Animal Behaviour 131 (2017) 23e32

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Animal Behaviour
journal homepage: www.elsevier.com/locate/anbehav

Accounting for syntax in analyses of countersinging reveals hidden

vocal dynamics in a songbird with a large repertoire
Richard W. Hedley a, *, Kaleda K. Denton a, Robert E. Weiss b
a
Department of Ecology and Evolutionary Biology, University of California Los Angeles, Los Angeles, CA, U.S.A.
b
Department of Biostatistics, UCLA Fielding School of Public Health, Los Angeles, CA, U.S.A.

a r t i c l e i n f o
Identifying the signalling strategies employed by animals during vocal interactions is a challenge,
Article history: especially for species with large vocal repertoires. We propose that efforts to study such vocal dynamics
Received 13 November 2016 can benet by integrating models of syntax into their analyses. In this study, we conducted playback
Initial acceptance 27 December 2016 experiments on Cassin's vireo, Vireo cassinii, to examine the role of syntax, and more specically, shared
Final acceptance 8 May 2017 syntactic patterns, in countersinging. We presented 11 males with song sequences ordered according to
population norms, and with sequences whose order deviated from population norms. We did not nd
MS. number: A16-00990R2 evidence that individuals markedly altered their responses based on the syntax of the playback, either in
their physical approach to the speaker or in the quantity of song they delivered in response. We did,
Keywords: however, nd evidence that syntax was important in governing their choice of phrase types in response
birdsong
to the playbacks. Subjects did not match the playback phrase types. Instead, they engaged in a vocal
Cassin's vireo
behaviour referred to as song advancing, where they responded to a stimulus phrase type by singing the
countersinging
syntax
phrase type that most often followed the stimulus in their own normal song sequences. When playback
vocal interaction sequences were ordered according to population norms, song advancing resulted in birds pre-empting
the upcoming playback phrase type or delivering another of the prior playback phrase types (i.e.
delayed matching) at higher rates than when playback sequences deviated from population norms. The
detection of song advancing was only possible with the explicit inclusion of syntax in our analysis,
suggesting that studies of the vocal interactions of species with repertoires of multiple vocalizations can
benet from consideration not only of a subject's repertoire, but also their syntax.
2017 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Countersinging interactions between neighbouring songbirds
facilitate the establishment and maintenance of territory bound-
aries (Yasukawa, 1981). In species that possess repertoires of
discrete song types, such interactions can involve complex dy-
namics and apparently sophisticated exchange of information
(Burt, Bard, Campbell, & Beecher, 2002; Burt, Campbell, & Beecher,
2001; Molles, 2006; Searcy & Beecher, 2009). Many species have
been shown to employ pattern-specic responses, meaning they
adjust their song pattern in response to the song of their rival (Todt
& Naguib, 2000). This can include subtly altering the acoustic
structure of an individual song type (Vehrencamp, Yantachka, Hall,
& de Kort, 2013), or altering their choice of song type altogether
(Falls, 1985; Krebs, Ashcroft, & Orsdol, 1981; Stoddard, Beecher,
Campbell, & Horning, 1992). Identifying and studying pattern-
* Correspondence: R. W. Hedley, 621 Charles E. Young Drive South, Room 3113,
Los Angeles, CA 90095, U.S.A.
E-mail address: rhedley@ucla.edu (R. W. Hedley).
specic responses is fundamental to our understanding of the
evolution of vocal complexity in birds. This is particularly true of
species with large song repertoires, in which the dynamics of vocal
interactions can be complex, and where the functional signicance
of large song repertoires remains a topic of debate (Byers &
Kroodsma, 2009).
A central challenge in the study of pattern-specic responses is
determining what factors lead a bird to deliver a particular song
type at a particular time. This aim makes clear that if we seek to
understand a singer's vocal behaviour, we should attempt to
identify any and all inuences on song choices, whether internal or
external. It is well established that many, and probably all, song-
birds abide by an internal syntax (Berwick, Okanoya, Beckers, &
Bolhuis, 2011), meaning they arrange repertoire elements into
nonrandom sequences characterized by frequent transitions be-
tween some pairs of elements and rare transitions between others.
In light of this, the answer to the question may sometimes be as
simple as to say that the bird delivered song type B at time t
because he delivered song type A at time t
1, and typically

