Research Article |
Corresponding author: Hume B. Douglas ( hume.douglas@agr.gc.ca ) Academic editor: Lyubomir Penev
© 2017 Hume B. Douglas.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Douglas HB (2017) World reclassification of the Cardiophorinae (Coleoptera, Elateridae), based on phylogenetic analyses of morphological characters. ZooKeys 655: 1-130. https://doi.org/10.3897/zookeys.655.11894
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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; Dicronychussensu Platia, 1994; Dicronychussensu Méquignon, 1931; Craspedostethussensu Schwarz, 1906 (i.e., including Tropidiplus Fleutiaux, 1903); Paracardiophorussensu 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.
Cardiophorinae , Negastriinae , Physodactylinae , Phylogeny, Biogeography
The Cardiophorinae are known from all continents except Antarctica and from most large temperate and tropical islands. While larvae of Horistonotus uhleri Horn, 1871 attack roots of corn, cotton, oats, peanuts and tobacco (
Prior to this study Cardiophorinae had 29 described extant genera worldwide including about 1100 extant species, and two fossil species (
The Cardiophorinae were divided into two tribes when
Although little-tested phylogenetically, the monophyly of the Cardiophorinae has never been questioned in the literature. Subfamily-level non-monophyly is however possible due to inconsistencies in the characters used to separate Cardiophorinae from Negastriinae. Several apparent synapomorphies unite Negastriinae and Cardiophorinae: closed mesocoxal cavities; hind wing without anal cell; and basally-fused parameres, articulated at their midlength (
Other putative evidence for cardiophorine monophyly comes from the distinctive cardiophorine larvae (
A previous study (
No one has phylogenetically tested hypotheses about the membership or internal groupings of the Cardiophorinae. Additionally, no work including keys and diagnoses for all cardiophorine genera has been published since
I present here the only phylogenetic analysis of the Cardiophorinae to date, using 80 exemplar-species including much of the available morphological variation. These include 56 species from 27 of 29 described cardiophorine genera. The type species of 27 genus-level cardiophorine taxa and 20 outgroup taxa were included to ensure that included species truly represent named genera. Some additional morphologically-divergent or geographically distant members of genera are added as preliminary tests of generic monophyly. Outgroups represented most elateroid taxa most expected to confound elaterid monophyly. This study also tests the hypothesis that Nyctor Semenov-Tian-Shanskij & Pjatakova, 1936 is sister to the remainder of the Cardiophorinae, and thereby also testing the validity of subfamily Nyctorini.
Objectives of this study are: to test the monophyly of Cardiophorinae, its tribes, and genera. These results are used to redescribe the Cardiophorinae and its tribes and provide keys and diagnoses to define all included genera. Taxa are transferred as required to reflect phylogenetic findings and accepted taxonomic concepts.
Specimens examined for morphological coding belonged to 29 insect collections (Table
Non-type specimens were identified by comparison with types (types were examined for 40 species) or specimens identified by experienced workers (three species,
AMNH | American Museum of Natural History, New York, New York, USA |
NHM | The Natural History Museum [formerly British Museum (Natural History), BMNH], London, England |
BPBM | Bernice P. Bishop Museum, Honolulu, Hawaii, USA |
CASC | California Academy of Science, Department of Entomology, San Francisco, California, USA |
CMNC | Canadian Museum of Nature, Ottawa, Canada |
CNCI | Canadian National Collection of Insects, Arachnids, and Nematodes, Agriculture and Agri-Food Canada, Ottawa, Canada |
CUIC | Cornell University Insect Collection, Department of Entomology, Cornell University, Ithaca, New York, USA |
DEBU | University of Guelph Insect Collection, Guelph, Ontario, Canada |
DEIC | Institut fur Pflanzenschutzforschung, Eberswalde, Germany |
FSCA | Florida State Collection of Arthropods, Division of Plant Industry, Florida Department of Agriculture and Consumer Services, Gainesville, Florida, USA |
ISNB | Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium |
LSUK | Linnean Society, London, United Kingdom |
MCZC | Museum of Comparative Zoology, Harvard University, Cambridge Massachusetts, USA |
MNHN | Muséum National d’Histoire Naturelle, Entomologie, Paris, France |
MSUC | Michigan State University, Department of Entomology Museum, East Lansing Michigan, USA |
MZHF | University of Helsinki, Zoological Museum, Helsinki, Finland |
MZSP | Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil |
NZAC | New Zealand Arthropod Collection, Landcare Research, Auckland, New Zealand |
SANC | South African National Collection of Insects, Pretoria, South Africa |
SEMC | Snow Entomological Museum, Kansas State Biological Survey, University of Kansas, Lawrence, Kansas, USA |
TAMU | Insect Collection, Department of Entomology, Texas A&M University, College Station, Texas, USA |
TARI | Taiwan Agricultural Research Institute, Taichung, Taiwan |
UCMP | University of California Museum of Palaeontology, Berkeley, California, USA |
USNM | United States National Museum (Natural History), Washington, District of Columbia, USA |
ZMAS | Russian Academy of Sciences, Zoological Institute, St. Petersburg, Russia |
ZMHB | Museum für Naturkunde der Humboldt-Universität, Berlin, Germany |
ZMUC | University of Copenhagen, Zoological Museum, Copenhagen, Denmark |
ZMUM | Zoological Museum of Moscow Lomonosov State University, Moscow, Russia |
For this study, 51 ingroup and 26 outgroup species were coded for phylogenetic analysis (
Ingroup taxa included 53 cardiophorine species, including 5 taxa found to be near Cardiophorinae by
The outgroup (24 taxa) includes subfamilies Negastriinae, Elaterinae, Agrypninae, and Dendrometrinae. Previous studies indicated the Negastriinae were the sister group to (
Specimens were relaxed for examination by placement in nearly boiling distilled water for 10–30 minutes. Wings were photographed in water under a cover slip on a glass microscope slide. Male and female genitalia were prepared and examined as outlined in
Morphological characters were coded using majority coding of polymorphisms in order to use all available information and avoid bias. For qualitatively defined characters, majority coding was practiced by coding the character state most commonly observed in each species. For 27 quantitatively coded morphometric characters (Table
Phylogenetic characters. Morphological characters used for phylogenetic analysis. Quantitative characters are indicated by the term “Quantitative.”; described here are the measurements and ratios used to obtain data for quantitative coding. Length refers to the portion of the distance between two points parallel to the longitudinal axis of the specimen’s body (e.g., measurements a–c, Fig.
1 | Antennae: 0) with 11 antennomeres [Cardiophorus gramineus]; 1) with 12 antennomeres [Pityobius anguinus] |
2 | Quantitative. Ratio of lengths of antennomeres: two: three |
3 | Quantitative. Ratio of lengths of antennomeres: four: three |
4 | Quantitative. Ratio of dimensions of antennomere 11: (length): (maximum height) |
5 | Antennomere 1 with dorsolateral carina: 0) absent [Elater ferrugineus]; 1) present [Zorochros demustoides] |
6 | Antennae with sensory elements beginning on antennomere: 0) 3 [Cardiophorus gramineus; 1) 4 [Elater ferrugineus] |
7 | Quantitative. Ratio of dimensions of antennal fossa: (width of fossa):(minimum distance from fossa to eye) |
8 | Head with area between antenna fossa and compound eye: 0) unsculptured [Cardiophorus gramineus]; 1) with carina connecting fossa and eye [Esthesopus castaneus], or with 2 pits with non-depressed area between them [Cardiotarsus mjobergi]; 2) with a single pit [Aptopus agrestis] |
9 | Mandibular apex: 0) unidentate (simple) [Cebrio gigas]; 1) bidentate [Paracardiophorus musculus] 2) tridentate [Buckelater argutus] (O) |
10 | Labrum: 0) evenly convex dorsally [Cardiophorus gramineus]; 1) character state not assigned; 2) flat [Cebrio gigas] or broadly concave [Craspedostethus rufiventris] |
11 | Supra-antennal carina with split next to eyes: 0) absent (Fig. |
12 | Quantitative. Ratio of dimensions of frontoclypeal region: (distance between supra-antennal carina and labrum at midline): (minimum distance between antennal fossae) |
13 | Frontoclypeal region with carinae from bases of mandibles extending meso-dorsad to supra-antennal carina: 0) absent [Cardiophorus gramineus]; 1) present [Dicrepidius ramicornis] |
14 | Fronto-clypeus at midline in side view, with concavity between antero-ventral edge (adjacent to labrum) and supra-antennal carina: 0) absent [Agriotes sputator]; 1) present [Cardiophorus gramineus] |
15 | Fronto-clypeus with crenulations on anterior margin: 0) absent [Elater ferrugineus]; 1) present [Neoarhaphes americanus] |
16 | Frons with mesal groove: 0) absent [Elater ferrugineus]; 1) present [Negastrius americanus] |
17 | Frons with supra-orbital groove: 0) absent [Elater ferrugineus]; 1) present (Figs |
18 | Quantitative. (Ocular index) Ratio of dimensions of compound eyes: (maximum distance between outer edges of compound eyes)/ (minimum distance between inner edges of eyes) |
19 | Apical segment of maxillary palp: 0) securiform to subtriangular (apex truncate, widest near apex) [Cardiophorus gramineus]; 1) oblong-ovate (apex rounded, sides equally curved, widest near midlength) [Aphricus australicus]; 2) lanceolate [Adrastus pallens]; 3) character state not assigned; 4) bottle shaped (lageniform) [Arhaphes diptychus] |
20 | Apical segment of labial palp: 0) securiform or subtriangular [Cardiophorus gramineus]; 1) oblong-ovate [Aphricus australicus]; 2) lanceolate [Adrastus pallens]; 3) bottle shaped [Arhaphes diptychus] |
21 | Mentum: 0) without macrosetae [Cardiophorus gramineus]; 1) with macrosetae [Elater ferrugineus] |
22 | Pronotum with space between punctures on disc: 0) flat [Cardiophorus gramineus]; 1) with tubercles [Zorochros demustoides], or ridges [Negastrius pulchellus] |
23 | Pronotum with scale-like setae, i.e., dorsoventrally compressed, and/or wider at midlength than at base: 0) absent [Cardiophorus gramineus]; 1) present [Agrypnella eburnea] |
24 | Head and pronotum with integument: 0) unicoloured [Negastrius pulchellus]; 1) with contrasting light and dark areas [Cardiophorus gramineus] |
25 | Pronotum with hind angles: 0) not truncate dorsally (Fig. |
26 | Pronotum with lateral edge of hind angle with tubercle: 0) absent [Cardiophorus gramineus]; 1) present [Platynychus indicus] |
27 | Hind angles of pronotum with dorsal carina (beside lateral pronotal carina): 0) absent [Athous vittatus]; 1) present, but not reaching anterior edge of pronotum [Elater ferrugineus]; 2) present, reaching anterior edge of pronotum [Quasimus minutissimus] (O) |
28 | Pronotum at middle of posterior edge with: 0) arcuate indentation between 2 apices [Elater ferrugineus]; 1) point or lobe between 2 apices (= 3 apices) [Cardiophorus gramineus]; 2) single arcuate lobe (= 1 apex) [Athous vittatus] |
29 | Pronotum with sublateral incisions: 0) absent, and longitudinal carinae absent [Elater ferrugineus]; 1) present, without longitudinal carinae [Dicrepidius ramicornis]; 2) present, and with longitudinal carinae [Cardiophorus gramineus] (O) |
30 | Pronotum with lateral carina: 0) reaching from anterior edge to posterior or lateral edge of hind angle [Elater ferrugineus]; 1) absent anteriorly (ventrad of edge of pronotum in some) [Cardiophorus gramineus] |
31 | Pronotum extending laterally beyond lateral carina: 0) for entire length (Figs |
32 | Posterior edges of hypomeron mesad of hind angles: 0) with rectangular or semicircular indentations (Fig. |
33 | Hypomera with loop shaped carinae near procoxae: 0) absent [Elater ferrugineus]; 1) present [Arhaphes diptychus] |
34 | Prosternum with anterior edge: 0) short, exposing labium [Physodactylus henningi]; 1) not short, produced as lobe, concealing labium when head not extended [Cardiophorus gramineus] |
35 | Prosternum with anterior edge at midline: 0) arcuate [Elater ferrugineus]; 1) notched [undescribed species, New Zealand, transferred in this study to Aphricus]; 2) with tubercle [Esthesopus castaneus] |
36 | Prosternum with sides near midlength: 0) straight [Cardiophorus gramineus] to weakly concave [Elater ferrugineus]; 1) convex [Negastrius pulchellus] |
37 | Pronotosternal sutures with anterior ends: 0) closed [Cardiophorus gramineus]; 1) open, produced into grooves large enough to guide, but not conceal, antennae [Agriotes sputator]; 2) excavated deeply enough to conceal part of length of antennae [Agrypnus murinus] (O) |
38 | Pronotosternal sutures: 0) not interrupted, pronotum and prosternum not fused [Elater ferrugineus]; 1) partly interrupted, pronotum and prosternum fused [Arhaphes diptychus] |
39 | Prosternal process with “V” shaped carina on sides of ventral surface in ventral view: 0) absent or interrupted [Elater ferrugineus]; 1) complete, connected basally to carinae surrounding procoxal cavities [Paracardiophorus musculus] |
40 | Quantitative. Ratio of dimensions of prosternal process: (length of portion of prosternal process extending posterad of procoxae): (maximum length of exposed part of procoxae in ventral view) |
41 | Quantitative. Ratio of dimensions of prosternal process (Fig. |
42 | Quantitative. Ratio of dimensions of prosternal process (Fig. |
43 | Quantitative. Ratio of dimensions of Prosternal process (Fig. |
44 | Prosternal process (anterad of ventral apex) with angle between ventral surface and ventral surface of middle of prosternum anterior to procoxae: 0) less than 30° (horizontal) [Elater ferrugineus]; 1) more than 30° (inclined dorsally) [Dima elateroides] |
45 | Procoxal cavities: 0) open posteriorly [Elater ferrugineus]; 1) fully closed posteriorly [Paracardiophorus musculus] |
46 | Protibiae near apex with posterior surface: 0) convex, not modified for digging [Cardiophorus gramineus]; 1) flattened, concave, or broadened apically, apparently modified for digging [Cebrio gigas] |
47 | Pronotum with punctures on disc: 0) elongate [Athous vittatus]; 1) not elongate [Cardiophorus gramineus]; 2) absent, setae on tubercles [Anelastes druryi] |
48 | Scutellum with middle of anterior edge: 0) straight [Pyrophorus noctilucus] or convex [Elater ferrugineus] (Fig. |
49 | Scutellum with anterolateral edges: 0) convex throughout [Cardiophorus gramineus]; 1) straight [Athous vittatus], or concave posterior to anterolateral corners [Agriotes sputator] |
50 | Scutellum with posterolateral edges: 0) with straight [Cardiophorus gramineus] (Fig. |
51 | Sides of mesosternal cavity with antero-ventral angles in lateral view: 0) prominent and rounded (Fig. |
52 | Sides of mesosternal cavity posterior to anterior edge of mesocoxae in ventral view: 0) U shaped [Cardiophorus gramineus]; 1) straight, forming a V [Elater ferrugineus]; 2) with anteromesal projection [Arhaphes diptychus]; 3) not assigned; 4) with 3 obtuse angles and 4 sides [Cardiophorus convexulus] |
53 | Sides of mesosternal cavity anterior to mesocoxae: 0) sinuate [Cardiophorus gramineus]; 1) straight (parallel or anteriorly convergent) [Semiotus furcatus] |
54 | Anterior edge of mesosternum in ventral view: 0) concave lateral to anterior protrusions of mesosternal fossa [Elater ferrugineus]; 1) convex lateral to anterior protrusions of mesosternal fossa [Negastrius pulchellus], or evenly convex, uninterrupted by mesosternal fossa [Zorochros demustoides] |
55 | Mesepisternum with anterior projection of anteromesal corners extending beyond junction with mesosternum: 0) concave mesally forming an acute point (Fig. |
56 | Mesepisternum with circular pit at anteromesal corner: 0) absent [Elater ferrugineus] (some with groove [Athous vittatus]); 1) present [Agrypnus murinus] |
57 | Angle of anterolateral corner of mesepisternum (angle between tangents of edge of sclerite at lowest part of concavity immediately mesad of angle, and an equidistant point on lateral edge): 0) approximately right angled (80°–100°, Fig. |
58 | Quantitative. Ratio of dimensions of mesepimeron: (maximum width, measured parallel to anterior edge)/ (maximum length, perpendicular to width) |
59 | Mesotrochantin: 0) visible [Elater ferrugineus], or partly concealed by mesosternum; 1) not visible [Cardiophorus gramineus] |
60 | Mesocoxal cavity: 0) open to both mesepimeron and mesepisternum (Fig. |
61 | Quantitative. Ratio of dimensions of midleg: (trochanter length): (femur length) |
62 | Quantitative. Ratio of dimensions of exposed portion of metepisternum: (length): (width). Width measured at midlength, not including portion covered by closed elytra |
63 | Metasternum behind mesocoxal cavities: 0) without postcoxal lines [Elater ferrugineus]; 1) with postcoxal lines, either arc-shaped [Lissomus bicolor] or loop-shaped [Quasimus minutissimus] |
64 | Metasternum with width ratio of lateral carina (raised flattened lateral edge of metasternum, Fig. |
65 | Edge of elytra in dorsal view between anterior-most point and humeral angle: 0) sinuate or with tubercle [Cardiophorus gramineus] (Fig. |
66 | Number of puncture rows or striae on elytra: 0) 0 (absent on basal half) [Cebrio gigas]; 1) 9 [Elater ferrugineus] |
67 | Elytra with intervals 1–8 on basal third: 0) flattened or rounded [Elater ferrugineus]; 1) partly or completely costate [Negastrius pulchellus] |
68 | Elytra with apical half of intervals 1–8: 0) flattened or rounded throughout[Elater ferrugineus]; 1) with at least some costate [Aphricus californicus] |
69 | Elytra with apical half of interval 9: 0) flattened or rounded [Elater ferrugineus]; 1) costate [Triplonychoidus trivittatus (Champion)] |
