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Assessing Vocal Variety in the Winter Wren, a Bird with a Complex Repertoire’

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Assessing Vocal Variety in the Winter Wren, a Bird with a Complex Repertoire’ The Condor 923949 0 The Cooper OrnithologicalSociety 1995 ASSESSING VOCAL VARIETY IN THE WINTER WREN, A BIRD WITH A COMPLEX REPERTOIRE’ BEATRICE VAN HORNE Department of Biology, ColoradoState University,Fort Collins, CO 80521 Abstract. The song of the Winter Wren (Troglodytestroglodytes) varies in a complex fashion in both time and space.An averageof 2 1 songtypes occurredin males recordedon at leasttwo mornings;these song types included many variants. Syllableswithin songswere not arrangedrandomly, as thosethat belongto certain syllabletypes based on acousticshape were more likely to be usedin the introductoryportion of the song.Probabilities of transition among syllable types for consecutivesyllables were uneven but consistentbetween years. The repertoireof songtypes is probably not fixed but varies in its constituentsong types both through the breedingseason and between years. Key words: Song structure;song complexity; song repertoire; Winter Wren; Troglodytes troglodytes. INTRODUCTION period (day, season, lifetime) is specified, rep- The complexity of bird song contributes to its ertoire size is difficult to estimate, as the prob- aestheticappeal to the human ear. Evolutionists ability of singing a given song type is often not and behaviorists have constructedhypotheses to independent of bout context [see Kroodsma account for differencesin songcomplexity within (1982)]. (Lambrechts and Dhondt 1988, Catchpole 1980) My objective here is to illustrate problems in and sometimes among (Searcyand Marler 1984) measuring complexity in the songsof a complex species. The notion that increased complexity songsterusing Winter Wrens, Troglodytestrog- has an evolutionary cost is implicit in evolu- lodytes, as an example. To characterize song tionary explanations. structuring, I describe patterning through time, If hypothesesabout song complexity are to be both within and among songs and within and tested, the term “complexity” requires a quan- among bouts. I then abstract several measures tifiable definition. However, the term has been of within-song complexity, among-song com- used variably in the avian acoustical literature. plexity, and complexity within a recording ses- It may represent the total number of different sion or within consecutive sessions.I also de- song types or song-type elements produced over scribe how the repertoire changeswithin season a specifiedperiod, or it may include the pattern- and between years, and how this change might ing of these elements within or among songs. intluence the description of complexity. Mathematically, we may define complexity as METHODS the inverse of predictability. That is, complexity should increase as the sum of the squared prob- CONTEXT OF RECORDINGS abilities of subsequentevents (song types or syl- The study area included 4 ha of early succes- lables) decreases.It is thus a function of the num- sional coastal spruce-hemlock forest (Greene ber of songsor syllables used, their “evenness” 1982) and an adjacent lo-ha stand of mature in frequency of occurrence, and the existence of timber in the CascadeHead Experimental Forest rules or constraints for patterning through time of the U.S. Forest Service (Siuslaw National For- (structuring). Most theory about songcomplexity est, Oregon). During March of 1984 and 1985, pertains to repertoire size, i.e., among-song com- singing male Winter Wrens were captured in mist plexity. Within-song complexity, however, de- nets and uniquely marked with a combination termines the amount of variation available for of three plastic colored leg bands and an alu- among-song complexity. Additionally, when the minum band. On mornings without rain I re- corded the songsof a single male. I usually began recording during the first songbout of the mom- ing (one bout includes all consecutive repetitions ’ ’ Received 3 June 1994. Accepted 24 August 1994. of one or more songs without a separation of 1391 40 BEATRICE VAN HORNE more than 30 set) and continued until I had re- added to the master catalogue. Data from each corded at least 100 songsor until the male began song was entered into a computer file as a single foraging, engagingin courtship, or was otherwise observation that included its duration, the in- interrupted. At least part of this singing occurred terval between it and the previous song, and the before there was adequate light for foraging, so numeric code sequence identifying consecutive it was not likely to be interrupted by foraging syllables along with the number of consecutive behavior. When recording sessionswere inter- repetitions of those syllables. Becauseintroduc- rupted by rain, I continued them on a second tory notes were often sung more softly than the and occasionally a third morning. Songsthat pro- remainder of the song, I used the best represen- duced unclear sonograms were not included in tation of the notes to identify the introductory the analysis. I attempted to record each male at syllables and assumed that repetitions of that least twice, separatedby