Larvae of Alaus myops, A. oculatus, Chalcolepidius porcatus, Hemirhipus apicalis and generic larval characterization (Elateridae, Agrypninae, Hemirhipini)

Larva of Hemirhipus Latreille, 1825 is herein described for the first time. Larvae of Chalcolepidius porcatus (Linnaeus, 1767), from Peru, and Hemirhipus apicalis Candèze, 1857, from Argentina, are described and of Alaus myops (Fabricius, 1801), from USA (Illinois and Maryland), and A. oculatus (Linnaeus, 1758), from USA (Illinois and Florida), are redescribed and illustrated. A comparison among the known larvae in each genus is presented. An historical review and the larval characterization of five genera and fourteen species of Hemirhipini genera are also included.

Review of the genus Chalcolepidius Eschscholtz, 1829 (Coleoptera, Elateridae, Agrypninae)

The genus Chalcolepidius is revised. Type specimens of 65 nominal species, except C. costatus Pjatakowa, 1941, C. fleutiauxi Pjatakowa, 1941 and C. viriditarsus Schwarz, 1906, are examined. Eighty five species are studied, of which 34 are synonymyzed and 12 new species described; three species, C. alicii Pjatakowa, 1941, C. haroldi Candèze, 1878 and C. unicus Fleutiaux, 1910, formely included in this genus, are not congeneric and are removed; C. validus Candèze, 1857 is revalidated. The genus is now formed by 63 species. Redescriptions, illustrations and a key for the examined species, and a cladistic analysis for groups of species are also included. New synonyms established: C. apacheanus Casey, 1891 = C. simulans Casey, 1907 syn. nov. = C. acuminatus Casey, 1907 syn. nov. = C. nobilis Casey, 1907 syn. nov.; C. approximatus Erichson, 1841 = C. aztecus Casey, 1907 syn. nov. = C. niger Pjatakowa, 1941 syn. nov.; C. attenuatus Erichson, 1841 = C. cuneatus Champion, 1894 syn. nov. = C. tenuis Champion, 1894 syn. nov.; C. aurulentus Candèze, 1874 = C. candezei Dohrn, 1881 syn. nov. = C. grossheimi Pjatakowa, 1941 syn. nov.; C. bomplandii Guérin, 1844 = C. humboldti Candèze, 1881 syn. nov.; C. chalcantheus Candèze, 1857 = C. violaceous Pjatakowa, 1941 syn. nov.; C. cyaneus Candèze, 1881 = C. scitus Candèze, 1889 syn. nov. = C. abbreviatovittatus Pjatakowa, 1941 syn. nov.; C. desmarestii Chevrolat, 1835 = C. brevicollis Casey, 1907 syn. nov.; C. gossipiatus Guérin, 1844 = C. erichsonii Guérin-Méneville, 1844 syn. nov. = C. lemoinii Candèze, 1857 syn. nov.; C. inops Candèze, 1886 = C. murinus Champion, 1894 syn. nov.; C. jansoni Candèze, 1874 = C. mucronatus Candèze, 1889 syn. nov.; C. lacordairii Candèze, 1857 = C. exquisitus Candèze, 1886 syn. nov. = C. monachus Candèze, 1893 syn. nov.; C. lenzi Candèze, 1886 = C. behrensi Candèze, 1886 syn. nov.; C. oxydatus Candèze, 1857 = C. jekeli Candèze, 1874 syn. nov.; C. porcatus (Linnaeus, 1767) = C. peruanus Candèze, 1886 syn. nov. = C. flavostriatus Pjatakowa, 1941 syn. nov. = C. herbstii multistriatus Golbach, 1977 syn. nov.; C. rugatus Candèze, 1857 = C. amictus Casey, 1907 syn. nov.; C. smaragdinus LeConte, 1854 = C. ostentus Casey, 1907 syn. nov. = C. rectus Casey, 1907 syn. nov.; C. sulcatus (Fabricius, 1777) = C. herbstii Erichson, 1841 syn. nov; C. virens (Fabricius, 1787) = C. perrisi Candèze, 1857 syn. nov.; C. virginalis Candèze, 1857 = C. championi Casey, 1907 syn. nov.; C. viridipilis (Say, 1825) = C. debilis Casey, 1907 syn. nov.; C. webbi LeConte, 1854 = C. sonoricus Casey, 1907 syn. nov.; C. zonatus Eschscholtz, 1829 = C. longicollis Candèze, 1857 syn. nov. New species described: C. albisetosus sp. nov. (Ecuador), C. albiventris sp. nov. (Mexico: Veracruz), C. copulatuvittatus sp. nov. (Venezuela), C. extenuatuvittatus sp. nov. (Venezuela), C. fasciatus sp. nov. (Mexico: Durango), C. ferratuvittatus sp. nov. (Ecuador), C. proximus sp. nov. (Mexico: Sinaloa), C. serricornis sp. nov. (Mexico: Veracruz), C. spinipennis sp. nov. (Mexico: Veracruz), C. supremus sp. nov. (Venezuela), C. truncuvittatus sp. nov. (Mexico: Tamaulipas) and C. virgatipennis sp. nov. (Mexico: Durango). Redescribed species: C. angustatus Candèze, 1857, C. apacheanus Casey, 1891, C. approximatus Erichson, 1841, C. attenuatus Erichson, 1841, C. aurulentus Candèze, 1874, C. bomplandii Guérin-Méneville, 1844, C. boucardi Candèze, 1874, C. chalcantheus Candèze, 1857, C. corpulentus Candèze, 1874, C. cyaneus Candèze, 1881, C. desmarestii Chevrolat, 1835, C. dugesi Candèze, 1886, C. erythroloma Candèze, 1857, C. eschscholtzi Chevrolat, 1833, C. exulatus Candèze, 1874, C. fabricii Erichson, 1841, C. forreri Candèze, 1886, C. fryi Candèze, 1874, C. gossipiatus Guérin-Méneville, 1844, C. inops Candèze, 1886, C. jansoni Candèze, 1874, C. lacordairii Candèze, 1857, C. lafargi Chevrolat, 1835, C. lenzi Candèze, 1886, C. limbatus (Fabricius, 1777), C. mexicanus Castelnau, 1836, C. mniszechi Candèze, 1881, C. mocquerysii Candèze, 1857, C. morio Candèze, 1857, C. obscurus Castelnau, 1836, C. oxydatus Candèze, 1857, C. porcatus (Linnaeus, 1767), C. pruinosus Erichson, 1841, C. rodriguezi Candèze, 1886, C. rostainei Candèze, 1889, C. rubripennis LeConte, 1861, C. rugatus Candèze, 1857, C. silbermanni Chevrolat, 1835, C. smaragdinus LeConte, 1854, C. sulcatus (Fabricius, 1777), C. tartarus Fall, 1898, C. validus Candèze, 1857, reval., C. villei Candèze, 1878, C. virens (Fabricius, 1787), C. virginalis Candèze, 1857, C. viridipilis (Say, 1825), C. webbi LeConte, 1854, C. zonatus Eschscholtz, 1829.