http://dx.doi.org/10.1016/j.anbehav.2017.06.021
0003-3472/ 2017 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.
24 R. W. Hedley et al. / Animal Behaviour 131 (2017) 23e32
transitions from A to B. Although this study focuses on a species in
which syntax governs the order of song types (or phrase types)
within a bout, the same reasoning can be applied to syllable types
in species where syntax governs the ordering of syllables within a
song.
When a bird interacts with a rival, it may abide by its internal
syntax, or its song choices may be inuenced by external factors
such as the rival's song. In black-throated grey warblers, Setophaga
nigrescens, for instance, the repertoire is partitioned into type I and
II songs, and playback of any song lead males to respond with type I
songs (Morrison & Hardy, 1983). In this case, the warbler's song
sequence was disrupted by an external stimulus, leading the bird to
deliver a type I song regardless of its prior singing behaviour. A
similar behaviour is song type matching, where a bird repeats a
perceived song type shortly after it was delivered by a rival (Falls,
1985; Krebs et al., 1981; Stoddard et al., 1992). Song type match-
ing can be thought of as an interruption of a bird's normal song
sequence.
Given that syntax is likely to play such a large role in deter-
mining song type choices, and that pattern-specic responses
constitute disruptions to a bird's normal song sequences, it is sur-
prising that few studies have assessed the role of syntax in coun-
tersinging interactions. Those that have done so have occasionally
documented additional behaviours. For example, Verner (1975)
noted that western marsh wrens, Cistothorus palustris, deliver
their songs by cycling through their large song repertoires in nearly
stereotyped orders, and that the sequences employed by neigh-
bouring males are similar. When Verner (1975) broadcast a
sequence to a bird, the subject anticipated each song type to be
played next by the recorder (emphasis his, page 283). The birds, he
proposed, cued in on the previous playback song, which led them to
jump ahead in a shared sequence. Similar observations have been
made in wood thrush, Hylocichla mustelina (Whitney, 1985), and
nightingales, Luscinia megarhynchos (Todt, 1971). We henceforth
refer to this behaviour as song advancing, dened as responding to
a perceived song with the song type that typically follows it in one's
own preferred song sequences. Song advancing would seem to be
challenging, if not impossible, to detect and study without prior
knowledge of an individual bird's syntax. The integration of syn-
tactic models into analyses of vocal interactions is therefore war-
ranted, and may even be necessary if we wish to identify an
exhaustive list of the vocal behaviours of any species, not only
because syntax itself may guide a bird's song choices, but also
because some pattern-specic responses may be syntactic in na-
ture, as in the case of song advancing.
We sought to take such an approach using playback experi-
ments to examine the countersinging dynamics of Cassin's vireo,
Vireo cassinii. Males of this species possess repertoires of ~50 phrase
types (equivalent to song types in other species) and sing according
to a structured syntax, wherein the identity of upcoming phrase
types can be predicted with >55% accuracy if the previous phrase
type is known (Hedley, 2016a). Song sequences in this species show
two additional phenomena. First, phrase types are arranged into
clusters that consistently appear together in sequences (Fig. 1a).
These clusters have been shown to be stable over time with respect
to the phrase types contained therein, and the song sequences can
be well described using Markov models (Hedley, 2016a). Second,
cluster composition is often shared between individuals (Fig. 1b).
That is, not only do neighbours overlap in their song repertoires
(Hedley, 2016b), their syntax appears to be shared to an extent as
well.
Our experiments examined the role of shared song syntax in
countersinging interactions. Observations of natural counter-
singing interactions, such as the interaction depicted in Fig. 1c,
suggest that birds interact nonrandomly using phrase types that are
shared between the two participants. To test the role of syntax in
these interactions, we presented each bird with a playback trial
containing phrase types that normally occurred adjacently in se-
quences (typical trials) and one containing phrase types that rarely
occurred adjacently in sequences (atypical trials), and examined
the dynamics of each bird's response to the playbacks.
Our statistical approach to analyse vocal responses was moti-
vated by the work of Kroodsma (1975), who proposed that the
probability of matching in response to a phrase type depends on
four factors: (1) the frequency of occurrence of that phrase type
overall; (2) the transition probability from the bird's most recent
phrase type to the playback phrase type; (3) the amount of time
since the bird most recently sang the phrase type in question; and
(4) the vocal behaviour of other males within earshot. We
employed a model that incorporated properties of syntax from the
songs of the subjects, and thereby effectively controlled for (1), (2)
and (3). The inuence of other males (4) is precisely what we hope
to understand with playback experiments, and while it is possible
that songs from distant background males may affect the subject,
such effects are likely to be minimal relative to the effect of a
playback speaker positioned within the territory to simulate a
territorial intrusion. Although Kroodsma (1975) referred only to
song type matching, our model was exible enough to be applied to
song advancing as well. We used this approach to show that Cas-
sin's vireos engage in song advancing at levels exceeding the rate
expected by chance.
METHODS
Study Site and Species
Research was conducted in a 1 km2 valley of mixed con-
iferedeciduous forest near Volcano, CA, U.S.A. (UTM: 10 S 706584
4262742, datum WGS 84). Experiments were approved by the
Animal Research Committee at the University of California Los
Angeles (ARC number 2013-041-03A) and conducted under U.S.
Fish and Wildlife Service bird banding permit number 23809 and
California Scientic Collecting Permit number 12750. Prior to the
initiation of this study, 11 male Cassin's Vireos were colour banded
for individual identication. These 11 birds were all of the males
present at our study site in 2015.
Males in our study population possess repertoires composed of
an average of 51 phrase types (range 31e60), which they deliver in
structured sequences with immediate variety, meaning phrase
types are rarely repeated consecutively (Hedley, 2016b). Phrase
types are short (<0.7 s long), highly stereotyped, and can be readily
identied by a trained observer with >99% accuracy (Hedley,
2016a). Phrase types are widely shared in this population, such
that the repertoires of any two males tend to overlap by about 50%
(Hedley, 2016b). Details regarding the singing style of this species
are examined in more detail elsewhere (Hedley, 2016a, 2016b).
Nonexperimental Recording Sets
We constructed a set of recordings of each of the 11 individuals
made under nonexperimental conditions. In general, these re-
cordings were made when birds were singing individually on their
territory and not interacting closely with another bird. The
nonexperimental recording sets contained a total of 62 395 phrases
(mean 5672 phrases per bird, range 1498e14 101) and 300 re-
cordings (mean 27 recordings per bird, range 9e60). The
nonexperimental recording sets of ve individuals spanned 3 years,
those of two individuals spanned 2 years and those of four in-
dividuals included only recordings from 2015.
 R. W. Hedley et al. / Animal Behaviour 131 (2017) 23e32 25

40
30 (a) 30 (b) (c)

25 25 30
Phrase type ID

Unique to bird 2
20 Shared cluster 1 20
20
15 15
Shared cluster 2 Shared phrase types
10 10
10
5 5
Unique to bird 1
0 0 0
0 20 40 60 80 100 120 0 20 40 60 80 100 120 0 50 100 150
Position in sequence

Figure 1. Syntactic patterns are shared between individuals. (a, b) Two separate sequences of solo singing by two individuals (read from left to right). Phrase type ID (ordinate) was
assigned to each phrase type on the order of appearance in (a). Phrase types unique to (b) were assigned numbers >23 as a continuation of the numbering scheme in (a). Phrase
types are often delivered in clusters, appearing as groups of points on the plot. Two groups of phrase types that are clustered similarly in the repertoires of both birds are highlighted
with red and blue backgrounds, illustrating that syntactic patterns, in addition to phrase types, are often shared between individuals in this species. We presented individuals with
sets of phrase types that were often observed clustered together in song sequences in our population (typical trials), and sets of phrase types that were never clustered together
(atypical trials). In this example, a typical trial could be composed of phrase types in either shared cluster 1 or shared cluster 2, while an atypical trial might combine phrase types
from shared cluster 1, shared cluster 2 and other phrase types to form an abnormal sequence. (c) A countersinging interaction between two birds across a territorial boundary (bird
1 open black circles; bird 2 lled red circles). Shared phrase types are shown in the middle of the plot, with unshared phrase types above and below this region. When one bird
delivered a shared phrase type, the other bird often sang that same phrase type shortly thereafter, suggesting that the song choices of the two birds are not independent.

