Dysarchus tuberculatus, from Matthews & Bouchard (2008). Scale bar = 5 mm.

Belongs within: Tenebrionidae.
Contains: Helea, Pterohelaeus, Celibe, Saragus, Boreosaragus, Sympetes, Nyctozoilus, Cyphaleus, Platyphanes.

The Heleini are an Australasian tribe of relatively large tenebrionid beetles with simple antennal sensilla and unlobed tarsomeres. Members of the subtribe Asphalina are very similar to the related tribe Tenebrionini, being non-metallic in coloration and usually with nine primary striae on the elytra (Matthews & Bouchard 2008).

Characters (from Matthews & Bouchard 2008): Oblong, ovoid or subcircular in outline, glabrous or pilose, piceous, fuscous or metallic. Total length 6–35 mm. Edge of clypeus either more or less straight in middle or excised. Eyes transverse, reniform. Labrum transverse, basal membrane visible or not, medial tormal arms of epipharynx without anterior and posterior mesal extensions or with latter only, epitorma present, setal pattern symmetrical. Antennae variable in length, from not reaching base of prothorax to reaching well beyond it, segments basally more or less elongate becoming shorter and subtriangular distally, only a little widened there, with simple sensilla. Corpotentorium straight. Mandibles apically bidentate or truncate, molar surface with or without fine striations. Maxilla with or without lacinial uncus, palpi with apical segment more or less triangular. Pronotum variable, sometimes explanate laterally. Prosternum moderately long before coxae. Procoxal cavities usually internally open. Mesocoxal cavities partly closed by mesepimeron. Metendosternite without laminae, with short stem. Elytra primitively with nine striae and scutellary striole, often becoming estriate or with secondary striae, epipleura complete to apex. Wings present or absent, venation either unmodified or with CuA1 interrupted and usually a new crossvein between MP3+4 and CuA1+2, medial fleck present or absent, with diffuse apical fleck. Legs either unmodified and cursorial or variously modified for digging, if the latter, then often with enlarged spur of protibia (forespur), tarsomeres subtriangular, plantar surfaces densely setose or tomentose; more coarsely bristled in genera that live in burrows or caves. Abdomen with defensive reservoirs usually short and conical with lateral stiffening sclerites or prongs of cuticle emerging from sternite VII, but frequently without prongs, allowing them to be everted, and becoming either reduced or increased in size, without common volume. Secretory gland fields vary considerably, although always associated with bases of reservoirs. Ovipositor with coxite lobe 4 shortened or fused with 3, not digitate, and if evident, articulated at end of lobe 3 or reduced to small lateral sclerite on lobe 3, gonostyles lateral, primitive ovipositor with coxites about equal in length to paraprocts, occasionally with paraprocts becoming more or less elongated or becoming strongly sclerotised and toothed. Female tract without bursa, with single coiled spermathecal tubule arising at base of filamentous accessory gland, or separately at anterior edge of vagina. Aedeagus with basal piece generally longer than apical, sclerotised rods of median lobe often becoming enlarged or distinctively shaped, apical piece often setose or spinulose.

