On the systematics of Noteridae (Coleoptera: Adephaga: Hydradephaga): Phylogeny, description of a new tribe, genus and species, and survey of female genital morphology

A phylogenetic analysis of Noteridae Thomson, 1860 is presented based on 33 adult morphological characters with focus on placement of a new genus and species from Venezuela, Tonerus wheeleri, gen. nov, sp. nov. Characters are derived from many used previously in Noteridae systematics with many new characters, especially from a survey of the female genitalia. Exemplars from ail noterid genera were included except Phreatodytes Ueno, 1957 and Synchortus Sharp, 1882 with outgroups from Paelobiidae, Amphizoidae and Dytiscidae. Relationships among the genera were found to be (Notomicrus Sharp + Speonoterus Spangler) + (Tonerus + (Neohydrocoptus Sato + (Pronoterus Sharp + (Mesonoterus Sharp + (Noterus Clairville + (Renotus Guignot + (Siolius Balfour-Browne + (Suphis Aube + (Canthydrus [Lioconthydrus] Guignot + Suphisellus Crotch)) + (Canthydrus Sharp sensu stricto + Hydrocanthus Say))))))))). Tonerus occupies a phylogenetic position that requires a new tribe, Tonerini tr. nov. characterised by the following character states: (1) the metacoxa and metafurca are fused laterally forming a complete ring, (2) the noterid platform extends anteriorly onto the metaventrite, (3) the female [aterotergites are short and anteriorly not broadly expanded and posteriorly extending well beyond the gonocoxae bases, (4) the gonocoxae are apically rounded, (5) the protarsus is attached at the apex of the protibia, (6) there is a distinct dorsoapical protibial angle, and (7) there are several stout, straight spines at the apex of the protarsus. Three subfamilies of Noteridae are recognised: (1) Phreatodytinae Ueno, including only Phreatodytes; (2) Notomicrinae Zimmermann, including Notomicrus and Speonoterus, and (3) Noterinae, including all other genera. Phreatodytinae and Notomicrinae each include a single tribe. Noterinae includes the tribes Tonerini (Tonerus), Neohydrocoptini Zalat, Saleh, Angus and Kaschef (Neohydrocoptus), Pronoterini Nilsson (Pronoterus) and Noterini (all other genera of Noterinae). Canthydrus is polyphyletic with the subgenus Liocanthydrus sister to Suphisellus and Canthydrus s. str. sister to Hydrocanthus, so Liocanthydrus is elevated from subgenus to genus rank (new status). Characters historically used to define Hydrocanthus (Sternocanthus) Guignot were determined to be unreliable, and Sternocanthus is placed in synonymy with Hydrocanthus (new synonym). The resulting phylogeny is compared with published hypotheses, and the evolution of characters is discussed.