Social Hierarchy within Communal Foraging Flocks of Great Cormorants Phalacrocorax carbo as Reflected by Differences in Prey Composition and Food Intake at the Roost

Day-fresh Great Cormorant pellets were collected at a huge single winter roost at Kreupel island in Lake IJsselmeer, The Netherlands. The roost is situated on a sandy island, 3 m above the water level and is regularly used by 1000–8000+ Cormorants. By examining pellets and thus reconstructing the previous day's meal of Cormorants gathering at the roost, we explored spatio-temporal differences in the diet of individual birds. We found that structural differences in pellets (size, colour and structure) are related to their content (fish species and fresh mass of a daily meal). Structurally larger and lighter coloured pellets contained larger and coarser fish and represented a higher total fish mass. Pellets of different size and structure were non-randomly distributed at the roost. Birds positioned centrally in the roost produced on average larger pellets than birds at the edges. Cormorants having few or no food remains in their pellets were more frequently recorded on the edge of the roost than in the centre and in particular, they were more likely to be located to the rear of the roost. The same was true for pellets showing a bloody signature which may be linked to either injury by sharp fin rays or to a parasite induced effect. Pellet data were corroborated by observations of differential distribution of faecal splashes of Cormorants, showing that individuals in the centre of the roost produced most guano. Clear diet differences existed within the roost between birds in the centre and elsewhere in the roost, whereas smaller differences existed between the front, edge and rear of the roost. Our data suggest a social hierarchy based on higher foraging success of birds returning earlier to the roost and settling in the centre, hereby forcing later arriving and less successful Cormorants towards the edge, rear and front position. We conclude that position at the roost is a reflection of previous foraging success, possibly caused by differences in dominance between Cormorants during the period of communal foraging. From a methodological point of view, our study demonstrates that, because of structural differences in pellets and spatial differences in occurrence, it is important to collect pellets randomly if one is to describe a proper diet sample by using pellets at any given site. Especially in marshlands and under trees and bush growth with a lot of litter, the likely collection of larger, more conspicuous pellets will cause a bias towards the larger fish species and sizes. Also sampling only at edge locations or specifically at the centre of a roost may give biased results, as this may result in an under- or overestimation in reconstructed fish uptake (e.g. fish size, mass, species) from the waterbody under study.