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Ecology, Behavior and Social Organization of Saffron-Cowled Blackbirds (Xanthopsar Flavus)

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Ecology, Behavior and Social Organization of Saffron-Cowled Blackbirds (Xanthopsar Flavus) ORNITOLOGIA NEOTROPICAL 16: 15–29, 2005 © The Neotropical Ornithological Society ECOLOGY, BEHAVIOR AND SOCIAL ORGANIZATION OF SAFFRON-COWLED BLACKBIRDS (XANTHOPSAR FLAVUS) Rosendo M. Fraga Tucumán 335, Piso 11C, 1049 Buenos Aires, Argentina. E-mail: [email protected] Resumen. – Ecología, comportamiento y organización social del Tordo Amarillo (Xanthopsar fla- vus). – La ecología de la conducta y la organización social del Tordo amarillo (Xanthopsar flavus) se inves- tigó entre 1995 y 2004, principalmente en Argentina y Paraguay; también se obtuvo información de una pequeña población cautiva. Según una reciente filogenia molecular, este ictérido y ambas especies del genero Pseudoleistes forman un clado monofilético bien definido (clado Pseudoleistes). El Tordo Amarillo muestra un mínimo dimorfismo sexual en tamaño, pero un marcado dimorfismo sexual en coloración, concordando y discordando, respectivamente, con los Pseudoleistes. El Tordo Amarillo se alimenta principal- mente de artrópodos obtenidos por “gapeo” en el suelo o en vegetación baja. Nidifica como parejas monógamas en la vegetación acuática emergente de humedales, o en la vegetación herbácea en sitios secos. La nidificación puede ser solitaria o en pequeñas colonias de hasta 40 parejas. Los machos alimentan a los pichones tanto como las hembras, y se observaron machos ayudantes (helpers) en dos de seis nidos o gru- pos aislados con volantones. El repertorio vocal del Tordo Amarillo es relativamente exiguo (menos de 10 vocalizaciones) y mayormente compartido por ambos sexos. En todos estos aspectos, el Tordo Amarillo se asemeja a los Pseudoleistes, aunque su organización parece menos compleja. Una hipótesis basada en la dis- tribución filogenética del uso de humedales y conducta parental en ictéridos predijo que tanto la poliginia como un cuidado paterno mínimo se deberían dar en el clado Pseudoleistes, pero la predicción se basa en una categorización floja del hábitat y conducta territorial de estas especies. Abstract. – The behavioral ecology and social organization of Saffron-cowled Blackbirds (Xanthopsar fla- vus) was investigated between 1995 and 2004, mostly in Argentina and Paraguay. In addition, some data were obtained from a small captive population. According to a new molecular phylogeny, this blackbird, plus the two Pseudoleistes marshbirds, forms a well-defined monophyletic clade (marshbird clade). Sexes of Saffron-cowled Blackbirds show minimal size dimorphism and marked color dimorphism, agreeing and disagreeing with the marshbirds. The blackbird feeds mostly on arthropods by gaping in the ground or in low vegetation, and breed as monogamous pairs either in marsh emergent vegetation, or in herbaceous vegetation in dry soil. Nesting occurs solitarily, or in small colonies of up to 40 pairs. Males feed chicks at similar rates than females. Male helpers were observed in two of six isolated nests or postfledging groups. The vocal repertory of Saffron-cowled Blackbirds is comparatively small (less than 10 vocalizations) and mostly shared between the sexes. Saffron-cowled Blackbirds closely resemble the marshbirds in all those aspects, but their social organization seems less complex. An hypothesis based on the phylogenetic distri- bution of marsh nesting and parental behavior in icterids predicted that polygynous mating and minimal male care should occur in the marshbird clade, but the prediction was based on weak categorizations of habitats and territorial behavior. Accepted 22 November 2004. Key words: Saffron-cowled Blackbird, Xanthopsar flavus, displays, vocalizations, breeding organization, behavioral evolution, molecular phylogeny. 15 FRAGA INTRODUCTION icterids. In the new phylogenetic tree (Lanyon & Omland 1999, Johnson & Lanyon 1999), The Saffron-cowled Blackbird (Xanthopsar fla- Saffron-cowled Blackbirds are ancestral to an vus), a red data book species, survives in frag- exclusively South American clade containing ments of its former distribution in Argentina, also the marshbirds of the genus Pseudoleistes Brazil, Paraguay and Uruguay (BirdLife Inter- [Brown-and-yellow (P. virescens), Yellow- national 2002). Recent literature on the spe- rumped (P. guirahuro)]. The clade containing cies is mostly concerned with conservation Saffron-cowled Blackbirds and the marsh- issues (Fraga et al. 1998, Azpiroz 2000, Dias & birds (marshbird clade) was classified as Maurício 2002). Natural history and behav- marsh-nesting and, by being monogamous, it ioral data is included in some of those papers, was a clear exception to the trend. On the as well as in Belton (1985) and in the older lit- other hand, the nearest icterids outside this