Abstract
Increasing ocean temperatures have widespread consequences for coral reefs, one of which is coral bleaching. We analyzed a global network of associations between coral species and Symbiodiniaceae for resistance to temperature stress and robustness to perturbations. Null networks were created by changing either the physiological parameters of the nodes or the structures of the networks. We developed a bleaching model in which each link, association, is given a weight based on temperature thresholds for specific host-symbiont pairs and links are removed as temperature increases. Resistance to temperature stress was determined from the response of the networks to the bleaching model. Ecological robustness, defined by how much perturbation is needed to decrease the number of nodes by 50%, was determined for multiple removal models that considered traits of the hosts, symbionts, and their associations. Network resistance to bleaching and robustness to perturbations differed from the null networks and varied across spatial scales, supporting that thermal tolerances, local association patterns, and environment play an important role in network persistence. Networks were more robust to attacks on associations than to attacks on species. Although the global network was fairly robust to random link removals, when links are removed according to the bleaching model, robustness decreases by about 20%. Specific environmental attacks, in the form of increasing temperatures, destabilize the global network of coral species and Symbiodiniaceae. On a global scale, the network was more robust to removals of links with susceptible Symbiodiniaceae than it was to removals of links with susceptible hosts. Thus, the symbionts convey more stability to the symbiosis than the hosts when the system is under an environmental attack. However, our results also provide evidence that the environment of the networks affects robustness to link perturbations. Our work shows that ecological resistance and robustness can be assessed through network analysis that considers specific biological traits and functional weaknesses. The global network of associations between corals and Symbiodiniaceae and its distribution of thermal tolerances are non-random, and the evolution of this architecture has led to higher sensitivity to environmental perturbations.