View source for Dung beetle

{{short description|Informal group of insects}}
{{other uses}}
{{Paraphyletic group
| name = Dung beetle
| image = Scarabaeus viettei 01.jpg
| image_alt = Scarabaeus viettei (syn. Madateuchus viettei, Scarabaeidae) in dry spiny forest close to Mangily, western Madagascar
| image_caption = ''{{ill|Scarabaeus viettei|sv}}'' (syn. ''Madateuchus viettei'', [[Scarabaeidae]]) in [[Madagascar spiny forests|dry spiny forest]] close to [[Mangily]], western [[Madagascar]]
| auto = yes
| parent = Scarabaeoidea
| includes = 
*[[Geotrupidae]]
*[[Scarabaeinae]]
*[[Aphodiinae]]
| excludes = 
*[[Termitoderini]]
}}

'''Dung beetles''' are [[beetle]]s that [[coprophagia|feed on feces]]. Some species of dung beetles can bury dung 250 times their own mass in one night.<ref>{{cite web|title=Some Less Known Fascinating Facts about Dung Beetle|url=http://news.biharprabha.com/2014/04/some-less-known-fascinating-facts-about-dung-beetle/|work=IANS|publisher=news.biharprabha.com|access-date=21 April 2014}}</ref>

Many dung beetles, known as ''rollers'', roll dung into round balls, which are used as a food source or breeding chambers. Other dung beetles like ''[[Euoniticellus intermedius]]'', known as ''tunnelers'', bury the dung wherever they find it. A third group, the ''dwellers'', neither roll nor burrow: they simply live in dung. They are often attracted by the feces collected by [[burrowing owl]]s. There are dung beetle species of various colors and sizes, and some functional traits such as body mass (or biomass) and leg length can have high levels of variability.<ref name=":0">{{Cite journal|last1=Griffiths|first1=Hannah M.|last2=Louzada|first2=Julio|last3=Bardgett|first3=Richard D.|last4=Barlow|first4=Jos|date=2016-03-03|title=Assessing the Importance of Intraspecific Variability in Dung Beetle Functional Traits|journal=PLOS ONE|volume=11|issue=3|pages=e0145598|doi=10.1371/journal.pone.0145598|pmid=26939121|pmc=4777568|issn=1932-6203|bibcode=2016PLoSO..1145598G|doi-access=free}}</ref>

All the [[species]] belong to the superfamily [[Scarabaeoidea]], most of them to the subfamilies [[Scarabaeinae]] and [[Aphodiinae]] of the family [[Scarabaeidae]] (scarab beetles). As most species of Scarabaeinae feed exclusively on feces, that subfamily is often dubbed ''true dung beetles''. There are dung-feeding beetles which belong to other families, such as the [[Geotrupidae]] (the ''earth-boring dung beetle''). The Scarabaeinae alone comprises more than 5,000 species.<ref>Frolov, A.V. [http://www.zin.ru/animalia/coleoptera/eng/afrsc_ru.htm "Subfamily Scarabaeinae: atlas of representatives of the tribes (Scarabaeidae)"]. Retrieved on 2007-08-02.</ref>

The [[nocturnal]] African dung beetle ''[[Scarabaeus satyrus]]'' is one of the few known invertebrate animals that navigate and orient themselves using the [[Milky Way]].<ref>{{cite news|url=https://www.bbc.co.uk/news/science-environment-21150721|author=Jonathan Amos|title=Dung beetles guided by Milky Way|work=[[BBC News]]|date=24 January 2013|access-date=28 January 2013}}</ref><ref>{{Cite journal | last1 = Dacke | first1 = M. | last2 = Baird | first2 = E. | last3 = Byrne | first3 = M. | last4 = Scholtz | first4 = C. H. | last5 = Warrant | first5 = E. J. | title = Dung Beetles Use the Milky Way for Orientation | doi = 10.1016/j.cub.2012.12.034 | journal = Current Biology | volume = 23 | issue = 4 | pages = 298–300 | year = 2013 | pmid =  23352694| doi-access = free }}</ref> The daily dung of one [[African elephant|elephant]] can support 2,000,000 beetles.<ref name="j480">{{cite journal |last=Krell |first=Frank-Thorsten |last2=Krell-Westerwalbesloh |first2=Sylvia |date=2024 |title=One elephant may sustain 2 million dung beetles in East African savannas on any given day |journal=The Science of Nature |volume=111 |issue=1 |page= |doi=10.1007/s00114-024-01894-9 |issn=0028-1042}}</ref> 

