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Revista Brasileira de Ornitologia 27(3): 149–152. ARTICLE September 2019 Evidence of breeding activity of subadult Turdus thrushes in Argentina 1,2 2,3 2,4 5,6 Exequiel Barboza , Patricia Capllonch , Fernando Diego Ortiz & Alex E. Jahn Museo de Ciencias Naturales, Fundación Miguel Lillo, San Miguel de Tucumán, Tucumán, Argentina. Centro Nacional de Anillado de Aves (CENAA), Facultad de Ciencias Naturales e Instituto Miguel Lillo, Universidad Nacional de Tucumán, San Miguel de Tucumán, Tucumán, Argentina. Cátedra de Biornitología Argentina, Facultad de Ciencias Naturales e Instituto Miguel Lillo, Universidad Nacional de Tucumán, San Miguel de Tucumán, Tucumán, Argentina. Centro de Rehabilitación de Aves Rapaces (CeRAR), Reserva Experimental Horco Molle, Facultad de Ciencias Naturales e Instituto Miguel Lillo, Universidad Nacional de Tucumán, San Miguel de Tucumán, Tucumán, Argentina. Environmental Resilience Institute, Indiana University, Bloomington, Indiana, USA. Corresponding author: alexjahn@iu.edu Received on 04 February 2019. Accepted on 05 August 2019. ABSTRACT: Little is known about the age at which many Neotropical bird species first attempt to bree d. We evaluated the breeding condition of 515 adult and subadult males and females among four species of Turdus thrushes (T. nigriceps, T. chiguanco, T. amaurochalinus and T. rufiventris) in Tucumán, Argentina during three breeding seasons (2015 to 2018). We registered a total of 126 individuals with brood patches and cloacal protuberances, which accounted for 24.5% of thrushes in breeding condition that we sampled. Forty thrushes had a brood patch (31.7% of those in breeding condition), of which 11 were subadults (8.7%). Eighty-six thrushes (68.3%) had a swollen cloacal protuberance, 18 of which were subadults (14.3%). Only a few of the subadults in breeding condition built nests, incubated eggs or raised nestlings. Of 130 nests we found, only 3 belonged to a subadult thrush (in all cases, a subadult female with an adult social mate), one of which was successful. Further research on these patterns among various Neotropical bird species could lead to important insights into the life history strategies that characterize different populations, and how these ultimately affect their population dynamics. KEY-WORDS: brood patch, cloacal protuberance, nest, reproduction, Tucumán. INTRODUCTION strategies and the causes of population demographic patterns (Clutton-Brock 1988). According to life history A substantial body of research has shown that a bird's theory, birds should begin breeding when the net benefits reproductive performance can vary considerably over are greater than those that promote delaying breeding its lifetime (reviewed by Newton 1989). Reproductive (Promislow & Harvey 1990). Breeding at an early age can success in passerines generally improves with age (e.g., be beneficial if it reduces the chance of an individual dying Förschler & Kalko 2006), with older birds initiating before producing offspring or if it maximizes lifetime nesting activities earlier, having larger broods, and reproductive success. However, breeding at an early age lower rates of nest failure (Robertson & Rendell 2001). can be risky if survival of the offspring or of the adults is Females that breed in their first year tend to raise fewer reduced due to inexperience. Numerous previous studies offspring per season than older females (Nol & Smith have shown support for the idea that reproductive success 1987), although nest success eventually diminishes after increases with age, due to higher breeding experience of peaking in middle-aged females (Robertson & Rendell older birds (Nol & Smith 1987). 2001). Reproductive success is generally lower when one Nevertheless, our understanding of the causes member of a breeding pair is younger, whereas experienced of variation in reproductive success with age remains pairs, consisting of two adults, show higher reproductive incomplete. It some cases, it is apparent that a positive performance (Geslin et al. 2004). Thus, the first year in relationship between reproductive success and age is a bird's life appears to be generally characterized by low due to age-related experience in such tasks as foraging reproductive success (Förschler & Kalko 2006). (Förschler & Kalko 2006) and the care of eggs and young Evaluating how age influences reproductive success is (Robertson & Rendell 2001). Lower reproductive success important for understanding the evolution of life history may also change with age due to senility and lower fertility Subadult Turdus thrushes Barboza et al. laetevirens, Myrsine ferruginea, Eugenia mato, Myrcianthes as a bird ages (Partridge 1989). Additionally, as the value mato, Eugenia pungens, Eugenia seudo-mato, and Prunus of current reproduction increases relative to that of future tucumanensis; 4) Quebrada del Río Blanquito, town of reproduction (as should be the case with older vs. younger o o birds), the stress response may be attenuated, with older Tafí del Valle, 26 52'S; 65 41''W, 2400 m elevation; this birds exhibiting higher stress‐induced prolactin levels - site is in Temperate Cloud forest and primarily composed of Alnus acuminata with scattered Sambucus peruviana, ensuring that reproduction is not suspended compared to Rosa rubiginosa; 5) Rancho del