Ecology of Fear in a Colonial Breeder: Colony Structure in Ground-Nesting Great Cormorants Phalacrocorax carbo Reflects Presence of Predators

Ground-nesting Great Cormorants were monitored in three neighbouring colonies at Lake IJsselmeer, The Netherlands. Using aerial photographs taken during peak breeding time, nest density and nearest neighbour distance were determined for four sequential years. In addition, species and number of predators were determined. In total, five mammalian and nine avian predatory species were associated with the Cormorant breeding colonies. Spatial distribution of nests mostly showed dispersed and random patterns rather than a contagious pattern. The latter distribution, with less distance between nests than expected both from a random and equal distribution pattern, was found in the colony of De Ven in 2013 during the last year of its existence. The predator Red Fox Vulpes vulpes arrived at the colony in 2010. In all three colonies, nest density was highest and nearest neighbour distance shortest in colonies with the highest number of predators. At low to moderate predatory pressure, ground-nesting Cormorants left free space between nests that was used by adult birds during take-off and landing. During the last years of its existence the shrinking colony of De Ven showed an almost circular shape, with an extreme nest density and the lowest edge-to-surface area ratio. But with Foxes present, breeding at the fringe still caused greater losses due to direct predation. Breeding success fluctuated synchronously between colonies but was lower in colonies where the number of predators was higher. The arrival of Red Foxes in De Ven caused extreme losses of young and over the years resulted in a strong decline in number of breeders, eventually leading to complete abandoning of the site in 2014. Large gulls formed another important group of predators but did not cause the Cormorants to abandon the breeding site. In the Vooroever colony, bush and tree cover supplied shelter and allowed birds to breed in greater density without causing nearest neighbour density to decrease, as was the case when no cover was available. Greater nest density and reduced nearest neighbour distances are considered to be a pro-active response by individual birds to the presence of predators. When predator numbers increased, the within-colony open spaces that normally exist under circumstances of moderate density were filled up with nests, leaving little or no room for landing and departure. This leads to reduced edge effects and a circular shape of the colony, thereby potentially limiting predation risk. As a consequence of extreme high nest densities, breeding success was lower due to interference by other Cormorants. This study is the first to show that colony structure in waterbirds is affected by forces of attraction and repulsion between founding birds that are predator driven.