We synthesize data on all known extant and fossil Coleoptera family-group names for the first time. A catalogue of 4887 family-group names (124 fossil, 4763 extant) based on 4707 distinct genera in Coleoptera is given. A total of 4492 names are available, 183 of which are permanently invalid because they are based on a preoccupied or a suppressed type genus. Names are listed in a classification framework. We recognize as valid 24 superfamilies, 211 families, 541 subfamilies, 1663 tribes and 740 subtribes. For each name, the original spelling, author, year of publication, page number, correct stem and type genus are included. The original spelling and availability of each name were checked from primary literature. A list of necessary changes due to Priority and Homonymy problems, and actions taken, is given. Current usage of names was conserved, whenever possible, to promote stability of the classification.

New synonymies (family-group names followed by genus-group names): Agronomina Gistel, 1848 syn. nov. of Amarina Zimmermann, 1832 (Carabidae), Hylepnigalioini Gistel, 1856 syn. nov. of Melandryini Leach, 1815 (Melandryidae), Polycystophoridae Gistel, 1856 syn. nov. of Malachiinae Fleming, 1821 (Melyridae), Sclerasteinae Gistel, 1856 syn. nov. of Ptilininae Shuckard, 1839 (Ptinidae), Phloeonomini Adam, 2001 syn. nov. of Omaliini MacLeay, 1825 (Staphylinidae), Sepedophilini Adam, 2001 syn. nov. of Tachyporini MacLeay, 1825 (Staphylinidae), Phibalini Gistel, 1856 syn. nov. of Cteniopodini Solier, 1835 (Tenebrionidae); Agronoma Gistel 1848 (type species Carabus familiaris Duftschmid, 1812, designated herein) syn. nov. of Amara Bonelli, 1810 (Carabidae), Hylepnigalio Gistel, 1856 (type species Chrysomela caraboides Linnaeus, 1760, by monotypy) syn. nov. of Melandrya Fabricius, 1801 (Melandryidae), Polycystophorus Gistel, 1856 (type species Cantharis aeneus Linnaeus, 1758, designated herein) syn. nov. of Malachius Fabricius, 1775 (Melyridae), Sclerastes Gistel, 1856 (type species Ptilinus costatus Gyllenhal, 1827, designated herein) syn. nov. of Ptilinus Geoffroy, 1762 (Ptinidae), Paniscus Gistel, 1848 (type species Scarabaeus fasciatus Linnaeus, 1758, designated herein) syn. nov. of Trichius Fabricius, 1775 (Scarabaeidae), Phibalus Gistel, 1856 (type species Chrysomela pubescens Linnaeus, 1758, by monotypy) syn. nov. of Omophlus Dejean, 1834 (Tenebrionidae). The following new replacement name is proposed: Gompeliina Bouchard, 2011 nom. nov. for Olotelina Baguena Corella, 1948 (Aderidae).

Reversal of Precedence (Article 23.9) is used to conserve usage of the following names (family-group names followed by genus-group names): Perigonini Horn, 1881 nom. protectum over Trechicini Bates, 1873 nom. oblitum (Carabidae), Anisodactylina Lacordaire, 1854 nom. protectum over Eurytrichina LeConte, 1848 nom. oblitum (Carabidae), Smicronychini Seidlitz, 1891 nom. protectum over Desmorini LeConte, 1876 nom. oblitum (Curculionidae), Bagoinae Thomson, 1859 nom. protectum over Lyprinae Gistel 1848 nom. oblitum (Curculionidae), Aterpina Lacordaire, 1863 nom. protectum over Heliomenina Gistel, 1848 nom. oblitum (Curculionidae), Naupactini Gistel, 1848 nom. protectum over Iphiini Schonherr, 1823 nom. oblitum (Curculionidae), Cleonini Schonherr, 1826 nom. protectum over Geomorini Schonherr, 1823 nom. oblitum (Curculionidae), Magdalidini Pascoe, 1870 nom. protectum over Scardamyctini Gistel, 1848 nom. oblitum (Curculionidae), Agrypninae/-ini Candeze, 1857 nom. protecta over Adelocerinae/-ini Gistel, 1848 nom. oblita and Pangaurinae/-ini Gistel, 1856 nom. oblita (Elateridae), Prosternini Gistel, 1856 nom. protectum over Diacanthini Gistel, 1848 nom. oblitum (Elateridae), Calopodinae Costa, 1852 nom. protectum over Sparedrinae Gistel, 1848 nom. oblitum (Oedemeridae), Adesmiini Lacordaire, 1859 nom. protectum over Macropodini Agassiz, 1846 nom. oblitum (Tenebrionidae), Bolitophagini Kirby, 1837 nom. protectum over Eledonini Billberg, 1820 nom. oblitum (Tenebrionidae), Throscidae Laporte, 1840 nom. protectum over Stereolidae Rafinesque, 1815 nom. oblitum (Throscidae) and Lophocaterini Crowson, 1964 over Lycoptini Casey, 1890 nom. oblitum (Trogossitidae); Monotoma Herbst, 1799 nom. protectum over Monotoma Panzer, 1792 nom. oblitum (Monotomidae); Pediacus Shuckard, 1839 nom. protectum over Biophloeus Dejean, 1835 nom. oblitum (Cucujidae), Pachypus Dejean, 1821 nom. protectum over Pachypus Billberg, 1820 nom. oblitum (Scarabaeidae), Sparrmannia Laporte, 1840 nom. protectum over Leocaeta Dejean, 1833 nom. oblitum and Cephalotrichia Hope, 1837 nom. oblitum (Scarabaeidae).

