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Tenebrioninae is the largest
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...
of the
darkling beetle Darkling beetle is the common name for members of the beetle family Tenebrionidae, comprising over 20,000 species in a cosmopolitan distribution. Taxonomy ''Tenebrio'' is the Latin generic name that Carl Linnaeus assigned to some flour beetles ...
s (Tenebrionidae), containing
flour beetle Flour beetles are members of several darkling beetle genera including '' Tribolium'' and ''Tenebrio''. They are pests of cereal silos and are widely used as laboratory animals, as they are easy to keep. The flour beetles consume wheat and other ...
s, among others. Tenebrioninae contains more than 20 tribes.


Description


Adults

Adults are robust, mid-sized
beetle Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
s that typically have
elytra An elytron (; ; : elytra, ) is a modified, hardened forewing of beetles (Coleoptera), though a few of the true bugs (Hemiptera) such as the family Schizopteridae are extremely similar; in true bugs, the forewings are called hemelytra (sometime ...
with some sort of corrugation on the upper side. They are typically black, dark brown or grey, and often have a
satin A satin weave is a type of Textile, fabric weave that produces a characteristically glossy, smooth or lustrous material, typically with a glossy top surface and a dull back; it is not durable, as it tends to snag. It is one of three fundamen ...
y sheen. The body is shaped like a medication capsule or like a
bullet A bullet is a kinetic projectile, a component of firearm ammunition that is shot from a gun barrel. They are made of a variety of materials, such as copper, lead, steel, polymer, rubber and even wax; and are made in various shapes and constru ...
; the legs can be short and stout or long and spindly. They eat both fresh and decaying vegetation, including
vegetable Vegetables are edible parts of plants that are consumed by humans or other animals as food. This original meaning is still commonly used, and is applied to plants collectively to refer to all edible plant matter, including edible flower, flo ...
produce, and several are commercially important pests of
flour Flour is a powder made by Mill (grinding), grinding raw grains, List of root vegetables, roots, beans, Nut (fruit), nuts, or seeds. Flours are used to make many different foods. Cereal flour, particularly wheat flour, is the main ingredie ...
and other
cereal A cereal is a grass cultivated for its edible grain. Cereals are the world's largest crops, and are therefore staple foods. They include rice, wheat, rye, oats, barley, millet, and maize ( Corn). Edible grains from other plant families, ...
products. The subfamily has been characterized as adults having mandibles with the back opposite the cutting edge, without margination and excavated opposite the molar pait; having
ocelli A simple eye or ocellus (sometimes called a pigment pit) is a form of eye or an optical arrangement which has a single lens without the sort of elaborate retina that occurs in most vertebrates. These eyes are called "simple" to distinguish the ...
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 The pygidium (: pygidia) is the posterior body part or shield of crustaceans and some other arthropods, such as insects and the extinct trilobites. In groups other than insects, it contains the anus and, in females, the ovipositor. It is compos ...
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: usually cylindrical to slightly flattened, occasionally short and broad, or strongly flattened. The head and all visible
tergites A ''tergum'' (Latin for "the back"; : ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod segment other than the head. The anterior edge is called the 'base' and posterior edge is called the 'apex' or 'margin'. ...
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 Mealworms are the larval form of the yellow mealworm beetle, ''Tenebrio molitor'', a species of darkling beetle. The yellow mealworm beetle prefers a warmer climate and higher humidity. Male mealworm beetles release a sex pheromone to attract ...
(''Tenebrio molitor'') are commonly used as feeder insects for
reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
s and
amphibian Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s. Other ''
Tenebrio ''Tenebrio'' is a genus of darkling beetles. Adults are long and can live for 1–2 years. The larvae are minor pests, but they are also widely reared and sold as pet food. Species The genus contains the following extant species: * '' Tenebrio ...
'' and '' Tribolium'' species are also bred as animal food. The
red flour beetle The red flour beetle (''Tribolium castaneum'') is a species of beetle in the family Tenebrionidae, the darkling beetles. The red flour beetle, and other closely related beetles like '' Gnatocerus cornutus'', are a worldwide pest of stored produc ...
(''Tribolium castaneum'') is a popular
genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinians, Augustinian ...
model organism A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workings of other organisms. Mo ...
, especially in studies of
intragenomic conflict Intragenomic conflict refers to the evolutionary phenomenon where genes have phenotypic effects that promote their own transmission in detriment of the transmission of other genes that reside in the same genome. The selfish gene theory postulates ...
and
population ecology Population ecology is a sub-field of ecology that deals with the dynamics of species populations and how these populations interact with the environment (biophysical), environment, such as birth rate, birth and death rates, and by immigration an ...
. Several species of ''Tenebrio'' and ''Tribolium'', such as the
confused flour beetle The confused flour beetle (''Tribolium confusum''), a type of darkling beetle known as a flour beetle, is a globally found, common pest insect known for attacking and infesting stored flour and grain. They are one of the most common and most destr ...
(''T. confusum'') and '' T. destructor'', and other genera such as '' Gnathocerus cornutus'', are pests of cereal and flour silos and other storage facilities.


