Ethology

Responses to Variation in Song Length by Male White-Crowned Sparrows

Douglas A. Nelson & Angelika Poesel
Borror Laboratory of Bioacoustics, Department of Evolution, Ecology and Organismal Biology, The Ohio State University, Columbus, OH, USA

Correspondence Douglas A. Nelson, Borror Laboratory of Bioacoustics, Department of Evolution, Ecology and Organismal Biology, The Ohio State University, Columbus, OH 43212, USA. E-mail: nelson.228@osu.edu

Abstract Bird song has the potential to convey a variety of information about the singer, including the singers current motivational state. A recent review by Searcy & Beecher (2009) (Anim. Behav. 78, 2009; 1281) emphasized how little is currently known about whether song functions as an aggressive signal in territorial interactions. A recent observational study of song length variation in the Puget Sound white-crowned sparrow reported that males shorten the terminal trill in their single song type while singing close by another male, and shorten the trill even further immediately before chasing the opponent. Here, we report the results of two song playback experiments designed to test whether males distinguish the differences in song length that occur in agonistic and nonagonistic contexts. We found that males gave stronger responses, as measured by close approach distances, to either a series of songs increasing in duration (compared to a series decreasing in duration), or to long loud songs (compared to short soft songs). In both experiments, males gave weaker responses to the stimuli hypothesized to be more aggressive. Males shortened their songs in response to both simulated intrusions. Combined with the observational data on song variation in different contexts, these experiments support the conclusion that song length variation contains information about aggressive motivation in the white-crowned sparrow.

Received: July 22, 2011 Initial acceptance: August 19, 2011 Final acceptance: September 28, 2011 (L. Ebensperger)
doi: 10.1111/j.1439-0310.2011.01979.x

Introduction In conicts over resources, animals commonly exchange signals rst before resorting, if necessary, to aggressive behaviours such as attacking or ghting (Maynard Smith & Harper 2003; Searcy & Nowicki 2005). The signals exchanged in these agonistic contexts, which may include both aggression and appeasement, may convey information about the animals quality, ghting ability and or motivational state. As long as there are costs associated with signalling that make a signal reliable to receivers, signalling may help avoid the costs (risk of injury, time spent) associated with aggression (Searcy & Nowicki 2005).
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Bird song plays a prominent role in territory acquisition and defence (Catchpole & Slater 2008), and various singing behaviours such as song type matching and song overlapping have been studied for their role in mediating territory disputes (Todt & Naguib 2000). While some aspects of song may provide information about the singers quality, which may remain fairly constant over periods of days to months, song can also vary on a moment-tomoment basis and can potentially provide information about the singers short-term motivation or intentions. In agonistic contexts, animals may benet by providing information about their aggressive intentions, their likelihood to attack, if this reduces the costs associated with escalated conicts. Searcy &
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D. A. Nelson & A. Poesel

