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Typical Males and Unconventional Females: Songs and Singing Behaviors of a Tropical, Duetting Oriole in the Breeding and Non-Breeding Season

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Typical Males and Unconventional Females: Songs and Singing Behaviors of a Tropical, Duetting Oriole in the Breeding and Non-Breeding Season ORIGINAL RESEARCH published: 25 February 2016 doi: 10.3389/fevo.2016.00014 Typical Males and Unconventional Females: Songs and Singing Behaviors of a Tropical, Duetting Oriole in the Breeding and Non-Breeding Season Karan J. Odom 1*, Kevin E. Omland 1, David R. McCaffrey 2, Michelle K. Monroe 1, Jennifer L. Christhilf 1, Natalie S. Roberts 1 and David M. Logue 3 1 Department of Biological Sciences, University of Maryland, Baltimore County, Baltimore, MD, USA, 2 Department of Geography, University of Lethbridge, Lethbridge, AB, Canada, 3 Department of Psychology, University of Lethbridge, Lethbridge, AB, Canada Edited by: Deseada Parejo, Universidad de Extremadura, Spain Recent research emphasizes that female song is evolutionarily important, yet there are Reviewed by: still few species for which we have quantified the similarities and differences between Ximena J. Nelson, male and female song. Comparing song rates and the structure of female and male University of Canterbury, New Zealand Christine Renee Dahlin, song is an important first step to forming hypotheses about functional and evolutionary University of Pittsburgh at Johnstown, differences that may exist between females and males, especially in year-round territorial USA species that may use their songs for breeding and non-breeding activities. We compared *Correspondence: Karan J. Odom female and male singing rates and song structure in a tropical New World oriole, the [email protected] Venezuelan troupial (Icterus icterus) during both the breeding and non-breeding season and between the dawn and day. Males sang solos at particularly high rates during the Specialty section: This article was submitted to breeding season before dawn. Females, however, sang at consistent rates year-round, Behavioral and Evolutionary Ecology, primarily during the day. Females answered 75% of male day songs, producing duets, a section of the journal whereas males answered only 42% of female songs. Duets were common year-round, Frontiers in Ecology and Evolution but occurred more often during the non-breeding season. Structurally, female songs Received: 16 November 2015 Accepted: 09 February 2016 were higher pitched and shorter than male songs. We detected no sex differences in Published: 25 February 2016 the number or order of syllables, however, interestingly, answers were shorter than duet Citation: initiations and solos, and, during the breeding season, songs that initiated duets were Odom KJ, Omland KE, McCaffrey DR, characterized by higher syllable diversity than were answers or solos. The fact that males Monroe MK, Christhilf JL, Roberts NS and Logue DM (2016) Typical Males sing more during the breeding season supports the classical hypothesis that male song and Unconventional Females: Songs is a sexually selected trait. However, our findings that females sing solos and answer the and Singing Behaviors of a Tropical, Duetting Oriole in the Breeding and majority of male songs to create duets year-round suggests that female song may have Non-Breeding Season. evolved to serve multiple functions not exclusively tied to breeding. Front. Ecol. Evol. 4:14. doi: 10.3389/fevo.2016.00014 Keywords: female song, duet, troupial, Icterus, behavior, selection Frontiers in Ecology and Evolution | www.frontiersin.org 1 February 2016 | Volume 4 | Article 14 Odom et al. Female and Male Troupial Songs INTRODUCTION et al., 2012). Two studies report female song rates that exceed male song rates (Price et al., 2008; Illes and Yunes-Jimenez, Most bird song research has been conducted on temperate 2009). However, no studies have examined how male and female male songbirds and supports the hypothesis that male song is song rates vary between the breeding and non-breeding season. shaped by sexual selection to attract mates and defend territories Here, we quantify the song rates, as well as song structure of (Beecher and Brenowitz, 2005; Catchpole and Slater, 2008). male and female Venezuelan troupials (Icterus icterus, hereafter Conversely, very little information exists on the function or even “troupials”) in both the breeding and non-breeding seasons to the structure of song in female songbirds (Langmore, 1998; Riebel provide insight about the function and selection pressures acting et al., 2005). However, recent research provides evidence that on male and female song. female song is common and was likely ancestral in songbirds Troupials, a New World oriole, are a sexually monochromatic, (Odom et al., 2014). Thus, to comprehensively investigate the socially monogamous, tropical-breeding songbird (Jaramillo and evolution of song in songbirds, we need to study both male and Burke, 1999). Mated pairs maintain territories together year- female song. Comparing the structure and timing of female and round and both males and females participate in territory defense male songs is a first step in understanding the selection pressures (Odom unpublished