Vocalizations of the Golden-Crowned Kinglet in Eastern North America

j. Field Ornithol., 64(3):346-351

VOCALIZATIONS OF THE GOLDEN-CROWNED KINGLET IN EASTERN NORTH AMERICA

CHRISTOPHER T. NAUGLER Departmentof Biology Queen'sUniversity KingstonOntario K7L $N6 Canada Abstract.--Songsand callsof the Golden-crownedKinglet (Regulussatrapa) from a number of locationsin Eastern North America were examined.Songs were comprisedof a variable number of six syllable types. The order of syllable presentation,however, was highly predictableboth within and among individuals.Song structureresembled that of the con- generic Firecrest (R. ignicapillus)of Europe. Two commoncalls of the Golden-crowned Kinglet (ti and zeet) occurredthroughout the geographicrange examined and strongly resembledcalls of the Firecrestand Goldcrest(R. regulus),as well as havingsimilar putative functions.

VOCALIZACI•)N DE REGULUS SATRAPA EN EL ESTE DE NORTE AMERICA Sinopsis.--Sccxamin6 cl cantoy las 11amadasdc individuosdc Regulus satrapa cn localidadcs dcl cstc dc Nortc Am(•rica. Sc cncontr6 una scric dc cantos variables dc scis silabas. El ordcn dc las silabasresult6 scr prcdcciblctanto cnun mismoindividuo como entre individuos.La cstructuradc la canci6ncs parccidaa ladc congOherescuropcos como R. ignicapillus.Dos dc las 11amadasdcl ave (ti y ziit) sc cncontrarona todo lo largo dc la cxtcnci6ngcogrfifica cstudiaday scparcccn mucho alas producidaspot R. ignicapillusy R. regulus.Las 11amadas parcccn tcncr una funci6n similar cn las tres cspccics. Despite extensivedescriptive work over the past severaldecades, the vocalizationsof most speciesof North American passetineshave yet to be adequatelydescribed. The Golden-crownedKinglet (Regulussatrapa) is one such species.In contrast,European membersof this genus,the Goldcrest(R. regulus)and the Firecrest(R. ignicapillus)have been studied extensively(Becker 1976, 1977a,b). An early descriptiongives the songof the Golden-crownedKinglet as "risingat first with fine, high-pitched,somewhat faltering notes, and then falling away into a short, rapid rather explosivewarble" (Forbush 1929: 376). Galati and Galati (1985) recognizedt.wo categories of song:simple song,consisting of a seriesof ascendingnotes, and complexsong, which beginslike simple songbut endsin a musical warble. Descriptionsof calls of the Golden-crownedKinglet have recognized from two to five call types (Forbush 1929:376, Galati and Galati 1985), but the absenceof sound spectrographsand variation in onomatopoeic descriptionsmake the work of previousauthors difficult to compare.Here I present descriptionsand sonagramsof the song and two call types of the Golden-crowned Kinglet.

METHODS I obtained tape-recordedGolden-crowned Kinglet vocalizationsfrom three sources.First, I recordedspontaneous songs and/or calls from 12 Golden-crownedKinglets at intervalsfrom 9 May-15 Oct. 1991 at the

346 Vol.64, No. 3 KingletVocalizations [347

Evelyn and Morrill Richardson Field Station property, Bon Portage Island, Nova Scotia, Canada, and five birds on 3-8 May 1991 near Bridgewater,Nova Scotia.These recordings were madewith a SonyWM- D60 cassetterecorder and a Sony PC-62 microphonemounted in a 30- cm parabolic reflector. Individuals were not color-marked,but rather singlerecordings were made from variousbirds believedto be different individualsbecause of their locations.Second, I obtainedthe following recordingsfrom the Library of Natural Sounds,Cornell University, Ith- aca, New York: one bird recordedin Michigan 14 Jun. 1956; one bird recordedin New Jersey29 May 1959; four birds recordedin New York State 28 May 1959 and 13 Aug. 1974; two birds recordedin Maine on 17 Sept. and 8 Oct. 1981; and two birds recordedin North Carolina on 11 May and 13 Jun. 1979. The Maine recordingsand two of the New York recordingswere of callsonly, the otherswere of songs.The breeding statusof singingbirds was not definitelyknown. All recordingsof singing birds were made within the usual breedingseason, however. Third, the Borror Laboratory of Bioacoustics,Department of Zoology, Ohio State University, Columbus,Ohio, providedme with recordingsof songsfrom two birds recordedin Ohio on 10 Apr. 1955 and 18 Apr. 1971 (likely beforebreeding had commenced),and one bird from New York State on 2 Jun. 1989. Although female Golden-crownedKinglets have been re- portedto singsimilar songs to thoseof the males(Galati and Galati 1985), all songson which this paper is basedwere recordedfrom males. I spectrographicallyanalyzed 131 songsfrom 19 individualsusing a Kay ElemetricsCorp. 5500 DSP Sona-Graphand a Kay ElemetricsCorp. 5509 Grey ScalePrinter. I measuredfrequency (defined as the frequency at peakamplitude) and duration parameters from songsyllables and notes. Note and syllabletypes were then classifiedby frequency,duration and note morphology.The small number of individualsfrom most localities did not permit statisticalcomparisons among geographic areas.