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Family-group names in Coleoptera (Insecta)

Insect diversity represents about 60% of the estimated million-and-a-half described eukaryotic species worldwide, yet comprehensive and well-resolved intra-ordinal phylogenies are still lacking for the majority of insect groups. This is the case especially for the most species-rich insect group, the beetles (Coleoptera), a group for which less than 4% of the known species have had their DNA sequenced. In this study, we reconstruct the first higher level phylogeny based on DNA sequence data for the species-rich darkling beetles, a family comprising at least 20 000 species. Although amongst all families of beetles Tenebrionidae ranks seventh in terms of species diversity, the lack of knowledge on the phylogeny and systematics of the group is such that its monophyly has been questioned (not to mention those of the subfamilies and tribes contained within it). We investigate the evolutionary history of Tenebrionidae using multiple phylogenetic inference methods (Bayesian inference, maximum likelihood and parsimony) to analyse a dataset consisting of eight gene fragments across 404 taxa (including 250 tenebrionid species). Although the resulting phylogenetic framework only encompasses a fraction of the known tenebrionid diversity, it provides important information on their systematics and evolution. Whatever the methods used, our results provide strong support for the monophyly of the family, and highlight the likely paraphyletic or polyphyletic nature of several important tenebrionid subfamilies and tribes, notably the polyphyletic subfamilies Diaperinae and Tenebrioninae that clearly require substantial revision in the future. Some interesting associations in several groups are also revealed by the phylogenetic analyses, such as the pairing of Aphtora Bates with Phrenapatinae. Furthermore this study advances our knowledge of the evolution of the group, providing novel insights into much-debated theories, such as the apparent relict distribution of the tribe Elenophorini.

Higher level molecular phylogeny of darkling beetles (Coleoptera: Tenebrionidae)

