title: Shadowed by scale: subtle behavioral niche partitioning in two sympatric, tropical breeding albatross species.
abstract: BACKGROUND
  To meet the minimum energetic requirements needed to support parents and their provisioned offspring, the timing of breeding in birds typically coincides with periods of high food abundance. Seasonality and synchrony of the reproductive cycle is especially important for marine species that breed in high latitudes with seasonal booms in ocean productivity. Laysan and black-footed albatrosses breeding in the northwestern Hawaiian Islands have a dual reliance on both seasonally productive waters of high latitudes and on nutrient-poor waters of low latitudes, because their foraging ranges contract during the short but critical brood-guard stage. Therefore, these species face an additional constraint of having to negotiate nutrient-poor waters during the most energetically-demanding stage of the breeding cycle. This constriction of foraging range likely results in a higher density of foraging competitors. Thus, our aim was to understand how Hawaiian albatross partition resources both between and within species in this highly constrained breeding stage while foraging in less productive waters and simultaneously experiencing increased competition. High-precision GPS dataloggers were deployed on black-footed (Phoebastria nigripes, n=20) and Laysan (Phoebastria immutabilis, n=18) albatrosses during the brood-guard stage of the breeding season in 2006 (n=8), 2009 (n=13), 2010 (n=16) and 2012 (n=1). We used GPS data and movement analyses to identify six different behavioral states in foraging albatrosses that we then used to characterize foraging trips across individuals and species. We examined whether variations in behavior were correlated with both intrinsic factors (sex, body size, body condition) and extrinsic factors (lunar phase, wind speed, year).
  
  
  RESULTS
  Behavioral partitioning was revealed both between and within species in Hawaiian albatrosses. Both species were highly active during chick-brooding trips and foraged across day and night; however, Laysan albatrosses relied on foraging at night to a greater extent than black-footed albatrosses and exhibited different foraging patterns at night. For both species, foraging along direct flight paths and foraging on the water in a "sit-and-wait" strategy were just as prevalent as foraging in a searching flight mode, indicating flexibility in foraging strategies in Hawaiian albatross. Both species strongly increased drift forage at night when the lunar phase was the darkest, suggesting Hawaiian albatross feed on diel vertically-migrating prey to some extent. Black-footed albatrosses showed greater variation in foraging behavior between individuals which suggests a higher level of intra-specific competition. This behavioral variability in black-footed albatrosses was not correlated with sex or body size, but differences in body condition suggested varying efficiencies among foraging patterns. Behavioral variability in Laysan albatrosses was correlated with sex, such that females exhibited greater flight foraging than drift foraging, had longer trip durations and flew farther maximum distances from the breeding colony, but with no difference in body condition.
  
  
  CONCLUSIONS
  Fine-scale movement data and an analysis of multiple behavioral states identified behavioral mechanisms that facilitate coexistence within a community of albatross during a critical life-history period when energetic demands are high, resources are limited, and competition for food is greatest.
authors.affiliation: Department of Ocean Sciences, University of California Santa Cruz, 100 Shaffer Road, Santa Cruz, CA 95060 USA.
authors.forename: Melinda G
authors.identifier: 0000-0003-0572-0026
authors.initials: MG
authors.lastname: Conners
authors.affiliation: Environmental Research Division, Southwest Fisheries Science Center, NOAA Fisheries, 1352 Lighthouse Avenue, Pacific Grove, CA 93950 USA ; Joint Institute for Marine and Atmospheric Research, University of Hawai'i at Manoa, 1000 Pope Road, Honolulu, HI 96822 USA.
authors.forename: Elliott L
authors.initials: EL
authors.lastname: Hazen
authors.affiliation: Department of Ocean Sciences, University of California Santa Cruz, 100 Shaffer Road, Santa Cruz, CA 95060 USA ; Institute of Marine Sciences, University of California, Santa Cruz, CA 95060-5730 USA.
authors.forename: Daniel P
authors.initials: DP
authors.lastname: Costa
authors.affiliation: Institute of Marine Sciences, University of California, Santa Cruz, CA 95060-5730 USA ; Department of Biological Sciences, San José State University, One Washington Square, San Jose, CA 95192 USA.
authors.forename: Scott A
authors.initials: SA
authors.lastname: Shaffer
keywords: Albatrosses; Behavioral plasticity; Coexistence; Foraging strategies; Intraspecific variability; Niche partitioning; Nocturnal foraging; Phoebastria; “Sit-and-wait” foraging