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Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Coleopterida
Ordo: Coleoptera
Subordo: Polyphaga
Infraordo: Cucujiformia
Superfamilia: Tenebrionoidea

Familia: Tenebrionidae
Subfamilia: Tenebrioninae
Tribus (30): Acropteronini - Alphitobiini - Amarygmini - Amphidorini - Apocryphini - Blaptini - Bolitophagini - Centronopini - Cerenopini - Cyphaleini - Dissonomini - Eulabini - Heleini - Helopini - Helopinini - Melanimini - Opatrini - Palorini - Paoligenini - Pedinini - Platyscelidini - Praeugenini - Rhysopaussini - Scaurini - Scotobiini - Tenebrionini - Titaenini - Toxicini - Triboliini - Ulomini - incertae sedis

Name

Tenebrioninae Latreille, 1802
Synonyms

Amarygminae

References
cited sources

Bouchard, P., Lawrence, J.F., Davies, A.E. & Newton, A.F. 2005. Synoptic classification of the world Tenebrionidae (Insecta: Coleoptera) with a review of family-group names. Annales Zoologici 55(4): 499–530. PDF. Reference page.

Links

Zicha, Ondřej et al. Tenebrioninae – Taxon details on Biological Library (BioLib).

Tenebrioninae is the largest subfamily of the darkling beetles (Tenebrionidae), containing flour beetles, among others. Tenebrioninae contains more than 20 tribes.

Description
Larvae of Alphitobius laevigatus (Alphitobiini)
Adults

Adults are robust, mid-sized beetles that typically have elytra with some sort of corrugation on the upper side. They are typically black, dark brown or grey, and often have a satiny sheen. The body is shaped like a medication capsule or like a bullet; the legs can be short and stout or long and spindly. They eat both fresh and decaying vegetation, including vegetable produce, and several are commercially important pests of flour and other cereal products.

The subfamily has been characterized[1] as adults having mandibles with the back opposite the cutting edge, without margination and excavated opposite the molar pait; having ocelli arranged in two transverse, crescent shaped or circular groups on each side of head, and with five more or less fused lenses; having antennae with basal articles noticeably longer than wide; having pygidium that is apically bicomute; and having abdominal spiracles that are oval and transverse, among other characteristics.
Larvae

Larvae of the tenebrioninae subfamily take after most other tenebrionid larvae:[2] usually cylindrical to slightly flattened, occasionally short and broad, or strongly flattened. The head and all visible tergites or only the head and abdominal apex are heavily sclerotized.

Diagnostic characters for larvae include the presence of a frontoclypeal suture, flat and dome-like antennal sensorium, simple malar apex which is not cleft, simple ninth sternum, annular or annular- multiforous spiracles, and the absence of an endocarina, mandibular prostheca, hypostomal rods, ventral prolegs, and patches or rows of tergal asperites.
Notable species

Larvae of the yellow mealworm beetle (Tenebrio molitor) are commonly used as feeder insects for reptiles and amphibians. Other Tenebrio and Tribolium species are also bred as animal food. The red flour beetle (Tribolium castaneum) is a popular genetics model organism, especially in studies of intragenomic conflict and population ecology.

Several species of Tenebrio and Tribolium, such as the confused flour beetle (T. confusum) and T. destructor, and other genera such as Gnathocerus cornutus, are pests of cereal and flour silos and other storage facilities.
Systematics
Amarygmini: Plesiophthalmus nigrocyaneus
Blaptini: Blaps cf. mucronata
Bolitophagini: Bolitophagus reticulatus
Mimopeus parvus
Helopini: Stenomax aeneus
Scaphidemini: Scaphidema metallicum
Triboliini: Tribolium destructor
Ulomini: Uloma culinaris

The Tenebrioninae are traditionally divided in some 20-30 tribes. The exact delimitation and validity of several of these is unclear. Molecular phylogenetic studies have yielded inconsistent results, perhaps due to hybridization, horizontal gene transfer by Wolbachia bacteria, and insufficient taxon sampling obfuscating the information contained in DNA sequence data.[3][4]