A review of genus-group names for darkling beetles in the family Tenebrionidae (Insecta: Coleoptera) is presented. A catalogue of 4122 nomenclaturally available genus-group names, representing 2307 valid genera (33 of which are extinct) and 761 valid subgenera, is given. For each name the author, date, page number, gender, type species, type fixation, current status, and first synonymy (when the name is a synonym) are provided. Genus-group names in this family are also recorded in a classification framework, along with data on the distribution of valid genera and subgenera within major biogeographical realms. A list of 535 unavailable genus-group names (e.g., incorrect subsequent spellings) is included. Notes on the date of publication of references cited herein are given, when known. The following genera and subgenera are made available for the first time: Anemiadena Bouchard & Bousquet, subgen. nov. (in Cheirodes Gene, 1839), Armigena Bouchard & Bousquet, subgen. nov. (in Nesogena Maklin, 1863), Debeauxiella Bouchard & Bousquet, subgen. nov. (in Hyperops Eschscholtz, 1831), Hyperopsis Bouchard & Bousquet, subgen. nov. (in Hyperops Eschscholtz, 1831), Linio Bouchard & Bousquet, subgen. nov. (in Nilio Latreille, 1802), Matthewsotys Bouchard & Bousquet, gen. nov., Neosolenopistoma Bouchard & Bousquet, subgen. nov. (in Eurynotus W. Kirby, 1819), Paragena Bouchard & Bousquet, subgen. nov. (in Nesogena Maklin, 1863), Paulianaria Bouchard & Bousquet, gen. nov., Phyllechus Bouchard & Bousquet, gen. nov., Prorhytinota Bouchard & Bousquet, subgen. nov. (in Rhytinota Eschscholtz, 1831), Pseudorozonia Bouchard & Bousquet, subgen. nov. (in Rozonia Fairmaire, 1888), Pseudothinobatis Bouchard & Bousquet, gen. nov., Rhytinopsis Bouchard & Bousquet, subgen. nov. (in Thalpophilodes Strand, 1942), Rhytistena Bouchard & Bousquet, subgen. nov. (in Rhytinota Eschscholtz, 1831), Spinosdara Bouchard & Bousquet, subgen. nov. (in Osdara Walker, 1858), Spongesmia Bouchard & Bousquet, subgen. nov. (in Adesmia Fischer, 1822), and Zambesmia Bouchard & Bousquet, subgen. nov. (in Adesmia Fischer, 1822). The names Adeps Gistel, 1857 and Adepsion Strand, 1917 syn. nov. [= Tetraphyllus Laporte & Brulle, 1831], Asyrmatus Canzoneri, 1959 syn. nov. [= Pystelops Gozis, 1910], Euzadenos Koch, 1956 syn. nov. [= Selenepistoma Dejean, 1834], Gondwanodilamus Kaszab, 1969 syn. nov. [= Conibius J.L. LeConte, 1851], Gyrinodes Fauvel, 1897 syn. nov. [= Nesotes Allard, 1876], Helopondrus Reitter, 1922 syn. nov. [= Horistelops Gozis, 1910], Hybonotus Dejean, 1834 syn. nov. [= Damatris Laporte, 1840], Iphthimera Reitter, 1916 syn. nov. [= Metriopus Solier, 1835], Lagriomima Pic, 1950 syn. nov. [= Neogria Borchmann, 1911], Orphelops Gozis, 1910 syn. nov. [= Nalassus Mulsant, 1854], Phymatium Billberg, 1820 syn. nov. [= Cryptochile Latreille, 1828], Prosoblapsia Skopin & Kaszab, 1978 syn. nov. [= Genoblaps Bauer, 1921], and Pseudopimelia Gebler, 1859 syn. nov. [= Lasiostola Dejean, 1834] are established as new synonyms (valid names in square brackets). Anachayus Bouchard & Bousquet, nom. nov. is proposed as a replacement name for Chatanayus Ardoin, 1957, Genateropa Bouchard & Bousquet, nom. nov. as a replacement name for Apterogena Ardoin, 1962, Hemipristula Bouchard & Bousquet, nom. nov. as a replacement name for Hemipristis Kolbe, 1903, Kochotella Bouchard & Bousquet, nom. nov. as a replacement name for Millotella Koch, 1962, Medvedevoblaps Bouchard & Bousquet, nom. nov. as a replacement name for Protoblaps G.S. Medvedev, 1998, and Subpterocoma Bouchard & Bousquet, nom. nov. is proposed as a replacement name for Pseudopimelia Motschulsky, 1860. Neoeutrapela Bousquet & Bouchard, 2013 is downgraded to a subgenus (stat. nov.) of Impressosora Pic, 1952. Anchomma J.L. LeConte, 1858 is placed in Stenosini: Dichillina (previously in Pimeliinae: Anepsiini); Entypodera Gerstaecker, 1871, Impressosora Pic, 1952 and Xanthalia Fairmaire, 1894 are placed in Lagriinae: Lagriini: Statirina (previously in Lagriinae: Lagriini: Lagriina); Loxostethus Triplehorn, 1962 is placed in Diaperinae: Diaperini: Diaperina (previously in Diaperinae: Diaperini: Adelinina); Periphanodes Gebien, 1943 is placed in