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

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


Introduction
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 (Gibson 1916), most species are probably carnivores. This, because of observed insectivorous behaviour (Devetak and Arnett 2012), and long paddle-like mandibles, which appear better adapted for locomotion and puncturing prey than chewing plant materials. Most larval cardiophorines inhabit soil (many in sandy soil), and dead or hollow trees (Palm 1972). Larvae move by pushing soil particles aside with their mandibles and maxilla-labial complex, while the thoracic legs and hydrostatic extension and contraction of the abdomen propel the larva forward. Traction for hydrostatic motion is partly by expansion and contraction of digitate anal lobes and smaller lateral abdominal projections (video available upon request). Cardiophorines are probably sometimes trophically important: adults are among the most abundant insects attracted to lights in some desert habitats (e.g. spp. of Horistonotus Candèze, 1860, Esthesopus Eschscholtz, 1829and Aptopus Eschscholtz, 1829 during the rainy season, Sonoran Desert, USA). They are also important pollinators: for example one South African orchid is pollinated primarily by a Cardiophorus Eschscholtz, 1829 species (Peter and Johnson 2005). Many Cardiophorinae are rare or localized to particular sand deposits or montane forests (e.g. Douglas 2003, Girard 2003, and Platia and Gudenzi 2000b and some are probably at extinction risk. Presently only Cardiophorus gramineus Scopoli, 1763 has formal conservation protection (as one of twelve beetle species protected by law in Sweden (Ljungberg et al. 2010).
Prior to this study Cardiophorinae had 29 described extant genera worldwide including about 1100 extant species, and two fossil species (Cockerell 1925, Eocene: USA;Hawkswood et al. 2009, Pleistocene: Madagascar), with one genus known only from fossils (Mionelater Becker, Miocene: Mexico). However, it remains unknown whether the subfamily and its genera are monophyletic. Eschscholtz (1829) named three cardiophorine genera (Aptopus, Cardiophorus, and Esthesopus) in his initial division of genus Elater Linnaeus, 1758. Fifteen additional Cardiophorine genera were described between 1800 and 1900, of which four to seven were in synonymy at the outset of this study according to various authors (listed in synonymy). Candèze (1860) wrote the first genus level revision of the Cardiophorinae, in his four-volume total revision of Elateridae (Candèze 1857-63). The most recent genus level revision was published by Schwarz (1906). The monophyly and membership of all genus level groups remain untested hypotheses because phylogenetic analysis has never been applied to any of the genera.

History of genera and tribes
The Cardiophorinae were divided into two tribes when Gurjeva (1974a) transferred the monotypic tribe Nyctorini from the Elaterinae into the Cardiophorinae. This transfer was made without comment, suggesting Gurjeva did not realize Nyctorini was incorrectly described as Elaterinae. So, division of the Cardiophorinae into tribes Cardiophorini and Nyctorini was perhaps accidental. Dolin (1975) placed Nyctorini in synonymy under Cardiophorini, effectively eliminating tribal level structure. Stibick (1979a) removed Nyctorini from synonymy because, although he dismissed two diagnostic characters as weak, he did not know of other Cardiophorinae with absent [short] adult prosternal lobes. This study will use phylogenetic results to assess whether tribe Nyctorini should be a synonym of Cardiophorini.

Phylogeny and monophyly of subfamily, tribes and genera
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 (Douglas 2011). The characters used to distinguish these two subfamilies since their description (Candèze 1860, Negastriinae as part of Cryptohypnites) are the short cardiophorine prosternal process, the broad prosternum of most Negastrii-nae, and the heart-shaped scutellum of most Cardiophorinae. However, Stibick (1979a) noticed these were not universal and has omitted the prosternal width character and qualified the other two characters with the terms "usually" and "normally." Other putative evidence for cardiophorine monophyly comes from the distinctive cardiophorine larvae (Hyslop 1921, Ôhira 1962, Stibick 1979a, Calder 1996. Potentially synapomorphic characteristics include: deeply cleft mandibles, thread-like abdomen with extra pseudosegmentation, and digitate anal lobes (Stibick 1979a). Although this larval type is known from Europe (Palm 1972), Central Asia (Atamuradov 1993), northeast Asia (Dolin and Gurjeva 1975), Japan (Ôhira 1962), Australia (Calder 1996), New Zealand (collections only, without digitate anal lobes), North America (Tenhet 1941) and South America (Costa et al. 1988), larvae remain unknown for most genera and species. Thus, it remains unknown whether these probable synapomorphies are of the Cardiophorinae alone, of the Cardiophorinae and other taxa, or of only some Cardiophorinae. Although these strong larval morphological characters exist and are possible evidence for cardiophorine monophyly, too few larvae are known for them to yet be used to test monophyly.
Douglas ' (2011) finding close relationship between the Cardiophorinae and the Negastriinae agrees with DNA sequence data-based results (Sagegami-Oba et al. 2007;Oba 2007;Bocak 2011 [with Platiana Schimmel, 1993 as sister to included Cardiophorinae, assigned to Dimini]; and Kundrata et al. 2016). Douglas (2011) also found strong support for Dendrometrinae: Hypnoidini as sister to Cardiophorinae + Negastriinae + Tropihypnus. This study uses Negastriinae, Hypnoidini, and Tropihypnus as outgroups for phylogenetic analysis of the Cardiophorinae, as the taxa identified as most likely (Douglas 2011) to render the Cardiophorinae non-monophyletic. Cardiophorine monophyly has not been demonstrated through analyses of larval or adult morphology to date, and requires testing.

