Stereotyped and Plastic Song in Adult Indigo Buntings, Passerina Cyanea
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Anim.Behav., 1991, A, 367-388 Stereotypedand plastic song in adult indigobuntings, Passerina cyanea DANIEL MARGOLIASH, CYNTHIA A. STAICER* & SUE A. INOUET Departmentof OrganismalBiology and Anatomy , The Universityof Chicago,Chicago, IL 60637, U.S.A. ( Received18 July 1990;initial acceptance4 September 1990; rtnalacceptance 10December 1990; MS. number t5835 ) Abstract.The vocal behaviourof indigo buntingswas studiedby monitoring individual malesin sound isolation boxeswith a computer-assistedsystem. Yearlings and older adults sang'adult plastic' songas well asstereotyped song. Distinguishing features ofadult plasticsong include: a largersyllable repertoire; variablesyllable order; emphasis of the syllablesthat appearin the terminalsequence of stereotypedsong; 'S-note' a high-frequency ofvariable morphology;calls; occasional production ofsyllables poorly formed, of intermediateform, or in combination;and low amplitude of delivery.Stereotyped and adult plastic songswere delivered in distinct bouts.Adult plasticsongs of indigo buntingsshare many of the character- istics ofearly plastic songsofyoung birds. Severalother extended-agelearners also exhibit a complex, variable,and low amplitudeadult singingbehaviour, sometimes described as subsong, in addition to loud stereotypedsong. Indigo buntings,and other speciesthat can learn new songsas adults,may retain into adulthood developmentalprocesses that in most specieslargely terminate with the onsetof stereotyped song. A dramatic event during song developmentin Thompson 1968;Williams & MacRoberts 1977; passerinebirds is the rapid transition from juvenile Feekes 1982). Such birds may require auditory plastic singing to a stereotyped,adult pattern feedback to maintain their songs (Nottebohm (reviewedin Marler & Peters 1982a).This tran- 1980).The similaritiesbetween juvenile and adult sition has important behaviouralimplications, plasticsong, and the possiblerole of adult plastic includingthe ability to competefor matesand terri- song in adult song learning have not beenexten- tories(e.g. McDonald 1989),as well as profound sively analysed.Because a bird with vocal experi- physiological implications, especially emanci- ence has developed an internal representation pation from auditory feedback(Konishi 1965a). betweenvocal gestureand produced sound, and Much of the vocalmaterial present in juvenilesing- has establisheda repertoire of sounds,this may ing is no longer expressedafter the emergenceof effect both the rate and process of subsequent crystallizedsong (Marler & Peters1982b). Whether songlearning. this material is completely lost remains unclear, To addressthese issues, we choseto studyindigo however,because birds often exhibit abbreviated buntings.This specieshas been the focusof exten- statesof variablesinging as they re-establishadult sivestudies examining the basicsinging behaviour, song in subsequentsprings (Marler 1956, 1970; song learning, adaptive significanceand cultural Mulligan 1966). evolution of song (Thompson 1972;Emlen 1972; The descriptionof song developmenthas been Payneet al. I 98l). In particular,yearlings (birds in largely derived from work with age-dependent their secondcalendar year of life) and, to a lesser learners,birds that usually do not change their extent,older adult birds sometimesengage in 'song songsas adults. In birds that modify song reper- matching' (Thompson 1970;Payne et al. 1981, toires as adults, there is evidencethat the plastic 1988).During songmatching a bird cancompletely song stageis extendedinto adulthood so that two changehis stereotypedsong to closelymimic the distinct types of song are maintained (Rice & song of a neighbouringadult (Payne l98l). This leadsto groupsof2-22 birds sharingthe samesong *Present address:Department of Biology, Dalhousie University,Halifax, Nova ScotiaB3H 4JI, Canada. (Payneet al. 1981,1988). fPresent address:712 Rose Lane, Matteson,lL 60M3, The singing of male indigo buntings has been U.S.A. classifiedinto threemajor typesof song.Territorial M03-3 472 I 9 | I 090367+ 22 503.00I 0 @l99l TheAssociation for theStudv of AnimalBehaviour 6 Z ″筋 α′』夕れαッわ″″,42 ,ヨ adultssing relatively simple stereotyped songs (also this time, birds caught as immatureswere in th€ir calledthe'advertising' song; Thompson 1968) and secondcalendar year and sowere called yearlings. occasionally sing more complex, longer songs. Sixadditional males, four yearlingsand two older Theselonger songs often incorporateextra syllable adults,were collected on 6-8 June 1989near Niles, typesbeyond those expressed in stereotypedsong. Michigan.The songsof all 1989birds were recorded Indigo buntings also sing low amplitude, long in thefield with standardtechniques prior to capture