ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 99(2): 5–94, 2014 Classification, natural history, and evolution of the Epiphloeinae (Coleoptera: Cleridae) Part XI. Generic taxonomy, intergeneric phylogeny, and catalogue of the subfamily WESTON OPITZ Kansas Wesleyan University, Department of Biology; 100 East Claflin Avenue, Salina, Kansas 67401-6196; USA; e-mail: [email protected] OPITZ W. 2014: Classification, natural history, and evolution of the Epiphloeinae (Coleoptera: Cleridae). Part XI. Generic taxonomy, intergeneric phylogeny, and catalogue of the subfamily. Acta Musei Moraviae, Scientiae biologicae (Brno) 99(2): 5–94. – This treatise is the author’s culmination efforts with the checkered beetle subfamily Epiphloeinae Kuwert. The work contains a summation of the natural history of epiphloeine species, discussion of characters and character states found useful for the discernment of epiphloeine genera and species, description of the subfamily, key to genera, synoptic description of the genera, intergeneric hypothesis of phylogeny, catalogue of the species with each species represented by a color photograph, and index of genera, their species, and their synonyms. Keywords. Coleoptera, Cleridae, Epiphloeinae, genera, phylogeny, catalogue Introduction Epiphloeines are New World arbophilic beetles that frequent tree trunks infested with wood decomposing insects such as bark beetles. Within this group of checkered beetles one finds considerable intrageneric diversity of integumental color and body form varies extensively across generic lines. The species of some genera blend beautifully with the dark scabrous characteristic of tree bark; as in members of Plocamocera Spinola and Silverasia Nemésio. In other epiphloeines the elytra are brightly colored, as one finds among species of the mimetic Epiphloeus Spinola and Acanthocollum Opitz. Commonly, body form varies from suboval, rectangulate, or narrow triangular, with Batesian lycid- like body shapes among members of Ichnea Laporte and cantharoid shapes in Opitzia Nemésio and Hapsidopteris Opitz. As defined herein, Epiphloeinae contains 271 species classified into 25 genera whose combined distribution extends from eastern Canada to northern Argentina. The genera, and their species and type localities, are listed in the taxonomic catalogue that forms the last component of this treatise. Three synapotypic characteristics define the monophyly of Epiphloeinae. One, there are two fully-developed pairs of pronotal trichobothria, one pair is situated paralaterally on the pronotal disc, the second is located on the pronotal sides. Two, the antenna are inserted along the lower frontal margin of the eyes. Three, the metendosternite lacks a furcal lamina. 5 W. OPITZ Cleridae subfamily classification issues “The systematics of the Cleridae should be revised to reflect molecular data” GUNTER et al. 2013: 8”. The authors of this statement may be correct when, and only when, adequate representations of species, of an adequate representation of genera, have been investigated from a molecular basis; and when it is possible to determine the apotypic or plesiotypic evolutionary condition of a set of genes. Commonality in genetic base pairs does not necessarily translate into phylogenetic relationship; no more than similarity in morphological characteristics translates into sister group relationships. Only synapotypic base pairs or synapotypic structural characteristics qualify for modern thoughts about phylogenetic kinships. In the abovementioned molecular work 148 species and 70 genera [out of about 3,629 species and 334 genera (OPITZ 2010: 55)] were considered. While the molecular study in question is an important augment to Cleridae systematic work, I submit that the study contains insufficient representation of Cleridae taxa for sustaining issues of Cleridae higher classification. Moreover, the molecular analysis bestows monophyly on some taxa, but their definition of monophyly is based on a computer program that does not distinguish between the apotypic and plesiotypic state of a gene or sets of genes. The molecular study does not define which base pairs are derived and which are primitive. Having a similar set of genes does not necessarily mean phylogenetic relationships. The human genome shares base pairs with the genome of bacteria, which does not mean that the two taxa in question are closely related. The sharing of similar base pairs could simply mean that two sets of organisms have evolved similar sets of genes to solve a common problem for survival; nuances in unproven techniques of phylogenetic analysis should not trump carefully prepared morphological analysis in accordance with the proven phylogenetic principles of HENNIG (1966). The exclusion of apotypic and