Displays of Ringed Teal 97 Displays and breeding behaviour of captive Ringed Teal CaLlonetta Leucophrys G. L. Brewer Department of Biology, 101 Braddock Road, Frostburg State University, Frostburg, Maryland, 21532-1099, USA gbrewer0frostburg.edu Observations of captive Ringed Teal allowed several previously undescribed displays to be noted, including specialised displays for maintaining ^ contact, reinforcing the pairbond and biparental care. Males and females gave loud calls during female-only and pair flights that were associated with nest searching. Pair bonds were reinforced by bouts of contact calling when pairs became separated, and mates and family members greeted each other and came together in group displays after aggressive encounters. Males defended their mates throughout the breeding season, including dur­ ing forced extra-pair copulation attempts. Males did not consistently defend nest boxes. Females defended a nest box only when they physically occu­ pied it, laid eggs in more than one box, and were the only sex to incubate. One pair re-nested when their ducklings were about four weeks old. Both male and female parents performed Distraction Displays and called to ducklings when they became separated from the brood. A preliminary com­ parison suggests similarities between Ringed Teal and four closely-related species for displays associated with nest searching and pairbond mainte­ nance, including possible homologies for female vocalizations. Differences were noted in male vocalizations, female inciting behaviour, copulatory dis­ plays, and family displays, but further information on these species and wild Ringed Teal is needed to investigate possible convergence or phylogenetic influences. Key Words: Vocalisations, pairbond maintenance, biparental care, homology, comparative study © Wildfowl & Wetlands Trust W ild fow l 12001) 52: 97-125 98 Displays of Ringed Teal The biology and behaviour of the and the pochards (tribe Aythyini], Ringed Teal Callonetta leucophrys, Johnsgard (1960, 1965, 1978) noted which breeds in flooded forest areas in similarities with Aix. Both Delacour and parts of Argentina, Paraguay and Brazil Johnsgard concluded that Ringed Teal (Olrog 1968; Sick 1993), are poorly should be placed in the monotypie known, and the systematic position of genus Callonetta on the basis of its this species is still uncertain. Skeletal behaviour, but they did not agree on and plumage characteristics suggest which species were its closest rela­ that its nearest living relative is proba­ tives. bly the Brazilian Teal A m azonetta Preliminary observations of captive brasiliensis (Johnsgard 1960; individuals suggested that the pub­ Woolfenden 1961; W eller 1968). lished descriptions of behaviour and However, the feather proteins of Ringed vocalisations of Ringed Teal were Teal resemble those of the genus Anas incomplete. The main objective of this (Brush 1976] and, along with the study was to document displays and Brazilian Teal, it has been classified as breeding behaviour (especially vocali­ closer to the dabbling ducks by Bottjer sations) to supplement previous (1983) on the basis of immunoelec- accounts. Ringed Teal displays were trophoretic data. More recently, on the also compared, whenever possible, to basis of comparative morphology, those of potential close relatives to Livezey (1991) recognized Ringed Teal begin to examine the behavioural affini­ and Brazilian Teal as sister-genera and ties of this species. separated from both the other former Cairinini and the true dabbling ducks. Methods Data from DNA sequencing of m ito­ Captive Ringed Teal were observed chondrial genes suggest that Ringed at Cedar Creek Natural History Area, Teal are grouped with Maned Duck Bethel, Minnesota, from 19 July to 2 Chenonetta jubata and that Brazilian September 1983 and from 16 May to 14 Teal are most closely related to August 1984. In 1983, three full-winged Bronze-winged Duck Anas iSpeculanasI pairs, two full-winged unpaired males, specularis (Johnson 1997; Johnson & and one pinioned unpaired male Ringed Sorenson 1998). Teal were housed in a flight pen (5 x1 8 Behavioural data can be useful in χ 3.6m) with two nest boxes at a height solving taxonomic problems and com­ of 2m (described in Brewer 1988). parative studies of behaviour have been From 25 July to 8 August, 17 Speckled especially valuable in research on Teal Anas flavirostris flavirostris were waterfowl (Delacour & Mayr 1945; housed with the Ringed Teal. Very few Johnsgard 1965, 1978; Lorenz 1971). interspecific interactions were Although Delacour (1959) noted behav­ observed. In 1984, five pairs of fu ll­ ioural similarities between Ringed Teal winged Ringed Teal were released into Displays of Ringed Teal 99 a Large flight pen (55 χ 27.5 χ 3.6m) Maned Duck (pinioned) were observed described in McKinney (1967). This pen casually at the Wildfowl & Wetlands contained six nest boxes spaced evenly Trust, Slimbridge, England from 5 around the pen at a height of about 2m September to 12 September 1985. (described