Scale-dependent topographic complexity underpins abundance and spatial distribution of ecosystem engineers on natural and artificial structures

In response to ongoing coastal urbanization, it is critical to develop effective methods to improve the biodiversity and ecological sustainability of artificial shorelines. Enhancing the topographic complexity of coastal infrastructure through the mimicry of natural substrata may facilitate the establishment of ecosystem engineering species and associated biogenic habitat formation. However, interactions between ecosystem engineers and their substratum are likely determined by organismal size and resource needs, thus making responses to topography highly scale-dependent. Here, we assessed the topographic properties (rugosity, surface area, micro-surface orientations) that underpin the abundance and distribution of two ecosystem engineers (fucoids, limpets) across six spatial scales (1-500 mm). Furthermore, we assessed the 'biogenic' rugosity created by barnacle matrices across fine scales (1-20 mm). Field surveys and 3D scanning, conducted across natural and artificial substrata, showed major effects of rugosity and associated topographic variables on ecosystem engineer assemblages and spatial occupancy, while additional abiotic environmental factors (compass direction, wave exposure) and biotic associations only had weak influences. Natural substrata exhibited <= 67 % higher rugosity than artificial ones. Fucoid-covered patches were predominantly associated with high-rugosity substrata and horizontal micro-surfaces, while homescars of limpets (>= 15 mm shell length) predominated on smoother