title: Vortex Formation and Foraging in Polyphenic Spadefoot Toad Tadpoles.
abstract: Animal aggregations are widespread in nature and can exhibit complex emergent properties not found at an individual level. We investigate one such example here, collective vortex formation by congeneric spadefoot toad tadpoles: Spea bombifrons and S. multiplicata. Tadpoles of these species develop into either an omnivorous or a carnivorous (cannibalistic) morph depending on diet. Previous studies show S. multiplicata are more likely to develop into omnivores and feed on suspended organic matter in the water body. The omnivorous morph is frequently social, forming aggregates that move and forage together, and form vortices in which they adopt a distinctive slowly-rotating circular formation. This behaviour has been speculated to act as a means to agitate the substratum in ponds and thus could be a collective foraging strategy. Here we perform a quantitative investigation of the behaviour of tadpoles within aggregates. We found that only S. multiplicata groups exhibited vortex formation, suggesting that social interactions differ between species. The probability of collectively forming a vortex, in response to introduced food particles, increased for higher tadpole densities and when tadpoles were hungry. Individuals inside a vortex moved faster and exhibited higher (by approximately 27%) tailbeat frequencies than those outside the vortex, thus incurring a personal energetic cost. The resulting environmental modification, however, suggests vortex behaviour may be an adaptation to actively create, and exploit, a resource patch within the environment.
authors.affiliation: Department of Zoology, University of Oxford, Tinbergen Building, South Parks Road, Oxford OX1 3PS, UK.
authors.forename: Sepideh
authors.initials: S
authors.lastname: Bazazi
authors.forename: Karin S
authors.initials: KS
authors.lastname: Pfennig
authors.forename: Nils Olav
authors.initials: NO
authors.lastname: Handegard
authors.forename: Iain D
authors.initials: ID
authors.lastname: Couzin