title: The intensity of horizontal and vertical search in a diving forager: the harbour seal.
abstract: BACKGROUND
  Free ranging foraging animals can vary their searching intensity in response to the profitability of the environment by modifying their movements. Marine diving animals forage in a three dimensional space and searching intensity can be varied in both the horizontal and vertical planes. Therefore understanding the relationship between the allocation of searching effort in these two spaces can provide a better understanding of searching strategies and a more robust identification of foraging behaviour from the multitude of foraging indices (FIs) available. We investigated the movement of a widespread marine coastal predator, the harbour seal (Phoca vitulina), and compared two sets of foraging indices reflecting searching intensity respectively in the horizontal plane (displacement speed, extensive vs. intensive movement types, residence time) and in the vertical dimension (time at the bottom of a dive). We then tested how several factors (dive depth, direction of the trip with respect to haul-out site, different predatory tactics, the presence of factors confounding the detection of foraging, and temporal resolution of the data) affected their relationships.
  
  
  RESULTS
  Overall the indices only showed a very weak positive correlation across the two spaces. However controlling for various factors strengthened the relationships. Resting at sea, a behaviour intrinsically static in the horizontal plane, was found to be strongly negatively related to the time spent at the bottom of the dives, indirectly weakening the relationship between horizontal and vertical foraging indices. Predatory tactic (benthic vs. pelagic) was found to directly affect the relationship. In benthic (as opposed to pelagic) foraging a stronger positive relationship was found between vertical and horizontal indices.
  
  
  CONCLUSIONS
  Our results indicated that movement responses, leading to an intensification of search, are similar in the two spaces (positive relationship), but additional factors need to be taken into account for this relationship to emerge. Foraging indices measuring residence in the horizontal plane tend to be inflated by resting events at sea, while vertical indices tend to distinguish mainly between periods of activity and inactivity, or of benthic and pelagic foraging. The simultaneous consideration of horizontal and vertical movements, as well as topographic information, allows additional behavioural states to be inferred, providing greater insight into the interpretation of foraging activity.
authors.affiliation: Institute of Marine Research, Tromsø, Norway ; University of Tromsø, Tromsø, Norway.
authors.forename: Virginie
authors.initials: V
authors.lastname: Ramasco
authors.affiliation: University of Tromsø, Tromsø, Norway.
authors.forename: Frédéric
authors.initials: F
authors.lastname: Barraquand
authors.affiliation: Akvaplan-niva, Tromsø, Norway.
authors.forename: Martin
authors.initials: M
authors.lastname: Biuw
authors.affiliation: Sea Mammal Research Unit, St. Andrews, UK.
authors.forename: Bernie
authors.initials: B
authors.lastname: McConnell
authors.affiliation: Institute of Marine Research, Tromsø, Norway.
authors.forename: Kjell T
authors.initials: KT
authors.lastname: Nilssen
keywords: Benthic diving; Foraging index; Movement response; Predatory strategy; Profitability; Resting