songswith 'squeaky'characteristics during staged (seePayne 1982).These birds were immediately territorial encounters (Emlen 1972; Thompson returned to the laboratory, housed in individual 1972).Squeaky songs are difficult to record in the sound isolation boxes and monitored. For 1989 field and have not beenextensively studied. Here, birds,the first songswere recorded within aslittle as we report that isolatedindigo buntingsin the lab- 3 daysofcapture, oratory commonly and spontaneouslysing an All birdsin IAC boxeswere housed in'standard' apparentlymore variable form of squeakysong. wire cagesmeasuring 45 x 35 x25 cm and main- Our analysisindicates that thesesongs have many tained on a 16:8h light:dark cycle. Any residual featuresin commonwith earlyplastic songs of criti- acoustic cross-talk betweenbooths during loud cal period learners,and we term it'adult plastic' vocalizations was masked by wide-band noise song.We note the similarity betweenthis and vari- ( - 80 dB) broadcastintothe room. Somebirds sang ous secondarysinging behaviour described for spontaneously,otherscame into full songonly after other speciesand discussthe possiblerole adult receivingtestosterone implants. The 1987 birds, plastic songmay play in the modification of adult movedfrom shorterdays (in the generalaviary) to stereotypedsongs. longerdays (in IAC boxes),werephotosensitive and presumablymaximally sensitiveto the effectsof METHODS testosteroneon singingrate, while the 1989birds, which had already experiencedlong days in the Bird Collection field, were presumably less sensitive (Nowicki & Two groupsof indigo buntingswere collected by Ball 1989).We did not analyseany songsfor 1987 mistnets.Twenty birdswerecollectedin Mississippi birds until a bird had achievedfull stereotyped during the October 1987southward migration, and songwhereas we recordedall songsfor 1989birds are termed 1987 birds. We collected 'immature' immediately after captur€. After recording indi- birds (a few months old), 'yearlings'(birds in their vidually isolated1987 birds, l0 socialpairings were secondcalendar year of life), and 'older adults' establishedthroughout the spring/summerof 1988 (birdsin their third calendaryear or later).These age by housing pairs in half-cages(a standard cage groupsare easilydistinguished on the basisofplu- divided in half by a hardwarecloth partition with mage,with yearlings(and Octoberimmatures) but I x I cm mesh) within IAC boxes. One pair of not older adults, exhibiting some brown feathers birds interactedvia an acousticlink but remained amongthe wing (primary)coverts (see Payne 1982). physicallyisolated in separateIAC boxes.Social Becausebuntings isolated during their first yearcan pairings were maintained up to severalmonths, developabnormal songs(Rice & Thompson 1968; after which birds wereisolated and recordedagain. Paynel98l), the1987 birdswerehousedthroughout Someindividuals were socially paired two or three theautumn of 1987inindividual wire cagesmeasur- times. In this paper, we report on the singing ing45 x 35 x 45 crn.The cages were kept in a general behaviourofisolated birds. Changes in songduring aviary where the birds could hear each other, and social pairings will be reported in a subsequent interactwithneighbours in adjacentcagesseparated paper. by 5 or more cm. The generalaviary, which could hold up to 75cages, also housed zebra finches, Tae- Song Recordingand Display niopygia guttam, and white-throated sparrows, Data Zonotrichia albicollis, and was maintained on a collection natural Chicago photoperiod. From January to Vocalizations were recorded by microphones August of 1988,small groups of thesebirds were (Realisticno. 33-201l) insidethe soundboxes, pro- movedinto soundattenuation booths (IAC no. AC- cessedlocally (AC coupling and gain), and driven l , with interior dimensionsof 57'5 x 40 x 35 crn)and differentially over 30-m cablesto the laboratory. In theirvocalbehaviourwasmonitored (see below). By the laboratory, the signalwas processed with four- Margoliash et al.: Adult plastic song in buntings 369 pole lowpassand highpassfilters, achievinga fre- coupled to a vector processor(CSPI Mini-Map quencyresponse ofthe electronicsof * 0.5dB from XL). Spectrogramswere generatedwith standard 1-9'5 kHz, and -24 dB at 500Hz and 20 kHz. The digital signalprocessing techniques (Oppenheim & incomplete high-frequency filtering resulted in Schafer 1975).Depending on choice of analysis someartefacts when the signalswere digitized (see parameters,it waspossible to createa displaysimi- below). Calibration was via a l-inch (2.5-cm) lar in physicalsize and quality to that producedby a Bruel & Kjaer microphone(no. 4145),a Hewlett- Kay analogSonagraph, or to displayspectrograms Packardwave analyser (no. 3581C),and computer with up to the full dynamicrange of the converter analysis.The frequencyresponse