Exangeltus gracilior, from Matthews & Bouchard (2008). Scale bar = 1 mm.

Belongs within: Tenebrionoidea.
Contains: Adesmia, Helops, Lagriinae, Phrenapatinae, Cossyphodini, Sepidiini, Pimeliini, Tentyria, Alleculinae, Diaperinae, Stenochiinae, Toxicini, Blaps, Amarygmini, Bolitophagini, Ulomini, Opatrini, Palorini, Heleini, Triboliini, Titaenini, Tenebrio.

The Tenebrionidae are a large and varied family of beetles, most of which are scavengers on dead plant and fungal material, and are one of the most successful families of beetles in arid environments. Many tenebrionids are able to discharge defensive chemicals from glands in the anal region, though these glands are absent in some significant subgroups such as the Pimeliinae. In those groups lacking defensive glands, the sternites of the abdomen hinge along the midline but those tenebrionids with defensive glands have sternites hinging laterally to allow the abdomen to expand before expelling the defensive fluid (Matthews & Bouchard 2008).

The Tenebrionidae may be divided between three major lineages, referred to by Matthews & Bouchard (2008) as the lagrioid, pimelioid and tenebrionoid branches, based primarily on features of the male and female reproductive systems. Members of the lagrioid branch include the Neotropical genus Nilio, false ladybird beetles, adults of which have a circular outline and a setose, nearly hemispherical dorsal surface (Bouchard 2014). The Pimeliinae are the most diverse subgroup of the tenebrionids in arid environments in most parts of the world except Australia (Matthews & Bouchard 2008).

Characters (from Lawrence & Britton 1991): Usually black or brown and subglabrous, with concealed antennal insertions and usually stout, moniliform or incrassate antennae. Eyes usually emarginate; lateral pronotal carinae usually simple and complete; prosternal process usually moderately broad, strongly curved dorsally behind coxae, neither strongly nor abruptly expanded apically; junctions of first three ventrites different from those separating last two, the basal ventrites being strongly connate and the apical ones more or less movable, often with exposed membrane. Externally open fore coxal cavities only occur in Tanylypa, internally open ones in some Tenebrioninae. Tarsal formula usually 5-5-4, sometimes 4-4-4. Most species with paired glandular reservoirs, associated with chemical defence sustem, located usually at base of fifth ventrite. Larvae almost always elongate (ovate in Leiochrini) and subcylindrical to slightly flattened. May be very lightly sclerotised, moderately sclerotised or yellow in colour, with or without darker head and urogomphi, or uniformly heavily sclerotised and darkly pigmented; vestiture consists of scattered fine hairs except in Lagriini whose larvae are densely setose.