Cladistic analysis of Hemirhipini with establishment of Propalaus gen. nov. (Coleoptera, Elateridae, agrypninae)

This article includes a cladistic analysis of the tribe Hemirhipini. Are included 20 Hemirhipini genera (sensu Casari-Chen 1994), Saltamartinus Casari (1996b) (Hemirhipini), 6 genera excluded from Hemirhipini and kept in Agrypninae (formerly Pyrophorinae) (Casari-Chen 1993) and also, Aphileus Candèze (1857), Pyrophorus Billberg (1820) and Thoramus Sharp (1877). The type-species of the majority of genera and all species of the American genera (except Saltamartinus viduus (Chevrolat 1867)) are included. This analysis demonstrates that 30 genera belong to Hemirhipini: Abiphis Fleutiaux (1926), Alaolacon Candèze (1865), Alaomorphus Hauser (1900), Alaus Eschscholtz (1829), Aliteus Candèze (1857), Anthracalaus Fairmaire (1888), Aphileus Candèze (1857), Austrocalais Neboiss (1967), Calais Castelnau (1836), Catelanus Fleutiaux (1942), Chalcolepidius Eschscholtz (1829), Chalcolepis Candèze (1857), Conobajulus Van Zwaluwenburg (1940), Coryleus Fleutiaux (1942), Cryptalaus Ôhira (1967), Eleuphemus Hyslop (1921), Eumoeus Candèze (1874), Fusimorphus Fleutiaux (1942), Hemirhipus Latreille (1829), Lacais Fleutiaux (1942), Lycoreus Candèze (1857), Mocquerysia Fleutiaux (1899), Neocalais Girard (1971), Pherhimius Fleutiaux (1942), Phibisa Fleutiaux (1942), Propalaus gen. nov., Pseudocalais Girard (1971), Saltamartinus Casari (1996), Tetrigus Candèze (1857) and Thoramus Sharp (1877). The species included in Alaus do not make a monophyletic group and Propalaus gen. nov. is established to include Alaus alicii (Pjatakowa 1941) and A. haroldi (Candèze 1878). A description of Propalaus gen. nov. (type-species: Chalcolepidius haroldi Candèze, 1878) and a new key to Hemirhipini genera are also presented.

