Avian Predator Buffers Against Variability in Marine Habitats with Flexible Foraging Behavior
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See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/323201633 Avian predator buffers against variability in marine habitats with flexible foraging behavior Article in Marine Biology · February 2018 DOI: 10.1007/s00227-018-3304-4 CITATIONS READS 3 190 10 authors, including: John F Piatt Mayumi L. Arimitsu United States Geological Survey United States Geological Survey 204 PUBLICATIONS 10,798 CITATIONS 32 PUBLICATIONS 450 CITATIONS SEE PROFILE SEE PROFILE Gary Drew Martin Renner United States Geological Survey 42 PUBLICATIONS 721 CITATIONS 33 PUBLICATIONS 590 CITATIONS SEE PROFILE SEE PROFILE Some of the authors of this publication are also working on these related projects: Reconnaissance of the intertidal and shallow subtidal benthic of Kasatochi Island following the 2008 volcanic eruption. View project Seabird ecology View project All content following this page was uploaded by John F Piatt on 19 February 2018. The user has requested enhancement of the downloaded file. Marine Biology (2018) 165:47 https://doi.org/10.1007/s00227-018-3304-4 ORIGINAL PAPER Avian predator bufers against variability in marine habitats with fexible foraging behavior Sarah K. Schoen1 · John F. Piatt1 · Mayumi L. Arimitsu2 · Brielle M. Hefin2 · Erica N. Madison1 · Gary S. Drew1 · Martin Renner3 · Nora A. Rojek4 · David C. Douglas2 · Anthony R. DeGange1 Received: 11 September 2017 / Accepted: 31 January 2018 © This is a U.S. government work and its text is not subject to copyright protection in the United States; however, its text may be subject to foreign copyright protection 2018 Abstract How well seabirds compensate for variability in prey abundance and composition near their breeding colonies infuences their distribution and reproductive success. We used tufted pufns (Fratercula cirrhata) as forage fsh samplers to study marine food webs from the western Aleutian Islands (53°N, 173°E) to Kodiak Island (57°N, 153°W), Alaska, during August 2012–2014. Around each colony we obtained data on: environmental characteristics (sea surface temperature and salinity, seafoor depth and slope, tidal range, and chlorophyll-a), relative forage fsh biomass (hydroacoustic backscatter), and seabird community composition and density at-sea. On colonies, we collected pufn chick-meals to characterize forage communities and determine meal energy density, and measured chicks to obtain a body condition index. There were distinct environmen- tal gradients from west to east, and environmental variables difered by ecoregions: the (1) Western-Central Aleutians, (2) Eastern Aleutians, and, (3) Alaska Peninsula. Forage fsh biomass, species richness, and community composition all difered markedly between ecoregions. Forage biomass was strongly correlated with environmental gradients, and environmental gradients and forage biomass accounted for ~ 50% of the variability in at-sea density of tufted pufns and all seabird taxa combined. Despite the local and regional variability in marine environments and forage, the mean biomass of prey delivered to pufn chicks did not difer signifcantly between ecoregions, nor did chick condition or pufn density at-sea. We conclude that pufns can adjust their foraging behavior to produce healthy chicks across a wide range of environmental conditions. This extraordinary fexibility enables their overall success and wide distribution across the North Pacifc Ocean. Introduction Seabirds inhabit nearly all marine environments from the Responsible Editor: Y. Cherel. tropics to the poles. Physiological and morphological adap- tations of individual species to those environments shape Reviewed by C. Cotté and an undisclosed expert. their behavior, ecology, and demography (Schreiber and Electronic supplementary material The online version of this Burger 2001), and ultimately determine the marine ecore- article (https://doi.org/10.1007/s00227-018-3304-4) contains gions in which they can survive. Seabird life history charac- supplementary material, which is available to authorized users. teristics have evolved to account for the energetic constraints they face in supplying food to their chicks at centralized * Sarah K. Schoen [email protected] colonies while foraging for patchily distributed prey in sur- rounding waters (Lack 1968). For example, generalists with 1 US Geological Survey Alaska Science Center, 4210 fexibility in diet and foraging location can compensate for University Ave, Anchorage, AK 99508, USA variation in prey availability by adjusting their time budg- 2 US Geological Survey Alaska Science Center, 250 Egan Dr, ets and provisioning strategies to optimize breeding success Juneau, AK 99801, USA (Piatt et al. 2007; Ainley et al. 2014). 