Song Length Variation in Sparrows

Beecher (2009) have suggested three criteria that must be met in order for a signal to qualify as an aggressive signal: (1) The signal should occur frequently in aggressive contexts, (2) the signal should reliably predict attack by the signaller, and (3) receivers should respond to the signal. These authors reviewed several types of singing behaviours that have often been considered to be aggressive (song type matching, frequency matching, song overlapping, low-amplitude song (soft song hereafter), song type switching and vocal performance) with respect to these criteria and concluded that the evidence is fairly weak that these singing behaviours, with the exception of soft song (Dabelsteen et al. 1998), are aggressive signals. Song length is another character that varies within a singing bout, and could potentially provide information about aggressive motivation, as evidence from a variety of species suggests. For example, Jarvi et al.(1980) suggested that short songs by willow warblers, Phylloscopus trochilus, may signal intention to attack. Similarly, Capp & Searcy (1990) observed in the bobolink, Dolichonyx oryzivorus, that shorter songs were sung just prior to attack. Observational evidence on variation in song length in male Puget Sound white-crowned sparrows, Zonotrichia leucophrys pugetensis, meets the rst two of the three criteria outlined by Searcy & Beecher (2009) for establishing that a song characteristic is aggressive. In this species, most adult males sing a single song type that consists of a series of phrases (whistle, buzz, note complex, trill) presented in an invariant order (Fig. 1). Males vary the length of their song by varying the number of repeated syllables in the trill phrase (Nelson & Poesel 2011). With respect to the rst criterion, the contextual use of song, males sing songs with terminal trills averaging four syllable repetitions (range = 07) when singing spontaneously prior to pairing early in the breeding season (Fig. 1). In contrast, in close malemale territory interactions, males sing songs with shorter trills, averaging one to two syllables (Nelson & Poesel 2011). During these interactions, males sing shorter trills when <10 m from the opponent than when further than 10 m, and the short songs are sometimes of low amplitude. Second, in malemale interactions, variation in the length of the short trill is associated with the singers subsequent behaviour. Males shorten the trill just prior to attacking the opponent, but do not change trill length prior to ying away from the opponent (Nelson & Poesel 2011). All but one attacks occurred when the two males were <10 m apart. Observational evidence therefore indicates that the trill is
Ethology 118 (2012) 2432 2011 Blackwell Verlag GmbH

(a)

(b)

Fig. 1: (a) Un-manipulated song (long song) of a Puget Sound whitecrowned sparrow illustrating terminology (trill length = ve syllables). Spectrogram generated in Signal (Engineering Design) with a 256 point FFT from a le digitized at 25 kHz, step size 3 ms (frequency resolution = 98 Hz, temporal = 10 ms). (b) Single examples of songs with different trill lengths (short, long, elongated: S, L, E) used in Experiment 1. See text for description of synthesis procedure. Intervals between songs have not been preserved in the Figure.

shorter in agonistic contexts and variation in trill length within agonistic interactions predicts subsequent aggressive behaviour. The aim of this study was to address the third criterion listed by Searcy & Beecher (2009): do male receivers respond to differences in trill length? We performed two playback experiments that varied trill length. In Experiment 1, we presented males with a series of songs that either increased or decreased in trill length while maintaining amplitude and the total duration of songs throughout the trial constant. The goal of this design was to simulate an escalating interaction (decreasing trill length) vs. a de-escalating interaction (increasing trill length). In Experiment 2, we presented males with either a constant series of long loud songs, or a series of short soft songs. Here, we contrasted responses to songs that are normally sung when birds are singing undisturbed on their territories (long loud songs) vs. songs sung while males are close together in a territorial conict (short soft songs). Methods
Experimental Methods

We tested Puget Sound white-crowned sparrow males on their territories in Bullards Beach State
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Song Length Variation in Sparrows

D. A. Nelson & A. Poesel

Park, Bandon, Oregon, USA between 19 April and 4 May, 2007 (n = 12, Experiment 1) and 19 May, 2008 (n = 11, Experiment 2). Most males were colour-banded, and no male was tested twice within or between experiments. In 2007, all but three males were un-paired, in 2008 all subjects were un-paired. Both experiments were within-subject designs where we presented the two treatments to each male with approximately 1 h between treatments. Territories were mapped by observing song posts on days prior to testing. To simulate an intrusion by a newly arrived male, we placed a loudspeaker (PAL; Tivoli Audio, Cambridge MA, USA) atop a 1.6-mhigh tripod, 1015 m inside the boundary of the subjects territory, adjacent to an area unoccupied by a white-crowned sparrow. We chose a location with trees or shrubs distributed as evenly as possible near the speaker for the male to perch in. The two stimuli were presented to a subject on one morning between 06:00 and 12:00 PST. Presentation order was randomized among males. At least 48 h elapsed between tests of males on neighbouring territories. One observer sat 10 m from the speaker and observed the males behaviour during three periods: a 2-min-long pre-playback period, a 2-min-long playback period, and a 5-min-long post-playback period. All songs by the subject were recorded, and behavioural observations were dictated into the recorder. We recorded songs using a Marantz PMD670 solid-state recorder (Marantz Professional, Kanagawa, Japan) sampling at 48 kHz, 16-bit amplitude resolution, and Sennheiser MKH70 or ME67 shotgun microphones (Sennheiser, Wedemark, Germany) with Rycote windscreens (Rycote Ltd, Stroud, UK). Recordings were examined using a real-time spectrogram in either Signal (Engineering Design; Berkeley, CA, USA) or Syrinx (John Burt).
Stimuli, Experiment 1

