HAT

HAT clade

The close relationship of Holisus ( Hyptiomina ), Atanygnathus ( Tanygnathinina ) and Amblyopinina was first recovered by Chani-Posse et al. (2018) using morphological and molecular data. This clade, named here as ‘HAT’ ( Hyptiomina – Amblyopinina – Tanygnathinina ) was corroborated by Schillhammer & Brunke (2018) and again by the present study. The highly derived morphology of Holisus previously led systematists to consider it as a member of Philonthina and therefore, it was not included in the analyses of, for example, Brunke & Solodovnikov (2013) and Brunke et al. (2016).

The HAT clade is supported by many notable morphological characters, some of which have been associated previously with Amblyopinina View in CoL alone: posterior ridge of the mesosternum absent; humeral spines of the elytron absent ( Holisus View in CoL with nonhomologous row positioned far anteriad); wing veins CuA and MP4 fused; base of paramere of aedeagus tightly fused to median lobe, often indistinguishable. Although Holisus View in CoL and Atanygnathus View in CoL are apomorphyrich clades and easily diagnosed, the diverse Amblyopinina View in CoL , at present, are recognized merely by the absence of these apomorphies. Unless diagnostic synapomorphies can be found for Amblyopinina View in CoL , it might be in the best interest of systematics to treat the entire HAT clade as subtribe Tanygnathinina View in CoL , the oldest family-group name. However, a better diversity of Amblyopinina View in CoL , including South African Natalignathus Solodovnikov (Solodovnikov, 2005) View in CoL and the enigmatic Andean Devilleferus ( Jenkins Shaw et al., 2017) View in CoL , should be included in the present dataset before such limits are defined.