<==Heleini [Helaeinae]
    |    |--Sloanea Carter 1916MB08
    |    |    `--*S. costata Carter 1916MB08
    |    |--Asphalus Pascoe 1868MB08
    |    |    |--*A. ebenicus Pascoe 1868MB08
    |    |    `--A. striatus Carter 1911MB08
    |    |--Bassianus Matthews & Doyen 1989MB08
    |    |    |--*B. colydioides (Erichson 1842) (see below for synonymy)MB08
    |    |    |--B. humilis (Erichson 1842) [=Tenebrio humilis]MB08
    |    |    |--B. rectibasis (Carter 1914) [=Menephilus rectibasis]MB08
    |    |    `--B. sydneyanus (Blackburn 1893) [=Menephilus sydneyanus]MB08
    |    `--Meneristes Pascoe 1869 [incl. Asiris Motschulsky 1872]MB08
    |         |--M. australis (Boisduval 1835) (see below for synonymy)MB08
    |         |--M. latior Carter 1914MB08
    |         |--M. porosus Carter 1918MB08
    |         |--M. proximus Carter 1914 [=M. tibialis var. proximus]MB08
    |         `--M. tibialis Carter 1914MB08
    |    |--HeleaB14
    |    |--PterohelaeusMB08
    |    |--CelibeMB08
    |    |--SaragusMB08
    |    |--BoreosaragusMB08
    |    |--SympetesMB08
    |    |--Camponotiphilus Lea 1914 [=Campanotiphilus (l. c.)]MB08
    |    |    `--*C. fimbricollis Lea 1914MB08
    |    |--Edylius Champion 1894MB08
    |    |    `--*E. canescens Champion 1894MB08
    |    |--Trichosaragus Blackburn 1890MB08
    |    |    |--*T. pilosellus Blackburn 1890MB08
    |    |    |--T. convexus (Carter 1919) [=Saragus convexus]MB08
    |    |    `--T. granulatus Carter 1926MB08
    |    |--MimopeusMB08
    |    |    |--M. costellusWH02
    |    |    |--M. elongatus (Brême 1842) [incl. M. amaroides Pascoe 1866]MB08
    |    |    |--M. lateralisWH02
    |    |    `--M. opaculusWH02
    |    |--Ospidus Pascoe 1866MB08
    |    |    |--*O. chyrysomeloides Pascoe 1866 [incl. O. paropsoides Carter 1909]MB08
    |    |    |--O. gibbus Blackburn 1897MB08
    |    |    |--O. major Carter 1920MB08
    |    |    `--O. vestitus Carter 1919MB08
    |    |--Cillibus Matthews 1993MB08
    |    |    |--*C. blackburni (Macleay 1888) [=Saragus blackburni]MB08
    |    |    |--C. convexicollis (Macleay 1888) [=Saragus convexicollis]MB08
    |    |    |--C. incisus (Pascoe 1870) [=Saragus incisus]MB08
    |    |    `--C. ovalis (Macleay 1872) [=Saragus ovalis]MB08
    |    |--Emcephalus Kirby 1828 (see below for synonymy)MB08
    |    |    |--*E. gibbosus Kirby 1828MB08 (see below for synonymy)
    |    |    |--E. camelus (Carter 1911) [=Encara camelus]MB08
    |    |    |--E. floccosus (Pascoe 1870) [=Saragus floccosus]MB08
    |    |    |--E. lacordairei (Macleay 1887) [=Encara lacordairei]MB08
    |    |    |--E. nigrus (Carter 1905) [=Encara nigra]MB08
    |    |    |--*Euhelaeus’ speculiferus Gebien 1920MB08
    |    |    `--E. submaculatus Brême 1842MB08 [=Encephalus submaculatusMB08, Encara submaculatumMas86]
    |    |--Brises Pascoe 1869 [incl. Ephidonius Pascoe 1869]MB08
    |    |    |--*B. trachynotoides Pascoe 1869MB08
    |    |    |--B. acuticornis (Pascoe 1869) (see below for synonymy)MB08
    |    |    |--B. blairi Carter 1914MB08
    |    |    |--B. caraboides Matthews 1986MB08
    |    |    |--B. katherinae Matthews 1986MB08
    |    |    |--B. nullarboricus Matthews 1986MB08
    |    |    |--B. occidentalis Matthews 1986MB08
    |    |    `--B. parvicollis (Blackburn 1895) [=Ephidonius parvicollis]MB08
    |    `--Dysarchus Pascoe 1866 [incl. Saragella Carter 1937, Saragodinus Bates 1872]MB08
    |         |--*D. odewahni Pascoe 1866MB08
    |         |--D. batesi (Haag-Rutenberg 1878) [=Saragodinus batesi]MB08
    |         |--D. browni Carter 1930MB08
    |         |--D. duboulayi (Bates 1872) [=*Saragodinus duboulayi]MB08
    |         |--D. howitti (Bates 1872) [=Saragodinus howitti]MB08
    |         |--D. irregularis Carter 1924MB08
    |         |--D. palpalis (Carter 1937) [=*Saragella palpalis]MB08
    |         |--D. percostatus Carter 1930MB08
    |         `--D. tuberculatus (Blackburn 1888) [=Saragodinus tuberculatus]MB08
    `--Cyphaleina [Cyphaleinae, Cyphaleini]MB08
         |--Amphianax Bates 1873MB08
         |    `--*A. subcoriaceus Bates 1873 [incl. Agasthenes euclensis Carter 1910, Ag. goudiei Carter 1909]MB08
         |--Amarygmimus Bates 1873 [=Amarygmomimus Rye 1875]MB08
         |    `--*A. duboulayi Bates 1873MB08
         |--Phanechloros Matthews & Bouchard 2008 [=Chlorophanes Matthews 1992 (preoc.)]MB08
         |    `--*P. punctipennis (Carter 1911) [=Platyphanes punctipennis. *Chlorophanes punctipennis]MB08
