Rooks (Corvus frugilegus) spontaneously attempt to vocally entrain to rhythmic stimuli

Rooks (Corvus frugilegus) spontaneously attempt to vocally entrain to rhythmic stimuli | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Rooks (Corvus frugilegus) spontaneously attempt to vocally entrain to rhythmic stimuli Killian Martin, Maëlan Tomasek, Agnès Hivet, Andrea Ravignani, and 2 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6785900/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 26 Jan, 2026 Read the published version in Animal Cognition → Version 1 posted 9 You are reading this latest preprint version Abstract Musicality is the predisposition to process and produce music. In human beings, processing and producing music often involves entrainment, the ability to synchronise behaviour to external rhythms. Non-human primates generally exhibit poor entrainment skills, which may be due to their relative lack of vocal learning. Focusing on non-primate species like songbird species, is one way to investigate further the evolution of musicality. Here, we investigate spontaneous vocal entrainment in rooks, a social corvid, using non-biologically relevant stimuli. We exposed individual rooks to non natural sound stimuli and tested the effect of various tempos and metrical structures on their willingness to sing along, and on their capacity to entrain (singing along the tempo or the meter, or both). Several individuals sang while listening to the stimuli. Among them, two individuals were influenced by particular tempi and/or metrical structures: one bird produced shorter vocalisations at slower tempo and another reduced the intervals between its vocalisations upon hearing isochronous sequences with a unary metre and slow tempo. Still, the timing of the start of their vocalisations did not match accurately the timing of the beat of the stimuli. Our results suggest that rooks attempted to vocally entrain, an as-yet rare demonstration of vocal flexibility, even among open-ended vocal learners. Despite their evolutionary distance from humans, rooks, and possibly other corvids and songbirds, are interesting species for future studies on rhythmic perception, and could help shed light on the evolution of musical abilities. corvid synchronisation vocalisation rhythm music Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction Musicality is the predisposition to process and produce music (Honing et al., 2015 ). Several hypotheses have been proposed to explain the evolutionary origins of musicality such as promoting social bonding (Music for Social bonding Hypothesis, Dunbar, 2023 ; Savage et al., 2021 ), credible signaling (Mehr et al., 2021 ), facilitating individual identification (Patel and von Rueden, 2021 ) or coordinating activities (Mogan et al., 2017 ). Most of these theories have in common a strong link between musicality and sociality in humans with a striking illustration in human children, where synchronizing to a common rhythm promotes prosocial behaviour and cooperation (Rabinowitch and Meltzoff, 2017 ; Wan and Zhu, 2021 ). Human musicality also involves a strong cultural component, which underlies its multiple forms in various societies. According to Podlipniak ( 2023 ), the evolution of human musicality is supported by cognitive plasticity at the basis of this cultural transmission (see also Killin, 2018 ; Patel 2021 ; Savage et al., 2021 ). Indeed, human musicality frequently involves coordinating to new rhythms, suggesting an open-ended ability to learn new behaviours to adapt to new stimuli. In addition to cognitive plasticity, several components of musicality are commonly found throughout human musical cultures and occasionally in non-human species (Bouwer et al., 2021 ; Honing et al., 2015 ). Beat induction, which allows us to perceive a regular pulse (Patel, 2009), together with relative pitch, or tonal encoding of pitch, are among these basic components. Investigating these basic components in other social species could lead to a better understanding of the evolutionary origins of musicality. Beat induction affords entrainment, i.e., the ability to synchronise behaviour to external rhythms, from tapping in time to a rhythmic stimulus to elaborate multi-participant dances and songs. Entrainment and other rhythm production may be either reactive, i.e. reacting only to past rhythmic events, or predictive, i.e. predicting and matching future rhythmic events, which can imply different internal representations of rhythm and thus indicate different cognitive processes (Bouwer et al., 2021 ; Kotz et al., 2018 ; Ravignani et al., 2013 ). Recent literature suggests that entrainment originates in neural oscillations, which may themselves be based on basic neural circuitry conserved throughout animal evolution (Rouse et al., 2016 ; Wilson and Cook, 2016 ). Yet, evidence for entrainment has remained scarce among mammals and birds (Wilson and Cook, 2016 ). Most studies have looked for entrainment in non-human primates (e.g. chimpanzees: Takeya et al., 2017 ; Hattori and Tomonoga 2021: bonobos: Large et al., 2015; macaques: Nagasaka et al., 2013 ; Selezneva et al., 2013 ; Takeya et al., 2018 ; Zarco et al., 2009 ), with comparatively few studies in other taxa (sea lion: Cook et al., 2013 ; Rouse et al., 2016 ; rat: Katsu et al., 2021 ; budgerigar: Hasegawa et al., 2011 ; cockatiel: Patel et al., 2009 ). Moreover, the vast majority of studies on entrainment has relied on prior conditioning for the specific task rather than on spontaneous entrainment. Non-human primates, despite their close phylogenetic relationship to humans and having some rhythmic abilities, do not appear to share the human capacity for entrainment. For instance, chimpanzees react to rhythms but perform poorly at actually keeping the beat (Hattori and Tomonaga, 2020 ) and show only loosely-coordinated displays, whether through vocalisations (Ghiglieri, 1984 ; Merker et al., 2009 ) or body movements (Hattori and Tomonaga, 2020 ; Lameira et al., 2019 ; Schweinfurth et al., 2022 ). Great apes are capable of social learning and thus of cultural transmission of behaviours (e.g. tool use: van Schaik et al., 1990:, context-specific gestures: Malherbe et al., 2025 ), including drumming (Eleuteri et al., 2025 ; van Loon et al., 2025 ), but unlike humans, they are not open-ended vocal learners, which may partly explain their poorer skills for sound entrainment. By contrast, some species of songbirds are open-ended vocal learners, and may thus be better candidates to study entrainment abilities (Rothenberg, 2014 ; Snyder and Creanza, 2021 ). Several parallels have been drawn between birdsong and human music or language. First, species with demonstrated entrainment tend to also be social species, in line with a purported link between sociality and musicality (Podlipniak, 2023 ; Ravignani et al., 2013 ; Savage et al., 2021 ; Wilson and Cook, 2016 ). Second, birdsong and human music share similarities in temporal organisation (Sainburg et al., 2019 ; Snyder and Creanza, 2021 ), and in the neural pathways activated in the respective species (i.e. humans for music, birds for birdsong) when listened to (Earp and Maney, 2012 ) or performed as a group (Stevenson et al., 2020 ; Riters et al., 2019 ). More precisely, when exposed to a rhythm the auditory sensory-motor areas of the human brain (Grahn and Rowe., 2009; Zatorre et al., 2007 ) and the songbird song circuit (Lampen et al., 2019 ) both show higher activity when synchronising to a rhythm, even at times when no outward behaviour occurs (i.e. no body movement, no vocal output). Some parrot species, including cockatoos (Patel et al., 2009 ) and budgerigars (Hasegawa et al., 2011 ; Seki et al., 2019), are able to extract the beat from music and adapt their movements to it (Fitch, 2009 ; Patel et al., 2009 ). Amongst all songbirds, parrots and corvids could be good models to test vocal entrainment in the avian clade. Both groups of species can imitate sounds (including human or other species voices), an ability that could be necessary for rhythmic learning and entrainment abilities (Dooling et al., 2002 ; Schachner et al., 2009 ; Schachner, 2010 ). A few studies have evidenced cultural transmission of calls and thus open-ended vocal learning in various corvids (Brown, 1985 ; Enggist-Dueblin and Pfister, 2002 ; Kondo, 2021 ). Corvids can also produce duetting behaviour demonstrating their ability to coordinate vocal production with conspecifics (Kondo et al., 2010 ; Seed et al., 2007 ). Corvids can also learn to produce particular vocalisations in response to particular stimuli, suggesting volitional control of their vocal output (Brecht et al., 2019 ; Liao et al., 2024 ; Tomasek et al., 2023 ). Some corvid species also produce undirected songs, i.e. a series of varied vocalisations (variously described as squawks, sneezes, snores or cackles) that may exhibit high structural variation between occurrences (Brown et al., 1985; Brown and Farabaugh, 1997 ; Coombs, 1960 ; Tomasek et al., 2023 ; Martin et al., 2024 ). Among corvids, rooks are well-known for their social cognition skills and complex social lives (Clayton and Emery 2004; Coombs, 1960 ). A previous study evidenced the ability of rooks to spontaneously sing in response to external stimuli, not only starting but also stopping vocal production within a few seconds of the stimulus starting and stopping, respectively (Tomasek et al., 2023 ). This result demonstrates that rooks are capable of coordinating their vocalisations to an external sound without conditioning or reinforcement; it does not give any indication of their capacity to entrain to a rhythm, also because of a timescale longer than that of music. The present study tested whether rooks can vocally entrain to a rhythmic, biologically non-relevant stimulus without prior conditioning. Birds were individually exposed to stimuli with different rhythms, varying in both tempo and metrical structure. Whenever this exposure led to the bird singing, we recorded these songs and evaluated whether birds adjusted their songs to the tempo or to the metrical structure. We predicted that if rooks have the ability to entrain spontaneously to the stimulus, they should adjust their song to the tempos and structures of the stimulus they hear. We additionally investigated whether any adjustment could be more compatible with reactive (if adjustment matches to the preceding stimulus onset) or predictive (if adjustment matches the closest stimulus onset, whether before or after) timing. Methods Study group We studied a group of eleven adult captive rooks (eight males, three females), housed in an outdoor aviary in Strasbourg (France). All birds were captured as juveniles from wild colonies in the region around Strasbourg, were hand-raised and had been housed together in a single aviary since their capture. Each bird was identified by permanent coloured leg rings and an individual name. Water and food were always provided before the experiment was conducted and were accessible ad libitum throughout the day. The study was carried out between September and October 2021 (outside of the breeding period), and between February and June 2022 (during the