70 | Elytra: 0) without spots or markings [Elater ferrugineus] (some with basal markings or longitudinal stripe); 1) with distinct transverse markings or spots not confluent with anterior edge [Negastrius pulchellus] |
71 | Upper edge of elytral epipleura: 0) with minute regular serrations [Cardiophorus gramineus]; 1) without serrations [Elater ferrugineus] |
72 | Elytral apex with punctures at least 1.5X diameter of largest on anterior half: 0) absent [Elater ferrugineus]; 1) present, single [Paracardiophorus subcruciatus], or multiple [Agrypnella eburnea] |
73 | Elytron with apical shelf like extension: 0) absent [Elater ferrugineus]; 1) present [Cardiophorus nigratissimus]. |
74 | Hind wing: 0) with venation well developed, wing area greater than elytral area [Elater ferrugineus]; 1) with veins weakly sclerotised, wing area less than half of elytral area [Dima elateroides]. |
75 | Hind wing membrane: 0) not notched in anal area [Elater ferrugineus]; 1) notched in anal area (between AA3+4 and AP) [Negastrius pulchellus] |
76 | Hind wing with apical concavity: 0) absent [Elater ferrugineus]; 1) present [Berninelsonius hyperboreus] |
77 | Quantitative. Ratio of dimensions of hind wing, radial cell: (length): (width) |
78 | Quantitative. Ratio of dimensions of Hind wing: proximal, posterior angle of radial cell (measured in degrees) between tangents of: anterior edge of cell, at 1/3 distance from posterior angle to anterior edge of cell; and posterior edge of cell at an equal distance from angle) |
79 | Hind wing with vein AA3: 0) joining CuA posterior to divergence from AA4 [Elater ferrugineus]; 1) joining CuA at divergence from AA4 [Esthesopus castaneus]; 2) joining CuA, AA4 not continuing posteriorly [Negastrius pulchellus] |
80 | Hind wing with wedge cell: 0) present (Fig. |
81 | Hind wing vein MP3 and MP4 separating: 0) distal to intersection with CuA1 [Cardiophorus gramineus]; 1) proximal [Elater ferrugineus] to or at same level as [Macropogon piceus] intersection with CuA1. (CuA1 interrupted in some) |
82 | Hind wing with vein CuA1: 0) uninterrupted (Fig. |
83 | Hind wing with CuA1 0) not forked [Elater ferrugineus]; 1) forked at junction with MP3+4 forming additional closed cell [Blaiseus bedeli] (Fig. |
84 | Hind wing with vein MP3+4 with proximal extension of crossvein mp1+2-mp3+4: 0) present [Elater ferrugineus]; 1) absent (Figs |
85 | Apex of hind wing with anterior field sclerotisation ( |
86 | Apex of hind wing with median field ( |
87 | Apex of hind wing with linear sclerites of posterior field ( |
88 | Quantitative. Ratio of dimensions of metacoxal plate: (width): (length at widest point mesad of trochanter attachment) |
89 | Metacoxae with posterior excavation: 0) sufficient to cover at least 2/3 area of trochanter with hind legs withdrawn (Figs |
90 | Metacoxal plate: 0) not reaching metepisternum, or shorter than 1/3 posterior width of metepisternum lateral to intersection with metepisternum (Fig. |
91 | Metacoxal plate with mesally directed hook: 0) absent [Elater ferrugineus]; 1) present [Cardiohypnus mirabilis] |
92 | Metatibia with number of apical spurs: 0) 2 [Elater ferrugineus]; 1) 0 [Lissomus bicolor] |
93 | Quantitative. Ratio of dorsal lengths of tarsomeres of hind leg: (1): (2) |
94 | Metatarsi with only the following tarsomeres lobed or lamellate: 0) none [Elater ferrugineus]; 1) not assigned; 2) 4 [Cardiotarsus capensis Candèze]; 3) 3, 4 [Monadicus sp.]; 4) 1, 2, 3 [Athous vittatus] |
95 | Metatarsal claws with basal setae: 0) absent [Elater ferrugineus]; 1) present, multiple in some [Agrypnus murinus] |
96 | Metatarsal claws with number of apices per side (including flange sensu |
97 | Urosternites 3–6 with size of adjacent punctures: 0) approximately uniform [Elater ferrugineus]; 1) of 2 size classes [Paracardiophorus musculus] |
98 | Urosternites 3–7 with elongate punctures: 0) absent [Athous vittatus] or limited to urosternites 6 and 7; 1) throughout [Ampedus sanguineus] |
99 | Lateral edges of urosternites 3–7 with serrations: 0) present on at least some [Cardiophorus gramineus]; 1) absent [Elater ferrugineus] |
100 | Urosternites 6 and 7 with multiple longitudinal ridges: 0) absent [Elater ferrugineus]; 1) present [Tropidiplus tellinii] |
101 | Urosternite 7 (and in some 6) with second carina mesad of lateral carina: 0) absent [Cardiophorus gramineus]; 1) present [Tropidiplus tellinii] |
102 | Male urosternite 8 with mesal third of anterior sclerotised band (anterior margin between bases of anterior lobes): 0) straight, bisinuate [Arhaphes diptychus] or arcuate [Paracardiophorus musculus], width uniform or interrupted mesally [Cardiophorus gramineus]; 1) anteriorly angulate, width uniform [Zorochros demustoides]; 2) broadened mesally, anteriorly arcuate [Ampedus sanguineus], interrupted laterally in some [Physorhinus erythrocephalus (Fabricius)]; 3) indistinguishable from posterior lobes, because sclerite of uniform length across width, or longest mesally [Aphricus australicus]; 4) indistinguishable from posterior lobes, because posterior lobes connate basally [Pachyelater madagascariensis]; 5) absent (entirely unsclerotised) although posterior lobes present [Dicrepidius ramicornis] |
103 | Sclerotised basal band of male urosternite 8: 0) not extending laterally beyond posterior lobes (i.e., posterior edge of sclerite convex from apex of lobes to sides of sclerite) [Elater ferrugineus]; 1) extending laterally beyond slope of lobes (posterior edge of sclerite sinuate from apex of lobes to sides of sclerite) [Paracardiophorus musculus] |
104 | Tergite and sternite of male abdominal segment 9 articulated at: 0) sides [Elater ferrugineus]; 1) base [Blaiseus bedeli] |
105 | Male urosternite 9 with anterior end: 0) not pointed [Elater ferrugineus]; 1) pointed [Paradonus pectoralis] |
106 | Quantitative. Ratio of lengths of male urotergites: (9 [at point of greatest length])/ (10 [at midline]) |
107 | Male urotergites 9 and 10: 0) not fused mesally [Elater ferrugineus]; 1) fused mesally [Fleutiauxellus maritimus (Curtis)] |
108 | Male urotergite 9 with shape of apical emargination: 0) “U” [Elater ferrugineus]; 1) between “U” and “V” [Ampedus sanguineus]; 2) “V” [Agrypnus murinus] |
109 | Aedeagus with antero-dorsal (basal) concavity of phallobase in dorsal view: 0) simple (Fig. |
110 | Aedeagus with posterior (apical) emargination of phallobase: 0) deep and evenly concave [Elater ferrugineus]; 1) reduced, concavity nearly absent [Pityobius anguinus]; 2) produced mesally [Athous vittatus] |
111 | Quantitative. Ratio of dimensions of aedeagus: (width of posterodorsal concavity of phallobase (concavity at apical end of phallobse) [0 if absent]): (maximum width of phallobase) |
112 | Quantitative. Ratio of dimensions of aedeagus: (length of phallobase): (maximum width of phallobase) |
113 | Quantitative. Ratio of dimensions of median lobe of aedeagus: (length of apical portion [portion posterad of concavity between basal struts]): (length of concavity between basal struts) |
114 | Quantitative. Ratio of dimensions of aedeagus: (length of entire median lobe [including basal struts]): (maximum width of phallobase) |
115 | Quantitative. Ratio of dimensions of aedeagus: (length of parameres): (maximum width of phallobase) |
116 | Quantitative. Ratio of dimensions of aedeagus: (length of overlap between phallobase and parameres): (maximum width of phallobase) |
117 | Aedeagus with parameres: 0) articulated with median lobe basally, pivoting at base [Elater ferrugineus] (Fig. |
118 | Parameres in species with parameres articulated beyond bases with abrupt narrowing immediately apicad of point of articulation: 0) absent [Cardiophorus gramineus], (Fig. |
119 | Aedeagus with parameres with pre-apical or apical expansions: 0) absent [Elater ferrugineus]; 1) present, lateral or ventral [Agriotes sputator]; 2) not assigned; 3) present, mesal side [Cardiophorus luridipes Candèze] |
120 | Parameres with ratio of lateral width to dorsoventral depth of free portion: 0) less than 2 [Cardiophorus gramineus]; 1) greater than 2 [Cardiophorus luridipes] |
121 | Parameres with number of apices: 0) 1 [Elater ferrugineus], (Fig. |
122 | Parameres in dorsal view with profile of mesal edge of apices: 0) not concave [Elater ferrugineus]; 1) concave [Dicrepidius ramicornis] |
123 | Parameres with number of setae on each: 0) 0 [Agriotes sputator]; 1) 1 [Hypnoidus riparius], (Fig. |
124 | Parameres with setae: 0) restricted to sides, basad of apicolateral expansions [Dicronychus cinereus] or of apical arc in species without expansions [Esthesopus parcus Horn]; 1) restricted to apex, apicad of apicolateral expansions if present [Athous vittatus] or to apical arc if expansions absent [Selonodon speratus]; 2) both apicad and basad of apicolateral expansions [Pyrophorus noctilucus] or apical arc; ?) with position uncertain because apical arc not distinguishable and apicolateral expansions absent |
125 | Parameres with apices: 0) opaque or otherwise not abruptly translucent [Cardiophorus gramineus]; 1) abruptly translucent [Dicrepidius ramicornis] |
126 | Aedeagus with apex of median lobe in lateral view: 0) without globular expansion [Elater ferrugineus], straight or down turned; 1) with globular expansion [Esthesopus castaneus], straight or down turned; 2) without expansion, but abruptly upturned [Agrypnella eburnea] |
127 | Female urosternite 8 with lateral sclerotisations: 0) joined at or near apex [Elater ferrugineus]; 1) not joined apically [Zorochros demustoides] |
128 | Female urosternite 8 with lateral sclerotisation: 0) joined to sclerotised spiculum [Elater ferrugineus]; 1) separated from sclerotised spiculum by membranous tissue or intermittent sclerotisation [Cardiophorus gramineus] |
129 | Female urosternite 8 with spiculum: 0) present [Elater ferrugineus]; 1) absent [Pachyelater madagascariensis] |
130 | Ovipositor with sclerites of coxites: 0) narrow, occupying less than 1/2 width of coxites at midlength [Cardiophorus gramineus], (Fig. |
131 | Quantitative. Ratio of dimensions of ovipositor: (paraproct length): (coxite length) |
132 | Ovipositor with apical stylus: 0) absent [Cardiophorus gramineus]; 1) present [Agriotes sputator], (Fig. |
133 | Vagina with colleterial glands: 0) absent [Cardiophorus gramineus], (Fig. |
134 | Vagina with colleterial glands attached: 0) on either side of common oviduct [Elater ferrugineus]; 1) anterior to common oviduct [Anisomerus sylvestris]; 2) posterior to common oviduct [Athous vittatus] |
135 | Bursa copulatrix with spermathecal gland duct opening (or shared spermatheca-spermathecal gland duct): 0) away from base of anterior blind sac [Cardiophorus gramineus]; 1) adjacent to base of blind sac [Paracardiophorus musculus], (Fig. |
136 | Spermathecal gland (or shared) duct opening: 0) at distal (anterior) end of bursa copulatrix [Elater ferrugineus], (Fig. |
137 | Bursa copulatrix with spermathecal gland duct attached to: 0) main bursa [Cardiophorus gramineus], (Fig. |
138 | Female spermathecal gland duct with single row of diverticulae: 0) present [Cardiophorus gramineus]; 1) absent (but duct present) [Elater ferrugineus] |
139 | Number of coil-type spermathecae: 0) 0 [Cardiophorus gramineus]; 1) 1 [Agriotes sputator]; 2) 2 [Pyrophorus noctilucus] |
140 | Female with number of sclerotised capsule-type spermathecae: 0) 0 [Cardiophorus gramineus]; 1) 1 [Elater ferrugineus] |
141 | Bursa copulatrix with blind anterior sac attached near: 0) apex of bursa [Cardiophorus gramineus], (Fig. |
142 | Bursa copulatrix with number of blind pedunculate sacs: 0) 0; 1) 1 [Cardiophorus gramineus], (Fig. |
143 | Bursa copulatrix with number of non-pedunculate blind tubular extensions: 0) 0 [Cardiophorus gramineus], (Fig. |
144 | Bursa copulatrix with free spines (i.e., not part of sclerite with multiple spines): 0) absent [Cardiophorus gramineus], (Fig. |
145 | Bursa copulatrix with free spines: 0) not combined into paired, discrete, ovoid patches [Melanotus castanipes]; 1) present and combined into paired discrete ovoid patches, with individual spines separated by membranous tissue [Exoeolus rufescens]; 2) present and combined into flexible paired discrete ovoid patches, but with some spines partially fused [Physorhinus erythrocephalus] (O) |
146 | Bursa copulatrix with rugose, spineless, partially sclerotised patches: 0) absent [Cardiophorus gramineus], (Fig. |
147 | Bursa copulatrix with single dorsal and ventral sclerites both: 0) absent [Negastrius pulchellus], (Fig. |
148 | Bursa copulatrix with single dorsal and ventral sclerites both: 0) bilaterally symmetrical [Oedostethus femoralis]; 1) not symmetrical [Quasimus minutissimus] |
149 | Bursa copulatrix, of species with single symmetrical dorsal and ventral sclerite of bursa copulatrix, with ventral sclerite: 0) fully sclerotized at midline [Oedostethus femoralis]; 1) divided or weakly sclerotised along midline [Neoarhaphes americanus] |
150 | Bursa copulatrix with single dorsal and ventral sclerites: 0) both not ring-like, opaque at center [Oedostethus femoralis]; 1) with at least dorsal sclerite ring-like, and transparent at center [Arhaphes diptychus] |
151 | Quantitative. (meristic) Ratio of dimensions of bursa copulatrix with paired proximal sclerites: (count # of rows of spines, including outer row, which surrounds most of sclerite in most species) |
152 | Bursa copulatrix with proximal sclerites (defined as pair of spine-bearing sclerites closest to vagina (for species with only 1 pair of sclerites: defined as proximal if not surrounding base of spermathecal gland duct): 0) absent [Craspedostethus rufiventris]; 1) present [Cardiophorus gramineus], (Fig. |
153 | Bursa copulatrix with points of proximal sclerites: 0) simple [Cardiophorus gramineus], (Fig. |
154 | Bursa copulatrix with proximal sclerites: 0) ovoid [Cardiophorus gramineus], (Fig. |
155 | Bursa copulatrix with placement of proximal sclerites: 0) symmetrical [Cardiophorus gramineus]; 1) asymmetrical [Cardiophorus brunnipennis] |
156 | Quantitative. Ratio of dimensions of bursa copulatrix: (length of largest spines of a proximal sclerite [measured as smallest possible distance between a line connecting 2 adjacent apices and the deepest part of the concavity between them]): (length of sclerite) (Fig. |
157 | Bursa copulatrix with paired distal sclerites (pair farthest from vagina, at base of spermathecal gland duct or shared duct): 0) absent [Elater ferrugineus]; 1) present [Cardiophorus inflatus], fused together in some at wall of bursa [Cardiophorus gramineus] (Figs |
158 | Bursa copulatrix with two distal sclerites: 0) separate [Cardiophorus inflatus], (Fig. |
159 | Bursa copulatrix with distal sclerites: 0) smooth [Cardiophorus gramineus], (Fig. |
160 | Bursa copulatrix with distal sclerites: 0) flexible and at least in part weakly sclerotised and membranous [Cardiophorus convexus] (Fig. |
161 | Bursa copulatrix with tube-like sclerotisation of base of spermathecal gland duct: 0) absent [Cardiophorus gramineus], (Fig. |
162 | Habitat: 0) restricted to riparian areas [Negastrius pulchellus]; 1) not restricted to riparian areas [Elater ferrugineus] (Not used to infer phylogeny) |
163 | Bursa copulatrix with multiple parallel linear sclerites: 0) absent [Elater ferrugineus]; 1) present [Athous vittatus] |
Morphological character selection and coding were performed together. All observed variation was evaluated as a potential character source following one procedure. To be considered suitable, variation between homologous structures must allow diagnosis between at least one pair of species. All 136 characters that could not be described as length ratios, or counts, were treated qualitatively as binary or multistate characters. Qualitative characters included the presence or absence of structures, or objective shape descriptors (e.g., notched vs. uniformly convex). An exemplar species was assigned for each qualitatively defined character state in an effort to produce repeatable, standardized character state definitions (many follow
Phylogenetic analysis was conducted using 163 characters (Table
Although both parsimony and Bayesian analyses (as implemented by MrBayes v.3.1.2,
Bayes factors were used, as outlined by
Phylogenetic analysis was performed using both parsimony and Bayesian criteria. For Bayesian analyses, prior probability distributions were at default values of MrBayes. Gamma distribution was approximated using the default setting of four rate classes. Settings for likelihood parameters used were Mkv (nst=1, coding=variable) and Mkv (nst=1, coding=variable, rates=gamma). Searches began with randomly selected starting trees and were run for 8 million cycles (until the average standard deviation of split frequencies between four parallel runs was below 0.01). Samples of trees from the MCMC chain were taken every 100 cycles, which resulted in 80 thousand trees. All but the first 20 thousand trees were used to compute a majority rule consensus tree assigning posterior probabilities of tree topology. The matrix was analysed three times to test repeatability. Because these differed slightly, the analysis with the highest average harmonic mean log likelihood was used for phylogenetic inference.
Parsimony analysis was performed using PAUP* (
Testing hypotheses of monophyly was done by determination of the PP of the focal clade. The hypotheses tested are either ones stated explicitly as such, or ones implied by the description of taxa.
A key to the genera of cardiophorine, and corresponding diagnoses were developed using: existing diagnostic characters, the phylogenetic matrix (
Bayesian analysis of the Cardiophorinae, Negastriinae and Hypnoidini matrix resulted in trees largely agreeing with results of morphological analysis of Elateridae (
Tests of hypotheses of monophyly for Cardiophorinae and Negastriinae based on Bayesian posterior probabilities (Fig.
