Abstract
Song type matching has been hypothesized to be a graded signal of aggression; however, it is often the case that variation in matching behavior is unrelated to variation in aggressiveness. An alternative view is that whether an individual matches a song is determined mainly by syntactic rules governing how songs are sequenced. In song sparrows (Melospiza melodia), two such rules are the cycling rule, which directs that a bird cycles through its song types in close to the minimum number of bouts, and the bout length rule, which directs that a long bout of a song type is followed by a long interval before that song type is sung again. The effect of these rules on matching is confirmed here for a population of eastern song sparrows. Territorial males were challenged at the end of a recording session with playback of one of their own song types. Logistic regression showed that the probability of matching the playback song type increased with the length of the interval since the subject had last sung that song type, as predicted by the cycling rule. The probability of matching decreased as prior bout length increased, as predicted by the bout length rule. In a multivariate logistic regression, interval length and prior bout length were both associated with matching and together correctly predicted matching in 81.3% of cases. The results support the syntactic constraints hypothesis, which proposes that matching is a non-signaling by-product of internal rules governing the ordering of song type sequences.
Significance statement
Vocal matching has attracted widespread interest in large part because it seems an effective method of directing an aggressive message at a particular recipient. Here, we show that in an eastern population of song sparrows, decisions on whether to match another bird are largely determined by internal rules of syntax governing how a singer sequences its song types, rather than by variation in aggressiveness or other individual traits. These results support the view that vocal matching is an incidental byproduct of internal mechanisms controlling the ordering of vocalization types and so is not a signal at all. This hypothesis may be broadly applicable to vocal matching in other species.
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Data are deposited on Figshare https://doi.org/10.6084/m9.figshare.20483550.
References
Akçay C, Beecher MD (2020) Song sparrows do not discriminate between their own song and stranger song. Behav Process 178:104184 https://doi.org/10.1016/j.beproc.2020.104184
Akçay C, Tom ME, Holmes D, Campbell SE, Beecher MD (2011) Sing softly and carry a big stick: signals of aggressive intent in the song sparrow. Anim Behav 82:377–382 https://doi.org/10.1016/j.anbehav.2011.05.016
Akçay C, Tom ME, Campbell SE, Beecher MD (2013) Song type matching is an honest early threat signal in a hierarchical animal communication system. Proc R Soc B 280:20122517 https://doi.org/10.1098/rspb.2012.2517
Alves A, Antunes R, Bird A, Tyack PL, Miller PJO, Lam FPA, Kvadsheim PHK (2014) Vocal matching of naval sonar signals by long-finned pilot whales (Globicephala melas). Mar Mammal Sci 30:1248–1257 https://doi.org/10.1111/mms.12099
Anderson RC, Searcy WA, Nowicki S (2005) Partial song matching in an eastern population of song sparrows, Melospiza melodia. Anim Behav 69:189–196 https://doi.org/10.1016/j.anbehav.2004.02.019
Anderson RC, Searcy WA, Nowicki S (2008) Testing the function of song matching in birds: responses of eastern male song sparrows Melospiza melodia to partial song matching. Behaviour 145:347–363 https://doi.org/10.1163/156853908783402876