3 Tern Again Consulting, 811 Ocean Drive Loop, Homer, Aspects of seabird breeding biology (e.g., diet, reproduc- AK 99603, USA tive success, chick growth, fedging success) and distribu- 4 US Fish and Wildlife Service Alaska Maritime National tion and density at sea have been correlated with changes in Wildlife Refuge, 95 Sterling Hwy, Homer, AK 99603, USA Vol.:(0123456789)1 3 47 Page 2 of 14 Marine Biology (2018) 165:47 food supply and direct or indirect (via food supply) efects To address this hypothesis, we studied tufted pufns at of marine climate (e.g., Gjerdrum et al. 2003; Frederiksen 25 colonies (each on a diferent island) located between et al. 2005; Piatt et al. 2007; Renner et al. 2012; Gladics the western Aleutian Islands and Kodiak Island, Alaska et al. 2015; Sydeman et al. 2017). The use of seabird diets (Fig. 1). Piatt and Springer (2007) proposed several ecore- to study forage fsh communities can be an efective method gion boundaries within this area based on mesoscale pat- for assessing the status of marine food webs and ecosystem terns in biological indicators and topographic, bathymetric, health over space and time (Sydeman et al. 2017; Piatt et al. and oceanographic features, including (Fig. 1): Amchitka 2018). Tufted pufns (Fratercula cirrhata) are a particularly Pass separating the Central Aleutians from the Western useful species to study because of their wide distribution Aleutians; Samalga Pass separating the Eastern Aleutians across a large range of environmental and prey conditions from the Central Aleutians; and Unimak Pass separating the from California to Japan, and from the subtropics to the Arc- Alaska Peninsula from the Eastern Aleutians. Samalga Pass, tic (Piatt and Kitaysky 2002; Drew et al. 2015). In addition, in particular, has been identifed as a demarcation of “pro- they are a colonial piscivore with a broad diet, and they feed found ecosystem change” in terms of oceanography (Hunt their chicks whole food items, which can be easily identi- and Stabeno 2005; Ladd et al. 2005a), zooplankton (Coyle fed and measured (Piatt and Kitaysky 2002). Unlike many 2005), fsh (Logerwell et al. 2005; Sydeman et al. 2017), seabirds that depend on just a few select forage species in seabirds (Byrd et al. 2005; Jahncke et al. 2005; Ladd et al. their breeding range (Ainley et al. 2002; Gladics et al. 2015), 2005b) and marine mammals (Sinclair et al. 2005). tufted pufns are generalist predators whose diverse diets Against this backdrop of distinct ecoregions, we assessed refect the availability of a variety of prey species (Piatt and local food webs from the diets of tufted pufns breeding Kitaysky 2002; Sydeman et al. 2017). Indeed, in the Gulf at 25 colonies across the entire area. We then character- of Alaska and the Aleutian Islands pufns feed their chicks ized local forage fsh communities with respect to marine more than 40 species of forage fsh, including 16 main prey habitats (e.g., Abookire and Piatt 2005; Logerwell et al. groups (Sydeman et al. 2017). We hypothesized that, along 2005). We collected data on environmental variables that with some key foraging adaptations (e.g., multiple prey load- have been shown to infuence upper trophic levels, including ers, burrow-nesters), the diversity in diet of tufted pufns seabird diets, hatch dates, growth rates, and fedging success during chick rearing may help them bufer against regional (Gjerdrum et al. 2003; Renner et al. 2012; Gladics et al. diferences in habitat and prey quality. 2015). We also estimated forage fsh biomass (hydroacoustic Fig. 1 Tufted Pufn colonies sampled during August, 2012–2014 dots. Important island passes are denoted with double lines and from in the Aleutian Islands and Gulf of Alaska. Colonies within difer- west to east are: (1) Buldir, (2) Amchitka, (3) Samalga, and (4) Uni- ent marine ecoregions [Western-Central Aleutians (WCA), Eastern mak passes. Bathymetry is displayed with darker colors representing Aleutians (EA), and Alaska Peninsula (AP)] are denoted by colored deeper depths 1 3 Marine Biology (2018) 165:47 Page 3 of 14 47 backscatter index) and seabird densities at sea around each Kitaysky 2002) and 70 km for Atlantic pufns (Harris et al. colony. At colonies, we collected data on tufted pufn chick 2012), although most birds probably forage much closer body condition. The specifc objectives of our study were to: to colonies (e.g., < 40 km) as adults try to minimize the (1) assess geographic variability in pufn diet in relation to amount of time spent commuting to provision chicks (Piatt variation in marine habitat, (2) relate the at-sea density of et al. 2007; Harris et al. 2012). We believe that our surveys, tufted pufns and marine birds to marine habitat characteris- conducted in close proximity to colonies, provide a reason- tics and prey availability, and, (3) assess which (if any) envi- able proxy for the marine habitats likely to be encountered ronmental and food factors infuence pufn chick condition. by pufns foraging at somewhat greater distances from their colonies. In addition to environmental data, we collected data on tufted pufn chicks at each colony. We selected colo- Materials and methods nies for sampling based on biological (colony size, historical data, etc.) and logistical (weather, accessibility, etc.) consid- Study area erations. We attempted to sample each of 23 colonies during the middle of chick-rearing (approximately August 7–22) in Our study area spanned 2400 km from the terminus of the 2012, 2013 and 2014 (Table 1; Fig.