breeding period); during these two periods, singing activity is generally high in this species. Sound stimulus construction Several sound stimuli were constructed. Each sound stimulus was constructed by assembling 6 sequences of 15 s each, separated by 5 s silence. Each sequence was composed of several short samples of noise (hereafter called tokens) separated by short silent intervals and arranged in predetermined tempo or metre (Fig. 1 ). The sequences could have one of four tempos: 36, 60, 84, or 120 bpm (Fig. 1 A), and one of three metres (patterns of noise tokens repeated over time): unary (i.e. an isochronous sequence of identical sounds), binary, and ternary (Fig. 1 B), resulting in twelve possible sequences. The tempos were defined by the inter-onset intervals (IOI) between successive tokens (e.g. 60 bpm corresponding to 1 s IOI, 120 bpm corresponding to 0.5 s IOI). The metrical sequences were created by clipping the token to the appropriate duration relative to the IOI, either long, where the noise token made up 80% of the IOI, or short, 50% of the IOI, (Mora et al., 2013 , Fitch 2013 ) The unary metre included only long tokens, the binary metre alternated long and short tokens, and the ternary metre repeated a long-short-short pattern. Two different noise tokens could be used to reduce the potential of habituation to a particular noise: a sound of running water tap, and a sound of toy drone (both known to elicit singing from a preliminary study and from Tomasek et al., 2023 ). Within one sound stimulus, only one type of token was used for all sequences, i.e. either water or drone noise. Each sound stimulus was then constructed by semi-randomly drawing six sequences without replacement from the twelve possibilities, then concatenating them with 5 s silences between sequences (Fig. 1 C). Additionally, we constrained the draw so that successive sequences must share either the same tempo or the same metre. A custom R script was used to generate the stimuli. Song elicitation trials A trial consisted in exposing a bird to one sound stimulus. In order to selectively expose individuals to the sound stimulus and avoid interference due to other individuals being exposed to the same stimulus, we built six “sound shower” apparatuses (Fig. 1 D). Each apparatus consisted of a rigid plastic hemisphere (inside dimensions: 27 cm diameter, 14 cm depth) lined with sound-absorbing foam on the outside, with a metal receptacle to hold a speaker (XRS-XB12 or XRS-XB13, Sony). This allowed stimuli to be played and be audible only in a cone under the shower, which was ascertained with a sound level-metre before the experiments: we tested that the sound level was not higher than background outside of the cone of exposure. This allowed selective exposure of single individuals at a time. The sound showers were then installed throughout the aviary at places where birds frequently perched when singing during normal activities. To habituate the birds to the apparatus, the sound showers were installed two weeks before the experiment and left in the aviary until the end of the experimental period. This design allowed us to maintain social dynamics while avoiding statistical dependence issues, enabling us to treat data from different birds as independent data points. Stimulus playback and song recording relied on a single computer running one Audacity session, meaning that triggering the playback simultaneously triggered the recording. The computer was connected to each speaker in the sound showers via Bluetooth at a range of 5 to 12 m. The birds were recorded with a directional microphone (NTG8, RØDE) connected to a sound card (Scarlett 2i2, Focusrite) itself plugged into the computer via USB, and the recordings were digitised at a 48kHz sampling rate. The microphone and computer were held by the same experimenter, outside of the reach of all apparatuses to minimise rerecording of the stimuli. A trial started when a bird spontaneously landed or walked under a sound shower. The bird’s vocalization (if any) were then recorded until the bird left on its own, without interference from the experimenter. For each trial, the experimenter noted whether the bird had vocalised and how long it remained under the shower. When a bird stayed until the end of the stimulus, a different stimulus was played, for as long as the bird remained under the sound shower. In cases where a bird flew away before the end of a stimulus, a different stimulus was selected the next time it arrived under a sound shower. Annotation of recorded songs Recordings in which a rook vocalised at the same time as the stimulus played were manually annotated in Audacity. The spectrogram of the recording was computed (settings: frequency range 1-24000 Hz, Blackman-Harris window, 1024-sample window, zero-padding factor of 2, on the Mel scale, a gain of 0 dB, a dB range of 70, and a high boost of 20 dB) and the beginning and end of each vocalisation were noted based on the spectrogram trace. The stimuli were annotated in the same manner. From these annotations, we computed four acoustic measures (Fig. 1 E): the vocalisation IOI (Ravignani et al., 2017), the duration of vocalisations, the latency between the onset of each vocalisation and the onset of the last preceding noise token (to account for the case where the bird reacts to the stimulus, Ravignani, 2019 ), and the absolute value of the latency between the onset of each vocalisation and the closest onset of a noise token (to account for the case where the bird predicts the stimulus instead of reacting to it, Anglada-Tort et al., 2022 ). Latencies were normalised to relative phases to avoid biases (since slower tempos might otherwise be associated with longer latencies just by virtue of longer intervals betw < een noise tokens), by dividing by the tempo of the respective sequence and multiplying by 60. Statistical analysis We analysed the effects of stimulus tempo and metre on each of the acoustic measures using generalised linear mixed models (GLMM) in R (R Core Team, 2024 ; v4.1.1). We considered the tempo as an ordinal variable, to avoid assuming a linear effect on the acoustic measures. We considered both tempo and rhythm at the same time, effectively combining them into one variable. Finally, we included the trial as a random factor. We fitted separate models for each individual, as we expected that different individuals might be influenced by the stimuli in different manners. One GLMM was fitted per measure, using the lme4 package (Bates et al., 2015 ; v1.1-35.4) with a Gamma family and log link function. The type of noise token (running water or drone) was not included because preliminary likelihood-ratio tests (LRT) showed no significant difference between models including this variable and models leaving it out. Finally, we only selected bouts where at least five vocalisations were produced during a single stimulus sequence, discarding shorter bouts and vocalisations produced between sequences. Each GLMM was compared to a null model including only the random effect using LRT tests with the lmtest package (Zeileis and Hothorn, 2002 ; v0.9-40). For models that were significantly better at fitting the data than their respective null models, pairwise post-hoc comparisons were conducted between values of tempo and metre. These post-hoc comparisons were conducted using the emmeans package (Lenth, 2022 ; v1.8.0) and corrected with Tukey’s method for multiple comparisons. Results Stimulus exposures and song production during trials All eleven rooks in the study group were exposed at least once to the stimuli, and eight sang at least once during a trial (Fig. 2 A). Of the individuals that sang during the trials, only three sang during 11 or 12 of the possible sequences (Balbo, Kafka, and Merlin, Fig. 2 B). Other individuals responded to too few different sequences (Brain: 6, Elie: 5, Tom: 8) or did not respond enough in general (Osiris: 3 responses total, Jolene: 2 responses total) to be included in the same statistical analysis (but see Supplementary Material for a reduced analysis of Brain, Elie, and Tom). Individual preferences for particular tempos or metrical structures For those individuals who sang during the trials, we evaluated whether some individuals might preferentially sing during particular tempos or metrical structures (Fig. 2 B). There was no statistical preference for singing during any particular sequence among the twelve possible ( χ 2 tests for each individual, all p > 0.90). Likewise, when testing each individual on either tempo or metre separately, no statistical preference emerged ( χ 2 tests for each individual, all p > 0.90). Table 1 Likelihood-ratio tests (LRT) comparing the effects of tempo and metre on each measure (IOI, duration, and latencies), compared to the null model. Each line corresponds to one individual bird. For each column, the test statistic is indicated with the associated p-value in parentheses. Significant tests are indicated in bold. Source Vocalisation IOI Vocalisation duration Latency (last onset) Latency (closest onset) Balbo 52.4 (< 0.001) 74.9 (< 0.001) 3.2 (0.99) 1.9 (1) Kafka 4.0 (0.95) 23.4 (0.01) 2.9 (0.98) 3.1 (0.98) Merlin 44.7 (< 0.001) 11.2 (0.43) 4.3 (0.96) 2.6 (1) Did rooks synchronise their songs to the stimuli? We evaluated the ability of the rooks to match the timing of their songs to the sequences played, measured by the latency between the onset of each vocalisation and the onset of the last preceding noise token within the sequence (Fig. 4 A and B). Likelihood-ratio tests indicated that the stimulus had no effect on either of the latency measures in every bird (Table 1 , both Latency columns). Therefore, no bird matched the timing of their song production to the stimuli. Influence of stimuli on vocalisation duration and IOI during songs Even if rooks did not match their song production to the stimuli, they might be influenced by the sequence tempo or metre. The LRT found effects of the sequence tempo and/or metre in two individuals (Balbo, Merlin) for the vocalisation IOI, and also in two individuals (Balbo, Kafka) for the vocalisation duration. Indeed, for IOI, the pairwise tests on the GLMM models showed that Balbo (Fig. 4 C, left) produced significantly longer IOI during sequences with the 36 bpm tempo and the unary metre compared to either the binary metre at the same tempo (ratio 1.61 ± 0.21, z = 3.66, p = 0.014) or the 60 bpm tempo at the same metre (ratio 1.67 ± 0.22, z = 3.85, p = 0.007). Moreover at the 36 bpm tempo, he produced significantly shorter IOI during a binary metre compared to a ternary metre (ratio 0.64 ± 0.08, z = -3.46, p = 0.027). Merlin (Fig. 4 C, right) produced significantly shorter IOI during sequences with an unary metre than with either a binary (ratio 0.58 ± 0.07, z = -4.33, p = 0.001) or ternary metre (ratio 0.59 ± 0.08, z = -4.16, p = 0.002), but only at the 36 bpm tempo. For vocalisation duration (Fig. 4 D), Balbo produced longer vocalisations when the tempo was slower, but not at all metres (Fig. 4 D, left); for instance, during an unary metre, his vocalisations at 36 bpm were longer than at 60 bpm (ratio 2.07 ± 0.29, z = 5.25, p < 0.001), and, during a ternary metre, his