The nonexperimental recording sets served three purposes: to
determine the phrase types in each bird's repertoire; to parame-
trize a Markov model describing each bird's syntax; and to identify
transitions between phrase types common in the sequences of
multiple individuals, for the purposes of designing playback se-
quences. Importantly, previous work has not found evidence of
marked reorganization of syntax within or between years in a given
bird's song sequences (Hedley, 2016a), so a nonexperimental set of
recordings is likely to give a good representation of a bird's syntax.
Playback Design
Each colour-banded bird was subjected to two playback trials
containing ve phrase types per trial. To assemble the playback
sequences, we rst selected 55 phrase types from the population.
We assembled 11 typical trial sequences and 11 atypical trial se-
quences such that each phrase type appeared in one typical and one
atypical sequence. To determine which phrase types to combine
into typical and atypical trials, we overlaid Markov transition
matrices for all birds calculated from the nonexperimental
recording set to identify transitions that were both common in the
nonexperimental recording set as a whole, and delivered by mul-
tiple individuals. Typical trials were assembled by identifying sets
of ve phrase types that frequently occurred adjacently in se-
quences, and atypical trials were assembled by identifying sets of
ve phrase types that rarely or never occurred adjacently.
As conrmation that the typical and atypical sequences had the
desired properties, we again examined the nonexperimental
recording set sequences. Each set of ve phrase types could give
rise to 20 possible unique transitions (excluding repetitions), for a
total of 440 unique transitions across the 22 trial sequences. In the
nonexperimental recording set of 62 395 phrases, transitions be-
tween phrase types assembled into typical trials occurred 19 697
times, whereas transitions between phrase types in atypical trials
occurred just 220 times. Similarly, each possible unique transition
in the typical sequences occurred in the sequences of 2.37 1.61
(mean SD) individuals in the nonexperimental recording set,
while each possible transition in the atypical trials occurred in the
sequences of only 0.26 0.37 individuals.
When assigning a typical and atypical sequence to each indi-
vidual, we applied two additional constraints. First, birds were
never subjected to the same phrase type in the typical and atypical
trials, to avoid habituation that might occur with multiple pre-
sentations of the same stimulus. Second, each bird was presented
with some phrase types that were present in their repertoires and
some that were not, to simulate a real intruder and examine
whether presentation of an unshared phrase type may elicit its
delivery from a bird. The number of shared and unshared phrase
types presented (either three or four shared phrase types and one
or two unshared) was balanced across the typical and atypical trials
for each bird.
The ve phrase types chosen for a given trial were arranged into
a sequence of 25 phrases, where each type occurred ve times.
Within this sequence, phrase order was randomized in such a way
that consecutive phrases were never of the same type. Phrases were
delivered every 2 s, as is typical for this species, so the sequence
took 50 s to complete, and was followed by 10 s of silence. This
1 min segment repeated ve times, comprising a 5 min playback
period. Because of the differences in the phrase types contained in
each sequence, and the randomization of the sequence, no two
sequences in the experiments were alike.
We sourced phrase type exemplars from high-quality re-
cordings of 12 individuals in the same study area during 2013 and
2014, but the 10 phrase types presented to each bird need not have
originated from the same individual. Instead, phrase types were
combined into articial sequences to simulate an unfamiliar
intruder. Three individuals were unintentionally presented with
one to three phrase type exemplars recorded from themselves at an
earlier date. Since subjects may respond differently to their own
songs and those of a rival (Falls, 1985; Stoddard et al., 1992), we ran
the main analyses again without these individuals. The main con-
clusions did not change upon removal of these three individuals,
but we included them in the analysis presented here to avoid un-
necessarily reducing the sample size. A summary of the results
from analysis without these three individuals is provided in
Supplementary material 1.
Playback Field Methods
We conducted playbacks from 28 April to 31 May 2015 between
0700 and 1100 hours. An attempt was made to conduct both trials
during the same phase of nesting, either during territory
26 R. W. Hedley et al. / Animal Behaviour 131 (2017) 23e32
establishment prior to nesting (N 2 individuals) or during incu-
bation (N 9 individuals) when the male was off the nest. Ten
subjects received the two trials separated by 1e3 days, but one in-
dividual's nest failed between trials, so the second trial was post-
poned until incubation began on a subsequent nest 23 days later. We
placed a Jambox speaker (Jawbone, Inc., https://jawbone.com/)
about 1.8 m up in a tree within the bird's known territory, with ags
10 m on each side to aid in distance estimation. The speaker was
placed in the same location for both trials, and presentation order of
the two trials was randomized. Phrase amplitude was normalized
and broadcast at ~80 dB measured 1 m from the speaker.
A single observer conducted all trials. The observer stood ~20 m
from the speaker, recorded the subject's songs with a Sennheiser
MKH20-P48 microphone and Telinga parabolic reector onto a
Marantz PMD661 recording device, and dictated into the micro-
phone the bird's estimated distance from the speaker. The experi-
mental period lasted 12 min: the rst 2 min were passive
observation, the subsequent 5 min coincided with the broadcast of
the playback le, and the last 5 min comprised a second period of
passive observation. Two observers examined the spectrogram of
the resulting .wav le in the program Praat (Boersma & Weenink,
2014). The observers annotated to phrase type the phrases from
the speaker and subject by assigning each phrase type a two-letter
code (see Hedley, 2016b), and noted the subject's estimated dis-
tance from the speaker when each phrase was delivered. When the
observers differed in their annotation of a phrase type (<1% of
phrases), they discussed the discrepancy to arrive at a consensus.
Statistical Analyses
Initially, we examined whether seven different pattern-specic
responses occurred above chance levels in the vocal responses of
the subjects. These seven behaviours were as follows: (1) imme-
diate matching (IM), dened as repeating the most recent playback
phrase type; (2) delayed matching (DM), dened as repeating any
of the phrase types that had previously emanated from the speaker
during that trial; and (3e7) ve forms of song advancing (SA1 to
SA5). The rst form of song advancing (SA1) was dened as
responding to the most recent stimulus phrase type with the
phrase type that most commonly follows it in the bird's normal
syntax, as determined from that bird's nonexperimental recording
set. The second form was dened as responding to a stimulus with
its second most common successor in the subject's nonexperi-
mental recording set, and so on until the fth form of song
advancing, SA5, dened as responding to a stimulus with its fth
most common successor.
For each of the above behaviours, we employed a logistic
regression model that took into consideration each bird's known
syntax determined from the nonexperimental recording set. The
model incorporates Markov transition probabilities to estimate the
probability that the vocal behaviour in question would have
occurred by chance each time the subject sang a phrase. Chance
probabilities varied throughout the experiment as a function of the
most recent phrase type delivered by the bird, so the model
effectively controlled for the inuence of syntax on these
probabilities.
We considered each of the subjects' phrases during the 5 min
playback trials to be an independent Bernoulli trial, where the
outcome can be either 1 or 0 with some probability of each
outcome. These Bernoulli trials were modelled using a logistic
regression model
yi jmi  Bernoullimi (1)
logitmi logitpyi 1jxi
1 a0 bj (2)
In this model, yi designated whether phrase i from the bird did
(yi 1) or did not (yi 0) constitute the behaviour in question (i.e.
IM, DM or SA1-5, which were each analysed separately). Independent
of this, mi was the estimated probability of engaging in the behaviour
at phrase i, which itself depended on the values on the right side of
equation (2). The rst of these values, p(yi 1jxi-1), was the probability
of the behaviour occurring by chance at phrase i given the subject's
previous phrase type xi-1. This chance probability p(yi 1jxi-1) was
calculated using a Markov model, smoothed with Backoff smoothing
(Jurafsky & Martin, 2000) and Witten-Bell discounting (Witten & Bell,
1991), and parametrized from the subject's nonexperimental
recording set (see Supplementary material 1). The value a0 was an
intercept, and the values bj were random effects to account for the fact
that multiple responses were derived from each subject.
In the above model, the value of the intercept is of primary in-
terest: an intercept (a0) with posterior density signicantly greater
than zero indicates that the behaviour in question occurred more
often than expected by chance. The model can be thought of as
assessing whether or not syntax alone could account for the
number of occurrences of a behaviour, or whether the bird
appeared to intentionally deviate from its syntax to increase the
rate of the behaviour in question.
In the IM model and SA1 to SA5 models, the chance probability
p(yi 1jxi-1) was simply the probability of singing the single phrase
type that would constitute a match or advance at phrase i given the
bird's previous phrase type. In the DM model, however, the chance
probability was the sum of the probabilities of singing each of the
phrase types that would constitute a delayed match, as dened above.
After assessing which of the seven vocal behaviours occurred
above chance levels, we added additional predictors to the models
to assess whether the treatment type (typical or atypical) or the
subject's distance from the speaker affected the tendency to engage
in each behaviour. The addition of a treatment parameter tested
whether individuals altered their vocal behaviour in response to
the syntax of the playback. The addition of a distance parameter
tested whether birds altered their vocal behaviours based on their
proximity to the speaker, as might be expected if these vocal be-
haviours are associated with the conveyance of aggression or if they
reect a bird's motivational state. Treatment (TMT) was coded as
0 for all phrases in atypical trials, and 1 in typical trials. Distance (D)
was the bird's estimated distance from the speaker when each
phrase was delivered. The treatment and distance variables were
not collinear (R2 0.039).
Upon adding these predictors to the model, the updated model
took on the following structure:
yi jmi  Bernoullimi (3)
logitmi logitpyi 1jxi
1 Zi a bj (4)
Where Zi is a vector containing a 1 followed by the values of the
two additional predictor variables described above, and a is a
vector of the intercept (a0) followed by two coefcients (aTMT, aD).
In this model, the value of the intercept is of little interest, since it
represents the tendency to engage in song advancing when all
other covariates are zero. Instead, we are interested in the other
coefcients. A signicantly nonzero value of one of these co-
efcients represents a signicant effect of that predictor on the
tendency of the subjects to engage in song advancing. The magni-
tude of a coefcient represents the inuence of a one-unit change
in the independent variable on the probability of engaging in song
advancing on the logit scale, a nonlinear scale. On the logit scale, a
 R. W. Hedley et al. / Animal Behaviour 131 (2017) 23e32 27

