Evidence for Vocal Learning by a Scrub Jay

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OHNO, S. 1967. Sex chromosomes and sex-linked ß J. S. QUINNß F. COOKE, & B. N. WHITE. 1987. genes. New York, Springer. DNA marker analysisdetects multiple maternity PAGE, D. C., M. E. HARPER, J. LOVE, & D. BOTSTEIN. and paternity in singlebroods of the LesserSnow 1984. Occurrence of a transpositionfrom the Goose. Nature 326: 392-394. X-chromosomelong arm to the Y-chromosome SOKAL,R. R., & F. J. ROHLF. 1969. Biometry. San short arm during human evolution. Nature 311: Franciscoß W. H. Freeman and Co. 119-123. SOUTHERN,E.g. 1975. Detection of specific se- ß B. DE MARTINvILLEßD. BARKER,A. WYMAN, R. quencesamong DNA fragmentsseparated by gel WHITEßU. FRANCKE,&D. BOTSTEIN.1982. Single- electrophoresis.J. Mol. Biol. 98: 503-517. copy sequencehybridizes to polymorphic and STRANGEL,P.W. 1986. Lackof effectsfrom sampling homologousloci on human X and Y chromo- blood from small birds. Condor 88: 244-245. somes. Proc. Natl. Acad. Sci. USA 79: 5352-5356. TABER,R. D. 1971. Criteria of sexand age. Pp. 325- PRus,S. E., & S.C. SCHMUTZ.1987. Comparative 401 in Wildlife managementtechniquesß 3rd ed. efficiencyand accuracyof surgical and cytoge- (R. H. Giles,Ed.). Washingtonß D.C., The Wildlife netic sexing in Psittacines.Avian Diseases31: Society. 420-424. TONEßM., Y. SAKAKI, T. HASHIGUCHI, & S. MIZUNO. QUINN, T. W., & B. N. WHITE. 1987. Identification of 1984. Genus specificityand extensive methyl- restriction-fragment-lengthpolymorphisms in ation of the W chromosome-specificrepetitive genomicDNA of the LesserSnow Goose(Anser DNA sequencesfrom the domestic fowl, Gallus caerulescenscaerulescens). Mol. Biol. Evol. 4: 126- gallusdomesticus. Chromosoma 89: 228-237. 143. ß J. C. DAVIESßF. COOKEß& B. N. WHITE. 1989. Received14 April 1989ßaccepted 9 September1989. Geneticanalysis of offspringof a female-female pair in the LesserSnow GooseChen caerulescens caerulescens).Auk 106: 177-184.

Evidence for Vocal Learning by a Scrub Jay

TOM WEBBER• AND ROSEMARY A. STEFANI2 •FloridaMuseum of NaturalHistoryß University of FloridaßGainesvilleß Florida 32611 USA, and 2Divisionof EnvironmentalStudiesß University of California,Davisß California 95616 USA

The vocalizations of corvids, other than their faint- while calling loudly. Green gave the rattle call (Web- ly uttered and seldom heard whisper song, are typ- ber 1984)in responseto Pink's flights.This behavior ically referred to as calls rather than songs(Hardy is typical of ScrubJays when they form pairs in the 1983, Goodwin 1986). This convention reflectsa wide- wild (Webber 1984).Other jays in the aviaryßinclud- spreadimpression that these calls are more simple ing Red, often called and flew in responseto Pink than the primary advertisingsongs of many other and Green'spair-forming behavior, as wild ScrubJays oscinesand are delivered in a different way. Corvids do when a new pair formsin a neighboringterritory mimic the soundsof other bird speciesand even the (Webber 1984). Red gave two kinds of calls (referred soundsof inanimate objects(Goodwin 1986),but the to as A [Fig. la] and B [Fig. ld]) that, to Webberß evidencethat they learn their distinctivespecies-spe- sounded identical to those of Pink (Fig. lb, e) and cific callsis sketchy.We presentevidence that a cap- anothermale FloridaScrub Jay ("Yellow") in the same tive ScrubJay (Aphelocoma coerulescens superciliosa) from aviary. Webber recordedthe calls of Red, Pinkßand California learned two calls from Florida Scrub Jays Yellow on six daysfrom 11 Januaryto 29 March 1981. (A. c. coerulescens). These recordingsinclude at least 138 calls of type A A Californiajay wascaptured near Carmichael, Sac- by Pink and 40 by Red, as well as at least 100 callsof ramentoCountyß as a late-stagenestling (23 June1979) type B by Pinkß36 by Red, and 7 by Yellow (Florida and hand-raised. On 5 August 1979, it was given to Museum of Natural History BioacousticsArchives the Florida Museumßwhere it lived in a large aviary mastertapes 905-907). with other wild-caught ScrubJays from Florida and Webber(1984) found no calls of typesA or B in a California (Webber and Cox 1987). In the following 12-month study of Scrub Jay calls in Los Angeles account,we refer to this California jay as "Red" be- County, California (FSM masters 633-637B, 639A- causeof its red leg bands. 641B).Stefani (pers. obs.) also found no callsof types In mid-January1981ß Webber noticed that a male A or B in a 3-month study of ScrubJay vocalizations FloridaScrub Jay (pink leg bands:"Pink") in the avi- in Davisß California, 28 miles from Carmichael. We ary was pairing with a female ("Green") from Cali- think it unlikely that wild ScrubJays in Carmichael fornia. Pink passedfood to Greenßcalled in response give calls A and B. to her flights, and flew back and forth in front of her Wild Scrub Jaysin Florida give calls (FM masters January1990] ShortCommunications 203