Research plan
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 Schwarz (1906). Thirteen new genera, six subgenera and hundreds of species have been described since Schenkling's (1925) catalog. For these reasons, it is difficult to identify many cardiophorines and the current nomenclature is unlikely to reflect evolutionary history. Additionally, inconsistencies in the use of genus level names also make genus level identifications difficult using literature alone.
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 morphologicallydivergent 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.

Taxon sampling and Specimens examined
Specimens examined for morphological coding belonged to 29 insect collections (Table  1). Codens listed here follow Arnett et al. (1993), except where collections preferred other codens. Among these specimens were 61 primary types, or paratype specimens, representing 41 species (Appendix III). Appendix III also includes lectotype designations for 29 species. These are designated to fix generic concepts and to ensure their universal and consistent interpretation. Types of 307 more species, which were not coded for phylogenetic analysis, were photographed at NHM (London), ISNB and MNHN. Types of 85 more North American cardiophorine species (listed in Douglas 2003) were also examined to ensure taxon sampling reflected much of the group's morphological variation and assess new taxonomic placements.
Non-type specimens were identified by comparison with types (types were examined for 40 species) or specimens identified by experienced workers (three species, Appendix I). All non-type specimens examined were labelled with unique identifier numbers (Appendix II). Three distinctive undescribed species were included to better represent the Cardiophorinae. All identifications of non-types were evaluated using published keys and descriptions (Appendix I), and five species were identified using literature alone. Information from type specimens was often used in coding species. In most cases where a single name-bearing type did not already exist, a lectotype was designated for each species name (Appendix III). viewed as from inside the bursa (internal view), unless stated otherwise. Sclerites of the bursa copulatrix are illustrated in internal view unless lateral view is specified.

Morphological character coding
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 2), the value entered for each species was the mean of ratios of length measurements or mean angle. These values were ranked assigned to character-state bins "0" or "1" based on whether the measured value for that character for each species was above or below the median value for the character among all species.
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 Douglas 2011). Following these criteria, all qualitative characters identified as showing non-overlapping variation between at least two species were considered for possible use. Subsequent characters with apparent developmental or genetic non-independence were then excluded. Autapomorpic characters were also encoded because they provide branch length information for Bayesian analysis and diagnostic characters.
Phylogenetic analysis was conducted using 163 characters (Table 2), of which 27 were coded quantitatively (into binary pairs) and 136 qualitatively. These characters included 376 character states (after binary coding of quantitative characters), of which 40 were autapomorphic (Table 2). Qualitatively coded characters 6,9,22,27,29,37,44,60,85,123,133,145, and 161 were treated as ordered multistate characters. Three characters, common to many fossorial Elateridae (Douglas 2011), were omitted from the analyses presented here, to avoid phylogenetic bias due to convergent evolution. These were characters 9, 34 and 46 which included the following character states apparently associated with fossorial adults: mandibular apex unidentate; prosternum with anterior edge short, exposing labium; and protibiae near apex with posterior surface flattened, concave, or broadened apically, apparently modified for digging. Character 162, riparian habitat association was also excluded from the analysis. Analyses including these characters (not presented) had similar topologies to those with them omitted but Bayesian posterior probability values (PP) were lower throughout the tree, supporting the hypothesis of convergence. Table 2. 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. 5). Species named in brackets are designated as references to typify character states. Reference species used to typify character states in Douglas (2011) were re-used here even where the reference species was not included in this study. O = ordered multistate character.

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 (

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° (

Choice of optimality criteria
Although both parsimony and Bayesian analyses (as implemented by MrBayes v.3.1.2, Ronquist 2001, Ronquist andHuelsenbeck 2003, using the model by Lewis 2001) were used here to infer phylogeny, results of the model-based Bayesian analyses were preferred for taxonomic inference. The major expected advantage of using Bayesian analyses for morphological data is that the Mkv model of Lewis uses branch length information while parsimony does not. Empirically, Wiens (2005) found that accuracy of Bayesian analyses equalled or exceeded that of parsimony. Parsimony analyses were also performed because parsimony remains widely accepted.

Model selection
Bayes factors were used, as outlined by Sikes et al. (2006), to infer which of two evolutionary models best fit the data. These were the generalized Jukes-Cantor model for k states (Mkv), corrected for acquisition bias (Lewis 2001), with or without gamma distributed rate variation between characters (Mkv vs. Mkv + Γ). A Bayes factor of 1118 (2X ln L = -12837 for -Gamma and = -11719 for +Gamma) showed strong support (assessed as outlined by Kass and Raftery 1995) for models including gammadistributed rate variation between characters over models that did not include gamma variation.