RESULTS Songs.--Golden-crownedKinglet songswere composedof a seriesof syllables,most of which were repeatedseveral times. Individual syllables were sortedinto six differenttypes with little ambiguity.The mostcommon syllabletype was designatedby the letter 'A' and subsequentletters were assignedon the basisof the mostcommon heterosyllabic transitions (Fig. 1). Frequencyand duration parametersof thesesyllable types are given in Table 1. All syllabletypes were presentin all of the geographic areas sampledwith two exceptions.Syllable type F was not observedin the New York sample,and syllabletypes A and B were not observedin the New Jerseysample. This, however,may havebeen a samplingartifact due to the small number of individuals from these areas. The number of syllabletypes used and the number of repetitionsof each type varied in subsequentsongs from the same individual. The simplestsongs contained only syllabletype A. If moresyllable types were present,the order of presentationwithin a songwas highly predictable. 348] C. T. Naugler J. Field Ornithol. Summer 1993

A B c D

12- kHz

E F 6 H

12- kHz

12- kHz 8

1 2 3

s FIGURE1. (A)-(F) syllable typescomprising Golden-crowned Kinglet song,three examples for each type, each from a different individual, (G) ti call note from three individuals, (H) zeet call note from three individuals, (I) typical Golden-crownedKinglet song.All sonagramsprinted with a 234 Hz analyzingfilter. Sonagrams marked with (õ) are ¸ 1991 all rightsreserved, Library of Natural Sounds,Cornell Laboratoryof Ornithology, Ithaca,New York. Sonagramsmarked with (ô) are ¸ Borror Laboratoryof BiGacoustics, Department of Zoology, Ohio State University, Columbus, Ohio, all rights reserved. Unmarked sonagramsare from birds recordedby the author in Nova Scotia. Vol.64, No. 3 KingletVocalizations [349

TABLE1. Parametersof Golden-crownedKinglet syllabletypes. Means were calculated for eachindividual, and the meansof means(grand means)are reportedbelow. Sample size is the number birds.

Syllable type Parameter A B C D E F

Mean frequency(kHz) 7195 7665 7768 7900 7349 4731 SE 19.0 33.5 127.0 53.1 56.6 488.0 n 11 6 9 6 6 4 Mean duration (s) 0.17 0.15 0.15 0.20 0.59 0.05 SE 3.6 6.0 21.6 7.0 3.1 1.0 n 11 6 9 6 6 4

Table 2 showsthe transition probabilities(i.e., the probability that type B followstype A) for the syllabletypes in the songsI examined.Matrices calculatedfor individual birds were very similar and so only the pooled matrix is shownhere. Most syllableswere repeatedseveral times, except syllablesB and D, which were more likely to be followedby a different syllabletype than repeated(Table 2). This table alsoshows that Golden- crowned Kinglets would occasionallyskip a syllable type within a song but would rarely go back to an earlier type. For example,one bird from Bridgewater, Nova Scotia, gave five successivesongs with the syllable patternsAAAAAC, AAABCCCCEFFFF, AACCCCDDE, AAABCCC and AAAABCCC. Birds switchedto an earlier syllabletype in only four of 1155 transitionsexamined, and two of theseinstances are from a single songrendition from the New Jerseybird. This bird is alsothe only one that did not beginits songswith syllabletype A andwas the only individual

TABLE2. Transition probabilitiesfor next syllabletype in Golden-crownedKinglet songs, basedon 131 songsfrom 19 individualsfrom Nova Scotia, New York, New Jersey, North Carolina, Michigan and Ohio. Number in parenthesesis number of observed instances.Some rows do not sum to one becauseof rounding errors.