(hereafter long song) using standard commands in Signal to create a song with a short trill (0 or 1 syllables, determined by coin ip) or an elongated trill (67 syllables, Fig. 1). The short trill song was created by removing terminal syllables. The elongated trill was created by rst duplicating one of the central syllables and then inserting one or two duplicates next to the central syllable while maintaining the same inter-syllable intervals. Prior to insertion, the amplitude of each duplicate syllable was adjusted, so that within the elongated trill, the amplitude of successive syllables gradually declined, as occurs in natural trills (Fig. 1). The shortened and elongated trills fell within the natural range of variation in trill length (Nelson & Poesel 2011). Each short, long and elongated song was duplicated four times at a constant song period (song onset to song onset) of 10 s. The two playback stimuli were (1) a series of songs of increasing trill duration (four short songs, four long songs, four elongated songs) and (2) a series of decreasing trill duration (four elongated, four long, four short). It should be noted that this manipulation controls the amplitude and total amount of song between the two treatments: all that differs is the order in which songs of different length occurred. The stimuli were then copied onto a Sony TC-D5ProII cassette tape recorder for playback. Songs were broadcast at a natural level (approximately 76 dB SPL at 8 m, measured with a Radio Shack digital sound level meter).
Stimuli, Experiment 2

In experiment 1, we compared responses to a series of songs with increasing trill length to a series with decreasing trill length. We used songs recorded in previous years (20042006) at Bullards Beach. Songs were digitized at or resampled to 44.1 kHz with 16-bit amplitude resolution, and high-pass-ltered >1500 Hz using Signal. For each subject (n = 12), we selected a different high-quality recording of a single song sung by a male in a previous year at least 500 m from the subjects territory. The other selection criterion was that the song contained 35 syllables in the terminal trill (Fig. 1). We then manipulated this natural-length prototype song
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In experiment 2, we compared responses to short soft songs to responses to long loud songs as used in spontaneous singing. Short songs sometimes sound much quieter than long songs in malemale interactions, although we have not measured sound amplitude in natural interactions as males move about rapidly (Nelson & Poesel 2011). To ensure that males detected short soft playback stimuli, we rst lured the subject to within 16 m of the loudspeaker by playing back repetitions of long loud song. As soon as the male landed within 16 m of the speaker, we either (1) played ten repetitions of the same loud long song, or (2) changed to ten repetitions of the same song with shortened trill and amplitude decreased by 12 dB. The )12 dB value produced a decrease in amplitude similar to what we perceived in the eld. Similarly, Anderson et al. (2007) played back soft song sparrow (Melospiza melodia) songs at an amplitude 11 dB lower than normal songs, an amplitude difference approximating the difference
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D. A. Nelson & A. Poesel

Song Length Variation in Sparrows

between songs designated soft and loud by a human observer. As in experiment 1, stimuli were high-quality recordings from previous years in the local population that were likely to be unfamiliar to the subjects. A different recording was used for each subject (n = 11). Using Signal, we created two stimuli for each male. First, we copied the long song ten times with inter-song intervals varying between  9.5 and 11 s (X = 10 s) to create a sequence of loud long songs. We then took the long song, shortened it so that it retained zero or one trill syllables (determined by coin-ip), and decreased the amplitude 12 dB by multiplying the amplitude values in the digital waveform by 0.25. This short soft song was then copied ten times maintaining the same song periods as in the long song sequence it was paired with. Digital les containing the song stimuli were played back from a Marantz PMD670 solidstate recorder connected to the PAL loudspeaker positioned as in Experiment 1. The loud long stimulus was played back at about 76 dB SPL at 8 m as measured by a Radio Shack sound pressure level meter; the short soft song was 12 dB below this. The short soft songs were audible to observers positioned 10 m away.
Data Analysis