Phylogenetic analyses
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* (Swofford 2001). The heuristic search procedure was used with 1000 random replications of stepwise-addition, with the branch-swapping algorithm (maxtrees set to auto-increase, multrees option in effect, Appendix 5). Bootstrap values were generated through 1000 replicates of bootstrapping using the same settings but with maxtrees set to 1000 and the number of replicates of stepwise addition reduced to 10 to reduce processing time. Decay index scores were calculated using PRAP (Müller 2004) for decay analysis of a strict consensus of all trees found in initial parsimony analysis. Zander (2004) found that Bayesian posterior probability (PP) values are predictably liberal at branch lengths typical for morphological studies (fewer than 35 changes). Because of this bias, PP values and other branch support metrics were corrected using Zander's table 4 when assessing clade credibility for hypothesis testing.

Tests of monophyly
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.

Generic diagnoses, figures, and key to genera
A key to the genera of cardiophorine, and corresponding diagnoses were developed using: existing diagnostic characters, the phylogenetic matrix (Appendix IV), and examination of other species from each genus.). Where phylogenetic results were informative, classification was revised to reflect phylogenetic history through synonymy, description of new genera, changes of rank, and new generic placements. Existing generic concepts were maintained where phylogenetic results were inconclusive.

Results
Bayesian analysis of the Cardiophorinae, Negastriinae and Hypnoidini matrix resulted in trees largely agreeing with results of morphological analysis of Elateridae (Douglas 2011). They agree in finding a well-supported monophyletic Cardiophorinae (as redefined below, Fig. 36, Node d, posterior probability PP = 0.96) and Cardiophorinae + Negastriinae (Fig. 36, Node a, PP = 1.00) which together render Hypnoidini paraphyletic (Fig. 36). Among these strongly supported clades, only Cardiophorinae + Negastriinae had support above 95% after correction for branch length (branch length = 28.2, correction according to Zander 2004, table 4). As in previous analyses, the most likely sister group of this hypnoidine-cardiophorine clade was Agriotes. However, unlike previous analyses, the Cardiophorinae here render the Negastriinae paraphyletic. Rejection of monophyly of the Negastriinae was strong (Table 3) although Negastriinae remains largely unresolved here (Fig. 36).
Within the Cardiophorinae, there were several clades with moderately high support (e.g. Fig. 36, Nodes e-i), but these do not subdivide the tree into even-sized major clades. The somewhat pectinate shape of this tree makes the terms basal and apical useful here to refer to taxa nearer to or farther from the root. Resolution was low in the tree's mid-region, especially within the paraphyletic Cardiophorus. Clades with more than 90% support within the Cardiophorinae include genus Blaiseus Fleutiaux (Blaiseus bedeli Fleutiaux, 1931plus B. nothoafricanus Douglas, 2009, Paracardiophorus + Cardiophorus cardisce (Say, 1834) + C. luridipes Candèze, 1860 (PP = 1.00, ≥ 95% after correction for branch length) and the Brazilian genera Globothorax Fleutiaux, 1891 + Teslasena (PP = 1.00, ≥ 95% after correction for branch length). Support for a priori hypotheses of monophyly was mostly weak to absent ( Table 3). Hypotheses of monophyly of the Negastriinae and the Hypnoidini were rejected ( Table 3).
Probabilities that the 11 genera tested were truly monophyletic ranged from <0.000008 to 0.90, and hypotheses of monophyly were rejected for all except Cardiophorus: Perrinellus, Horistonotus and Blaiseus (Table 3). Similarly, all tested published hypotheses of generic synonymy were found to have low support and three of four were clearly rejected Parsimony analysis mainly corroborated results of Bayesian analysis, also with low resolution near Cardiophorus (Fig. 37), except where branch support was low. Unlike the Bayesian tree, Adrastus pallens was included in the hypnoidine-cardiophorine clade, between the Negastriinae (here monophyletic excluding Negastrius americanus, D =1, BS < 50%) and the Hypnoidini (monophyletic D =1, BS < 50%). The Cardiophorinae were again monophyletic (D=2, BS < 50%) in the parsimony analysis with the addition of Negastrius americanus, Margogastrius, Pachyelater, and Teslasena. In both trees Blaiseus, Aphricus and Patriciella, an undescribed species from New Zealand, Pachyelater, Negastrius americanus, Nyctor, Neocardiophorus, and Margogastrius are near the base of Cardiophorinae. The remainder of the Cardiophorinae were weakly resolved by parsimony analysis, except that as in the Bayesian analysis Paracardiophorus grouped with Cardiophorus cardisce, and C. luridipes. Both analyses also included a clade of 11-13 genera (Node j of Bayesian analysis), whose genera are entirely or mainly in the southern hemisphere or northern tropical regions. These "southern clade" taxa are: Esthesopus; Odontocardus; Triplonychoidus; Aptopus agrestis Erichson, 1840;Paraplatynychus Fleutiaux, 1931;Triplonychus Candèze, 1860;Cardiotarsus mjobergi (Elston, 1930); Cardiodontulus Van Zwaluwenburg, 1963;Craspedostethus Schwarz, 1898; Paracardiophorus species from Australia and Chile; and Buckelater Costa, 1973. Only parsimony analysis included genera Globothorax and Teslasena in the southern clade.

