{{Short description|Family of beetles}} {{Automatic taxobox | fossil_range = {{fossil range|Late Jurassic|Recent}} | image = Onymacris unguicularis MHNT.jpg | image_caption = ''[[Onymacris unguicularis]]'' ([[Tenebrioninae]]: [[Adesmiini]])<br />Scale bar (bottom left) is 10 mm | taxon = Tenebrionidae | display_parents = 3 | authority = [[Pierre André Latreille|Latreille]], 1802 | subdivision_ranks = Subfamilies | subdivision = See text | synonyms = Alleculidae }}

'''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 [[genus|generic]] name that [[Carl Linnaeus]] assigned to some [[flour beetle]]s in his ''[[10th edition of Systema Naturae]]'' 1758–59.<ref>{{cite web|url=https://archive.org/details/carolilinniani00linn |last=Linnaeus |first=Carl |author-link=Carl Linnaeus |title=Caroli Linnæi ... Animalium specierum in classes, ordines, genera, species, methodica dispositio .. |page=134 |place=Leiden |publisher=Theodor Haak |language=Latin |year=1759}}</ref> The name means "lover of darkness";<ref name= "Jaeger">{{cite book |last=Jaeger |first=Edmund Carroll |author-link=Edmund Jaeger|title=A source-book of biological names and terms |publisher=Charles C. Thomas |location=Springfield, IL |orig-date=1959 |date=1978 |edition=6th printing, 3rd |page=259 |isbn=0398009163 |url=https://archive.org/details/sourcebookofbiol0000jaeg/page/258/mode/2up?view=theater |url-access=registration}}</ref> the English language term 'darkling' means "characterised by darkness or obscurity";<ref name= "isbn0-19-861271-0">{{cite book |editor-last=Brown |editor-first=Lesley |title=The New shorter Oxford English dictionary on historical principles, Vol. 1, A–M |publisher=Clarendon Press |location=Oxford |year=1993 |page=594 |isbn=0198612710 |url=https://archive.org/details/newshorteroxford00lesl |url-access=registration }}</ref> see also English 'tenebrous', figuratively "obscure, gloomy."<ref name=Onions>{{cite book |editor-last=Onions |editor-first=C. T. |editor-link=C. T. Onions |title=The shorter Oxford English dictionary on historical principles, Vol. 2, N-Z |page=2261 |edition=Reset with revised etymologies and addenda, 3rd |place=Oxford |publisher=Clarendon Press |url=https://archive.org/details/shorteroxfordeng0000litt/page/2260/mode/2up |url-access=registration |date=1973|isbn=978-0-19-861116-5 }}<!--Apparently Vol 2 of the New shorter OED is not on archive.org--></ref>

Many Tenebrionidae species inhabit dark places; in [[genera]] such as ''Stenocara'' and ''Onymacris'', they 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.<ref name=Bouchard2021/><ref name="BLDN">Bouchard, Patrice; Lawrence, John F.; Davies, Anthony E.; Newton, Alfred F. (2005) [https://www.miiz.waw.pl/periodicals/annales-zoologici/abstracts/AZ_55-4abs.pdf "Synoptic Classification of the World Tenebrionidae (Insecta: Coleoptera) with a Review of Family-Group Names"]. ''Annales Zoologici'' (Warszawa), 55(4): 499–530.</ref> * [[Alleculinae]] <small>Laporte, 1840</small> * [[Blaptinae]] <small>Leach, 1815</small> * [[Diaperinae]] <small>Latreille, 1802</small> * [[Kuhitangiinae]] <small>G.S. Medvedev, 1962</small> * [[Lagriinae]] <small>Latreille, 1825</small> * [[Nilioninae]] <small>Oken, 1843</small> * [[Phrenapatinae]] <small>Solier, 1834</small> * [[Pimeliinae]] <small>Latreille, 1802</small> * [[Stenochiinae]] <small>Kirby, 1837</small> * [[Tenebrioninae]] <small>Latreille, 1802</small> * [[Zolodininae]] <small>Watt, 1975</small>

Ongoing phylogenetic studies are showing that some taxonomic changes are needed. For instance the tribal classification of tribe [[Pedinini]] has recently been altered.<ref name="Kaminski2019">{{Cite journal |doi=10.1093/zoolinnean/zly033|title=Molecular phylogeny of Pedinini (Coleoptera, Tenebrionidae) and its implications for higher-level classification|journal=Zoological Journal of the Linnean Society|volume=185|issue=1|pages=77–97|year=2019|last1=Kamiński |first1=M.J. |last2=Kanda |first2=K. |last3=Lumen |first3=R. |last4=Smith |first4=A.D. |last5=Iwan |first5=D.}}</ref>

