate each other (W.-C. Liu unpubl. data). Sometimes during an interaction, a male consistently overlapped the song of his neighbor, beginning his song before the end of the neighbor's song; this type of countersinging was often followed by fighting, chasing, or high-pitched calls. Overall, 42 overlapping and alternating bouts (from 23 pairs of neighboring males) were recorded at dawn throughout the breeding season (April to late July) from 1997 to 1999 (n = 68.5 h of observations). During the day, however, we recorded only five close-range countersinging duels during territorial encounters; they all occurred during the early breeding season (April-May), when males were competing for their territories (n = 115.5 h of observations). Singing behavior of polyterritorial males.— From 1996 to 1998, we found six polyterritorial males that defended two separate territories. Each male first settled on a territory and acquired a mate there before settling on a second territory when his female was incubating (for definition, see Ford 1996). No matter where the female or nest was located, each male spent more time singing his dawn chorus on the first territory he settled. During the dawn chorus, four polyterritorial males sang only on the first territory (even though the female for one male was on his second territory), but two males also sang in their second territory. During the day, each polyterritorial male sang as if it was unpaired. These males thus sang continuously, with relatively constant song duration and singing rate, only when their females were not nearby (song rate at territory without female: 5.9 ± 0.8 songs min−1; at territory with female: 0.2 ± 0.7 songs min−1; t = 1, P = 0.036, n = 5 males). Discussion Our study reveals that Chipping Sparrows modify their simple songs in a continuous fashion for use at different times of day. As sunrise approaches, the short and rapidly delivered dawn songs gradually increase in duration and intersong interval. During the day, however, little variation was found in these features. Male Chipping Sparrows also change their song duration under other circumstances. During incubation, for example, males occasionally sing during the day, and these daytime songs are especially long. Or, males who alternate songs at close range adjust the song duration and intersong interval to accommodate each other (Liu 2001). Such variable singing behavior occurs in other songbird species as well. Male Great Tits (Parus major), for example, also have a song with a highly repetitive note; an individual may vary its song length in a singing bout, and song length may provide a short-term signal related to his willingness to respond (McGregor and Horn 1992). Kentucky Warblers (Oporornis formosus) and Black-capped Chickadees (Poecile atricapillus) also have a single song type. These birds, however, can lower or raise the frequency of the song in response to playback experiments (Morton and Young 1986, Weisman and Ratcliffe 1989, Otter et al. 2002), and patterns of singing seem to indicate relative male quality (Christie et al. 2004). These examples reveal that birds with simple songs can modify their songs in subtle ways to deal with different social situations, and these modifications may signal a male's aggressiveness, attention, or willingness to interact with others. Intrasexual function of the dawn chorus.— For several reasons, the dawn chorus of Chipping Sparrows appears to function primarily for close-range communication among neighboring males. (1) At dawn, neighboring males frequently come to the boundary of their territories where they face each other and sing at close range. Such close-range countersinging was common at dawn but not during the day. (2) Singing near the ground is more suitable for short-range than for long-range communication, because signals suffer increased reverberation and attenuation, particularly for species like Chipping Sparrows with high-frequency songs (Wiley and Richards 1978, 1982). (3) Territorial males sing few if any dawn songs during the early breeding season, when males are establishing territories and attracting females. The dawn chorus arises only after most neighboring males have settled on their territories. The dawn chorus thus seems to mediate social relationships among males, beginning about the time when females begin to arrive. (4) Neighbor-removal experiments show that a solitary male sings no songs at dawn after all neighboring males have been removed, but does not change his daytime singing behaviors. After the neighbors were released, the previously solitary male resumes his intensive dawn chorus (Liu 2004). Neighboring males thus have a strong influence on a male's dawn, but not daytime, singing behavior. Why do these neighboring Chipping Sparrows interact at dawn? Territorial relationships among neighboring birds are dynamic and complex (McGregor 1993, Smith 1997). Territorial neighboring males, for example, may compete for each other's territorial resources — such as nest sites, food, or females—throughout the breeding season. Neighbors can perhaps also cooperate in defending a territory against a stranger or predator, or in attracting females (Beletsky and Orians 1989). A territorial male may thus communicate frequently with his close neighbors as they announce or assess each other's strength or willingness to defend the territory or as they challenge each other. One might thus expect neighboring male Chipping Sparrows to use the close-range dawn chorus to communicate in this dynamic territorial system (the “social-dynamic” hypothesis; Staicer et. al. 1996). Perhaps male Chipping Sparrows use the dawn chorus as an honest signal to evaluate each other's quality, and neighboring females may use this signal to choose a high-quality male (“honest advertisement” hypothesis; Otter et al. 1997, 1998; Mennill et al. 2002). Intersexual function of the dawn chorus.