Review of Pharaxonotha Reitter (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting the cycad genus Dioon Lindl. (Cycadales), with descriptions of nine new species and comments on P. kirschii Reitter

The beetle genus Pharaxonotha Reitter (Coleoptera: Erotylidae: Pharaxonothinae) is found in the cones of cycad genera in the New World, including species of Dioon Lindl., Ceratozamia Brongn., Microcycas (Miq.) A.DC and Zamia L. In this paper nine new species found in Dioon are described by Skelley, Tang and Pérez-Farrera: Pharaxonotha bicolor, P. dimorpha, P. fawcettae, P. gigantea, P. novoai, P. occidentalis, P. sclerotiza, P. woodruffi, P. vovidesi. A key to described species of Pharaxonotha inhabiting Dioon is presented, along with an account of Pharaxonotha kirschii Reitter for comparison. The New World beetle genus Pharaxonotha Reitter (Erotylidae: Pharaxonothinae) has a close association with cycads (Cycadales), an ancient lineage of gymnosperms (Norstog and Nichols 1997). Except for the type species of Pharaxonotha, P. kirschii Reitter, which is a minor stored products pest, all other species of the genus that have been described are inhabitants of cycad cones (Pakaluk 1988; Chaves and Genaro 2005; Franz and Skelley 2008; Xu et al. 2015; Skelley et al. 2017; Santiago-Jiménez et al. 2019; Skelley and Segalla 2019; Skelley and Tang 2020). In the New World, almost all cycad populations studied have an associated species or two of Pharaxonotha, rang­ing from the southern US and Caribbean to Bolivia and Brazil. Exclusion experiments on the cycad genus Zamia L. indicate that they are pollinators (Tang 1987; Valencia-Montoya et al. 2017; Segalla et al. 2021). As part of a series of papers describing species on New World cycads, here we describe nine species of Pharaxonotha inhabiting cones of the cycad genus Dioon Lindl. Although Vovides (1991) and Navarrete-Heredia (2018) reported on the presence of Pharaxonotha in Dioon cones, this is the first paper to our knowledge, where species inhabiting Dioon have been scientifically described. To aid with their identification, an addendum to the key of Skelley and Tang (2020) for species of Pharaxonotha found on Dioon is pr