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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
Subfamiliae (11): Alleculinae - Blaptinae - Cossyphodinae - Diaperinae - KuhitangiinaeLagriinae - Nilioninae - Phrenapatinae - Pimeliinae - Stenochiinae - Tenebrioninae - Zolodininae

[source: Bouchard et al. (2005)]
Overview of genera

Accanthopus – Centorus – Echinotus – Foranotum – Leptoderis – Pulposipes – Stenocara – Sternotrigon – … Syachis –
Name

Tenebrionidae Latreille, 1802
Synonyms

Alleculidae
Lagriidae

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.
Kamiński, M.J., Kanda, K., Lumen, R., Ulmer, J.W., Wirth, C.C., Bouchard, P., Aalbu, R., Mal, N. & Smith, A.D. 2019. A catalogue of the tribe Sepidiini Eschscholtz, 1829 (Tenebrionidae, Pimeliinae) of the world. ZooKeys 844: 1–121. DOI: 10.3897/zookeys.844.34241 Open access. Reference page.
Latreille, P.A. 1802: Histoire naturelle, générale et particulière des Crustacés et des Insectes. Tome 3. F. Dufart, Paris. Internet Archive BHL
Nabozhenko, M.V. & Sadeghi, S. 2017. Foranotum perforatum gen. et sp. nov.—a new troglobitic darkling beetle (Coleoptera: Tenebrionidae: Kuhitangiinae: Foranotini trib. nov.) from a cave in Southern Zagros, Iran. Zootaxa 4338(1): 163–172. DOI: 10.11646/zootaxa.4338.1.9. Reference page.

Additional references

ANDO, K. 2003: A New Tenebrionid Genus from Borneo, with Description of a New Species (Coleoptera: Tenebrionidae). The Entomological Review of Japan 58 (2): 107–112.
Ando, K. & Ruzzier, E. 2016. New species of Tenebrionidae Latreille, 1802 (Coleoptera: Tenebrionoidea) from the Philippines. Zootaxa 4175(5): 480–486. DOI: 10.11646/zootaxa.4175.5.6. Reference page.
Ba, Y.-B. & Ren, G.-D. 2013. Taxonomy of Syachis Bates (Coleoptera, Tenebrionidae) and description of two new species from China. Acta zootaxonomica sinica 38(1): 104–108. Abstract Reference page.
Becvar, S. & L. Purchart, 2008: Revision of the genus Hexarhopalus Fairmaire, 1891 (Coleoptera: Tenebrionidae: Cnodaloninae), with description of Malaysphena gen. nov. Acta Zoologici 58 (1): 35–70.
Bouchard, P.; Smith, A.D. (eds.) 2014: Proceedings of the Third International Tenebrionoidea Symposium, Arizona, USA, 2013. ZooKeys, (415) Reference page.
Bousquet, Y., Thomas, D.B., Bouchard, P., Smith, A.D., Aalbu, R.L., Johnston, M.A. & Steiner Jr., W.E. 2018. Catalogue of Tenebrionidae (Coleoptera) of North America. ZooKeys 728: 1–455. DOI: 10.3897/zookeys.728.20602, PMC: 5799738 Open access. PMID: 29416389. Reference page.
Carrara, R. & Flores, G.E. 2015. Endemic epigean Tenebrionids (Coleoptera: Tenebrionidae) from the Andean Region: exploring the patagonian–diversification hypothesis. Zootaxa 4007(1): 47–62. DOI: 10.11646/zootaxa.4007.1.3. Preview (PDF) Reference page.
Doyen, J.T.; Lawrence, J.F. 1979: Relationships and higher classification of some Tenebrionidae and Zopheridae (Coleoptera). Systematic entomology, 4: 333–377.
Ferrer, J. 2015. Revision of the genus Leptoderis Billberg, 1820 with comments about the origin, composition, anatomy and necrophagy of the tribu Elenophorini (Coleoptera, Tenebrionidae). Boletín de la Sociedad Entomológica Aragonesa 57(2): 19–38. Reference page.
Iwan, D., Löbl, I., Bouchard, P., Bousquet, Y., Kamiński, M., Merkl, O., Ando, K., & Schawaller, W. (2020) Family Tenebrionidae Latreille, 1802, In: Iwan, D. & Löbl, I. (eds.), Catalogue of Palaearctic Coleoptera, 5, Tenebrionidea Revised and Updated Second Edition. Brill, Leiden, pp. 104-475. Reference page.
Johnston, M.A., Aalbu, R.L. & Franz, N.M. 2018. An updated checklist of the Tenebrionidae sec. Bousquet et al. 2018 of the Algodones Dunes of California, with comments on checklist data practices. Biodiversity Data Journal 6: e24927. DOI: 10.3897/BDJ.6.e24927. Reference page.
Kamiński, M.J. 2015. A taxonomic revision of the Quadrideres simplicipes species-group (Coleoptera: Tenebrionidae: Pedinini), including description of a new species from Tanzania. Zootaxa 3986(2): 217–226. DOI: 10.11646/zootaxa.3986.2.5 Preview (PDF). Reference page.
Kergoat, G.J. et al. 2014: Higher level molecular phylogeny of darkling beetles (Coleoptera: Tenebrionidae). Systematic entomology 39(3): 486–499. DOI: 10.1111/syen.12065 Reference page.
Kirejtshuk, A.G.; Merkl, O.; Kernegger, F. 2008: A new species of the genus Pentaphyllus Dejean, 1821 (Coleoptera, Tenebrionidae, Diaperinae) from the Baltic amber and checklist of the fossil Tenebrionidae. Zoosystematica Rossica, 17(1): 131–137.
Labrique H. 2009: Une nouvelle espèce de Scaurus F. du Sud marocain (Coleoptera, Tenebrionidae). Société entomologique de Mulhouse 65 (1): 9–13, 5 photos coul. - Scaurus girardi n. sp.
Lawrence, J.F.; Spilman, T.J. 1991: Tenebrionidae (Tenebrionoidea) (including Alleculidae, Cossyphodidae, Lagriidae, Nilionidae, Rhysopaussidae, Tentyriidae). pp. 520–524 in: Lawrence, J.F. (coordinator) Order Coleoptera, in: Stehr, F.W. (ed.) Immature insects, 2. Kendall/Hunt Publishing Company, Iowa, USA.
I.Löbl & A.Smetana (eds). 2008 Catalogue of Palearctic Coleoptera. Vol. 5: Tenebrionoidea. Apollo Books, Stenstrup, Denmark
ISBN 87-88757-84-6, p. 105
Masumoto, K. and K. Akita, 2012: A Revision of the Genus Hoploedipinus Kaszab (Coleoptera, Tenebrionidae) Part 2. Japanese Journal of Systematic Entomology, 18 (2):
Matthews, E.G.; Lawrence, J.F.; Bouchard, P.; Steiner, W.E., jr.; Ślipiński, A. 2010: 11.14. Tenebrionidae Latreille, 1802. pp. 574–659 in: Leschen, R.A.B.; Beutel, R.G.; Lawrence, J.F. (volume eds.) Coleoptera, beetles. Volume 2: Morphology and systematics (Elateroidea, Bostrichiformia, Cucujiformia partim). In: Kristensen, N.P. & Beutel, R.G. (eds.) Handbook of zoology. A natural history of the phyla of the animal kingdom. Volume IV. Arthropoda: Insecta. Part 38. Berlin, New York: Walter de Gruyter. ISBN 3110190753 ISBN 9783110190755
Wagner, G.K. & Gosik, R. 2016. Comparative morphology of immature stages of two sympatric Tenebrionidae species, with comments on their biology. Zootaxa 4111(3): 201–222. DOI: 10.11646/zootaxa.4111.3.1. Reference page.
Watt, J.C. 1975(1974): A revised subfamily classification of Tenebrionidae (Coleoptera). New Zealand journal of zoology 1(4): 381-452. DOI: 10.1080/03014223.1974.9517846 Reference page.