— Whether or not the dawn chorus in Chipping Sparrows has an intersexual function is still unknown. Several lines of evidence, however, suggest that the dawn chorus of a male is not a signal to his own female. First, a male sings his dawn chorus throughout the breeding season, whether he is paired or unpaired. During the early breeding season, about two weeks after arriving, males start singing the dawn chorus even if they are still unpaired. Also, a male continues to sing the dawn chorus at the end of the breeding season, even if his female has left or shows no breeding activity. Second, males curtail singing at dawn, when their females are most fertile. Thus, males do not use dawn song to stimulate their fertile female, and the fertile female does not use the dawn song at that time to evaluate the quality of her mate. These observations thus exclude the “mate-guarding” hypothesis (Mace 1987). Perhaps silence during this period is a mate-guarding strategy, so that neighbors cannot locate this silent, defending male, or the male can more efficiently guard his mate from extra-pair copulations (see also Klit 1999). Third, males come to their territory boundary and sing an intense dawn chorus at neighboring males, and the singing locations are not close to the nest site of their females (females stayed in the nest during the incubation stage). Although the dawn chorus is not directed at a male's own female, neighboring females might eavesdrop on a male's dawn chorus by listening to the dawn countersinging of contesting neighboring males, and thereby be able to choose a high-quality male for extrapair fertilizations (Otter et al. 1999; see above). Function of the daytime song.— In contrast to dawn songs, the daytime songs of Chipping Sparrows appear to function primarily for long-distance communication for female attraction. (1) During the day, territorial males sing on a high perch and rarely interact closely with neighboring birds while singing. They also sing constantly, even if no neighbor is around (Liu 2004). The main receiver of daytime singing seems not to be the singing male's close neighbors. (2) After pairing, a male reduces daytime singing (but not the dawn chorus) significantly. Also, after a male loses his female, he resumes daytime singing that is typical of all unpaired males. These observations suggest that daytime song is used for female attraction. (3) All six polyterritorial males rarely sang daytime songs in a territory with females, just like paired mono-territory males. However, in their second territory, with no female around, each paired male constantly sang daytime song like an unpaired male. These singing patterns suggest that paired males sing daytime songs at the second territory to attract another female. Similar singing patterns can be found in other polyterritorial species, such as Wood Warblers (Phylloscopus sibilatrix; Temrin et al. 1984) and American Redstarts (Setophaga ruticilla; Secunda and Sherry 1991). Overall, our field observations reveal that male Chipping Sparrows can adjust their simple songs for singing under different circumstances. Particularly, songs used at dawn and during daytime differ in song duration or singing rate, or both, thereby permitting males to use this variable singing behavior to convey different messages in particular behavioral contexts or social circumstances. Perhaps the relatively stereotyped song during the day provides a stronger stimulus for female perception, whereas the variable song at dawn can provide each male with more plasticity to interact with his neighbor or to evaluate his strength. Interestingly, the dawn chorus and daytime singing show different gene-expression patterns in the Chipping Sparrow's forebrain song system (Liu and Nottebohm 2005), which suggests that different neural mechanisms are involved in these diel patterns of singing behavior. In the future, an integrative approach may help us better understand the evolution of the complexity of the dawn chorus. Acknowledgments We thank B. Byers, D. Spector, T. Miller, and an anonymous reviewer for their helpful comments on this manuscript. We are grateful to J. Fair of Wildwood Cemetery and to the Metropolitan District Commission at the Quabbin Reservoir for kindly allowing us to study there. Literature Cited Beecher , M. D. , P. K. Stoddard, S. E. Campbell, and C. L. Horning. 1996 . Repertoire matching between neighbouring Song Sparrows. Animal Behaviour 51 : 917 – 923 . Google Scholar Crossref Search ADS WorldCat Beletsky , L. D. , and G. H. Orians. 1989 . 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TI - Dawn and Daytime Singing Behavior of Chipping Sparrows (Spizella Passerina)
JF - Ornithology
DO - 10.1093/auk/124.1.44
DA - 2007-01-01
UR - https://www.deepdyve.com/lp/oxford-university-press/dawn-and-daytime-singing-behavior-of-chipping-sparrows-spizella-ECO5OLL50j
SP - 44
EP - 52
VL - 124
IS - 1
DP - DeepDyve
ER -