Tenebrionidae [Cossyphores, Pimeliariae, Tenebrionites, Ulominae]
    |--+--Lagriinae MW15
    |  |--Phrenapatinae MB08
    |  `--Nilio Latreille 1802 L02 [=Nilion L02; Nilionidae, Nilioninae MB08]
    |       |--*N. villosus [=Coccinella villosa] L02
    |       `--N. lanatus Germar 1824 B14
    `--+--+--Zolodininae [Zolodinini] MW15
       |  |    |--Zolodinus zelandicus LB91
       |  |    `--Tanylypa Pascoe 1869 MB08
       |  |         `--*T. morio Pascoe 1869 MB08
       |  `--Pimeliinae [Tentyriinae] MW15
       |       |  i. s.: Onymacris B14
       |       |           |--O. bicolor (Haag-Butenberg 1875) B14
       |       |           |--O. laevigatus B14
       |       |           `--O. unguicularis B14
       |       |         Bothroles C68
       |       |--Cossyphodini B14
       |       |--Sepidiini B14
       |       |--Ceratanisini LB91
       |       |--Pimeliini B14
       |       |--Lachnogyini MB08
       |       |--Salax Guérin-Méneville 1834 [Trilobocarini] MB08
       |       |    `--*S. lacordairei Guérin-Méneville 1834 MB08
       |       |--Nyctoporis [Nyctoporini] MB08
       |       |    `--N. cristata Eschscholtz 1831 [incl. Emeax sculpturatus Pascoe 1866] MB08
       |       |--Coelus [Coniontini] B14
       |       |    |--C. ciliatus MW15
       |       |    |--C. globosus LeConte 1852 B14
       |       |    |--C. gracilis B14
       |       |    `--C. maritimus B14
       |       |--Cryptochilini B14
       |       |    |--Vansonium B14
       |       |    `--Calognathus chevrolati B14
       |       |         |--C. c. chevrolati B14
       |       |         `--C. c. eberlanzi Koch 1950 B14
       |       |--Nycteliini B14
       |       |    |--Entomobalia B14
       |       |    `--Nyctelia B14
       |       |         |--N. geometrica Fairmaire 1905 B14
       |       |         `--N. westwoodi B14
       |       |--Tentyriini MB08
       |       |    |--Tentyria K03
       |       |    `--Stenosida Solier 1835 MB08
       |       |         |--‘Tagenia’ striatopunctata Wiedemann 1821 [incl. *Stenosida tenuicollis Solier 1835] MB08
       |       |         |--S. australis Medvedev 1995 MB08
       |       |         `--S. darevskii MB08
       |       |--Cnemeplatini MB08
       |       |    |  i. s.: Actizeta MB08
       |       |    `--Thorictosomatina MB08
       |       |         |--Wattiana Matthews & Lawrence 2005 MB08
       |       |         |    `--*W. greensladei Mattthews & Lawrence 2005 MB08
       |       |         `--Thorictosoma Lea 1919 GWW10, L19
       |       |              |--*T. ectatommae Lea 1919 L19
       |       |              `--T. tibiale Lea 1919 L19
       |       `--Vacronini MB08
       |            |--Eupsophulus MB08
       |            |--Lixionica Blackburn 1896 MB08
       |            |    `--*L. costatipennis Blackburn 1896 MB08
       |            `--Exangeltus Blackburn 1897 [=Exangelutus (l. c.)] MB08
       |                 |--*E. angustus Blackburn 1897 MB08
       |                 |--E. brevicollis Kulzer 1964 MB08
       |                 |--E. gracilior Blackburn 1903 MB08
       |                 `--E. rufipennis Carter 1919 MB08
       `--+--Trachelostenus [Trachelostenidae] MW15
          |--Alleculinae MB08
          |--Diaperinae MB08
          |--Stenochiinae MB08
          `--Tenebrioninae [Opatrinae] MW15
               |  i. s.: Merinus C68
               |         Eleodes B14
               |           |--E. (Eleodes) acutus (Say 1824) B14
               |           |--E. caudifera C90
               |           |--E. hispilabris C90
               |           `--E. obscurus B14
               |         Rhyzodina mniszechii Chevrolat 1873 B14
               |         Idiobates MB08
               |--Toxicini LB91
               |--Blaps LB91
               |--Amarygmini LB91
               |--Bolitophagini B14
               |--Ulomini LB91
               |--Opatrini LB91
               |--Palorini MB08
               |--Cheirodes Gené 1839 [Melanimini] MB08
               |    `--*C. sardous Gené 1839 [incl. Anemia caulobioides Carter 1920] MB08
               |--Platynotini [Platynotina] MB08
               |    |  i. s.: ‘Tenebrio’ dispar Herbst 1797 [incl. Menearchus impressosulcatus Carter 1920] MB08
               |    `--Anomalipus B14
               |         |--A. elephas Fåhraeus 1870 B14
               |         `--A. expansicollis G91
               |--Pedinini B14
               |    |--Leichenum Dejean 1834 [incl. Endothina Carter 1924, Endothyna (l. c.); Leichenina, Leichenini] MB08
               |    |    |--*L. pictum (Fabricius 1801) [=Opatrum pictum] MB08
               |    |    |--L. canaliculatum (Fabricius 1798) (see below for synonymy) MB08
               |    |    `--L. pulchellum G89
               |    `--Pedinus Latreille 1802 L02
               |         |--P. dermestoides [=Blaps dermestoides] L02
               |         |--P. femoralis [=Blaps femorale] L02
               |         |--P. glaber G20
               |         `--P. strigosus C01
               `--+--Heleini MB08
                  |--Triboliini MB08
                  |--Titaenini MB08
                  |--Acropteron Perty 1830 FS90 [Acropteronini MB08]
                  |    |--A. chabrieri Fleutiaux & Sallé 1890 FS90
                  |    `--A. transversicolle FS90
                  |--Tenebrionini MB08
                  |    |--Tenebrio B14
                  |    `--Paratoxicum Champion 1894 [incl. Schizophthalmotribolium Kaszab 1940] MB08
                  |         |--*P. iridescens Champion 1894 MB08
                  |         |--P. australiae (Kaszab 1940) [=*Schizophthalmotribolium australiae] MB08
                  |         `--P. nigricans Oke 1932 MB08
                  `--Alphitobiini MB08
                       |--Diaclina Jacquelin du Val 1861 MB08
                       |    |--*D. chrysomelina (Carter 1908) [=Tenebrio chrysomelinus] MB08
                       |    `--D. nitida (Carter 1908) [=Heterocheira nitida; incl. D. immaculata Gebien 1920] MB08
                       `--Alphitobius Stephens 1829 [incl. Microphyes Macleay 1872] MB08
                            |--A. laevigatus (Fabricius 1781) (see below for synonymy) MB08
                            |--A. diaperinus (Panzer 1797) [=Tenebrio diaperinus] MB08
                            `--A. piceus Ol. 1795 [incl. A. convexus, A. fagi, A. manillensis, A. orizae] FS90

