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 Eciton army ants, with a focus on Eciton burchellii. 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 E. burchellii, 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.

California State Collection of Arthropods California Department of Food and Agriculture Sacramento California USA

Department of Biological Sciences National Sun Yat sen University Kaohsiung Taiwan

Department of Biology Technical University of Darmstadt Darmstadt Germany

Department of Ecology Faculty of Environmental Sciences Czech University of Life Sciences Prague Praha Suchdol Czech Republic

Department of Entomology National Museum Prague Czech Republic

Division of Science Yale NUS College Singapore Singapore

Howard Hughes Medical Institute New York New York USA

Instituto Argentino de Investigaciones de las Zonas Aridas Mendoza Argentina

Laboratory of Social Evolution and Behavior The Rockefeller University New York New York USA

Museum and Institute of Zoology of the Polish Academy of Sciences Warsaw Poland

Museum of Nature Hamburg Leibniz Institute for the Analysis of Biodiversity Change Hamburg Germany

NUS College National University of Singapore Singapore Singapore

The Hokkaido University Museum Hokkaido University Sapporo Japan

The Kyushu University Museum Hakozaki Fukuoka Japan

University of Arizona Insect Collection Tucson Arizona USA

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