title: Daily activity budgets reveal a quasi-flightless stage during non-breeding in Hawaiian albatrosses.
abstract: BACKGROUND
  Animals adjust activity budgets as competing demands for limited time and energy shift across life history phases. For far-ranging migrants and especially pelagic seabirds, activity during breeding and migration are generally well studied but the "overwinter" phase of non-breeding has received less attention. Yet this is a critical time for recovery from breeding, plumage replacement and gaining energy stores for return migration and the next breeding attempt. We aimed to identify patterns in daily activity budgets (i.e. time in flight, floating on the water's surface and active foraging) and associated spatial distributions during overwinter for the laysan Phoebastria immutabilis and black-footed P. nigripes albatrosses using state-space models and generalized additive mixed-effects models (GAMMs). We applied these models to time-series of positional and immersion-state data from small light- and conductivity-based data loggers.
  
  
  RESULTS
  During overwinter, both species exhibited a consistent 'quasi-flightless' stage beginning c. 30 days after initiating migration and lasting c. 40 days, characterized by frequent long bouts of floating, very little sustained flight, and infrequent active foraging. Minimal daily movements were made within localized areas during this time; individual laysan albatross concentrated into the northwest corner of the Pacific while black-footed albatross spread widely across the North Pacific Ocean basin. Activity gradually shifted toward increased time in flight and active foraging, less time floating, and greater daily travel distances until colony return c. 155 days after initial departure.
  
  
  CONCLUSIONS
  Our results demonstrate that these species make parallel adjustments to activity budgets at a daily time-scale within the overwinter phase of non-breeding despite different at-sea distributions and phenologies. The 'quasi-flightless' stage likely reflects compromised flight from active wing moult while the subsequent increase in activity may occur as priorities shift toward mass gain for breeding. The novel application of a GAMM-based approach used in this study offers the possibility of identifying population-level patterns in shifting activity budgets over extended periods while allowing for individual-level variation in the timing of events. The information gained can also help to elucidate the whereabouts of areas important at different times across life history phases for far-ranging migrants.
authors.affiliation: Biology Department, Dalhousie University, Halifax, NS Canada.
authors.forename: Sarah E
authors.initials: SE
authors.lastname: Gutowsky
authors.affiliation: Fish Ecology & Conservation Physiology Lab, Carleton University, Ottawa, ON Canada.
authors.forename: Lee Fg
authors.initials: LF
authors.lastname: Gutowsky
authors.affiliation: Department of Biological Sciences, Macquarie University, Sydney, NSW Australia.
authors.forename: Ian D
authors.initials: ID
authors.lastname: Jonsen
authors.affiliation: Biology Department, Dalhousie University, Halifax, NS Canada.
authors.forename: Marty L
authors.initials: ML
authors.lastname: Leonard
authors.affiliation: USFWS, Pacific Region, Migratory Birds and Habitat Programs, Portland, OR USA.
authors.forename: Maura B
authors.initials: MB
authors.lastname: Naughton
authors.affiliation: USFWS, Pacific Region, Migratory Birds and Habitat Programs, Portland, OR USA.
authors.forename: Marc D
authors.initials: MD
authors.lastname: Romano
authors.affiliation: Department of Biological Sciences, San Jose State University, San Jose, CA USA ; Institute of Marine Sciences, University of California, Santa Cruz, CA USA.
authors.forename: Scott A
authors.initials: SA
authors.lastname: Shaffer
keywords: Activity budget; Behaviour; Biologging; Flight; Migration; Moult; Non-breeding, Overwinter; North Pacific; Seabirds