Tenebrionidae incertae sedis:
  Zeadelium gratiosum ED04
  Stomium BHM08
  Parastizopus armaticeps GE05
  Ulomites GE05
  Pseudognaptorina Kaszab 1977 K77
    `--*P. nepalica Kaszab 1977 K77
  Freudeia Kaszab 1961 K77
    |--F. granulipleuris Kaszab 1977 K77
    |--F. martensi Kaszab 1977 K77
    |--F. nepalica Kaszab 1961 K77
    `--F. punctipleuris Kaszab 1977 K77
  Blaptyscelis Koch 1961 [=Blaptyscellis Pierre 1961] K77
    |--B. alpinus (Kaszab 1965) K77
    |--B. costulatus (Fairmaire 1901) K77
    |--B. dreuxi Pierre 1961 K77
    |--B. nepalicus (Kaszab 1973) [=Tagenoides nepalicus] K77
    |--B. subcarinatus (Blair 1927) K77
    `--B. zurstrasseni Kaszab 1977 K77
  Platolenes nepalensis Kaszab 1977 K77
  Heterotarsus K77
    |--H. auriculatus Marseul 1876 K77
    `--H. pustulifer F89
  Agnaptoria Reitter 1887 K77
    |--A. rubripes Reitter 1887 K77
    |--A. seidlitzi Reitter 1893 K77
    `--A. subcarinata Blair 1927 K77
  Platynoscelis (Leipopleura) politicollis Kaszab 1940 K77
  Hexarhopalus entomogonoides (Allard 1896) [=Ptilonix entomogonoides] K77
  Zophosis Latreille 1802 L02
    |--*Z. testudinaria [=Erodius testudinarius] L02
    |--Z. abbreviata Sol. 1834 G89
    |--Z. acuticosta Fairm. 1885 G89
    |--Z. plana [=Erodius planus] G89
    |--Z. punctata Brullé 1832 G89
    |--Z. pygmaea Sol. 1834 G89
    |--Z. quadricostata Sol. 1834 G89
    |--Z. sabaea Baudi 1881 G89
    `--Z. sulcata Deyr. 1867 G89
  Arthrodes G89
    |--A. impressifrons Baudi 1881 G89
    `--A. rotundatus Sol. 1834 G89
  Erodius G89
    |--E. gibbus L02
    `--E. puncticollis Sol. 1834 G89
  Adesmia G89
  Rhytidonota praelonga (Reiche 1850) [=Axumia praelonga] G89
  Mesostena G89
    |--M. laevicollis Sol. 1835 G89
    |--M. puncticollis Sol. 1835 G89
    `--M. punctipennis Sol. 1835 G89
  Mesostenopa arabica Gestro 1881 G89
  Micipsa philistina Reiche 1857 G89
  Oxycara G89
    |--O. hegeterica (Reiche 1857) [=Melanerus hegetericus] G89
    `--O. pygmaea (Reiche 1857) [=Melanerus pygmaeus] G89
  Sceleodis castaneus [=Cratopus castaneus] G89
  Phaeotribon pulchellus G89
  Himatismus villosus Haag 1870 G89
  Pogonobasis ornata Sol. 1837 G89
  Peristeptus gestroi Haag 1875 G89
  Stenosis costulata Baudi 1881 G89
  Eutagenia arabs (Baudi 1881) [=Stenosis arabs] G89
  Thriptera G89
    |--T. crinita (Klug 1830) [=Pimelia crinita] G89
    `--T. murina Baudi 1881 G89
  Gedeon abyssinicus G89
  Hopatrinus G89
    |--H. corvinus Muls 1852 G89
    `--H. niloticus Muls 1852 G89
  Scleron G89
    |--S. orientale (Fabr. 1775) [=Opatrum orientale] G89
    `--S. sulcatum Baudi 1876 G89
  Hopatrum G89
    |--H. famelicum Oliv. 1811 G89
    |--H. hispidum Brullé 1838 G89
    |--H. lugens G89