vocalisations at 36 bpm were longer than at 84 bpm (ratio 1.82 ± 0.24, z = 4.63, p = 0.0002). Conversely, at 60 bpm he produced significantly longer vocalisations during a binary metre than during either an unary metre (ratio 0.64 ± 0.07, z = -4.10, p < 0.001) or a ternary metre (ratio 1.63 ± 0.23, z = 3.50, p = 0.002). Finally, at 84 bpm his vocalisations were longer during an unary metre than during a ternary metre (ratio 1.66 ± 0.20, z = 4.14, p = 0.002) and also during a binary metre than during a ternary metre (ratio 1.71 ± 0.20, z = 4.52, p = 0.0004), with no difference between unary and binary metres (ratio 0.97 ± 0.11, z = -0.28, p = 1). For Kafka however, no paired comparison reached statistical significance after Tukey correction. Brain, Elie, and Tom were analysed in a similar manner (albeit only with subsets of the possible stimuli) and also adjusted their song production to the stimuli (Table S1 , Fig. S1 ). Discussion We investigated the vocal entrainment abilities of rooks to non-biologically-relevant rhythmic stimuli. When exposed to rhythmic stimuli, a majority of subjects responded vocally to these stimuli albeit heterogeneously. All three males that sang frequently enough to be robustly analysed (Balbo, Kafka, Merlin) sometimes adjusted their song production while exposed to the stimuli in various manners, e.g., by slowing down when exposed to some of the slower tempos (Balbo) or by singing faster when exposed to unary rhythms (Merlin). However, none of them showed precise entrainment to the stimuli, whether in the reactive (i.e. latency with respect to the preceding stimulus onset) or predictive (i.e. latency with respect to the closest stimulus onset) sense. The three other males that could be analysed in a similar but more restricted manner (Brain, Elie, and Tom) also modified their song production. The remaining individuals in the group responded too rarely to analyse at all. A first question is whether birds responded in an automatic manner to the sound stimuli. If that was the case, we would expect most birds to respond frequently when exposed to the sound stimuli. However, we observed a large heterogeneity in vocalizing responses (both within and between individuals). Behaviourally speaking, the rooks showed clear interest in the stimulus, often approaching and looking at the speaker, yet most individuals did not produce songs. Note also that during the experiment, singing was never rewarded nor praised by observers to avoid potential conditioning effects. Thus, responses were not reinforced as was done in most other studies on motor entrainment (e.g. cockatiel: Patel et al., 2009 ; sea lion: Rouse et al., 2016 ) and likely reflect the natural tendency of birds to respond to this type of stimulus. A second question is whether rooks perceived the variations in tempo and metrical structures. The best indication that they did perceive the variations in tempo and metre is the ability of the two individuals who responded most often (Balbo and Brain) to modify their vocalization when listening to particular tempo and/or metres. The use of several types of metre structures was implemented to investigate whether birds react to higher order acoustic patterns, and whether they can perceive interval groups. In this study, one subject vocalized differently when exposed to unary metres compared to when exposed to non-unary metres, suggesting that rooks might perceive and discriminate different metrical structures. There was also a trend for more frequent responses to some metres than others (e.g. Merlin responded most often to the 84 bpm ternary sequences, Elie responded most often to the 120 bpm sequences in general), possibly another indicator of metre discrimination in this species. However, none of the birds adjusted their vocalisations perfectly to these metres, or appeared to discriminate between the binary and ternary metres. We can thus not conclude further about their perception of higher order acoustic patterns. Humans can group together rhythmic sequences varying in emphasis (iambic grouping) or duration (trochaic grouping) and assemble series of pulses into corresponding groups (de la Mora et al., 2013 ; Fitch, 2013 ). The perception of rhythmically structured sounds by humans and other animals remains an active topic of research (see e.g. Kondoh et al., 2021 ), particularly in terms of the origin of musicality. Some species can discriminate between different styles of sounds, such as different composers (pigeon, Porter and Neuringer, 1984 ) or genres (carp, Chase, 2001 ), though the mechanisms behind this discrimination are only rarely known (Celma-Miralles and Toro, 2020 ; Bouwer et al., 2021 ; Kriengwatana et al., 2022 ). Birds, for instance, appear to use features based on temporal oscillations and sound trajectory (i.e. whether sound pitch increases or decreases over time) instead of absolute pitch (Dooling and Prior, 2017 ; Bregman et al., 2016 ; ten Cate et al., 2016 ). Several species have demonstrated rhythm perception (Bouwer et al., 2021 ; Kriengwatana et al., 2022 ), although only between unary and non-unary metres, and not between different non-unary metres (Hagmann and Cook, 2010 ; Humpal and Cynx, 1984 ; Celma-Miralles and Toro, 2020 ; Merchant and Honing, 2014 ; Rouse et al., 2020; Verga et al., 2022 ). In rooks, additional studies are needed to examine this question more thoroughly. A third question is whether birds modulated their songs simply because they were aroused by particular metres or tempos or because they attempted to entrain to those stimuli (even if failing at doing so). If birds modified their songs because of arousal, we would expect a general acceleration of their productions with faster tempos, and/or with more complex metres compared to unary metres. The two birds who modified their song in response to the stimuli in a significant manner indeed shortened their vocalisations or the inter-onset interval between two vocalisations but these adjustments mostly occurred at the slower tempos, i.e., when it is likely easier to adjust. Thus, we cannot exclude that both birds attempted to adjust to the stimulus, rather than “simply” singing faster due to general arousal produced by exposure to the stimuli. An interesting development would be to investigate the vocal flexibility of rooks when singing (a phenomenon that has extremely rarely been described, even in other open-ended vocal learners like parrots or starlings) and whether they shorten vocalisations or shift to selecting shorter vocalisations from their repertoire to better adjust to the beat. Additional studies with more birds, and slower tempos are needed to investigate these questions. In birds, some authors suggest that vocal learning and vocal mimicry are two factors that enhance the likelihood that a species will be capable of entrainment (Wilson and Moehlis, 2014 ). Corvids are known to mimic sounds (Brown, 1985 ), some species may acquire new calls throughout their lifespan (e.g. Kondo, 2021 ) and some have been described as highly variable in their vocalisations (e.g. Roskaft and Epsmark, 1982; Brown and Farabaugh 1997 ; Ellis, 2008 ; Martin et al., 2024 ). Their general vocal aptitudes might enable the flexible song production needed to entrain to rhythmic sounds. Rhythmic entrainment is the ability to perceive a pulse that marks equally spaced points in music or a sequence of auditory stimuli and then to align the motor actions to that pulse or beat (Merchant and Honing, 2014 ). We do not find evidence of vocal entrainment, although two birds may have attempted to entrain. Singing in synchrony with an external pulse is not a standard example of motor entrainment, but it can be considered as a sonorant movement (Chauvigné et al., 2014 ). Some of the rooks occasionally moved at the same time as they sang, although these movements appeared connected to the production of vocalisations (e.g. leaning forward, spreading their wings, raising their head) and they never produced movements such as head-bobbing or foot tapping, like observed in parrots (Patel et al., 2009 ; Keehn et al., 2019 ). We observed no difference in body movements when comparing the same individuals in the experimental conditions here and in their normal life in the aviary, although we did not specifically investigate this aspect. Vocal entrainment is rarely investigated and may be a rare ability in non-human animals. To date, among non-human vertebrates, parrots may be the sole species able to vocally entrain, as shown by scores of amateur online videos of pet parrots singing along to their owners playing music. However, to our knowledge, there is no further empirical evidence of vocal entrainment to an external rhythmic stimulus in any other species. While the recent literature on entrainment and on the evolutionary origins of musicality has skyrocketed in recent years, only a handful of these publications are presenting new data (but see e.g. Eleuteri et al., 2025 ; van Loon et al., 2025 for recent studies in chimpanzees). It remains to see how easily rooks can be conditioned to entrain, as, contrary to most studies, we neither rewarded nor encouraged the birds to sing along with the stimulus. In general, vocal control should be investigated further in rooks and other species; only carrion crows have recently been investigated and successfully learned to vocalise or stay silent on specific cues (Brecht et al., 2019 ) and vocalise specific numbers of times (Liao et al., 2024 ), demonstrating self-control abilities. If rooks can perceive rhythms and adjust their vocal production accordingly, could they also have an internal representation of rhythm? If so, they might be able to predict future pulses, i.e. predictive timing, instead of simply reacting to past pulses, i.e. reactive timing (Ravignani et al., 2014 ). Finding predictive timing in such a phylogenetically distant species as the rook may provide insight into the capabilities and, indeed, the conditions in which musicality may have arisen in humans. Corvids may be phylogenetically distant to humans but, as songbirds with highly developed cognitive and vocal abilities, they could bring much needed comparative perspectives in the study of the evolutionary origins of musicality. Declarations Compliance with Ethical Standards. This study was based on non-invasive procedures and voluntary participation of the birds by way of approaching the apparatus, and we followed ASAB animal care guidelines. French guidelines required no further approval. Author Contribution K.M and V.D . wrote the main manuscript including all figures and tables. K.M and A.H. performed the experiments and analysis. All authors reviewed the manuscript. Data Availability The data and code used for the analysis are provided as supplementary information files. References Anglada-Tort, M., Harrison, P. M. C., & Jacoby, N. (2022). REPP: A robust cross-platform solution for online sensorimotor synchronization experiments. 