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 Aphricus LeConte, 1853; Aptopus Eschscholtz, 1829; Cardiophorus Eschscholtz, 1829; Cardiotarsus Eschscholtz, 1836; Paracardiophorus Schwarz, 1895; Phorocardius Fleutiaux, 1931; Dicronychus sensu Platia, 1994; Dicronychus sensu Méquignon, 1931; Craspedostethus sensu Schwarz, 1906 (i.e., including Tropidiplus Fleutiaux, 1903); Paracardiophorus sensu Cobos, 1970, although well-supported alternative classifications were available for only some. Based on taxonomic interpretation of phylogenetic results: Nyctorini is syn. n. of Cardiophorini; Globothorax Fleutiaux, 1891 (Physodactylinae), Margogastrius Schwarz, 1903 (Physodactylinae), and Pachyelater Lesne, 1897 (Dendrometrinae) are transferred to Cardiophorinae. The following changes are proposed for cardiophorine genera: Aptopus Eschscholtz, 1829 is redefined to exclude Horistonotus-like species; Coptostethus Wollaston, 1854 is subgenus of Cardiophorus; Dicronychus Brullé, 1832 and Diocarphus Fleutiaux, 1947, Metacardiophorus Gurjeva, 1966, Platynychus Motschulsky, 1858, and Zygocardiophorus Iablokoff-Khnzorian and Mardjanian, 1981 are placed at genus rank; Paracardiophorus Schwarz, 1895 is redefined based on North American and Eurasian species only; Horistonotus Candèze, 1860 redefined to include species with multiple apices on each side of their tarsal claws; Patriciella Van Zwaluwenburg, 1953 is syn. n. of Aphricus LeConte, 1853; Teslasena Fleutiaux, 1892 (Physodactylinae) is syn. n. of Globothorax Fleutiaux, 1891. The following new genera are described: Austrocardiophorus (type species: Cardiophorus humeralis Fairmaire and Germain, 1860); Chileaphricus (type species: Aphricus chilensis Fleutiaux, 1940); Floridelater (type species: Coptostethus americanus Horn, 1871, transferred from Negastriinae to Cardiophorinae). Paradicronychus (nomen nudum), is syn. n. of Cardiophorus Eschscholtz, 1829. Generic reassignments to make Cardiodontulus, Cardiophorus, Cardiotarsus, Paracardiophorus consistent with phylogenetically revised genus concepts resulted in 84 new combinations. Lectotypes are designated for 29 type species to fix generic concepts: Anelastes femoralis Lucas, 1857; Aphricus chilensis Fleutiaux, 1940; Athous argentatus Abeille de Perrin, 1894; Cardiophorus adjutor Candèze, 1875; Cardiophorus florentini Fleutiaux, 1895; Cardiophorus inflatus Candèze, 1882; Cardiophorus luridipes Candèze, 1860; Cardiophorus mirabilis Candèze, 1860; Cardiophorus musculus Erichson, 1840; Cardiotarsus capensis Candèze, 1860; Cardiotarsus vitalisi Fleutiaux, 1918; Craspedostethus rufiventris Schwarz, 1898; Elater cinereus Herbst, 1784; Elater minutissimus Germar, 1817; Elater sputator Linnaeus, 1758; Elater thoracicus Fabricius, 1801; Eniconyx pullatus Horn, 1884; Esthesopus castaneus Eschscholtz, 1829; Gastrimargus schneideri Schwarz, 1902; Globothorax chevrolati Fleutiaux, 1891; Horistonotus flavidus Candèze, 1860; Horistonotus simplex LeConte, 1863; Lesnelater madagascariensis Fleutiaux, 1935; Oedostethus femoralis LeConte, 1853; Phorocardius solitarius Fleutiaux, 1931; Platynychus indicus Motschulsky, 1858; Platynychus mixtus Fleutiaux, 1931; Triplonychus acuminatus Candèze, 1860; Tropidiplus tellinii 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.