Monophyly of Cardiophorinae
Bayesian (Fig. 36, Table 3) and also parsimony analyses (Fig. 37) show that the Cardiophorinae are a well-supported clade if several taxa are transferred into Cardiophori-nae. Here, the Cardiophorinae can be corrected by adding a few species and genera from subfamilies Physodactylinae, Dendrometrinae and Negastriinae (Margogastrius, Negastrius americanus, Teslasena, and the undescribed species from New Zealand, and Pachyelater Lesne). Three of these also require further taxonomic alterations, as discussed below.
The resulting Bayesian tree (Fig. 36) also showed Negastriinae as paraphyletic (Nodes a-c) and strong support for monophyly of Cardiophorinae + Negastriinae (Node a). However parsimony analysis found weak support for a monophyletic core Negastriinae that is sister to the Cardiophorinae (Fig. 37). In the Bayesian analysis, support for negastriine monophyly was only 0.0001 (Table 3). The strength of this rejection is surprising, given that Bayesian analysis (Fig. 36) left Negastriinae mostly unresolved (Node a), and that support for the two nodes showing paraphyly of the Negastriinae was only 0. 65 & 0.72 (Nodes b & c). Further phylogenetic analysis, including more genera of the Negastriinae, is important to further test the validity of the Negastriinae and membership of both subfamilies.

Tribal classification of the Cardiophorinae
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 3, Fig. 36). The only diagnostic characters of Nyctorini (Semenov-Tian-Shanskij and Pjatakova 1936), i.e., the short anterior prosternal lobe and sexual size dimorphism, are homoplastic characters found in many fossorial elaterids. For these reasons Nyctorini should be a junior synonym of Cardiophorini, effectively eliminating tribal level classification of the Cardiophorinae. Because of the generally pectinate shape of trees for Cardiophorinae (Figs 36,37), no natural divisions were found for a convenient tribal level classification, and all Cardiophorinae should be placed in tribe Cardiophorini.

