Army ant middens - Home and nursery of a diverse beetle fauna.

Army ants provide nourishment to a large variety of animals. This includes birds that feed on animals flushed out by army ant raids, symbiotic arthropods that consume the ants' prey or their brood, and other arthropods that scavenge on army ant refuse deposits. The latter have not received much attention, and the few published studies lack detailed species identifications. Here we provide a first systematic inventory of the beetle fauna associated with refuse deposits of army ants, with a focus on . We collected 8364 adult beetles, 511 larvae, and 24 eggs from 34 deposits at La Selva Biological Station, Costa Rica. We used a combination of DNA barcoding and morphology to identify a subset of 436 specimens to species level. The samples included several new species, and we here formally describe two water scavenger beetles (Hydrophilidae). Refuse deposits harbored a diverse beetle fauna. The identified subset consisted of 91 beetle species from 12 families, with rove beetles being the most abundant and diverse visitors. Of the 85 species found with , 50 species were collected from only one or two refuse deposits. Conversely, seven species were found in 10 or more refuse deposits, indicating a certain level of habitat specialization. We matched adults and immatures for 22 beetle species via DNA barcodes, demonstrating that army ant middens also serve as a beetle nursery. The present survey highlights the significant ecological function of army ants as promoters of biodiversity and their status as keystone species in tropical rainforests.