Hombre Solo, El younger birds (Angelier et al. 2007). Infiernillo, km 78 on Route 307, 2800 m elevation; this The genus Tur dus is one of the largest avian genera on the planet, comprising 85 species (Gill & Donsker site is located 20 km north of the town of Tafí del Valle o o 2016). Substantial information on how to estimate the (26 49'S; 65 43'W) and is primarily composed of grasses (Festuca hieronymeus) and patches of bushes (Iochroma age of Turdus thrushes is already available, facilitating australe, Baccharis sp. and Satureja parvifolia). research on their age-specific behaviors. Turdus nigriceps and T. rufiventris a cquire definitive adult plumage after 14 months and T. amaurochalinus do so in at least 16 Data collection months, although they do not necessarily begin to breed We classified a thrus h as a juvenile (i.e., hatched during at that age (Howell et al. 2003). Molt cycle information the current season), as a subadult (i.e., hatched during is lacking for T. chiguanco; nevertheless, subadults of all the previous season) or as an adult (i.e., hatched at least the species we studied are in their formative plumage in spring, which is similar to adult plumage except that it two seasons prior to capture) using the amount of skull is more opaque and heavily worn. They may also exhibit ossification, the color and texture of the gape (i.e., soft, swollen and yellow in juveniles) and plumage/molt some retained juvenile body feathers and wing coverts characteristics (e.g., shine, wear, retained juvenile feathers (e.g., Ortiz et al. 2017). or recently molted; Wolfe et al. 2009, Ortiz et al. 2017). Previous research has shown that subadult Turdus thrushes that are one year old may have a well-developed Turdus can be reliably classified as subadults because cloacal protuberance, which raises the possibility that these they retain the juvenile wing coverts, which exhibit buffy tips, and exhibit swollen gapes for several months during thrushes reproduce before acquiring their adult plumage the first year of life (e.g., Ortiz et al. 2017). After breeding (Capllonch 2015). Given the gap in information on and before migrating in fall, they molt body feathers, age-related breeding effort in Neotropical passerines, we evaluated the incidence of breeding activity of subadult remiges and rectrices, molting into their basic definitive Turdus thrushes in northwestern Argentina. plumage, which is much brighter and without buffy tips (Capllonch 2015). For example, T. amaurochalinus begins molting immediately after breeding in late summer or in early autumn and continues molt during fall migration METHODS (Soria et al. 2008). We captured thrushes using mist nets during their breeding We classified a thrus h as being in breeding condition by the presence of a swollen cloacal protuberance and season, from September–February 2015, 2016, 2017, a developed brood patch (Ralph et al. 1996) and also and January–February 2018 at sites located within four determined sex by the presence of a swollen cloacal major ecosystem types of Tucumán province, Argentina: Montane Chaco Woodland, Temperate Cloud Forest, protuberance (males) or developed brood patch (females; Subtropical Montane Forest and Lowland Foothill Forest Ralph et al. 1996). For T. nigriceps, we also used sexual dimorphism in plumage as an indicator of sex (Narosky (Cabrera 1976). The sites we worked at are: 1) Ti cucho, o o & Yzurieta 2010). 26 31'S; 64 53'W, 600 m a.s.l.; this site is found in Montane Chaco Woodland, and common trees here are Schinopsis marginata, Acacia caven, Prosopis nigra, Geoffroea Nest monitoring decorticans, Ceiba insignis as well as several types of cactus We conducted nest searches by walking along paths, (e.g., Thrichocereus terscheckii, Opuntia sp.); 2) Botanical o o paying particular attention to forks in branches of trees Garden, Reserva de Horco Molle, 26 55'S; 65 05'W, 550 and shrubs, which are typical locations where thrushes m a.s.l.; this site is located in Lowland Foothill Forest, with common plants being Heliocarpus popayanensis, place their cup nests. We also observed the behavior of Tecoma stans, Juglans australis, Anadenanthera colubrine, adult thrushes to find nests, such as escape movements from the nest when we approached. Females will also Cedrela lilloi, Parapiptadenia excelsa and Allophyllus edulis; make a distinct “cluck” call when an observer is near 3) El Rulo, Yerba Buena, 1000 m a.s.l.; this site is in the nest, which helped detect the presence of an active Subtropical Montane Forest and is primarily composed of Cinnamomum porphyrium, Cupania vernalis, Myrsine nest. When nests have nestlings, both parents are often Revista Brasileira de Ornitologia 27(3): 2019 Subadult Turdus thrushes Barboza et al. near the nest and make distinct vocalizations, which aids 2017. The eggs were predated and t he female did not re- in finding the nest. We checked nest contents using a use the nest. The thir d was located in the basin of the ladder or with a video camera attached to a long pole. Blanquito River near Tafí del Valle and which belonged We classified a nest as successful if it fledged at least one to a T. nigriceps subadult female that was captured on 03 nestling. We classified a nest as failed if there were missing Jan 2018 in a mist net placed near the nest. The three eggs (i.e., which were present in a previous visit