A phylogenetic study of the subtribe Dicrepidiina (Elateridae, Elaterinae, Ampedini)

It is presented a cladistic analysis of the Dicrepidiina aiming to test the monophyletism of the subtribe and to establish the relationships among the genera. The subtribe is composed by 36 genera and all of them, except Asebis, Lamononia, Neopsephus, Semiotopsis and Spilomorphus were included in the analysis. Fifty two species, especially the type-species of each genus were studied: Achrestus flavocinctus (Candèze, 1859), A. venustus Champion, 1895, Adiaphorus gracilis Schwarz, 1901, A. ponticerianus Candèze, 1859, Anoplischiopsis bivittatus Champion, 1895, Anoplischius bicarinatus Candèze, 1859, A. conicus Candèze, 1900, A. haematopus Candèze, 1859, A. pyronotus Candèze, 1859, Atractosomus flavescens (Germar, 1839), Blauta cribraria (Germar, 1844), Calopsephus apicalis (Schwarz, 1903), Catalamprus angustus (Fleutiaux, 1902), Crepidius flabellifer (Erichson, 1847), C. resectus Candèze, 1859, Cyathodera auripilosus Costa, 1968, C. lanugicollis (Candèze, 1859), C. longicornis Blanchard, 1843, Dayakus angularis Candèze, 1893, Dicrepidius ramicornis (Palisot de Beauvois, 1805), Dipropus brasilianus (Germar, 1824), D. factuellus Candèze, 1859, D. laticollis (Eschscholtz, 1829), D. pinguis (Candèze, 1859), D. schwarzi (Becker, 1961), Elius birmanicus Candèze, 1893, E. dilatatus Candèze, 1878, Heterocrepidius gilvellus Candèze, 1859, H. ventralis Guérin-Méneville, 1838, Lampropsephus cyaneus (Candèze, 1878), Loboederus appendiculatus (Perty, 1830), Olophoeus gibbus Candèze, 1859, Ovipalpus pubescens Solier, 1851, Pantolamprus ligneus Candèze, 1896, P. mirabilis Candèze, 1896, P. perpulcher Westwood, 1842, Paraloboderus glaber Golbach, 1990, Proloboderus crassipes Fleutiaux, 1912, Propsephus beniensis (Candèze, 1859), P. cavifrons (Erichson, 1843), Pseudolophoeus guineensis (Candèze, 1881), Rhinopsephus apicalis (Schwarz, 1903), Sephilus formosanus Schwarz, 1912, S. frontalis Candèze, 1878, Singhalenus gibbus Candèze, 1892, S. taprobanicus Candèze, 1859, Sphenomerus antennalis Candèze, 1859, S. brunneus Candèze, 1865, Spilus atractomorphus Candèze, 1859, S. nitidus Candèze, 1859, Stenocrepidius simonii Fleutiaux, 1891 and Trielasmus varians Blanchard, 1846. Chalcolepidius zonatus (Hemirhipini, Agrypninae), Ctenicera silvatica (Prosternini, Prosterninae), and species of the other subtribes of Ampedini (Elaterinae): Ampedus sanguineus (Ampedina), Melanotus spernendus (Melanotina) and Anchastus digittatus and Physorhinus xanthocephalus (Physorhinina) were used as outgroups. The results of the phylogenetic analysis demonstrated that Dicrepidiina, as formerly defined, does not form a monophyletic group. One genus, represented by Ovipalpus pubescens, was removed from the subtribe. The subtribe is characterized by presence of lamella under 2nd and 3rd tarsomeres of all legs. Also, it was revealed that the genera Achrestus, Anoplischius, Dipropus and Propsephus are not monophyletic. Due to the scarcity of information, all the studied species are redescribed and illustrated.

A phylogenetic study of the subtribe Dicrepidiina (Elateridae, Elaterinae, Ampedini)

