Foraging Tactics of Chick-Rearing Crozet Shags: Individuals Display Repetitive Activity and Diving Patterns Over Time
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Polar Biol (2006) 29: 562–569 DOI 10.1007/s00300-005-0089-y ORIGINAL PAPER Timothe´e R. Cook Æ Yves Cherel Æ Yann Tremblay Foraging tactics of chick-rearing Crozet shags: individuals display repetitive activity and diving patterns over time Received: 27 June 2005 / Revised: 2 November 2005 / Accepted: 5 November 2005 / Published online: 30 November 2005 Ó Springer-Verlag 2005 Abstract It is in the interest of resident and long-lived of the ocean ecosystem. To understand how they man- benthic foragers to learn to apply efficient foraging age foraging in the marine environment, researchers tactics throughout their lifetime, thus increasing their have devoted considerable effort in recent years to individual efficiency. To test whether individuals are understanding the foraging ecology of marine mammals capable of applying an individual-specific foraging pat- and seabirds (Jouventin and Weimerskirch 1990; Wilson tern, we checked for the existence of established foraging et al. 1992; Costa 1993). routines. Using ventrally attached time-depth recorders, Nevertheless, individual foraging behaviour of mar- we studied the individual foraging tactics of chick-rear- ine top predators has been poorly investigated, mainly ing Crozet shags (Phalacrocorax melanogenis, Blyth because it implies studying individuals over time (lon- 1860), as measured by the consistency in individual daily gitudinal studies), and researchers generally focus their activity patterns and diving profiles over time. Individ- work at a species or population level (cross-sectional uals displayed a fidelity to the time of first daily trip to studies: Bolnick et al. 2002; Tremblay and Cherel 2003). sea and also a strong fidelity to one, two or three depth Studies on foraging thus tend to define the range of the ranges day after day. We suggest foraging area fidelity, a niche that is optimal for the population (Krebs and behaviour that could help increase foraging efficiency Davies 1987). One of the reasons for this bias toward thanks to the memorization of the bottom’s topography cross-sectional studies comes from the statistical sword and the habits of its fauna, as a hypothesis for explaining of Damocles: ‘‘individual observations are meaning- some of these patterns. We propose the question of less’’. Yet, many species are in fact composed of indi- foraging area fidelity should be more specifically ad- vidual specialists that use small subsets of the dressed in the future. population’s niche. A key characteristic of individual specialization is the repetition of a specific behaviour and/or diet over time (Amundsen et al. 1997; Estes et al. 2003). This was described in a variety of species, on both a dietary and/or a behavioural level (see Bolnick et al. Introduction 2003 for review). Specialization by individuals has been shown not only to avoid competition between conspe- Seabirds and non-cetacean marine mammals have very cifics, but also to considerably increase individual effi- distinct areas for breeding (on land), and for feeding (at ciency in finding food through learning processes (e.g. in sea). Consequently, they face a permanent challenge, insects and fish: Heinrich 1976; Werner et al. 1981). which has shaped their life histories (Ashmole 1971; The main challenge for an individual in finding food Fiedler 2002): how to find mobile food in the immensity in the ocean is suspected to be food predictability. Sea- birds generally rely on more or less predictable oceanic features associated with food availability such as fronts, T. R. Cook (&) Æ Y. Cherel upwellings, or continental shelves (Hunt 1991). These Centre d’Etudes Biologiques de Chize´, CEBC - CNRS UPR 1934, features are obviously more predictable when they are Villiers-en-bois, 79360 Beauvoir-sur-Niort, France E-mail: [email protected] static rather than when they are dynamic. Some species Tel.: +33-5-49096111 of marine mammals and diving seabirds target benthic Fax: +33-5-49096526 or epibenthic prey (e.g. cormorants, penguins, sea lions, respectively, Orta 1992; Tremblay and Cherel 2000; Present address: Y. Tremblay Long Marine Laboratory, University of California Santa Cruz, Costa and Gales 2003), and thus it is reasonable to as- 100 Shaffer Road, Santa Cruz, CA 95060, USA sume they may rely on topographic cues to find their 563 prey. Topographic cues give individuals the possibility The TDRs were 8.6·2.0·1.1 cm3, and weighed 27 g, of memorizing the location and quality of distinct for- corresponding to 1.1% of the birds’ mean body mass. aging grounds on a very precise scale, which is virtually Recorder tips were streamlined to reduce drag. The impossible to do using dynamic cues. pressure and light sensors reacted immediately to Crozet shags (Phalacrocorax melanogenis, Blyth changes in the environment, while the temperature sen- 1860) are non-migratory benthic foragers, which