to the nestlings of that nest were killed in an intense storm, nest) or when nestlings that were too young to fledge likely due to excessive water in the nest, which was placed were missing. If a nest failed, it was checked at least once in a cavity. The nest was not re-used. more to determine if it was re-used. If a nestling fledged from a nest, the nest was checked again ten days later to determine if the nest was re-used. DISCUSSION We found that subadult Turdus thrushes attempt to breed, RESULTS although our results suggest that only a small proportion of subadult Turdus thrushes do so. We only found three We caught a total of 515 Turdus thrushes from four nests belonging to subadults, such that we cannot yet species (T. nigriceps, T. chiguanco, T. amaurochalinus and make definitive conclusions regarding the probability T. rufiventris), 126 of which were in bree ding condition, of success between adults and subadults; nevertheless, accounting for 24.5% of all thrushes captured. Forty subadults of both sexes were found in breeding condition. thrushes had a brood patch (31.7% of those in breeding Although we did not find any nests belonging to subadult condition), of which 11 were subadults (8.7%). Eighty- males, the males we sampled that had a swollen cloacal six thrushes (68.3%) had a swollen cloacal protuberance, protuberance may have pursued extra-pair copulations. 18 of which were subadults (14.3%). Previous research has shown that adults may benefit We monitored 130 nests, 38 of which belonged to from holding territories in higher quality habitat with T. amaurochalinus, 12 to T. nigriceps, 38 of T. chiguanco, better nest sites and by having more experience with and 42 to T. rufiventris ( Table 1). None of these nests breeding and foraging, allowing them to maintain a belonged to a subadult male, three belonged to a subadult proper energy balance, which is key to physiologically female, and the rest belonged to adults. Success rates of preparing for reproduction (Robertson & Rendell 2001). nests belonging to adults (i.e., excluding the three nests In contrast, subadult thrushes may use their first year belonging to a subadult female) ranged between 28.6 and to obtain the necessary skills necessary to successfully 50% (Table 1). In all three cases in which the nest belonged breed, such as nest building and foraging. Notably, we to a subadult female, their social mate was an adult male. found several adult nests abandoned before laying eggs, The first belonged to a T. nigriceps subadult female, which whereas all nests of subadults had eggs. Further research was captured next to the nest on 08 November 2016 in comparing nest outcomes between adults and subadults El Rulo, San Javier. This nest was successful, fledging at different stages of the nesting cy cle may shed light three nestlings; the nest was not re-used. The second was on how these and other bird species develop the skills a nest of a T. amaurochalinus subadult female, which was necessary to nest successfully. captured leaving the nest in Ticucho on 18 November Further studies on age-related breeding effort Table 1. Nest outcomes for four species of Turdus thrushes monitored in Tucumán province, Argentina. Abandoned Failed at Failed at Total Adult nests Successful % Successful before laying egg stage nestling stage nests T. amaurochalinus 1 14 6 16 37 43.2 T. nigriceps 05 0 5 10 50.0 T. chiguanco 4 13 9 12 38 31.6 T. rufiventris 1 24 5 12 42 28.6 Total 6 56 20 45 127 Subadult nests T. amaurochalinus 0 1 0 0 1 0.0 T. nigriceps 0 0 1 1 2 50.0 Total 01 1 1 3 Revista Brasileira de Ornitologia 27(3): 2019 Subadult Turdus thrushes Barboza et al. variation in contrasting breeding systems. Chicago: University of in Neotropical passerines not only offers a better Chicago Press understanding of the poorly-understood first year of life Fay R., Barbraud C., Delord K. & Weimerskirch H. 2016. Variation of Neotropical birds, it can also provide novel insights in the age of first reproduction: different strategies or individual into the drivers of population dynamics within and across quality? 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Journal of Animal Ecology 71: 55–64. ACKNOWLEDGEMENTS Ortiz D., Jahn A.E., Capllonch P., Cereghetti J. & Barboza E. 2017. Determinación de edad y muda del Zorzal Plomizo (Turdus We are grateful to three anonymous reviewers, whose nigriceps) en Tucumán, Argentina. Cochabamba: Abstracts of the IX Congreso de Ornitología. comments greatly improved the manuscript. We thank Partridge L. 1989. Lifetime reproductive success and life-history the Universidad Nacional de Tucumán for allowing us evolution, p. 349–362. In: Newton I. (ed.). Lifetime reproduction access to the Horco Molle and Ticucho properties. We of birds. London: Academic Press. thank the Municipality of Tafí del Valle for allowing Promislow D.E.L. & Harvey P.H. 1990. Living fast and dying young: access to Quebrada del Blanquito, and we thank Ing. Juan a comparative analysis of life-history variation among mammals. Journal of Zoology, London 220: 417–437. Manuel Paz for letting us access Rancho del Hombre Solo. 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Ornithology Research – Springer Journals
Published: Sep 1, 2019
Keywords: brood patch; cloacal protuberance; nest; reproduction; Tucumán
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