duration of the period to yield the mean distance. In experiment 1, we also measured the males approach distance to the speaker during the rst 40 s, the middle 40 s and last 40 s of playback. These intervals correspond to when males heard short, long, or elongated songs depending on the treatment. The eight variables were entered into a principal components analysis, and the rst two principal components (PCs) were used as the dependent response variables in statistical testing. We compared the principal component scores for the two stimuli in each experiment using Wilcoxon matched-pairs signedranks tests. We also compared trill length between the pre-trial period and the playback period. For the few males that did not sing in the pre-trial period, we substituted trill length estimated from a 20-minlong focal animal watch conducted while the male was unpaired earlier in the breeding season. We report exact two-tailed probabilities for all tests as calculated in SPSS 19 (Chicago, IL, USA). Results
Experiment 1: Increasing vs. Decreasing Trill Length

Data were collected and analysed similarly in both experiments. During the 2-min-long pre-trial period, we counted the number of songs and the number of trill notes (hereafter trill length) in each song. We measured four response variables in both the playback and post-playback periods: the number of songs, trill length of each song, the mean approach distance and closest approach distance. We counted the number of trill notes in each song by viewing the recording of the trial in a real-time spectrogram program. The number of trill notes is strongly correlated with trill and total song length (Nelson & Poesel 2011). Mean distance from the speaker during each period was calculated by noting the males position continuously relative to ve categories: 0 2 m (median: 1 m), 24 m (3 m), 48 m (6 m), 8 16 m (12 m), and 16+ m (24 m). Males typically used a few perches, and horizontal distances between perches and speaker were measured after a trial with a marked rope. We ignored vertical distance to the speaker because most trees were <3 m tall. The number of seconds spent in each category was multiplied by the respective median distance, the products summed, and the sum divided by the
Ethology 118 (2012) 2432 2011 Blackwell Verlag GmbH

The composite response measure PC1 was strongly correlated with the four approach distance variables, and negatively correlated with the number of songs in the post-playback period (Table 1). PC2 was positively correlated with the number of trill notes in playback and post-playback songs. Response to playback, as measured by PC1, differed signicantly between the increasing trill length and decreasing trill length series of songs (Fig. 2a; Wilcoxon matched-pairs signed-ranks test T = 9, n = 12, p = 0.016). As measured by PC2, birds sang shorter trills in response to the increasing than decreasing trill length stimulus (Fig. 2a, c; T = 7, n = 12, p = 0.009). The signicant difference in PC1 appeared to come about because males approached somewhat closer at the end of the playback period and remained near the speaker during the post-trial period in response to the increasing trill length stimulus (Fig. 2b). Males approached both playback series gradually: they were further away in the rst 40 s of playback irrespective of whether the songs broadcast then contained long or short trills than they were in the last 40 s of playback (Fig. 2b). Males decreased their trill length during playback in response to both stimuli from a median of 3.9 trill notes in spontaneous singing to 1.9 notes during playback, but the decrease was signicant only in response to the increasing trill length stimulus (Fig. 2c: T = 0, n = 9,
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Song Length Variation in Sparrows

D. A. Nelson & A. Poesel

Table 1: Correlations between the rst two principal components and the eight original response variables measured in Experiments 1 and 2
Experiment 1 PC 1 Variance explained Playback closest approach Mean distance post-playback Mean distance playback Post-playback closest approach Post-playback song rate Playback song rate Playback trill length Post-playback trill length 46% 0.90 0.88 0.87 0.81 )0.71 0.31 )0.01 )0.03 PC 2 23% 0.07 0.05 0.03 0.35 0.38 0.29 0.89 0.82 Experiment 2 PC1 40% 0.68 0.69 0.89 0.77 )0.63 0.22 0.49 0.44 PC2 17% )0.22 )0.39 )0.03 )0.22 0.10 )0.02 0.73 0.73