Ansell D, Magrath RD, Haff TM (2020) Song matching in a long-lived, sedentary bird with a low song rate: the importance of song type, song duration and intrusion. Ethology 126:1098–1110 https://doi.org/10.1111/eth.13090
Arak A (1983) Vocal interactions, call matching and territoriality in a Sri Lankan treefrog, Philautus leucorhinus (Rhacophoridae). Anim Behav 31:292–302 https://doi.org/10.1016/S0003-3472(83)80199-7
Balsby TJS, Bradbury JW (2009) Vocal matching by orange-fronted conures (Aratinga canicularis). Behav Process 82:133–139 https://doi.org/10.1016/j.beproc.2009.05.005
Beecher MD, Campbell SE, Stoddard PK (1994) Correlation of song learning and territory establishment strategies in the song sparrow. P Natl Acad Sci USA 91:1450–1454 https://doi.org/10.1073/pnas.91.4.1450
Beecher MD, Campbell SE, Burt JM, Hill CE, Nordby JC (2000) Song-type matching between neighbouring song sparrows. Anim Behav 59:21–27 https://doi.org/10.1006/anbe.1999.1276
Bertram B (1970) The vocal behaviour of the Indian hill mynah, Gracula religiosa. Anim Behav Monogr 3:79–192 https://doi.org/10.1016/S0066-1856(70)80003-6
Burt JM, Campbell SE, Beecher MD (2001) Song type matching as threat: a test using interactive playback. Anim Behav 62:1163–1170 https://doi.org/10.1006/anbe.2001.1847
Burt JM, Bard SC, Campbell SE, Beecher MD (2002) Alternative forms of song matching in song sparrows. Anim Behav 63:1143–1151 https://doi.org/10.1006/anbe.2002.3011
Catchpole CK, Slater PJB (2008) Bird song: biological themes and variations, 2nd edn. Cambridge University Press, Cambridge
DuBois AL, Nowicki S, Searcy WA (2016) A test for repertoire matching in eastern song sparrows. J Avian Biol 47:146–152 https://doi.org/10.1111/jav.00811
Falls JB (1985) Song matching in western meadowlarks. Can J Zool 63:2520–2524 https:/doi.org/10.1139/z85-373
Falls JB, Krebs JR (1975) Sequence of songs in repertoires of western meadowlarks (Sturnella neglecta). Can J Zool 53:1165–1178 https://doi.org/10.1139/z75-135
Falls JB, Krebs JR, McGregor PK (1982) Song matching in the great tit (Parus major): the effect of similarity and familiarity. Anim Behav 30:997–1009 https://doi.org/10.1016/S0003-3472(82)80188-7
Gammon DE, Hendrick MC, Baker MC (2008) Vocal communication in a songbird with a novel song repertoire. Behaviour 145:1003–1026 https://doi.org/10.1163/156853908784474524
Gerhardt HC, Roberts JD, Bee MA, Schwartz JJ (2000) Call matching in the quacking frog (Crinia georgiana). Behav Ecol Sociobiol 48:243–251 https://doi.org/10.1007/s002650000226
Hedley RW, Denton KK, Weiss RE (2017) Accounting for syntax in analyses of countersinging reveals hidden vocal dynamics in a songbird with a large repertoire. Anim Behav 131:23–32 https://doi.org/10.1016/j.anbehav.2017.06.021
Hill CE, Campbell SE, Nordby JC, Burt JM, Beecher MD (1999) Song sharing in two populations of song sparrows (Melospiza melodia). Behav Ecol Sociobiol 46:341–349 https://doi.org/10.1007/s002650050628
Hinde RA (1958) Alternative motor patterns in chaffinch song. Anim Behav 6:211–218 https://doi.org/10.1016/0003-3472(58)90053-8
Hughes M, Nowicki S, Searcy WA, Peters S (1998) Song-type sharing in song sparrows: implications for repertoire function and song learning. Behav Ecol Sociobiol 42:437–446 https://doi.org/10.1007/s002650050458
Hurd PL, Enquist M (2005) A strategic taxonomy of biological communication. Anim Behav 70:1155–1170 https://doi.org/10.1016/j.anbehav.2005.02.014
Hyman J, Hughes M, Searcy WA, Nowicki S (2004) Individual variation in the strength of territory defense in male song sparrows: correlates of age, territory tenure, and neighbor aggressiveness. Behaviour 141:15–27 https://doi.org/10.1163/156853904772746574