in Brewer 1988). Also Captive, pinioned North American housed in this pen were eight adult, Wood Duck and Mandarin Aix galericu- and later ten young, C h i loe Wigeon A. lata were observed occasionally at the sibitatrix and four adult, and later ten Como Park Zoo, St. Paul, Minnesota in young, White-cheeked Pintail A. May 1986. bahamensis bahamensis. Few interac­ Recordings of Ringed Teal, Brazilian tions were observed between these Teal, Maned Duck and North American species and the Ringed Teal. Wood Duck were made with a Sony Adult Ringed Teal were marked with TCM 5000-EV cassette tape recorder. coloured plastic nasal saddles (males) Additional recordings of Ringed Teal or discs (females) for individual identi­ were made with a Uher Report-L tape fication. A total of fifteen birds was recorder (tape speed of 9.5 cm/s) and a observed, with two pairs, one male and directional microphone. Recordings one female observed in both years. The were analysed with Kay sound spectro­ male was unpaired in 1983 and paired graphs 6061 B and 7800. in 1984, and the female was paired in An Elmo super 8 sound 10125-XL both years, but to different males. All macro camera was used to film Ringed birds were at least one year old. Teal behaviour at a speed of 18 All behavioural interactions were frames/s and films were analysed with noted on data sheets during 135.5 a Timelapse Data Analyser 3420 projec­ hours of scheduled observation, mostly tor. Film fram es were traced to during three-hour watches starting just illustrate major changes in movements before sunrise when the birds were during displays, including some transi­ especially active. Nests were checked tions between movements to indicate every few days during the laying period their fluidity. and only a few times during incubation Data were analysed using nonpara- when females were off their nests. metric statistical tests due to smalL Casual observations were made sever­ sample sizes. Differences in male and al times a week outside the breeding female behaviour were analysed using season (21 November 1984 to 30 April the Wilcoxon matched-pairs signed- 1985) on ten birds (two pairs, four ranks test, and associations between unpaired males, one unpaired female post-copulatory displays and copula­ and one juvenile female) in an animal tion success were analysed using the holding facility located at the University McNemartest [Conover 1980). of Minnesota, St. Paul, Minnesota. Captive Brazilian Teal (pinioned) and 100 Displays of Ringed Teal Results and Discussion a comparison of closely-related species can be useful to investigate In this paper, displays (stereotyped these factors. The identities of the behaviour patterns that are specialised Ringed Teal's closest relatives are con­ as signals; Moynihan 1955) are capi­ troversial, but four spcecies have been talised, and presented in boldface type proposed: North American Wood Duck, when they are described for the first Mandarin, Maned Duck and Brazilian time. Display terminology is that of Teal. Although information on display Johnsgard (1965) and McKinney (1965) repertoires is incomplete for several of unless otherwise noted. Most data on these species, identification similari­ breeding activities are from 1984. In ties and differences in comparison to both years, there was limited opportu­ the Ringed Teal has been attempted nity to observe unpaired birds. Further (Table 1). FI omologies are tentatively details, additional sonagrams and proposed only for female vocalisations, measured call characteristics, and although signalling needs to be associ­ tracings from film are presented in ated with breeding in cavities in wooded Brewer (1988). wetlands, maintenance of long-term Display repertoires are shaped by pairbonds, and biparental care may phylogenetic, ecological and social explain some display similarities. influences (McKinney 1975, 1992), and Displays of Ringed Teal 101 Table 1. Comparison of common Ringed Teal di splays to those of proposed close relatives.1 All known Ringed Teal displays are listed in the first colum. Y = display sim ilar to Ringed Teal, N = dis- play present but not sim ilar to Ringed Teal, - = d isplay not present, ? = unknown or uncertain (sonogram not available]. N orth Total Number Ringed B razilian A m erican Mandarin Maned of displays: Teal Teal Wood Duck Duck Y/Y? N/N? -/? Male displays: Flight call Y? N Y? Y? 3 1 0 Long whistle Y N N N 1 3 0 Short calls N N N N 0 4 0 Pre-copulatory: Head-bobbing Y Y 2 0 2 Head-jerking --- N? 0 1 3 Post-copulatory: Long whistle Y 1 03 Alert-beside - N - N 0 2 2 Female displays: Flight call Y? Y Y N 3 1 0 Houii Y? Y Y Y 4 0 0 Alarm call Y Y Y Y 4 0 0 "Inciting": Rotary-head- movements N N 0 2 2 Peep calls ---- 0 0 4 Soft calls Y Y Y Y 4 0 0 Displays of both sexes: Pre-flight: Neck-craning YYY 3 0 1 Greeting N N N N 0 4 0 Post-aggression N N N N 0 4 0 Distraction display Y ?? Y 2 0 2 Duckling displays: Contentment calls ? Y Y? ? 2 0 2 Distress calls ? Y Y? ? 2 0 2 'See text for sources of display information.