Hypothesis and citation | posterior probability |
---|---|
Subfamilies | |
Cardiophorinae auctorum | <0.000008 |
Cardiophorinae (w Negastrius americanus and Physodactylinae spp.) | <0.000008 |
Negastriinae (sensu Stibick, 1979a) | 0.000108 |
Tribes | |
Cardiophorini Candèze, 1859 (excluding Nyctorini) | 0.000267 |
Genera of Cardiophorinae | |
Aphricus LeConte, 1853 (+/- undescribed sp. from New Zealand) | 0.0017 |
Aptopus Eschscholtz, 1829 | 0.000033 |
Blaiseus Fleutiaux, 1931 | 0.9 |
Cardiophorus Eschscholtz, 1829 | <0.000008 |
C.: subgenus Cardiophorus | <0.000008 |
C.: subgenus Perrinellus Buysson, 1899 | 0.065 |
Cardiotarsus Eschscholtz, 1836 (+/- Cardiotarsus mjobergi) | 0.0036 |
Esthesopus Eschscholtz, 1829 | 0.016 |
Horistonotus Candèze, 1860 | 0.33 |
Paracardiophorus Schwarz, 1895b | <0.000008 |
Phorocardius Fleutiaux, 1931 | 0.00058 |
Synonymies (putative synonyms of Cardiophorus were not tested because its generic monophyly was rejected) | |
Dicronychus (= Paradicronychus (nomen nudum)): |
0.069549 |
Dicronychus (= Platynychus): |
<0.000008 |
Craspedostethus (= Tropidiplus): |
0.000008 |
Paracardiophorus (= Craspedostethus): |
<0.000008 |
Within the Cardiophorinae, there were several clades with moderately high support (e.g. Fig.
Support for a priori hypotheses of monophyly was mostly weak to absent (Table
Parsimony analysis mainly corroborated results of Bayesian analysis, also with low resolution near Cardiophorus (Fig.
Bayesian (Fig.
The resulting Bayesian tree (Fig.
The existing tribal classification of the Cardiophorinae is incorrect according to both Bayesian and parsimony analyses. This is because the monotypic Nyctorini rendered the only other tribe, the Cardiophorini, paraphyletic (Table
The only taxonomic change to Negastriinae is the transfer of Negastrius americanus from Negastrius Thomson, to Cardiophorinae (as a new genus). Within Negastriinae, three genera previously transferred from Cardiophorinae form a well-supported clade in both Bayesian (PP > 0.95 after correction for branch length, length = 27, uncorrected probability = 1.00) and parsimony (D = 5, BS = 75) analyses (Figs
The required changes of classification among the Cardiophorinae are discussed beginning at the root of Cardiophorinae in the Bayesian tree. Some paraphyletic and polyphyletic genera are recognised here, in cases where phylogenetic results did not provide well-supported alternative to the prior classification. The most basal cardiophorine node is an eight-way polytomy (Fig.
Blaiseus Fleutiaux, another basal cardiophorine, was found monophyletic here. This genus has 10 species distributed in Southeast Asia, South Africa, and Central and North America, (
Pachyelater Lesne, 1897 is a robust-bodied fossorial elaterid genus from Madagascar with sexually dimorphic males and females (Figs
The remaining taxa in the polytomy of the Bayesian analysis (Fig.
Among the genera historically confounded with Cardiophorus, the most basal is Paracardiophorus. This genus was found polyphyletic (Table
Beyond confusion with Paracardiophorus, genus Cardiophorus: subgenus Cardiophorus was paraphyletic at four nodes (Fig.
Two other Cardiophorus subgenera, Coptostethus Wollaston and Perrinellus Buysson are each based on a single evolutionarily labile character (reduction of flight wings, and narrowed base of scutellum respectively), and are probably not monophyletic (although not synonymised here). The first, Coptostethus, is a name historically applied to various short-winged Cardiophorinae. Some Cardiophorus from Africa and Eurasia possibly adapted for fossorial life have been grouped into the subgenus Perrinellus, which was not recovered as monophyletic in either analysis (PP = 0.06, Table
Globothorax Fleutiaux and Teslasena Fleutiaux (Physodactylinae) are a strongly supported (Fig.
Dicronychus Brullé was coded here based on D. cinereus Brullé, which was considered the senior synonym of the type species at the beginning of this study. Although within the Paraphyletic Cardiophorus according to both analyses (Figs
Genus Cardiophorellus Cobos also falls within the paraphyletic nominate subgenus of Cardiophorus (Fig.
Although the hypothesis of Aptopus Eschscholtz monophyly was rejected (Table
Genus Phorocardius Fleutiaux was described to include Cardiophorus-like species with apically bidentate tarsal claws (Fig.
Genus Cardiotarsus includes species from Africa, Mauritius, S. and E. Asia and Australia. These analyses included the type species (C. capensis Candèze, 1860, known here from females only), another (undescribed) African species and Cardiotarsus mjobergi, Australia’s only known species. Bayesian hypothesis testing (Table
The remaining apical southern clade (PP = 0.81, Fig.
Of the five species in the weakly supported apical clade (Fig.
This section outlines some character state changes implied by the trees (Figs
Three characters unite the Negastriinae + Cardiophorinae. The fusion of the parameres at their midlength into a tube (Char. 117, Figs
No clear evidence was found for basal synapomorphies of Negastriinae not also shared by Cardiophorinae. As found by
Several synapomorphies unite three brightly patterned riparian negastriine genera from the Neotropics (Agrypnella), the Himalayan foothills (Cardiohypnus), and Australia (Rivulicola). These are the only Negastriinae with sublateral pronotal incisions and carinae (Char. 29). They are also the only Negastriinae, except for Monadicus, with: scale-like setae (Char. 23); and the posterior edges of hypomeron mesad of hind angles with rectangular or semicircular indentations (Char. 32). Two of these, Agrypnella and Cardiohypnus, also have sides of pronotum overhanging the lateral carinae like in Cardiophorus.
Quasimus Gozis, 1886 + Yukoana Kishii 1959 (both Negastriinae, Quasimusini) share several possible synapomorphies: tarsomere 4 (and no others) is lobed on all legs (shared in Negastriinae by only Neoarhaphes
The Cardiophorinae have only two apparent synapomorphies not shared with at least some Negastriinae: the straight-sided prosternum (Char. 36); and presence of paired proximal sclerites in the bursa copulatrix (absent in Blaiseus, Craspedostethus, Floridelater (formerly Negastrius americanus), and Pachyelater, Char. 152). A third possible synapomorphy, the presence of one or two pedunculate anterior sacs of the bursa copulatrix (Char. 143) is shared by all examined Cardiophorinae and their apparent sister-taxon, Fleutiauxellus.
Within Cardiophorinae, only a few groups were united by moderate to high branch support. Pachyelater + Aphricus + undescribed species from New Zealand + Patriciella share straight sides of the mesosternal cavity posterior to anterior edge of mesocoxae (Char. 52). The Palaearctic Paracardiophorus + the Nearctic Cardiophorus cardisce + C. luridipes all share dorsally truncate pronotal hind angles.
Additional phylogenetic research with more taxon sampling is needed throughout Cardiophorinae to test generic monophyly and better understand intergeneric relationships. Additional collecting and taxon sampling would be useful among the basal cardiophorines, for which only two of nine genera are known from both sexes.
Some areas of the tree have low clade support and short branch lengths. These may approximate a hard polytomy, and thus it might be impossible to infer branching patterns using morphology alone. Combined analysis of multiple gene regions plus morphology might resolve these regions, once specimens suitable for DNA sequencing have been collected. Discovery of undescribed females or males from several genera would also provide useful data. Meanwhile I recommend continuing to recognize some heterogeneous genera until phylogenetic knowledge improves.
1 | Prosternum with sides near midlength convex, or partly fused with pronotum; if scutellum emarginate anteromesally, then with dorsal vestiture of scale-like or apically broadened setae; some with tubercles between setal punctures on pronotum; bursa copulatrix with symmetrical pair of spine-bearing sclerites absent, or connected to each other by semi-sclerotised tissue | Not Cardiophorinae: Negastriinae (revised by |
– | Prosternum with sides near midlength straight or concave, not fused with pronotum. Most with scutellum emarginate anteromesally; setae evenly tapered in all; none with tubercles between pronotal setae; most with minute serrations along upper edge of elytral epipleurae and/or at sides of urosternites 3–7; bursa copulatrix of most with symmetrical pair of separate, spine-bearing sclerites (proximal sclerites); Cardiophorinae | 2 |
2 (1) | Pronotum with complete carina at lateral edge or on hypomeron, reaching from hind angle to anterior edge (reaches only 9/10 to anterior edge in two Southeast Asian Paraplatynychus species). From Oriental and Ethiopian realms) | 3 |
– | Pronotum with lateral carina not reaching anterior edge (ventrad of lateral edge of pronotum in some, called submarginal line in earlier publications), [rest of Cardiophorinae] | 4 |
3 (2) | Tarsal claws with or without basal point; tarsal claws with basal setae (Fig. |
Tropidiplus Fleutiaux, 1903 |
– | Tarsal claws with both basal and apical points, without basal setae; bursa copulatrix with proximal sclerites bilobed (Fig. |
Paraplatynychus Fleutiaux, 1931 |
4 (2) | Pronotum with lateral carina present (short in some) but below edge of dorsal part of pronotum (Fig. |
5 |
– | Pronotum with lateral carina extending anterad from hind angles following lateral edge (Fig. |
20 |
5 (4) | Tarsomere 4 with ventral lobe or pad extending beyond base of tarsomere 5 (Fig. |
6 |
– | Tarsomere 4 without ventral lobe or pad extending beyond base of tarsomere 5 | 7 |
6 (5) | Tarsal claws one apex per side (Africa, Mauritius, S. and E. Asia, Japan, Taiwan) |
Cardiotarsus |
– | Tarsal claws with both basal and apical points (Figs |
Odontocardus Fleutiaux, 1931 |
7 (5) | Tarsal claws with 3 or more points per side (as in Fig. |
Aptopus Eschscholtz, 1829 |
– | Tarsal claws with single apical point or both basal and apical points on each side | 8 |
8 (7) | Tarsal claws with 2 points per side (Figs |
9 |
– | Tarsal claws with only 1 point per side | 14 |
9 (8) | Procoxal cavities open | 10 |
– | Procoxal cavities closed | 11 |
10 (9) | Tarsal claws with second point near apex on each side (Fig. |
Phorocardius Fleutiaux, 1931 |
– | Tarsal claws with second point at base of each side (Fig. |
Dicronychus Brullé, 1832. Some brachypterous spp. of Cardiophorus s.g. Coptostethus key to here |
11 (9) | Head with area between antennal fossa and compound eye unsculptured; bursa copulatrix with paired distal sclerites (pair farthest from vagina) present and fused into a “U” shape (Fig. |
Platynychus Motschulsky, 1858 (monophyly unknown) |
– | Head with area between antennal fossa and compound eye with carina connecting fossa and eye, or with 2 pits with non-depressed area between them or with a single pit; bursa copulatrix without distal sclerites (i.e. a second pair, farther from vagina, at base of spermathecal gland duct); base of spermathecal gland duct with tube-like sclerotisation (Figs |
12 |
12 (11) | Head with area between antennal fossa and compound eye with carina connecting fossa and eye, or with 2 pits with non-depressed area between them; tarsal claws with ventral surface convex mesad of basal apex (as in Fig. |
Displatynychus Ôhira, 1987 |
– | Head with area between antennal fossa and eye with a single pit; tarsal claws with ventral surface concave mesad of basal apex (Fig. |
Diocarphus Fleutiaux, 1947 |
14 (8) | Mandibular apex unidentate (simple) | 15 |
– | Mandibular apex bidentate or tridentate | 17 |
15 (14) | Head with supra-orbital groove absent; posterior edges of hypomeron mesad of hind angles without indentations (Uzbekistan, only males known) | Metacardiophorus Gurjeva, 1966 |
– | Head with supra-orbital groove (Fig. |
16 |
16 (15) | Scutellum with anterior edge broadly concave (Fig. |
Cardiophorellus Cobos, 1970 (3 spp., Congo). Subgenus Parapleonomus not examined here. |
– | Scutellum with anterior edge abruptly emarginate (Fig. |
Margogastrius Schwarz, 1903 (monotypic, M. schneideri Schwarz) |
17 (14) | Edge of elytra in dorsal view between anterior-most point and humeral angle arcuate or straight, without sinuation (Fig. |
18 |
– | Edge of elytra in dorsal view between anterior-most point and humeral angle sinuate or tuberculate (Fig. |
19 |
18 (17) | Head with supra antennal carina not elevated, with area between carina and base of labrum not concave in lateral view, carina not forked beside compound eye (Fig. |
Zygocardiophorus Iablokoff-Khnzorian & Mardjanian, 1981, stat. n. (monotypic, Z. nigratissimus ( |
– | Head with supra antennal carina elevated, with area between carina and base of labrum concave in lateral view, carina forked beside compound eye (Fig. |
Paracardiophorus Schwarz, 1895, part |
19 (17) | Pronotum with dorsal hind angle carinae extending to anterior quarter; dorsal surface of labrum flat in side view; bursa copulatrix with a pair of flexible concave sclerites (Fig. |
Craspedostethus Schwarz, 1898 |
– | Pronotum with dorsal hind angle carina not reaching anterior third (Fig. |
Cardiophorus Eschscholtz, 1829 (paraphyletic) |
20 (4) | Tarsomere 4 lobed or lamellate (Fig. |
21 |
– | No tarsomeres lobed or lamellate, apex of tarsomere 4 vertical | 26 |
21 (20) | Tarsal claws with 2 points per side (Fig. |
22 |
– | Tarsal claws with 1 point per side | 24 |
22 (21) | Scutellum with middle of anterior edge straight (Fig. |
Esthesopus Eschscholtz, 1829 |
– | Scutellum with middle of anterior edge concave: broadly or emarginate (Figs |
23 |
23 (22) | Elytra with apical half of interval 9 flattened or rounded (Australia, Papua New Guinea) | Cardiodontulus Van Zwaluwenburg, 1963 |
– | Apical half of elytral interval 9 costate |
Triplonychoidus |
24 (21) | Mandibles with apices simple and aedeagus with free portion of parameres split vertically into dorsal and ventral lobes (Fig. |
Blaiseus Fleutiaux, 1931, part |
– | Mandibles with 2–3 points; tergite and sternite of male abdominal segment 9 articulated at sides; aedeagus with parameres not split (Fig. |
25 |
25 (24) | Distance between antennae only equal to ¼ width of head (across of compound eyes), nasale facing ventrally; procoxal cavities closed; hind wing notched in anal area (only males known, Brazil) |
Buckelater |
– | Distance between antennae more than ¼ head width; nasale facing anteroventrally; procoxal cavities open; hind wing not notched in anal area (South Africa) | Cardiotarsus part |
26 (20) | Tarsal claws with multiple points per side (including basal tooth); bursa copulatrix with 1 pair of sclerites | 27 |
– | Tarsal claws with one apex per side; bursa copulatrix with 0–4 sclerites | 31 |
27 (26) | Tibiae flattened and broadened apically (Figs |
Globothorax Fleutiaux, 1891 |
– | Tibiae not modified for digging; some with apical half of elytral intervals 1–9 costate (Fig. |
28 |
28 (27) | Tarsal claws with only 2 points per side; bursa with proximal sclerites ovoid or bilobed | 29 |
– | Tarsal claws with more than two points per side; known females with bursa with proximal sclerites elongate, parallel sided | 30 |
29 (28) | Bursa copulatrix with proximal sclerites not bilobed (Fig. |
Horistonotus Candèze, 1860, part (type species keys here) |
– | Bursa copulatrix with proximal sclerites bilobed (Fig. |
Ryukyucardiophorus Ôhira, 1973 |
30 (28) | Tarsal claws with 3 points per side; elytra with apical half of intervals 1–8 costate (Fig. |
Triplonychus Candèze, 1860 |
– | Tarsal claws with 7 or more points per side (Fig. |
Aptopus agrestis (Erichson, 1840, within expanded concept of Horistonotus) |
31 (26) | Scutellum with middle of anterior edge convex (Fig. |
32 |
– | Scutellum with middle of anterior edge emarginate (Fig. |
36 |
32 (31) | Procoxal cavities closed; mesepisternum with projection of anteromesal corners concave mesally; anterior edge of elytra in dorsal view between anterior-most point and humeral angle sinuate or tuberculate (Fig. |
Austrocardiophorus gen. n. |
– | Procoxal cavities open; mesepisternum with anteromesal corners rounded; anterior edge of elytra evenly rounded (Fig. |
33 |
33 (32) | Scutellum with posterior apex bilobed (Fig. |
Floridelater gen. n. |
– | Scutellum with posterior apex not bilobed; prosternum with anterior edge short, exposing labium; pronotum with setae on disc not on tubercles; hind wing with venation well developed, some or all with wing area greater than elytral area | 34 |
34 (33) | Aedeagus with paramere apices forked (Fig. |
Blaiseus Fleutiaux, 1931 (part) |
– | Aedeagus with paramere apices not forked | 35 |
35 (34) | Labrum flat in lateral view; tibiae modified for digging, or not (Fig. |
Aphricus LeConte, 1853 (part) |
– | Labrum convex dorsally in lateral view; tibiae modified for digging (Figs |
Pachyelater Lesne, 1897 |
36 (31) | Prosternum with anterior edge short, exposing labium | 37 |
– | Prosternum with anterior edge not short, produced as lobe, concealing labium when head not extended | 39 |
37 (36) | Head with area between antenna fossa and edge of compound eye unsculptured; hind wing notched in anal area (between AA3+4 and AP); parameres with 2 setae each (central Asia) | Nyctor expallidus Semenov-Tian-Shanskij & Pjatakova, 1936 |
– | Head with area between antenna fossa and compound eye with carina connecting fossa and eye or with 2 pits with non-depressed area between; hind wing notched or not in anal area; parameres of aedeagus each with 3 (Fig. |
38 |
38 (37) | Labrum flat in lateral view; frons without supra-orbital groove; prosternum with anterior edge notched at midline; mesocoxal cavity closed to mesepimeron and mesepisternum by extension of mesosternum; mesotrochantin hidden …. | Aphricus LeConte, 1853 (part) |
– | Labrum convex in side view; frons with supra-orbital groove; prosternum with anterior edge at midline arcuate; mesocoxal cavity open to mesepimeron and mesepisternum (Figs |
Chileaphricus gen. n. |
39 (36) | Mandibles with apices simple, scutellum pointed at posterior apex; urosternites 3–7 with or without serrations along sides (Uzbekistan, monotypic, only males known) | Metacardiophorus Gurjeva, 1966, stat. n. (monotypic, M. sogdianus Gurjeva) |
– | Mandibles with 2 points, scutellum pointed or rounded at posterior apex; urosternites 3–7 without serrations along sides | 40 |
40 (39) | Pronotum with carina along lateral edge reaching more than halfway to anterior edge; bursa copulatrix with paired proximal sclerites partially membranous between spines (Ryukyu Islands, Japan) | Allocardiophorus Ôhira, 1989 |
– | Pronotum with carina along lateral edge restricted to hind angles, or reaching less than halfway to anterior edge; female bursa copulatrix with proximal sclerites solid (Fig. |
41 |
41 (40) | Scutellum with anterolateral edges evenly rounded (Fig. |
42 |
– | Scutellum with anterolateral edges straight or concave posterior to anterolateral corners (Fig. |
43 |
42 (41) | Scutellum with posterior apex evenly rounded (central Asia, only males known) | Neocardiophorus Gurjeva, 1966 |
– | Scutellum with posterior apex pointed; wings incapable of flight (cave inhabiting species, Porto Santo Island, Madeira Archipelago) | Cardiophorus (Coptostethus) Wollaston, 1854 (part). Some Cardiophorus: Coptostethus spp. from the Canary Islands have tarsal claws with 2 apices per side. |
43 (41) | Supra-antennal carina without longitudinal split next to eyes (Fig. |
Cardiophorus (Perrinellus) argentatus Buysson, 1899 (other spp from N. Africa, Ceylon, Central Asia, probably not monophyletic with this). Cardiophorus (Lasiocerus) du Buysson, described from Azerbaijan for a species with long antennae with dense setae, and later synonymised under s.g. Perrinellus was not located for examination, and may not match these key characteristics. |
– | Supra-antennal carina with longitudinal split next to eyes (Fig. |
Paracardiophorus Schwarz, 1895 (part) |
Descriptions of new and redefined genera.