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Supplementary Files vocalentrainmentrooksupplementary.pdf stimuluscreationcode.zip rhythmtempoanalysis.r fulltablewithlatency.tsv examplesessionMerlin.mp4 Cite Share Download PDF Status: Published Journal Publication published 26 Jan, 2026 Read the published version in Animal Cognition → Version 1 posted Editorial decision: Revision requested 07 Aug, 2025 Reviews received at journal 07 Aug, 2025 Reviews received at journal 07 Jul, 2025 Reviewers agreed at journal 07 Jun, 2025 Reviewers agreed at journal 06 Jun, 2025 Reviewers invited by journal 03 Jun, 2025 Editor assigned by journal 03 Jun, 2025 Submission checks completed at journal 31 May, 2025 First submitted to journal 30 May, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6785900","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":466042343,"identity":"1a08a952-963f-49c9-b370-2ba736a2c6d3","order_by":0,"name":"Killian Martin","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABEklEQVRIiWNgGAWjYBACCWbmxgMJMF4Cgw2IYjwAItkbcGlhbIBpYWxIYEgDs8BaeA7g0AJUCJNiBBp7mLAWyXaglgc1dxh0288+f/Bwx3k5c/4DDId5Ku4w8Ehj1yMNdtixZwxmZ9INGxLP3Da2nJEA1HLmGQMPXwJWLXJgLWyHGcwOpDE2JLbdTtxwg//DYd62wwz2PNgdBtHyD6jl/DOQlnP1G84DHQbSwoNDC9hhiUAFZjfAthxIMAAGIF4tks0gLX2HecxuPGOckdiWbLjhRgLDwTlnDvPg0iJx/vDBhz++HZYzO5/G8PFnm528wfkDjA/eVByWw6UFBlClmXjQRQgCxh+kqR8Fo2AUjILhDQAIS2XYNxSbUAAAAABJRU5ErkJggg==","orcid":"","institution":"École Normale Supérieure de Lyon","correspondingAuthor":true,"prefix":"","firstName":"Killian","middleName":"","lastName":"Martin","suffix":""},{"id":466042344,"identity":"b98f8fdc-d592-4a37-b51d-567d2beed697","order_by":1,"name":"Maëlan Tomasek","email":"","orcid":"","institution":"École Normale Supérieure de Lyon","correspondingAuthor":false,"prefix":"","firstName":"Maëlan","middleName":"","lastName":"Tomasek","suffix":""},{"id":466042345,"identity":"42ee2629-b0b4-426b-b3b1-a9c69d19814b","order_by":2,"name":"Agnès Hivet","email":"","orcid":"","institution":"École Normale Supérieure de Lyon","correspondingAuthor":false,"prefix":"","firstName":"Agnès","middleName":"","lastName":"Hivet","suffix":""},{"id":466042346,"identity":"c68b8dfb-7b9f-496c-ac83-1ce33d46c4dc","order_by":3,"name":"Andrea Ravignani","email":"","orcid":"","institution":"Center for Music in the Brain, Department of Clinical Medicine, Aarhus University \u0026 The Royal Academy of Music","correspondingAuthor":false,"prefix":"","firstName":"Andrea","middleName":"","lastName":"Ravignani","suffix":""},{"id":466042347,"identity":"b8992a77-a0a4-4fbc-9d79-e785706ac07b","order_by":4,"name":"Nicolas Obin","email":"","orcid":"","institution":"Sciences et Technologies de la Musique et du Son","correspondingAuthor":false,"prefix":"","firstName":"Nicolas","middleName":"","lastName":"Obin","suffix":""},{"id":466042348,"identity":"fe547c9b-ccaa-4371-a613-efc0019bea95","order_by":5,"name":"Valérie Dufour","email":"","orcid":"","institution":"Physiologie de la Reproduction et des Comportements","correspondingAuthor":false,"prefix":"","firstName":"Valérie","middleName":"","lastName":"Dufour","suffix":""}],"badges":[],"createdAt":"2025-05-30 15:08:26","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6785900/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6785900/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s10071-025-02038-w","type":"published","date":"2026-01-26T15:59:10+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":84052688,"identity":"7625afab-d2a1-44d9-be5d-5c23e0db84c8","added_by":"auto","created_at":"2025-06-06 08:45:11","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":803630,"visible":true,"origin":"","legend":"\u003cp\u003eStudy protocol and stimuli. A-B) Schematic representation of one sequence with each of the different tempos (A) and metres (B). Each black filled rectangle represents one noise token, white spaces represent silent intervals. C) An example stimulus used for a song elicitation trial, illustrating the draw-without-replacement rule and the rule that successive sequences must share either tempo or metre.. D) Schematic representation of a song elicitation trial. When a rook stays within the sound exposure cone under the sound shower, a stimulus is played from the speaker. Arrows indicate the direction of the stimulus sound. The rook’s vocalisations are recorded by a directional microphone and stored on a computer, with a soundcard connecting the two. The disposition of the sound showers within the aviary meant that the distance between the microphone and the rook was between 5 and 12 m (depending on the particular sound shower). E) Schematic representation of each acoustic measure taken from the annotated songs and stimuli (annotations are shown by the dashed vertical lines: black for the stimulus, red for the song): the inter-onset interval between successive vocalisations (Vocalisation IOI), the duration of the vocalisation (Vocalisation duration), the latency between the onset of each vocalisation and the onset of the last noise token (Latency - last onset), and the latency between the onset of each vocalisation and the onset of the closest noise token (Latency - closest onset).\u003c/p\u003e","description":"","filename":"fig1protocole.tiff.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6785900/v1/abc63f22bf71c090ad766b87.jpg"},{"id":84052690,"identity":"19a37a85-a50d-4cb0-abe9-9bf210c086d5","added_by":"auto","created_at":"2025-06-06 08:45:11","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":242586,"visible":true,"origin":"","legend":"\u003cp\u003eStimulus expositions and song production. A) Number of sequences during which an individual was present (red) and during which it sang (blue) over the course of the experiment. B) Number of times each individual sang during a sequence, for each tempo and metre. Only individuals who sang at least 5 times in total are shown, and only sequences during which an individual sang at least once are shown.\u003c/p\u003e","description":"","filename":"fig2responses.tiff.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6785900/v1/54c5365c2e5dd24df3a5d62f.jpg"},{"id":84052691,"identity":"14a45674-35a9-498a-aa4c-e0fe3638da9e","added_by":"auto","created_at":"2025-06-06 08:45:11","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":542375,"visible":true,"origin":"","legend":"\u003cp\u003eDistribution of all four acoustic measures for each individual, per combination of tempo and metre in stimulus sequences. (A) Latency between the onset of each vocalisation and the onset of the last preceding noise token. (B) Latency between the onset of each vocalisation and the onset of the closest preceding noise token. (C) Vocalisation IOI. (D) Vocalisation duration. Each panel corresponds to one of the individuals included in the analysis. Horizontal bars correspond to statistical tests, with the corresponding p-value above the bar. Only significant differences are shown. Note that the graph uses the log-scale on the y-axis for legibility.\u003c/p\u003e","description":"","filename":"fig3results.tiff.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6785900/v1/ab23f9b9eb4928e81907603a.jpg"},{"id":84053311,"identity":"719b9d60-ec9e-4a20-99af-ef30af6918cc","added_by":"auto","created_at":"2025-06-06 08:53:11","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":475993,"visible":true,"origin":"","legend":"\u003cp\u003eSummary of significant results. All significant pairwise differences found between different stimuli for Balbo and Merlin are summarised. The direction of each difference (from greater to lower) is indicated by the direction of the corresponding arrow (e.g. upper left: Balbo produced vocalisations at significantly longer IOI at 36 bpm and an unary metre than at 60 bpm and an unary metre). Lines represent differences between non-adjacent cells, with the line ending in the cell corresponding to the comparison. Non-significant results are omitted.\u003c/p\u003e","description":"","filename":"fig4recap.tiff.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6785900/v1/77e8ba3265240b08a7ff5e46.jpg"},{"id":101690762,"identity":"c706128d-e76f-4c6f-bc34-b1c8436e7001","added_by":"auto","created_at":"2026-02-02 16:08:15","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2690650,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6785900/v1/d591435e-bc66-481c-93b4-0037576168f5.pdf"},{"id":84052696,"identity":"f66a115d-eb8a-4c94-b3e5-14a6ae8e1fa4","added_by":"auto","created_at":"2025-06-06 08:45:11","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":352132,"visible":true,"origin":"","legend":"","description":"","filename":"vocalentrainmentrooksupplementary.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6785900/v1/28c5bab7cdd6808eff919d0d.pdf"},{"id":84052693,"identity":"9d127461-5065-43f8-9813-cbe44ada3da7","added_by":"auto","created_at":"2025-06-06 08:45:11","extension":"zip","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":716537,"visible":true,"origin":"","legend":"","description":"","filename":"stimuluscreationcode.zip","url":"https://assets-eu.researchsquare.com/files/rs-6785900/v1/a7832bfb48d8136b5a85c598.zip"},{"id":84053310,"identity":"ae2ac427-d9dc-437f-94cb-b2c847bbe947","added_by":"auto","created_at":"2025-06-06 08:53:11","extension":"r","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":7527,"visible":true,"origin":"","legend":"","description":"","filename":"rhythmtempoanalysis.r","url":"https://assets-eu.researchsquare.com/files/rs-6785900/v1/9e495f205fd3ab26d2bcd139.r"},{"id":84052698,"identity":"17608eda-4cfc-4a33-abd5-95e4bbc2df06","added_by":"auto","created_at":"2025-06-06 08:45:11","extension":"tsv","order_by":3,"title":"","display":"","copyAsset":false,"role":"supplement","size":883589,"visible":true,"origin":"","legend":"","description":"","filename":"fulltablewithlatency.tsv","url":"https://assets-eu.researchsquare.com/files/rs-6785900/v1/248d121755670c47b28ae7dd.tsv"},{"id":84052737,"identity":"3cd99b19-6463-4667-aa2e-60fd2584e6cd","added_by":"auto","created_at":"2025-06-06 08:45:56","extension":"mp4","order_by":4,"title":"","display":"","copyAsset":false,"role":"supplement","size":845301143,"visible":true,"origin":"","legend":"","description":"","filename":"examplesessionMerlin.mp4","url":"https://assets-eu.researchsquare.com/files/rs-6785900/v1/dfece5242addaf6245e9699c.mp4"}],"financialInterests":"No competing interests reported.","formattedTitle":"Rooks (Corvus frugilegus) spontaneously attempt to vocally entrain to rhythmic stimuli","fulltext":[{"header":"Introduction","content":"\u003cp\u003eMusicality is the predisposition to process and produce music (Honing et al., \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Several hypotheses have been proposed to explain the evolutionary origins of musicality such as promoting social bonding (Music for Social bonding Hypothesis, Dunbar, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Savage et al., \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), credible signaling (Mehr et al., \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), facilitating individual identification (Patel and von Rueden, \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) or coordinating activities (Mogan et al., \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Most of these theories have in common a strong link between musicality and sociality in humans with a striking illustration in human children, where synchronizing to a common rhythm promotes prosocial behaviour and cooperation (Rabinowitch and Meltzoff, \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Wan and Zhu, \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Human musicality also involves a strong cultural component, which underlies its multiple forms in various societies. According to Podlipniak (\u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), the evolution of human musicality is supported by cognitive plasticity at the basis of this cultural transmission (see also Killin, \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Patel \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Savage et al., \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Indeed, human musicality frequently involves coordinating to new rhythms, suggesting an open-ended ability to learn new behaviours to adapt to new stimuli.