The misspelling "Te'''<u>r</u>'''ebrionidae" occurs frequently enough to be easily overlooked.<ref name="Hill1997">{{cite book|author=Dennis S. Hill|title=The Economic Importance of Insects|url=https://books.google.com/books?id=RKQIAqMyBJgC&pg=PA109|year=1997|publisher=Springer Science & Business Media|isbn=978-0412498008|pages=109–}}</ref><ref>{{Cite web |title=Egyptian Beetle (Blaps polychresta) - by Graeme Ruck - JungleDragon |url=https://www.jungledragon.com/image/12213/egyptian_beetle_blaps_polychresta.html |access-date=2023-02-02 |website=www.jungledragon.com |language=en}}</ref> The error appears to have no particular significance, but to be the product of misreadings, mis-scans and mis-typings. [[File:Tenebrionidae Beetle, Little Stsimons Island, Georgia, face 2016-02-03-17.04 (24717881181).jpg|thumb|''Tenebrionidae'' head]]

The oldest known member of the family is ''[[Jurallecula]]'' from the [[Late Jurassic]] [[Karabastau Formation]] of Kazakhstan, assigned to the subfamily Alleculinae.

==Characteristics== The Tenebrionidae may be identified by a combination of features, including: * Their eleven-segmented [[Antenna (biology)|antenna]]e that may be [[wikt:filiform|filiform]], [[Antenna (biology)|moniliform]] or weakly clubbed * First abdominal [[sternite]] is entire and not divided by the hind [[Arthropod coxa|coxa]]e * Eyes notched by a frontal ridge * Four segments in the hind pair of [[Arthropod leg|tarsi]] and five in the fore and mid-legs (5–5–4), with simple claws

==Biology and ecology== Tenebrionid beetles occupy [[ecological niche]]s in mainly deserts and forests as plant [[scavenger]]s. Most species are generalistic [[omnivores]], and feed on decaying leaves, rotting wood, fresh plant matter, dead insects, and fungi as larvae and adults.<ref>{{cite web|url=http://bugguide.net/node/view/7187|title=Species Bolitotherus cornutus – Forked Fungus Beetle}}</ref> Several genera, including ''Bolitotherus'', are specialized [[fungivores]] which feed on [[polypore]]s. Many of the larger species are flightless, and those that are capable, such as [[Tenebrio molitor|''T. molitor'']], often rarely do so.<ref>{{YouTube|id=rGDd3S8XOYI|title=Flying Mealworm Beetle (Tenebrio molitor)}}</ref><ref name="bugguide.net">{{Cite web |title=Family Tenebrionidae - Darkling Beetles |url=https://bugguide.net/node/view/152 |access-date=2023-02-02 |website=bugguide.net}}</ref><ref>{{cite web|url=http://www.americaninsects.net/b/bolitotherus-cornutus.html|title=Bolitotherus cornutus|access-date=2015-08-14|archive-url=https://web.archive.org/web/20160304122536/http://www.americaninsects.net/b/bolitotherus-cornutus.html|archive-date=2016-03-04|url-status=dead}}</ref> [[File:Eleodes (Eleodes) tribulus larva.jpg|thumb|A tenebrionid larva (''[[Eleodes]]'' sp.)]] The [[larvae]], known as [[mealworms]] or false wireworms, are usually [[fossorial]], heavily [[Sclerotization|sclerotized]] and nocturnal. They may possibly be an important resource for certain [[invertebrates]] and small [[mammals]]. Adults of many species have chemical defenses and are relatively protected against predators.<ref name="bugguide.net"/> 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.

