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Foraging Behavior and Diet of Streaked Shearwaters Calonectris Leucomelas Rearing Chicks on Mikura Island

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Foraging Behavior and Diet of Streaked Shearwaters Calonectris Leucomelas Rearing Chicks on Mikura Island Ornithol Sci 11: 9 – 19 (2012) ORIGINAL ARTICLE Foraging behavior and diet of Streaked Shearwaters Calonectris leucomelas rearing chicks on Mikura Island Kei MATSUMOTO1,*, Nariko OKA2, Daisuke OCHI1,**, Fumihito MUTO3,***, Takashi P. SATOH3,**** and Yutaka WATANUKI1,# 1 Graduate School of Fisheries Sciences, Hokkaido University, 3–1–1 Minato-cho, Hakodate, Hokkaido 041–8611, Japan 2 Research Division, Yamashina Institute for Ornithology, 115 Konoyama, Abiko, Chiba 270–1145, Japan 3 Atmosphere and Ocean Research Institute, the University of Tokyo, Kashiwanoha 5–1–5, Kashiwa, Chiba 277–8564, Japan Abstract Foraging behavior and diet of Streaked Shearwaters Calonectris leucomelas ORNITHOLOGICAL rearing chicks on Mikura Island was studied using depth and temperature recording SCIENCE data-loggers and stomach contents. Water mass where birds were foraging was esti- © The Ornithological Society mated using sea surface temperature experienced by birds. Birds spent 76–96% of of Japan 2012 their time at sea flying, 4–24% for landing on the water, and made a few (0.5–17.0 per day) shallow (<6 m) dives. Birds made many short (<2 days) and some long (4–10 days) trips. During short trips, birds stayed in the warmer Kuroshio and Kuroshio-Oyashio mixed regions, and fed on Japanese Anchovy Engraulis japonicus, Common Squid Todarodes pacificus and Flying Fish Cypselurus hiraii. During long trips, birds stayed in the colder Oyashio region and fed on anchovy and Pacific Saury Cololabis saira. Birds made more dives during short trips than they did during long trips. Streaked Shearwaters breeding on the island in low—productive Kuroshio water, therefore, adopted dual foraging strategies, and changed their diet and dive frequencies in relation to water masses. Key words Anchovy, Data-logger, Dual foraging, Oyashio During chick rearing, parents of some species of (Chaurand & Weimerskirch 1994; Weimerskirch et albatrosses and shearwaters often adopt dual foraging al. 1997, 2003). strategies, i.e. making short and long foraging trips to Streaked Shearwater Calonectris leucomelas is foraging areas close to and distant from their breed- endemic to East Asia, breeding only around Japan ing site (Weimerskirch et al. 1994; Weimerskirch and southern Korea. Mikura Island (33°52′N, 1998). To feed chicks quickly, parents make many 139°14′E) supports one of the largest colonies short trips to low-productive waters near the colony, (1,750,000–3,500,000 birds, Oka 2004) and is but they make a few long trips to distant and high- located in a low-productive Kuroshio region 300– productive waters to restore their own energy reserves 800 km south of the high-productive Kuroshio- Oyashio mixed and the Oyashio region (Odate 1994; (Received 12 September 2011; Accepted 27 December 2011) Sugiura & Tsunogai 2005). Therefore, birds breeding # Corresponding author, E-mail: [email protected] * Present address: System Engineering Division, Kitami Insti- on this island may adopt dual foraging strategies tute of Technology, Koen-cho 165, Kitami, Hokkaido 090– (Ochi et al. 2010), though foraging behavior and diet 8507, Japan during long and short trips are unknown. ** Present address: National Research Institute of Far Seas Fish- To understand the pattern of exploitation of marine eries, Fisheries Research Agency, 5–7–1 Orido, Shimizu, Shizuoka 424–8633, Japan resources during the chick-rearing period, we studied *** Present address: School of Marine Science and Technology, the foraging behavior of Streaked Shearwaters rear- Tokai University, 3–20–1 Orido, Shimizu, Shizuoka 424– ing chicks on Mikura Island, using depth and tem- 8610, Japan perature recording data-loggers. We also examined **** Present address: Collection Center, National Museum of Nature and Sciences, 3–23–1 Hyakunin-cho, Shinjuku-ku, prey items found in their stomach contents in relation Tokyo 169–0073, Japan to foraging habitats. 