Song Variation in a Population of Alder Flycatchers
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J. Field Ornithol. 75(2):146±151, 2004 Song variation in a population of Alder Flycatchers Scott F. Lovell and M. Ross Lein1 Department of Biological Sciences, University of Calgary, Calgary, Alberta T2N 1N4, Canada Received 17 February 2003; accepted 30 April 2003 ABSTRACT. Song variation has been studied extensively over the past 50 yr but almost entirely in oscine passerines. Although learning is an essential component of song development in most, if not all, oscines, there is no de®nitive evidence for song learning in suboscine passerines. This suggests that the patterns and extent of individual and geographic variation may differ between these groups as well. We examined individual variation in the ``fee-bee-o'' song in a population of Alder Flycatchers (Empidonax alnorum) in southwestern Alberta. Songs of individual males were recorded during the breeding season in 2001. We measured temporal and frequency variables of songs and conducted univariate and multivariate statistical analyses to characterize variation within and among individuals. There was little variation among the songs of an individual male during single recording sessions and across recordings made over the breeding season. All measured variables varied signi®cantly more among than within individuals. Discriminant function analysis assigned 91±100% of songs to the correct individual. Therefore, there was suf®cient variation among individual males to identify them statistically and, potentially, to permit individual discrimination by the birds. SINOPSIS. VariacioÂn en el canto de una poblacioÂn de Empidonax alnorum La variacioÂn en el canto de las aves ha sido estudiada extensivamente en los ultimos 50 anÄos, particularmente en paserinos oscinos. Aunque el aprendizaje es un componente esencial en el desarrollo del canto de la mayorõÂa, si no en todos los oscinos, no hay evidencia para el aprendizaje del canto en paserinos suboscinos. Esto sugiere que el patroÂn y la amplitud de la variacioÂn individual y geogra®ca pudiera ser diferente entre estos dos grupos. Examinamos la variacioÂn individual en el canto ``®i-bii-o'' de una poblacioÂn de individuos de Empidonax alnorum en el suroeste de Alberta. Se grabo el canto de individuos particulares durante la temporada reproductive de 2001. Medimos variables temporales y de frecuencia y utilizamos anaÂlisis estadõÂstico univariable y multivariable para caracterizar la variacioÂn en y entre individuos. Se encontro muy poca variacioÂn entre individuos machos en una grabacioÂn hecha en un momento dado y en grabaciones obtenidas lo largo de la eÂpoca reproductiva. Todas las variables estudiadas, variaron sign®cativamente maÂs entre individuos que en individuos particulares. El anaÂlisis discriminativo de fun- ciones asigno de 91±100% de las canciones al individuo correcto. Por tanto, hubo su®ciente variacioÂn entre indi- viduos machos para poder identi®carlos, utilizando estadõÂsticas, y potencialmente para permitir que las propias aves puedan discriminar entre individuos. Key words: Alberta, Empidonax alnorum, individual variation, song, suboscine The avian order Passeriformes includes ap- in suboscines (Kroodsma 1984). A potential ef- proximately 5700 species or about 60% of liv- fect of learning versus non-learning modes of ing species of birds (Sibley and Monroe 1990). song development may be greater geographic Suboscine passerines (suborder Tyranni) make and individual variation in song structure ex- up approximately 20% (1151 species) of the hibited by oscines than by most suboscines order and are a dominant part of Neotropical (Kroodsma 1996). avifaunas (Sibley and Monroe 1990). Despite Although little, if any, geographic or individ- this, our knowledge of passerine song and its ual variation has been described in songs of functions is biased heavily toward studies of os- suboscines (Stein 1963; Lanyon 1978; Payne cine passerines (suborder Passeri). However, it and Budde 1979; Johnson 1980), visual in- is known that species in the two suborders ap- spection of published audiospectrograms (e.g., parently differ in mechanisms of song devel- Stein 1963; Payne and Budde 1979; Kroodsma opment. Learning and auditory feedback play a 1984) suggests measurable variation. However, major role in the development of songs in all large sample sizes of songs and quantitative oscine passerines studied (33 families; Kroods- analyses have been lacking in most studies of ma 1982, 1996; Slater 1989). In contrast, song variation in suboscine passerine songs (see ontogeny is apparently genetically-programmed Sedgwick 2001 for an exception). Individual variation in the vocalizations of 1 Corresponding author. Email: [email protected] birds has been documented in many species (re- 146 Vol. 75, No. 2 Song Variation in Alder Flycatchers 147 Fig. 1. Audiospectrogram of a ``fee-bee-o'' song of the Alder Flycatcher. Lines and symbols indicate time (T) and frequency (F) measurements. See Appendix 1 for