plesiotypis parameters in molecular phylogenetics is akin to the idea the commonality of characteristics translates into evolutionary kinships. Such methodology is similar the the dicredited canons of Numerical Taxomy. These considerations becomes particularly relevant when results from a molecular study are used to minimize the significance of taxa-rich morphological analyses. In a recent contribution BARTLETT (2013: 412) writes...” The subfamilies of OPITZ’S (2010) ‘split’ classification (of Korynetinae, sensu latu), though satisfying the need for a more detailed classification, are not all well-defined in terms of synapomorphies and most, with the exception of Epiphloeinae, are not well-supported by molecular evidence.” I presume that this statement relates to Bartlett’s belief that the synapotypies given in OPITZ’S (2010) for subfamilies Enopliinae, Tarsosteninae, Korynetinae, Neorthopleurinae and Peloniinae are manifestations of the use of “continuous characters as defined by SMITH & HENDRICKS (2013: 367). If this assumption is correct I vehemently disagree with Bartlett’s assessment. For example, Enopliinae is clearly defined by a partial commissure (a result of a meeting of the pronotal hem with the dorsolateral carina partially mesad to the hind corner of the pronotum), or in Tarsosteninae where the capitulum is consistently shorter than the combined length of funicular antennomeres. In 6 Acta Musei Moraviae, Sci. biol. (Brno), 99(2), 2014 Epiphloeinae (Coleoptera: Cleridae): Taxonomy, phylogeny, catalogue the taxa that I have examined, there is no in-between of these characteristic among taxa with a reduced 4th tarsomere. The meeting of the pronotal hem with the dorsolateral carina either meets at the corner or it does not. Moreover, elsewhere (OPITZ 2011:142) I have addressed the issue of “homoplastic apotypy” being a viable consideration in establishing phylogenetic relationships (on that basis... “homoplastic apotypies may be a manifestation of selection of (distantly related) different ancestral genomes to solve a common (functional) problem”. Similarity of morphological character states based on functional parameters would of course assure similar nucleic acid base pairs. However, similar structure, and their corresponding similar base pairs, does not necessarily translate into close evolutionary kinships. The molecular study of GUNTER et al. (2013: 634), and the works of KOLIBÁČ (1997: 358; 2010: 260), LESCHEN (2010: 5, 257), BOUCHARD et al. (2011: 348), and LAWRENCE et al. (2010: 5; 2011: 8) place a high value on the phylogenetic significance of the reduction of the 4th tarsomere in the Cleridae. The result of such an interpretation has prompted some of the above authors to place OPITZ’s (2010: 117) Epiphloeinae, Enopliinae, Neorthopleurinae, Peloniinae, Tarsosteninae, and Korynetinae under one subfamily, the Korynetinae. I found the length, and therefore expression, of this tarsomere variable within the family; some of this variation can be seen in illustrations by OPITZ (1998: Fig. 68; 2006: Fig. 39a; 2010: Fig. 92). Moreover, the complete reduction of the metatarsus to four tarsomeres in Anthicoclerinae [synonymized under Clerinae by BOUCHARD et al. (2011: 348) but resurrected herein] although a significant evolutionary change, represents, and gives an indication of, a magnitude of evolutionary plasticity in the expression of the tarsomere structure. Moreover, the molecular surfacing of the Clerinae (sensu latu) – Korynetinae (sensu latu) gap is bound to become apparent in any comprehensive morphologic or molecular study. This has always been considered an obvious break in the family, but the real issue is what manner of discontinuity (characteristic gap) is worthy of subfamily status. If we regard the molecular discontinuities, exhibited in GUNTER et al. (2013), worthy of subfamily status then we might suggest that in the Cleridae we have only two subfamilies, Clerinae and Korynetinae; an idea that would produce a very unbalanced classification in the family. Epiphloeinae taxonomic history The history of the Epiphloeinae taxon begins with Thomas Say who in 1825 described Enoplium dislocatum (SAY 1825: 176). Then, Laporte described Ichnea lycoides (LAPORTE 1836: 55). Laporte was followed by SPINOLA (1841: 75) who classified Ichnea Laporte and Epiphloeus Spinola under the informal name Ichnoïdes. AGASSIZ (1846: 193) Latinized Ichnoïdes to Ichneoidea and credited the name to Spinola. This action by Agassiz made Ichneoidea available for subfamily nomenclatural consideration; however as pointed out by OPITZ & HERMAN (2009: 183), the correct name for the subfamily under consideration is Epiphloeinae Kuwert as governed by the provisions of Articles 23.9.1 and 23.9.2 of the International Commission