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The Relationships Between Prey Size, Nestling Age, Provisioning Rate, and Elevation in the Varied Tit Parus Varius

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The Relationships Between Prey Size, Nestling Age, Provisioning Rate, and Elevation in the Varied Tit Parus Varius Ornithol Sci 15: 29 – 36 (2016) ORIGINAL ARTICLE The relationships between prey size, nestling age, provisioning rate, and elevation in the Varied Tit Parus varius Jong Koo LEE, Woongsoon JANG, Ok-Sik CHUNG#,* and Woo-Shin LEE Department of Forest Sciences, Seoul National University, Seoul 151–921, Republic of Korea ORNITHOLOGICAL Abstract To determine the major factor of change in provisioning rates to Varied Tit Parus varius nestlings in relation to increasing elevation, we used video record- SCIENCE ings to investigate the number of parent visits and the size of prey carried by parents © The Ornithological Society to their nestlings during the breeding process. Video cameras were installed at the of Japan 2016 entrances of 35 nests (17 at 300 m a.s.l., 7 at 900 m a.s.l., and 11 at 1,400 m a.s.l.). We also measured air temperature during the breeding season at three study sites. The provisioning rates per nestling showed a significantly negative relationship with prey size provisioned, except during the early nestling stage (days 1 to 4), when small prey were selectively provisioned by the parents. Prey at the higher elevation during days 5 to 15, also, were significantly smaller than at the lower elevation. The mean tempera- ture at each breeding site during the breeding period did not differ significantly due to the delayed onset of breeding at higher elevation. Therefore, we conclude that the increase in the provisioning rate per Varied Tit nestling at higher elevation is associ- ated with a decrease in prey sizes at that elevation, rather than with a difference in the temperature. Key words Elevation, Nestling age, Prey size, Provisioning rate per nestling The provisioning rate of adult birds to their off- during the early nestling stage, in response to their spring differs even within the same species resulting physiological and nutritional demand (Moreno 1987; from various factors such as: adult body size; habitat Naef-Daenzer et al. 2000; Slagsvold & Wiebe 2007). quality; distance between the nest and prey resources; However, provisioning by parents may represent a the size of available prey; the capability or body size trade-off between parents and offspring, in effect of the parent birds; and, the number of nestlings parent-offspring conflict. More frequent provision- (Nur 1984b; Nour et al. 1998; Grieco 2001; Grieco ing of chicks, while aiding nestling survival, repre- 2002; Kilner 2002; Tremblay et al. 2005; Strauss et sents an extra cost for parents due to their increased al. 2005; Nakagawa et al. 2007; Mägi et al. 2009; flight distance and their missed opportunity costs Wilkin et al. 2009). In particular, the quantity and (Strauss et al. 2005). Parent birds should determine quality of prey with which the parents supply their the frequency of provisioning taking into account chicks play a critical role in determining nestling sur- the needs of their offspring. Nestlings express their vival. Nestlings are completely dependent on the food nutritional needs to their parent birds by means of supplied by their parents (Bengtsson & Rydén 1983; their begging calls; these represent an honesty signal Nur 1984a), thus parents rapidly respond to envi- of offspring energy state, and parents respond imme- ronmental changes by collaborating to enhance the diately to the begging call (Bengtsson & Rydén 1983; survival of their brood (Gibb 1950; Wright & Cuthill Grieco 2001; Sacchi et al. 2001). Therefore, prey 1989). Parents, also, tend to selectively feed specific quality and quantity fed to nestlings may be one of types of prey, such as small, soft insects to nestlings the most important factors affecting the provisioning rate, because it directly affects the energy state of the offspring (Martin 1987; Boutin 1990; Grieco 2001; (Received 1 March 2015; Accepted 15 August 2015) Grieco 2002). Therefore, to determine the main cause # Corresponding author, E-mail: [email protected] * Present address: Chungnam Institute, 73-26 Yeonsuwon-gil, of change in provisioning rate, it might be necessary Gongju-si, Chungcheongnam-do, Korea to look into the prey size provisioned to nestlings 29 JK. LEE et al. during the nestling period. ambient temperature could be a major factor contrib- Elevation is considered to be a factor that can uting to differences in provisioning rates to nestlings impact provisioning rates because it is an environ- by Varied Tit Parus varius parents. Our hypothesis mental gradient that affects a range of character- is that differences in provisioning rates result from istics including: behavior, morphology, and breed- differences in prey size, rather than from cold stress ing habits. In practice, birds that breed at