Resumen
Heterogeneity can boost biodiversity, as well as increase the resilience of an ecosystem to changing environmental conditions; therefore, it is important to understand how topographic heterogeneity in ecosystems is formed. Sandy tidal marshes have a repetitive pattern of higher elevated hummocks surrounded by lower elevated depressions, representing topographic heterogeneity at the scale of a few square meters. The aims of this study were to determine when this topographic heterogeneity forms, how it is structured, and whether it persists during marsh development. The soil topography of marshes consists of coarse-grained sediment formed before marsh vegetation development, with an overlaying fine-grained sediment layer formed after initial marsh development. To gain insight into the formation of topographic heterogeneity, we studied the underlying soil topography of four European sandy marshes, where topographic heterogeneity at a scale of a few square meters was present. The differences in elevation between hummocks and depressions can either be caused by heterogeneity in the coarse-grained sediment or by heterogeneity in the top layer containing the fine-grained sediment. Our results showed that the largest percentage of elevational differences between hummocks and depressions could be attributed to heterogeneity in the underlying coarse-grained substratum. Therefore, we conclude that the patterns in all four marshes were primarily formed before marsh development, before fine-grained sediment was deposited on top of the coarse-grained sediment. However, a smaller percentage of the elevational difference between hummocks and depressions can also be explained by the presence of thicker fine-grained sediment layers on top of hummocks compared with depressions. This implies that marsh accretion rates were higher on hummocks compared with depressions. However, this result was limited to very early stages of marsh development, as marsh accretion rates estimated on marshes ranging between 15- and 120-years-old showed that depressions actually accreted sediments at a significantly faster rate than hummocks. Eventually, the patterns of heterogeneity stabilized and we found similar marsh accretion rates on hummocks and in depressions in the 120-year-old marsh, which resulted in the persistency of these topographic patterns.