Genera of the Cardiophorinae and Negastriinae
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 36,37). These distinctive genera, Agrypnella, Cardiohypnus and Rivulicola live in riparian habitats in the Neotropics, South Asia and Australia respectively. Rivulicola is unusual as the only Negastriinae known from Australia. These are recognizable among the Negastriinae because of their scale-like setae. All three genera were once placed in the Cardiophorinae because of their heart shaped scutella (with emarginate anterior edge), ovoid pronota and elytra, and short prosternal processes. They were transferred independently to the Negastriinae by three different authors (Dolin 1992, Golbach 1994, Calder 1996, at least in part, because of their convex prosternal sides. However, despite the similarities that these three genera share with Cardiophorinae, this group was not found sister to the Cardiophorinae here. Fleutiauxellus Méquignon, a negastriine appearing less like Cardiophorus is the most likely sister group to the Cardiophorinae according to Bayesian analysis, sharing with many Cardiophorinae the pedunculate anterior sac of the bursa copulatrix. 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. 36, Node d). Here, genus Aphricus is made paraphyletic in both analyses by at least the fossorial Australian genus Patriciella Van Zwaluwenburg, 1953 and an undescribed species similar to Aphricus from New Zealand. In order to avoid recognising a non-monophyletic genus, new genus Chileaphricus is established for Aphricus chilensis Fleutiaux. Since Aphricus (from California, USA) + Patriciella and the undescribed species from New Zealand, form a clade with moderate support (PP = 0.82, D = 1) they should be treated as a single genus (by synonymising Patriciella under Aphricus, its type species becoming Aphricus australicus Van Zwaluwenburg, 1947).
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, (Douglas 2009). The type species (B. Bedeli Fleutiaux, 1931), and a male of the South African B. nothoafricanus Douglas, 2009 were included here. Although support for their monophyly was only 0.90 (also supported by parsimony, Fig. 37. Bremer support, (D) = 3, bootstrap support, (BS) = 81), the characters uniting them are distinctive. One such synapomorphy is their unique, split parameres (Fig. 25). The widespread distribution of the few known species, and the basal position of this genus within Cardiophorinae suggest Blaiseus is a long-separated lineage with a possibly relictual distribution.
Pachyelater Lesne, 1897 is a robust-bodied fossorial elaterid genus from Madagascar with sexually dimorphic males and females . Because this genus falls within the Cardiophorinae in both analyses, it should be transferred from Dendrometrinae to Cardiophorinae. Because females of Pachyelater are flightless and fossorially adapted the undiscovered females of the closely-related Aphricus may also share these traits (Fig. 36). Furthermore, females of Aphricus spp. may also be similarly larger than males, with reduced eyes and have the ovipositor and bursa copulatrix without sclerites. Margogastrius Schwarz, a genus known from only two damaged female type specimens from coastal Tanzania (also flightless and fossorial) was found with weak branch support as the sister to Blaiseus (although they are not closely related according to parsimony, Fig. 37). Examining internal genitalia of the remaining undissected type specimen might yield further phylogenetic information. No associated males have been identified with external morphology or distribution like these females. While the historically enigmatic species Negastrius americanus clearly belongs to the Cardiophorinae, the characters examined in both male and female specimens did not suggest placement in any other genus (Figs 36,37). Therefore I propose to place it in the new monotypic genus Floridelater gen. n.
The remaining taxa in the polytomy of the Bayesian analysis (Fig. 36, Node d): Neocardiophorus Gurjeva, 1966;Nyctor Semenov-Tian-Shanskij & Pjatakova, 1936;and Cardiophorus subgenus Metacardiophorus Gurjeva, 1966 are all known from central Asia. Among these, only Nyctor is known from both sexes, so the discovery of females of the other two genera would provide important data for improved phylogenetic placement and on the evolution of flightlessness in the Cardiophorinae. Since subgenus Metacardiophorus is distantly related to subgenus Cardiophorus (Fig. 36, Node d, not h; Fig. 37), it should be raised to genus rank.
Among the genera historically confounded with Cardiophorus, the most basal is Paracardiophorus. This genus was found polyphyletic (Table 3, Fig. 36: Nodes e, f & i, Fig. 37) because it includes superficially similar species from Australia and Chile. It is argued below that those should be part of a new genus. The type species of Paracardiophorus forms a fully supported clade with two North American Cardiophorus species (PP > 0.95 after correction for branch length (12), uncorrected probability =1.00), which is also indicated by parsimony analysis (Fig. 27, D = 2, BS = 57). These and all other North American species with the same apparent synapomorphies should be transferred to Paracardiophorus. These are the North American Cardiophorus with the base of the female spermathecal gland duct sclerotised (Figs 34,35), some species also have truncate pronotal hind angles (Fig. 4), and the aedeagal parameres spatulate (Fig.  65). Paracardiophorus is the most likely (PP = 0.67) sister group of the remainder of Cardiophorinae.
Beyond confusion with Paracardiophorus, genus Cardiophorus: subgenus Cardiophorus was paraphyletic at four nodes (Fig. 36,. It is also paraphyletic at 2 or more nodes in parsimony analysis, with most forming a polytomy in the parsimony analysis ( Fig. 37). This large genus, which contains half the described cardiophorine species, is paraphyletic because it also includes 21 other genera (Fig. 36, Node h). Unfortunately, because of this poor phylogenetic resolution, there is little basis yet for an improved definition of Cardiophorus. The monotypic Cardiophorus: subgenus Zygocardiophorus Iablokoff-Khnzorian & Mardjanian, 1981 was found to be sister to Cardiophorus + the remainder of Cardiophorinae (Nodes f, g) and thus should be raised to genus rank. The position of Cardiophorus: subgenus Lasiocerus Buysson is unknown, because no specimens were available for examination. Dolin and Gurjeva (1975) described genus Paradicronychus based on larval characters only (although conspecific adults were also known), and without a formal designation of a type species. Because of IZCN regulations for genera described after 1930 (Art. 13.3), Paradicronychus is not an available name. Although larvae of many cardiophorines from the former USSR are known, larval morphology of the world fauna remains too poorly documented to define genera based on larvae alone. Both analyses placed C. inflatus Candèze, 1882, (considered Paradicronychus by Dolin andGurjeva (1975)) within the broadly paraphyletic Cardiophorus. Because of this result, and because no adult characters were identified to distinguish it from Cardiophorus, the nomen nudum name Paradicronychus should be placed as synonym of the nominate subgenus of Cardiophorus (with its included species to Cardiophorus as C. inflatus Candèze, 1882, andC. nothus Candèze, 1865).
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 3). Evidence that numerous other cardiophorines have similar modifications for digging may be further evidence these characters are convergent and this assemblage is artificial. While these genera remain non-monophyletic and weakly defined, I do not recommend taxonomic changes until their positions are better resolved.
Globothorax Fleutiaux and Teslasena Fleutiaux (Physodactylinae) are a strongly supported ( Fig. 36. Uncorrected PP = 1.00, branch length = 2; Fig. 37, Bremer support = 2) clade within Cardiophorus and the other genera rendering it paraphyletic (Node g). Like the other genera here, these two should not be synonymised under Cardiophorus. Teslasena should be considered a junior synonym of Globothorax, because of the well-supported monophyly of these two species. This synonymy means that included species Teslasena femoralis (Lucas, 1857), Teslasena foucarti Chassain, 2005, andTeslasena lucasi Fleutiaux, 1899 are transferred to Globothorax as Globothorax femoralis (Lucas, 1857, Anelastes); G. foucarti Chassain, 2005;and G. lucasi Fleutiaux, 1899 respectively. Their sympatry in Brazil further supports the hypothesis that the known specimens of Teslasena and Globothorax are dimorphic males and females of one genus (although not necessarily conspecific). I recommend this despite characters presented by Rosa (2014) distinguishing the two genera: these may be variation between species, or sexual dimorphism but their presence does not refute the hypothesis that they are best understood as congeneric.
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 36, 37), Dicronychus should not be a synonym of Cardiophorus, at least until a monophyletic Cardiophorus can be defined. However, since Dicronychus is only distinguished from Cardiophorus by the presence of a second tarsal claw tooth (Fig. 21), there may be no basis on which to distinguish it from Cardiophorus even at the species level because of apparent intraspecific dimorphism. Such dimorphic claws may underlie the sympatric Cardiophorus aptopoides Candèze, 1865;and C. brevis (Candèze, 1859) from Mexico, which appear identical except the presence or absence of a basal claw tooth (including aedeagal shape and regional colour variants). A similar otherwise apparently identical Cardiophorus-Dicronychus pair of species (Cardiophorus varius, Cate et al., 2002 andD. hoberlandti Cate et al., 2002) from Iran also may be a single species with dimorphic claws. There is no evidence for the monophyly of genera Dicronychus and Platynychus Motschulsky, 1858 (PP = 0.07, Figs 36, 37, Table 3), therefore Platynychus should be removed from synonymy under Dicronychus, where it has been placed by some authors. Platynychus is distinguished from both Cardiophorus and Dicronychus by its closed procoxal cavities.
Genus Cardiophorellus Cobos also falls within the paraphyletic nominate subgenus of Cardiophorus (Fig. 36, Node h, not contracted by parsimony, Fig. 37). The type species of Cardiophorellus is much like Cardiophorus except the anterior edge of its scutellum is broadly concave and not angulately emarginate, and its mandibles are simple. Due to phylogenetic uncertainty, there is no evident best taxonomic placement for Cardiophorellus. Because of this uncertainty, and because Cardiophorellus is readily diagnosed, it seems best to continue to consider Cardiophorellus a valid genus. The type specimen of the monotypic subgenus Cardiophorellus (Parapleonomus) Cobos, 1970 was not found at MNHN (Paris), so I cannot comment on its validity or rank.
Although the hypothesis of Aptopus Eschscholtz monophyly was rejected (Table 3, Fig. 37), this only affects the placement of the species A. agrestis (Erichson). Apart from its pectinate claws, this species is like Horistonotus species with costate elytral intervals. Because parsimony phylogenetic analysis suggested A. agrestis was the most likely sister taxon to Horistonotus simplex LeConte, such Aptopus species with carinae following the lateral edge of the pronotum should be transferred to Horistonotus. However, because the type specimen of A. agrestis was not examined, this species is not transferred to Horistonotus here. The concept of Aptopus used here is from modern authors (e.g. Aranda 1998, also Section 1 of Candèze 1860) because the type specimens of the type species, A. tibialis Eschscholtz, 1829 are lost or were unavailable for examination and because the only published species description lacks detail (eight words only).
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 2) rejected the hy-pothesis that even the two African species were monophyletic (also not recovered by parsimony, Fig. 37). C. mjobergi was placed at Node k of the Bayesian tree ( Fig. 36) within the southern clade ( Fig. 36 node j). I propose transfer of Cardiotarsus mjobergi to genus Cardiodontulus, from Papua New Guinea, because of this non-monophyly and it matches the Van Zwaluwenburg's definition of that genus. This placement is also plausible, because both are from the Australian biogeographic region. Although the type specimen of C. mjobergi was not examined, I am confident in the identification of the specimens examined because they were from near the type locality, which is in a well-collected area near a major insect collection, and this species was also illustrated in Calder's (1996) guide to Australian Elateridae. Otherwise, I propose no changes to the biologically inaccurate (but easily diagnosable) genus Cardiotarsus until the phylogeny of Cardiophorinae is better resolved.
The remaining apical southern clade (PP = 0.81, Fig. 36: Node j) is composed mainly of Australian and Neotropical species, plus two South Asian genera and one from Africa. This clade was also inferred by parsimony ( Fig. 37, but with Globothorax and Teslasena added), and includes mostly species with bilobed or multilobed proximal sclerites of the bursa copulatrix, and many of the species with closed procoxal cavities, and lacking lateral expansions of the parameres. Among these, the monophyly of each of Odontocardus Fleutiaux, 1931;Triplonychoidus Schwarz, 1906;Paraplatynychus Fleutiaux, 1931;Triplonychus Candèze, 1860;Cardiodontulus Van Zwaluwenburg, 1963; Craspedostethus; and Buckelater were not tested. These genera remain unaltered, except as discussed for Cardiodontulus. Two large, mainly Neotropical genera (extending into temperate North America) Esthesopus and Horistonotus are both not monophyletic (Table 3). However their definitions and status should be maintained until better resolution is available. The definition of Horistonotus is broadened here to include species with multiple claw points.
Of the five species in the weakly supported apical clade ( Fig. 36, Node l), two belong to the polyphyletic genus Paracardiophorus. These two species from Australia and Chile are rendered paraphyletic (also at low posterior probability) by Buckelater, from Brazil. Because the included Australian and South American Paracardiophorus are identical in most characters including the male and female genitalia, I propose placement of them in a new genus along with other species from both continents sharing their diagnostic characters. The type species of this new genus, Austrocardiophorus, is Cardiophorus humeralis Fairmaire & Germain, 1860 from Chile (recently in Paracardiophorus). This solution is considered preferable to placement in the currently monotypic Buckelater because its female genitalic characters remain unknown, which contributes to taxonomic uncertainty.

