World reclassification of the Cardiophorinae (Coleoptera, Elateridae), based on phylogenetic analyses of morphological characters

Abstract

The prior genus-level classification of Cardiophorinae had never been assessed phylogenetically, and not revised since 1906. A phylogeny for Cardiophorinae and Negastriinae is inferred by Bayesian analyses of 163 adult morphological characters to revise the generic classification. Parsimony analysis is also performed to assess the sensitivity of the Bayesian results to the choice of optimality criterion. Bayesian hypothesis testing rejected monophyly for: Negastriinae; Cardiophorinae (but monophyletic after addition of four taxa); Cardiophorini; cardiophorine genera <i>Aphricus</i> LeConte, 1853; <i>Aptopus</i> Eschscholtz, 1829; <i>Cardiophorus</i> Eschscholtz, 1829; <i>Cardiotarsus</i> Eschscholtz, 1836; <i>Paracardiophorus</i> Schwarz, 1895; <i>Phorocardius</i> Fleutiaux, 1931; <i>Dicronychus</i><i>sensu</i> Platia, 1994; <i>Dicronychus</i><i>sensu</i> Méquignon, 1931; <i>Craspedostethus</i><i>sensu</i> Schwarz, 1906 (i.e., including <i>Tropidiplus</i> Fleutiaux, 1903); <i>Paracardiophorus</i><i>sensu</i> Cobos, 1970, although well-supported alternative classifications were available for only some. Based on taxonomic interpretation of phylogenetic results: Nyctorini is <b>syn. n.</b> of Cardiophorini; <i>Globothorax</i> Fleutiaux, 1891 (Physodactylinae), <i>Margogastrius</i> Schwarz, 1903 (Physodactylinae), and <i>Pachyelater</i> Lesne, 1897 (Dendrometrinae) are transferred to Cardiophorinae. The following changes are proposed for cardiophorine genera: <i>Aptopus</i> Eschscholtz, 1829 is redefined to exclude <i>Horistonotus</i>-like species; <i>Coptostethus</i> Wollaston, 1854 is subgenus of <i>Cardiophorus</i>; <i>Dicronychus</i> Brullé, 1832 and <i>Diocarphus</i> Fleutiaux, 1947, <i>Metacardiophorus</i> Gurjeva, 1966, <i>Platynychus</i> Motschulsky, 1858, and <i>Zygocardiophorus</i> Iablokoff-Khnzorian and Mardjanian, 1981 are placed at genus rank; <i>Paracardiophorus</i> Schwarz, 1895 is redefined based on North American and Eurasian species only; <i>Horistonotus</i> Candèze, 1860 redefined to include species with multiple apices on each side of their tarsal claws; <i>Patriciella</i> Van Zwaluwenburg, 1953 is <b>syn. n.</b> of <i>Aphricus</i> LeConte, 1853; <i>Teslasena</i> Fleutiaux, 1892 (Physodactylinae) is <b>syn. n.</b> of <i>Globothorax</i> Fleutiaux, 1891. The following new genera are described: <i>Austrocardiophorus</i> (type species: <i>Cardiophorus humeralis</i> Fairmaire and Germain, 1860); <i>Chileaphricus</i> (type species: <i>Aphricus chilensis</i> Fleutiaux, 1940); <i>Floridelater</i> (type species: <i>Coptostethus americanus</i> Horn, 1871, transferred from Negastriinae to Cardiophorinae). <i>Paradicronychus</i> (<i>nomen nudum</i>), is <b>syn. n.</b> of <i>Cardiophorus</i> Eschscholtz, 1829. Generic reassignments to make <i>Cardiodontulus</i>, <i>Cardiophorus</i>, <i>Cardiotarsus</i>, <i>Paracardiophorus</i> consistent with phylogenetically revised genus concepts resulted in 84 new combinations. Lectotypes are designated for 29 type species to fix generic concepts: <i>Anelastes femoralis</i> Lucas, 1857; <i>Aphricus chilensis</i> Fleutiaux, 1940; <i>Athous argentatus</i> Abeille de Perrin, 1894; <i>Cardiophorus adjutor</i> Candèze, 1875; <i>Cardiophorus florentini</i> Fleutiaux, 1895; <i>Cardiophorus inflatus</i> Candèze, 1882; <i>Cardiophorus luridipes</i> Candèze, 1860; <i>Cardiophorus mirabilis</i> Candèze, 1860; <i>Cardiophorus musculus</i> Erichson, 1840; <i>Cardiotarsus capensis</i> Candèze, 1860; <i>Cardiotarsus vitalisi</i> Fleutiaux, 1918; <i>Craspedostethus rufiventris</i> Schwarz, 1898; <i>Elater cinereus</i> Herbst, 1784; <i>Elater minutissimus</i> Germar, 1817; <i>Elater sputator</i> Linnaeus, 1758; <i>Elater thoracicus</i> Fabricius, 1801; <i>Eniconyx pullatus</i> Horn, 1884; <i>Esthesopus castaneus</i> Eschscholtz, 1829; <i>Gastrimargus schneideri</i> Schwarz, 1902; <i>Globothorax chevrolati</i> Fleutiaux, 1891; <i>Horistonotus flavidus</i> Candèze, 1860; <i>Horistonotus simplex</i> LeConte, 1863; <i>Lesnelater madagascariensis</i> Fleutiaux, 1935; <i>Oedostethus femoralis</i> LeConte, 1853; <i>Phorocardius solitarius</i> Fleutiaux, 1931; <i>Platynychus indicus</i> Motschulsky, 1858; <i>Platynychus mixtus</i> Fleutiaux, 1931; <i>Triplonychus acuminatus</i> Candèze, 1860; <i>Tropidiplus tellinii</i> Fleutiaux, 1903. A key to genera and diagnoses are provided for all genera and subgenera. A bibliographic synonymy includes references for all taxonomic changes to genera and new species through 2015.