Janik VM (2000) Whistle matching in wild bottlenose dolphins (Tursiops truncatus). Science 289:1355–1357 https://doi.org/10.1126/science.289.5483.1355
King SL, McGregor PK (2016) Vocal matching: the what, the why and the how. Biol Lett 12:20160666 https://doi.org/10.1098/rsbl.2016.0666
Krebs JR (1976) Habituation and song repertoires in the great tit. Behav Ecol Sociobiol 1:215–227 https://doi.org/10.1007/BF00299198
Krebs JR, Ashcroft R, Van Orsdol K (1981) Song matching in the great tit Parus major L. Anim Behav 29:918–923 https://doi.org/10.1016/S0003-3472(81)80029-2
Lapierre JM, Mennill DJ, MacDougall-Shackleton EA (2011) Spatial and age-related variation in use of locally common song elements in dawn singing of song sparrows Melospiza melodia: old males sing the hits. Behav Ecol Sociobiol 65:2149–2160 https://doi.org/10.1007/s00265-011-1223-1
Machlis L, Dodd PWD, Fentress JC (1985) The pooling fallacy: problems arising when individuals contribute more than one observation to the data set. Z Tierpsychol 68:201–214 https://doi.org/10.1111/j.1439-0310.1985.tb00124.x
Marler P, Peters S (1987) A sensitive period for song acquisition in the song sparrow, Melsopiza melodia: A case of age-limited learning. Ethology 76:89–100 https://doi.org/10.1111/j.1439-0310.1987.tb00675.x
Marler P, Peters S (1988) The role of song phonology and syntax in vocal learning preferences in the song sparrow, Melospiza melodia. Ethology 77:125–149 https://doi.org/10.1111/j.1439-0310.1988.tb00198.x
Martin DJ (1990) Songs of the fox sparrow. III. Ordering of song. Wilson Bull 102:655–671
Miller PJO, Shapiro AD, Tyack PL, Solow AR (2004) Call-type matching in vocal exchanges of free-ranging resident killer whales, Orcinus orca. Anim Behav 67:1099–1107 https://doi.org/10.1016/j.anbehav.2003.06.017
Nice MM (1943) Studies in the life history of the song sparrow. II. The behavior of the song sparrow and other passerines. Trans Linn Soc NY 6:1–328
Nordby JC, Campbell SE, Beecher MD (1999) Ecological correlates of song learning in song sparrows. Behav Ecol 10:287–297 https://doi.org/10.1093/beheco/10.3.287
Nowicki S, Podos J, Valdés F (1994) Temporal patterning of within-song type and between-song type variation in song repertoires. Behav Ecol Sociobiol 34:329–335 https://doi.org/10.1007/BF00197003
Nowicki S, Searcy WA, Krueger T, Hughes M (2002) Individual variation in response to simulated territorial challenge among territory-holding song sparrows. J Avian Biol 33:253–259 https://doi.org/10.1034/j.1600-048X.2002.330307.x
Rogers D (2004) Repertoire size, song sharing and type matching in the rufous bristlebird (Dasyornis broadbenti). Emu 104:7–13 https://doi.org/10.1071/MU01070
Saunders AA (1924) Recognizing individual birds by song. Auk 41:242–259 https://doi.org/10.2307/4074619
Schroeder DJ, Wiley RH (1983) Communication with shared song themes in tufted titmice. Auk 100:414–424 https://doi.org/10.1093/auk/100.2.414
Searcy WA, Beecher MD (2009) Song as an aggressive signal in songbirds. Anim Behav 78:1281–1292 https://doi.org/10.1016/j.anbehav.2009.08.011
Searcy WA, McArthur PD, Peters SS, Marler P (1981) Response of male song and swamp sparrows to neighbour, stranger, and self songs. Behaviour 77:152–163 https://doi.org/10.1163/156853981X00202
Searcy WA, McArthur PD, Yasukawa K (1985) Song repertoire size and male quality in song sparrows. Condor 87:222–228 https://doi.org/10.2307/1366886
Searcy WA, Anderson RC, Nowicki S (2006) Bird song as a signal of aggressive intent. Behav Ecol Sociobiol 60:234–241 https://doi.org/10.1007/s00265-006-0161-9
Searcy WA, DuBois AL, Rivera-Cáceres K, Nowicki S (2013) A test of a hierarchical signalling model in song sparrows. Anim Behav 86:309–315 https://doi.org/10.1016/j.anbehav.2013.05.019
