The importance of heuweltjie patch isolation, size, and quality for arthropods in the Succulent Karoo, South Africa

A characteristic feature of Earth's drylands is the patchy nature of the vegetation, often referred to as a two-phase mosaic landscape, comprised of a homogenous matrix containing distinctive vegetated patches. The latter are considered vital for ecosystem functioning as they provide refuge to biota from unsuitable conditions. Ground-living (epigeic) and foliage arthropods contribute to dryland biodiversity patterns and processes, but little is known of how their richness, abundance, and composition varies between patches and the matrix in these systems. Throughout the Succulent Karoo, South Africa, such patches (earthen mounds referred to as heuweltjies) are hotspots for both floral and faunal diversity. We investigate how epigeic and foliage-dwelling arthropod species richness, abundance, and community composition respond to heuweltjie patch characteristics, particularly isolation (distance to nearest neighbor, average distance to nearest neighbor, patch density), size (area), and quality (average plant height, dead plant cover, leaf litter cover, termite frass) during a severe drought. Patch isolation and quality were significantly correlated with arthropod richness, abundance, and community composition. More specifically, average proximity of sampled heuweltjies to other neighboring heuweltjies, termite frass, and vegetation structure (height, leaf litter and dead plant cover) were key determinants for epigeic and foliage-dwelling arthropods' species richness, abundance, and community composition. The uniqueness of these mounds as landscape features increases niche and microhabitat availability for arthropods. This emphasizes their importance as promoters of landscape heterogeneity and highlight heuweltjies as key to the spatial distribution of arthropod assemblages in the Succulent Karoo.