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Vocal Behaviour of White-Eared Ground-Sparrows (Melozone Leucotis) During the Breeding Season: Repertoires, Diel Variation, Beha

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Vocal Behaviour of White-Eared Ground-Sparrows (Melozone Leucotis) During the Breeding Season: Repertoires, Diel Variation, Beha Author's personal copy J Ornithol DOI 10.1007/s10336-015-1237-y ORIGINAL ARTICLE Vocal behaviour of White-eared Ground-sparrows (Melozone leucotis) during the breeding season: repertoires, diel variation, behavioural contexts, and individual distinctiveness 1,2 2 1 Luis Sandoval • Carolina Me´ndez • Daniel J. Mennill Received: 7 March 2014 / Revised: 9 October 2014 / Accepted: 13 May 2015 Ó Dt. Ornithologen-Gesellschaft e.V. 2015 Abstract There are relatively few quantitative descriptive variation in each category of vocalisation, and found that the studies of the vocalisations and vocal behaviour of tropical Ground-sparrows produced all three vocalisations at higher bird species, in spite of the tropic’s rich avian biodiversity output at dawn (between 0500 and 0600 hours) compared to and the extensive variety of vocalisations produced by the rest of the day. This study allowed us to conduct the first tropical birds. This lack of information inhibits our under- comparisons of vocalisations between White-eared Ground- standing of tropical animals, including our ability to perform sparrows and North American species in the genus Melo- comparative analyses on vocal behaviours from an evolu- zone, and revealed both similarities and differences between tionary perspective. In this study, we present the first quan- the species groups. Our investigation also showed that titative description of the vocal repertoire and daily vocal vocalisations related to communication within pairs and to activity of White-eared Ground-sparrows (Melozone leuco- territory defence (calls and duets) exhibited lower levels of tis), using focal and autonomous recordings collected during individual distinctiveness than vocalisations related mainly two consecutive breeding seasons in Costa Rica. We clas- to female attraction (male solo songs). Our observations sified vocalisations into categories based on their visual suggest that each of the three types of vocalisations have appearance on sound spectrograms to create a library of multiple functions in White-eared Ground-sparrows, vocalisations for this species. We found that White-eared revealing diverse communication functions with a small Ground-sparrows produce three main categories of vocali- vocal repertoire in this tropical songbird. sations: solo songs, calls, and duets. Solo songs were pro- duced only by males. Each male sang a repertoire of solo Keywords Calls Á Diel variation Á Duets Á Emberizidae Á song types, which all shared the same general structure with Songs Á Tropics short introductory notes, a frequency-modulated middle section, and a terminal trill. Both sexes produce calls and Zusammenfassung coordinated vocal duets. We quantified patterns of diel Gesangsverhalten von Weißohrammern (Melozone leu- cotis)wa¨hrend der Brutzeit: Repertoires, Tages- Communicated by C. G. Guglielmo. variation, Verhaltenskontext und individuelle Charak- teristik Electronic supplementary material The online version of this article (doi:10.1007/s10336-015-1237-y) contains supplementary material, which is available to authorized users. Trotz des Vogelartenreichtums der Tropen und der großen Vielfalt an Lauta¨ußerungen tropischer Vo¨gel, gibt es & Luis Sandoval verha¨ltnisma¨ßig wenige quantitativ-beschreibende Studien zu [email protected] Vokalisation und Gesangsverhalten tropischer Vogelarten. 1 Department of Biological Sciences, University of Windsor, Dieser Mangel an Informationen vermindert unser 401 Sunset Avenue, Windsor, ON N9B3P4, Canada Versta¨ndnis tropischer Vo¨gel, einschließlich der Fa¨higkeit, 2 Laboratorio de Bioacu´stica Escuela de Biologı´a, Universidad vergleichende Analysen zum Gesangsverhalten aus Sicht der de Costa Rica, San Pedro, Montes de Oca, Costa Rica Evolution durchzufu¨hren. In dieser Studie pra¨sentieren wir 123 Author's personal copy J Ornithol erstmals eine quantitative Beschreibung des Stimmenreper- species. Moreover, with the recent importance assigned to toires und der ta¨glichen Gesangsaktivita¨t von Weißohram- vocalisations as a tool for resolving taxonomic issues mern (Melozone leucotis) mittels gezielter und unabha¨ngiger between closely related taxa (Price and Lanyon 2002; Aufnahmen, die in zwei aufeinander folgenden Brutsaisons in Stiles 2009; Cadena and Cuervo 2010; Millsap et al. 2011), Costa Rica gesammelt wurden. Wir teilten die Lauta¨ußerun- studies of this type are becoming increasingly informative. gen in Kategorien ein, um auf Basis des Aussehens der Our restricted information on the vocal behaviour of birds Klangspektrogramme eine Sammlung der Lauta¨ußerungen also limits or delays the ability to