Two invaders instead of one: the true identity of species under the name Ceracis cucullatus (Coleoptera: Ciidae).

The Neotropical obligate fungivorous beetle Ceracis cucullatus (Mellié) has attracted attention of coleopterists due to the increasing number of records of populations in Africa. Although its disjunct populations have been interpreted as a cohesive taxonomic unity, previous comparisons between African and Neotropical specimens revealed differences in their external morphology, causing uncertainty about the true unity of the species. Here, we compare the external morphology of specimens named Cer. cucullatus from several localities of the Neotropical, Palearctic, Afrotropical, Afrotemperate and Oriental regions. As results, we reverse three previous junior synonymies of Cer. cucullatus, proposing Cer. lamellatus (Pic) and Cer. tabellifer (Mellié), both reinstated status and new combinations, as separate species. We also propose Enn. bilamellatum Pic as a new synonym of Cer. tabellifer. In face of these taxonomic changes, we identify Cer. tabellifer as the actual invasive species on African lands, instead of Cer. cucullatus as was previously accepted. Then, through historical records gathered from scientific collections and literature, and through examination of recently collected specimens from South Africa and Brazil, we provide data on host fungi and geographic distribution of Cer. tabellifer. Based on these data, we discuss possible explanations to the successful invasion of Cer. tabellifer in Africa and elsewhere and its potential threat to native faunas of ciids. This study helps to fulfil an old gap in the literature on biological invasions, with considerably more studies on predatory species, disease vectors or potential pests of agricultural crops, than on non-pest fungivorous organisms.