Article: Coptoclavid beetles (Coleoptera: Adephaga) from the Lower Cretaceous of Spain: a new feeding strategy in beetles

Currently the beetle family Coptoclavidae consists of four subfamilies known from the Upper Triassic-Lower Cretaceous (Aptian). We describe two new subfamilies, three new genera and five new species from the Las Hoyas (Cuenca Province) and El Montsec (Lleida Province) localities of Barremian (Early Cretaceous) age from Spain: the first new subfamily, Hispanoclavinae, is from Las Hoyas and comprises one new genus, Hispanoclavina, and two new species, H. diazromerali and H. gratshevi; the second, Coptoclaviscinae, is represented at El Montsec by one new species of Coptoclavella (C. inexpecta). In addition, at Las Hoyas the subfamilies Necronectinae and Coptoclavinae are each represented by one new genus and species: Ovonectes pilosum and Hoyaclava buscalionae, respectively. All beetles previously assigned to the family Coptoclavidae have been interpreted as active hunters. However, we consider Hispanoclavina and Hoyaclava to be filter-feeding, with forelegs adapted for filtering plankton (probably zooplankton because other members of the family are carnivorous) on and beneath the water surface. This represents a new feeding strategy in beetles (extant and extinct). The five new species extend the known geographical distribution of the Coptoclavidae into the western part of European Barremian deposits. The Las Hoyas locality now has the highest known diversity of coptoclavid species.