Character evolution
This section outlines some character state changes implied by the trees (Figs 36, 37), which may be diagnostically helpful. While these characters may be true synapomor-phies of their groups, Bayesian analysis does not rely on identifying them unambiguously as such.
Three characters unite the Negastriinae + Cardiophorinae. The fusion of the parameres at their midlength into a tube (Char. 117,Figs 24, 25) appears unique among the Coleoptera (Iablokoff-Khnzorian and Mardjanian 1981), and universal among Cardiophorinae and Negastriinae. Examination of two other possible synapomorphies revealed more intrageneric variability than found by Douglas (2011). Firstly, the hindwing membrane has an anal notch (Fig. 17, at AA4) in all examined Negastriinae except Migiwa Kishii, 1966, but this notch is present in only most Cardiophorinae (Char. 75). Secondly, all included Negastriinae, except Arhaphes Candèze, 1860, but only most cardiophorine genera had a tridentate lobe at the midline of the posterior edge of the pronotum. The only character to distinguish the Cardiophorinae from the Negastriinae, was an apparent reversal to straight-sided prosternum (alternative = convex, Char. 36). No variation from this character-state was found in Cardiophorinae.
No clear evidence was found for basal synapomorphies of Negastriinae not also shared by Cardiophorinae. As found by Douglas (2011), they were distinguished from Cardiophorinae by their convex lateral edges of the prosternum (near midlength). However, this character is an apparent symplesiomorphy shared with Hypnoidus and Tropihypnus according to the most likely topologies identified by Douglas (2011).
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.
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.