In some treatments, the Pimeliinae are included as yet another tribe. This may be correct, but as far as can be told they are a closely related but well distinct lineage of Tenebrionidae. The Opatrini are sometimes elevated to familial rank as Opatrinae. The Crypticini, Pentaphyllini, and Scaphidemini are sometimes placed here, but are more commonly located in the Diaperinae.[3][4]

In research by Kamiński et al. published in 2021, the following tribes were moved from Tenebrioninae into the newly resurrected subfamily Blaptinae. These tribes contained 281 genera and about 4000 species, about 50% of Tenebrioninae. The new classification was followed by Bouchard et al. the same year.[5][6]

Amphidorini LeConte, 1862
Blaptini Leach, 1815
Dendarini Mulsant & Rey, 1854
Opatrini Brullé, 1832
Pedinini Eschscholtz, 1829
Platynotini Mulsant & Rey, 1853
Platyscelidini Lacordaire, 1859

The resulting Tenebrioninae, according to "Review of genus-group names in the family Tenebrionidae" (Bouchard et al. 2021), includes the following tribes:[6]

Acropteronini Doyen, 1989
Alphitobiini Reitter, 1917
Amarygmini Gistel, 1848
Apocryphini Lacordaire, 1859
Bolitophagini W. Kirby, 1837
Centronopini Doyen, 1989
Cerenopini Horn, 1870
Dissonomini G.S. Medvedev, 1968
Eulabini Horn, 1870
Falsocossyphini Ferrer, 2006
Heleini Fleming, 1821
Helopini Latreille, 1802
Melanimonini Seidlitz, 1894 (1854)
Metaclisini Steiner, 2016
Palorini Matthews, 2003
Paoligenini Ferrer, 2013
Praeugenini De Moor, 1970
Rhysopaussini Wasmann, 1896
Scaurini Billberg, 1820
Scotobiini Solier, 1838
Tenebrionini Latreille, 1802
Titaenini Fauvel, 1905
Toxicini Oken, 1843
Trachelostenini Lacordaire, 1859
Triboliini Gistel, 1848
Ulomini Blanchard, 1845

In addition, the following genera are of uncertain placement in this subfamily:

Anophthalmolamus Ferrer, 1993
Hangaya Matthews & Merkl, 2015
Penichrus Champion, 1885

References

St. George, R. A. (1924). STUDIES ON THE LARVAE OF NORTH AMERICAN BEETLES OF THE SUBFAMILY TENEBRIONINAE WITH A DESCRIPTION OF THE LARVA AND PUPA OF MERINUS LAEVIS (OLIVIER). Proceedings of the United States National Museum: Bureau of Entomology, United States Department of Agriculture. pp. 2–3.
"Introduction to the Darkling beetles of Eastern United States and Florida". entnemdept.ufl.edu. Retrieved 2021-12-31.
Angelini, D. R. & E. L. Jockusch. (2008). Relationships among pest flour beetles of the genus Tribolium (Tenebrionidae) inferred from multiple molecular markers. Molecular Phylogenetics and Evolution 46(1) 127-41. doi:10.1016/j.ympev.2007.08.017 (HTML abstract)
Bouchard, P.; et al. (2005). "Synoptic classification of the world Tenebrionidae (Insecta: Coleoptera) with a review of family-group names" (PDF). Annales Zoologici. 55 (4): 499–530. Archived from the original (PDF) on 2016-03-03.
Kamiński, Marcin; Lumen, Ryan; Kanda, Kojun; Iwan, Dariusz; et al. (2021). "Reevaluation of Blapimorpha and Opatrinae: addressing a major phylogeny-classification gap in darkling beetles (Coleoptera: Tenebrionidae: Blaptinae)". Systematic Entomology. 46: 140–156. doi:10.1111/syen.12453. S2CID 224888924.
Bouchard, Patrice; Bousquet, Yves; Aalbu, Rolf L.; Alonso-Zarazaga, Miguel A.; et al. (2021). "Review of genus-group names in the family Tenebrionidae (Insecta, Coleoptera)". ZooKeys (1050): 1–633. doi:10.3897/zookeys.1050.64217. hdl:10261/250214. PMC 8328949. PMID 34385881.

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