Cardiophorus humeralis Fairmaire & Germain, 1860
Prothorax. Pronotum with carina along lateral edge visible in dorsal view, not reaching anterior edge; procoxal cavities closed. Pterothorax. Scutellum with anterior edge broadly concave. Legs. Tarsi non-lobed and tarsal claws with one apex per side.
Length 3–10 mm. Integument black, brown, or red, some with white, yellow or red markings on elytra or contrasting pronotum and elytra. Head: Antennal sensory elements beginning on antennomere 4; mandibles with apices bidentate or tridentate on each side. Labrum evenly convex; area between antennal fossa and compound eye unsculptured, or with carina connecting them. Frons with supra-antennal carina forked near juncture with compound eye (Fig.
Masculine. Named for a genus of Cardiophorinae known only from the southern hemisphere.
Please see text of discussion above for argumentation for new genus. No unique synapomorphies of this genus were identified. Known from Chile and Australia, 58 spp.
All Australian species are transferred from Paracardiophorus to Austrocardiophorus as: Austrocardiophorus alternatus Carter, 1939, A. amabilis Carter, 1939; A. antennalis Schwarz, 1907; A. assimilis Carter, 1939; A. atronotatus Carter, 1939; A. attenuatipennis Elston, 1930; A. australis (Candèze, 1860, Horistonotus); A. bicolor (Candèze, 1878, Cardiophorus); A. carissimus Carter, 1939; A. compactus (Candèze, 1882, Cardiophorus); A. consobrinus (Candèze, 1878, Horistonotus); A. consputus (Candèze, 1878, Cardiophorus); A. cooki Carter, 1939; A. despectus (Candèze, 1882, Cardiophorus); A. dimidiatus Schwarz, 1902; A. dissimilis Schwarz, 1903; A. divisus (Candèze, 1865, Horistonotus); A. dulcis Carter, 1939; A. elevatus (Van Zwaluwenburg, 1947, Cardiophorus); A. elisus (Candèze, 1865, Horistonotus); A. eucalypti (Blackburn, 1892, Cardiophorus); A. flavipennis (Candèze, 1878, Cardiophorus); A. flavopictus (Carter, 1939, Hypnoidus); A. fulvosignatus (Candèze, 1878, Cardiophorus); A. hamatus (Candèze, 1878, Cardiophorus); A. humilis (Candèze, 1865, Horistonotus); A. jugulus Elston, 1930; A. lenis (Candèze, 1865, Horistonotus); A. litoralis Carter, 1939; A. longicornis (Candèze, 1878, Horistonotus); A. macleayi (Schwarz, 1907, Cardiophorus); A. malkini (Van Zwaluwenburg, 1947, Cardiophorus); A. mastersii (Macleay, 1872, Elater); A. minimus (Candèze, 1878, Cardiophorus); A. mjobergi Elston, 1930; A. moseri Schwarz, 1902; A. nigrosuffusus Carter, 1939; A. occidentalis Carter, 1939; A. octavus (Candèze, 1878, Cardiophorus); A. octosignatus Carter, 1939; A. pallidipennis (Candèze, 1878, Cardiophorus); A. quadripunctatus (Blanchard, 1853, Agriotes); A. quadristellatus Carter, 1939; A. rufopictus Carter, 1939; A. sexnotatus Carter, 1939; A. stellatus Carter, 1939; A. subcruciatus Carter, 1939; A. subfasciatus Carter, 1939; A. tumidithorax (Schwarz, 1907, Cardiophorus); A. vagus Schwarz, 1907; A. varians Carter, 1939; A. variegatus Schwarz, 1902; A. venustus (Candèze, 1860, Cardiophorus); A. victoriensis (Blackburn, 1892, Cardiophorus); A. vittipennis Carter, 1939; A. xanthomus (Candèze, 1865, Horistonotus). The following Chilean species (all) are also transferred from Paracardiophorus to Austrocardiophorus: Paracardiophorus delfini (Fleutiaux, 1907, Cardiophorus); A. elegans (Solier, 1851, Cardiophorus); A. humeralis (Fairmaire & Germain, 1860, Cardiophorus).
Chileaphricus chilensis (Fleutiaux, 1940)
Head. Labrum convex in lateral view. Prothorax. Pronotum with lateral carina absent or restricted to basal ¼ or absent; procoxal cavities open. Pterothorax. Scutellum with middle of anterior edge emarginate, posterior apex not bilobed; mesocoxal cavity open to both mesepimeron and mesepisternum (Fig.
Length 9.5 mm. Integument brown. Head: Antennal sensory elements beginning on antennomere 4 (Fig.
Masculine. Named for a genus of Cardiophorinae known only from Chile.
Please see text of discussion above for argumentation for new genus. The exposed mesotrochantin is unique among Cardiophorinae. Known from: Chile, 1 sp. The type species is transferred from Aphricus to Chileaphricus as: Chileaphricus chilensis (Fleutiaux, 1940)
Cardiophorus musculus Erichson, 1840: 299.
Head. Mandibular apices bidentate; supra antennal carina elevated with area between carina and base of labrum concave in lateral view. Prothorax. Pronotum with lateral carina not reaching anterior edge, hidden in dorsal view by overhanging edge of dorsal part of pronotum (= submarginal line). Pterothorax. Elytra all-black, or with pale markings, area between anterior-most point and humeral angle arcuate or straight in dorsal view (not sinuate) (Fig.
Length 4–8 mm. Integument black, some with pale spots or lines on elytra. Head: Antennal sensory elements beginning on antennomere 3; antennae not reaching pronotal hind angles; mandibles with apices bidentate. Labrum evenly convex; area between antennal fossa and compound eye unsculptured. Frons with supra-antennal carina forked near juncture with compound eye (Fig.
Please see text of discussion above for argumentation for new genus. No unique synapomorphies of this genus were identified. Known from throughout the Holarctic region, 49 spp.
Genus membership revised here to include North American spp. and exclude Australian and South American spp.
The following North American species are transferred from Cardiophorus to Paracardiophorus as: Paracardiophorus abbreviatus Blanchard, 1889; P. acutus Lanchester, 1971; P. amplicollis Motschulsky, 1859; P. aquilis Lanchester, 1971; P. bifasciatus Blanchard, 1889; P. breviatus Lanchester, 1971; P. cardisce (Say, 1839, Elater); P. catskillensis Douglas, 2003; P. columbianus Lanchester, 1971; P. coxalis Blanchard, 1889; P. fenestratus LeConte, 1859; P. gemmifer Blanchard, 1889; P. ignotus Lanchester, 1971; P. kooskooskiensis Lanchester, 1971; P. luridipes Candèze, 1860; P. plebejus Lanchester, 1971; P. propinquus Lanchester, 1971; P. pullus Blanchard, 1889; P. snakensis Lanchester, 1971; P. spurius Lanchester, 1971; P. stigmaticus Candèze, 1869; P. tumidicollis LeConte, 1853.
Coptostethus americanus Horn, 1871.
Pterothorax. Scutellum with anterior edge broadly concave; posterior apex bilobed (Fig.
Length 3.8–4.5 mm. Integument grey-brown to red-brown; setae simple. Head: Antennal sensory elements beginning on antennomere 3 (Fig.
Masculine. Named for a genus of Cardiophorinae known only from southeastern USA.
Please see text of discussion above for argumentation for new genus. The posteriorly bilobed scutellum is unique among Elateridae examined. Known from: USA, coastal dunes by Gulf of Mexico, 1 sp. Often collected by sifting loose sand among dune vegetation.
The type species is transferred from Negastrius to Floridelater as: Floridelater americanus (Horn, 1871, Coptostethus).
Combined diagnosis of Cardiophorinae + Negastriinae
If procoxal cavities not closed to mesepisternum and mesepimeron, then scutellum emarginate anteriorly; hind wing without wedge cell; male aedeagus with paramere bases fused together into tube both dorsally and ventrally, articulated apicad of bases, or rigid (Figs
This template includes much of the described morphological range of genera of Cardiophorinae and outlines variable characters for describing new genera or species of uncertain generic assignment.
Length 3–15 mm. Integument black, brown, yellow and/or red, some with spots or lines on pronotum or elytra; setae simple. Body without concavities for reception of tarsi. Head: Antennae weakly serrate; antennal sensory elements beginning on antennomere 3 or 4; mandibles with apices unidentate to tridentate. Labrum flat or evenly convex; area between each antennal fossa and adjacent compound eye unsculptured, with carina joining fossa and eye, or with groove or pit(s). Frons convex; with supra-antennal carinae complete across frontoclypeal region (Fig.
If procoxal cavities not closed to mesepisternum and mesepimeron, then scutellum emarginate anteriorly. Prosternum with sides near midlength straight or concave, (pronotum and prosternum also not fused). Hind wing without wedge cell; all setae evenly tapered; males with aedeagus with paramere bases fused together into tube both dorsally and ventrally, articulated apicad of bases or rigid (Figs
Diagnoses presented here distinguish each genus from all other Cardiophorinae. Diagnoses for newly described and redescribed genera are provided above with corresponding genus description.
Allocardiophorus
Head. Mandibles bidentate. Prothorax. Pronotum with lateral carina reaching more than halfway to anterior edge, not hidden by lateral expansion of pronotum in dorsal view; prosternum with anterior edge produced as lobe, concealing labium when head not extended; procoxal cavities open. Pterothorax. Scutellum with anterior edge emarginate (Fig.
Aphricus LeConte, 1853: 501. Type species: A. californicus LeConte, 1853: 502.
Patriciella Van Zwaluwenburg, 1953: 20. Syn. n.
Replacement name for Patricia Van Zwaluwenburg.
Patricia
Van Zwaluwenburg, 1947: 113. Type species: P. australica
Head. Labrum flat in lateral view. Prothorax. Pronotum with lateral carina absent or restricted to basal ¼; procoxal cavities open. Pterothorax. Scutellum with middle of anterior edge convex to broadly concave posterior apex not bilobed. Legs. Tarsomeres without ventral lobes; tarsal claws with one apex per side. Aedeagus. Aedeagus with paramere apices not forked (Fig.
Aptopus Eschscholtz, 1829: 32. Type species: A. tibialis Eschscholtz, 1829: 32.
Definition restricted here to exclude species near Aptopus agrestis (Erichson).
Prothorax. Pronotum with lateral carina not reaching anterior edge, hidden in dorsal view by overhanging edge of dorsal part of pronotum (= submarginal line). Legs. Tarsi without apically extending lobes or pads, tarsal claws with 5 or more points per side. Also. Bursa copulatrix with both proximal (Fig.
Blaiseus
Fleutiaux, 1931: 307. Type species: B. bedeli
Mandibles simple; aedeagus with parameres split vertically into dorsal and ventral lobes (Fig.
Buckelater Costa, 1973: 33. Type species: B. argutus Costa, 1973: 35.
Distance between antennae equal to only ¼ width of head (measured across outsides of compound eyes), nasale oriented ventrally (Fig.
Cardiodontulus Van Zwaluwenburg, 1963: 341. Type species: C. brandti Van Zwaluwenburg, 1963: 341.
Prothorax. Pronotum with lateral carina extending anterad from hind angles following lateral edge, not reaching anterior edge; procoxal cavities closed. Pterothorax. Scutellum with middle of anterior edge concave: broadly or abruptly emarginate. Legs. Tarsomere 4 lobed or lamellate; Tarsal claws with 2 points per side (Fig.
Cardiophorellus Cobos, 1970a: 222. Type species: C. gracilicornis Cobos, 1970a: 223.
Head. Mandibular apex unidentate (simple). Prothorax. Pronotum with lateral carina present but hidden in dorsal view by swollen pronotum (= submarginal line). Pterothorax. Scutellum with middle of anterior edge broadly concave. Also: Head with supra-orbital groove present; posterior edges of hypomeron mesad of hind angles with rectangular or semicircular indentations; prosternum with anterior edge short, exposing labium; tarsomere 4 without ventral lobe or pad extending beyond base of tarsomere 5; tarsal claws with 1 point per side; tibiae with posterior surfaces convex, weakly modified for digging (Figs
Cardiophorellus : subgenus Parapleonomus Cobos, 1970a: 222. Type species: C. inermis Cobos, 1970a: 222.
Diagnosis inferred from Cobos, 1970a. Insufficient information available for inclusion in key to genera.
Head. Mandibular apex bidentate or multidentate; supra–orbital groove present. Prothorax. Pronotum without lateral carina, or apparently hind-angle carina; prosternum with anterior edge short, exposing labium. Legs. Tarsomere 4 without ventral lobe or pad extending beyond base of tarsomere 5; tarsal claws with 1 point per side; tibiae with posterior surfaces convex, modified for digging. Aedeagus. Parameres with apices undivided. Females undescribed or unassociated. Known from: South Africa, Gauteng Province, 1 sp.
Cardiophorus Eschscholtz, 1829: 34. Type species: Elater thoracicus Fabricius, 1801: 236, now a synonym of Cardiophorus gramineus (Scopoli, 1763).
Caloderus Stephens, 1830: 269. Type species: Elater thoracicus Fabricius, 1801: 236.
Melanotus Gistel, 1834: 11, not Eschscholtz, 1829: 32. Type species Elater thoracicus Fabricius, 1801: 236, designated by Sánchez (1996) [not in References].
Paradicronychus Dolin and Gurjeva, 1975: 116 (nomen nudum) —Here placed as syn. n. of Cardiophorus.
Head. Mandibles with two or three apices. Prothorax. Pronotum with lateral carina not reaching anterior third (Fig.
Coptostethus Wollaston, 1854: 238. Type species: C. femoratus Wollaston, 1854: 240.
Head. Mandibles with 2 points. Prothorax. Pronotum with carina along lateral edge hidden or not in dorsal view; restricted to hind angles, or reaching less than halfway to anterior edge. Prosternum with anterior edge not short, produced as lobe, concealing labium when head not extended. Pterothorax. Scutellum with middle of anterior edge abruptly emarginate, anterolateral edges evenly rounded, and posterior apex pointed. Legs. No tarsomeres lobed or lamellate; tarsal claws with one apex per side. Also. Pterothorax. Brachypterous. Abdomen. Urosternites 3–7 without serrations along sides.
Porto Santo Island, Madeira Archipelago, from a cave, females unknown. The relatedness of this species to brachypterous Cardiophorinae from the Canary Archipelago. and South Africa also placed in subgenus Coptostethus has not been demonstrated. Some Coptostethus spp. from the Canary Archipelago have tarsal claws with 2 apices (Fig.
Cardiophorus : subgenus Perrinellus Buysson, 1899: 282. Type species: Athous argentatus Abeille de Perrin, 1894: 92.
Cardiophorus : subgenus Lasiocerus Buysson, 1912: 129. Type species: C. schusteri Buysson, 1912: 128.
Head. Mandibles with 2 apices; supra-antennal carina without longitudinal split next to compound eyes. Prothorax. Pronotum with lateral carina reaching less than halfway to anterior edge (in type species not distinguishable from the dorsal hind angle carina), not hidden by lateral expansion of pronotum in dorsal view; prosternum with anterior edge produced as lobe, concealing labium when head not extended. Pterothorax. Scutellum with anterior edge emarginate, anterolateral edges broadened posterior to anterolateral corners, posterior end pointed. Legs. tarsi without ventral lobes and tarsal claws with one apex per side. Also. Pronotum with hind angles not truncate dorsally; procoxal cavities open. Aedeagus with parameres approximately cylindrical pre-apically (Fig.
Israel, other spp from North Africa, Ceylon, Central Asia, probably not monophyletic, 12 spp. Cardiophorus (Lasiocerus) du Buysson was not located for examination, and may not match these key characteristics. Subgenus Lasiocerus was described from Azerbaijan, and distinguished by its long antennae with dense setae.
Cardiotarsus Eschscholtz, 1836: published in identification table opposite p.5, without associated spp. Type species: C. capensis Candèze, 1860: 226.
Pronotum. Lateral carina not reaching anterior edge (Fig.
Craspedostethus Schwarz, 1898b: 414. Replacement name for Craspedonotus Schwarz.
Craspedonotus Schwarz, 1898a: 148. Type species: C. rufiventris Schwarz, 1898a: 148.
Head. Mandibular apex bidentate or tridentate. Prothorax. Pronotum with lateral carina reaching ¾ of distance to anterior edge, hidden in dorsal view by overhanging edge of dorsal part of pronotum (Fig.
Dicronychus
Brullé, 1832: 138. Type species: Elater obesus
Paramecus Dillwyn, 1829: 32. Type species: Paramecus cordiger Dillwyn, 1829 (= Elater equiseti Herbst, 1784: 114.
Prothorax. Pronotum with lateral carina not reaching anterior edge, hidden in dorsal view by overhanging edge of dorsal part of pronotum (= submarginal line); procoxal cavities open. Legs. Tarsomere 4 without ventral lobe or pad extending beyond base of tarsomere 5; tarsal claws with second apex at base on each side (Fig.
Diocarphus Fleutiaux, 1947a: 364. stat. n., raised to genus rank.