at least two weeks, be- song contained the same introductory notes. fore he began tending nestlings (usually in late Although there were too many different syl- April or early May). In some cases the males lables for analysis of transitions between sylla- disappeared before a second recording could be bles, it seemed from visual inspection that there made. might be rules governing placement of certain Singing males were classified into 1 of 5 phe- shapesof syllables within the song. Consequent- nological stageson the day of each recording ses- ly, after all syllables had been identified, the syl- sion: 1 = prior to egg-laying by any females on lable catalogue was used to assign each syllable the territory, 2 = during laying by the primary to one of 11 groups, based on the general shape or first female on the territory, 3 = during in- of the syllable and ignoring its frequency (e.g., cubation by the primary female, 4 = nestlings in inverted ‘Iv,” simple ascending,“m” shape,etc.). the nest of any female on the territory, and 5 = Using the terminology of Lynch et al. 1989, I after young had fledgedfrom all nests or all nests refer to these groups of like syllables as “syllable had been desertedby both parents. Where exact types.” Songswere re-written to a new computer dates of a given nesting event were not known, file with syllable type numbers substituted for mean values for these events over the course of syllable numbers. This file was used to analyze the three-year study were used, seven days for the locations of syllable types in songs and the laying, 14 days for incubation, and 16 days for frequency of transitions among syllable types the nestling period. within songs. All transitions among syllable types in each RECORDING OF VOCALIZATIONS AND songwere summed acrossall birds within a given DESCRIPTION OF STRUCTURE year to obtain a matrix whose rows represented WITHIN SONGS the number of all transitions from each possible In 1984, songs were recorded on a Uher 4000 syllable type, and whosecolumns representedthe Rep-L reel-to-reel recorder with a Sennheiserdi- number ofall transitions to each possiblesyllable rectional microphone (K3-U/ME 80); in 1985 a type. Thus, the row and column totals repre- Sony Pro Walkman WM-D6C cassetteplayer was sentedthe number oftimes syllablesfrom a given used. Songswere digitized using a UNISCAN II syllable type were sung, lessthe number of times real time analyzer and printed on a dot matrix these were the ending and beginning syllables in printer. Songswere defined as vocalizations that songs,respectively. contained no gapsof ~0.5 set; gapswithin songs To investigate whether there were “rules” of were always much smaller than this (see Fig. 1). transition among syllable types, independent of Within a song,a syllable was a single continuous the commonness of the syllable types, I com- mark on the sonograph. Occasionally, normally pared observed patterns of transition to random continuous marks appeared to separateinto syl- patterns of transition with the same frequencies lables in certain recordings; these retained their of occurrence of the syllable types. I constructed classification as single syllables. Syllables were random matrices by beginning with an empty assignednumbers using transparencies of pages matrix of the same dimension as the observed of a master catalogue that included all syllables matrix for that year. A cell was randomly selected arranged by shape and frequency. Syllable iden- and a single transition added to it. For example, tification was thus independent of context. Each the cell selectedat random might representtran- new syllable was given a unique number and sitions from syllable 3 to syllable 11, and it would WINTER WREN VOCAL VARIETY 41 (ZHY) kmnbaq 42 BEATRICE VAN HORNE be incremented by 1. This processwas repeated, for preferred “blocks” of syllables, as any such with transitions being added one at a time (unless blocks would be broken and joined in a nonran- the addition would violate row or column totals, dom fashion as new variants of the song were which represented syllable type commonness) introduced. That is, a variant of a songmay have until the total number oftransitions in the matrix a new block of syllables inserted or substituted matched the total number of transitions in the for the ending of the original song, while a sub- observed matrix for that year. sequent variant may use part of that block of A measure of dissimilarity between matrices syllables combined with part of another block of was obtained using a modified distance measure: syllables that had been used previously. In es- sence,the patterning through time can be likened to a very complex version of the sequencecircles used by Bakker et al. (1983) to describeBobolink (Dolichonyx oryzivorus) song, although Winter where x and y are the cells of transition matrices Wren songsare more discrete. X and Y, respectively, and N is the number of syllable types in each matrix. D values of pairs This pattern, with its insertions, substitutions, consisting of 1 random and 1 observed matrix and deletions, is analogous to the structuring of were compared to D values for pairs of random long strings of DNA and RNA.
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