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Nesting Success and Productivity of Tucuman Parrots (Amazona Tucumana) in High-Altitude Forests of Argentina: Do They Differ from Lowland Amazona Parrots?

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Nesting Success and Productivity of Tucuman Parrots (Amazona Tucumana) in High-Altitude Forests of Argentina: Do They Differ from Lowland Amazona Parrots? CSIRO PUBLISHING Emu http://dx.doi.org/10.1071/MU12062 Nesting success and productivity of Tucuman Parrots (Amazona tucumana) in high-altitude forests of Argentina: do they differ from lowland Amazona parrots? Luis RiveraA,D, Natalia Politi A, Enrique H. Bucher B and Anna Pidgeon C AFacultad de Ciencias Agrarias, Universidad Nacional de Jujuy, CONICET, Alberdi 47, S. S. de Jujuy (4600), Jujuy, Argentina. BCentro de Zoología Aplicada, Universidad Nacional de Córdoba, CONICET, Casilla de Correo 122, Córdoba (5000), Argentina. CDepartment of Forest and Wildlife Ecology, University of Wisconsin – Madison, 120 Russell Laboratories, 1630 Linden Drive, Madison, Wisconsin, WI 53706-1598, USA. DCorresponding author. Email: [email protected] Abstract. Most of our knowledge of reproduction of wild parrots in the Neotropics comes from studies of tropical lowland species, with few studies addressing species of high-altitude forests. We studied the reproductive biology of Tucuman Parrots (Amazona tucumana) in north-western Argentina between 2004 and 2009. We obtained data on reproductive output for 86 nests and on causes of mortality for 94 nests. Mean clutch-size per nesting attempt was 3.6 eggs Æ 1.0 (s.d.). Hatching success (proportion of eggs laid that hatch) was 0.77 Æ 0.17. Fledging success (proportion of nestlings that fledge) was 0.83 Æ 0.13. The overall breeding success (mean number of fledglings per laying female per year) was 2.3 Æ 0.8. Overall finite nesting success (daily survival rate to the power of the nesting length) was 0.53 Æ 0.27, and chick finite nesting success rate was 0.74 Æ 0.22. We did not find differences in reproductive rate between Tucuman Parrots and other species of Amazona parrot from lowland habitats. Productivity and nesting success of Tucuman Parrots had high values in some years and low values in others. This was probably related to fruiting events of Podocarpus parlatorei – a critical food item. The main causes of nesting failure were predation (16%) and abandonment (12%). Our results suggest that for several species of Amazona in lowland habitats, predation and poaching may be the main limiting factors whereas climatic factors and food availability may contribute most to nesting failure at higher altitudes. Additional keywords: breeding biology, mortality factors, reproductive output, subtropical cloud forests. Received 1 October 2012, accepted 10 May 2103, published online 20 September 2013 Introduction parrots that occur in high-altitude forests, i.e. the Thick-billed Detailed knowledge of breeding biology is necessary for under- Parrot (Rhynchopsitta pachyrhyncha) and the Indigo-winged standing variation in avian reproductive strategies, because it Parrot (Hapalopsittaca fuertesi), there has been only one study provides critical natural history data that are useful for generating that has described reproductive rates (Thick-billed: Monterrubio new hypotheses and testing old ones (Auer et al. 2007). The lack et al. 2002) and another on nestling growth and plumage devel- of information on the natural history, ecology and demography opment (Indigo-winged: Tovar-Martinez 2009). of many parrot species precludes an assessment of the mechan- There is a well-established pattern of variation in bird life- isms that regulate population dynamics (Koenig 2001). There history traits along latitudinal gradients in which life-history have been a number of studies of the nesting and reproductive strategies of tropical and southern hemisphere birds differ behaviour of several lowland tropical forest species of Amazona markedly from those of northern temperate birds (Martin parrot (Gnam 1991; Rojas Suárez 1991; Enkerlin-Hoeflich 1995; 2004). Elevation has long been considered an important factor Koenig 2001; Fernandes Seixas and Mourao 2002; Renton and in shaping the evolution of life-history traits (Cody 1966; Salinas Melgoza 2004; Sanz and Rodríguez Ferraro 2006; Rodrí- Badyaev 1997). High-elevation species have been thought to guez Castillo and Eberhard 2006; Berkunsky and Reboreda have higher reproductive rates, potentially compensating for 2009). However, the behaviour and reproductive rates of Ama- the less predictable climate, and lower population density and zona parrots that nest in temperate and high-altitude forests to experience generally more limited availability of food (Cody (above 1500 m above sea level), specifically the Scaly-naped 1966; Badyaev 1997). Congeners whose young are altricial Parrot (A. mercenaria) and the Tucuman Parrot (A. tucumana), have larger clutches at high elevation than at lower elevations