Phorocardius : subgenus Diocarphus Fleutiaux, 1947a: 364. Type species: P. solitarius Fleutiaux, 1931: 309.
Head. Head with single pit between antennal fossa and eye. Prothorax. Pronotum with lateral carina not reaching anterior edge, hidden in dorsal view by overhanging edge of dorsal part of pronotum (= submarginal line); procoxal cavities closed. Pterothorax. Anterior edge of scutellum emarginate. Legs. Tarsomere 4 without ventral lobe or pad extending beyond base of tarsomere 5; tarsal claws with two apices per side.
Also. Legs. tarsal claws with ventral surface concave mesad of basal apex (Fig.
Displatynychus Ôhira, 1987: 92.
Platynychus : subgenus Displatynychus Ôhira, 1987: 92. Type species: Cardiophorus adjutor Candèze, 1875: 17.
Head. Area between antennal fossa and compound eye with carina connecting fossa and eye, or with 2 pits with non-depressed area between them. Prothorax. Pronotum with lateral carina not reaching anterior edge, hidden in dorsal view by overhanging edge of dorsal part of pronotum (= submarginal line); procoxal cavities closed. Pterothorax. Anterior edge of scutellum emarginate. Legs. Tarsomere 4 without ventral lobe or pad extending beyond base of tarsomere 5; tarsal claws with two apices per side. Also. Legs. Tarsal claws with ventral surface convex mesad of basal apex. Bursa Copulatrix. Proximal (largest) sclerites ovoid; paired distal sclerites absent (i.e. a second pair, farther from vagina, at base of spermathecal gland duct); base of spermathecal gland duct with tube-like sclerotisation (Fig.
Esthesopus Eschscholtz, 1829: 32. Type species: E. castaneus Eschscholtz, 1829: 32.
Prothorax. Pronotum with carina extending anterad from hind angles following lateral edge (ie, not below lateral edge of prothorax, but in some species not distinguishable from the dorsal hind angle carina), not reaching anterior edge. Pterothorax. Scutellum with middle of anterior edge straight (Fig.
Globothorax Fleutiaux, 1891: ccxxxii. Type species: G. chevrolati Fleutiaux, 1891: ccxxxiii.
Teslasena Fleutiaux, 1892: 410. Syn. n. Type species, Anelastes femoralis Lucas, 1857: 71.
Legs. Tibiae flattened and broadened apically (Figs
Horistonotus Candèze, 1860: 243. Type species: H. flavidus Candèze, 1860: 250.
Prothorax. Pronotum with carina extending anterad from hind angles following lateral edge (ie, not below lateral edge of prothorax, but in some species not distinguishable from the dorsal hind angle carina), not reaching anterior edge. Legs. Tibiae not modified for digging; tarsi with tarsomere 4 not lobed or lamellate, tarsal claws with 2 or 7 points per side (Fig.
Margogastrius Schwarz, 1903b: 80. Replacement name for Gastrimargus Schwarz, 1902.
Head. Mandibular apex unidentate (simple); supra-orbital groove present. Prothorax. Pronotum with lateral carina not reaching anterior edge, hidden in dorsal view by overhanging edge of dorsal part of pronotum (= submarginal line). Pterothorax. Scutellum with middle of anterior edge abruptly emarginate. Legs. Tarsal claws with one point per side. Also. Posterior edges of hypomeron mesad of hind angles with rectangular or semicircular indentations; prosternum with anterior edge not short, produced as lobe, concealing labium when head retracted; tibiae with posterior surfaces flattened and broadened apically (Fig.
Metacardiophorus Gurjeva, 1966: 91. stat. n., raised to genus rank.
Cardiophorus : subgenus Metacardiophorus Gurjeva, 1966: 91. Type species: C. sogdianus Gurjeva, 1966: 91.
Head. Mandibles with apices unidentate. Prothorax. Pronotum with lateral carina not reaching anterior edge, extending anterad from hind angles following lateral edge (Fig.
Mionelater Becker, 1963: 125. Type species: M. planatus Becker, 1963: 126.
Head. Supra antennal carina porrect in lateral view; eyes large; antennae serrate with acute apicoventral angles. Prothorax. Hind angles elongate, pronotum abruptly constricted at base. Pterothorax. Mesocoxal cavity possibly open to mesepisternum and mesepimeron. Legs. No tarsomeres lobed or lamellate; tarsal claws with one apex per side; tarsal claws with one apex per side. Not in key to species.
Comment: the serrate antennae, large eyes, shelf-like supra-antennal carina, elongate pronotal hind angles, and open mesocoxal cavities suggest this genus might belong to Dendrometrinae or another subfamily. One known species.
Neocardiophorus Gurjeva, 1966: 95. Type species: N. mamajevi Gurjeva, 1966: 95.
Head. Mandibles with 2 points. Prothorax. Pronotum with carina along lateral edge not hidden in dorsal view, and restricted to hind angles, or not reaching more than halfway to anterior edge. Prosternum with anterior edge not short, produced as lobe, concealing labium when head not extended. Pterothorax. Scutellum with middle of anterior edge abruptly emarginate, anterolateral edges evenly rounded, and posterior apex evenly rounded. Legs. No tarsomeres lobed or lamellate; tarsal claws with one apex per side; tarsal claws with one apex per side. Also. Pterothorax. Abdomen. Urosternites 3–7 without serrations along sides. Two species, known from males from Turkmenistan and Uzbekistan.
Nyctor Semenov-Tian-Shanskij & Pjatakova, 1936: 101 N. expallidus Semenov-Tian-Shanskij & Pjatakova, 1936: 102.
Head. Head with area between antenna fossa and compound eye unsculptured. Prothorax. Pronotum with lateral carina extending anterad from hind angles following lateral edge, not reaching anterior edge; prosternum with anterior edge short, exposing labium. Pterothorax. Scutellum with middle of anterior edge abruptly emarginate. Legs. No tarsomeres lobed or lamellate; tarsal claws with one apex per side; tarsal claws with one apex per side. Also. Hind wing. Membrane notched in anal area (between AA3+4 and AP). Not reduced in female. Female genitalia. Proximal (largest) sclerites of bursa copulatrix ovoid; distal sclerites absent. Male genitalia. Parameres with 2 setae each. Known from Tajikistan, Turkmenistan, and Uzbekistan, 2 spp.
Odontocardus Fleutiaux, 1931: 332. Type species: Cardiotarsus vitalisi Fleutiaux, 1918b: 231.
Prothorax. Pronotum with lateral carina not reaching anterior edge, hidden in dorsal view by overhanging edge of dorsal part of pronotum (= submarginal line). Legs. Tarsomere 4 with ventral lobe or pad extending beyond base of tarsomere 5; tarsal claws with both basal and apical points on each side. Also. Bursa copulatrix with paired proximal (largest) sclerites of bursa copulatrix ovoid. Known from Cambodia, Vietnam, Laos, Philippines, 6 spp.
Pachyelater Lesne, 1897b: 117 Replacement name for Parelater Lesne, 1897a. Transferred here to Cardiophorinae.
Lesnelater
Fleutiaux, 1935a: 116. Type species Lesnelater madagascariensis
Parelater
Head. Mandibles bidentate; labrum convex in lateral view. Prothorax. Lateral carina not reaching anterior edge, not hidden in dorsal view; procoxal cavities open; prosternum with anterior edge short, exposing labium; pronotum with setae on disc not on tubercles. Mesothorax. Scutellum with anterior edge broadly concave, posterior end not bilobed; mesepisternum with anteromesal corners rounded; anterior edge of elytra rounded or straight in dorsal view. Legs. Protibiae flattened and broadened apparently for digging (Figs
Platynychus : subgenus Paraplatynychus Fleutiaux, 1931: 315. Type species: Paraplatynychus mixtus Fleutiaux, 1931: 326.
Prothorax. Pronotum with complete carina at lateral edge (Fig.
Phorocardius Fleutiaux, 1931: 308. Type species: Cardiophorus florentini Fleutiaux, 1895a: 687.
Prothorax. Pronotum with lateral carina not reaching anterior edge (Fig.
Platynychus Motschulsky, 1858: 58. Type species: P. indicus Motschulsky, 1858: 59.
Head. Head with area between antenna fossa and compound eye unsculptured. Prothorax. Pronotum with lateral carina not reaching anterior edge (Fig.
Ryukyucardiophorus
Ôhira, 1973a: 32. Type species: Paracardiophorus loochooensis,
Pronotum with carina extending anterad from hind angles following lateral edge (ie, not below lateral edge of prothorax, but in some species not distinguishable from the dorsal hind angle carina), not reaching anterior edge. Legs. Protibiae not modified for digging; tarsi with tarsomere 4 not lobed or lamellate, claws with 2 points per side. Bursa copulatrix with paired proximal (largest) sclerites bilobed with attached semi-sclerotised membrane with spines (Fig.
Triplonychoidus Schwarz, 1906: 181. Type species: Triplonychus trivittatus Champion, 1895: 427.
Prothorax. Pronotum with lateral carina extending anterad from hind angles following lateral edge, not reaching anterior edge; procoxal cavities open. Pterothorax. Scutellum with middle of anterior edge concave: (broadly or abruptly emarginate). Legs. Tarsomere 4 lobed or lamellate; tarsal claws with two apices per side. Also. Apical half of elytral interval 9 costate. Known from Mexico to South America, females unknown, 2 spp.
Triplonychus Candèze, 1860: 236. Type species: T. acuminatus Candèze, 1860: 238.
Prothorax. Pronotum with carina extending anterad from hind angles following lateral edge (ie, not below lateral edge of prothorax, but in some species not distinguishable from the dorsal hind angle carina), not reaching anterior edge. Legs. Protibiae not modified for digging; tarsi with tarsomere 4 not lobed or lamellate, claws with 3 points per side. Also. Head with area between antenna fossa and compound eye with either carina connecting fossa and eye, or with 2 pits with non-depressed area between. Elytra with apical half of intervals 1–8 costate (Fig.
Tropidiplus Fleutiaux, 1903: 251. Type species: T. tellinii Fleutiaux, 1903: 251.
Prothorax. Pronotum with complete lateral carina (Fig.
Zygocardiophorus Iablokoff-Khnzorian & Mardjanian, 1981: 247. Type species, C. nigratissimus Buysson, 1891: 134. Stat. n. raised to genus rank.
Head. Mandibular apices tridentate; supra antennal carina low with area between carina and base of labrum not concave in lateral view, carina not forked beside compound eye (Fig.
1–2 Frontoclypeal area Cardiophorus Scale bar = 0.5 mm). 1 C. convexus (Say), (anteroventral view) 2 C. gramineus, (dorsal view). CE = compound eye; SC = supra-antennal carina; SG = supra-orbital groove. Figures captions include post-revision names 3–4 Latero-ventral view of hypomeron Scale bar = 1 mm) 3 Cardiophorus gagates Erichson 4 Paracardiophorus propinquus Lanchester. Ant = anterior; LC = lateral carina 5 Lateral view of prosternal process of Cardiophorus erythropus Erichson Scale bar = 0.5 mm). A length from procoxa to dorsal apex; Ant = Anterior B length from procoxa to end at halfway between dorsal and ventral apices C length from procoxa to ventral apex D vertical distance between dorsal and ventral apices. Dorsal and ventral apices are considered points where profile of respective surface is 45º from horizontal 6–9 Dorsal view of scutellum of Cardiophorinae Scale bar = 0.5 mm) 6 Esthesopus castaneus 7 Blaiseus bedeli 8 Cardiophorus gramineus 9 Floridelater americanus 10 Latero-ventral view of mesepimeron and mesepisternum of Cardiophorus fenestratus (LeConte) showing measurement of angle (a) of anterolateral corner of mesepisternum Scale bar = 0.5 mm). MST = mesepisternum; MRN = mesepimeron 11–13 Ventral view of left side of mesocoxal cavity of Elateridae (After
14 Lateral view of metasternum of Zygocardiophorus nigratissimus Buysson Scale bar = 0.5 mm). LC = lateral carina; MTP = metepisternum; MTS = metasternum15–16 Dorsal view of anterior end of left elytron of Cardiophorinae Scale bar = 0.5 mm). 15 Cardiophorus gramineus 16 Paracardiophorus cardisce (Say) 17–18 Hind wing of Elateroidea Scale bar = 1 mm) 17 Anelastes druryi (Kirby) 18 Blaiseus bedeli. Vein names follow
23–25 Dorsal view of aedeagus of Elateridae Scale bar = 0.5 mm). 23 Hypnoidus riparius Fabricius, 1792 [not Cardiophorinae] 24 Paracardiophorus cardisce 25 Blaiseus bedeli 28–31 Proximal sclerites of the bursa copulatrix of Cardiophorinae, interior view (top end of sclerite extends furthest into bursa) Scale bar = 0.5 mm). 28 Paracardiophorus cardisce 29 Paraplatynychus mixtus 30 Globothorax chevrolati 31 Esthesopus parcus Horn 32–33 Distal sclerites of the bursa copulatrix of Cardiophorinae, interior view Scale bar = 0.5 mm). 32 Cardiophorus inflatus 33 Cardiophorus gramineus 34–35 Sclerotised base of spermathecal gland duct of Cardiophorinae, lateral view Scale bar = 0.5 mm). 34 Paracardiophorus cardisce 35 Paracardiophorus musculus. SD = spermathecal gland duct. Captions reflect the revised classification.
Inferred phylogeny of Cardiophorinae and Negastriinae based on 159 adult morphological characters. Tree is a 50% majority-rule phylogram with branch lengths estimated by MrBayes, of 120000 post-burnin trees. Model = Mkv+G. Values are posterior probabilities. Scale bar indicates 0.1 changes per site. Several nodes are labelled a-l to simplify discussion. Taxon labels D, P, and N are members of Dendrometrinae, Physodactylinae and Negastriinae within the Cardiophorine. Genus name “C.” indicates Cardiophorus.Names are pre-revision. Type species of genera are marked with “*”.
Inferred phylogeny of Cardiophorinae and Negastriinae based on 139 parsimony-informative adult morphological characters (left), with Bayesian topology (adapted from Fig.
38–41 Chileaphricus chilensis lectotype male 38, 39 pterothorax 40 head and prothorax 41 aedeagus 42–44 Margogastrius schneideri paralectotype female. 42, 43 habitus 44 spermatheca, lateral view 45–47 Floridelater americanus. 45 aedeagus 46, 47 male 48–49 Blaiseus bedeli lectotype male. Scale bars: 1 mm, 0.5 mm for detail photos. Captions reflect the revised classification.
50–53 Pachyelater madagascariensis 50 female 51, 52 male 53 aedeagus 54–56 Aphricus sp. 54 aedeagus 55, 56 male 57–58 Neocardiophorus mamajevi. 57 male 58 aedeagus 59–61 Nyctor expallidus. 59 aedeagus 60, 61 male 62 Metacardiophorus sogdianus, male paratype. Scale bars: 1 mm for habiti, 0.5 mm for detail photos. Captions reflect the revised classification.
63–64 Metacardiophorus sogdianus, paratype 63 male 64 aedeagus 65–68. Paracardiophorus musculus. 65 aedeagus 66 proximal sclerite 67, 68 female 69–73. Zygocardiophorus nigratissimus. 69, 70 male 71 elytral apex 72 proximal sclerite 73 aedeagus 74–76 Platynychus indicus, female lectotype 74–75 adult 76 distal and proximal sclerites of bursa copulatrix, lateral view 77, 78 Globothorax chevrolati lectotype female. Scale bars: 1 mm for full habiti, 0.5 mm for detail photos. Captions reflect the revised classification.
79, 80 Globothorax femoralis lectotype male 81–84 Cardiophorus gramineus 81, 82 female 83 bursa copulatrix with distal and proximal sclerites, lateral view 84 aedeagus 85–87 Cardiophorellus gracilicornis paratype. 85, 86 male 87 aedeagus 88–89 Dicronychus cinereus female. Scale bars: 1 mm for habiti, 0.5 mm for detail photos. Captions reflect the revised classification.
90–92 Dicronychus cinereus. 90 proximal sclerite 91 distal sclerites 92 aedeagus 93–97 Aptopus rugiceps 93, 94 adult 95 proximal sclerite 96 distal sclerite 97 aedeagus 98–99Cardiophorus (Perinellus) argentatus, Lectotype. 98 male 99 aedeagus 100Cardiophorus (Coptostethus) femoratus, Lectotype 101–103 Phorocardius florentini 101, 102 Lectotype 103 bursal sclerites 104 Tropidiplus tellinii, female. Scale bars: 1 mm for habiti, 0.5 mm for detail photos. Captions reflect the revised classification.
105–108 Tropidiplus tellinii 105 female 106 tarsal claw with basal seta, lateral view (Lectotype male) 107 distal & proximal sclerites, internal view 108 aedeagus (Lectotype male) 109–112 Displatynychus adjutor 109, 110 female 111 distal sclerite 112 aedeagus. 113–115 Diocarphus solitarius. 113 distal and proximal sclerites in bursa copulatrix, internal view 114, 115 female 116–118 Cardiotarsus capensis 116, 117 male 118 bursal sclerites lectotype 119–121 Odontocardus vitalisi 119 distal sclerites of bursa copulatrix, internal view 120, 121 male. Scale bars: 1 mm for habiti, 0.5 mm for detail photos, 0.1 mm for tarsal claw. Captions reflect the revised classification.
122 Odontocardus vitalisi. Aedeagus 123–125 Triplonychoidus trivittatus paralectotype. 123 aedeagus 124, 125 male 126–131 Esthesopus castaneus 126, 127 male 128 ventral view of tarsomeres 4&5 129 tarsal claw 130 proximal sclerite of bursa copulatrix 131 aedeagus 132–135 Aptopus agrestis 132, 133 male 134 tarsal claw 135 aedeagus 136–139 Horistonotus flavidus 136, 137 lectotype female 138 proximal sclerite of lectotype 139 aedeagus Scale bars: 1 mm for habiti, 0.5 mm for detail photos, 0.1 mm for tarsal claw. Captions reflect the revised classification.
140–143 Paraplatynychus mixtus 140, 141 male 142 proximal sclerite 143 aedeagus 144–146 Triplonychus acuminatus, Lectotyp. 144, 145 male 146 aedeagus 147–150 Cardiodontulus brandti, male paratype 147 aedeagus 148, 149 adult 150 tarsal claw 151–153 Craspedostethus rufiventris 151 female Lectotype 152 lateral view of pronotal hind angle of female Lectotype 153 aedeagus 154 Craspedostethus culcarius , bursal sclerites of female labeled as type. Scale bars: 1 mm for habiti, 0.5 mm for detail photos, 0.25 for 142, 143. Captions reflect the revised classification.
