Revision of the genus Attaphila (Blattodea: Blaberoidea), myrmecophiles living in the mushroom gardens of leaf-cutting ants

The genus Attaphila, comprising minute myrmecophilous cockroaches, is revised, including now six previously known (A. aptera, A. bergi, A. flava, A. fungicola, A. schuppi, A. sexdentis) and three new species (A. multisetosasp. nov. Bohn and Klass, A. paucisetosasp. nov. Bohn and Klass, A. sinuosocarinatasp. nov. Bohn and Klass). All species are described or redescribed and depicted with their main characteristics; determination keys allow the identification of males and females. Especially the male characters allow a distribution to two species-groups with differing host specificity: bergi-group associated with Acromyrmex (and possibly Amoimyrmex) ants, fungicola-group associated with Atta ants; the former appears paraphyletic, the latter monophyletic. The genus Attaphila is characterised emphasising its unique features: (1) insertion of antennae at the bottom of a wide funnel-shaped deepening; (2) antenna with the possibility of a rectangular bending between scapus and pedicellus (associated with a distal excavation of the scapus) and (3) with an unusual shape and low number of antennomeres; (4) femora of legs with a ventral groove allowing a close spacing of femur and tibia during a strong flexion; (5) a complex and unusual shape of the laterosternal shelf area of the female genitalia (lack of shelf, presence of a pair of complicated tubular invaginations); and (6) lateral parts of abdominal tergite T9 of male ending in a pair of ventromesally directed arms, which contact the lateral margins of the subgenital plate. Functional aspects and the possible biological roles of these features are discussed. Older biological data are summarised and new observations are presented. The position of Attaphila within Blattodea is discussed. Like a recent molecular study, the morphology of the male genitalia places the genus in the Blaberoidea. The molecular result of Attaphila being closest to three particular blattellid genera, however, is conflictual from the morphological perspective.