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Applying citizen science data to quantify differences in song between controversial avian taxa, the sagebrush and timberline subspecies of the Brewer’s Sparrow (Spizella breweri)

Abstract

Divergence in the acoustic structure of mate-attraction signals between closely related taxa can contribute to reproductive isolation and speciation, and acoustic differences are often useful for identifying otherwise similar taxa from recordings. The Brewer’s Sparrow (Spizella breweri) is a migratory oscine songbird with 2 recently diverged subspecies. The nominate subspecies (S. b. breweri) primarily breeds in sagebrush (Artemisia spp.) shrublands in the western United States and southwestern Canada, whereas the Timberline Sparrow (S. b. taverneri) breeds in alpine shrubs and conifer krummholz near treeline in mountain ranges from east-central Alaska to northwestern Montana. Quantitative comparison of the acoustic structure of Brewer’s Sparrow mate-attraction songs is needed to help resolve the outstanding debate regarding the taxonomic status of S. b. taverneri and improve subspecific identification from song recordings. I compared the acoustic structure of 181 short song types from 180 male S. b. breweri and 22 short song types from 20 male S. b. taverneri recorded from across each subspecies’ breeding range. Despite extreme variation in acoustic structure of short songs among individuals, 19 acoustic variables differed between subspecies. Short song types of S. b. taverneri averaged 19.3%–22.1% lower maximum frequency, 4.9%–15.9% higher minimum frequency, 36.7%–42.6% narrower frequency bandwidth, and 12.9%–26.0% lower aggregate entropy (a measure of sound complexity) than those of S. b. brewer across acoustic elements (song, section, syllable, and note). Random forest classification analysis identified 10 acoustic variables important for distinguishing between short songs of the 2 subspecies and correctly classified 89.2% (95% CI 84.1% to 93.1%) of songs overall. Principal component scores representing overall acoustic structure showed an abrupt change near the breeding boundary between S. b. breweri and S. b. taverneri but overlapped sufficiently that they did not meet an established “75%–99% rule” for reciprocal diagnosability of subspecies. An easy-to-measure logistic regression threshold based on song bandwidth and mean note bandwidth accurately classified >90% of short songs to subspecies. Additional recordings of S. b. taverneri are needed to improve our understanding of geographic variation in the song of that subspecies. Data on responses of males and females to songs of the other subspecies would inform whether acoustic differences are sufficient to impede mate recognition and attraction in areas of potential breeding overlap.


La diferencia en la estructura acústica de las señales de atracción de pareja entre taxones estrechamente emparentados puede contribuir al aislamiento reproductivo y a la especiación, y las diferencias acústicas suelen ser útiles para identificar entre taxones similares a partir de grabaciones. El gorrión de Brewer (Spizella breweri) es un ave canora oscina y migratoria con dos subespecies que divergieron recientemente. La subespecie nominada (S. b. breweri) cría principalmente en matorrales de artemisa (Artemisia spp.) en el oeste de EE.UU. y el suroeste de Canadá, mientras que el gorrión de las maderas (S. b. taverneri) cría en matorrales alpinos y krummholz de coníferas cerca del límite arbóreo en cordilleras desde el centro-este de Alaska hasta el noroeste de Montana. La comparación cuantitativa de la estructura acústica de los cantos de atracción de pareja del gorrión de Brewer es necesaria para ayudar a resolver el debate pendiente sobre el estado taxonómico de S. b. taverneri y mejorar la identificación subespecífica a partir de grabaciones de cantos. Comparé la estructura acústica de 181 tipos de cantos cortos de 180 machos de S. b. breweri y 22 tipos de cantos cortos de 20 machos de S. b. taverneri grabados en toda el área de reproducción de cada subespecie. A pesar de la extrema variación en la estructura acústica de los cantos cortos entre individuos, 19 variables acústicas difirieron entre subespecies. Los tipos de canto corto de S. b. taverneri mostraron en promedio un 19.3%–22.1% menos frecuencia máxima, un 4.9%–15.9% más frecuencia mínima, un 36.7%–42.6% menos ancho de banda de frecuencia y un 12.9%–26.0% menos entropía agregada (una medida de la complejidad del sonido) que los de S. b. breweri en todos los elementos acústicos (canto, sección, sílaba y nota). El análisis de clasificación de bosque aleatorio identificó 10 variables acústicas importantes para distinguir entre los cantos cortos de las dos subespecies y clasificó correctamente el 89.2% (IC 95% 84.1%–93.1%) de los cantos en general. Las puntuaciones de los componentes principales que representan la estructura acústica global mostraron un cambio abrupto cerca del límite de reproducción de S. b. breweri y S. b. taverneri, pero se superpusieron lo suficiente como para no cumplir la regla establecida del “75%–99%” para el diagnóstico recíproco de las subespecies. Un umbral de regresión logística fácil de medir basado en el ancho de banda del canto y el ancho de banda medio de las notas clasificó con precision >90% de los cantos cortos en subespecies. Se necesitan grabaciones adicionales de S. b. taverneri para mejorar nuestra comprensión de la variación geográfica del canto en esta subespecie. Los datos acerca de las señales de respuesta de machos y hembras a los cantos de las otras subespecies podrían brindar información acerca de si las diferencias acústicas son suficientes para impedir el reconocimiento y la atracción de la pareja en zonas de potencial sobrelapamiento reproductivo.