difference of one is equivalent to an increase in probability from matching or song advancing relative to chance levels. We ran the
0.005 to 0.0125, from 0.05 to 0.135, or from 0.5 to 0.731. A more model for each of the seven behaviours (IM, DM, SA1-5), and results
extensive description of logistic regression and the logit scale is are shown in Table 1 and Fig. 2. Only SA1 occurred signicantly
given by Sokal and Rohlf (1995). more often than was expected by chance (P 0.0026), indicating
To identify the best t model to explain variation in each that birds often responded to perceived phrase types by singing the
behaviour, we also conducted model selection using deviance in- phrase type that would most commonly follow it in their own
formation criteria (DIC). DIC is an approach commonly employed in normal song sequences. SA1 remained signicant at a Bonferroni-
conjunction with Markov chain Monte Carlo methods corrected signicance threshold of a 0.05/7 0.007.
(Spiegelhalter, Best, Carlin, & van der Linde, 2002). It measures the
t of models with different numbers of parameters while penal- Effects of Treatment and Distance on Vocal Behaviours
izing more complex models. In our case, we compared models with
and without the treatment and distance coefcients. Lower DIC Next we sought to examine whether the treatment type or a
values indicated models that better t the data. subject's distance from the speaker inuenced the tendency to
The playback sequences contained shared and unshared phrase engage in song advancing (SA1). To do so, we added treatment type
types, but subjects never responded to a playback with an unshared and distance as predictors in the model. Results for the rst form of
phrase type, indicating that our prior assessment of which phrase song advancing (SA1) are shown in Table 2. Treatment type had a
types were in each bird's repertoires was accurate. As a result, our signicant negative effect on the tendency to engage in song
analysis only considered the subject able to engage in immediate advancing, indicating a stronger tendency to engage in song
matching or song advancing if (1) the most recent playback phrase advancing in atypical trials. The effect of distance was not signi-
type was shared and (2) the subject had not delivered any intervening cant. Model selection conrmed these results: the best model for
phrases since the most recent playback phrase. Delayed matching SA1 was a model including treatment, but not distance, as a pre-
was less constrained, since the bird could engage in delayed matching dictor. Model selection results for SA1 are shown in Table 3.
at any time as long as at least one shared phrase type had been
delivered by the playback. Immediate matching and song advancing Table 1
Analysis of the behaviours that occurred at levels greater than expected by chance in
models were therefore based on lower sample sizes (N 1125
the vocal responses of Cassin's vireos
phrases) than the delayed matching model (N 2438 phrases).
Analyses were conducted using a Bayesian framework where Vocal behaviour Coefcient (a0) SD P
coefcients were assigned uninformative priors (normal with Immediate matching (IM) 0.04 0.34 0.45
mean 0). Fixed effects were assigned a wide prior SD of 10. Delayed matching (DM) 0.11 0.18 0.27
Random effect SDs were assumed to have unknown variance, and Song advancing (SA1) 1.03 0.33 0.0026
Song advancing (SA2) 0.45 0.31 0.072
the precision (i.e. 1/(SD2)) was assigned a prior based on a gamma Song advancing (SA3) 0.29 0.31 0.16
distribution with shape and scale parameters of 1, per the recom- Song advancing (SA4)
0.20 0.33 0.27
mendations made by Dey, Ghosh, and Mallick (2000). Analysis was Song advancing (SA5)
0.43 0.38 0.12
carried out via 25 000 Markov chain Monte Carlo simulations in A signicantly positive coefcient estimate represents a behaviour that occurred
Jags (Plummer, 2003), version 3.3.0, implemented in R (R Core more often than expected by chance. P values represent the proportion of the
Team, 2016) with the package R2Jags (Su & Yajima, 2015). All posterior density that was smaller than zero (for positive coefcient estimates) or
data for this experiment is available on Figshare (http://dx.doi.org/ larger than zero (for negative coefcient estimates).

