Long-term ecological changes in fishes and macro-invertebrates in the world's warmest coral reefs.

The Arabian Gulf is a natural laboratory for examining the consequences of large-scale disturbances due to global warming on coral reef ecosystems because of its extreme temperature regime. Using a coral reef monitoring time series extending from 1985 to 2015, we examined the long-term ecological changes in fish and macro-invertebrate communities as these habitats suffered heat shocks. We used a GLMM modelling framework to obtain clean annual signals in community indicators from noisy data. We also visualized temporal change in the taxonomic composition of fishes and macro-invertebrates. A phase shift from predominantly reef-building corals to barren grounds occurred between 1996 and 2000. Macro-invertebrates responded rapidly, and most of associated indicators recovered to pre-shift levels in 15 years. Fishes generally had lagged responses to the phase shift and had shifted to a new state with lower abundance, as well as different species composition. Increased levels of herbivory first by macro-invertebrates, mostly sea urchins, and then fishes, could have suppressed macro-algae expansion and consequently led to the dominance of barren ground. When the phase shift occurred, most of the 14 fish families declined in abundance while macro-invertebrate groups increased. Fish families able to utilize non-coral habitats appeared more resilient to the disturbances and subsequent coral degradation. Unlike other regions, we observed high resilience of the coral-dependent butterflyfishes to coral loss, possibly due to local migration from other less-impacted coral reefs. We hypothesized a top-down control mechanism mediated by predation by fishes has contributed to shaping the temporal and spatial patterns of the macro-invertebrates. Our results also revealed differences in spatial preferences among fishes and macro-invertebrate groups, which could be used to set priorities and develop effective conservation and management strategies.