83.4.10 Supplementary Material 1.pdf (335 kB)
Metadata for Brewer’s Sparrow (Spizella breweri) short song recordings by subspecies, source, and recording ID.

83.4.10 Supplementary Material 2.pdf (20 kB)
Pearson correlation coefficients among 10 acoustic variables measured on 181 short songs of “sagebrush” Brewer’s Sparrows (Spizella breweri breweri) and 22 short songs of Timberline Sparrows (S. b. taverneri) that were included in principal component analysis.

83.4.10 Supplementary Material 3.pdf (8174 kB)
Jittered boxplots of song-level acoustic variables measured on short WALKER ♦ SONGS OF BREWER’S SPARROW SUBSPECIES 565 songs of male “sagebrush” Brewer’s Sparrows (Spizella breweri breweri) and Timberline Sparrows (S. b. taverneri).

83.4.10 Supplementary Material 4.pdf (6033 kB)
Jittered boxplots of section-level acoustic variables measured on short songs of male “sagebrush” Brewer’s Sparrows (Spizella breweri breweri) and Timberline Sparrows (S. b. taverneri).

83.4.10 Supplementary Material 5.pdf (6033 kB)
Jittered boxplots of syllable-level acoustic variables measured on short songs of male “sagebrush” Brewer’s Sparrows (Spizella breweri breweri) and Timberline Sparrows (S. b. taverneri).

83.4.10 Supplementary Material 6.pdf (6033 kB)
Jittered boxplots of note-level acoustic variables measured on short songs of male “sagebrush” Brewer’s Sparrows (Spizella breweri breweri) and Timberline Sparrows (S. b. taverneri).

83.4.10 Supplementary Material 7.pdf (862 kB)
Relative importance and significance of 19 acoustic variables (standardized to Z scores) for correctly classifying short songs of male “sagebrush” Brewer’s Sparrows (Spizella breweri breweri) and Timberline Sparrows (S. b. taverneri) as measured by the mean decrease in accuracy and the mean decrease in the Gini index in random forest analysis.

83.4.10 Supplementary Material 8.pdf (18 kB)
Eigenvalues and percent variance explained by each dimension from principal component analysis of 10 acoustic variables measured on 181 short songs of “sagebrush” Brewer’s Sparrows (Spizella breweri breweri) and 22 short songs of Timberline Sparrows (S. b. taverneri).

83.4.10 Supplementary Material 9.pdf (1670 kB)
Scree plot showing percentage explained by each dimension in principal component analysis of 10 acoustic variables measured on 181 short songs of “sagebrush” Brewer’s Sparrows (Spizella breweri breweri) and 22 short songs of Timberline Sparrows (S. b. taverneri).

83.4.10 Supplementary Material 10.pdf (19 kB)
Rotated loadings from principal component analysis of 10 acoustic variables measured on 181 short songs of “sagebrush” Brewer’s Sparrows (Spizella breweri breweri) and 22 short songs of Timberline Sparrows (S. b. taverneri).

83.4.10 Supplementary Material 11.pdf (1682 kB)
Contributions of 10 acoustic variables measured on 181 short songs of “sagebrush” Brewer’s Sparrows (Spizella breweri breweri) and 22 short songs of Timberline Sparrows (S. b. taverneri) to the first 2 principal components in principal component analysis.

83.4.10 Supplementary Material 12.pdf (8496 kB)
Brewer’s Sparrow (Spizella breweri) short songs recorded in northwestern Montana ranked by principal component (PC1) score. Map inset shows the location in relation to state and provincial boundaries.

83.4.10 Supplementary Material 13.pdf (1108 kB)
Histograms of principal component (PC1) scores representing the overall acoustic structure of Brewer’s Sparrow (Spizella breweri) short songs by subspecies.

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