10.6084/m9.gshare.3410110).

RESULTS
Expected
Strength of Response to Playback Treatments
Observed
***
0.1
Individuals exhibited a strong physical response, approaching
towards the speaker in every trial (Supplementary material 2,
Proportion of phrases

Fig. S2), and responded vocally with an average of 111 phrases

(range 12e145) per 5 min playback interval. There were no clear
differences in the physical response to the trials with typical phrase
type sequencing patterns compared to those with atypical
sequencing. The minimum distance from the subject to the speaker
did not differ between the two treatments (Wilcoxon signed-ranks 0.05
test: T 7, N 11, P 1), nor did the amount of time spent within
10 m of the speaker (T 35, N 11, P 0.16), the latency to
approach within 10 m of the speaker (T 13, N 7, P 0.94), or the
number of changes in position, either towards or away from the
speaker (T 24.5, N 11, P 0.80).
In addition, we found no differences in the total number of
phrases (T 40.5, N 11, P 0.53) or in the number of different 0
phrase types (T 33, N 11, P 0.61) delivered in response to the IM DM SA1 SA2 SA3 SA4 SA5
two treatments during the 5 min playback periods. Vocal behaviour

Matching or Advancing Relative to Chance Levels
When we ran our model without additional predictor variables,
the intercept represented the tendency for the birds to engage in
Figure 2. Summary of song type matching and song advancing in response to play-
back. Expected and observed rates of immediate matching (IM), delayed matching
(DM) and song advancing (SA1-5) are shown. Expected values were calculated using the
P
formula for the Poisson binomial distribution: Ex ni1 pi (see Supplementary
material 1; Johnson, Kemp, & Kotz, 2005). ***P 0.003.
28 R. W. Hedley et al. / Animal Behaviour 131 (2017) 23e32

