Paracardiophorus musculus, copyright Hungarian Natural History Museum.

Belongs within: Elateridae.

The Cardiophorinae are a group of click beetles united by the presence of two sublateral carinae and incisions on the base of the pronotum, a truncate prosternal spine, a cordate scutellum, and larvae with exodont mandibles. Members of the Old World genus Paracardiophorus have an incomplete lateral margin on the pronotum and simple tarsal segments (Calder 1996).

Characters (from Calder 1996): Head more or less convex anteriorly. Frontoclypeal region raised above plane of labrum forming wide transverse region between basal articulations of antennae. Frontal carina complete across front between eyes. Pronotum usually wider than long, occasionally as wide as long. Base of pronotum with two sublateral carinae and incisions. Prosternal spine (posterior to procoxae) about same length as coxal diameter, truncate apically. Scutellum cordate with strongly pointed apex. Meso-trochantin not visible. Mesocoxal cavity closed to mesepisternum and mesepimeron. Hind wing with short radial cell; vein MP4 without apparent cross vein to CuA2; wedge cell absent; apex of wing membrane (without membrane) fairly long occupying at least one-quarter of length of wing membrane; apical sclerotisations various. Tarsomeres usually simple, tarsomere 4 strongly expanded laterally and cordiform in Cardiotarsus. Tarsal claws without basal setae on outer flat portion. Female genitalia with large glandular reservoir entering bursa copulatrix on mid-ventral surface.

<==Cardiophorinae [Cardiophorini, Esthesopinae]
    |--Aphricus LeConte 1853J02
    |--Aptopus Eschscholtz 1829 [incl. Eniconyx Horn 1884]J02
    |--Patriciella Van Zwaluwenburg 1953C96
    |    `--*P. australica (Van Zwaluwenburg 1953) [=*Patricia australica]C96
    |--Esthesopus Eschscholtz 1829J02
    |    `--E. poedicus Cand. 1860 [incl. E. unifasciatus]FS90
    |--Horistonotus Candèze 1860J02
    |    `--H. asthenicusFS90
    |--Dicronychus [Dicronychidae]KB11
    |    |--D. cinereusKB11
    |    `--D. rubripesKB11
    |--Cardiotarsus Eschscholtz 1836C96
    |    |--*C. capensisC96
    |    |--C. housaianus Kishii 1994PS07
    |    |--C. mjobergi Carter 1939C96
    |    `--C. taiwanus Kishii 1994PS07
    |--Cardiophorus Eschscholtz 1829J02
    |    |--C. atramentariusB90
    |    |--C. cinereusC01
    |    |--C. conductusF89
    |    |--C. dilutusG89
    |    |--C. ebeninus [incl. C. curtulus]B90
    |    |--C. equisetiC01
    |    |--C. erichsoniKB11
    |    |--C. fairmairei Cand. 1888F89
    |    |--C. funebrisG89
    |    |--C. hedenborgiiG89
    |    |--C. javanusF89
    |    |--C. joculariusG89
    |    |--C. musculusB90
    |    |--C. notatus (Fabricius 1781) [=Elater notatus]B14
    |    |--C. pallidulus Cand. 1888F89
    |    |--C. pallipesF89
    |    |--C. rubripesC01
    |    |--C. rufipesC01
    |    |--C. servilisF89
    |    |--C. sexmaculatus Lucas 1846E12
    |    `--C. soricinusF89
    `--Paracardiophorus Schwarz 1895C96
         |--*P. musculus [=Cardiotarsus musculus]C96
         |--P. alternatus Carter 1939C96
         |--P. amabilis Carter 1939C96
         |--P. antennalis Schwarz 1907C96
         |--P. assimilis Carter 1939C96
         |--P. atronotatus Carter 1939C96
         |--P. attenuatipennis Elston 1930C96
         |--P. australis (Candèze 1860) [=Horistonotus australis; incl. H. bicolor Rainbow 1904]C96
         |--P. bicolor (Candèze 1878) [=Cardiophorus bicolor]C96
         |--P. carissimus Carter 1939C96
         |--P. compactus (Candèze 1882) [=Cardiophorus compactus]C96
         |--P. consobrinus (Candèze 1878) [=Horistonotus consobrinus]C96
         |--P. consputus (Candèze 1878) [=Cardiophorus consputus]C96
         |--P. cooki Carter 1939C96
         |--P. despectus (Candèze 1882) [=Cardiophorus despectus]C96
         |--P. dimidiatus Schwarz 1902C96
         |--P. dissimilis Schwarz 1903C96
         |--P. divisus (Candèze 1865) [=Cardiophorus divisus]C96
         |--P. dulcis Carter 1939C96
         |--P. elevatus (Van Zwaluwenburg 1947) [=Cardiophorus elevatus]C96
         |--P. elisus (Candèze 1865) [=Horistonotus elisus]C96
         |--P. eucalypti (Blackburn 1892) [=Cardiophorus eucalypti]C96
         |--P. flavipennis (Candèze 1878) [=Cardiophorus flavipennis]C96
         |--P. flavopictus (Carter 1939) [=Hypnoidus flavopictus]C96
         |--P. fulvosignatus (Candèze 1878) [=Cardiophorus fulvosignatus]C96
         |--P. hamatus (Candèze 1878) [=Cardiophorus hamatus]C96
         |--P. humilis (Candèze 1865) [=Cardiophorus humilis]C96
         |--P. jugulus Elston 1930C96
         |--P. lenis (Candèze 1865) [=Horistonotus lenis]C96
         |--P. litoralis Carter 1939C96
         |--P. longicornis (Candèze 1878) [=Horistonotus longicornis]C96
         |--P. macleayi (Schwarz 1907) [=Cardiophorus macleayi, C. quadrimaculatus Macleay 1888 non Motschulsky 1861]C96
         |--P. malkini (Van Zwaluwenburg 1947) [=Cardiophorus malkini]C96
         |--P. mastersii (Macleay 1872) [=Elater mastersii]C96
         |--P. minimus (Candèze 1878) [=Cardiophorus minimus]C96
         |--P. mjobergi Elston 1930C96
         |--P. moseri Schwarz 1902C96
         |--P. nigrosuffusus Carter 1939C96
         |--P. occidentalis Carter 1939C96
         |--P. octavus (Candèze 1878) [=Cardiophorus octavus]C96
         |--P. octosignatus Carter 1939C96
         |--P. pallidipennis (Candèze 1878) [=Cardiophorus pallidipennis]C96
         |--P. quadripunctatus (Blanchard 1853) [=Agriotes quadipunctatus]C96
         |--P. quadristellatus Carter 1939C96
         |--P. rufopictus Carter 1939C96
         |--P. sexnotatus Carter 1939C96
         |--P. stellatus Carter 1939C96
         |--P. subcruciatus Carter 1939C96
         |--P. subfasciatus Carter 1939C96
         |--P. tumidithorax (Schwarz 1907) [=Cardiophorus tumidithorax, C. tumidicollis Schwarz 1907]C96
         |--P. vagus Schwarz 1907C96
         |--P. varians Carter 1939C96
         |--P. variegatus Schwarz 1902C96
         |--P. venustus (Candèze 1860) [=Cardiophorus venustus]C96
         |--P. victoriensis (Blackburn 1892) [=Cardiophorus victoriensis]C96
         |--P. vittipennis Carter 1939C96
         `--P. xanthomus (Candèze 1865) [=Horistonotus xanthomus]C96