Links

Australian Faunal Directory
EoL
Tenebrionidae in SYNOPSIS OF THE DESCRIBED COLEOPTERA OF THE WORLD
Bouchard, Patrice 2008. Tenebrionidae. Darkling beetles. Version 06 November 2008 (temporary) in The Tree of Life Web Project
tenebrioniDBase
Zoological Institute of Russian Academy of Sciences: Atlas of darkling beetles (Tenebrionidae) of Russia

Vernacular names
беларуская: Чарнацелкі
čeština: Potemníkovití
Deutsch: Schwarzkäfer
English: darkling beetles
magyar: Gyászbogárfélék
日本語: ゴミムシダマシ科
한국어: 거저리과
lietuvių: Juodvabaliai
македонски: Брашнари
polski: Czarnuchowate
русский: Чернотелки
svenska: Svartbaggar
中文: 擬步行蟲科

Darkling beetle is the common name of the large family of beetles Tenebrionidae. The number of species in the Tenebrionidae is estimated at more than 20,000 and the family is cosmopolitan in distribution.

Taxonomy

Tenebrio is the Latin generic name that Carl Linnaeus assigned to some flour beetles in his 10th edition of Systema Naturae 1758-59.[1] The word means "seeker of dark places"[2] (or figuratively a trickster); an English language analogy is "darkling".[3]

Numerous Tenebrionidae species do inhabit dark places, however, there are many species in genera such as Stenocara and Onymacris, which are active by day and inactive at night.