         |--Paraphanes Macleay 1887MB08
         |    `--*P. nitidus Macleay 1887MB08
         |--Onotrichus Carter 1911MB08
         |    |--*O. lateralis Carter 1911MB08
         |    `--O. minor Carter 1914MB08
         |--Mithippia Pascoe 1869MB08
         |    |--*M. aurita Pacoe 1869 [incl. Tarpela catenulata Allard 1877]MB08
         |    `--M. jansoni Bates 1873MB08
         |--Mitrothorax Carter 1914 (see below for synonymy)MB08
         |    |--*M. convexicollis (Carter 1913) [=*Mitrephorus convexicollis]MB08
         |    `--M. breweri (Bates 1873) [=*Ctimene breweri, *Timeneca breweri]MB08
         |--Agasthenes Bates 1873MB08
         |    |--*A. westwoodi Bates 1873MB08
         |    |--A. championi Carter 1910MB08
         |    `--A. frenchi Carter 1909MB08
         |--Lepispilus Westwood 1841 (see below for synonymy)MB08
         |    |--*L. sulcicollis (Boisduval 1835) [=*Pachycoelia sulcicollis]MB08
         |    |--L. ocularis Carter 1932MB08
         |    |--L. rotundicollis Blackburn 1889MB08
         |    `--L. stygianus Pascoe 1869 [incl. *Tyndarisus longitarsis Pascoe 1869]MB08
         |--Aglypta Gebien 1908 [=Onoglypta Carter 1926]MB08
         |    |--*A. octocostata Gebien 1908 [=*Onoglypta octocostata]MB08
         |    |--A. gebieni Carter 1911MB08
         |    |--A. parva (Carter 1911) [=Onosterrhus parvus]MB08
         |    `--A. rugosa (Carter 1937) [=Onoglypta rugosa]MB08
         |--Atoreuma Gebien 1941 [=Eutoreuma Carter 1914 (preoc.), Toreuma Carter 1913 (preoc.)]MB08
         |    |--*A. cupreum (Carter 1913) [=*Toreuma cupreum, *Eutoreuma cupreum]MB08
         |    |--A. minor (Carter 1924) [=Eutoreuma minor]MB08
         |    |--A. rufoaeneum (Carter 1914) [=Olisthaena rufoaeneum]MB08
         |    `--A. turneri (Carter 1933) [=Eutoreuma turneri]MB08
         |--Olisthaena Erichson 1842 (see below for synonymy)MB08
         |    |--*O. nitida Erichson 1842MB08 [=Chartopteryx nitidaMas86; incl. Decialma erichsoni Champion 1894MB08]
         |    |--O. anthracina (Pascoe 1869) [=*Hectus anthracinus, *Aphectus anthracinus]MB08
         |    |--O. sparsa Carter 1933MB08
         |    `--O. tenuitarsis (Pascoe 1869) [=*Decialma tenuitarsis; incl. D. pascoei Bates 1873]MB08
         |--Styrus Bates 1873MB08
         |    |--S. batesi Haag-Rutenberg 1879 [=*S. elongatulus Bates 1873 non Nyctozoilus elongatulus Macleay 1872]MB08
         |    |--S. clathratus Blackburn 1890MB08
         |    |--S. elongatulus (Macleay 1872) [=Nyctozoilus elongatulus]MB08
         |    |--S. latior Carter 1909MB08
         |    `--S. puncticollis Carter 1919MB08
         |--Prophanes Westwood 1849 [incl. Lygestira Pascoe 1866, Maerodes Waterhouse 1877, Moerodes Rye 1879]MB08
         |    |--*P. aculeatus Westwood 1849 [incl. P. westwoodi Macleay 1872, *Maerodes westwoodi]MB08
         |    |--P. cupreipennis Macleay 1886Mac86
         |    |--P. latus (Waterhouse 1877) [=Lygestira lata]MB08
         |    |--P. mastersiLB91
         |    |--P. simplex Westwood 1849MB08 [=Lygestira simplexMas86, incl. *L. funerea Pascoe 1866MB08]
         |    `--P. submetallicus Macleay 1886Mac86
         |--Hemicyclus Westwood 1841 [incl. Cyclophanes Carter 1913]MB08
         |    |--H. reaumuri (Laporte 1840) (see below for synonymy)MB08
         |    |--H. flavipes Carter 1913MB08
         |    |--H. punctulatus Pascoe 1866MB08
         |    |--H. sphaeroides Carter 1929MB08
         |    |--H. splendens (Carter 1913) [=Cyclophanes splendens]MB08
         |    `--H. variegatus (Carter 1913) [=*Cyclophanes variegatus]MB08
         |--Bolbophanes Carter 1913 [incl. Pseudobolbophanes Kulzer 1954]MB08
         |    |--*B. dumbrelli (Lea 1895) [=Paraphanes dumbrelli]MB08
         |    |--B. finschi Kulzer 1964MB08
         |    |--B. malaisei (Kulzer 1954) [=*Pseudobolbophanes malaisei]MB08
         |    |--B. pallidipes Carter 1929MB08
         |    |--B. rugatus Carter 1913MB08
         |    `--B. varicolor Carter 1913 [incl. B. sericatus Carter 1919]MB08
         `--Byallius Pascoe 1869MB08
              |--*B. reticulatus Pascoe 1869MB08
              |--B. angustatus Carter 1925MB08
              |--B. kosciuskoanus Carter 1908MB08
              |--B. laticollis Carter 1914MB08
              |--B. oberonius Carter 1933MB08
              |--B. ovensensis Carter 1909MB08
              |--B. punctatus Carter 1919MB08
              `--B. revolutus (Carter 1914) [=Styrus revolutus]MB08