(a)

Response difference stronger to increasing length

(b) 25
10 20 8

(c)
6

Songs/min

Meters

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6 4

10

1 5

2 2
PC 1 PC 2

0
S L E E L S Mean distance Mean distance

Pre play post

Pre play post

Playback

Post-playback

Trial period

Fig. 2: Experiment 1: Responses of male white-crowned sparrows to series of songs of either increasing or decreasing trill length. (a) Difference in the PC scores [calculated from eight behaviour measurements (Table 1)] between the increasing and decreasing trill length series calculated within males. Negative differences correspond to shorter mean approach distances, and higher song rate (PC1) and shorter trill lengths (PC2) to the increasing trill length series relative to the decreasing trill length series. (b) Response variables (mean distance from loudspeaker during each of the 40-s-long intervals during playback (S = short trill, L = long trill, E = elongated trill); and mean approach distance and mean song rate during post-playback) measured in each trial. Close approach distance during and after playback, and song rate during playback are not shown. Hatched bars are responses to increasing trill length, open bars are responses to decreasing trill length. (c) Mean trill length during the pre-trial, playback and post-playback periods. The lower and upper edges of the boxes represent the rst and third quartiles, the horizontal line within each box represents the median. The vertical lines (whiskers) include the range of values within 1.5 times the inter-quartile range.

p = 0.004; decreasing length: T = 10, n = 9, p = 0.16).

Experiment 2: Long Loud vs. Short Soft Songs

The pattern of correlations between response variables and PCs was similar to that found in Experiment 1 (Table 1), except that the strength of correlations was somewhat weaker, so that only 57% of variance in the original eight variables was accounted for by the rst two PCs. After being lured
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in with playback of loud long songs, males gave signicantly different responses to long loud and short soft songs. As measured by PC1, they remained closer to the speaker in response to playback of long loud songs than to playback of short soft songs (Fig 3a; Wilcoxon matched-pairs signed-ranks test, T = 5, n = 11; p = 0.01). Trill length responses represented by PC2 did not differ between treatments (T = 193, n = 11; p = 0.24; Fig. 3a). Males decreased the trill length in their songs relative to the pre-trial trill length in response to both stimuli (long loud:
Ethology 118 (2012) 2432 2011 Blackwell Verlag GmbH

D. A. Nelson & A. Poesel

Song Length Variation in Sparrows

(a)

(b)

(c)

Response difference (long-short trill) stronger to long, lloud

*
1

25

6 5

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15 0

Meters

Trill length (notes)

Song rate

4 3 2 1 0
Pre play post Pre play post

10

1 5 2

2
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Mean Close distance Playback Mean distance

Post-playback

Fig. 3: Experiment 2: Responses of male white-crowned sparrows to either long loud songs or short soft songs. (a) Difference in the PC scores between the response to long loud and short soft songs calculated within males. Negative differences correspond to closer approach and higher song rate (PC 1) and shorter trill lengths (PC2) to long loud songs than to short soft songs. (b) Four of eight response variables (mean distance and close approach distance during playback, and mean approach distance and mean song rate during post-playback) measured in each trial. (c) Mean trill length during the pre-trial, playback and post-playback periods. Hatched bars are responses to long loud songs, open bars are to short soft songs. Box plots as described in Fig. 2.