    |--H. micans G89
    |--H. murinum Baudi 1876 G89
    |--H. rusticum Oliv. 1811 G89
    `--H. sericeum Baudi 1876 G89
  Halonomus subplumbeus (Fairm. 1856) [=Platydema subplumbea] G89
  Hopatroides punctulatus G89
  Calcar raffrayi Fairm. 1873 G89
  Dilamus pictus Baudi 1881 G89
  Micrantereus G89
    |--M. gerstaeckeri G89
    |--M. rugulosus Gestro 1878 G89
    `--M. tentyrioides G89
  Helopinus G89
    |--H. elegans Baudi 1881 G89
    `--H. psalidiformis Ancey 1881 G89
  Praeugena gagatina Mäklin 1864 G89
  Boletoxenus bellicum K98
  Philolithus densicollis [=Pelecyphorus densicollis] C90
  Stenomorpha C90
  Cryptoglossa C90
  Neatus tenebrioides LKW09
  Anatolica C01
    |--A. iduna C01
    |--A. paradoxa C01
    `--A. sulcipennis Reitt. in Csiki 1901 C01
  Melanesthes C01
    |--M. ciliata C01
    |--M. heydeni Csiki 1901 C01
    |--M. maxima C01
    `--M. mongolica Csiki 1901 C01
  Melaxumia angulosa C01
  Scytosoma arcibasis C01
  Microdera interrupta C01
  Dordanea kraatzi C01
  Trigonoscelis C01
    |--T. kraatzi C01
    |--T. laeviuscula C01
    |--T. reitteri Csiki 1901 C01
    |--T. seidlitzi C01
    `--T. zichyi Csiki 1901 C01
  Platyscelis rugifrons C01
  Dendarus C01
    |--D. barbarus Lucas 1847 E12
    |--D. extensus C01
    |--D. rotundicollis Lucas 1847 E12
    `--D. subvariolosus Lucas 1847 E12
  Scleropatrum C01
    |--S. carinatum C01
    `--S. tuberculiferum C01
  Praociini B14
  Ammophorus [Scotobiini] MB08
    `--A. bifoveatus PP72
  Gonocnemis B14
  Asida Latreille 1802 L02
    |--*A. grisea [=Opatrum griseum] L02
    |--A. affinis Lucas 1847 E12
    |--A. complanata Lucas 1847 E12
    |--A. lapidaria Lucas 1847 E12
    `--A. luctosa Boisduval 1835 B35
  Akis L02
    |--A. (Eurychora) ciliata G20
    |--A. (Akis) collaris G20
    `--A. reflexa L02
  Tagenia Latreille 1802 L02
    |--*T. filiformis [=Akis filiformis] L02
    `--T. algirica Lucas 1846 E12
  Scaurus striatus L02
  Hegeter Latreille 1802 L02
    |--*H. elongatus [=Blaps elongata] L02
    `--H. striatus G20
  Orthocerus Latreille 1802 L02
    |--O. histricornis G20
    `--O. muticus [=Sarrotrium muticum] L02
  ‘Silvanus’ costatus Steinheil 1869 (n. d.) H73
  Metaclisa [Centronopini] MB08
  Damatris MB08
  Baryscelis MB08
    |--B. laticollis Boisduval 1835 (n. d.) MB08
    `--B. politus Boisduval 1835 (n. d.) MB08 [=Meneristes politus M86]
  Chrysobalus fulgidipennis Boisduval 1835 (n. d.) MB08 [=Cyphaleus fulgidipennis M86]
  Helops MB08
  Pseudostrongylium viridipenne Kraatz 1880 (n. d.) MB08
  Tetraphyllus sumptuosus Hope 1843 (n. d.) MB08
  Brysax M86
    |--B. egenus Pascoe 1866 M86
    |--B. macleayi Pascoe 1866 M86
    `--B. saccharatus Pascoe 1870 M86