Table 2 expected by chance. These values ranged from 0.75 to 3.38. Inter-
Factors affecting the tendency to engage in song advancing estingly, the two birds that engaged in song advancing at the
Parameter Coefcient SD P highest rates were the two that were exposed to playbacks during
Intercept (a0) 1.46 0.42 0.0012
territory establishment. These two individuals engaged in song
Treatment (aTMT)
0.62 0.30 0.021 advancing at 3.38 and 2.41 times chance levels, while the remain-
Distance (aD)
0.0097 0.024 0.34 ing nine birds that were tested later in the nesting cycle engaged in
Signicantly nonzero coefcient estimates represent factors that affected the ten- song advancing between 0.75 and 2.13 times chance levels.
dency for the subjects to engage in song advancing, and the sign (positive or Although suggestive of an inuence of breeding stage on the ten-
negative) indicates whether this was a positive or negative effect. P values represent dency to engage in song advancing, the small sample sizes preclude
the proportion of the posterior density that was smaller than zero (for positive
meaningful statistical analysis of this trend, so we leave this to
coefcient estimates) or larger than zero (for negative coefcient estimates).
future studies.
We also ran similar analyses for the six other vocal behaviours
The negative treatment coefcient for SA1 indicates that there (IM, DM, SA2-5) by adding treatment and distance as predictors of
was a stronger tendency to engage in song advancing in the atypical those behaviours. Results are provided in Supplementary material
trial than in the typical trial. Closer inspection, however, reveals 1. Neither treatment nor distance showed a signicant effect on
that this was driven not by higher absolute rates of song advancing any of these behaviours, with the exception of the DM model,
in the atypical trials, but by lower expected rates of song advancing which showed a signicant positive effect of treatment. Delayed
in the atypical trials; song advancing occurred at approximately matching occurred at a rate of 10.4% in atypical trials and 14.2% in
equal rates between the two treatments (Fig. 3). It may not be the typical trials. Model selection gave broadly similar results: the top
case that the treatment itself was salient to the birds. An alternative model for the other behaviours was the model containing only the
interpretation is that subjects simply tended to engage in song intercept, again with the exception of DM, where the top model
advancing about 10% of the time, regardless of the treatment and included the intercept and the treatment parameter. Results of
their prior singing behaviour. model selection for these six behaviours are provided in
At the individual level, 10 of 11 subjects engaged in song Supplementary material 1.
advancing (SA1) above chance levels. We quantied the overall Since DM did not occur above chance levels overall, we sus-
strength of the tendency to engage in song advancing for each in- pected that the positive treatment coefcient in the DM might be a
dividual by dividing the observed rate of song advancing by the rate by-product of song advancing, rather than an active behaviour on
the part of the birds. In typical trials, the similarity between the
subject's own syntax and the syntax of the playback would mean
that advancing in response to a playback phrase would likely result
Table 3 in the delivery of another playback phrase type (i.e. a delayed
Results of model selection to examine the best model for the rst form of song
match). This would not be expected to be the case in atypical trials,
advancing (SA1)
because the atypical trials were composed of phrase types that do
Parameters DIC DDIC not normally occur near each other in sequences within this
Intercepttreatment 615.74 0 population.
Intercept 617.91 2.17 To investigate whether song advancing could explain the dif-
Intercepttreatmentdistance 618.26 2.52
ferences in delayed matching rates between typical and atypical
Interceptdistance 620.07 4.34
treatments, we calculated rates of delayed matching when birds
Lower deviance information criteria (DIC) values indicate a better model t, while engaged in song advancing, and when they did not, in typical and
accounting for the complexity of the model. DDIC reects the t of the model
atypical trials. Results are illustrated in Fig. 4a. Rates of delayed
relative to the top model.
matching were signicantly different between these four contexts
(Fisher's exact test: P < 0.001). Post hoc comparisons revealed that
song advancing in typical trials led to greater rates of delayed
0.12 matching than in the other three contexts (P < 0.001 for all com-
Expected
Observed parisons). Song advancing in atypical trials had the opposite effect,
by decreasing the rate of delayed matching relative to the other
0.1 three contexts (P < 0.005 for all comparisons). When birds did not
engage in song advancing, delayed matching occurred at rates that
Rate of song advancing

did not signicantly differ between typical and atypical trials

0.08 (P 0.78). All signicant comparisons remained signicant at a
Bonferroni-corrected signicance threshold of a 0.05/6 0.008.
These results support the idea that the difference in delayed
0.06 matching rates between typical and atypical trials can be explained
as a by-product of song advancing.

0.04 Pre-emption of Upcoming Playback Phrases

Given that song advancing involves responding with the phrase

type most likely to follow the stimulus phrase in a preferred
0.02
sequence, a reasonable hypothesis regarding this behaviour is that
it might lead to the pre-emption of a rival's upcoming phrase type
that has yet to be delivered. Pre-emption of a rival, however, would
0 only be expected to be successful if the two participants deliver
Atypical trials Typical trials
their songs in similar orders. To test this, we investigated the rate of
Figure 3. Rates of song advancing in typical and atypical trials. these pre-emptive matches of the upcoming playback phrase type
 R. W. Hedley et al. / Animal Behaviour 131 (2017) 23e32 29

(a) (b)
0.8
a b c c 0.15 a b b b
37/55 7/52

Rate of pre-emptive matching

Rate of delayed matching
0.6

0.1

0.4

0.05
0.2 16/482
62/488 71/528 14/521

0/54 0/54
0 0
Song Song Other Other Song Song Other Other
advancing, advancing, responses, responses, advancing, advancing, responses, responses,
typical atypical typical atypical typical atypical typical atypical
trials trials trials trials trials trials trials trials

Figure 4. Rates of (a) delayed matching and (b) pre-emptive matching of the upcoming playback phrase when the subjects engaged in song advancing (SA1) versus when they did
not (other responses) in typical and atypical trials. Numbers above bars indicate sample sizes of matches (delayed or pre-emptive) divided by the number of instances of the
behaviour (song advancing or other responses) in each trial type (typical versus atypical). Letters above bars indicate signicant differences, assessed with Fisher's exact tests. Bars
with the same letter were not signicantly different from one another (all pairwise P > 0.35); bars with different letters were signicantly different (all pairwise P  0.005).