9 K. MATSUMOTO et al. STUDY AREA AND METHODS to record depth (0.1 m resolution, 0.5 m accuracy) and temperature (0.018°C resolution, 0.1°C accu- 1) Study site and year racy) every second. D2GTs recorded tail-head and Fieldwork was conducted in the Nango area on ventral-dorsal accelerations at 16 Hz also. Data were Mikura Island, Tokyo (Permit No. 462, Ministry of downloaded onto a laptop computer using Logger the Environment) in 2003, 2005, and 2006. We Tools ver. 3.1 (Marine Micro Technology) and were deployed data-loggers on birds to study foraging analyzed using Igor Pro ver. 4.04 (Wave Metrics Inc.). behavior in 2003 and 2006 and collected stomach To assess the impacts of logger attachments on contents in all three years. We measured trip duration chick growth (slopes of linear regressions of chick using behavioral data collected by data-loggers in mass against date), chicks of parents carrying loggers 2003 and 2006 and using the departure/arrival dates and those without loggers were weighed in the late determined by trapping birds in 2005 and 2006. afternoon every 3–5 days. 2) Data-logger deployment 3) Analyses of pressure and temperature In September and October, data-loggers were The loggers were attached on the middle of the attached to the shearwater breast feathers using Tesa bird’s abdomen. We considered the birds to dive if tape (Tesa AG, Germany). Parents were caught by the depth of loggers was deeper than half of the total hand when they were staying with chicks (<26-day- length of birds (0.5 m). The temperature differences old) in nest boxes during the night, and weighed with between the air and the surface seawater were 0.4– a Pesola spring balance (±5 g). Sexes were deter- 6.3°C during the study period. Thus, we estimated mined using sex-specific calls (Arima & Sugawa that the birds were “landing on the water” during the 2004; Appendix), though we combined male and period between consecutive abrupt changes in tem- female data in this study since the sample size was perature (Fig. 1 b) following Tremblay et al. (2003). small. The birds were released in the nest boxes soon Fluctuation of the tail-head surge acceleration indi- after and recaptured in the nests at night 1–19 days cated that the birds were flapping their wings during later. these landing activities (Fig. 1a). The data-loggers (Little Leonardo, Tokyo) we used The birds made successive water-landings (bouts were DT (14–16 g weight, 47–53 mm length, 15–16 A, C, D, E, in Fig. 1a) or an extended period of land- mm diameter) and D2GT (20 g weight, 60 mm ing (bouts B, F in Fig. 1a) and both of these were length, 5–16 mm diameter). Both were programmed defined as “water-landing bouts”. When birds were Fig. 1. Tail to head surge acceleration (a, in G), recorded temperature (b), and depth (c). Thick horizontal bar in (a) indicates water-landing bouts A–F. Horizontal lines in (b) represents the time when sea surface tempera- ture (SST) was given. 10 Foraging of Streaked Shearwater on the water for an extended period, the recorded their nests, trap doors were set at burrow entrances depths were 0.1–0.2 m (bouts B, F, Fig. 1a, c). Cri- and then the nests were inspected at 21:00–22:00 teria to separate water-landing bouts were calculated and 01:00–03:00 every day to recapture the parents. for each bird using the log-survivorship curves A stone was also placed at the nest chamber entrance (Gentry & Kooyman 1986) of the flight time between to interrupt the returning parents and allow their cap- water landing bouts and were 5.3–6.0 min. ture before they fed their chicks. The resolution of depth allowed us to identify Oils on the surface of stomach samples were col- pressure differences of 0.01 atm (10 hPa) in the air: lected and weighed to the nearest 0.1 g using an approximate pressure difference between sea level electric balance. Then stomach contents were sieved and 100 m altitude. Shearwaters usually fly close (<0.5 mm mesh) and weighed. Samples were pre- (<20 m) to the sea surface (Warham 1996; Pennycuick served in 60% isopropyl alcohol (in 2003) or frozen 2002). As the study site was about 300 m above sea (–20°C in 2005 and 2006). Prey items were identi- level, the rapid decrease and increase of the pressure fied using key external characteristics later in a labo- by 30–35 hPa (corresponding to about 0.3 m change ratory. For well-digested samples, hard parts of the in depth) recorded in the evening and morning, prey (otoliths, vertebrae, fish scales, and squid beaks) respectively, should relate to their arrival to and were used for identification. departure from the colony. To aid in the identification of prey species, partial Sea surface temperatures (SSTs) were recorded sequence data of mitochondrial DNA was extracted when birds were landing on the water. Since the tem- from the muscle tissue of fish in the stomach samples perature-response time (90% time constant) of log- using the phenol-chloroform method or an AquaPure gers between 0°C and 20°C was 12–30 s, we defined Genomic DNA Isolation kit (Bio-Rad). 413 bp in the the SST as the mean temperature during the period cytochrome-b and 320 bp in the 16S ribosomal RNA when the temperature was stable for at least 30 s genes were amplified using PCR. Double-stranded while birds were on the water (Fig. 1b). As Streaked PCR products of both genes, purified using the Shearwaters were rarely observed in waters east of ExoSAP-IT (USB), were subsequently used for 145°E in September and October (Kuroda 1991), we direct-cycle sequencing with dye-labeled terminators then estimated the water masses where birds were (Applied Biosystems).
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