Searcy WA, Akçay C, Nowicki S, Beecher MD (2014) Aggressive signaling in song sparrows and other songbirds. Adv Study Behav 46:89–125 https://doi.org/10.1016/B978-0-12-800286-5.00003-1
Searcy WA, Ocampo D, Nowicki S (2019) Constraints on song type matching in a songbird. Behav Ecol Sociobiol 73:102 https://doi.org/10.1007/s00265-019-2708-6
Searcy WA, Soha J, Peters S, Nowicki S (2022) Long-distance dependencies in birdsong syntax. Proc R Soc B 289:20212473 https://doi.org/10.1098/rspb.2021.2473
Stoddard PK, Beecher MD, Campbell SE, Horning CL (1992) Song-type matching in the song sparrow. Can J Zool 70:1440–1444 https://doi.org/10.1139/z92-200
Sugiura H (1998) Matching of acoustic features during the vocal exchange of coo calls by Japanese macaques. Anim Behav 55:673–687 https://doi.org/10.1006/anbe.1997.0602
Templeton CN, Ríos-Chelén AA, Quirós-Guerrero E, Mann NI, Slater PJB (2013) Female happy wrens select songs to cooperate with their mates rather than confront intruders. Biol Lett 9:20120863 https://doi.org/10.1098/rsbl.2012.0863
Todt D (1981) On functions of vocal matching: effect of counter-replies on song post choice and singing. Z Tierpsychol 57:73–93 https://doi.org/10.1111/j.1439-0310.1981.tb01313.x
Vehrencamp SL (2001) Is song-type matching a conventional signal of aggressive intentions? Proc R Soc Lond B 268:1637–1642 https://doi.org/10.1098/rspb.2001.1714
Verner J (1975) Complex song repertoire of male long-billed marsh wrens in eastern Washington. Living Bird 14:263–300
Wheeldon A, Szymanski P, Surmacki A, Osiejuk TS (2021) Song type and song type matching are important for joint territorial defense in a duetting songbird. Behav Ecol 32:883–894 https://doi.org/10.1093/beheco/arab030
Whitney CL (1985) Serial order in wood thrush song. Anim Behav 33:1250–1265 https://doi.org/10.1016/S0003-3472(85)80185-8
Yuan DH, Price JJ (2011) Song-type sharing and matching in a bird with very large song repertoires, the tropical mockingbird. Behaviour 148:673–689 https://doi.org/10.1163/000579511X573908
Ballentine B, Searcy WA, Nowicki S (2008) Reliable aggressive signalling in swamp sparrows. Anim Behav 75:693-703. https//doi.org/https://doi.org/10.1016/j.anbehav.2007.07.025
Acknowledgements
We thank Susan Peters for independently classifying answer songs into song types. We are grateful to Dr. Corinne Richards-Zawacki and the staff of the University of Pittsburgh’s Pymatuning Laboratory of Ecology for logistical support, to the Pennsylvania Game Commission for access to study sites, and to the Robert E. Maytag Research Fund of the University of Miami and Duke University’s Trinity College of Arts and Sciences for financial support. We thank Jeff Podos and two anonymous reviewers for helpful advice on the manuscript.
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All procedures used in this study were approved by the Institutional Animal Care and Use Committees of the University of Miami (protocols #14-049 and #20-055-LF), Duke University (protocols #A004-17-01 and #A260-19-12) and the University of Pittsburgh (protocols #18052783 and #21059264). All applicable national and institutional guidelines for the ethical treatment of animals were followed. The research was conducted with free-living birds that were not captured or otherwise handled for this study.
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Searcy, W.A., Chronister, L.M. & Nowicki, S. Syntactic rules predict song type matching in a songbird. Behav Ecol Sociobiol 77, 12 (2023). https://doi.org/10.1007/s00265-022-03286-3
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DOI: https://doi.org/10.1007/s00265-022-03286-3