Facilitation cascades across space: Monitoring estuarine foundation species from satellites to the microscope

Estuaries are productive ecosystems that are vulnerable to human impacts and environmental disturbances. Estuaries are often inhabited by foundation species that facilitate other organisms by creating and modifying habitats. However, spatiotemporal variability and underpinning drivers of facilitation from estuarine foundation species are poorly studied, especially in the context of monitoring and conservation. Here, we combined the analyses of satellite, drone, and camera images, close-up field sampling of quadrats and individuals, and laboratory microscopy, to quantify co-occurrences between the cockle Austrovenus stutchburyi and green macroalga Ulva spp., and their associated invertebrate communities, across spatial resolutions and environmental gradients in a New Zealand estuary. We found higher abundance of Ulva in winter and higher abundance of Austrovenus at a site near the ocean. Furthermore, Ulva was, across space and time, often attached to Austrovenus, either live or dead shells, suggesting a hierarchy between a primary (cockle) and secondary (macroalgae) foundation species. Finally, Ulva provided a better habitat for mobile invertebrates than Austrovenus, and thereby increased estuarine diversity through a density-dependent facilitation cascade, which also was consistent in space and time. Overall, our study showed that facilitation cascades can be indirectly inferred and monitored through remote sensing of foundation species using supplementary imaging methods, and that integration of data across scales allows researchers to better understand the drivers and consequences of facilitation from foundation species. These insights may aid in the development of efficient monitoring strategies and ultimately improve the management of estuarine ecosystems.