Cholevinae are well founded as a monophylum, but their internal phylogenetic relationships constitute a matter of longstanding dispute. The morphology has been the main source of information in most available phylogenetic studies on the subfamily. Most of the characters used, however, were limited to easily visible external structures and genitalia. Here we investigate the informative power of an unexplored character system for the higher-level systematics of Cholevinae: the morphology of the pretarsus and distal margin of the terminal tarsomere. We analysed and documented these structures in representatives of the five most species-rich tribes of Cholevinae (encompassing 13 subtribes) using scanning electron microscopy. We identified several diagnostic features and recognize potential synapomorphies at the tribal, subtribal and generic levels. The architecture of the median and empodial sclerites (including the empodial setae), the shape and composition of the medial projection of the distal margin of the terminal tarsomere, and the armature of the claws were considered a promising source of information for delimiting tribes and subtribes. Our data challenge the traditional view of a close proximity of Eucatopini and Ptomaphagini as well as a previously suggested synapomorphy of Leptodirini, and reinforce the monophyly of Ptomaphagini. This contribution is one of the rare comparative studies on the coleopteran pretarsus and one of very few evaluating the systematic usefulness of the distal margin of the terminal tarsomere in insects. 392

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The current state of knowledge of the suprageneric relationships in Cholevinae is either derived from informal evaluations of putative synapomorphies or based on molecular studies with limited taxonomic sampling. Here we assessed the higher-level relationships in this subfamily based on a phylogenetic analysis of 97 morphological characters scored for 93 terminals, representing all tribes. Both parsimony and Bayesian analyses were used. The monophyletic origin of Cholevinae was corroborated, except for the unexpected inclusion of Leptinus in the implied weighting analysis. Eucatopini + Oritocatopini were retrieved as basal branches in the evolution of Cholevinae. The monophyletic origin of all remaining Cholevinae was confirmed, which is consistent with molecular evidence. Anemadini was non-monophyletic, in accordance with earlier hypotheses. Cholevini was rendered non-monophyletic by the uncertain inclusion of Prionochaeta and the consistent exclusion of Cholevinus. A close affinity of Ptomaphagini to Sciaphyini and Leptodirini was suggested, although the position of Sciaphyes remains uncertain. The phylogenetic hypothesis of Cholevinae provided here is the most comprehensive presently available. The list of characters shows that a substantial part of the data was obtained from the ventral side. This is a strong argument for a detailed pictorial documentation of the ventral body parts in taxonomic descriptions, in contrast to the common practice of only illustrating the dorsal habitus of the beetles.

Detailed studies of microstructure has recently been shown to provide phylogenetic signals at several supraspecific levels within leiodid coleopterans, as well as in other insects. The tribe Ptomaphagini (Leiodidae: Cholevinae), with a Holarctic-Neotropical-Oriental distribution, has been characterized, among other things, by having a comb of equal-sized, flat spines around the apex of the tibiae of all legs, with a row of spines extending along the outer edge of the protibiae in the subtribes Baryodirina and Ptomaphaginina (but not in Ptomaphagina). A pattern similar to the one in Ptomaphaginina also occurs in the Neotropical cholevine tribe Eucatopini, and this has been used to indicate a phylogenetic relationship between the two tribes (but recent phylogenetic studies have not supported such a close relationship). We here review and revise the presence and structure of periapical (here called an ‘apical crown’) and marginal (here called an ‘external comb’) combs of spines on tibiae in Ptomaphagini, using other cholevines (with and without apical tibial combs) for comparison. We find a phylogenetic signal in an apical crown of tibial spines not interrupted at the outer spur, which seems to be an additional synapomorphy of Ptomaphagini, differing from the pattern in Eucatopini and remaining cholevines with an apical comb of spines, in which the comb is interrupted. We highlight differences not previously noticed between the apical protibial armature of Ptomaphaginina and Eucatopini.