Systematics

The Tenebrioninae are traditionally divided in some 20-30
tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide use of the term in English is in the discipline of anthropology. The definition is contested, in part due to conflict ...
s. The exact delimitation and validity of several of these is unclear.
Molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
studies have yielded inconsistent results, perhaps due to
hybrid Hybrid may refer to: Science * Hybrid (biology), an offspring resulting from cross-breeding ** Hybrid grape, grape varieties produced by cross-breeding two ''Vitis'' species ** Hybridity, the property of a hybrid plant which is a union of two diff ...
ization,
horizontal gene transfer Horizontal gene transfer (HGT) or lateral gene transfer (LGT) is the movement of genetic material between organisms other than by the ("vertical") transmission of DNA from parent to offspring (reproduction). HGT is an important factor in the e ...
by ''
Wolbachia ''Wolbachia'' is a genus of gram-negative bacteria infecting many species of arthropods and filarial nematodes. The symbiotic relationship ranges from parasitism to obligate mutualism. It is one of the most common parasitic microbes of arthrop ...
'' bacteria, and insufficient
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
sampling obfuscating the information contained in
DNA sequence A nucleic acid sequence is a succession of bases within the nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. This succession is denoted by a series of a set of five different letters that indicate the order of the nu ...
data.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 ''Molecular Phylogenetics and Evolution'' is a peer-reviewed scientific journal of evolutionary biology and phylogenetics. The journal is edited by E.A. Zimmer. Indexing The journal is indexed in: * EMBiology *Journal Citation Reports *Scopus ...
'' 46(1) 127-41. (HTML abstract)
In some treatments, the
Pimeliinae Pimeliinae is a subfamily of beetles in the family Tenebrionidae. Selected genera * Tribus Adelostomini ** '' Adelostoma'' ** '' Carinosella'' ** '' Eurychora'' ** '' Prunaspila'' * Tribus Adesmiini ** '' Adesmia'' ** '' Alogenius'' ** ' ...
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 Darkling beetle is the common name for members of the beetle family Tenebrionidae, comprising over 20,000 species in a cosmopolitan distribution. Taxonomy ''Tenebrio'' is the Latin generic name that Carl Linnaeus assigned to some flour beetles ...
. The
Opatrini Opatrini is a tribe of darkling beetles (Tenebrionidae) in the subfamily Tenebrioninae. In research by Kamiński et al. published in 2021, Opatrini and six other tribes were moved from Tenebrioninae to a newly resurrected subfamily,Blaptinae. T ...
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 Diaperinae is a subfamily of darkling beetles in the family Tenebrionidae Darkling beetle is the common name for members of the beetle family Tenebrionidae, comprising over 20,000 species in a cosmopolitan distribution. Taxonomy ''Tenebrio'' ...
. In research by Kamiński et al. published in 2021, the following tribes were moved from Tenebrioninae into the newly resurrected subfamily
Blaptinae Blaptinae is a subfamily of darkling beetles in the family Tenebrionidae Darkling beetle is the common name for members of the beetle family Tenebrionidae, comprising over 20,000 species in a cosmopolitan distribution. Taxonomy ''Tenebrio'' i ...
. 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. *
Amphidorini Amphidorini is a tribe of darkling beetles in the family Tenebrionidae. There are six genera in Amphidorini. In research by Kamiński et al. published in 2021, Amphidorini and six other tribes were moved from Tenebrioninae into the newly resurre ...
LeConte, 1862 *
Blaptini Blaptini is a tribe of darkling beetles in the family Tenebrionidae. There are more than 30 genera recognised in the tribe Blaptini. In research by Kamiński et al. published in 2021, Blaptini and six other tribes were moved from Tenebrioninae i ...
Leach, 1815 *