\u003c/p\u003e \u003cp\u003eIn addition to cognitive plasticity, several components of musicality are commonly found throughout human musical cultures and occasionally in non-human species (Bouwer et al., \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Honing et al., \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Beat induction, which allows us to perceive a regular pulse (Patel, 2009), together with relative pitch, or tonal encoding of pitch, are among these basic components. Investigating these basic components in other social species could lead to a better understanding of the evolutionary origins of musicality. Beat induction affords entrainment, i.e., the ability to synchronise behaviour to external rhythms, from tapping in time to a rhythmic stimulus to elaborate multi-participant dances and songs. Entrainment and other rhythm production may be either reactive, i.e. reacting only to past rhythmic events, or predictive, i.e. predicting and matching future rhythmic events, which can imply different internal representations of rhythm and thus indicate different cognitive processes (Bouwer et al., \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Kotz et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Ravignani et al., \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eRecent literature suggests that entrainment originates in neural oscillations, which may themselves be based on basic neural circuitry conserved throughout animal evolution (Rouse et al., \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Wilson and Cook, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Yet, evidence for entrainment has remained scarce among mammals and birds (Wilson and Cook, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Most studies have looked for entrainment in non-human primates (e.g. chimpanzees: Takeya et al., \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Hattori and Tomonoga 2021: bonobos: Large et al., 2015; macaques: Nagasaka et al., \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Selezneva et al., \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Takeya et al., \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Zarco et al., \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e2009\u003c/span\u003e), with comparatively few studies in other taxa (sea lion: Cook et al., \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Rouse et al., \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; rat: Katsu et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; budgerigar: Hasegawa et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; cockatiel: Patel et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). Moreover, the vast majority of studies on entrainment has relied on prior conditioning for the specific task rather than on spontaneous entrainment. Non-human primates, despite their close phylogenetic relationship to humans and having some rhythmic abilities, do not appear to share the human capacity for entrainment. For instance, chimpanzees react to rhythms but perform poorly at actually keeping the beat (Hattori and Tomonaga, \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) and show only loosely-coordinated displays, whether through vocalisations (Ghiglieri, \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e1984\u003c/span\u003e; Merker et al., \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) or body movements (Hattori and Tomonaga, \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Lameira et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Schweinfurth et al., \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Great apes are capable of social learning and thus of cultural transmission of behaviours (e.g. tool use: van Schaik et al., 1990:, context-specific gestures: Malherbe et al., \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), including drumming (Eleuteri et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; van Loon et al., \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), but unlike humans, they are not open-ended vocal learners, which may partly explain their poorer skills for sound entrainment. By contrast, some species of songbirds are open-ended vocal learners, and may thus be better candidates to study entrainment abilities (Rothenberg, \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Snyder and Creanza, \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSeveral parallels have been drawn between birdsong and human music or language. First, species with demonstrated entrainment tend to also be social species, in line with a purported link between sociality and musicality (Podlipniak, \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Ravignani et al., \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Savage et al., \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Wilson and Cook, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Second, birdsong and human music share similarities in temporal organisation (Sainburg et al., \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Snyder and Creanza, \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), and in the neural pathways activated in the respective species (i.e. humans for music, birds for birdsong) when listened to (Earp and Maney, \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) or performed as a group (Stevenson et al., \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Riters et al., \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). More precisely, when exposed to a rhythm the auditory sensory-motor areas of the human brain (Grahn and Rowe., 2009; Zatorre et al., \u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e2007\u003c/span\u003e) and the songbird song circuit (Lampen et al., \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) both show higher activity when synchronising to a rhythm, even at times when no outward behaviour occurs (i.e. no body movement, no vocal output). Some parrot species, including cockatoos (Patel et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) and budgerigars (Hasegawa et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Seki et al., 2019), are able to extract the beat from music and adapt their movements to it (Fitch, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Patel et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). Amongst all songbirds, parrots and corvids could be good models to test vocal entrainment in the avian clade. Both groups of species can imitate sounds (including human or other species voices), an ability that could be necessary for rhythmic learning and entrainment abilities (Dooling et al., \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Schachner et al., \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Schachner, \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). A few studies have evidenced cultural transmission of calls and thus open-ended vocal learning in various corvids (Brown, \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e1985\u003c/span\u003e; Enggist-Dueblin and Pfister, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Kondo, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Corvids can also produce duetting behaviour demonstrating their ability to coordinate vocal production with conspecifics (Kondo et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Seed et al., \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). Corvids can also learn to produce particular vocalisations in response to particular stimuli, suggesting volitional control of their vocal output (Brecht et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Liao et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Tomasek et al., \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Some corvid species also produce undirected songs, i.e. a series of varied vocalisations (variously described as squawks, sneezes, snores or cackles) that may exhibit high structural variation between occurrences (Brown et al., 1985; Brown and Farabaugh, \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Coombs, \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1960\u003c/span\u003e; Tomasek et al., \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Martin et al., \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Among corvids, rooks are well-known for their social cognition skills and complex social lives (Clayton and Emery 2004; Coombs, \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1960\u003c/span\u003e). A previous study evidenced the ability of rooks to spontaneously sing in response to external stimuli, not only starting but also stopping vocal production within a few seconds of the stimulus starting and stopping, respectively (Tomasek et al., \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). This result demonstrates that rooks are capable of coordinating their vocalisations to an external sound without conditioning or reinforcement; it does not give any indication of their capacity to entrain to a rhythm, also because of a timescale longer than that of music.\u003c/p\u003e \u003cp\u003eThe present study tested whether rooks can vocally entrain to a rhythmic, biologically non-relevant stimulus without prior conditioning. Birds were individually exposed to stimuli with different rhythms, varying in both tempo and metrical structure. Whenever this exposure led to the bird singing, we recorded these songs and evaluated whether birds adjusted their songs to the tempo or to the metrical structure. We predicted that if rooks have the ability to entrain spontaneously to the stimulus, they should adjust their song to the tempos and structures of the stimulus they hear. We additionally investigated whether any adjustment could be more compatible with reactive (if adjustment matches to the preceding stimulus onset) or predictive (if adjustment matches the closest stimulus onset, whether before or after) timing.