perform comparative fu¨r diese Art anzulegen. Weißohrammern erzeugen drei analyses on vocal behaviours that could enhance our Hauptkategorien von Lauta¨ußerungen: Sologesang, Rufe und understanding of taxonomic relationships (e.g. Price and Duette. Sologesa¨nge werden ausschließlich von Ma¨nnchen Lanyon 2002; Cadena and Cuervo 2010) and facilitate erzeugt. Jedes Ma¨nnchen sang ein Repertoire aus Sologe- better modeling of the evolution of diversity in tropical sangstypen, die alle die gleiche Grundstruktur hatten mit birds (Price et al. 2011). For example, if singing behaviour, kurzen Einleitungsto¨nen, einem frequenzmoduliertem Mit- repertoire size, and patterns of diel variation are consistent telteil und einem abschließendem Triller. Beide Geschlechter between individuals across a species distribution, this may erzeugen Rufe und aufeinander abgestimmte Stimmduette. reflect close genetic relationships (Price and Lanyon 2002; Wir quantifizierten Muster in der ta¨glichen Variation der Cadena and Cuervo 2010). einzelnen Lauta¨ußerungskategorien. Dabei konnten wir fest- In this study, we present the first quantitative description stellen, dass Grundammern in der Morgenda¨mmerung (zwi- of the vocal repertoire, singing behaviour, and pattern of schen 5:00 und 6:00) eine ho¨here Gesangs- und diel variation in White-eared Ground-sparrows (Melozone Stimma¨ußerungsleistung erbringen im Vergleich zum Rest leucotis), a species found between 500 and 2000 m a.s.l., des Tages. Mit Hilfe dieser Studie ko¨nnen die ersten Ver- from Chiapas, Mexico, in the north, to Costa Rica’s Central gleiche von Lauta¨ußerungen zwischen Weißohrammern und Valley in the south (Stiles and Skutch 1989; Howell and anderen nordamerikanischen Arten der Gattung Melozone Webb 1995). Across their disjunct distribution, this angestellt werden, um sowohl Gemeinsamkeiten als auch Ground-sparrow inhabits thickets, secondary forest edges, Unterschiede zwischen den Artengruppen aufzuzeigen. and shaded coffee plantations (Stiles and Skutch 1989; Unsere Untersuchungen zeigen auch, dass Lauta¨ußerungen Howell and Webb 1995; Sandoval and Mennill 2012). im Zusammenhang mit der Kommunikation zwischen Brut- White-eared Ground-sparrow pairs use vocalisations to partnern und zur Revierverteidigung (Rufe und Duette) ein defend their territories throughout the year (Sandoval and geringeres Maß an individueller Charakteristik aufweisen, als Mennill 2012; Sandoval et al. 2013). They are one of seven solche, die vor allem der Weibchenanwerbung (Sologesa¨nge recognized species in the genus Melozone (Chesser et al. der Ma¨nnchen) dienen. Unsere Beobachtungen lassen erken- 2010), and one of three species in the genus where our nen, dass jeder der drei Lauta¨ußerungstypen verschiedene knowledge of vocal behaviour is restricted to brief and Funktionen bei Weißohrammern hat, was wiederum darauf anecdotal descriptions (Stiles and Skutch 1989; Howell and hin deutet, dass das kleine Stimmenrepertoire eines tropischen Webb 1995; Sandoval and Mennill 2012; see Sandoval Singvogels diverse Kommunikationsfunktionen haben kann. et al. 2014, for an exception). Historically, the Melozone genus included only three Mesoamerican species—Pre- vost’s Ground-sparrow (M. biarcuata), Rusty-crowned Ground-sparrow (M. kieneri), and White-eared Ground- Introduction sparrow (M. leucotis)—but the taxonomy has recently been changed as a result of genetic studies (DaCosta et al. 2009; A vocal repertoire is the complete library of vocalisations an Chesser et al. 2010; Klicka et al. 2014). Four Towhee individual or species can produce (Catchpole and Slater species formerly categorized as part of the Pipilo genus 2008). Oscines learn their vocalisations, and the result of this (Tweit and Finch 1994; Howell and Webb 1995; Johnson learning process is that each population or individual produces and Haight 1996; Benedict et al. 2011) are now classified new vocal characters by learning, making mistakes, or as Melozone: Abert’s Towhee (M. aberti), California improvising (Beecher and Brenowitz 2005). Oscines in the Towhee (M. crissalis), Canyon Towhee (M. fusca), and tropics exhibit several vocal behaviours that are rare or absent White-throated Towhee (M. albicollis). All four of these in the north temperate zone, including female solo songs species inhabit dense brush and understories as well as (Langmore 1998; Beecher and Brenowitz 2005), duets (Hall areas of semi-arid montane shrubland in Mexico and the 2004, 2009), and choruses (e.g. Baker 2004, 2009;Hale2006). southeastern United States (Tweit and Finch 1994; Howell The number of quantitative descriptive studies that have and Webb 1995; Johnson and Haight 1996; Benedict et al. been conducted on the vocalisations of tropical birds is 2011). The three Mesoamerican Ground-sparrows inhabit limited considering the impressive diversity of tropical bird mainly dense habitats such as thickets, young secondary 123 Author's
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