The formation of social groups has important impacts on fitness for many animal species, with differences in group compositions resulting in a range of fitness outcomes for individuals. Recent interest in mixed‐species grouping, which extends from a large body of literature invested in understanding single‐species grouping, highlights novel complexities of group formation which relate to phenotypic, behavioural and physiological differences that naturally exist between species. Among fishes, mixed‐species shoaling is a common form of social grouping behaviour displayed across a range of marine and freshwater ecosystems. Research explaining mixed‐species shoaling shows some overlap with explanations for single‐species shoaling; however, it also demonstrates that distinct differences between species give rise to unique cost‐benefit trade‐offs which need to be incorporated into conceptual models of mixed‐species shoaling behaviour. Unique predation related trade‐offs may arise from inefficiency of the confusion effect, variation in vigilance between species and unequal species‐preferences shown by predators, whilst unique foraging‐related trade‐offs may arise from diet partitioning, variations in foraging behaviour and differences in competitive abilities between species. We review the literature on fitness outcomes associated with mixed‐species shoaling and present a new theoretical framework to explain the cost‐benefit trade‐offs for individuals within mixed‐species shoals. The framework incorporates both trade‐offs arising from differences between species and those arising from group size, the former having been largely ignored due to a focus on single‐species shoaling. Our framework is designed to inform future research striving to explain mixed‐species shoaling behaviour.
The formation of mixed‐species social groups, whereby heterospecifics form and maintain either transient or stable groups with each other, can confer substantial fitness benefits to individuals. Such benefits may arise via multiple mechanisms associated with both predation avoidance and foraging efficiency. In fishes, mixed‐species shoaling reportedly occurs where displaced tropical species (known as “vagrants”) interact with resident temperate species, although little is known about the nature and frequency of such interactions. To investigate this phenomenon, we used displaced tropical Indo‐pacific Sergeant Abudefduf vaigiensis settling in temperate south‐eastern Australia as a model system. Underwater visual surveys revealed shoal composition and size differed significantly between open‐water and reef habitats, with shoals in open habitats being larger and more speciose. Shoals containing A. vaigiensis were mainly mixed‐species, and larger and more speciose in open habitats than nearer to reef. Since both foraging efficiency (via access to plankton) and predation threat likely increase with increasing distance from reef habitat, we suggest that mixed‐species shoaling mitigates predation risk whilst allowing increased foraging opportunities for A. vaigiensis in open areas. These findings provide support for the importance of mixed‐species shoaling to the persistence of tropical reef fishes in temperate regions.
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