\u003c/p\u003e"},{"header":"Methods","content":"\u003cp\u003e \u003cem\u003eStudy group\u003c/em\u003e \u003c/p\u003e\u003cp\u003eWe studied a group of eleven adult captive rooks (eight males, three females), housed in an outdoor aviary in Strasbourg (France). All birds were captured as juveniles from wild colonies in the region around Strasbourg, were hand-raised and had been housed together in a single aviary since their capture. Each bird was identified by permanent coloured leg rings and an individual name. Water and food were always provided before the experiment was conducted and were accessible \u003cem\u003ead libitum\u003c/em\u003e throughout the day. The study was carried out between September and October 2021 (outside of the breeding period), and between February and June 2022 (during the breeding period); during these two periods, singing activity is generally high in this species.\u003c/p\u003e\u003cp\u003e \u003cem\u003eSound stimulus construction\u003c/em\u003e \u003c/p\u003e\u003cp\u003eSeveral sound stimuli were constructed. Each sound stimulus was constructed by assembling 6 sequences of 15 s each, separated by 5 s silence. Each sequence was composed of several short samples of noise (hereafter called tokens) separated by short silent intervals and arranged in predetermined tempo or metre (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The sequences could have one of four tempos: 36, 60, 84, or 120 bpm (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eA), and one of three metres (patterns of noise tokens repeated over time): unary (i.e. an isochronous sequence of identical sounds), binary, and ternary (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eB), resulting in twelve possible sequences. The tempos were defined by the inter-onset intervals (IOI) between successive tokens (e.g. 60 bpm corresponding to 1 s IOI, 120 bpm corresponding to 0.5 s IOI). The metrical sequences were created by clipping the token to the appropriate duration relative to the IOI, either long, where the noise token made up 80% of the IOI, or short, 50% of the IOI, (Mora et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2013\u003c/span\u003e, Fitch \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) The unary metre included only long tokens, the binary metre alternated long and short tokens, and the ternary metre repeated a long-short-short pattern. Two different noise tokens could be used to reduce the potential of habituation to a particular noise: a sound of running water tap, and a sound of toy drone (both known to elicit singing from a preliminary study and from Tomasek et al., \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Within one sound stimulus, only one type of token was used for all sequences, i.e. either water or drone noise.\u003c/p\u003e\u003cp\u003eEach sound stimulus was then constructed by semi-randomly drawing six sequences without replacement from the twelve possibilities, then concatenating them with 5 s silences between sequences (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eC). Additionally, we constrained the draw so that successive sequences must share either the same tempo or the same metre. A custom R script was used to generate the stimuli.\u003c/p\u003e\u003cp\u003e \u003cem\u003eSong elicitation trials\u003c/em\u003e \u003c/p\u003e\u003cp\u003eA trial consisted in exposing a bird to one sound stimulus. In order to selectively expose individuals to the sound stimulus and avoid interference due to other individuals being exposed to the same stimulus, we built six “sound shower” apparatuses (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eD). Each apparatus consisted of a rigid plastic hemisphere (inside dimensions: 27 cm diameter, 14 cm depth) lined with sound-absorbing foam on the outside, with a metal receptacle to hold a speaker (XRS-XB12 or XRS-XB13, Sony). This allowed stimuli to be played and be audible only in a cone under the shower, which was ascertained with a sound level-metre before the experiments: we tested that the sound level was not higher than background outside of the cone of exposure. This allowed selective exposure of single individuals at a time. The sound showers were then installed throughout the aviary at places where birds frequently perched when singing during normal activities. To habituate the birds to the apparatus, the sound showers were installed two weeks before the experiment and left in the aviary until the end of the experimental period. This design allowed us to maintain social dynamics while avoiding statistical dependence issues, enabling us to treat data from different birds as independent data points.\u003c/p\u003e\u003cp\u003eStimulus playback and song recording relied on a single computer running one Audacity session, meaning that triggering the playback simultaneously triggered the recording. The computer was connected to each speaker in the sound showers via Bluetooth at a range of 5 to 12 m. The birds were recorded with a directional microphone (NTG8, RØDE) connected to a sound card (Scarlett 2i2, Focusrite) itself plugged into the computer via USB, and the recordings were digitised at a 48kHz sampling rate. The microphone and computer were held by the same experimenter, outside of the reach of all apparatuses to minimise rerecording of the stimuli. A trial started when a bird spontaneously landed or walked under a sound shower. The bird’s vocalization (if any) were then recorded until the bird left on its own, without interference from the experimenter. For each trial, the experimenter noted whether the bird had vocalised and how long it remained under the shower. When a bird stayed until the end of the stimulus, a different stimulus was played, for as long as the bird remained under the sound shower. In cases where a bird flew away before the end of a stimulus, a different stimulus was selected the next time it arrived under a sound shower.\u003c/p\u003e\u003cp\u003e \u003cem\u003eAnnotation of recorded songs\u003c/em\u003e \u003c/p\u003e\u003cp\u003eRecordings in which a rook vocalised at the same time as the stimulus played were manually annotated in Audacity. The spectrogram of the recording was computed (settings: frequency range 1-24000 Hz, Blackman-Harris window, 1024-sample window, zero-padding factor of 2, on the Mel scale, a gain of 0 dB, a dB range of 70, and a high boost of 20 dB) and the beginning and end of each vocalisation were noted based on the spectrogram trace. The stimuli were annotated in the same manner. From these annotations, we computed four acoustic measures (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eE): the vocalisation IOI (Ravignani et al., 2017), the duration of vocalisations, the latency between the onset of each vocalisation and the onset of the last preceding noise token (to account for the case where the bird reacts to the stimulus, Ravignani, \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), and the absolute value of the latency between the onset of each vocalisation and the closest onset of a noise token (to account for the case where the bird predicts the stimulus instead of reacting to it, Anglada-Tort et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Latencies were normalised to relative phases to avoid biases (since slower tempos might otherwise be associated with longer latencies just by virtue of longer intervals betw \u0026lt; een noise tokens), by dividing by the tempo of the respective sequence and multiplying by 60.\u003c/p\u003e\u003cp\u003e \u003cem\u003eStatistical analysis\u003c/em\u003e \u003c/p\u003e\u003cp\u003eWe analysed the effects of stimulus tempo and metre on each of the acoustic measures using generalised linear mixed models (GLMM) in R (R Core Team, \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; v4.1.1). We considered the tempo as an ordinal variable, to avoid assuming a linear effect on the acoustic measures. We considered both tempo and rhythm at the same time, effectively combining them into one variable. Finally, we included the trial as a random factor. We fitted separate models for each individual, as we expected that different individuals might be influenced by the stimuli in different manners. One GLMM was fitted per measure, using the \u003cem\u003elme4\u003c/em\u003e package (Bates et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; v1.1-35.4) with a Gamma family and log link function. The type of noise token (running water or drone) was not included because preliminary likelihood-ratio tests (LRT) showed no significant difference between models including this variable and models leaving it out. Finally, we only selected bouts where at least five vocalisations were produced during a single stimulus sequence, discarding shorter bouts and vocalisations produced between sequences.\u003c/p\u003e\u003cp\u003eEach GLMM was compared to a null model including only the random effect using LRT tests with the \u003cem\u003elmtest\u003c/em\u003e package (Zeileis and Hothorn, \u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; v0.9-40). For models that were significantly better at fitting the data than their respective null models, pairwise post-hoc comparisons were conducted between values of tempo and metre. These post-hoc comparisons were conducted using the \u003cem\u003eemmeans\u003c/em\u003e package (Lenth, \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; v1.8.0) and corrected with Tukey’s method for multiple comparisons.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e \u003cem\u003eStimulus exposures and song production during trials\u003c/em\u003e \u003c/p\u003e\u003cp\u003eAll eleven rooks in the study group were exposed at least once to the stimuli, and eight sang at least once during a trial (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eA). Of the individuals that sang during the trials, only three sang during 11 or 12 of the possible sequences (Balbo, Kafka, and Merlin, Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eB). Other individuals responded to too few different sequences (Brain: 6, Elie: 5, Tom: 8) or did not respond enough in general (Osiris: 3 responses total, Jolene: 2 responses total) to be included in the same statistical analysis (but see Supplementary Material for a reduced analysis of Brain, Elie, and Tom).\u003c/p\u003e\u003cp\u003e \u003cem\u003eIndividual preferences for particular tempos or metrical structures\u003c/em\u003e \u003c/p\u003e\u003cp\u003eFor those individuals who sang during the trials, we evaluated whether some individuals might preferentially sing during particular tempos or metrical structures (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eB). There was no statistical preference for singing during any particular sequence among the twelve possible (\u003cb\u003eχ\u003c/b\u003e\u003csup\u003e\u003cb\u003e2\u003c/b\u003e\u003c/sup\u003e tests for each individual, all p \u0026gt; 0.90). Likewise, when testing each individual on either tempo or metre separately, no statistical preference emerged (\u003cb\u003eχ\u003c/b\u003e\u003csup\u003e\u003cb\u003e2\u003c/b\u003e\u003c/sup\u003e tests for each individual, all p \u0026gt; 0.90).\u003c/p\u003e\u003cdiv class=\"gridtable\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eLikelihood-ratio tests (LRT) comparing the effects of tempo and metre on each measure (IOI, duration, and latencies), compared to the null model. Each line corresponds to one individual bird. For each column, the test statistic is indicated with the associated p-value in parentheses. Significant tests are indicated in bold.