Shapes of krill swarms and fish schools emerge as aggregation members avoid predators and access oxygen

Abstract

Many types of animals exhibit aggregative behavior: birds flock, bees swarm, fish shoal, and ungulates herd. Terrestrial and aerial aggregations can be observed directly, and photographic techniques have provided insights into the behaviors of animals in these environments and data against which behavioral theory can be tested. Underwater, however, limited visibility can hamper direct observation, and understanding of shoaling remains incomplete. We used multibeam sonar to observe three-dimensional structure of Antarctic krill shoals acoustically. Shoal size and packing density varied greatly, but surface area:volume ratios (roughnesses) were distributed narrowly about ∼3.3 m(-1). Shoals of clupeid fish (e.g., sardine, anchovy) from geographically and oceanographically diverse locations have very similar roughnesses. This common emergent shape property suggests common driving forces across diverse ecosystems. Group behavior can be complex, but a simple tradeoff--that we model--in which individual fish and krill juggle only their access to oxygen-replete water and exposure to predation can explain the observed shoal shape. Decreasing oxygen availability in a warming world ocean may impact shoal structure: because structure affects catchability by predators and fishers, understanding the response will be necessary for ecological and commercial reasons.