Crustose lichens form a crust that strongly adheres to the substrate (soil, rock, tree bark, etc.), making separation from the substrate impossible without destruction. The basic structure of crustose lichens consists of a cortex layer, an algal layer, and a medulla. The upper cortex layer is differentiated and is usually pigmented. The algal layer lies beneath the cortex. The medulla fastens the lichen to the substrate and is made up of fungal hyphae. The surface of crustose lichens is characterized by branching cracks that periodically close in response to climatic variations such as alternate wetting and drying regimes.