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e\u003ccolgroup cols=\"5\"\u003e\u003c/colgroup\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSource\u003c/p\u003e \u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eVocalisation\u003c/p\u003e \u003cp\u003eIOI\u003c/p\u003e \u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eVocalisation\u003c/p\u003e \u003cp\u003eduration\u003c/p\u003e \u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eLatency\u003c/p\u003e \u003cp\u003e(last onset)\u003c/p\u003e \u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eLatency\u003c/p\u003e \u003cp\u003e(closest onset)\u003c/p\u003e \u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBalbo\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003e52.4 (\u0026lt; 0.001)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e74.9 (\u0026lt; 0.001)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e3.2 (0.99)\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1.9 (1)\u003c/p\u003e \u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKafka\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e4.0 (0.95)\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e23.4 (0.01)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e2.9 (0.98)\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3.1 (0.98)\u003c/p\u003e \u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMerlin\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003e44.7 (\u0026lt; 0.001)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e11.2 (0.43)\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e4.3 (0.96)\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2.6 (1)\u003c/p\u003e \u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/table\u003e\u003c/div\u003e\u003cp\u003e \u003cem\u003eDid rooks synchronise their songs to the stimuli?\u003c/em\u003e \u003c/p\u003e\u003cp\u003eWe evaluated the ability of the rooks to match the timing of their songs to the sequences played, measured by the latency between the onset of each vocalisation and the onset of the last preceding noise token within the sequence (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eA and B). Likelihood-ratio tests indicated that the stimulus had no effect on either of the latency measures in every bird (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, both Latency columns). Therefore, no bird matched the timing of their song production to the stimuli.\u003c/p\u003e\u003cp\u003e \u003cem\u003eInfluence of stimuli on vocalisation duration and IOI during songs\u003c/em\u003e \u003c/p\u003e\u003cp\u003eEven if rooks did not match their song production to the stimuli, they might be influenced by the sequence tempo or metre. The LRT found effects of the sequence tempo and/or metre in two individuals (Balbo, Merlin) for the vocalisation IOI, and also in two individuals (Balbo, Kafka) for the vocalisation duration.\u003c/p\u003e\u003cp\u003eIndeed, for IOI, the pairwise tests on the GLMM models showed that Balbo (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eC, left) produced significantly longer IOI during sequences with the 36 bpm tempo and the unary metre compared to either the binary metre at the same tempo (ratio 1.61 ± 0.21, z = 3.66, p = 0.014) or the 60 bpm tempo at the same metre (ratio 1.67 ± 0.22, z = 3.85, p = 0.007). Moreover at the 36 bpm tempo, he produced significantly shorter IOI during a binary metre compared to a ternary metre (ratio 0.64 ± 0.08, z = -3.46, p = 0.027). Merlin (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eC, right) produced significantly shorter IOI during sequences with an unary metre than with either a binary (ratio 0.58 ± 0.07, z = -4.33, p = 0.001) or ternary metre (ratio 0.59 ± 0.08, z = -4.16, p = 0.002), but only at the 36 bpm tempo.\u003c/p\u003e\u003cp\u003eFor vocalisation duration (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eD), Balbo produced longer vocalisations when the tempo was slower, but not at all metres (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eD, left); for instance, during an unary metre, his vocalisations at 36 bpm were longer than at 60 bpm (ratio 2.07 ± 0.29, z = 5.25, p \u0026lt; 0.001), and, during a ternary metre, his vocalisations at 36 bpm were longer than at 84 bpm (ratio 1.82 ± 0.24, z = 4.63, p = 0.0002). Conversely, at 60 bpm he produced significantly longer vocalisations during a binary metre than during either an unary metre (ratio 0.64 ± 0.07, z = -4.10, p \u0026lt; 0.001) or a ternary metre (ratio 1.63 ± 0.23, z = 3.50, p = 0.002). Finally, at 84 bpm his vocalisations were longer during an unary metre than during a ternary metre (ratio 1.66 ± 0.20, z = 4.14, p = 0.002) and also during a binary metre than during a ternary metre (ratio 1.71 ± 0.20, z = 4.52, p = 0.0004), with no difference between unary and binary metres (ratio 0.97 ± 0.11, z = -0.28, p = 1). For Kafka however, no paired comparison reached statistical significance after Tukey correction.\u003c/p\u003e\u003cp\u003eBrain, Elie, and Tom were analysed in a similar manner (albeit only with subsets of the possible stimuli) and also adjusted their song production to the stimuli (Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e, Fig. \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e).\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eWe investigated the vocal entrainment abilities of rooks to non-biologically-relevant rhythmic stimuli. When exposed to rhythmic stimuli, a majority of subjects responded vocally to these stimuli albeit heterogeneously. All three males that sang frequently enough to be robustly analysed (Balbo, Kafka, Merlin) sometimes adjusted their song production while exposed to the stimuli in various manners, e.g., by slowing down when exposed to some of the slower tempos (Balbo) or by singing faster when exposed to unary rhythms (Merlin). However, none of them showed precise entrainment to the stimuli, whether in the reactive (i.e. latency with respect to the preceding stimulus onset) or predictive (i.e. latency with respect to the closest stimulus onset) sense. The three other males that could be analysed in a similar but more restricted manner (Brain, Elie, and Tom) also modified their song production. The remaining individuals in the group responded too rarely to analyse at all.\u003c/p\u003e\u003cp\u003eA first question is whether birds responded in an automatic manner to the sound stimuli. If that was the case, we would expect most birds to respond frequently when exposed to the sound stimuli. However, we observed a large heterogeneity in vocalizing responses (both within and between individuals). Behaviourally speaking, the rooks showed clear interest in the stimulus, often approaching and looking at the speaker, yet most individuals did not produce songs. Note also that during the experiment, singing was never rewarded nor praised by observers to avoid potential conditioning effects. Thus, responses were not reinforced as was done in most other studies on motor entrainment (e.g. cockatiel: Patel et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; sea lion: Rouse et al., \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) and likely reflect the natural tendency of birds to respond to this type of stimulus.\u003c/p\u003e\u003cp\u003eA second question is whether rooks perceived the variations in tempo and metrical structures. The best indication that they did perceive the variations in tempo and metre is the ability of the two individuals who responded most often (Balbo and Brain) to modify their vocalization when listening to particular tempo and/or metres. The use of several types of metre structures was implemented to investigate whether birds react to higher order acoustic patterns, and whether they can perceive interval groups. In this study, one subject vocalized differently when exposed to unary metres compared to when exposed to non-unary metres, suggesting that rooks might perceive and discriminate different metrical structures. There was also a trend for more frequent responses to some metres than others (e.g. Merlin responded most often to the 84 bpm ternary sequences, Elie responded most often to the 120 bpm sequences in general), possibly another indicator of metre discrimination in this species. However, none of the birds adjusted their vocalisations perfectly to these metres, or appeared to discriminate between the binary and ternary metres. We can thus not conclude further about their perception of higher order acoustic patterns. Humans can group together rhythmic sequences varying in emphasis (iambic grouping) or duration (trochaic grouping) and assemble series of pulses into corresponding groups (de la Mora et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Fitch, \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). The perception of rhythmically structured sounds by humans and other animals remains an active topic of research (see e.g. Kondoh et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), particularly in terms of the origin of musicality. Some species can discriminate between different styles of sounds, such as different composers (pigeon, Porter and Neuringer, \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e1984\u003c/span\u003e) or genres (carp, Chase, \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2001\u003c/span\u003e), though the mechanisms behind this discrimination are only rarely known (Celma-Miralles and Toro, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Bouwer et al., \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Kriengwatana et al., \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Birds, for instance, appear to use features based on temporal oscillations and sound trajectory (i.e. whether sound pitch increases or decreases over time) instead of absolute pitch (Dooling and Prior, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Bregman et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; ten Cate et al., \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Several species have demonstrated rhythm perception (Bouwer et al., \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Kriengwatana et al., \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), although only between unary and non-unary metres, and not between different non-unary metres (Hagmann and Cook, \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Humpal and Cynx, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1984\u003c/span\u003e; Celma-Miralles and Toro, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Merchant and Honing, \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Rouse et al., 2020; Verga et al., \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In rooks, additional studies are needed to examine this question more thoroughly.