data). Territories are predominantly non- that may be acting on the songs of each sex (Hall, 2004; Dahlin overlapping and nest sites are solitary (Odom unpublished data; and Benedict, 2014). Jaramillo and Burke, 1999). No prior studies have been conducted Female song is particularly common in the tropics and unlike on banded populations of troupials and there is no formal in temperate regions tropical songbirds may sing throughout the description of their vocalizations. Previous accounts describe breeding and non-breeding seasons (Morton, 1996; Stutchbury troupial songs as comprising 2–3 repeated, alternating low and and Morton, 2001). While temperate male songbirds use song to high “whistles” (pure tones) with distinct space between notes gain access to mates, female animals may use elaborate traits to (Jaramillo and Burke, 1999). Both males and females sing solos compete for resources that increase their survival and fecundity and mated pairs often overlap each other’s songs to form duets (Emlen and Oring, 1977; West-Eberhard, 1983). Elaborate female (Odom, pers. obs.). Existing accounts also mention high rates traits, therefore, may evolve through broader selection pressures, of singing at dawn, but suggest that males are the primary such as social or natural selection to compete for and maintain singers (although others have not had banded populations; multi-purpose territories or non-breeding resources (Lebas, Jaramillo and Burke, 1999). Sex-specific seasonal variation in 2006; Tobias et al., 2012; Clutton-Brock and Huchard, 2013). We dawn singing has not been documented in tropical songbirds propose, one way to assess whether a trait has been shaped by where female song is common. Therefore, we were also interested sexual selection is to examine whether the trait is used exclusively to quantitatively compare troupial songs and song rates between during the breeding season. Traits that are used well outside of the dawn and day. Troupials are native to NE Colombia, the breeding season may have evolved in response to social or most of Venezuela, Aruba, Curacao, and Isla Margarita, but natural selection rather than solely sexual selection (Tobias et al., are considered introduced to islands throughout the Caribbean 2012). including Puerto Rico, St. Thomas, the U.S. Virgin Islands, and Most studies that have directly compared male and female Bonaire (Raffaele et al., 1989; Jaramillo and Burke, 1999). songs have compared structural variation (Logue and Gammon, We offer the first comprehensive description of variation 2004; Mennill et al., 2005; Rogers, 2005; Brunton and Li, 2006; in male and female troupial songs (Figure 1). Our paper has Hall, 2006; Molles et al., 2006; Dowling and Webster, 2013; Hahn two goals: (1) quantify the singing behavior and song structure et al., 2013; exceptions: Price et al., 2008; Topp and Mennill, 2008; variation in male and female troupials, and (2) determine Illes and Yunes-Jimenez, 2009). Comparing structure of male whether male and female singing behavior and song structure and female song has revealed interesting trends in the overall varies as a function of season (breeding vs. non-breeding) or complexity or repertoire sizes of male vs. female song. This has time of day (dawn vs. day). We discuss our results in the context been used to assess the relative role of sexual selection in shaping of existing information on the function and selection pressures elaboration of male vs. female traits (Mennill et al., 2005; Brunton acting on male and female songs and duets. and Li, 2006). Similarly, structural similarity and sex-specific song features have been useful for evaluating sex-role similarity Methods and individual motivation, especially in duetting species (Logue We collected data over three field seasons in the dry forests of and Gammon, 2004; Rogers, 2005; Hall, 2006). Cabo Rojo National Wildlife Refuge in southwestern Puerto Rico ◦ ◦ In contrast to song structure, singing behavior and song rates (17 590 N, 67 100 W), constituting two breeding seasons and may be especially informative when considering the breadth of one non-breeding season. We recorded Troupials from 10 May selection pressures that impact tropical species that live in the to 30 June 2013, 22 April to 01 July 2014, and 09 November to 18 same location year-round (Price et al., 2008; Illes and Yunes- December 2014. Late April to early May is the beginning of the Jimenez, 2009). Outside of the breeding season, singing is more wet season in Puerto Rico and the start of the breeding season likely to mediate competition for resources or social interactions for troupials in the dry forests of the island. October to April is not related to mating. Comparing male and female song rates the dry season, which is the non-breeding season for troupials in between the breeding and non-breeding seasons is valuable our study area. Our non-breeding season research was conducted for determining the relative importance of male and female in the first half of the non-breeding season, and thereby avoiding songs in regulating breeding vs. non-breeding activities (Tobias any confounding effects of pre-breeding activities. Frontiers in Ecology and Evolution | www.frontiersin.org 2 February 2016 | Volume 4 | Article 14 Odom et al.
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