5 phylogenetic revision of the genus Mesynodites (Coleoptera: Histeridae : Hetaeriinae) with descriptions of new tribes, genera and species

Mesynodites Reichardt, 1924 is the largest genus within the exclusively inquilinous (social insect-associated) histerid beetle subfamily Hetaeriinae (Insecta: Coleoptera: Histeridae). The 44 described species are systematically revised, and new taxa are described based on newly discovered species and the results of a cladistic analysis. The work was based on available type specimens representing 42 species and approximately 2500 specimens of non-type material. Phylogenetic hypotheses were based on maximum parsimony analysis of 150 morphological characters derived from a selection of Mesynodites species and representatives of related genera. This analysis confirmed that, in its current concept, Mesynodites is not a monophyletic taxon. Species included currently in Mesynodites were scattered among 11 distinct lineages, all but one of them placed within a single informal subdivision of the Hetaeriinae known informally as Group E ofHelava etal. (1985). As a result, in the revised sense, Mesynodites includes only nine species, two of which are of uncertain affinities (type specimens of these species were not located). Revision of Mesynodites species and consideration of representative ingroup and outgroup species in the context of the phylogenetic analysis allowed numerous taxonomic changes to the Hetaeriinae. Two new tribes are described. The monotypic Synoditulini apparently represents the most basal lineage of the hetaeriines. The Nymphysterini (expanded and revised 'Group E') includes most of Mesynodites species and genera related to it, based on the analysis presented. All genera of Hetaeriinae, excluding 12 genera of doubtful affinities, received tribal assignments. The following new genera are described, mostly to accommodate species previously included into Mesynodites: Alienodites, Bruchodites, Guianahister, Helavadites, Microsynodites, Mutosynodites, Nicolasites and Reichenspergerites. Three new species are described: : Alienodites amazonicus (Brazil), Guianahister ashei (French Guiana, Guyana and Suriname) and Mesynodites degallieri (Brazil). The following new combinations are proposed: Aemulister hirsuta (Helava), Alienodites bifurcatus (Mann), Bruchodites praeclusus (Reichensperger), Daitrosister irregularis (Reichensperger), D. longipilus (Reichensperger), D. reticulatus (Bruch), D. setulosus (Reichensperger), Euclasea arcanus (Reichensperger), E. acamati (Reichensperger), E. detritus (Schmidt), E. diadochus (Reichensperger), E. inops (Reichensperger), E. novaeteutoniae (Reichensperger), E. pumilis (Reichensperger), E. sphaeroides (Reichensperger), E. splendens (Reichensperger), E. verruculosus (Reichensperger), Helavadites elegantulus (Reichensperger), H. gibbidorsum (Schmidt), H. manicus (Reichensperger), H. nanus (Reichensperger), H. obscurus (Reichensperger), Microsynodites ciliatus (Bruch), Microsynodites schmidti (Lewis), Microsynodites strigilatus (Reichensperger), Monotonoditespauperella (Lewis), Monotonodites raptantis (Reichensperger), Mutodites evanescens (Reichensperger), Mutodites megacantha (Reichensperger), Mutodites semistriatus (Bruch), Mutodites speculum (Reichensperger), Nicolasites graniformis (Schmidt), Reichenspergerites robustus (Reichensperger), Trichoreninus exclamationis (Reichensperger), T.geminus (Reichensperger), T. major (Bruch), T. schwarzmaieri (Reichensperger). Following new synonymies are established: Gallaster Helava in Helava etal. , 1985 = Aemulister Reichensperger, 1938; Euclasea obliqua Lewis, 1893 = Euclasea detritus (Schmidt, 1893); Mesynodites collaris Reichensperger, 1939 = Mesynodites aciculatus (Schmidt, 1893), Mesynodites eidmanni Reichensperger, 1935 = Mutodites semistriatus (Bruch, 1933), Mesynodites scaber Reichensperger, 1931 = Microsynodites schmidti (Lewis, 1893).