Stenochiinae: Cnodalonini (previously in Tenebrioninae: Helopini); Zadenos Laporte, 1840 is downgraded to a subgenus (stat. nov.) of the older name Selenepistoma Dejean, 1834. The type species [placed in square brackets] of the following available genus-group names are designated for the first time: Allostrongylium Kolbe, 1896 [Allostrongylium silvestre Kolbe, 1896], Auristira Borchmann, 1916 [Auristira octocostata Borchmann, 1916], Blapidocampsia Pic, 1919 [Campsia pallidipes Pic, 1918], Cerostena Solier, 1836 [Cerostena deplanata Solier, 1836], Coracostira Fairmaire, 1899 [Coracostira armipes Fairmaire, 1899], Dischidus Kolbe, 1886 [Helops sinuatus Fabricius, 1801], Eccoptostoma Gebien, 1913 [Taraxides ruficrus Fairmaire, 1894], Ellaemus Pascoe, 1866 [Emcephalus submaculatus Breme, 1842], Epeurycaulus Kolbe, 1902 [Epeurycaulus aldabricus Kolbe, 1902], Euschatia Solier, 1851 [Euschatia proxima Solier, 1851], Heliocaes Bedel, 1906 [Blaps emarginata Fabricius, 1792], Hemipristis Kolbe, 1903 [Hemipristis ukamia Kolbe, 1903], Iphthimera Reitter, 1916 [Stenocara ruficornis Solier, 1835], Isopedus Stein, 1877 [Helops tenebrioides Germar, 1813], Malacova Fairmaire, 1898 [Malacova bicolor Fairmaire, 1898], Modicodisema Pic, 1917 [Disema subopaca Pic, 1912], Peltadesmia Kuntzen, 1916 [Metriopus platynotus Gerstaecker, 1854], Phymatium Billberg, 1820 [Pimelia maculata Fabricius, 1781], Podoces Peringuey, 1886 [Podoces granosula Peringuey, 1886], Pseuduroplatopsis Pic, 1913 [Borchmannia javana Pic, 1913], Pteraulus Solier, 1848 [Pteraulus sulcatipennis Solier, 1848], Sciaca Solier, 1835 [Hylithus disctinctus Solier, 1835], Sterces Champion, 1891 [Sterces violaceipennis Champion, 1891] and Teremenes Carter, 1914 [Tenebrio longipennis Hope, 1843]. Evidence suggests that some type species were misidentified. In these instances, information on the misidentification is provided and, in the following cases, the taxonomic species actually involved is fixed as the type species [placed in square brackets] following requirements in Article 70.3 of the International Code of Zoological Nomenclature: Accanthopus Dejean, 1821 [Tenebrio velikensis Piller & Mitterpacher, 1783], Becvaramarygmus Masumoto, 1999 [Dietysus nodicornis Gravely, 1915], Heterophaga Dejean, 1834 [Opatrum laevigatum Fabricius, 1781], Laena Dejean, 1821, [Scaurus viennensis Sturm, 1807], Margus Dejean, 1834 [Colydium castaneum Herbst, 1797], Pachycera Eschscholtz, 1831 [Tenebrio buprestoides Fabricius, 1781], Saragus Erichson, 1842 [Celibe costata Solier, 1848], Stene Stephens, 1829 [Colydium castaneum Herbst, 1797], Stenosis Herbst, 1799 [Tagenia intermedia Solier, 1838] and Tentyriopsis Gebien, 1928 [Tentyriopsis pertyi Gebien, 1940]. The following First Reviser actions are proposed to fix the precedence of names or nomenclatural acts (rejected name or act in square brackets): Stenosis ciliaris Gebien, 1920 as the type species for Afronosis G.S. Medvedev, 1995 [Stenosis leontjevi G.S. Medvedev, 1995], Alienoplonyx Bremer, 2019 [Alienolonyx], Amblypteraca Mas-Peinado, Buckley, Ruiz & Garcia-Paris, 2018 [Amplypteraca], Caenocrypticoides Kaszab, 1969 [Caenocripticoides], Deriles Motschulsky, 1872 [Derilis], Eccoptostira Borchmann, 1936 [Ecoptostira], †Eodromus Haupt, 1950 [†Edromus], Eutelus Solier, 1843 [Lutelus], Euthriptera Reitter, 1893 [Enthriptera], Meglyphus Motschulsky, 1872 [Megliphus], Microtelopsis Koch, 1940 [Extetranosis Koch, 1940, Hypermicrotelopsis Koch, 1940], Neandrosus Pic, 1921 [Neoandrosus], Nodosogylium Pic, 1951 [Nodosogilium], Notiolesthus Motschulsky, 1872 [Notiolosthus], Pseudeucyrtus Pic, 1916 [Pseudocyrtus], Pseudotrichoplatyscelis Kaszab, 1960 [Pseudotrichoplatynoscelis and Pseudotrichoplatycelis], Rhydimorpha Koch, 1943 [Rhytimorpha], Rhophobas Motschulsky, 1872 [Rophobas], Rhyssochiton Gray, 1831 [Ryssocheton and Ryssochiton], Sphaerotidius Kaszab, 1941 [Spaerotidius], Stira Agassiz, 1846 (Mollusca) [Stira Agassiz, 1846 (Coleoptera)], Sulpiusoma Ferrer, 2006 [Sulpiosoma] and Taenobates Motschulsky, 1872 [Taeniobates]. Supporting evidence is provided for the conservation of usage of Cyphaleus Westwood, 1841 nomen protectum over Chrysobalus Boisduval, 1835 nomen oblitum.