[[Namib Desert beetle|Some species]] live in intensely dry deserts such as the [[Namib desert|Namib]], and have evolved adaptions by which they collect droplets of fog that deposit on their [[Elytron|elytra]]. As the droplets accumulate the water drains down the beetles' backs to their mouthparts, where they swallow it.<ref>{{Cite web|title=Desert beetles inspire aircraft design that doesn't freeze|url=http://www.zmescience.com/research/technology/desert-beetles-aicraft-freeze-042352/|website=ZME Science|access-date=2016-01-27|language=en-US|date=2016-01-25}}</ref>

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 types== The [[larva]]l stages of several species are cultured as [[feeder insect]]s for captive [[insectivore]]s or as laboratory subjects: * ''[[Tenebrio molitor]]'', or [[mealworm]], is commonly used to feed terrestrial [[amniote]]s kept in [[terrarium|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<ref name=alphdi/> * Many tenebrionids are pests of [[cereal]] and [[flour]] [[Storage silo|silo]]s and other storage facilities, including ''T. castaneum'', other ''[[Tribolium (beetle)|Tribolium]]'' species such as ''[[Tribolium confusum]]'' and ''[[Tribolium destructor]]'', and ''[[Gnatocerus cornutus]].'' * In southwestern North America, species of the genus ''[[Eleodes]]'' (particularly ''[[Eleodes obscura|E. obscura]]'') are well known as "[[pinacate beetle]]s" 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. * ''[[Luprops tristis]]'' is found in India, where it is known as the Mupli beetle. It is notorious for a defensive secretion that causes skin burns. Since they congregate in large numbers, they can easily become a pest. <!-- ==Identification== Important works include: * Aliquò, M., Rastelli, M., Rastelli, S., Soldati, F. "Coleotteri Tenebrionidi d'Italia – Darkling Beetles of Italy". The CD shows the pictures of all the Italian species of darkling beetles (Coleoptera, Tenebrionidae - excepting Allecuninae), photographic tables for recognizing the species, and Italian distribution maps for all species and subspecies. Published by the autors,"Associazione Naturalistica Piemontese", Natural History Museum of Carmagnola (Turin) Project Biodiversty (Rome). The texts of the species file are written both in Italian and in English. See http://web.tiscali.it/tenebrionidae. -->

== Gallery == {{Gallery |mode=packed |Diaperis boleti MHNT Fronton.jpg|''[[Diaperis boleti]]'' under bark of [[oak]]. |Lagria sp.jpg|''[[Lagria]]'' sp. ([[Lagriinae]]) in [[Tanzania]] |Tenebrionid sal.jpg|''[[Platynotus excavatus]]'', India |Stenocara dentata.jpg|''[[Stenocara dentata]]'' in southern Africa |Gonopus tibialis (Fabricius, 1798) (3989862005).jpg|''[[Gonopus tibialis]]'' |Piedishbeetle.jpg|Pie dish beetle, ''[[Helea]]'' sp. in Australia |Hegemona sp. Darkling Beetle, Caves Branch Jungle Lodge, Belmopan, Belize.jpg|''Hegemona'' sp. (De Laporte, 1840) from Belize |Neomida bicornis male.jpg|''[[Neomida bicornis]]'', male |Neomida bicornis female.jpg|''Neomida bicornis'', female |Meracantha contracta.jpg|''[[Meracantha contracta]]'', USA }}

==References== {{Reflist|refs= <ref name=alphdi>[http://entomology.ifas.ufl.edu/creatures/livestock/poultry/lesser_mealworm.htm ''Alphitobius diaperinus'', lesser mealworm.] [[University of Florida]] [[Institute of Food and Agricultural Sciences|IFAS]]</ref> <ref name=Bouchard2021> {{Cite journal | title = Review of genus-group names in the family Tenebrionidae (Insecta, Coleoptera) | date = 2021 | last1 = Bouchard | first1 = Patrice | last2 = Bousquet | first2 = Yves | last3 = Aalbu | first3 = Rolf L. | last4 = Alonso-Zarazaga | first4 = Miguel A. | last5 = Merkl | first5 = Ottó | last6 = Davies | first6 = Anthony E. | display-authors = 4 | journal = ZooKeys | issue = 1050 | pages = 1–633 | url = https://zookeys.pensoft.net/article/64217/ | doi = 10.3897/zookeys.1050.64217 | pmid = 34385881 | pmc = 8328949 | hdl = 10261/250214 | hdl-access = free | doi-access = free | bibcode = 2021ZooK.1050....1B }}</ref>

}}

==External links== * [http://www.tenebrionidae.net Tenebrionidae.net- information and pictures about darkling beetles]

{{Coleoptera|4}} {{Taxonbar|from=Q615799}} {{Authority control}}

[[Category:Tenebrionidae|*]] [[Category:Beetle families]] [[Category:Taxa named by Pierre André Latreille]]