When the "selfish herd" becomes the "frozen herd": spatial dynamics and population persistence in a colonial seabird

Aggregations are common in ecological systems at a range of scales and may be driven by exogenous constraints such as environmental heterogeneity and resource availability or by "self-organizing" interactions among individuals. One mechanism leading to self-organized animal aggregations is captured by Hamilton's "selfish herd" hypothesis, which suggests that aggregations may be driven by an individual's effort to minimize their risk of predation by surrounding themselves with conspecifics. We demonstrate that aggregations observed in Adélie Penguin (Pygoscelis adeliae) colonies are a convolution of both self-organized dynamics and external forcing arising from landscape terrain. In fluid, highly mobile aggregations, individuals are constantly moving in response to changing environmental conditions, the locations of predators, or the movements of conspecifics. However, when the ability to rearrange is limited and spatial reconfiguration occurs on slower time scales than changes in population size, systems may become trapped in suboptimal arrangements. We use simulated annealing to demonstrate that Adélie Penguin colonies are frozen in suboptimal spatial arrangements, and employ an individual-based modeling approach to demonstrate that this suboptimal spatial configuration is driven by a convolution of nest site fidelity and stochastic events at the level of individual nests. The resulting spatial dynamics are responsible for a hysteretic response to long-term changes in abundance. We find that declining abundance leads to fragmentation even in a homogeneous environment, which has population-level consequences for reproductive success because predation is biased towards colony edges. Strong edge effects from heterogeneous predation coupled with fragmentation in response to population declines create a positive feedback cycle that can accelerate population decline. This work provides a mechanistic understanding of complex spatial structuring in penguin colonies, provides a link between current spatial patterning and past dynamics, and suggests the possibility of critical collapse in seabird populations.