T = 3, n = 9, p = 0.02; short soft: T = 0, n = 9, p = 0.008; Fig. 3c), and most birds approached within 4 m during playback, indicating they could localize and hear the short soft songs (Fig. 3b). Discussion The results of two playback experiments show that male white-crowned sparrows respond differently to the long songs typical of solo spontaneous singing and the short songs common in close malemale interactions. The experimental results complement observational evidence (Nelson & Poesel 2011) that changes in song length are an aggressive signal in malemale interactions in the Puget Sound whitecrowned sparrow. In our playback tests, males gave weaker responses to the signal associated with an increased likelihood of attack (decreasing length series in Experiment 1, short soft songs in Experiment 2). In both the experiments, they approached less closely, and in experiment 1, males also sang longer trills in response to the decreasing song length series. In white-crowned sparrows, a long trill is less predictive of aggression than is a shorter trill (Nelson & Poesel 2011). It is difcult to attach a functional interpretation to these response patterns. It could be argued that our results show that males refrain from

approaching a highly threatening signal. Searcy et al. (2000; Searcy & Beecher 2009) have argued that response strength to territorial playback in male birds is an imperfect guide to whether a song is threatening or not, as some species respond strongly to songs judged to be aggressive, while other species show the opposite response. The response to playback is not sufcient by itself to show that a signal is aggressive, because we cannot predict a priori which way a given species will respond. Searcy et al. suggest that playback experiments test whether birds can distinguish signals, but we should not assume that strongly aggressive signals necessarily evoke strong responses, or vice versa. A comparison of playback studies in other species of song birds that vary song length in agonistic contexts illustrates the difculty in correlating response strength with inferred aggressive signal content. At least 17 species of song birds vary the length of their songs across contexts and have been tested in playback experiments with short and long songs. If there is a predictable relationship between message content and response, then songs judged to be aggressive based on the context they occur in should elicit similar responses (either relatively stronger or weaker compared to controls) across species. Contrary to expectation, it appears that song length is a better predictor of the strength of response to play29

Ethology 118 (2012) 2432 2011 Blackwell Verlag GmbH

Song Length Variation in Sparrows

D. A. Nelson & A. Poesel

back than is the contextual usage of the song type or variant. First, there is no consistent association between context and variation in song length in this set of species: in ve of the nine species for which observational data are available, shorter songs are given in malemale agonistic interactions (Table 2). In 12 species tested with playback, including all ve species in which short songs are used in agonistic contexts, males approach playback of long songs more closely than short songs (Table 2). No species shows the opposite pattern of approaching closer to short songs, and ve species approach similarly to long and short songs. Thus, long songs elicit stronger responses regardless of whether long songs are preferentially used in agonistic contexts. In addition to containing information about motivational tendencies, bird song also conveys information about the species, dialect, and individual identities of the singer, and potentially also information about the singers quality (Gil & Gahr 2002). It is well known that males give stronger responses to songs of their own species and local dialect (Becker

1982). It may be that long songs are more likely to contain the cues to species and dialect identities, thereby evoking strong recognition responses. In the white-crowned sparrow, the terminal trill presented alone elicits stronger responses from males than does any other phrase presented alone (Soha & Whaling 2002), and the trill encodes information about dialect identity (Nelson & Soha 2004). Long song may also be a general cue to male quality, as has been suggested in some species (great tit, Parus major (Lambrechts & Dhondt 1987) and blue tit, Cyanistes caeruleus (Kempenaers et al. 1997; Poesel et al. 2001). Experimental manipulation of a simple parameter like song length may simultaneously alter the cues to several types of information and complicate efforts to infer which cue(s) inuence response strength. In both of our experiments, males sang signicantly shorter trills in response to playback than they did in undisturbed singing prior to playback. The trills sung by subjects during these simulated territorial intrusions are similar in length to those observed in natural territorial interactions when males are within 10 m of

