A vast 4,000-year-old spatial pattern of termite mounds.

The origins of many large-scale 'biogenic' earthen structures are controversial, because often the species that built them have vanished. This is especially true when they form regular (over-dispersed), self-organized vegetation patterns [1]. Here, we describe a vast array of soil mounds constructed by termites (Syntermes dirus) that has persisted for up to 4000 years and covers an estimated 230,000 km2 of seasonally dry tropical forest in a relatively undisturbed and climatically stable region of Northeast Brazil. The mounds are not nests, but rather they are generated by the excavation of vast inter-connecting tunnel networks, resulting in approximately 10 km3 of soil being deposited in 200 million conical mounds that are 2.5 m tall and approximately 9 m in diameter. S. dirus termites are still present in the soil surrounding the mounds and we found that intra-specific aggression occurred at a scale much larger than an individual mound. We suggest that the complex network of tunnels built to access episodic leaf-fall has allowed for the optimization of waste soil removal, which over thousands of years has formed an over-dispersed spatial pattern of mounds.