Notholopha (Ecuadion), Moret & Toledano, 2002

Maddison, David R., 2014, An unexpected clade of South American ground beetles (Coleoptera, Carabidae, Bembidion), ZooKeys 416, pp. 113-155 : 131

Taxon classification Animalia Coleoptera Carabidae

Subgenus Ecuadion Moret & Toledano, 2002

Ecuadion Moret & Toledano, 2002; type species Bembidion fulvocinctum Bates, by original designation.

Remarks.

Ecuadion is the largest subgenus in the Antiperyphanes Complex, with over 50 described species ( Maddison and Toledano 2012; Moret and Toledano 2002; Toledano 2008; Vigna Taglianti and Toledano 2008) and likely many undescribed. It occurs from Costa Rica south to the mountains near Mendoza, Argentina. There are no known exoskeletal synapomorphies of the group, but it is well-supported by the molecular data, with bootstrap support in six of the seven genes examined (Table 3).

Ecuadion falls into two distinct clades among the sampled species: (1) the Bembidion chimborazonum group (clade m in Fig. 8), consisting mostly of larger, long-legged species (Figs 3A, B); (2) remaining Ecuadion (clade n in Fig. 8), consisting of mostly smaller species with shorter appendages (Figs 3 C–F). The Bembidion chimborazonum group is supported by all genes examined (Table 3); support for the complementary clade is not quite as strong, with the clearest evidence coming from ribosomal genes (Table 3).

Unlike most Bembidion , this subgenus has radiated in habitats away from water. Some occur in leaf litter in cloud forests (e.g., Bembidion andersoni , Bembidion georgeballi , Bembidion onorei , Bembidion sp. “Papallacta”; Fig. 4D), in habitats that would typically be occupied by the genus Trechus Clairville in North America. A number of species are found in open, high-elevation grasslands (e.g., Bembidion chimborazonum , Bembidion guamani , Bembidion humboldti ; Fig. 4C). Some species occur on the upper banks of creek shores (e.g., Bembidion sanctaemarthae , Bembidion ricei ); others are found on clay or silt cliffs (e.g., Bembidion agonoides , Bembidion walterrossii ).