Table 2: Species of birds tested for response to playback of long and short versions of species-specic song
Species Willow warbler, Phylloscopus trochilus Great reed warbler, Acrocephalus arundinaceus Redwing, Turdus iliacus White-throated sparrow, Zonotrichia albicollis White-crowned sparrow, Zonotrichia leucophrys Coal tit, Periparus ater Field sparrow, Spizella pusilla Indigo bunting, Passerina cyanea Yellow-headed blackbird, Xanthocephalus xanthocephalus Bonellis Warbler, Phylloscopus bonelli Whitethroat, Sylvia communis Golden-winged Warbler, Vermivora chysoptera Chafnch, Fringilla coelebs Marsh tit, Poecile palustris Aquatic warbler, Acrocephalus paludicola Great tit, Parus major Goldcrest, Regulus regulus Firecrest, Regulus ignicapillus Chiffchaff, Phylloscopus collybita Context S S S S S in in in in in ## ## ## ## ## Playback Appr Appr Appr Appr Appr dist dist dist dist dist L L L L L < < < < < S S S S S Source Jarvi et al. (1980) Catchpole (1983) Lampe (1991) Falls (1969) Nelson & Poesel (2011), this study Adhikerana & Slater (1993) Nelson & Croner (1991) Shiovitz (1975) Cosens & Falls (1984) Bremond (1976) Balsby & Dabelsteen (2001) Ficken & Ficken (1973) Leitao & Riebel (2003) Romanowski (1979) Catchpole & Leisler (1989) McGregor & Horn (1992) Becker (1976) Becker (1976) Becker (1982)

L in ## L, soft in ## L, soft in ## L in ##

Increase song rate and switching to S songs Appr dist L < S LS Sing L to playback L < Sa Appr dist L < S Appr dist L < S Appr dist L < S Appr dist L < S Appr dist L < S; Sing S to playback L = Sa; Match length Appr dist L = S Appr dist L = S Appr dist L = S; Sing S to playback

Appr dist, Approach distance. Context column describes whether short and long songs are given in different contexts. Playback column describes responses to songs of different length. ## indicates close-range ## interaction. S = short song, L = long song. Appr dist L < S indicates that males came in closer to playback of L than S. Species are sorted by the context in which songs occur and by their response to playback. a Response measure based partly on approach distance.

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Ethology 118 (2012) 2432 2011 Blackwell Verlag GmbH

D. A. Nelson & A. Poesel

Song Length Variation in Sparrows

one another (1.9 and 1.8 trill notes in experiments 1 and 2 respectively vs. 1.6 notes in natural interactions (Nelson & Poesel 2011)). The change in trill length was transient, as trill length increased during the post-playback period to a length similar to that in the pre-trial period (Figs 2 and 3). These experimentally induced changes in trill length support the conclusion reached from observations that short trills occur in close malemale interactions, a context in which aggression by either territory resident or intruder is equally likely (Nelson & Poesel 2011). Our two experiments manipulated different aspects of song structure. In experiment 1, the series of songs with decreasing song length was designed to simulate an increased probability of escalation while the increasing trill length series simulated deescalation. We controlled both song amplitude and the total duration of stimulation across all 12 songs in each series because it might be expected that louder and longer stimuli would elicit stronger responses by virtue of being more easily detectable. This control is biologically reasonable in this case because short songs are not always soft in close malemale interactions (Nelson & Poesel 2011). Males did not tend to approach closely until the middle of the playback period, and if they heard elongated songs at the end of playback, they remained closer to the speaker during the subsequent 5-min-long post-playback period than if they heard short songs during the last 40 s of playback (Fig. 2c). In experiment 2, song amplitude and length co-varied, as they do in some natural male male interactions, and males approached closer to long loud songs. The low amplitude of the short songs in experiment 2 may have contributed to the response difference, as there is evidence from other species that low amplitude songs are associated with aggression (Searcy & Beecher 2009), but experiment 1 indicates that low amplitude is not necessary to elicit a response difference. Both experiments demonstrate that males respond differently to songs that occur in agonistic vs. non-agonistic contexts, the third criterion listed by Searcy & Beecher (2009) for establishing that a signal is aggressive. Acknowledgements We thank the Oregon State Parks Department, the staff at Bullards Beach State Park and the Port of Bandon for granting permission to work on their properties. We thank the NSF for nancial support (IBN04-15842). Procedures approved by Ohio State University IACUC Protocol 2000A005.
Ethology 118 (2012) 2432 2011 Blackwell Verlag GmbH

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Ethology 118 (2012) 2432 2011 Blackwell Verlag GmbH