are not known (Juniper and Parr 1998). Of the other two (Stewart et al. 1977; Weathers et al. 2002; Johnson et al. 2006). Journal compilation Ó BirdLife Australia 2013 www.publish.csiro.au/journals/emu B Emu L. Rivera et al. However, there is also evidence suggesting that birds shift to a mountain range ~100 km long, between the Cordillera Oriental to slower life-history strategy with increasing elevation, investing the west and the Chaco plain to the east. The Southern Yungas less in reproduction and producing smaller clutches than birds at supports a semi-evergreen subtropical montane cloud forest that lower altitudes (Krementz and Handford 1984; Badyaev 1997; forms the southernmost part of the Andean tropical forest Badyaev and Ghalambor 2001). (Cabrera 1976). The cloud forest is dominated by Podocarpus Some reproductive parameters of Amazona parrots from parlatorei, Alnus acuminata and trees of the family Myrtaceae. lowland tropical forest are well known. Most Amazona parrots The local climate has a marked dry season from April to October have rates of nesting success of 35–57% (Gnam and Rockwell and a rainy season from November to March. Annual rainfall is 1991; Enkerlin-Hoeflich 1995; Koenig 2001; Fernandes Seixas 800–1500 mm and mean annual temperature is 11.7C (Mendoza and Mourao 2002; Renton and Salinas Melgoza 2004; Sanz and 2005). Rodríguez Ferraro 2006). However, island species tend to have Within the central sector of the Southern Yungas, we focussed higher rates of success, of 46–82% (Snyder et al. 1987; Gnam and on two areas where we knew of active nests: (1) El Rey National Rockwell 1991) and in studies of two Amazona species with the Park (24430S, 64380W, 44 000 ha) and (2) Portal de Piedra lowest documented rates of nesting success (i.e. 0–13%), rates of Private Reserve (24050S, 64260W, 400 ha). Both areas are nest poaching were high (Martuscelli 1995; Rodríguez Castillo strictly protected and controlled, no people live within the and Eberhard 2006). Most Amazona species have a mean clutch- reserves and, to the best of our knowledge, there is no poaching size of 2.6–3.6 eggs and produce 0.3–2.5 fledglings per female (in 5 years of field-work we did not detect any signs or tracks of (Snyder et al.1987; Gnam and Rockwell 1991; Enkerlin-Hoeflich people in the nesting areas). The study was conducted within 1995; Martuscelli 1995; Koenig 2001; Renton and Salinas accessible areas only. These comprised ~170 ha in Portal de Melgoza 2004; Rodríguez Castillo and Eberhard 2006; Sanz and Piedra Private Reserve and 50 ha in El Rey National Park. Rodríguez Ferraro 2006). Elevation of these areas is between 1450 and 2100 m above sea In contrast to many lowland forest species of Amazona,the level. Tucuman Parrot has a small geographical range in the narrow strip of cloud forest (1400–2200 m above sea level) of the Southern Yungas on the eastern slopes of the Andes from south-eastern Data collection Boliviatonorth-westernArgentina(FjeldsåandKrabbe1990).The Field-work was conducted from November to March over five species is categorised as Vulnerable on the IUCN Red List, and breeding seasons between 2004 and 2009. We recorded pairs of is included in CITES Appendix I prohibiting international trade Tucuman Parrots prospecting for nesting cavities in November, (BirdLife International 2011). A recent assessment of the status of laying eggs from early to mid-December, and young fledging the species estimated that current population is probably ~33% from mid- to late February. Daily searches for active nests were smaller than in the 1980s (Rivera et al. 2007, 2010). conducted for 35–40 days each breeding season during the Food availability is a major factor affecting annual produc- laying and incubation periods (December–mid-January). We tivity of parrots (Renton and Salinas Melgoza 2004; Díaz et al. used a hand-held GPS unit (Garmin Etrex HCx, Garmin Inter- 2012). The Tucuman Parrot relies heavily on a single food item, national Inc., Olathe, KS, USA) to record the boundaries of the the seeds of Podocarpus parlatorei (Pinales : Podocarpaceae), for area surveyed, which were later imported into a geographical feeding nestlings (Rivera 2011). The reproductive biology of information system (ArcGIS 9, ESRI, Redlands, CA, USA). the Tucuman Parrot is virtually unknown, with only one record Active nests were found by following males to the nesting area of a nest, from January in the 1940s in Chuquisaca Department, and locating the cavity when the female left to be fed by the male Bolivia (Bond and Meyer de Schauensee 1943). (González Elizondo 1998). Nests were found throughout the An understanding of the reproductive biology of the Tucuman incubation period and the early nestling period, when females Parrot may provide insights into the mechanisms that regulate were incubating or brooding. We did not find nests later in the population dynamics of this species and the associated limiting nesting cycle because
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