Fine-scale associational effects: Single plant neighbours can alter susceptibility of focal plants to herbivores.

The neighbourhood of plants in a patch can shape vulnerability of focal plants to herbivores, known as an associational effect. Associational effects of plant neighbourhoods are widely recognised. But whether a single neighbouring plant can exert an associational effect is unknown. Here, we tested if single neighbours indeed do influence the likelihood that a focal plant is visited and eaten by a mammalian herbivore. We then tested whether any refuge effect is strengthened by having more neighbours in direct proximity to a focal plant. We used native plant species and a browser/mixed feeder mammalian herbivore (swamp wallabies (Wallabia bicolor)) free-ranging in natural vegetation. We found that a single neighbouring plant did elicit associational effects. Specifically, plant pairs consisting of one high-quality seedling next to a single low-quality plant were visited and browsed by wallabies later and less than pairs of two high-quality seedlings. Having more neighbours did not strengthen these associational effects. Compared with no neighbours, one or five low-quality neighbours had the same effect in delaying time taken for wallabies to first visit a plot and browse on a high-quality focal seedling. While traditionally a 'patch' refers to a broad sphere-of-influence neighbouring plants have on a focal plant, our findings suggest the influence of plant neighbours can range from the nearest individual neighbour to the entire plant neighbourhood. Such fine-scale associational effects are fundamentally important for understanding intricate plant-herbivore interactions, and ecologically important by potentially having knock-on effects on plant survival, in turn influencing plant community structure.