Burrowing mammals are often considered to be ecosystem engineers as burrowing disturbs the soil, thereby potentially changing resource availability and affecting habitat conditions for other species. After their excavation, burrows may strongly impact local plant communities through several mechanisms, including resource trapping, altered chemical and physical soil properties, and amelioration of microclimatic conditions. We studied ecosystem engineering by aardvark (Orycteropus afer) burrowing by comparing soil and vegetation characteristics between three microsites (burrow entrances, excavated soil mounds and adjacent control sites). We were able to identify several engineering effects and distinguish between potential mechanisms. Burrow soils were cooler, drier and less compact than the other microsites, with all three microsites representing unique combinations of abiotic conditions. Mean species richness was higher at older burrows than mounds and non-burrowed controls, despite burrows having a smaller seedbank and not differing in soil fertility from mounds and control sites. However, the opposite was observed at fresh burrows and mounds, where control plots contained more species on average than the other two types of microsites. Burrow age and microsite type also affected species composition, although only a small proportion of species were significantly associated with specific microsites and just two species were limited to a single microsite type. We suggest that trampling and the physical digging action at burrow entrances, and burial by deposited soil at mounds, prevents the establishment of many plant species at active burrows. However, once abandoned, burrow entrances provide good physical conditions for seedling survival, allowing the establishment of more species. Therefore, as suggested previously for other ecosystem engineers, it is important to explicitly consider the age and degradation processes of engineered structures. In addition, our results highlight biologically-important differences in engineering impacts between burrow entrances, where soil is removed, and mounds, where soil is deposited. Such microscale differences are important to consider when examining bioturbation or, more generally, ecosystem engineering.