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d e f

o 1 2 TIME IN SECONDS Fig.1. (a)Call A givenby Red,1 March1981; FM master905B cut 3. (b)Call A givenby Pink,27 March 1981;FM master907A cut 1. (c) Closestmatch of call A by a wild ScrubJay, Cedar Key, Levy County,Florida, 5 March 1976;FM master904 cut 6. (d) Call B given by Red, 29 March 1981;FM master907B cut 1. (e) Call B given by Pink, 27 March 1981;FM master907A cut 1. (f) Closestmatch of call B by a wild ScrubJay, Cedar Key, Levy County, Florida, 17 March 1981;FM master906B cut 1. (Kay Sona-Graph7029A, wide band.)

904 and 906B) that sound the same as A and B. The differences between the calls of the Florida and var- sonogramsof thesewild ScrubJay calls (Fig. lc, e) ious westernpopulations, as well as variation within match those of A and B more closely than do sono- Florida (Webber pets. obs.)populations. Red's acqui- gramsof any known California ScrubJay calls.Red, sition of severalnew callssuggests that the geograph- the California jay, seemsto have learned two of its ical variation in some call types may be traditional callsand, to the extent that the confinesof the aviary and that if wild jays move from one area to another allowed, to have used them like wild ScrubJays. where the callsare different, they may learn the local We havefound no accountof similarvocal learning calls. by a corvid. Kroodsmaand BayIls (1982) infer that We are grateful for advice and assistancefrom J. BlueJays (Cyanocitta cHstata) learn their bell calls(Kra- Cox, J. W. Hardy, S. Hope, C. van Riper III, the late mer and Thompson 1979) becausethese calls vary R. Warner, G. E. Woolfenden, and an anonymousref- over short distancesin a dialect-like pattern (seealso eree. Racineand Thompson1983). Brown (1985)found that captive American Crows (Corvusbrachyrhynchos) ira- Rated elements in the whisper songsof other crows in their socialgroup. Choughs(Pyrrhocorax pyrrho- coraxand P. graculus)can imitate the individual vari- BROWN,E.D. 1985. The role of songand vocal im- ants in their associates'calls (Sitasuwan and Thaler itation among Common Crows (Corvusbrachy- 1985), as can Common Ravens (Corvuscorax; Gwinner rhynchos).Z. Tierpsychol.68: 115-136. and Kneutgen 1962). GOODWIN, D. 1986. Crows of the world, 2nd ed. The extentand nature of geographicalvariation in British Museum (Natural History). ScrubJay calls is little known, but there certainly are GWINNER,E., & J. KNœUTGEN.1962. Uber die biolo- 204 ShortCommunications [Auk,Vol. 107

gischebedeutung der zweckdienlichenAnwen- ganizationof wintering Blue Jays.Behaviour 87: dung erlernter Laute bei Vogeln. Z. Tierpsychol. 237-255. 19: 692-696. $ITASUWAN,•'•., & E. THALER. 1985. Lautinventarund HARDY,J. W. 1983. Insertbooklet for Voices of the Verstandigungbei Alpenkrahe (Pyrrhocoraxpyr- New World jays, crows,and their allies, family rhocorax),Alpendohle (Pyrrhocoraxgraculus) und Corvidae. LP disc, ARA-9. Gainesville, Florida, deren Hybriden. J. Ornithol. 126: 181-193. ARA records. WEBBER,T. 1984. Form and function of the long- KRAMER,I-I. G., & N. $. THOMPSON.1979. Geographic range calls of ScrubJays, Aphelocoma coerulescens variation in the bell calls of the Blue Jay (Cya- obscura. Ph.D. dissertation, Gainesville, Univ. nocittacristata). Auk 96: 423-425. Florida. KROODsMA,D. E., & J. E. BAYLIS.1982. Appendix: a , & J. A. Cox. 1987. Breedingand behaviour world survey of evidencefor vocal learning in of Scrub Jaysin captivity. Avicultural Magazine birds. In Acoustic communication in birds, vol. 2 93: 6-14. (D. E. Kroodsma and E. H. Miller, Eds.). New York, Academic Press. Received24 April 1989, accepted9 September1989. RACINE, R. N., & N. $. THOMPSON. 1983. Social or-