155–157 Austrocardiophorus humeralis. 155 adult 156 proximal sclerite 157 aedeagus 158–160 Buckelater argutus 158 male 159 anterior view of head capsule of male 160 aedeagus 161–162 Ryukyucardiophorus loochooensis 161 female 162 proximal sclerite. Scale bars: 1 mm for habiti, 0.5 mm for detail photo. Captions reflect the revised classification.
A complete bibliographic synonymy is presented here with references in chronological order to accurately document the nomenclatural history of the group through 2015. All family-group names in synonymy under Cardiophorinae. Several references were unavailable for examination, as is noted in the text. The synonymy began with a draft catalog provided by Prof. Paul Johnson (South Dakota State University, USA) and Schenkling’s most recent (1925) world catalog for all historically and currently recognized cardiophorine names. The Genera Insectorum (
The following faunal studies and lists were examined to find taxonomic changes within the Cardiophorinae since
Cardiophorinae Candèze, 1859.
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Aptopina Jakobson, 1913: 760. Type genus: Aptopus Eschscholtz, 1829. Synonymy verified by inclusion of Cardiophorina in synonymy under Aptopina.
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Esthesopinae Fleutiaux, 1919: 76 (incorrectly includes senior synonym, Cardiophorites Candèze, in synonymy). Type genus: Esthesopus Eschscholtz, 1829.
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Dicronychidaesensu Chûjô & Ôhira 1965: 28 (not Fleutiaux). Elevation, in error of Dicronychinae
Cardiophorini Candèze, 1859.
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Nyctorini Semenov-Tian-Shanskij & Pjatakova, 1936: 101. Syn. n. Type genus: Nyctor Semenov-Tian-Shanskij & Pjatakova, 1936. Described in Elaterinae
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—Stibick 1979: 181 (as valid without explanation)
Allocardiophorus Ôhira, 1989: 79 (species included: monotypic). Type species: Paracardiophorus nigroapicalis Miwa, 1927: 109. Fixed by original designation.
Aphricus LeConte, 1853: 501. Type species: A. californicus LeConte, 1853: 502. Type species fixed by original monotypy. Described in Cebrionites.
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Patriciella Van Zwaluwenburg, 1953: 20. syn. n. (Replacement name for Patricia Van Zwaluwenburg).
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Patricia Van Zwaluwenburg, 1947: 113 (species included: monotypic). Type species: P. australica
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Aptopus Eschscholtz, 1829: 32 (species included: ephippiger, tibialis). Type species: A. tibialis Eschscholtz, 1829: 32. Type species fixed by
—Dejean 1836: 99 (catalog)
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Eniconyx Horn, 1884: 51. Type species: Eniconyx pullatus Horn, 1884: 52 Type species fixed by
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Austrocardiophorus gen. n. Type species Cardiophorus humeralis Fairmaire & Germain, 1860: 5.
Blaiseus Fleutiaux, 1931: 307. Type species: B. bedeli Fleutiaux, 1931: 308. Type species fixed by original monotypy.
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Buckelater Costa, 1973: 33. Type species: B. argutus Costa, 1973: 35 (species included: monotypic). Type species fixed by original designation.
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Caloderus Stephens, 1830, see: Cardiophorus Eschscholtz
Cardiodontulus Van Zwaluwenburg, 1963: 341. Type species: C. brandti Van Zwaluwenburg, 1963: 341 (species included: monotypic). Type species fixed by original designation.
—Cardiotarsus mjobergi Carter is here transferred to this genus as Cardiodontulus mjobergi (Carter, 1939)
Cardiophorellus Cobos, 1970a: 222. Type species: C. gracilicornis
Cardiophorellus: subgenus Parapleonomus Cobos, 1970a: 222 (species included: monotypic). Type species: C. inermis Cobos, 1970a: 222. Type species fixed by original designation.
Cardiophorus Eschscholtz, 1829: 34 (species included: biguttatus, charactericus, discicollis, ebininus, equisiti, exaratus, latiusculus, luzonicus, ornatus, ruficollis, rufipes, suturalis, thoracicus). Type species: Elater thoracicus
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—Platia and Gudenzi 2009: 121 (spp.nov, Turkey, Russia)
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—Platia 2014: 72 (sp.n. Yemen)
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Caloderus Stephens, 1830: 269 (species included: equisiti, ruficollis, thoracicus). Type species: Elater thoracicus Fabricius, 1801: 236. Type species fixed by
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Paradicronychus Dolin & Gurjeva, 1975: 116 (nomen nudum, species included inflatus, nothus). No type species designated for this name, published after 1930, therefore name is not available.
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—Syn. n. of Cardiophorus here.
Cardiophorus: subgenus Coptostethus Wollaston, 1854: 238. Rank lowered to subgenus by
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Coptostethus Wollaston, 1854: 238. Type species: C. femoratus Wollaston, 1854: 240. Type species fixed by original monotypy.
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Cardiophorus: subgenus Perrinellus Buysson, 1899: 282 (species included: argentatus, bonnairei, bousaadensis). Type species: Athous argentatus
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Cardiophorus: subgenus Lasiocerus Buysson, 1912: 129. Type species: C. schusteri
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Cardiotarsus Eschscholtz, 1836: published in identification table opposite p.5, without associated spp. Type species: C. capensis Candèze, 1860: 226. Type species fixed by
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—Cardiotarsus mjobergi Carter is here recommended for transfer to Cardiodontulus Van Zwaluwenburg.
Chileaphricus gen. n. Type species, Aphricus chilensis Fleutiaux, 1940: 103 (species included: monotypic).
Coptostethus Wollaston, 1854, see: Cardiophorus: subgenus Coptostethus Wollaston
Craspedonotus Schwarz, 1898, see: Craspedostethus Schwarz
Craspedostethus Schwarz, 1898b: 414. Replacement name for Craspedonotus Schwarz, 1898.
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Craspedonotus Schwarz, 1898a: 148 (species included: rufiventris, semirufus, minutissimus). Type species: C. rufiventris Schwarz, 1898a: 148. Type species fixed by
Dicronychus Brullé, 1832: 138 (species included: obesus, messenicus). Type species: Elater obesus Brullé, 1832: 138, now referred to by replacement name Dicronychus brullei Platia & Gudenzi, 2003. Type species fixed by
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—Al Dhafer and Platia 2013: 19 (spp.n. Saudi Arabia)
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—Platia 2014: 72 (sp.n. Yemen)
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Eudicronychus Méquignon sensu Chûjô & Ôhira, 1965: 28. Error verified by inclusion of spp. assigned to Dicronychus Brullé.
Gauroderus Thomson, 1859: 104 (species included: monotypic). Type species: Elater cinereus Herbst, 1784: 114. Type species fixed by original designation.
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Paramecus Dillwyn, 1829: 32. Type species: Paramecus cordiger Dillwyn, 1829 (= Elater equiseti Herbst, 1784: 114). Type species fixed by original monotypy. Objective junior generic synonym of Dicronychus.
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Diocarphus Fleutiaux, 1947a: 364. stat. n., raised to genus rank. Phorocardius: subgenus Diocarphus Fleutiaux, 1947a: 364. Type species: P. solitarius Fleutiaux, 1931: 309. Type species fixed by original monotypy.
Displatynychus Ôhira, 1987: 92.
Platynychus: subgenus Displatynychus Ôhira, 1987: 92 (species included: monotypic). Type species: Cardiophorus adjutor Candèze, 1875: 17. Type species fixed by original designation.
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Eniconyx
Esthesopus Eschscholtz, 1829: 32. Type species: E. castaneus Eschscholtz, 1829: 32. Type species fixed by original monotypy.
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Floridelater gen. n. Type species Coptostethus americanus Horn, 1871: 303 (species included: monotypic).
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Gauroderus Thomson, 1859, see: Dicronychus Brullé
Gastrimargus Schwarz, 1902, see: Margogastrius Schwarz
Globothorax Fleutiaux, 1891: ccxxxii. Type species: G. chevrolati Fleutiaux, 1891: ccxxxiii. Type species fixed by original monotypy.
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Teslasena Fleutiaux, 1892: 410. Syn. n. Type species, Anelastes femoralis Lucas, 1857: 71, fixed by original monotypy. Described in Elaterinae: Physodactylini.
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Horistonotus Candèze, 1860: 243 (39 species included). Type species: H. flavidus Candèze, 1860: 250. Type species fixed by
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Lasiocerus Buysson, 1912, see: Cardiophorus: subgenus Lasiocerus Buysson
Lesnelater Fleutiaux, 1935, see: Pachyelater Lesne
Margogastrius Schwarz, 1903b: 80. Replacement name for Gastrimargus Schwarz. Transferred here to Cardiophorinae.
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Gastrimargus Schwarz, 1902: 309. Type species: G. schneideri Schwarz, 1902: 310. Type species fixed by original monotypy. Name preoccupied by Gastrimargus Spix, 1823.
Metacardiophorus Gurjeva, 1966: 91. stat. n., raised to genus rank. Cardiophorus: subgenus Metacardiophorus Gurjeva, 1966: 91. Type species: C. sogdianus Gurjeva, 1966: 91 (species included: ineptus, sogdianus). Type species fixed by original designation.
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Mionelater
Neocardiophorus Gurjeva, 1966: 95 (species included: fausti, mamajevi). Type species: N. mamajevi Gurjeva, 1966: 95. Type species fixed by original designation.
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Nyctor Semenov-Tian-Shanskij & Pjatakova, 1936: 101 (species included: monotypic). Type species: N. expallidus Semenov-Tian-Shanskij & Pjatakova, 1936: 102. Type species fixed by original designation.
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Odontocardus Fleutiaux, 1931: 332 (species included: lateralis, vitalisi, harmandi). Type species: Cardiotarsus vitalisi Fleutiaux, 1918b: 231. Type species fixed by original designation.
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Pachyelater Lesne, 1897b: 117 Replacement name for Parelater Lesne.
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Lesnelater Fleutiaux, 1935a: 116 (species included: madagascariensis, dubius, singularis, unicus). Described in Physodactylinae from males assigned to Pachyelater by
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Parelater
Paracardiophorus Schwarz, 1895b: 40 (species included: musculus, sequens, pullatus, granarius, erythrurus, subaeneus, fuscipennis, humeralis, australis, longicornis). Type species: Cardiophorus musculus Erichson, 1840: 299. Type species fixed by
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Paradicronychus Dolin & Gurjeva, 1975, see: Cardiophorus Eschscholtz
Paramecus Dillwyn, 1829, see: Dicronychus Brullé
Paraplatynychus Fleutiaux, 1931: 315.
Platynychus: subgenus Paraplatynychus Fleutiaux, 1931: 315 (species included: mixtus, costatus, fouqueti, incostatus). Type species: Platynychus mixtus Fleutiaux, 1931: 326. Type species fixed by original designation.
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Parelater Lesne, see: Pachyelater Lesne
Patriciella Van Zwaluwenburg, see: Aphricus LeConte
Patricia Van Zwaluwenburg, see: Aphricus LeConte
Perrinellus Buysson, see: Cardiophorus: subgenus Perrinellus Buysson
Phorocardius Fleutiaux, 1931: 308 (species included: astutus, bifidus, florentini, magnus, melanopterus, solitarius, unguicularis,). Type species: Cardiophorus florentini Fleutiaux, 1895a: 687. Type species fixed by original designation.
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—Platia 2015: 184 (sp.n. Maldives)
Platynychus Motschulsky, 1858: 58 (species included: pictus, curiatus, mauritanicus, indicus, nebulosus, crucifer, axillaris). Type species: P. indicus Motschulsky, 1858: 59. Type species fixed by
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Ryukyucardiophorus Ôhira, 1973a: 32 (species included: monotypic). Type species: Paracardiophorus loochooensis Miwa, 1934: 255. Type species fixed by original designation.
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—Platia 2015: 182 (sp.n. Maldives)
Teslasena Fleutiaux, see: Globothorax Fleutiaux
Triplonychoidus Schwarz, 1906: 181 (species included: trivittatus, parvulus). Type species: Triplonychus trivittatus Champion, 1895: 427. Type species fixed by
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Triplonychus Candèze, 1860: 236 (species included: ephippiger, acuminatus, longicollis, cayennensis, ventralis, plagiatus, lebasii, costatus, rufus, debilis). Type species: T. acuminatus Candèze, 1860: 238. Type species fixed by
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Tropidiplus Fleutiaux, 1903: 251. Type species: T. tellinii Fleutiaux, 1903: 251. Type species fixed by original monotypy.
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Zygocardiophorus Iablokoff-Khnzorian & Mardjanian, 1981: 247 (species included: nigratissimus, alienus). Stat. n. raised to genus rank. Type species, C. nigratissimus Buysson, 1891: 134, type species fixed by original designation. Described as subgenus of Cardiophorus.
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I thank Professor Stewart Peck, for his mentorship, and support. Thanks to R. Anderson (CMNC), E.T. Arias (EMEC), M. Barclay (NHM), P. Bouchard (CNCI), Y. Bousquet (CNCI), R. Brett (CASC), R. Brooks (SEMC), D. Drugmand (ISNB), M. Fitton (LSUK), C. Girard (MNHN), L. Herman (AMNH), E.R. Hoebeke (CUIC), M. Kerley (NHM), A. Konstantinov (USNM), B. Korotyaev (ZMAS), S. Laplante (CNCI), R. Leschen (NZAC), S.A. Marshall (DEBU), H. Mendel (NHM), J. Mertlik (WWW.elateridae.com), N. Nikitsky (ZMUM), J. Nunn (JNIC), G. Parsons (MSUC), P. Perkins (MCZC), G. Prinsloo (SANC), A. Rasnitsyn (PIN), E.G. Riley (TAMU), A. Samuelson (BPBM), P. Skelly (FSCA), M. Uhlig (ZMHB), N. Vandenberg (USNM), L. Vilhelmsen (ZMUC), and L. Zerche (DEIC) for specimen loans and help finding literature. I thank R. Anderson, E. Becker, A. Brunke, F. Chapleau, J. Cook, E. Jendek, P. Johnson, J. Kukalova-Peck, A.B.T. Smith, M.L. Smith, L. Packer, D. Sikes, J. Skevington for systematics advice. Thanks to K. Savard for help with figures. I thank E. Fuller, D. Ahrens, and an anonymous reviewer for detailed reviews of the manuscript. Most of this research was done at Carleton University. Financial support for this study and publication costs were provided by a Natural Sciences and Engineering Research Council of Canada (NSERC) Discovery Grant to S. Peck, and an NSERC postgraduate scholarship, two years of Ontario Graduate Scholarships and a Scholarship from the Canadian Museum of Nature to H. Douglas. Special thanks to my wife Daphne Uras.
