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Female Performance of Male Courtship Display in Northern Cardinals
M. Susan DeVries, Caitlin P. Winters, and Jodie M. Jawor

Southeastern Naturalist, Volume 13, Issue 2 (2014): N13–N17

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N13 2014 Southeastern Naturalist Notes Vol. 13, No. 2 M.S. DeVries, C.P. Winters, and J.M. Jawor Female Performance of Male Courtship Display in Northern Cardinals M. Susan DeVries1,2,*, Caitlin P. Winters1, and Jodie M. Jawor1 Abstract - Courtship behaviors in birds are often considered male-specific, as males compete for mates through displays that exhibit individual quality. Several courtship displays have been described for male Northern Cardinals including the complex song-dance display. We observed female cardinals performing the song-dance display on two separate occasions in south Mississippi within pre-breeding and breeding periods. Female performance of the display was very similar as reported for males. Given the behavioral attributes of cardinal mating pairs, it is plausible that bi-directional mate choice exists for this species and females are demonstrating aspects of individual quality to males through the song-dance display. Additional monitoring of courtship behavior is needed to determine the function of female performance display that was previously thought to be male specific. Introduction. Behavioral displays are essential for effective communication among organisms. Many taxa commonly use displays that convey information about territory ownership (i.e., territorial display) or coordinate reproductive events (i.e., courtship display; reviewed in Andersson 1994). Birds are well known for the performance of elaborate courtship. Such displays might accentuate a bird’s ornamentation (e.g., Amundsen et al. 1997), performance skill (e.g., Kelley and Endler 2012), and/or vocal repertoire (e.g., Dalziell et al. 2013). Cardinalis cardinalis (L.) (Northern Cardinal; hereafter Cardinal) are non-migratory, socially monogamous passerines that range from Central America to southern Canada (Halkin and Linville 1999). Adult plumage of this species is sexually dichromatic, with males having red body feathers as opposed to the tan body coloration of females. Both sexes are additionally ornamented, with a tall head crest, a robust orange bill, and a black/gray face mask of variable size and darkness. Unusual for most temperate songbirds, both male and female Cardinals sing (Halkin 1997, Yamaguchi 1998) and participate in prolonged defense of general-use territories (9+ months; DeVries 2013, Halkin and Linville 1999, Jawor 2007). This bi-parental species has a lengthy breeding season (6+ months) and behavioral displays are performed within reproductive periods by males (Lemon 1968) and females (Jawor and Breitwisch 2003, Lemon 1968, Shaver and Roberts 1933). However, reproductive behaviors of male Cardinals have been documented more extensively that those performed by females (e.g, Halkin and Linville 1999, Lemon 1968). Courtship displays have been described in detail for male Cardinals, most of which involve simple twisting and rotating of the body axis while in the presence of a female (e.g., “lopsided /asymmetrical display”; Lemon 1968). A more complex courtship male display of this species is the “song-dance display”, which has both visual and aural components (Lemon 1968). During this display, a male Cardinal faces a female and assumes an upright posture, with an erect head crest, a fanned tail, and smoothed body feathers (Lemon 1968). Maintaining this posture, the male shifts his weight from one foot to the other on a perch while singing, resulting in a swaying motion (Lemon 1968). Longitudinal twisting of the body axis can also occur (Lemon 1968). To our knowledge, performance of the song-dance display has only been described in detail 1University of Southern Mississippi, 118 College Drive #5018, Hattiesburg, MS 39406. 2University of Wisconsin-Madison, Department of Zoology, 250 N. Mills Street, Madison, WI 53706. *Corresponding author - msdevries@wisc.edu. Manuscript Editor: Frank R. Moore Notes of the Southeastern Naturalist, Issue 13/2, 2014 2014 Southeastern Naturalist Notes Vol. 13, No. 2 N14 M.S. DeVries, C.P. Winters, and J.M. Jawor for male Cardinals. Here, we report the first detailed observati ons of female Cardinals performing a very similar behavior to this presumably male-specific display. Observations. We observed female Cardinals performing the song-dance display within the pre-breeding and early breeding seasons at two different locations in Hattiesburg, MS. Both observations were seen following the broadcast of male Cardinal song (Observation 1: speaker broadcast of recorded song; Observation 2: live birdsong). Copulation attempts were not witnessed at the conclusion of the female display in which a male Cardinal was present (at least within the area of observation). The first observation was detected visually and aurally during the pre-breeding season for Cardinals at the Lake Thoreau Environmental Center in Lamar County, Hattiesburg, MS (31°21'1"N, 89°24'57"W). At approximately 0900 CST on 15 February 2010, a female Cardinal was observed performing the song-dance display in apparent response to hearing a pre-recorded male Cardinal song (Cornell Lab of Ornithology). No visual stimulus (e.g., male decoy, live bird, etc.) was present. Within 2 minutes of the onset of song broadcast (approximately 8–10 male songs), a female Cardinal flew onto the territory from an unknown location, perched within 3 m of the speaker, and began performing the song-dance display while oriented towards the speaker, which played continuously throughout her display. Consistent with Lemon’s (1968) description of the male song-dance display, the observed female’s crest was erect, wings were pressed against her body, feathers of her breast, head, and neck remained sleeked, and her tail was fully spread to expose most or all retrices. She maintained this posture while singing and shifting her body weight from foot to foot in an alternate fashion on a branch for approximately 10 seconds. It appeared as though the feet were being slightly raised from the branch as she rocked back and forth, which is a slight deviation of Lemon’s (1968) description of male song-dance display. Further, to the observers, the longitudinal axis of the female did not appear to be twisted to a great degree. The female’s song sounded fairly simple and consisted of repetitious syllables that overlapped with the male song broadcast. However, the song type of the female did not appear to be matched to the song type of the male song broadcast. Once she completed the display, the female left our observation area (presumably due to the lack of visual stimulus). It was unknown if the female was part of a mating pair located elsewhere on the study site or an unpaired individual attempting to acquire a mate (and territory). This female was not the owner of this territory from the previous breeding season and did not become the owner of this territory in the breeding season following our observation. Low and high temperatures on the date of this observation were 0.5 °C and 9.4 °C, respectively; temperature at time of observation was 2.2 °C (Weather Underground 2013), and skies were overcast. Apparent sunrise on 15 February 2010 was 0649 CST, and apparent sunset was 1733 CST (NOAA Solar Calculator 2013). The second observation of a female Cardinal performing the song-dance display was a visual and aural detection on the campus of the University of Southern Mississippi, Forrest County, Hattiesburg, MS (31°19'43"N, 89°20'2"W). This observation occurred at approximately 1130 CST on 22 March 2012. On this occasion, a female cardinal began performing the song-dance display in a tree (perched approximately 10 m high) while orienting towards a male Cardinal singing in a neighboring tree 4–5 m away (perched at approximately the same height as the female). The display lasted approximately 5–10 seconds before the female flew and perched in the tree in which the male was singing and began displaying within 1 m of him. While facing the male, she then resumed performance of the song-dance display for an additional 5–10 seconds before the male flew from the tree. The female followed the male, and both were quickly out of our range of N15 2014 Southeastern Naturalist Notes Vol. 13, No. 2 M.S. DeVries, C.P. Winters, and J.M. Jawor observation. Performance of the song-dance display by the female observed on this occasion was exactly as witnessed on 15 February 2010, only the display was now directed towards a male Cardinal as opposed to a sound speaker. Similar to the first observation, the female’s crest was erect, her tail was fully fanned, and her other feathers were flattened to her body as she swayed to and fro on a branch while singing in the direction of the male for approximately 5–10 seconds. Song of the female consisted of repeated syllables that overlapped with the song of the male, but the songs of the two individuals did not sound identical (to the observer). Reproductive status of the male and female was unknown, but both individuals were probably in breeding condition as this observation occurred within the early breeding season of Cardinals in this region (late March–early April; M.S. Devries, unpubl. data). It was also unknown if the male and female were a mated pair. Low and high temperatures on the date of this observation were 13.9 °C and 20 °C, respectively; temperature at time of observation was 15 °C (Weather Underground 2013), and skies were partly cloudy. Apparent sunrise on 22 March 2012 was 0558 CST, and apparent sunset was 1810 CST (NOAA 2013). Discussion. Documented observation of female Cardinals performing the song-dance display is of interest because, to our knowledge, it is the first detailed description of female Cardinals demonstrating a previously described male-specific behavioral display. Further, the song-dance display is an additional reproductive behavior performed by female Cardinals (for other examples, see Jawor and Breitwisch 2003, Shaver and Roberts 1933) that is clearly not the female pre-copulatory display reported by Lemon (1968). Courtship displays are often used to demonstrate aspects of individual quality to potential mates (reviewed in Andersson 1994). The display of male-typical courtship behavior by female Cardinals implies that females could be advertising information about their individual quality to males. Considering that the song-dance display has been observed in both male and female Cardinals, it is plausible that both sexes could use information from this display when forming breeding pairs. Bi-directional mate choice is potentially important in species in which pairs remain mated for multiple breeding seasons, jointly defend long-term territories, and contribute equally to offspring care, such as the