when the subjects engaged in song advancing (SA1) versus when increase as the birds become more familiar with the ordering of
they did not in typical and atypical trials. Each phrase was classied phrase types in the playback sequences, which in our experiments
as either an instance of song advancing or not. Not mutually contained ve repeated 1 min segments. We used logistic regres-
exclusively, each phrase was also classied as either a pre-emptive sion to test whether the rate of pre-emptive matching changed as a
match, if it was the same phrase type as the subsequent playback function of the time (in seconds) of each response phrase from the
phrase type, or not. start of the playback. The relationship was not signicant
Pre-emptive matches were observed 37 times throughout the (btime SE
0.0032 0.0022, P 0.141), indicating that the rate
experiments, and were observed at least one time in the responses of pre-emptive matching did not change markedly as the experi-
of eight of the 11 subjects. Fig. 4b depicts the rate of pre-emption ments progressed.
when birds engaged in song advancing, and when they did not,
in typical and atypical trials. Rates of pre-emptive matching were DISCUSSION
signicantly different between these four contexts (Fisher's exact
test: P 0.003). Post hoc comparisons revealed that song Responses to Typical and Atypical Playback Sequences
advancing in typical trials led to greater rates of pre-emptive
matching than in the other three contexts (P  0.005 for all com- Cassin's vireos did not alter their response to playbacks on the
parisons), but that the other three contexts did not differ signi- basis of the syntactic characteristics of the playback sequence. In
cantly from one another (P > 0.35 for all comparisons). All terms of their physical response, various metrics intended to
signicant comparisons remained signicant at a Bonferroni- measure the strength of their approach did not differ in response to
corrected signicance threshold of a 0.05/6 0.008. Thus, song the typical and atypical playback sequences. Similarly, the quantity
advancing sometimes led birds to sing the upcoming playback of their song output or the number of phrase types delivered in
phrase type before it had emanated from the speaker, but only response did not differ between treatments. Overall, song se-
when the playback phrase types were ordered according to locally quences arranged according to population norms and those with
typical syntax. When playback phrase types were arranged ac- abnormal ordering both appear to be perceived as a strong terri-
cording to locally atypical sequencing patterns, song advancing still torial intrusion and do not appear to evoke markedly different re-
occurred at similar rates, but did not affect the tendency for sub- sponses in this species.
jects to pre-empt the playback phrase type. In previous experiments that have examined responses to
The similar patterns regarding delayed and pre-emptive altered song sequences, results have been mixed. Skylarks, Alauda
matching shown in Fig. 4a and b can be explained based on the arvensis, for instance, exhibited heightened levels of aggression in
structure of typical and atypical playback sequences. Since play- response to altered song sequences (Briefer, Rybak, & Aubin, 2013).
backs were composed of repeated clusters of ve phrase types, an California thrashers, Toxostoma redivivum, and Eurasian wrens,
upcoming phrase type was likely to have already occurred earlier in Troglodytes troglodytes, showed the opposite effect, responding less
the playback. Therefore, instances of pre-emptive matching, in our strongly to altered sequences than to unaltered ones (Holland,
experiments, constituted a subset of delayed matching outcomes. Dabelsteen, & Paris, 2000; Taylor, Brumley, Hedley, & Cody, 2017).
In much the same vein as we proposed for delayed matching, In other species, including European robins, Erithacus rubecula
these results suggest the possibility that pre-emptive matching (Bremond, 1968) and indigo buntings, Passerina cyanea (Emlen,
may result as a by-product of the tendency for birds in this popu- 1972), alterations to song order did not appear to affect the
lation to deliver their songs in similar orders and to engage in song response of receivers.
advancing. Alternatively, pre-emptive matching may be an active It appears doubtful, given these disparate results from different
process, perhaps facilitated through familiarity with another bird's species, that syntax has the same function in all species. The precise
preferred song sequences. In that case, it might be expected to information encoded by syntax is an active area of ongoing study. It
30 R. W. Hedley et al. / Animal Behaviour 131 (2017) 23e32
is worth noting that in Cassin's vireos, syntax governs the order of
phrases (i.e. songs) within an extended bout of song, where suc-
cessive phrases are typically separated by two or more seconds.
From a theoretical standpoint, syntactic patterns spanning such
long gaps would seem poorly suited to encode information (e.g.
individual identity or species identity), since a listener would have
to wait several seconds to hear even a few consecutive phrases. In
contrast, in the species mentioned above where differential re-
sponses to altered and unaltered song sequences have been
observed, syntax governs transitions between elements within a
continuous burst of song, where elements are separated by gaps on
the order of a fraction of a second in duration. In those species,
differences between altered and unaltered song sequences would
likely be apparent within a second or two, which may facilitate
discrimination based on sequence properties. Whether the rate of
delivery of song elements is related in a systematic way to the
function of syntax is speculative, at present, and awaits additional
exploration in a wider suite of species.
Countersinging Dynamics in Cassin's Vireo
Although subjects did not give markedly different responses to
the two playback treatments, we nevertheless found evidence of a
role for the subject's own syntax in guiding vocal responses,
through the vocal behaviour called song advancing. Males engaged
in song advancing by frequently responding to a playback phrase
with the phrase type that would most commonly follow the
stimulus in the subject's normal song sequences (Fig. 2). Engaging
in song advancing resulted in increased rates of pre-emptive
matches, where the subject delivered the next playback phrase
type before it was broadcast by the speaker, and delayed matches,
where the subject delivered a playback phrase type that had come
from the speaker previously. Pre-emptive matching and delayed
matching were facilitated by shared syntactic patterns present in
the playback, and song advancing did not lead to pre-emptive
matches or delayed matches when the playback contained se-
quences whose syntax was atypical for the population under study
(Fig. 4). These results hint towards a central role for song syntax in
determining song choices during countersinging in this species and
help to shed light on the dynamics depicted in Fig. 1c, dynamics
that we have observed frequently in vocal interactions in this
species. While the patterns appear to show that one bird delivers a
shared phrase type that is soon copied by the other bird, our results
now lead us to suspect that the dynamics instead involve one bird
engaging in song advancing and pre-empting the song of the other
by jumping ahead in a sequence that is common to both
participants.
A conclusive appraisal of this view of countersinging dynamics
in Cassin's vireo will require analyses focusing on natural in-
teractions, to complement our experimental results. Notably, some
authors have proposed that singing behaviours comprise a graded
signalling system (Burt et al., 2001), where low or moderate levels
of threat might be met with song type matching, and higher levels
of threat with more aggressive behaviours such as soft song
(Ballentine, Searcy, & Nowicki, 2008; Searcy, Anderson, & Nowicki,
2006; Searcy & Beecher, 2009). If this paradigm can be applied to