erature (e.g., Gibson 1885, Hudson 1920). clade, the Yellow-hooded (Chrysomus ictero- However, a detailed account of the behavioral cephalus )and Chestnut-capped Blackbirds (C. ecology (including vocalizations) of this ict- ruficapillus), agreed with the hypothesis, as erid has not been published. This paper pre- both species (formerly included in Agelaius) sents new behavioral data on Saffron-cowled are marsh-nesting, polygynous and have Blackbirds, obtained in Argentina, Uruguay unequal parental care (Lowther et al. 2004). and Paraguay, and summarizes our knowledge The poverty of information on South Ameri- on the social organization of the species. can icterids, even on basic subjects like habi- The behavioral ecology of some icterid tat, food and general ecology, made more species has been intensively studied, with difficult the analysis of Searcy et al. (1999). important reviews that summarize data and The subject is discussed here using more propose ideas on the evolution of social updated comparative data. behavior and organization within the family (e.g., Orians 1985a, 1985b; Robinson 1986). MATERIALS AND METHODS The information on behavior and ecology of Saffron-cowled Blackbirds is discussed in this Saffron-cowled Blackbirds were observed at context. Thanks to a new molecular phylog- 36 localities in Argentina (provinces of Corri- eny of the family (Lanyon & Omland 1999, entes and Entre Ríos), Uruguay (departa- Johnson & Lanyon 1999), more rigorous mento Rocha) and Paraguay (departamentos approaches to behavioral evolution in the Ict- Misiones, Caazapá and Itapúa). Coordinates eridae are now possible (Searcy et al 1999). of localities where the species was observed Scientific names in this paper follow the last were obtained with a GPS receiver, and dis- version of the South American Checklist tances between localities and occupied areas Committee of the American Ornithologists’ were estimated with Arc View GIS. The area Union (Remsen et al. 2004). Lowther et al. most frequently visited was located around (2004) cover the nomenclatural history of the Perdices (33º16´S, 58º34´W) in southeastern genus Agelaius (formerly including Xanthopsar) Entre Ríos. A total of 127 days and approxi- and state the reasons for the recent generic mately 500 h of field observations (from changes. October 1995 to March 2004) are included in The phylogenetic analysis of Searcy et al. this report. A total of 17 nestlings and fledg- (1999) suggested that a switch to marsh-nest- lings were color banded for individual identi- ing was crucial for the evolution of polygy- fication. nous mating and unequal parental care in the Data were also obtained from a group of 16 SAFFRON-COWLED BLACKBIRD TABLE 1. Body masses (g), sexual size dimorphism and color dimorphism in Saffron-cowled Blackbirds and their four closely related species, according to molecular phylogeny. Sample sizes are given for males and females. Size dimorphism is expressed as the ratio of mean male body mass/mean female body mass. Species Mass N Sexual dimorphism Male Female Male Female Size Color Chestnut-capped Blackbird (Chrysomus ruficapillus) 41.5 32.0 5 3 1.297 Yes Yellow-headed Blackbird (C. icterocephalus) 34.5 26.7 11 14 1.292 Yes Safron-cowled Blackbird (Xanthopsar flavus) 41.1 38.3 11 10 1.073 Yes Yellow-rumped Marshbird (Pseudoleites guirahuro) 91.2 81.9 6 3 1.113 No Brown-and-yellow Marshbird (P. virescens) 79.6 72.9 62 56 1.092 No Sources: Personal unpubl. data; publications: Belton (1985), Darrieu & Camperi (1998), Magalhaes (1999), Fraga & Di Giacomo (2004); label data on specimens at the American Museum of Natural History, New York, the Field Museum of Natural History, Chicago, and, for Yellow-headed Blackbirds, the Colección Ornitológica Phelps, Caracas. nine captive blackbirds (seven males and two from the literature (Belton 1985, Darrieu & females, all color banded for individual recog- Camperi 1998, Magalhaes 1999, Fraga & Di nition) donated by bird catchers in Buenos Giacomo 2004) and label data on specimens Aires, and captured at unprecise localities of at the American Museum of Natural History Corrientes and Entre Ríos provinces. This (New York), Field Museum of Natural His- captive population was kept in a 64 m2 out- tory (Chicago), and the Colección Orni- door aviary near Luján, Buenos Aires Prov- tológica Phelps ( Caracas). ince, Argentina, from 15 May to 23 October I recorded bird vocalizations with an 1998, with a special permission from the pro- AKG C568 EB shotgun microphone and a vincial government. The rehabilitated black- Sony Walkman Professional cassette recorder. birds were afterwards released in a private All vocalizations used in this paper were nature reserve in Buenos Aires Province spontaneous, i.e., no playback was used. One (BirdLife International 2000), and monitored vocalization of each recognizable type, and for some weeks. with a minimum
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