Dendarini Dendarini is a tribe of darkling beetles in the family Tenebrionidae. There are more than 30 genera in Dendarini. In research by Kamiński et al. published in 2021, Dendarini and six other tribes were moved from Tenebrioninae into the newly res ...
Mulsant & Rey, 1854 *
Opatrini Opatrini is a tribe of darkling beetles (Tenebrionidae) in the subfamily Tenebrioninae. In research by Kamiński et al. published in 2021, Opatrini and six other tribes were moved from Tenebrioninae to a newly resurrected subfamily,Blaptinae. T ...
Brullé, 1832 *
Pedinini Pedinini is a tribe of darkling beetles in the family Tenebrionidae. There are about 19 genera in Pedinini. In research by Kamiński et al. published in 2021, Pedinini and six other tribes were moved from Tenebrioninae Tenebrioninae is the lar ...
Eschscholtz, 1829 *
Platynotini Platynotini is a tribe of darkling beetles in the family Tenebrionidae. There are more than 70 genera in Platynotini. In research by Kamiński et al. published in 2021, Platynotini and six other tribes were moved from Tenebrioninae into the newl ...
Mulsant & Rey, 1853 *
Platyscelidini Platyscelidini is a tribe of darkling beetles in the family Tenebrionidae. There are about eight genera in Platyscelidini. In research by Kamiński et al. published in 2021, Platyscelidini and six other tribes were moved from Tenebrioninae into ...
Lacordaire, 1859 The resulting Tenebrioninae, according to "Review of genus-group names in the family Tenebrionidae" (Bouchard et al. 2021), includes the following tribes: *
Acropteronini ''Acropteryx'' is a genus of darkling beetles in the family Tenebrionidae. They are found in the Neotropics The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropic ...
Doyen, 1989 *
Alphitobiini Alphitobiini is a tribe of darkling beetles in the family Tenebrionidae. There are about eight genera in Alphitobiini. Genera These genera belong to the tribe Alphitobiini: * ''Alphitobius ''Alphitobius'' is a genus of darkling beetles in the ...
Reitter, 1917 *
Amarygmini Amarygmini is a tribe of darkling beetles in the family Tenebrionidae. There are more than 80 genera in Amarygmini. Description Adult Amarygmini are active long-legged beetles that fly readily. They have an unhidden membrane between the front ...
Gistel, 1848 *
Apocryphini Apocryphini is a tribe of darkling beetles in the family Tenebrionidae. There are at least four genera in Apocryphini. Genera These genera belong to the tribe Apocryphini: * ''Apocrypha (genus), Apocrypha'' Eschscholtz, 1831 (North America and ...
Lacordaire, 1859 *
Bolitophagini Bolitophagini is a tribe of darkling beetles in the family Tenebrionidae. There are more than 20 genera in Bolitophagini. Genera These genera belong to the tribe Bolitophagini: * ''Afrobyrsax'' Ardoin, 1973 (tropical Africa) * ''Atasthalomorph ...
W. Kirby, 1837 *
Centronopini Centronopini is a tribe of darkling beetles in the family Tenebrionidae Darkling beetle is the common name for members of the beetle family Tenebrionidae, comprising over 20,000 species in a cosmopolitan distribution. Taxonomy ''Tenebrio'' is ...
Doyen, 1989 *
Cerenopini Cerenopini is a tribe of darkling beetles in the family Tenebrionidae Darkling beetle is the common name for members of the beetle family Tenebrionidae, comprising over 20,000 species in a cosmopolitan distribution. Taxonomy ''Tenebrio'' is t ...
Horn, 1870 *
Dissonomini Dissonomini is a tribe of darkling beetles in the family Tenebrionidae. There are at least two genera in Dissonomini, found in the Palearctic The Palearctic or Palaearctic is a biogeographic realm of the Earth, the largest of eight. Confine ...
G.S. Medvedev, 1968 *
Eulabini Eulabini is a tribe of darkling beetles in the family Tenebrionidae Darkling beetle is the common name for members of the beetle family Tenebrionidae, comprising over 20,000 species in a cosmopolitan distribution. Taxonomy ''Tenebrio'' is the ...
Horn, 1870 *
Falsocossyphini Falsocossyphini is a tribe of darkling beetles in the family Tenebrionidae. There are at least three genera in Falsocossyphini. Genera These genera belong to the tribe Falsocossyphini: * ''Blatticephalus'' Heller, 1918 (tropical Africa) * ''Fal ...
Ferrer, 2006 *