higher on nestlings at higher elevation. elevations have different breeding characteristics from those that breed at lower elevations due to the MATERIALS AND METHODS colder weather, stronger seasonality, and wider daily temperature ranges associated with high elevation 1) Study sites (Badyaev 1997). In a previous study on the Varied We selected three study sites facing in similar direc- Tit (Lee et al. 2011), we showed that the provision- tions and with similar angles of inclination: Piagol ing rates per nestling increased at higher elevation. (approximately 300 m above sea level), Siamjae However, the factors driving parents to feed their (900 m a.s.l.), and Nogodan (about 1,400 m a.s.l.) nestlings more often at higher elevations remain (Table 1). All three sites have similar vegetation unclear. Badyaev and Ghalambor (2001) suggested (broad-leaved forest), but are at different elevations that increased parental visits at high elevations reflect on Mt. Jiri, which is located in the southern part of a strategy to increase offspring survival in response South Korea (Table 1). Forty-eight nest boxes were to the colder environment at high elevation. How- installed at each of the three sites during November ever, this may only be true under the assumption 2006 and research was conducted from April to July that prey quality was identical at different elevations. 2007 during the breeding season. The Varied Tit is a Boyce (1979), Janes (1994), and Chown and Klok monogamous species that inhabits a wide range of (2003) have shown that in fact prey sizes, at loca- elevations from sea level to beyond 1,500 m a.s.l., tions where there is high seasonality, were inclined making it an appropriate species to study provision- to be smaller at higher elevations due to the short ing rate in relation to elevation. In order to compare spring season. Thus, it is possible that the difference the daily provisioning rates by parents, we used the in provisioning rates at higher elevations results from provisioning rate per nestling which was calculated smaller prey size, not lower temperature. by dividing the number of parent visits per hour by We conducted this study to investigate the key fac- the number of nestlings in the nest. We set up two tors that might explain the differences in provisioning HOBO Pro Series data loggers (On-set computer rates to nestlings in relation to elevation, focusing on Corporation, Porasset, MA, USA) at each site, which the size of prey and nestling age. Also, by comparing measured and stored ambient temperature data every the temperature during the breeding season at study 30 minutes. sites with different elevations, we tested whether Table 1. Description of study sites Study site Piagol (n=17) Siamjae (n=7) Nogodan (n=11) 35°15′37.1″N, 35°18′14.6″N, 35°17′36.9″N, Longitude/latitude 127°35′00.1″E 127°20′46.6″E 127°31′42.4″E Elevation (m) 310–380 904–977 1,346–1,443 Aspect South South Southeast Styrax japonica Quercus mongolica Quercus mongolica Dominant tree Carpinus laxiflora Acer pseudosieboldianum Acer pseudosieboldianum Quercus serrata Rhododendron mucronulatum Rhododendron schlippenbachii Quercus acutissima Quercus mongolica Dominant understory Rhododendron mucronulatum Carpinus cordata Rhododendron schlippenbachii 30 Effect of prey size on provisioning rate 2) Observations of provisioning rates and prey size study areas. The mean body lengths of male parents Video cameras (Sony Handycam HDR-SR1) were: 13.78 cm (SD=0.13 cm) in Piagol (n=17), were installed in the entrances of each nest box in 13.77 (SD=0.16 cm) in Siamjae (n=7), and 13.71 order to record parental visits. Seventeen cameras cm (SD=0.10 cm) in Nogodan (n=11) (ANOVA, were installed in Piagol, seven in Siamjae, and 11 P=0.383, F=0.781). The mean body lengths of in Nogodan. Video recordings were made for 4–8 female parents were: were 13.49 cm (SD=0.11 cm) hours each day from hatching until fledging of the in Piagol, 13.54 cm (SD=0.10 cm) in Siamjae, and nestlings. A total of 1,204 hours of recordings were 13.55 cm (SD=0.07 cm) in Nogodan (ANOVA, made at nests at Piagol, 571 hours at Siamjae, and P=0.283, F=1.119). On average, male tarsus length 976 hours at Nogodan. Video recordings allowed was 1.917 cm (SD=0.031 cm) in Piagol, 1.912 cm prey size to be estimated in comparison with the beak (SD=0.046 cm) in Siamjae, and 1.908 cm (SD=0.055 width and beak depth of parents (measured manually) cm) in Nogodan (ANOVA, P=0.868, F=0.142), (Grieco 2001). whereas mean female tarsus length was 1.845 cm (SD=0.052 cm) in Piagol, 1.834 cm (SD=0.065 cm) 3) Measurements of parents and their nestlings in Siamjae, and 1.852 cm (SD=0.037 cm) in Nogodan Parents were captured during incubation, measured (ANOVA, P=0.691, F=0.374). The average number (total body length, tarsus length, beak depth and beak of eggs laid per clutch was 6.69 (SD=0.75) at Piagol, width) and individually marked with colored leg 6.26 (SD=1.15) at Siamjae, and 5.90 (SD=0.70) at bands.
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