capture sor had a longer reaction time: TDRs were therefore essentially benthic fish during the breeding season (Ri- programmed to record depth and light every 1 s and doux 1994). At Possession Island, Crozet Archipelago, external temperature every 5 s. Depth and temperature they dive very close to the colony (within a mean range resolution were ±1 m and ±0.1°C, respectively. Light of 1–2 km), where the sea bottom falls away quickly (arbitrary scale) was linearly related to log10lx (Wanless from the coast, reaching 200 m in depth after only et al. 1999). Memory was 2.03 MB. 5 km. The Crozet shags work hard during chick- This study was approved by the ethics committee of rearing compared to other species of cormorants, and the French Polar Institute (Institut Paul Emile Victor, exhibit sexual differences in their foraging ecology, IPEV). All animals in this study were cared for in males diving deeper, foraging in different areas, eating accordance with its guidelines. different prey sizes, spending daily less time underwater, or having different activity schedules compared to fe- Data analysis and statistics males (Tremblay et al. 2005; Cook et al. unpublished), possibly as a consequence of intra-specific competition Time budget analysis was performed using the advan- or food limitation. tages inherent in the ventral attachment technique Being resident and long-lived benthic foragers, indi- (Tremblay et al. 2003). Foraging trips and flights were vidual Crozet shags should learn to apply efficient for- determined by simultaneous reading of depth, light, and aging tactics throughout their lifetime, thus increasing temperature profiles in relation to time (for details on their individual efficiency when foraging under situations the procedure at Crozet, see Tremblay et al. 2005). Using of competition or food limitation (Estes et al. 2003). In our own designed software for dive analysis (see for this perspective, we tested whether individuals were in- example Tremblay and Cherel 2000, 2003), analysis of deed capable of elaborating a foraging strategy (or tac- dive profiles was performed in order to gather dive tic), in the sense of applying a particular foraging pattern. depths for every individual. Mean bird flight speeds were To do so, we checked for the existence of established assumed 58 km/h (Spear and Ainley 1997). foraging routines in individual chick-rearing Crozet Individual foraging behaviours were compared from shags by measuring day to day individual activity pat- 1 day to the next, in order to determine possible estab- terns and diving depths using time-depth recorders (TDR lished and repeated patterns, suggesting individual tac- MK7 Wildlife Computers, Woodinville, Washington, tics. Time of first departure to sea was compared day to USA). day, over a period of 4–6 days. Mean daily flight dura- tion for all daily trips (to commute to the sea surface and Materials and methods to come back to the colony) was compared to daily mean dive depth over 4–5 days (in order to use days with full daily data on foraging activity). Daily mean and Study site, birds and general procedure maximum dive depths were examined in the same manner, and dive frequencies in relation to dive depth Field work was conducted between the 28 of January were compared on a daily basis. and the 9 of February 1999 at Pointe Basse (46°21¢S, Finally, the data were analysed statistically using the 51°42¢E), Possession Island, Crozet Archipelago (Local non-parametric Spearman rank correlation test (to test Time: UTC+3h20) in a small colony of Crozet shags (13 the effect of different variables on the departure time of nests). birds), and regressions otherwise, when data permitted it. Twelve individuals (six males and six females from six Software used was STATISTICA 6.1 (CopyrightÓ Stat- pairs) of Crozet shags were studied during the chick- Soft, Inc. 1984–2004) and SIGMAPLOT 8.0 (CopyrightÓ rearing period (1.8±0.7 chicks per nest, age of chicks SPSS, Inc. 1986–2001), with an a-level of significance of between 10 and 50 days). The birds were captured at P<0.05. The mean values are given ±SD. night by hand (both partners are then present at the nest), and weighed (precision ±25 g) using a spring balance, measured for sexing (Malacalaza and Hall 1988), and a TDR was attached ventrally according to Results Tremblay et al. (2003) for a 4–6 day deployment period, using cyanoacrylate glue (Loctite 401) and plastic ties At sea behaviour was successfully recorded for 4–6 (total initial handling time 5 min). At the end of the consecutive days in 12 Crozet shag individuals, days study period, loggers were retrieved at a moment when with full daily data ranging from 4 to 5. Detailed birds came back from the sea (immediately after return observation of several foraging parameters for each bird to the nest). separately showed remarkable consistency over time. 564 males over time were not explained by the daily time spent at sea (no relation between schedule and time 2 14 spent at sea: R =0.0045, N=21, P=0.773).