\u003c/p\u003e\u003cp\u003eA third question is whether birds modulated their songs simply because they were aroused by particular metres or tempos or because they attempted to entrain to those stimuli (even if failing at doing so). If birds modified their songs because of arousal, we would expect a general acceleration of their productions with faster tempos, and/or with more complex metres compared to unary metres. The two birds who modified their song in response to the stimuli in a significant manner indeed shortened their vocalisations or the inter-onset interval between two vocalisations but these adjustments mostly occurred at the slower tempos, i.e., when it is likely easier to adjust. Thus, we cannot exclude that both birds attempted to adjust to the stimulus, rather than “simply” singing faster due to general arousal produced by exposure to the stimuli. An interesting development would be to investigate the vocal flexibility of rooks when singing (a phenomenon that has extremely rarely been described, even in other open-ended vocal learners like parrots or starlings) and whether they shorten vocalisations or shift to selecting shorter vocalisations from their repertoire to better adjust to the beat. Additional studies with more birds, and slower tempos are needed to investigate these questions.\u003c/p\u003e\u003cp\u003eIn birds, some authors suggest that vocal learning and vocal mimicry are two factors that enhance the likelihood that a species will be capable of entrainment (Wilson and Moehlis, \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Corvids are known to mimic sounds (Brown, \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e1985\u003c/span\u003e), some species may acquire new calls throughout their lifespan (e.g. Kondo, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and some have been described as highly variable in their vocalisations (e.g. Roskaft and Epsmark, 1982; Brown and Farabaugh \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Ellis, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Martin et al., \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Their general vocal aptitudes might enable the flexible song production needed to entrain to rhythmic sounds. Rhythmic entrainment is the ability to perceive a pulse that marks equally spaced points in music or a sequence of auditory stimuli and then to align the motor actions to that pulse or beat (Merchant and Honing, \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). We do not find evidence of vocal entrainment, although two birds may have attempted to entrain. Singing in synchrony with an external pulse is not a standard example of motor entrainment, but it can be considered as a sonorant movement (Chauvigné et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Some of the rooks occasionally moved at the same time as they sang, although these movements appeared connected to the production of vocalisations (e.g. leaning forward, spreading their wings, raising their head) and they never produced movements such as head-bobbing or foot tapping, like observed in parrots (Patel et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Keehn et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). We observed no difference in body movements when comparing the same individuals in the experimental conditions here and in their normal life in the aviary, although we did not specifically investigate this aspect. Vocal entrainment is rarely investigated and may be a rare ability in non-human animals. To date, among non-human vertebrates, parrots may be the sole species able to vocally entrain, as shown by scores of amateur online videos of pet parrots singing along to their owners playing music. However, to our knowledge, there is no further empirical evidence of vocal entrainment to an external rhythmic stimulus in any other species.\u003c/p\u003e\u003cp\u003eWhile the recent literature on entrainment and on the evolutionary origins of musicality has skyrocketed in recent years, only a handful of these publications are presenting new data (but see e.g. Eleuteri et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; van Loon et al., \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2025\u003c/span\u003e for recent studies in chimpanzees). It remains to see how easily rooks can be conditioned to entrain, as, contrary to most studies, we neither rewarded nor encouraged the birds to sing along with the stimulus. In general, vocal control should be investigated further in rooks and other species; only carrion crows have recently been investigated and successfully learned to vocalise or stay silent on specific cues (Brecht et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) and vocalise specific numbers of times (Liao et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), demonstrating self-control abilities.\u003c/p\u003e\u003cp\u003e If rooks can perceive rhythms and adjust their vocal production accordingly, could they also have an internal representation of rhythm? If so, they might be able to predict future pulses, i.e. predictive timing, instead of simply reacting to past pulses, i.e. reactive timing (Ravignani et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Finding predictive timing in such a phylogenetically distant species as the rook may provide insight into the capabilities and, indeed, the conditions in which musicality may have arisen in humans. Corvids may be phylogenetically distant to humans but, as songbirds with highly developed cognitive and vocal abilities, they could bring much needed comparative perspectives in the study of the evolutionary origins of musicality.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003cstrong\u003eCompliance with Ethical Standards.\u003c/strong\u003e \u003cp\u003e This study was based on non-invasive procedures and voluntary participation of the birds by way of approaching the apparatus, and we followed ASAB animal care guidelines. French guidelines required no further approval.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eK.M and V.D . wrote the main manuscript including all figures and tables. K.M and A.H. performed the experiments and analysis. All authors reviewed the manuscript.\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eThe data and code used for the analysis are provided as supplementary information files.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAnglada-Tort, M., Harrison, P. M. C., \u0026amp; Jacoby, N. (2022). REPP: A robust cross-platform solution for online sensorimotor synchronization experiments. \u003cem\u003eBehavior Research Methods\u003c/em\u003e, \u003cem\u003e54\u003c/em\u003e(5), 2271\u0026ndash;2285. https://doi.org/10.3758/s13428-021-01722-2\u003c/li\u003e\n\u003cli\u003eBates, D., M\u0026auml;chler, M., Bolker, B., \u0026amp; Walker, S. (2015). Fitting Linear Mixed-Effects Models Using lme4. \u003cem\u003eJournal of Statistical Software\u003c/em\u003e, \u003cem\u003e67\u003c/em\u003e(1), 1\u0026ndash;48. https://doi.org/10.18637/jss.v067.i01\u003c/li\u003e\n\u003cli\u003eBouwer, F. 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URL https://CRAN.R-project.org/doc/Rnews/\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"animal-cognition","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"anco","sideBox":"Learn more about [Animal Cognition](http://link.springer.com/journal/10071)","snPcode":"10071","submissionUrl":"https://submission.nature.com/new-submission/10071/3","title":"Animal Cognition","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"corvid, synchronisation, vocalisation, rhythm, music","lastPublishedDoi":"10.21203/rs.3.rs-6785900/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6785900/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eMusicality is the predisposition to process and produce music. In human beings, processing and producing music often involves entrainment, the ability to synchronise behaviour to external rhythms. Non-human primates generally exhibit poor entrainment skills, which may be due to their relative lack of vocal learning. Focusing on non-primate species like songbird species, is one way to investigate further the evolution of musicality. Here, we investigate spontaneous vocal entrainment in rooks, a social corvid, using non-biologically relevant stimuli. We exposed individual rooks to non natural sound stimuli and tested the effect of various tempos and metrical structures on their willingness to sing along, and on their capacity to entrain (singing along the tempo or the meter, or both). Several individuals sang while listening to the stimuli. Among them, two individuals were influenced by particular tempi and/or metrical structures: one bird produced shorter vocalisations at slower tempo and another reduced the intervals between its vocalisations upon hearing isochronous sequences with a unary metre and slow tempo. Still, the timing of the start of their vocalisations did not match accurately the timing of the beat of the stimuli. Our results suggest that rooks attempted to vocally entrain, an as-yet rare demonstration of vocal flexibility, even among open-ended vocal learners. Despite their evolutionary distance from humans, rooks, and possibly other corvids and songbirds, are interesting species for future studies on rhythmic perception, and could help shed light on the evolution of musical abilities.\u003c/p\u003e","manuscriptTitle":"Rooks (Corvus frugilegus) spontaneously attempt to vocally entrain to rhythmic stimuli","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-06-06 08:45:06","doi":"10.21203/rs.3.rs-6785900/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-08-07T08:49:43+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-08-07T08:06:52+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-07-07T04:22:48+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"76587550650265994929536209354627371999","date":"2025-06-07T12:37:18+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"152622362331042667156234182050824330078","date":"2025-06-06T08:40:29+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-06-03T14:02:02+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-06-03T12:50:34+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-05-31T06:35:02+00:00","index":"","fulltext":""},{"type":"submitted","content":"Animal Cognition","date":"2025-05-30T15:00:06+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"animal-cognition","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"anco","sideBox":"Learn more about [Animal Cognition](http://link.springer.com/journal/10071)","snPcode":"10071","submissionUrl":"https://submission.nature.com/new-submission/10071/3","title":"Animal Cognition","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"f3bbbc1f-9899-495e-9b3a-d7dea24ca6b0","owner":[],"postedDate":"June 6th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2026-02-02T16:05:09+00:00","versionOfRecord":{"articleIdentity":"rs-6785900","link":"https://doi.org/10.1007/s10071-025-02038-w","journal":{"identity":"animal-cognition","isVorOnly":false,"title":"Animal Cognition"},"publishedOn":"2026-01-26 15:59:10","publishedOnDateReadable":"January 26th, 2026"},"versionCreatedAt":"2025-06-06 08:45:06","video":"","vorDoi":"10.1007/s10071-025-02038-w","vorDoiUrl":"https://doi.org/10.1007/s10071-025-02038-w","workflowStages":[]},"version":"v1","identity":"rs-6785900","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6785900","identity":"rs-6785900","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}