Heleini Heleini is a tribe of darkling beetles in the family Tenebrionidae. There are more than 40 genera in Heleini, found in Australasia. Genera These genera belong to the tribe Heleini: * ''Agastenes'' R. Lucas, 1920 * ''Aglypta'' Gebien, 1908 * ''A ...
Fleming, 1821 * Helopini Latreille, 1802 *
Melanimonini Melanimonini is a tribe of darkling beetles in the family Tenebrionidae Darkling beetle is the common name for members of the beetle family Tenebrionidae, comprising over 20,000 species in a cosmopolitan distribution. Taxonomy ''Tenebrio'' is ...
Seidlitz, 1894 (1854) *
Metaclisini ''Metaclisa'' is a genus of darkling beetles in the family Tenebrionidae, the sole genus of the tribe Metaclisini. There are at least four described species in ''Metaclisa'', found in North America, the Neotropical Realm, Neotropics, the Palearct ...
Steiner, 2016 *
Palorini Palorini is a tribe of darkling beetles in the family Tenebrionidae. There are about 12 genera in Palorini. Genera These genera belong to the tribe Palorini: * ''Astalbus'' Fairmaire, 1900 (tropical Africa) * ''Austropalorus'' Halstead, 1967 ...
Matthews, 2003 *
Paoligenini ''Paoligena'' is a genus of darkling beetles in the family Tenebrionidae, found in tropical Africa The Afrotropical realm is one of the Earth's eight biogeographic realms. It includes Sub-Saharan Africa, the southern Arabian Peninsula, the i ...
Ferrer, 2013 *
Praeugenini Praeugenini is a tribe of darkling beetles in the family Tenebrionidae. There are about seven genera in Praeugenini, found in tropical Africa. Genera These genera belong to the tribe Praeugenini: * ''Anarmostodera'' Fairmaire, 1897 * ''Dysgena' ...
De Moor, 1970 *
Rhysopaussini Rhysopaussini is a tribe of darkling beetles in the family Tenebrionidae. There are at least four genera in Rhysopaussini. Genera These genera belong to the tribe Rhysopaussini: * ''Mimoxenotermes'' Pic, 1931 (Indomalaya) * ''Rhysopaussus'' Was ...
Wasmann, 1896 * Scaurini Billberg, 1820 *
Scotobiini Scotobiini is a tribe of darkling beetles in the family Tenebrionidae. There are about six genera in Scotobiini, found in the Neotropics. Genera These genera belong to the tribe Scotobiini: * ''Ammophorus'' Guérin-Méneville, 1831 * ''Diastoleu ...
Solier, 1838 *
Tenebrionini Tenebrionini is a tribe of darkling beetles in the family Tenebrionidae. There are at least 20 genera in Tenebrionini. Genera These genera belong to the tribe Tenebrionini: * ''Ariarathus'' Fairmaire, 1891 (the Palearctic and Indomalaya) * ''A ...
Latreille, 1802 *
Titaenini Titaenini is a tribe of darkling beetles in the family Tenebrionidae. There are at least eight genera recognised in the tribe Titaenini, all found in Australasia. Genera These genera belong to the tribe Titaenini: * ''Artystona'' Bates, 1874 * ' ...
Fauvel, 1905 *
Toxicini Toxicini is a tribe of darkling beetles in the family Tenebrionidae. There are about 17 genera in Toxicini. Genera These genera belong to the tribe Toxicini: * ''Calymmus'' Montrouzier, 1860 (Australasia) * ''Chalcostylus'' Fairmaire, 1898 (t ...
Oken, 1843 *
Trachelostenini Trachelostenini is a tribe of darkling beetles in the family Tenebrionidae. There are at least three genera in Trachelostenini. It was historically ranked as the family Trachelostenidae, but a 2015 study sunk the family into the tenebrionid (dar ...
Lacordaire, 1859 *
Triboliini Triboliini is a tribe of darkling beetles in the family Tenebrionidae Darkling beetle is the common name for members of the beetle family Tenebrionidae, comprising over 20,000 species in a cosmopolitan distribution. Taxonomy ''Tenebrio'' is t ...
Gistel, 1848 *
Ulomini Ulomini is a tribe of darkling beetles in the family Tenebrionidae. There are more than 20 genera among Ulomini. Genera These genera belong to the tribe Ulomini: * ''Achthosus'' Pascoe, 1863 (Australasia) * ''Alegoria'' Laporte, 1840 (the Neo ...
Blanchard, 1845 In addition, the following genera are of uncertain placement in this subfamily: * '' Anophthalmolamus'' Ferrer, 1993 * '' Hangaya'' Matthews & Merkl, 2015 * '' Nycterinus'' Eschscholtz, 1829 * '' Penichrus'' Champion, 1885


References

* * {{Taxonbar, from1=Q914904 Beetle subfamilies Articles containing video clips Taxa named by Pierre André Latreille