  Ulodica hispida Pascoe 1869 M86
  Elascus M86
    |--E. crassicornis Pascoe 1860 M86
    `--E. lunatus Pascoe 1860 M86
  Ennebaeus ovalis Waterh. 1878 M86
  Typhobia Pascoe 1869 M86, BR05
  Nyctobates crenata Boisd. 1835 M86
  Menephilus M86
    |--M. colydioides Erichs. 1842 M86
    |--M. convexiusculus Hope 1842 M86
    |--M. corvinus Erichs. 1842 M86
    |--M. cyanipennis Hope 1842 M86
    |--M. humilis Erichs. 1842 M86
    |--M. lonipennis Hope 1842 M86
    `--M. nigerrimus [incl. M. australis] M86
  Acrothymus caenosus Pascoe 1866 M86
  Embaphion RD77
  Misolampus hofmannseggii G20
  Hesiodobates antiquus Kaszab & Schawaller 1984 P92
  Stibia hannai Blaisdell 1925 A72
  Stomion PP72
    |--S. galapagoensis PP72
    |--S. laevigatum PP72
    `--S. rugosum PP72
  Pedonoeces PP72
    |--P. bauri PP72
    `--P. opacus PP72
  Trientoma Sol. 1835 FS90
    |--T. guadeloupensis Fleutiaux & Sallé 1890 FS90
    `--T. sallei FS90
  Diastolinus Muls & Rey 1859 FS90
    `--D. perforatus Sahlb. 1823 FS90
  Opatrinus Latr. 1829 FS90
    `--O. gemellatus Ol. 1795 [incl. O. clathratus] FS90
  Blapstinus Latr. 1829 FS90
    `--B. opacus Muls. 1859 FS90
  Ulus Horn 1870 [incl. Scaptes] FS90
    `--U. squamulatus Champ. 1886 FS90
  Hesiodus Champ. 1885 FS90
    |--H. caraibus Fleutiaux & Sallé 1890 FS90
    `--H. longitarsis FS90
  Allegoria Cast. 1840 FS90
    |--A. castelnaui Fleutiaux & Sallé 1890 FS90
    `--A. dilatata FS90
  Antimachus Gistl 1829 FS90
    |--A. divergens FS90
    `--A. roudeni Petit in Fleutiaux & Sallé 1890 FS90
  Zophobas Blanch. 1845 FS90
    `--Z. morio Fab. 1777 [incl. Z. elongatus, Z. infuscatus, Z. nigritus] FS90
  Sitophagus Muls. 1854 FS90
    `--S. hololeptoides Cast. 1840 (see below for synonymy) FS90
  Anaedus Blanch. 1845 FS90
    `--A. delauneyi Fleutiaux & Sallé 1890 FS90
  Cyrtosoma Perty 1830 FS90
    `--C. lherminieri Chv. 1838 FS90
  Pyanisia Cast. 1840 FS90
    `--P. undata Fab. 1792 [incl. P. melanura] FS90
  Talanus Duv. 1857 FS90
    |--T. guadeloupensis Fleutiaux & Sallé 1890 FS90
    `--T. subexaratus FS90
  Lobopoda Sol. 1835 FS90
    |--L. jalapensis FS90
    `--L. tarsalis Chv. in Fleutiaux & Sallé 1890 FS90

Alphitobius laevigatus (Fabricius 1781) [=Opatrum laevigatum; incl. Helops picipes Panzer 1794, *Alphitobius picipes, *Microphyes rufipes Macleay 1872] MB08

Leichenum canaliculatum (Fabricius 1798) [=Opatrum canaliculatum; incl. *Endothina squamosa Carter 1924] MB08

Sitophagus hololeptoides Cast. 1840 [incl. S. castaneus Reitt. 1877, S. complanatus, S. farinarius Woll. 1858, S. solieri Muls. 1854] FS90

*Type species of generic name indicated


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