Specimens coded and identification information sources used for species used in phylogenetic analysis. Entries include information in the following sequence: species name, photo codes of specimens coded (
Outgroups
Adrastus pallens (Fabricius, 1792), M: P134, F: P135, lit.:
Agriotes sputator (Linnaeus, 1758), T069 (T070 to T072 not conspecific) M: P2C2; F: P2B6, P2C1, type species (Elaterinae: Agriotini)
Agrypnus murinus (Linnaeus, 1758), M: P072; F: P073, lit.:
Athous vittatus (Fabricius, 1792), M: P104, P105; F: P106, P107, lit.:
Berninelsonius hyperboreus (Gyllenhal, 1827), M: P203; F: P204, lit.:
Elater ferrugineus Linnaeus, 1758, M: P046; F: P047, T066, T067, lit.:
Hypnoidus riparius (Fabricius, 1792), M: P133; F: P131, P132, lit.:
Tropihypnus bimargo Reitter, 1896, M: P2D2; F: P2D1, lit.:
Physodactylinae
Margogastrius schneideri Schwarz, 1902, F: T058, T059, det.: none, type species
Pachyelater madagascariensis (Lesne, 1897), M: T024; F: T026, T027, lit.:
Teslasena femoralis Lucas, 1857, M: P061, T039, lit.:
Negastriinae
Agrypnella eburnea Champion, 1895, M: T008, P025, det.: E.C. Becker 1985, type species
Agrypnella squamifer Candèze, 1895, F: P029, det.: none
Arhaphes diptychus Candèze, 1860, M: P015; F: P016, det.: E. Candèze, type species,
Cardiohypnus mirabilis (Candèze, 1860), M: P028a, c; F: P028b, T009, lit:
Fleutiauxellus maritimus (Curtis, 1840), M: P185; F: P186, lit.:
Migiwa curatus Candèze, 1875, M: P179; F: P180, lit.: Kishii 1956, det.: unknown, non-type species
Monadicus sp., M: P181; F: P182, lit.:
Negastrius americanus (Horn, 1871), M: P157, F: P156, T095, det.: none, non-type species
Negastrius pulchellus Fabricius, 1801, M: P045; F: P013, P014, lit.:
Neoarhaphes americanus Champion, 1895, M: P187; F: P188, lit.:
Neohypdonus gentilis (LeConte, 1866), M: P177; F: P178, lit.:
Oedostethus femoralis LeConte, 1853, M: T016, P012; F: P010, lit.:
Paradonus pectoralis (Say, 1836), M: P175; F: P176, lit.:
Pronegastrius humeralis (Candèze, 1863), F: P183, P184, lit.:
Quasimus minutissimus Germar, 1817, M: P017; F: P018, T081, lit.:
Rivulicola variegatus (Macleay, 1872), M: P021a, b; F: P022, det.: Anonymous, type species
Yukoana carinicollis Lewis, 1894, M: P191; F: P192, lit.:
Zorochros demustoides Herbst, 1806, M: P2C4; F: P2C3, T082, lit.:
Cardiophorinae
Aphricus californicus LeConte, 1853, M: P034, T020, lit.:
Aphricus chilensis Fleutiaux, 1940, M: T036; F: none, det.: none, non-type species
Aptopus agrestis Erichson, 1840, M: P142, P143; F: none, det.: none, non-type species
Aptopus pullatus (Horn, 1884), M: P7A1–3; F: P036, T017, T018, lit.:
Blaiseus bedeli Fleutiaux, 1931, M: T034; F: none, det.: none, type species
Blaiseus nothoafricanus Douglas, 2009, M: P170, P171, lit.:
Buckelater argutus Costa, 1973, M: P098–102, T045– T049; F: none, det.: none, type species
Cardiodontulus brandti Van Zwaluwenburg, 1963, M: T001, T002; F: none, det.: none, type species
Cardiophorellus gracilicornis Cobos, 1970, M: T028, T029, det.: none, type species
Cardiophorus (Coptostethus) brunneipennis Wollaston, 1864, M: P155; F: P154, lit.:
Cardiophorus (Metacardiophorus) sogdianus Gurjeva, 1966, M: T056; F: none, det.: none, type species
Cardiophorus (Perrinellus) angustatus Fleutiaux, 1933, M: P059, P060; F: none, T038; F: none, det.: none, non-type species
Cardiophorus (Perrinellus) argentatus Abeille de Perrin, 1894, M: P069, T043, lit.:
Cardiophorus (Zygocardiophorus) nigratissimus Buysson, 1891, M: P149, P151; F: P150, P152, lit.:
Cardiophorus brevis Candèze, 1887, M: P168; F: P169, lit.: Champion 1895, det.: none, non-type species
Cardiophorus cardisce (Say, 1834), M: P6A5, P2A12, P2C6, P2B1, P158, P159; F: P5A2, P5A3, FAB8, T074, lit.:
Cardiophorus convexulus LeConte, 1853, M: P167, P1H3; F: P1H1, P3H3, T062, lit.:
Cardiophorus convexus (Say, 1823), M: P1A5, P1C10, P3B6, PBA1, PBA2, F: P1F4, P1F6, P4A3, P5D1, PBA3, PBA4, T063, lit.:
Cardiophorus gramineus (Scopoli, 1763), M: PBB1; F: P020, P023, P141, T012, T013, lit.:
Cardiophorus inflatus Candèze, 1882, F: T005; M: none, det.: none, non-type species
Cardiophorus luridipes Candèze, 1860, M: P165; F: P164, T065, lit:
Cardiophorus sp. undescribed, Congo, M: PBC4; F: C5, lit.:
Cardiophorus togatus Horn, 1871, M: P163, P1G9, P1G10; F: P5B2, T064, lit.:
Cardiotarsus capensis Candèze, 1860, P144–146 & 2M &3F, T075, det.: none, type species
Cardiotarsus mjobergi Carter, 1939, M: P172; F: P173, det.: none, non-type species
Cardiotarsus sp., M: HDCU222, HDCU223
Craspedostethus rufiventris Schwarz, 1898, M: P053; F: T060, T061, det.: none, type species
Dicronychus cinereus (Herbst, 1784), M: PBB2; F: P153, T080, lit.:
Esthesopus castaneus Eschscholtz, 1829, M: P049; F: P050, P051, T076, lit.:
Esthesopus parcus Horn, 1884, M: P7B2, P7B4; F: P7B3, P116, P117, T090, det.: none, non-type species
Globothorax chevrolati Fleutiaux, 1891, F: T037; M: none, det.: none, type species
Horistonotus flavidus Candèze, 1860, M: P097; F: T050 (=P095), T051 (=P096), det.: none, type species
Horistonotus simplex LeConte, 1863, M: P7A5, P7A6; F: P7A4, P7B1, P160, T088, lit.:
Neocardiophorus mamajevi
Nyctor expallidus Semenov-Tian-Shanskij & Pjatakova, 1936, M: T052, T053; F: T054, det.: none, type species
Odontocardus vitalisi Fleutiaux, 1918, M: P067; F: P068, T042, lit.:
Paracardiophorus humeralis (Fairmaire & Germain, 1860), M: PBD2; F: PBD3, P190, HDCU215, det.: Candèze, von Hayek, non-type species
Paracardiophorus musculus (Erichson, 1840), F: PBB3; M: PBB4, T077–T079, lit.:
Paracardiophorus subcruciatus Carter, 1939, M: PBA5; F: PBA6, P189, type not found, det.: none, non-type species
Patriciella australica Van Zwaluwenburg, 1947, M: T044, det.: none, type species
Phorocardius florentini Fleutiaux, 1895a, M: P056; F: P057; sex?: T035, lit.:
Phorocardius solitarius Fleutiaux, 1931a, F: T033, lit.:
Platynychus adjutor Candèze, 1875, M: P003, F: P002, T093, lit.:
Platynychus indicus Motschulsky, 1858, F: T041; M: none, det.: none, type species
Paraplatynychus mixtus Fleutiaux, 1931a, M: P065, P066; F: P064, T094, lit.:
Ryukyucardiophorus loochooensis (Miwa, 1934), M: P272, P273; F: P271, det.:
sp. undescribed, New Zealand, M: P005, lit.:
Triplonychoidus trivittatus (Champion, 1895), M: T010, T011, det.: none, type species
Triplonychus plagiatus (Erichson, 1840), M: P148 & 6 others; F: P147, T083,
Tropidiplus tellinii Fleutiaux, 1903, M: T040; F: P063, det.: von Hayek 1965, type species
Label data from non-type specimens of species coded for phylogenetic analyses, including information in the following order: scientific name and author, photo code number, identification code number from voucher label (begins with HDCU), label data, the sex of the specimen (M/F), and the coden representing the specimen depository. Text from multiple labels, beginning with uppermost label, are listed in a separate set of quotation marks, separated by semicolons. Text from each line of a label with multiple lines of text is separated by a “/”. Notes about the appearance of the label appear in brackets before quotation.
Adrastus pallens (Fabricius), P134, HDCU138, “Germany/ Holstein, Plön/ 26. Vii. 1964/ G. G. E. Scudder”, [partly handwritten] “Adrastus/ pallens F. 1967/ Wellschmeid det.”, M, CNCI; P135, HDCU139, [partly handwritten] “22.VII.1924 Finl./ Kuolemajärvi/ M.Ivaschinzeff”, [partly handwritten] “Adrastus F#/ pallens Fabr./ Dr. Wellschmeid det. 1962”, F, CNCI
Agriotes sputator (Linnaeus), P2C2 “Sydney, N.S./ VI–7–1965/ W.J.Brown”, M, CNCI; P2B6 “Sydney, N.S./ VI–7–1965/ W.J.Brown”, F, CNCI; P2C1 “North Sydney/ N.S. VI–19–1965/ W.J. Brown”, F, CNCI
Agrypnella eburnea Champion, P025, HDCU026, “Santarem, Para/Brazil IV–27–/1963 F.Werner”; [partly handwritten] “Agrypnella/ nr. eburnea/ MCZ has 3/ Det. E.C. Becker 1985” [partly handwritten] “Agrypnella/ eburnea/ BMNH –Homotype/ Det.HDouglas 2004” M, CNCI
Agrypnella squamifer Candèze, P029, HDCU033, [blue disc, handwritten] “Santar/ em”; [handwritten] “squamifer/ Cdz./ (named from type).”; [partly handwritten] “mentioned/ B.C.A.III.(1)415”, F, NHM
Agrypnus murinus Linnaeus, P072, HDCU076, [Partly handwritten] “Suisse-Lausanne/ Vidy/ 6 VI 42/ J. Aubert”, M, CNCI; P073, HDCU077, “France, Landes/ Parentis/ VI. 1977/ Vuillaume”, F, CNCI
Aphricus californicus LeConte, P034, HDCU038, “Congress Jct./ Ar. 7–28–33/ R.H.Beamer”; [partly handwritten] “Aphricus/ californicus/ LeC./ Det.HDouglas 2004”, M, CNCI
Aptopus agrestis Erichson, P142, HDCU146, “Faz.Aceiro/ Jatai, Goiás-Brasil/ X. 1962/ Exp.Dep.Zool.”, M, MZSP; P143, HDCU147, “Faz.Aceiro/ Jatai, Goiás-Brasil/ X. 1962/ Exp.Dep.Zool.”, M, MZSP
Aptopus pullatus (Horn), P036, HDCU040, “Baboquivari, Mts./ Ariz. VII–24–41/ E.L. Todd”; [partly handwritten] “Aptopus ♀?/ pullatus (Horn)/ Det.H
Arhaphes diptychus Candèze, P015, HDCU015, “Coll. R.I.Sc.N.B/ Inde/ Tenasserim/ Thagata/ Fea. Apr. 1887”; “Collection/ E. Candèze”; [handwritten] “Arrhaphes/ diptychus Cd. Dét. E. Candèze”, F, ISNB; P016, HDCU016, same data, M, ISNB
Athous argentatus Perrin, P069, HDCU073, [male symbol]; [handwritten] “Jerusalem/ D David”; MUSEUM PARIS/ Collection Léon Fairmaire/ 1906”, M, MNHN
Athous vittatus (Fabricius), P104, HDCU108, [partly handwritten] “CZECHOSLOVAKIA/ Mor. Bor. 24.v.1963/ Moravicany/ leg.V.Korbel”, M, CNCI; P105, HDCU109, same data, M, CNCI; P106, HDCU110, same data, F, CNCI; P107, HDCU111, same data, F, CNCI
Cardiophorinae sp., P005, HDCU005, “Rika Valley/ Clarence River/KA 12 Mar 69/ R.J.B. Power” “Swept grass Coprosoma” “Brounaeolus Hyslop” “N Z Arthropod/ Collection, NZAC/ Private Bag 92170/ AUCKLAND/ New Zealand” “nr Brounaeolus/ det. R.A. Leschen”, M, NZAC
Berninelsonius hyperboreus Gyllenhal, P203, HDCU207, “Y.T., British Mts./ “June Cr.”, 320m/ 3km NW Firth R./ 69°13’ , 140°05’/ J.M.Campbell”; “25.VI.1984/ 84–32 , misc./ beetles ex/ around camp”, M, CNCI; P204, HDCU208, same data, “29.VI.1984/ 84–37 , sifting/ Salix & Alnus/ litter”, F, CNCI
Cardiohypnus mirabilis Candèze, P028, HDCU029, 6 specimens on a pin labelled 6X “Haldwani Div./ Kumaon,/ India. H.G.C.”; 1X “H.G.Champion Coll./ B.M. 1953–156.”, now 4 on pin and 4 labels, NHM; P028a, HDCU030, same data, M, NHM; P028b, HDCU031, same data, F, NHM; P028c, HDCU032, top specimen of remaining 4 on pin, M, NHM
Cardiophorus brevis Champion, P168, HDCU172, “Mex: Oaxaca/ 20 mi. W. Pinotepa/ Nacional/VI.23.1983, M, CNCI; P169, HDCU173, “Mex: Oaxaca/ 20 mi. W. Pinotepa/ Nacional/VI.23.1983, F, CNCI
Cardiophorus cardisce (Say), P158, HDCU162, “Canada, ON, Lanark Co./ Almonte, on sand 16.V.04/ Hdouglas, Mlarivee, M?, CNCI; P159, HDCU163, “Canada, QC, Co. Pontiac, Bristol Mines, 45°30’ 00” N 76°21’00”W, 20.IV.03 SandDune/Pit, on sand , 11 AM HDouglas, Mlarivee”, M, CNCI
Cardiophorus convexulus LeConte, P167, HDCU171, [handwritten] “ON, Sandbanks/ PP. VI.16.2000/ Beating, H.Douglas”; Cardiophorus/ convexulus LeC./ Det.Hdouglas 2002”, M, CNCI
Cardiophorus (Coptostethus) brunneipennis Wollaston, P154, HDCU158, [Spain, handwritten] “IS. TENERIFE/ MASS. TENO/ Mt. DE L’AGUA/ 5.2.1991 m.800/ I. Gudenzi”; “COPTOSTETHUS/ brunneipennis Wol/ det. Platia 2002”, F, CNCI; P155, HDCU159, same data, M, CNCI
Cardiophorus gramineus Scopoli, P020, HDCU020, “Schleissheim/ 28.5.04”; München/ k.Ku zer”; [handwritten] “Cardiophorus/ gramineus Scop.”, F, CNCI; P023, HDCU024, [handwritten] “ROMA: E.U.R./ 16.V.’58.PARENTI”, F, CNCI; P141, HDCU145, [handwritten partly illegible] “Cerva U________”; [green square]; “BROOKLYN/ MUSEUM/ COLL. 1929”; USNM 2036286”, F, USNM
Cardiophorus luridipes Candèze, P164, HDCU168, “CALIFORNIA:/ Riverside Co.,/ 16 May 1924/ W. Benedict”, F, CNCI; P165, HDCU169, “CALIFORNIA:/ Riverside Co.,/ 16 May 1924/ W. Benedict”; [partly handwritten] “Cardiophorus/ luridipes/ Cand./ Det.H
Cardiophorus nigratissimus Buysson, P149, HDCU153, “Turkey-Gaziantep/ 18 km NW Kilis/ Kara Deresi river”; “m 420, 30.V.2002/ C.Giusto-S.Zoia”; “CARDIOPHORUS/ nigratissimus Buys./ det Platia 002”, M, CNCI; P150, HDCU154, same data, F, CNCI; P151, HDCU155, same data, M, CNCI; P152, HDCU156, same data, F, CNCI
Cardiophorus sp. P166, HDCU170, “Rep. Of Congo/Voka/V.1978 G. Onore”, F, CNCI
Cardiophorus togatus Horn, P163, HDCU167, “MEXICO: Nuevo Leon/ 37 km Linares, 1545 m/ 20 March 1991, Brooks/ Leschen. #29. Ex: beating”; [Partly Handwritten] “Cardiophorus/ togatus/ Horn/ Det.Hdouglas 2005”, M, SEMC
Cardiophorus (Perinellus) angustatus Fleutiaux, P059, HDCU063, “MAHATSINJO/ près Tananarive”, M, MNHN; P060, HDCU064, “MAHATSINJO/ près Tananarive”, M, MNHN
Cardiotarsus capensis Candèze, P144, HDCU148, “Winburg, OFS./ Aug., 1918./C.W. Mally./SN. 2543.”; [partly hadwritten] “ac.c.2450/ Ag Dp S Afr”; “NATIONAL COLL./ OF INSECTS/ Pretoria, S.Afr” M, SANC; P145, HDCU149, same data, M, SANC; P146, HDCU150, same data, F, SANC
Cardiotarsus mjobergi Carter, P172, HDCU176, “Australia, Qld. Mt./ Glorious 27°19'S,/ 152° 45'E, 1–5.XII.1997/ T. Hiller Malaise”, M, CNCI; HDCU177, same data, F, CNCI
Cardiotarsus sp. (nr. philautus), HDCU222 “Pietermaritzburg,/ Natal, S. Afr./ 23.XI.–5.XII.70/ H. & M. Townes”, F, CNCI; HDCU223, same data, M, CNCI
Craspedostethus rufiventris Schwarz, P053, HDCU057, “Kamerun/ Conradt”; “Coll. Kraatz”, M, Eberswalde; P054, HDCU058, “Kamerun/ Conradt”; “Coll. Kraatz”, M, Eberswalde
Dicronychus cinereus (Herbst), P153, HDCU157, [Italy, partly handwritten] “RAVENNA/ P. S. VITALE/ 16vi84/ LEG. CALLEGARI”; “DICRONYCHUS/ cinereus (Hbst.)/ det. Platia 002”, f, CNCI
Elater ferrugineus (Linnaeus), P046, HDCU050, unlabelled, M, CNCI; P047, HDCU051, [handwritten] “L. ferugi/ neus. 20.6/ Tero. Ital”; “343”, F, CNCI
Esthesopus castaneus Eschscholtz, P049, HDCU053, [handwritten] “Brazil”; “Sammlung/ CI Müller”; [partly handwritten] “Esthesopus/ castaneus Esch./ E.C. Becker 1959”, M, CNCI; P050, HDCU054, [handwritten] “Fry/ Rio. Jan/”; [partly handwritten]; “Fry Coll./ 1905. 100.”; “Esthesopus/ castaneus Er/ C.M.F. von Hayek det. 1985”; [handwritten] “Mandibles open”, F, NHM; P051, HDCU055, [handwritten] “Bahia”; “Coll. Janson./ Ex Mniszech.”; [handwritten] “Esthesopus/ castanueus Esch./ Cand. Cdz./ ex. Coll. De Mniszech.””, F, NHM
Esthesopus parcus Horn, P116, HDCU120, “AZ, Santa Cruz Co./ Peña Blanca Campgr./ 6 Aug 2002/ H. Douglas BL”, F, CNCI; P117, P161, HDCU165, same data, F, CNCI; P162, HDCU166, same data, M, CNCI; HDCU121, “AZ, Pima Co., Green Valley, on tree bark, 860m, 5 AUG/ 2002 H.Douglas”, ?, CNCI;
Fleutiauxellus maritimus (Candèze), P185, HDCU189, [partly handwritten] “Grünwald/ 23.5.49.”; “München/ Bühlmann”; [handwritten] “Hypnoidus/ maritimus Cand.”, M, CNCI; P186, HDCU190, Zakarpatny,/ Skope/ Pes Na Gorah / vii. 1966”; [partly handwritten] “Negastrius/ maritimus (Cand.)/ W. Dolin det. 1969”, F, CNCI
