Relationships between bioturbation by Tylos spinulosus (Crustacea, Isopoda) and its distribution on sandy beaches of north-central Chile

The semi-terrestrial isopod, Tylos spinulosus Dana, is a common inhabitant of the upper shore levels of sandy beaches of north-central Chile (ca. 26-30°S). During daylight hours, this isopod remains buried in the sand, while during the night emerges for feeding on stranded organic detritus, leaving exit holes on the beach surface. After feeding, isopods return to dig in their burrowing zones leaving surface irregularities such as cone-shaped mounds of sand. The burrowing preference of T. spinulosus was studied in the field, by: (i) releasing 30 isopods on artificially prepared sand circles (2 m diameter) having exit holes and mounds similar to those left by the isopods and on circles without holes and mounds, and (ii) counting active and buried isopods 15 min after their release in the experimental arenas. The circles had two densities of holes and mounds: treatments 1 and 2 had 100 and 200 holes, respectively, while treatments 3 and 4 had 100 and 200 mounds, respectively. Other 30 isopods were released on sand circles without these holes and mounds (treatment 5). A significantly higher number of isopods buried in circles with holes and mounds (either inside or outside them), compared with experimental arenas without such structures. These results show that the beach surface heterogeneity resulting from holes and mounds would be one of the processes explaining the patchiness of T. spinulosus and thus, its zonation on the intertidal zones of sandy beaches of north-central Chile.