Cassin's vireos, it is possible that our playback experiments, which
simulated a strong territorial intrusion, may have elicited only a
subset of the full suite of singing strategies employed by this spe-
cies. Additional behaviours, such as immediate matching, may
become apparent as analyses are expanded to a greater variety of
contexts.
Another question that might be claried through investigations
of natural interactions is whether pre-emptive matching is an
active process, rather than a by-product of song advancing. We did
not nd evidence for increased rates of pre-emptive matching to-
wards the end of playbacks, as might be expected if familiarity with
a sequence facilitated pre-emptive matching. While our results are
consistent with the view that pre-emptive matching is a by-
product of song advancing, it is also possible that the 5 min play-
back period was too short for familiarity with a sequence to
develop. This would not be problematic in natural vocal in-
teractions, since neighbours repeatedly interact with each other
over the course of days and weeks during the breeding season. If
pre-emptive matching is an active behaviour, birds might alter their
choice of phrase type based on prior familiarity with their rival's
syntax, and their choices may vary as a function of the identity of
the bird with which they are interacting. If it is instead a by-
product, birds would only be expected to select their response
with reference to their own syntax, rather than their rival's syntax,
through the process of song advancing. Research is currently
ongoing to examine whether our experimental results and in-
terpretations generalize to natural countersinging interactions.
Descriptions of song advancing in at least three bird species in
the laboratory (nightingale: Todt, 1971; marsh wren: Kroodsma,
1979; wood thrush: Whitney, 1985) and one species in the wild
(Cassin's vireo, this study) suggest that this behaviour is more
widespread than currently appreciated. Despite evidence of this
behaviour dating back over 40 years, few studies have focused on
understanding its role in communication or documenting it in
additional species. In stark contrast, vocal matching behaviours
have been documented in dozens of species as distantly related as
songbirds, parrots and cetaceans over the same time frame
(reviewed in King & Mcgregor, 2016). The critical difference be-
tween these two behaviours is that matching involves a stimulus
and response that share obvious acoustic structure, while song
advancing involves a stimulus and response that can differ arbi-
trarily in their acoustic structure. This difference leads to a vast
divergence in the detectability of these two behaviours. For
example, vocal matching is often apparent to the human ear
(Kroodsma, 1971) and readily discernible on a spectrogram
(Stoddard et al., 1992), while song advancing would seemingly be
impossible to detect by ear, by spectrogram, or by any method that
does not consider a subject's vocal responses in light of their syntax.
We propose, on the basis of our ndings, that animal communi-
cation research can benet greatly by incorporating and studying
syntax, and that a full understanding of the complexities and nu-
ances of the vocal interactions of animals may require such an
approach.
Potential Functions of Song Advancing
A signicant question not addressed by our results is the
question of the role of song advancing in communication. One
possibility is that song advancing may convey aggression from
sender to receiver. Searcy and Beecher (2009) outlined criteria for
identifying aggressive signals in avian communication. The rst of
these is to demonstrate that the behaviour occurs more often than
expected by chance, a criterion that is supported in the case of song
advancing in Cassin's vireos. Another criterion is that the behaviour
should be more prevalent during aggressive contexts. Previous
authors have argued that aggressive encounters are expected to be
more frequent and intense prior to the initiation of nesting. Higher
rates of song type matching early in the season has been put forth
as evidence that matching acts as a signal of aggression (Beecher,
Campbell, Burt, Hill, & Nordby, 2000). We noted that the birds
tested during the territory establishment phase in our experiment
showed higher rates of song advancing than those tested during
incubation. Although this is suggestive of a trend, and worthy of
further investigation, our sample of just two individuals in the pre-
 R. W. Hedley et al. / Animal Behaviour 131 (2017) 23e32
nesting phase is not sufcient to draw a conclusion. We also did not
observe a clear relationship between a bird's distance from the
speaker and the tendency to engage in song advancing, but again
our modest sample of just 11 individuals makes interpretation of
negative results challenging.
Perhaps the most critical consideration if a signal is to be of
biological importance is that it be salient to other birds, the third
criterion of Searcy and Beecher (2009). Our results shed little light
on this point. As mentioned above, the behaviour is difcult to
detect, so salience to a receiver cannot be assumed. It seems
plausible, however, that neighbouring birds may be familiar with
each other's syntax, either because their neighbour's syntax is
similar to their own, or because they have interacted with that
neighbour previously. The tendency for song advancing to result in
pre-emption of a rival's song may be another means by which it can
be detected by a receiver. Assessing receiver responses to song
advancing would require interactive playbacks and would be
another productive avenue of inquiry into this behaviour. On the
whole, our results neither fully satisfy nor contradict the criteria for
aggressive signals outlined by Searcy and Beecher (2009), but op-
portunities for follow-up experiments are plentiful.
Another possible function of song advancing is that it may
encode dominance relationships. Following up on the eld studies
of Kroodsma (1979), Verner (1975) demonstrated that, among two
captive marsh wrens, the dominant bird sought to maintain a
leading position in the delivery of shared song sequences, while the
subordinate bird trailed. Song advancing might achieve a similar
end, by allowing a bird to jump ahead of his rival in a shared
sequence, thereby establishing a leading role in a countersinging
interaction. The trailing bird may match the leader simply by
continuing to sing its normal song sequence. Whether this behav-
iour is actually related to dominance, as proposed by Kroodsma
(1979), remains an open question.
An alternative is that such vocal interactions are performances
whose evolution is mediated by the responses of eavesdropping
conspecics (Logue & Forstmeier, 2008). Results of two-speaker
playback experiments on nightingales showed that leaders in
matched countersinging interactions elicit more attention than
followers from male and female eavesdroppers alike (Bartsch,
Wenchel, Kaiser, & Kipper, 2014). If these results extend to Cas-
sin's vireos, asymmetrical costs and benets of song advancing may
be imposed, not by the participants themselves, but by eaves-
droppers. The extent to which leaderefollower dynamics apply to
the specic case of song advancing is one of many questions that
remains to be addressed in future studies.
AUTHOR CONTRIBUTIONS
R.W.H. designed and conducted experiments, processed and
analysed the data and wrote the manuscript. K.K.D. assisted with
postprocessing and annotation of data and helped draft the
manuscript. R.E.W. assisted with statistical analysis.
COMPETING INTERESTS
We have no competing interests.
Acknowledgments
This work was funded by National Science Foundation Award
Number 1125423 and the Ralph W. Schreiber Ornithology Research
Award from the Los Angeles Audubon Society. We thank Charles
31
Taylor and David Logue for feedback on the manuscript and Martin
Cody for input on experimental design.
Supplementary material
Supplementary material associated with this article is available,
in the online version, at http://dx.doi.org/10.1016/j.anbehav.2017.
06.021.
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