Cardinal. Cardinal pairs also mate assortatively by ornaments (Jawor et al. 2003) indicative of body condition (potentially diet-dependent bill color: males [Jawor and Breitwisch 2004], females [Jawor et al. 2004]) and parental care (diet-dependent carotenoid plumage: males [Jawor and Breitwisch 2004], females [Linville et al. 1998, Jawor et al. 2004]), suggesting that potential mates assess multiple aspects of individual quality. Further, female Cardinals with larger face masks demonstrate greater aggression towards intrasexual intruders (Jawor et al. 2004) and higher testosterone throughout molt (J.M. Jawor and C.P. Winters, unpubl. data), which could be important qualities for a species participating in prolonged territorial defense. The upright posture and rocking motion of the song-dance display might allow a female Cardinal to simultaneously display many of these ornaments to a potential mate. Song produced during the song-dance display could also convey information about the individual quality of a female Cardinal to a male (Nowicki et al. 1998, 2002; Yamaguchi 1998). Collectively, the vocal and visual components of this display might provide a male Cardinal with more information about a female’s overall condition than mutually exclusive demonstrations of just vocal capabilities or ornamentation (e.g., Galeotti et al. 1997, Hill and Montgomerie 1994). Future investigation of similarities between male and female song elements performed during the song-dance display might provide useful information as to how this species establishes and maintains pair bonds. 2014 Southeastern Naturalist Notes Vol. 13, No. 2 N16 M.S. DeVries, C.P. Winters, and J.M. Jawor This is not the first report of a female Cardinal performing a reproductive display that differs from the described pre-copulatory display of this species (Lemon 1968). Shaver and Roberts (1933) reported a single observation of a breeding pair of Cardinals simultaneously swaying together while singing within the month of March. While some features of this apparent courtship behavior are similar to those described in detail for the song-dance display (Lemon 1968, this report), Shaver and Roberts’ (1933) observation did not mention the birds shifting their weight from one foot to another or the spreading of the tail as detailed by Lemon (1968) and this report. Considering the Cardinals simultaneously performing this behavior were presumed to be a breeding pair, it is possible that the observation witnessed by Shaver and Roberts (1933) is a behavior to strengthen a previously established pair bond that subtly differs from the song-dance display as described by Lemon (1968) and this report. Jawor and Breitwisch (2003) also detail an ornament display demonstrated by female Cardinals in the breeding season that did not result in immediate copulation, suggesting that a function of this ornament display could be to establish a pair bond. Even though we have limited observations of female Cardinals executing the song-dance display, it is plausible that this behavior has a similar function. One of our observations occurred during the prebreeding period (February–early March) when pair bonds were still being established in our Cardinal population, but breeding had not yet begun (breeding onset for this population is late March–early April; M.S. DeVries, unpubl. data). The second observation occurred at the beginning of the breeding season for Cardinals in this region, but no copulation attempt occurred before the birds flew out of sight. In addition, Lemon (1968) reported no copulation attempts after witnessing males perform the song-dance display for females, providing further support that the song-dance display in general could function more for pair-bond establishment in Cardinals than copulation. Additional observation of female courtship behavior is needed to determine why a female Cardinal would perform a display formerly thought to be male-specific. However, as we have observed, this could be challenging, as courtship behavior of Cardinals is elusive to investigator observation (Halkin and Linville 1998). Female Cardinals are of duller coloration and are more furtive than males, making their visual displays more difficult to see. However, the song-dance display has been recently witnessed in female Cardinals of another population (Austin, TX; C.P. Winters, pers. observ.), suggesting that female demonstration of this display is not limited to populations within south Mississippi. If this display is present in multiple populations throughout the Cardinal’s range, and investigators are aware that female performance of this male-specific behavior is a possibility, determining the function of the song-dance display in female Cardinals might be accomplished through additional monitoring and examination of courtship behavior. Acknowledgments. We thank Aaron Holbrook for field assistance and the Lake Thoreau Development Committee for accommodating our research. We also thank Will Selman and two anonymous reviewers for providing valuable feedback on an earlier version of this manuscript. Funding was provided by an Animal Behavior Society Student Research Award to M.S. DeVries and a National Science Foundation grant (award #0947944) “GK-12 Connections in the Classroom: Molecules to Muscles” awarded to the University of Southern Mississippi. Literature Cited Amundsen T., E. Forsgren, and L.T.T. Hansen. 1997. 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