*Type species of generic name indicated


[B14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature’s gems. Ivy Press: Lewes (United Kingdom).

[B90] Buysson, H. du. 1890. Cardiophorus curtulus. Annales de la Société Entomologique de France, 6e série 9: cxlvii–cxlviii.

[C96] Calder, A. A. 1996. Click beetles: genera of the Australian Elateridae (Coleoptera). CSIRO Australia: Collingwood.

[C01] Csiki, E. 1901. Bogarak [Coleopteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 75–120. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l’Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[F89] Fleutiaux, E. 1889. Contributions a la faune Indo-Chinoise. Cicindelidae et Elateridae. 1er mémoire. Annales de la Société Entomologique de France, 6e série 9: 137–146.

[FS90] Fleutiaux, E., & A. Sallé. 1890. Liste des coléoptères de la Guadeloupe et descriptions d’espèces nouvelles. Annales de la Société Entomologique de France, 6e série 9: 351–484.

[G89] Gestro, R. 1889. Viaggio ab Assab nel Mar Rosso dei signori G. Doria ed O. Beccari con il R. Avviso “Esploratore” dal 16 Novembre 1879 al 26 Febbraio 1880.—IV. Coleotteri. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a, 7: 5–72.

[J02] Johnson, P. J. 2002. Elateridae Leach 1815. In: Arnett, R. H., Jr, M. C. Thomas, P. E. Skelley & J. H. Frank (eds) American Beetles vol. 2. Polyphaga: Scarabaeoidea through Curculionoidea pp. 160–173. CRC Press.

[KB11] Kundrata, R., & L. Bocak. 2011. The phylogeny and limits of Elateridae (Insecta, Coleoptera): is there a common tendency of click beetles to soft-bodiedness and neoteny? Zoologica Scripta 40: 364–378.

[PS07] Platia, G., & R. Schimmel. 2007. Click beetles of Taiwan collected by the expeditions of the Hungarian Natural History Museum in the years 1995 to 2003 (Coleoptera: Elateridae). Annales Historico-Naturales Musei Nationalis Hungarici 99: 49–91.

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