explanation of measurements. viewed in Beer 1970; Falls 1982). The few song repertoires consist of one relatively simple studies that have attempted to characterize in- song type (Fig. 1) described verbally as ``fee-bee- dividual variation in songs of suboscines (Kel- o'' (Stein 1963). logg and Stein 1953; Payne and Budde 1979; Individual songs were acquired from the Kroodsma 1984) did not measure enough original tapes as digital sound ®les using RTSD songs to describe variation within and among Ver. 1.10 and SIGNAL Ver. 3.1 bioacoustical individuals in a population adequately and thus software (Engineering Design, Belmont, MA) were not able to provide a comprehensive quan- with a sample rate of 25 kHz. We wrote a pro- titative analysis of variation. We recorded and gram in the SIGNAL language to assist in mea- analyzed songs of Alder Flycatchers (Empidonax surement of 12 temporal variables and six fre- alnorum) to determine whether variation was quency variables for each ``fee-bee-o'' song (Fig. suf®ciently restricted within individuals, but 1; measurements de®ned in Appendix 1). Tem- suf®ciently great among individuals, to permit poral variables were measured on wideband us to identify individuals by song. spectrograms with a resolution of 2.6 ms. Fre- quency variables were measured on a spectral METHODS contour generated from the spectrogram. Spec- tral contours track the frequency with the max- Songs of 13 territorial male Alder Flycatchers imum sound energy at a given time. This pro- were recorded during the breeding season of cedure avoided the subjectivity associated with 2001 at Bryant Creek (518029N, 1148479W) in measurement of frequency on a wideband spec- the foothills of the Rocky Mountains, approx- trogram. Two data sets were generated. The ®rst imately 80 km west of Calgary, Alberta. Alder included measurements of 10 songs recorded Flycatchers breed in willow (Salix spp.) and bog on a single day for each male (within-recordings birch (Betula glandulosa) thickets bordering data set). The second included measurements streams and beaver ponds in the area. Ten males of single examples of songs from each of a series were captured using mist nets, banded, and of recordings made on 4±9 different dates for marked with hair dye on their breast feathers each male (among-recordings data set). for individual identi®cation. Three unmarked SYSTAT 9.0 software (SPSS Inc., Chicago, individuals were identi®ed by territory position IL) was used in all statistical analyses. We used and their persistent use of speci®c song perches. coef®cients of variation (CV) to quantify the Songs were recorded between 03:00 to 08:00 magnitude of variability in each measured var- MST using a Nagra 4.2 reel-to-reel tape re- iable in the within-recordings data set. We cal- corder and a Sennheiser K6-P microphone in a culated within-male coef®cients of variation Telinga parabolic re¯ector. Alder Flycatchers (CVw) for each male to measure the amount of J. Field Ornithol. 148 S. F. Lovell and M. R. Lein Spring 2004 Fig. 2. Audiospectrograms of ``fee-bee-o'' songs of Alder Flycatchers indicating constancy of songs within individuals and variation among individuals. Each row has two songs of an individual. Note that examples for males FISP and WH2O were recorded in different years. variation during a recording session and calcu- si®cation technique and as a way to cross-vali- lated among-male coef®cients of variation date our results (Manly 1994). We also com- (CVa) from the variable means for each male. bined the two data sets into a cross-classi®ca- An ANOVA was conducted on each of the 18 tion data set, conducting an additional DFA to variables in the within-recordings data set to determine whether discriminant functions de- compare within-male and among-male variabil- rived from the among-recordings data set could ity in these variables. correctly identify the songs in the within-re- Many of the variables measured were corre- cordings data set. lated with one another. To reduce the number of variables and to eliminate problems caused RESULTS by colinearity of variables, we performed a prin- cipal components analysis (PCA) on each data Visual inspection of audiospectrograms sug- set. We then used discriminant function anal- gests differences in timing, ``shape,'' and pres- ysis (DFA) of principal component scores of ence or absence of different elements that char- songs in each of the two data sets to determine acterize songs of different individuals (Fig. 2). whether songs of individuals could be identi®ed Values of mean CVw ranged from 0.9% for reliably. Results of jack-knifed classi®cations, in maximum frequency of the terminal note of the which each song was classi®ed using discrimi- ``fee-bee-o'' song (F6) to 57.8% for rate of fre- nant functions computed from all songs in the quency change in the ``fee'' introduction (F2). data set except the case being classi®ed, are re- Values of CVa ranged from 5.2% for the max- ported as percentages of songs classi®ed cor- imum frequency of the ``fee'' chevron (F3) to rectly.