*Bassianus colydioides (Erichson 1842) [=Tenebrio colydioides; incl. Menephilus armstrongi Carter 1933, M. parvulus Macleay 1872]MB08

Brises acuticornis (Pascoe 1869) [=*Ephidonius acuticornis; incl. E. duboulayi Bates 1872, Brises granulatus Carter 1921]MB08

Emcephalus Kirby 1828 [=Ellaemus Pascoe 1866, Encara Gemminger & Harold 1870, Encephalus Brême 1842 non Kirby 1832; incl. Euhelaeus Gebien 1920]MB08

*Emcephalus gibbosus Kirby 1828MB08 [=*Ellaemus gibbosusMB08, *Encara gibbosaMB08; incl. Cilibe westwoodi Boisduval 1835MB08, En. westwoodiMas86]

Hemicyclus reaumuri (Laporte 1840) [=Tetraphyllus reaumuri; incl. *H. grandis Westwood 1841, H. metallicus Westwood 1841]MB08

Lepispilus Westwood 1841 [=Lepidospilus Agassiz 1846, Pachycoelia Boisduval 1835 (preoc.); incl. Tyndarisus Pascoe 1869]MB08

Meneristes australis (Boisduval 1835) [=Tenebrio australis; incl. *Asiris angulicollis Motschulsky 1872, Meneristes intermedius Pascoe 1869, *M. laticollis Pascoe 1869, Tenebrio nigerrimus Blanchard 1853 non Boisduval 1835]MB08

Mitrothorax Carter 1914 [=Mitrephorus Carter 1913 (preoc.); incl. Ctimene Bates 1873 (preoc.), Timeneca Carter 1914]MB08

Olisthaena Erichson 1842 [=Olisthoena (l. c.); incl. Aphectus Carter 1926, Decialma Pascoe 1869, Hectus Pascoe 1869]MB08

*Type species of generic name indicated


[B14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature’s gems. Ivy Press: Lewes (United Kingdom).

[LB91] Lawrence, J. F., & E. B. Britton. 1991. Coleoptera (beetles). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2nd ed. vol. 2 pp. 543–683. Melbourne University Press: Carlton (Victoria).

[Mac86] Macleay, W. 1886. The insects of the Fly River, New Guinea, “Coleoptera”. Proceedings of the Linnean Society of New South Wales, series 2, 1 (1): 136–157.

[Mas86] Masters, G. 1886. Catalogue of the described Coleoptera of Australia. Part IV. Proceedings of the Linnean Society of New South Wales, series 2, 1 (2): 259–380.

[MB08] Matthews, E. G., & P. Bouchard. 2008. Tenebrionid Beetles of Australia: Descriptions of tribes, keys to genera, catalogue of species. Australian Biological Resources Study: Canberra.

[WH02] Worthy, T. H., & R. N. Holdaway. 2002. The Lost World of the Moa: Prehistoric life of New Zealand. Indiana University Press: Bloomington (Indiana).


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