Globothorax chevrolati Fleutiaux, P058, HDCU062, “261”; [handwritten] “Brésil”; “Collection Chevrolat”; F, MNHN
Horistonotus flavidus Candèze P097, HDCU101, “Poá/ SP Brasil/ 10.XII.1966/ E.X.Rabello Col.”, M, MZSP;
Horistonotus simplex LeConte P160, HDCU164, “AZ Santa Cruz Co./ Pena Blanca Campgr./ 6 Aug 2002/ H.Douglas BL”, F, CNCI;
Hypnoidus riparius Fabricius P131, HDCU135, [partly handwritten]”Jilmeica/Boh. 16.V.48/A. Olexa”; [handwritten] “Hypnoidus/ riparius/ F./ Det. Olexa ‘73”, F, CNCI; P132, HDCU136, [partly handwritten] “Le Sancy/ P. Ole D./ 1.VII.72/ G. Minet”, F, CNCI; P133, HDCU137, [partly handwritten] “Le Sancy/ P. Ole D./ 1.VII.72/ G. Minet”, ?, CNCI
Migiwa curatus (Candèze), P179, HDCU183, [partly handwritten] “Mt. Fuji. JAPAN/ 3.VIII.1964/ M. Suda leg.”; [handwritten] “Negastrius/ curatus/ [name also written in Japanese characters], M, CNCI; P180, HDCU184, “Mt.Koya, Nara-ken/ Japan 17–VII–55/ S. Mizobe”, F, CNCI
Monadicus sp., P181, HDCU185, “Argentina, Prov.Bs.As/ Bs.As., San Isidro/ 10–15.I.1982/ H & A Howden”, M, CNCI; P182, HDCU186, F, same data
Negastrius americanus (Horn), P156, HDCU160, “USA, FL, St. Theresa,/ 7.V.2000, sifting sand &/ grass, sheltered beach,/ H.Douglas”, F, CNCI; P157, HDCU161, same data, M, CNCI
Negastrius pulchellus (Linnaeus), P013, HDCU013, “Jarov/ Boh.v.54./A.Olexa” [handwritten] “Hypnoidus/ pulchellus/ L./ Det.Olexa 73”, F, CNCI; P014, HDCU014, “589”; “Fennia/ Ik Ollila/ 12/6 1932/ K Lahtivirta” [handwritten] “Hypnoidus/ pulchellus L.” F, CNCI; P045, HDCU049, [handwritten] “Europe”, M, CNCI
Neoarhaphes americanus (Champion), P187, HDCU191, “Paraiso CZ/ PanApr 22 11/ EASchwarz”, M, CNCI; P188, HDCU192, “COSTA RICA. Punt./ 3 km N. Guacimal/ 800m 26.V.1979/ H & A Howden”, F, CNCI
Neohypdonus gentilis (LeConte), P177, HDCU181, “Saskatoon/ Saskatchewan/ VI–22–1974/ E.J.Kitely”, M, CNCI; P178, HDCU182, same data, F, CNCI
Odontocardus vitalisi Fleutiaux, P067, HDCU071, [partly handwritten] “MUSEUM PARIS/ Laos/ ou Cambodge/ Bonette 1911”, M, MNHN; P068, HDCU072, same data, “Cardiotarsus/ Vitalisi/ Fleut./ FLEUTIAUX det.”; F, MNHN
Oedostethus femoralis LeConte, P010, HDCU010, “CANADA Québec/ Brome Co.; Glen-/ Sutton 24.VII.1985/ Larochelle, Larivière”; [handwritten] “Oedostethus/ femoralis LeC./ Dét.: P. Bélanger, 1987”, F, CNCI; P011, HDCU011, same data, F, CNCI; P012, HDCU012, “CANADA Québec/ Brome Co.; Glen-/ Sutton 24.VII.1985/ Larochelle, Larivière”, M, CNCI
Paracardiophorus humeralis Fairmaire & Germain, P190, HDCU194, [handwritten] “CACHAGUA/ W. Aconcagua/ 18–IX–1974/ Col.P.VIDAL”, F, CNCI; TQ 1, HDCU215, [purple, partly handwritten] “Coll. R.I.Sc.N.B/ santiago/ ex. Coll. Mnizech” “Collection/ E. Candèze”; “(Bitactus cdz.)/ Humeralis/ Frm./ Chili. C. mn”; [partly handwritten] “Cardiophorus humeralis Fairm./ dét. E. Candèze”; [partly handwritten] “Paraplatynychus/ C.M.F. von Hayek/ det. 1960”; [handwritten] “cardiophorus/ humeralis/ no6i5”, F, ISNB
Paracardiophorus subcruciatus Carter, P189, HDCU193, “Australia, Qld. Mt./ Glorious 27°19'S,/ 152° 45'E, 1–5.XII.1997/ T. Hiller Malaise”, F, CNCI
Paradonus pectoralis (Say), P174, HDCU178, “USA, NY, Essex Co./ Crown Pt, splashing gravel beach under L. Champlain/ bridge, 19.VII.03 Hdouglas”, F, CNCI; P175, HDCU179, same data, M, CNCI; P176, HDCU180, “Canada, ON, Carleton,/ Crown Pt. 45°31’00”N–/ 76°08’00”W, 8.VI.03,/ sand/rock beach hand/ coll. J. Holland Leg.”, F, CNCI
Phorocardius florentini Fleutiaux, P056, HDCU060, “Tonkin/ Florentin”; “MUSÉUM PARIS/ Coll. E. FLEUTIAUX”, M, MNHN; P057, HDCU061, same data, F, MNHN
Platynychus adjutor Candèze, P001, HDCU001, [handwritten] “KU.79” “Japan./G.Lewis./1910–320., M, BM(NH); P002, HDCU002, on Card, under card [difficult to read] “Nowaba/ 6.80”; “DATA/ under card” Japan./ G. Lewis/ 1910–320.”, F, BM(NH); P003, HDCU003, “Yoyogi/ (Tokyo)/ 9–vi–1953/ A.Kato”; “9”; “Adjutor Cand/ ex Ôhira”, M, BM(NH)
Paraplatynychus mixtus Fleutiaux, P064, HDCU068, “MUSEUM PARIS/ HAUT-TONKIN/ CAO BANG/ CHRISSEMENT VILLAIN/ 1925”; [partly handwritten] “mixtus fleut./ FLEUTIAUX det.”; “paraplatynychus/ C.M.F. von Hayek/ det. 1965”, F, MNHN; P065, HDCU069, “Hué”; [handwritten] “Fl 200 mx”, M, MNHN; P066, HDCU070, “Hué”; [handwritten], M, MNHN
Pronegastrius humeralis (Candèze), P183, HDCU187, “Mimune(Mie)/ 1955–VI.18–21/ H.Ôhira.Coll”, F, CNCI; P184, HDCU188, same data; [handwritten] “Negastrius/ (Pronegastrius)/ humeralis/ Cand.”, F, CNCI
Quasimus minutissimus (Germar), P017, HDCU017, “Zbraslav/ Boh. C. VI53./ A. Olexa”; [handwritten] “Quasimus/ minutissimus/ Germ./ Det. Olexa ‘73”, M, CNCI; P018, HDCU018, same data, F, CNCI
Rivulicola variegatus Macleay, P021a, HDCU021, Specimen on left: [partly handwritten] “N.S. Wales/ Minto (W.W.F.)/ on/ apple trunk/ 16–VI–1903”, M, NHM; P021b, HDCU022, Specimen on right, same data, M, NHM; P022, HDCU023, [handwritten] “Kuranda/ 26–6–38/ C.G.C.”; [handwritten] “kuranda”; “A.E. Clarke/ Collection./ B.M.1957–24”; “♀ genitalia in/ water soluble resin/ Dimethyl Hydrantoin/ Formaldyhyde)”; [handwritten] “abdomen missing 19 III 85.”, F, NHM; P024, HDCU025, same data [handwritten] “Cryptohypnus/ variegatus Macl”, F, NHM
Ryukyucardiophorus loochooensis (Miwa), P271, “S. Ryukyu Is.:/ Ishigaki I./ Omoto-dake, 100-250m, 22.v.1964”; “Malaise Trap/ J.L. Gressitt”; “Paracardiophorus/ loochooensis/ Det H.
Teslasena femoralis Fleutiaux, P061, HDCU065, “MUSEUM PARIS/ GOYAZ à CUYUBA/ DE CASTELNAU 6–47”; [white disc, handwritten] “6/47”; [partly handwritten] “Teslasena/ FLEUTIAUX det.”, M, MNHN
Triplonychus plagiatus Erichson, P147, HDCU151, [handwritten] “35935”; [handwritten] “Brasil/ Puru”;”Fry Coll./ 1905.100.”; [handwritten] “35935=/ Brazil Para/ Higgins” [partly handwritten] “Triplonychus/ plagiatus/ Er. Det.Hdouglas 2005”F, NHM; P148, HDCU152, “Coll./ Janson.”; “Ega,/ Amazons./ Bates.” [partly handwritten] “Triplonychus/ plagiatus/ Er. Det.Hdouglas 2005”M, NHM
Tropidiplus tellinii Fleutiaux, [handwritten] P063, HDCU067, “Bir Donan/ 20.X.49”; [partly handwritten] “Tropidiplus/ telinii Fleut/ C.M.F.von Hayek det/ 1965”.F, MNHN
Yukoana carinicollis (Lewis), P191, HDCU195, [handwritten] “Mt. Mikuni/ 14–V–1966/ K.Baba”; [partly handwritten] “Yukoana/ carinicollis/ (Lewis, 1894)/ H.Ôhira Det. 1969”, M, CNCI; P192, HDCU196, [handwritten] “Kurokawa/ N. Echigo/ 21.–V, 1967/ K. Baba”; [handwritten] “Y. carinicollis/ Det. Ôhira 1967”, F, CNCI
Label data from type specimens of species coded for phylogenetic analyses. Entries include information in the following order: Type code (number attached to photos), scientific name, kind of type specimen (A = allotype, H = holotype, L = lectotype, N = neotype, P = paratype, PL = paralectotype, S = syntype, T = type of unknown kind), label data and any lectotype designation (text from multiple labels, beginning with uppermost label, are listed in a separate set of quotation marks, separated by semi-colons. Text from each line of a label with multiple lines of text is separated by a “/”. Notes about the appearance of a label appear in square brackets before quotation), the sex of the specimen (M/F), and the coden representing the specimen depository.
T001, Cardiodontulus brandti Van Zwaluwenburg, H, “NEW GUINEA: PAPUA/ Kiunga, Fly River/ IX–10–17–1957”; “Wm. W. Brandt/ Collector”; “♂”; [red] “Holotype”; “Cardiodontulus/ brandti/ ♂/ Van Zwal./ Holotype/ No.”; [pink] “HT–6047/ BISHOP MUSEUM”, M, BPBM
T002, Cardiodontulus brandti Van Zwaluwenburg, P, [genitalia in microvial, and left wing and urosternites 3–7 mounted on a card] “NEW GUINEA: PAPUA/ Kiunga, Fly River/ IX–10–17–1957”; “Wm. W. Brandt/ Collector”; “♂”; [yellow] “Paratype”; “Cardiodontulus/ brandti/ ♂/ Van Zwal.”, M, BPBM
T005, Cardiophorus inflatus Candèze, L, Lectotype designated here. [handwritten] “Mantchouria”; “Collection/ E. Candèze”; [handwritten] “n.sp./ inflatus/ Cdz/ Mandchuria”; [partly handwritten] “Cardiophorus/ inflatus cd./ dét. E. Candèze”; “♀”; [partly handwritten] “Holotype. C. inflatus/ C.M.F.von Hayek. Cand/ det., 1957”; [red] “Holotype”; and with the author’s red designation label “LECTOTYPE/ Cardiophorus/ inflatus/ Candèze desig./ Douglas, 2006” [with ♀ genitalia on slide on pin]. Label mailed separately to ISNB in 2006., F, ISNB
T008, Agrypnella eburnea Champion, H, [circular with red margin] “TYPE”; [handwritten] “Santarem”; “Coll./ Janson”; “Janson coll./ 1903–130.”; [handwritten] “Agrypnella/ eburnea, Ch./ type”, M, NHM
T009, Cardiophorus mirabilis Candèze, L, Lectotype designated here. [circular with blue margin] “SYN/ TYPE”; [handwritten] “Cardiophorus/ mirabilis/ Inde or. Cdz”; and with the author’s red designation label “LECTOTYPE/ Cardiophorus/ mirabilis/ Candèze desig./ Douglas 2015”, ?, NHM
T010, Triplonychus trivittatus Champion, L, Left hand specimen on card labeled: [disk with blue border] “SYN_/ TYPE”; “Tolé,/ Panama./ Champion.”; “♂”; [partly handwritten] “B.C.A.Coll.III[1]./ Triplonychus/ trivittatus/ Ch.”; [red label, partly handwritten] “LECTOTYPE Spcm on/ left/ Triplonychus/ trivittatus/ Champion/ desig. P.J.
T011, Triplonychus trivittatus Champion, PL, Right hand specimen [formerly left] on card labeled: [disk with blue border] “SYN_/ TYPE”; [disk with red border] “TYPE”; “Bugaba/ 800–1500 ft./ Champion”; “♂”; [partly handwritten] “B.C.A.Coll.III[1]./ Triplonychus/ trivittatus/ Ch.”, M, NHM
T012, Elater thoracicus Fabricius, PL, Pin unlabeled, in box with card: “Elater thoracicus F./ 2 syntypes/ UNIVERSITES/ ZOOLOGISKE MUSEUM/ UNIVERSITETSPARKEN 15/ DK–2100 KØBENHAVN Ø”, ?, ZMUC
T013, Elater thoracicus Fabricius, L, Lectotype designated here. Pin unlabeled, in box with card: “Elater thoracicus F./ 2 syntypes/ UNIVERSITES/ ZOOLOGISKE MUSEUM/ UNIVERSITETSPARKEN 15/ DK–2100 KØBENHAVN Ø”; and with the author’s red designation label “LECTOTYPE/ Elater/ thoracicus/ Fabricius desig./ Douglas 2015”, ?, ZMUC
T016, Oedostethus femoralis LeConte, L, Lectotype designated here, [green disc without writing]; [red label, partly handwritten] “Type/ 2461”; [handwritten] “Oedostethus/ femoralis/ Kansas. Lec.” “; and with the author’s red designation label “LECTOTYPE/ Oedostethus/ femoralis/ LeConte desig./ Douglas 2015”, M, MCZC
T017, Eniconyx pullatus Horn, L, Lectotype designated here. “Ari”; [partly handwritten red] “LectoTYPE/ 3405”; [HW] “Eniconyx/ pullatus/ Horn”; [partly handwritten, red] “MCZ/ SynType/33774”; and with the author’s red designation label “LECTOTYPE/ Eniconyx/ pullatus/ Horn desig./ Douglas 2015”, M, MCZC
T018, Eniconyx pullatus Horn, PL, “Ari”; [partly handwritten blue] “PARA-TYPE/ 3405”; and with the author’s orange label “PARALECTOTYPE/ Eniconyx/ pullatus/ Horn label/ Douglas 2015”, M, MCZC
T020, Aphricus californicus LeConte, H, [gold disc without writing]; [red] “Type./ 2623”; [handwritten] “Aphricus/ californicus/ S.D. Lec.”, M, MCZC
T024, Lesnelater madagascariensis Fleutiaux, L, Lectotype designated here. “Madagascar; Ambohibeloma.”; [handwritten] “Pachyelater/ madagascariensis/ ♂ presummé, type/ syn./ P. Lesne vid. 1906”; MUSÉUM PARIS/ 1952/ COLL A. OBERTHUR”; and with the author’s orange label “LECTOTYPE/ Lesnelater/ madagascariensis/ Fleutiaux label/ Douglas 2015”, M, MNHN
T026, Pachyelater madagascariensis Lesne, PL, “Madagascar; Ambohibeloma.”; [handwritten] “Pachyelater/ madagascariensis/ P. Lesne vid. 06”; MUSÉUM PARIS/ 1952/ COLL A. OBERTHUR” [with 6 antennomeres attached mounted on card attached to pin]. Abdomen on card, genitalia on separate glass slide HD–01, F, MNHN
T027, Pachyelater madagascariensis Lesne, PL, “Madagascar; Ambohibeloma.”; [handwritten] “Pachyelater/ madagascariensis/ Lesne/ P. Lesne vid. 06”; MUSÉUM PARIS/ 1952/ COLL A. OBERTHUR”. Abdomen on card, genitalia on separate glass slide HD–02, F, MNHN
T028, Cardiophorellus gracilicornis Cobos, P, “Odzala/ Congo/ Octobre”; “MUSÉUM PARIS/ MISSION/ A. DESCARPENTRIES/ ET. A. VILLIERS/ 1963–1964”; [red] “PARATYPUS/ A. COBOS”, M, MNHN
T029, Cardiophorellus gracilicornis Cobos, P, “Odzala/ Congo/ Octobre”; “MUSÉUM PARIS/ MISSION/ A. DESCARPENTRIES/ ET. A. VILLIERS/ 1963–1964”; [red] “PARATYPUS/ A. COBOS”, M, MNHN
T033, Phorocardius solitarius Fleutiaux, L, Lectotype designated here. [handwritten] “Tonkin/ Zhan Moi/ Vitalis Juin ##”; [red] “TYPE”; “MUSÉUM PARIS/ Coll. E. FLEUTIAUX”; [partly handwritten] “Phoroc/ solitarius/ Fleut. Type/ COLLECTION FLEUTIAUX”; [handwritten] “griffes très/ __lement bifides/4’ art, tarses/ tronqué/ carrément_”; and with the author’s red designation label “LECTOTYPE/ Phorocardius/ solitarius/ Fleutiaux desig./ Douglas 2015”. Body on card with abdomen, genitalia on separate glass slide, HD–03., F, MNHN
T034, Blaiseus bedeli Fleutiaux, L, “RÉG. DE LUC-NAM/ (TONKIN) L.BLAISE”; “MUSEUM PARIS/ (COLL. PH.FRANÇOIS)/ COLL. L. BEDEL 1922”; “TYPE”; [partly handwritten] “Blaiseus/ Bedeli Fleut./ type/ FLEUTIAUX det.”; “LECTOTYPE/Blaiseus/ bedeli/ Fleutiaux desig./Douglas 2006”, M, MNHN
T035, Cardiophorus florentini Fleutiaux, L, Lectotype designated here. “Tonkin/ Florentin”; [red] “Type”; “MUSÉUM PARIS/ Coll. E. FLEUTIAUX”; [handwritten] “Cardiophorus/ Florentini/ Fleut. Type”; [partly handwritten] “Fleut., Ann./ Soc. Ent. Fr./ 1894. P. 687/ Collection FLEUTIAUX”; [partly handwritten] “C. Florentini/ Fleut. type/ Collection FLEUTIAUX”; and with the author’s red designation label “LECTOTYPE/ Cardiophorus/ florentini/ Fleutiaux desig./ Douglas 2015”, ? , MNHN
T036, Aphricus chilensis Fleutiaux, L, Lectotype designated here. “TYPE”; [handwritten] “chili/ coll. Vienna”; [partly handwritten] “aphricus/ chilensis/ type/ COLLECTION FLEUTIAUX”. Head and thorax in capsule on separate pin labeled: [handwritten] “found loose/ in box/ CHM. 3.81”; and with the author’s red designation label “LECTOTYPE/ Aphricus/ chilensis/ Fleutiaux desig./ Douglas 2015”, M, MNHN
T037, Globothorax chevrolati Fleutiaux, L, Lectotype designated here. “Type”; “Collection Chevrolat”; [handwritten] “Globothorax/ chevrolati/ Fleut. type/ Bresil”; [partly handwritten] “Globothorax/ chevrolati/ Fleut. type/ Bresil/ Collection FLEUTIAUX”; [partly handwritten] “Fleut. E.R./ Soc. Ent. Belge/ 1891 p233/ Collection FLEUTIAUX”; and with the author’s red designation label “LECTOTYPE/ Globothorax/ chevrolati/ Fleutiaux desig./ Douglas 2015”, F, MNHN
T038, Cardiophorus (Perrinellus) angustatus Fleutiaux, T, “MAHATSINJO/ près Tananarive”; [handwritten] “Mahatsinjo/ Près Beforona”; “Type”; [partly handwritten] “Cardiophorus/ s.g. Perrinellus/ angustatus/ type/ Collection FLEUTIAUX”, M, MNHN