The family covers a varied range of forms, such that classification presents great difficulties. These eleven subfamilies were listed in the 2021 review by Bouchard, Bousquet, et al., updating a similar catalog from 2005.[4][5]

Alleculinae Laporte, 1840
Blaptinae Leach, 1815
Diaperinae Latreille, 1802
Kuhitangiinae G.S. Medvedev, 1962
Lagriinae Latreille, 1825
Nilioninae Oken, 1843
Phrenapatinae Solier, 1834
Pimeliinae Latreille, 1802
Stenochiinae Kirby, 1837
Tenebrioninae Latreille, 1802
Zolodininae Watt, 1975

Ongoing phylogenetic studies are showing that some taxonomic changes are needed. For instance the tribal classification of tribe Pedinini has recently been altered.[6]

The misspelling "Terebrionidae" occurs frequently enough to be easily overlooked.[7][8] The error appears to have no particular significance, but to be the product of misreadings, mis-scans and mis-typings.
Tenebrionidae head

The oldest known member of the family is Jurallecula from the Late Jurassic Karabastau Formation of Kazakhstan, assigned to the stepfamily Alleculinae.
Characteristics

The Tenebrionidae may be identified by a combination of features, including:

Their 11-segmented antennae that may be filiform, moniliform or weakly clubbed
First abdominal sternite is entire and not divided by the hind coxae
Eyes notched by a frontal ridge
The tarsi have four segments in the hind pair and five in the fore and mid legs (5-5-4), tarsal claws are simple

Biology and ecology

Tenebrionid beetles occupy ecological niches in mainly deserts and forests as plant scavengers. Most species are generalistic omnivores, and feed on decaying leaves, rotting wood, fresh plant matter, dead insects, and fungi as larvae and adults.[9] Several genera, including Bolitotherus, are specialized fungivores which feed on polypores. Many of the larger species are flightless, and those that are capable, such as T. molitor, only do so when necessary, such as when dispersing or malnourished.[10][11][12]
A tenebrionid larva (Eleodes sp.)

The larvae, known as mealworms or false wireworms, are usually fossorial, heavily armored and nocturnal. They may possibly be an important resource for certain invertebrates and small mammals. However, the adults of many species have chemical defenses and are relatively protected against predators.[11] Adults of most species, except grain pests, have slow metabolisms, and live long lives compared to other insects, ranging from approximately six months to two years.

Some species live in intensely dry deserts such as the Namib, and have evolved adaptions by which they collect droplets of fog that deposit on their elytra. As the droplets accumulate the water drains down the beetles' backs to their mouthparts, where they swallow it.[13]

Humans spread some species such that they have become cosmopolitan, such as Tribolium castaneum, the red flour beetle, which was spread through grain products.
Notable species

The larval stages of several species are cultured as feeder insects for captive insectivores or as laboratory subjects:

Tenebrio molitor is commonly used to feed terrestrial amniotes kept in terraria.
Tribolium castaneum is a laboratory animal useful as a model organism, especially in studies of intragenomic conflict and population ecology.
Zophobas morio, or superworm, is valued as a feed for captive reptiles; it contains less chitin than Tenebrio molitor.
Alphitobius diaperinus, lesser mealworm[14]
Many tenebrionids are pests of cereal and flour silos and other storage facilities, including T. castaneum, other Tribolium species such as Tribolium confusum and Tribolium destructor, and Gnathocerus cornutus.
In southwestern North America, species of the genus Eleodes (particularly E. obscurus) are well known as "pinacate beetles" or "desert stink beetles".
Several genera, such as Stenocara and Onymacris, are of interest in ecological studies of arid conditions and their associated adaptations.
Ulomoides dermestoides, known as "chinese weevil", "peanut beetle", "cancer beetle", or "asthma beetle", is eaten in Argentina where it is thought to be a treatment for cancer, asthma, and other illnesses.

References

"Caroli Linnæi ... Animalium specierum in classes, ordines, genera, species, methodica dispositio ." Internet Archive. 1759.
Jaeger, Edmund Carroll (1959). A source-book of biological names and terms. Springfield, IL: Thomas. ISBN 0398061793.
Brown, Lesley (1993). The New shorter Oxford English dictionary on historical principles. Oxford [Eng.]: Clarendon. ISBN 0198612710.
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.
Bouchard, Patrice. Lawrence, John F. Davies, Anthony E. Newton, Alfred F. Synoptic Classification of the World Tenebrionidae (Insecta: Coleoptera) with a Review of Family-Group Names. Annales Zoologici (Warszawa), 2005, 55(4): 499–530
Kamiński, M.J.; Kanda, K.; Lumen, R.; Smith, A.D.; Iwan, D. (2019). "Molecular phylogeny of Pedinini (Coleoptera, Tenebrionidae) and its implications for higher-level classification". Zoological Journal of the Linnean Society. 185 (1): 77–97. doi:10.1093/zoolinnean/zly033.
Dennis S. Hill (1997). The Economic Importance of Insects. Springer Science & Business Media. pp. 109–. ISBN 978-0412498008.
"Egyptian Beetle (Blaps polychresta)".
"Species Bolitotherus cornutus – Forked Fungus Beetle".
Flying Mealworm Beetle (Tenebrio molitor) on YouTube
"Family Tenebrionidae – Darkling Beetles".
"Bolitotherus cornutus". Archived from the original on 2016-03-04. Retrieved 2015-08-14.
"Desert beetles inspire aircraft design that doesn't freeze". ZME Science. 2016-01-25. Retrieved 2016-01-27.
Alphitobius diaperinus, lesser mealworm. University of Florida IFAS

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