Future research
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.  Stibick (1971), with subsequent changes by Calder 1996, Dolin 1976, Dolin 1992, Dolin and Girard 1998, Golbach 1994, Kishii 1976 Tarsal claws with or without basal point; tarsal claws with basal setae ( (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. 21 (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. 1). Elytra all-black, with or without apical shelf-like apical extensions (Fig. 71) (17) (40) Scutellum with anterolateral edges evenly rounded (Fig. 8)  . 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. 2); most species with pronotal hind angles truncate dorsally (Fig. 4) so apex is composed of only the narrow hypomeral portion; most species with posterior edges of hypomeron mesad of hind angles straight or sinuate; procoxal cavities open or closed; free portion of parameres cylindrical to flattened (Fig. 65); bursa copulatrix with proximal sclerites ovoid (Figs 28, 66 Description. 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. 4); frons with supra-orbital groove present (Fig. 1). Prothorax: Pronotum with punctures circular; sublateral incisions present, carinae present in some; posterior edge of pronotum with 3 low apices mesally; hind angles with a single carina reaching to near midlength, it is unknown whether this is the hind angle carina or the lateral carina, single carina not situated ventrad of lateral edge of pronotum; hind angles not truncate dorsally; hypomeral hind edge rectangularly emarginate (Fig. 3) immediately meso-ventrad of hind angles; procoxal cavities closed. Prosternum with sides concave in ventral view; anterior prosternal lobe covering labium; prosternal process not curved dorsad (less than 30°), ventral surface carinate laterally, or not. Mesothorax: Scutellum with anterior edge weakly concave, posterior apex rounded to pointed (Figs 7,8). Mesosternum with anterior edges weakly concave lateral to mesosternal cavity in lateral view; mesosternal cavity with lateral edges sinuate anterad of mesocoxae. Elytral intervals not costate. Hind wings, notched in anal area. Legs: Tarsi without apically extending lobes or pads; tarsal claws each with 1 apex; metacoxal plate covers 1/2-2/3 of metatrochanter with legs withdrawn. Male genitalia: Abdominal segment 9 with tergite and sternites articulated at sides; parameres without apicolateral or apicomedial expansions, apices not forked, sides with 2 setae; aedeagus with basal struts approximately 1 times median lobe length, median lobe simple, tapered. Female genitalia: Ovipositor with baculae present; coxites flexible. Bursa copulatrix with colleterial glands indiscernible; without sclerotised spermathecae; bilobed spine-bearing sclerites present (Fig. 156); spermathecal gland duct without row of diverticulae, base not sclerotised; anterior end of bursa with 2 pedunculate sacs sharing common attachment to bursa.
Etymology. Masculine. Named for a genus of Cardiophorinae known only from the southern hemisphere.