Syllable N + 1 A B C D E F

A 0.82 0.16 0.02 0 0 0 (427) (82) (13) B 0 0.31 0.57 0.12 0 0 (33) (60) (13) Syllable N C 0 0.01 0.66 0.19 0.11 0.03 (1) (127) (36) (21) (6) D 0 0 0 0.35 0.63 0.02 (16) (29) (1) E 0 0 0.01 0.01 0.75 0.24 (1) (1) (139) (44) F 0 0 0 0 0.01 0.99 (1) (104) 350] C.T. Naugler J.Field Ornithol. Summer 1993 that incorporateda zee! call into its song(see below). The incorporation of calls in songshas been previouslyreported for this species(Forbush 1929:376). The meanfrequency of syllabletypes within a songgenerally increased from syllabletypes A to B to C to D, and decreasedfrom syllabletypes D to E to F (Table 1). The structureof Golden-crownedKinglet songclosely resembles that of the congenericFirecrest of Europe, describedby Becker(1977a). The songsof both speciesconsist of groupsof repeatednotes, with remarkable similarity in note structurebetween the two species(compare Fig. 1 and Becker1977a:Fig. 2). In contrast,Goldcrest song is composedof a series of repeatedphrases each consisting of severalsyllables (Becker 1977a). Calls.--The two most commonlyobserved calls were the ti and zeet (Fig. 1). In Nova Scotia, the only location in which field observations were made, both of these calls were given by either sex. The ti note is apparentlythe tse call referredto by Galati and Galati (1985). I chose one of thesenotes at random from eight different individuals to measure frequencyand duration.This sampleconsisted of two birds from Ohio, two from New York, and four from Nova Scotia.The mean frequency was 7992 Hz (SE = 96.1), and the mean duration was 0.04 s (SE = 0.003). This call may functionas a contactnote, as it was often heard when two or more birds were together.This call has alsobeen reported to functionin parent-offspringinteractions (Galati 1991:41). The mean frequencyof zee! calls from 12 individualsfrom the samelocations was 7728 Hz (SE = 62.8), and the mean duration of the same calls was 0.18 s (SE -- 0.04). During territorial conflictsor in responseto human dis- turbance,birds oftengave zeet calls or a seriesof ti callsrapidly repeated. The "close-contactnote" and "alarm note" (Becker 1977b) of the Gold- crestand Firecreststrongly resemble the ti and zee!calls, respectively, of the Golden-crownedKinglet (Fig. 1), and appear to have similar functions.

DISCUSSION This paper presentsthe first quantitativeanalysis of vocalizationsof the Golden-crownedKinglet. The most extensivequalitative analysisto date is that of Galati and Galati (1985) and Galati (1991) on Golden- crowned Kinglets in Michigan. These authors classifiedsong as either simple or complex,depending on duration and the presenceof a lower "musical warble" at the end of the song. In the present study, songs containingnote types A to D probably correspondto the simple song reportedby theseauthors. Likewise, songs containing all notetypes probably correspondto complexsong. The functionalsignificance of song variation in this speciesis unclear.It is possible,however, that songswith differentnumbers of syllabletypes are usedin differentbehavioral con- texts.For example,Galati (1991:41-42) statesthat simplesongs are often usedin direct agonisticexchanges with anotherindividual and complex songsare usedfor "proclaimingterritory." There is oftena large amount Vol.64, No. 3 KingletVocalizations [3 51 of variance,however, in consecutivesongs from the sameindividual while presumablyengaged in the samebehavior. My observationssupport the suggestionof Galati (1991:41) that the ti call note is used as a contact call. Galati does not refer to the zeet call, however,although I found this vocalizationwas commonlyused. The similarity in structure of Golden-crownedKinglet and Firecrest songsmay indicaterelatedness between these species, although Mayr and Short (1970) suggestedthat the Golden-crownedKinglet was actually moreclosely related to the Goldcrest.With two sympatricspecies in North America and two in Europe, this genus may prove to be an interesting group in which to examine songevolution if the phylogenetichistory can be determined(see also Ingold et al. 1988). Future researchon vocalizationsof Golden-crowned Kinglets should include a more detailed analysis of among-individual song variation in specificgeographic locations, and shouldalso examine the possiblefunc- tions of within-individual songvariation.

ACKNOWLEDGMENTS

I thank L. Ratcliffe for the use of the Sona-Graph and P. C. Smith for help in the field. R. Kassen helped to analyze some of the recordings.S. A. Shackleton,K. Otter, A. Horn and three anonymousreviewers commented on earlier draftsof the manuscript.The Library of Natural Sounds,Cornell Laboratory of Ornithology and the Borror Laboratory of Or- nithology,Ohio StateUniversity providedrecordings from their collections.S. Leechassisted with manuscriptpreparation. While completingthis research,I was supportedby an NSERC PostgraduateScholarship.

LITERATURE CITED

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