Review of genus-group names in the family Tenebrionidae (Insecta, Coleoptera).

This study demonstrates the power of DNA barcoding to detect overlooked and newly arrived taxa. Sixty-three species of Coleoptera representing 25 families are studied based on DNA barcode data and morphological analysis of the barcoded specimens. Three of the species involve synonymies or previous taxonomic confusion in North America, while the first Canadian records are published for 60 species. Forty-two species are adventive in North America, and 40 of these adventive species originate from the Palaearctic region. Three genera are recorded from the Nearctic region for the first time: Coelostoma Brulle, 1835 (Hydrophilidae), Scydmoraphes Reitter, 1891 (Staphylinidae), and Lythraria Bedel, 1897 (Chrysomelidae). Two new synonymies are established: Mycetoporus triangulatus Campbell, 1991 (Staphylinidae) is a junior synonym of Mycetoporus reichei Pandelle, 1869, syn. nov. while Bledius philadelphicus Fall, 1919 (Staphylinidae) is a junior synonym of Bledius gallicus (Gravenhorst, 1806), syn. nov. The previously suggested move of Ctenicera tigrina (Fall, 1901) to the genus Pseudanostirus Dolin, 1964 (Elateridae) is formalized, resulting in Pseudanostirus tigrinus (Fall, 1901), comb. nov.

DNA barcodes reveal 63 overlooked species of Canadian beetles (Insecta, Coleoptera).

The taxonomic concepts of Blapimorpha and Opatrinae (informal and traditional, morphology-based groupings among darkling beetles) are tested using molecular phylogenetics and a reassessment of larval and adult morphology to address a major phylogeny-classification gap in Tenebrionidae. Instead of a holistic approach (family-level phylogeny), this study uses a bottom-up strategy (tribal grouping) in order to define larger, monophyletic lineages within Tenebrioninae. Sampling included representatives of 27 tenebrionid tribes: Alleculini, Amarygmini, Amphidorini, Blaptini, Bolitophagini, Branchini, Cerenopini, Coniontini, Caenocrypticini, Dendarini, Eulabini, Helopini, Lagriini, Melanimini, Opatrini, Pedinini, Phaleriini, Physogasterini, Platynotini, Platyscelidini, Praociini, Scaurini, Scotobiini, Tenebrionini, Trachyscelini, Triboliini and Ulomini. Molecular analyses were based on DNA sequence data from four non-overlapping gene regions: carbamoyl-phosphate synthetase domain of rudimentary (CAD) (723 bp), wingless (wg) (438 bp) and nuclear ribosomal 28S (1101 bp) and mitochondrial ribosomal 12S (363 bp). Additionally, 15 larval and imaginal characters were scored and subjected to an ancestral state reconstruction analysis. Results revealed that Amphidorini, Blaptini, Dendarini, Pedinini, Platynotini, Platyscelidini and Opatrini form a clade which can be defined by the following morphological features: adults—antennae lacking compound/stellate sensoria; procoxal cavities externally and internally closed, intersternal membrane of abdominal ventrites 3–5 visible; paired abdominal defensive glands present, elongate, not annulated; larvae—prolegs enlarged (adapted for digging); ninth tergite lacking urogomphi. To accommodate this monophyletic grouping (281 genera and ∼4000 species), the subfamily Blaptinae sens. nov. is resurrected. Prior to these results, all of the tribes within Blaptinae were classified within the polyphyletic subfamily Tenebrioninae. The non-monophyletic nature of Terebrioninae has already been postulated by previous authors, yet no taxonomic decisions were made to fix its status. The reinstatement of Blaptinae, which groups ∼50% of the former Tenebrioninae, helps to clarify phylogenetic relations among the whole family and is the first step towards a complete higher-level revision of Tenebrionidae. The Central Asian tribe Dissonomini (two genera, ∼30 species) was not included in Blaptinae due to a lack of representatives in the performed phylogenetic analyses; however, based on morphological features, the tribe is listed as a potential addition to the subfamily. © 2020 The Royal Entomological Society

J Milton Campbell

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TENEBR10NIDAE Latreille 1802