Discussion.
Please see text of discussion above for argumentation for new genus. not hidden. Legs. Tarsomeres without ventral lobes; tarsal claws with one apex per side. Aedeagus. Aedeagus with paramere apices not forked (Fig. 41).
Etymology. Masculine. Named for a genus of Cardiophorinae known only from Chile.
Discussion. 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.
Etymology. Masculine. Named for a genus of Cardiophorinae known only from southeastern USA.
Discussion. 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.

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 24, 25); female ovipositor without styli.

Description of Cardiophorinae
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.

Diagnosis of Cardiophorinae
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 24, 25); females with ovipositor lacking styli.

Diagnoses of genera
Diagnoses presented here distinguish each genus from all other Cardiophorinae. Diagnoses for newly described and redescribed genera are provided above with corresponding genus description.
Known from South and North America, and the Greater and Lesser Antilles, 50 spp.  (Figs 78, 80), apparently for digging, tarsi without apically extending lobes or pads, tarsal claws with 2-3 apices per side. Also, posterior edge of pronotum bidentate mesally; anterior edge of scutellum straight, females with compound eyes nearly flat and antennae reaching only 2/3 of distance to pronotal hind angles; bursa copulatrix with proximal (only) sclerites parallel sided (Fig. 30) Diagnosis. 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. 134). Also. Bursa copulatrix with paired proximal (largest) sclerites ovoid (Fig. 138). Known from South and North America, and the Antilles, 106 spp. Schwarz, 1903Figs 42-44 Margogastrius Schwarz, 1903b. Replacement name for Gastrimargus Schwarz, 1902. Diagnosis. 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. 42), apparently strongly modified for digging; Tarsomere 4 without ventral lobe or pad extending beyond base of tarsomere 5; proximal (largest) sclerites of bursa copulatrix reduced, capsule-like spermatheca attached to ventral surface of bursa by short duct (Fig. 44). Known from Tanzania, females only, 1 sp.

Metacardiophorus Gurjeva, 1966 Figs 62-64
Metacardiophorus Gurjeva, 1966: 91. stat Gurjeva, 1966: 95. Type species: N. mamajevi Gurjeva, 1966 Diagnosis. 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.  (Figs 50, 51); tarsi without ventral lobes and tarsal claws with one apex per side. Also, males with aedeagus parameres undivided (Fig.  53). Females with compound eyes reduced (Fig. 50); antennae reaching only halfway to pronotal hind angles; ovipositor reduced, with baculae shorter than coxites; bursa copulatrix without sclerites. Known from: Madagascar, Southern Africa, 6 spp. Diagnosis. Prothorax. Pronotum with complete carina at lateral edge (Fig. 141), reaching from hind angle to anterior edge of prothorax. Pterothorax. Scutellum with middle of anterior edge broadly concave. Also. Tarsal claws with both basal and apical points on each side, without basal setae; bursa copulatrix with proximal sclerites bilobed (Fig. 142). Diagnosis is based on type species. P. fuscipennis Candèze, 1860, andP. incostatus Fleutiaux, 1931  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 2 points per side. Bursa copulatrix with paired proximal (largest) sclerites bilobed with attached semi-sclerotised membrane with spines (Fig. 162). Also. Procoxal cavities apparently closed; aedeagus parameres apices without lateral expansions. Known from Japan, and Taiwan, 4 spp. Diagnosis. 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. 145) Diagnosis. Prothorax. Pronotum with complete lateral carina (Fig. 105), reaching from hind angle to anterior edge of prothorax (displaced ventrad in some). Legs. Tarsal claws with or without basal point on each side. Also. Scutellum with anterior edge abruptly emarginate; tarsal claws with basal setae (Fig. 106, possibly absent in some); bursa copulatrix with proximal sclerites elongate-ovoid (Fig. 107). In type species (T.

Diagnosis.
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). 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, area between anterior-most point and humeral angle arcuate or straight in dorsal view (not sinuate), apices with or without shelf-like epipleural extensions (Fig. 71). Legs. Tarsi without ventral lobes or pads extending beyond base; claws with only 1 point per side. Also. Bursa copulatrix with paired proximal (largest) sclerites ovoid with long spines; base of spermathecal gland duct sclerotised, but without paired plate-like appendages. Known from Georgia, and Turkey, east to Turkmenistan and Iran, 1 sp.

Revised synonymy of Cardiophorinae
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 (Schwarz 1906) and the Biologia Centrali-Americana (Champion 1895) were also consulted. Since earlier names and applications were sometimes not cited in these works, a general search was made through them to track names and their origins. Following this procedure, the following monographs, reviews, faunal lists and faunal works were also examined (Candèze 1860(Candèze , 1891aGistel 1834Gistel , 1848Gistel , 1856Harold 1869;Heyden et al. 1891Heyden et al. , 1908Jakobson 1913; Jacquelin du Val 1859; Kiesenwetter 1858; Lacordaire 1857; Stein and Wiese 1877). In addition, the following lists of genus-group names and the works they cited were also examined (Agassiz 1846;Scudder 1882, Waterhouse 1902, 1912, Lucas 1920.

Type specimens
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.