It is presented a cladistic analysis of the Dicrepidiina aiming to test the monophyletism of the subtribe and to establish the relationships among the genera. The subtribe is composed by 36 genera and all of them, except Asebis, Lamononia, Neopsephus, Semiotopsis and Spilomorphus were included in the analysis. Fifty two species, especially the type-species of each genus were studied: Achrestus flavocinctus (Candèze, 1859), A. venustus Champion, 1895, Adiaphorus gracilis Schwarz, 1901, A. ponticerianus Candèze, 1859, Anoplischiopsis bivittatus Champion, 1895, Anoplischius bicarinatus Candèze, 1859, A. conicus Candèze, 1900, A. haematopus Candèze, 1859, A. pyronotus Candèze, 1859, Atractosomus flavescens (Germar, 1839), Blauta cribraria (Germar, 1844), Calopsephus apicalis (Schwarz, 1903), Catalamprus angustus (Fleutiaux, 1902), Crepidius flabellifer (Erichson, 1847), C. resectus Candèze, 1859, Cyathodera auripilosus Costa, 1968, C. lanugicollis (Candèze, 1859), C. longicornis Blanchard, 1843, Dayakus angularis Candèze, 1893, Dicrepidius ramicornis (Palisot de Beauvois, 1805), Dipropus brasilianus (Germar, 1824), D. factuellus Candèze, 1859, D. laticollis (Eschscholtz, 1829), D. pinguis (Candèze, 1859), D. schwarzi (Becker, 1961), Elius birmanicus Candèze, 1893, E. dilatatus Candèze, 1878, Heterocrepidius gilvellus Candèze, 1859, H. ventralis Guérin-Méneville, 1838, Lampropsephus cyaneus (Candèze, 1878), Loboederus appendiculatus (Perty, 1830), Olophoeus gibbus Candèze, 1859, Ovipalpus pubescens Solier, 1851, Pantolamprus ligneus Candèze, 1896, P. mirabilis Candèze, 1896, P. perpulcher Westwood, 1842, Paraloboderus glaber Golbach, 1990, Proloboderus crassipes Fleutiaux, 1912, Propsephus beniensis (Candèze, 1859), P. cavifrons (Erichson, 1843), Pseudolophoeus guineensis (Candèze, 1881), Rhinopsephus apicalis (Schwarz, 1903), Sephilus formosanus Schwarz, 1912, S. frontalis Candèze, 1878, Singhalenus gibbus Candèze, 1892, S. taprobanicus Candèze, 1859, Sphenomerus antennalis Candèze, 1859, S. brunneus Candèze, 1865, Spilus atractomorphus Candèze, 1859, S. nitidus Candèze, 1859, Stenocrepidius simonii Fleutiaux, 1891 and Trielasmus varians Blanchard, 1846. Chalcolepidius zonatus (Hemirhipini, Agrypninae), Ctenicera silvatica (Prosternini, Prosterninae), and species of the other subtribes of Ampedini (Elaterinae): Ampedus sanguineus (Ampedina), Melanotus spernendus (Melanotina) and Anchastus digittatus and Physorhinus xanthocephalus (Physorhinina) were used as outgroups. The results of the phylogenetic analysis demonstrated that Dicrepidiina, as formerly defined, does not form a monophyletic group. One genus, represented by Ovipalpus pubescens, was removed from the subtribe. The subtribe is characterized by presence of lamella under 2nd and 3rd tarsomeres of all legs. Also, it was revealed that the genera Achrestus, Anoplischius, Dipropus and Propsephus are not monophyletic. Due to the scarcity of information, all the studied species are redescribed and illustrated.

Evolutionary chromosomal differentiation among four species of Conoderus Eschscholtz, 1829 (Coleoptera, Elateridae, Agrypninae, Conoderini) detected by standard staining, C-banding, silver nitrate impregnation, and CMA(3)/DA/DAPI staining

The speciose Brazilian Elateridae fauna is characterized by high karyotypic diversity, including one species (Chalcolepidius zonatus Eschscholtz, 1829) with the lowest diploid number within any Coleoptera order. Cytogenetic analysis of Conoderus dimidiatus Germar, 1839, C. scalaris (Germar, 1824,) C. ternarius Germar, 1839, and C. stigmosus Germar, 1839 by standard and differential staining was performed with the aim of establishing mechanisms of karyotypic differentiation in these species. Conoderus dimidiatus, C. scalaris, and C. ternarius have diploid numbers of 2n(male) = 17 and 2n(female) = 18, and a X0/XX sex determination system, similar to that encountered in the majority of Conoderini species. The karyotype of C. stigmosus was characterized by a diploid number of 2n=16 and a neoXY/neoXX sex determination system that was highly differentiated from other species of the genus. Some features of the mitotic and meiotic chromosomes suggest an autosome/ancestral X chromosome fusion as the cause of the neoXY system origin in C. stigmosus. C-banding and silver impregnation techniques showed that the four Conoderus species possess similar chromosomal characteristics to those registered in most Polyphaga species, including pericentromeric C band and autosomal NORs. Triple staining techniques including CMA(3)/DA/DAPI also provided useful information for differentiating these Conoderus species. These techniques revealed unique GC-rich heterochromatin associated with NORs in C. scalaris and C. stigmosus and CMA(3)-heteromorphism in C. scalaris and C. ternarius.