==Taxonomy==
Dung beetles are not a single taxonomic group; dung feeding is found in a number of families of beetles, so the behaviour cannot be assumed to have evolved only once.
* [[Coleoptera]] (order), beetles
** [[Scarabaeoidea]] (superfamily), scarabs (most families in the group do not use dung)
*** [[Geotrupidae]] (family), "earth-boring dung beetles"<ref>{{cite web|title=Geotrupidae (earth-boring dung beetles)|url=http://www.kerbtier.de/cgi-bin/enFSearch.cgi?Fam=Geotrupidae|publisher=kerbtier.de|access-date=28 March 2015}}</ref>
*** [[Scarabaeidae]] (family), "scarab beetles" (not all species use dung)
**** [[Scarabaeinae]] (subfamily), "true dung beetles"<ref>{{cite web|title=Scarabaeinae dung beetles|url=http://scarabaeinae.lifedesks.org/|access-date=28 March 2015|archive-url=https://web.archive.org/web/20150319101341/http://scarabaeinae.lifedesks.org/|archive-date=2015-03-19|url-status=dead}}</ref>
**** [[Aphodiinae]] (subfamily), "small dung beetles" (not all species use dung)<ref name=skelley>Skelley, P. E. [http://museum.unl.edu/research/entomology/Guide/Scarabaeoidea/Scarabaeidae/Aphodiinae/Aphodiinae-Overview/AphodiinaeO.html Aphodiinae.] {{Webarchive|url=https://web.archive.org/web/20180214210508/http://www.museum.unl.edu/research/entomology/Guide/Scarabaeoidea/Scarabaeidae/Aphodiinae/Aphodiinae-Overview/AphodiinaeO.html |date=2018-02-14 }} ''Generic Guide to New World Scarab Beetles.'' University of Nebraska State Museum. 2008 Version.</ref>

==Ecology and behavior==
[[File:Dung beetle working-001.ogv|thumb|Dung beetle rolling a ball of dung in the [[Addo Elephant National Park]], [[South Africa]]]]

Dung beetles live in many [[habitat]]s, including desert, [[grassland]]s and [[savanna]]s,<ref>{{Cite journal|last1=França|first1=Filipe M.|last2=Korasaki|first2=Vanesca|last3=Louzada|first3=Julio|last4=Vaz-de-Mello|first4=Fernando Z.|date=2016|title=First report on dung beetles in intra-Amazonian savannahs in Roraima, Brazil|journal=Biota Neotropica|volume=16|issue=1|doi=10.1590/1676-0611-BN-2015-0034|issn=1676-0603|doi-access=free}}</ref> [[farm]]lands, and native and planted [[forest]]s.<ref>{{Cite journal|last1=Beiroz|first1=Wallace|last2=Slade|first2=Eleanor M.|last3=Barlow|first3=Jos|last4=Silveira|first4=Juliana M.|last5=Louzada|first5=Julio|last6=Sayer|first6=Emma|date=2017-01-01|title=Dung beetle community dynamics in undisturbed tropical forests: implications for ecological evaluations of land-use change|journal=Insect Conservation and Diversity|language=en|volume=10|issue=1|pages=94–106|doi=10.1111/icad.12206|s2cid=88666069|issn=1752-4598|url=http://eprints.lancs.ac.uk/83810/1/BeirozEtAlPreprint.pdf}}</ref> They are highly influenced by the environmental context,<ref name=":0" /> and do not prefer extremely cold or dry weather. They are found on all continents except [[Antarctica]]. They eat the dung of [[herbivore]]s and [[omnivore]]s, and prefer that produced by the latter.<ref>{{cite web|last1=Dell'Amore|first1=Christine|title=Dung Beetles' Favorite Poop Revealed|url=http://voices.nationalgeographic.com/2012/04/17/weird-wild-dung-beetles-favorite-poop-revealed/|website=National Geographic|publisher=National Geographic Society|access-date=22 September 2016|archive-url=https://web.archive.org/web/20160923045212/http://voices.nationalgeographic.com/2012/04/17/weird-wild-dung-beetles-favorite-poop-revealed/|archive-date=2016-09-23|url-status=dead}}</ref> Many of them also feed on [[mushrooms]] and [[decomposition|decaying]] [[leaf|leaves]] and [[fruit]]s. The [[Neotropical]] ''[[Deltochilum valgum]]'', ''[[Deltochilum kolbei|D. kolbei]]'' and ''[[Deltochilum viridescens|D. viridescens]]'' are carnivores with a strong preference for preying upon [[millipedes]].<ref name=Silva2012>{{cite journal| last1=Silva | first1=F.A.B.  | last2=Vidaurre | first2=T. | last3=Vaz-de-Mello | first3=F. | last4=Louzada | first4=J. | year=2012 | title=Predatory behavior in Deltochilum: convergent evolution or a primitive character within this clade | journal=Journal of Natural History | volume=46 | issue=21–22 | pages=1359–1367 | doi=10.1080/00222933.2012.658584 | s2cid=85229138 }}</ref> Two other species from Brazil, ''[[Canthon dives]]'' and ''[[Canthon virens]]'', prey on queens and other winged forms of [[leafcutter ant]]s.<ref>{{Cite journal|last1=Forti|first1=Luiz Carlos|last2=Rinaldi|first2=Isabela Maria Piovesan|last3=Camargo|first3=Roberto da Silva|last4=Fujihara|first4=Ricardo Toshio|date=2012-03-06|title=Predatory Behavior of Canthon virens (Coleoptera: Scarabaeidae): A Predator of Leafcutter Ants|journal=Psyche: A Journal of Entomology|volume=2012|pages=1–5|doi=10.1155/2012/921465|language=en|doi-access=free|hdl=11449/73288|hdl-access=free}}</ref> One species from the [[Iberian Peninsula]], ''[[Thorectes lusitanicus]],'' feeds on acorns.<ref>{{Cite journal |last1=PéRez-Ramos |first1=Ignacio M. |last2=MarañóN |first2=Teodoro |last3=Lobo |first3=Jorge M. |last4=Verdú |first4=José R. |date=August 2007 |title=Acorn removal and dispersal by the dung beetle Thorectes lusitanicus : ecological implications |url=https://resjournals.onlinelibrary.wiley.com/doi/10.1111/j.1365-2311.2007.00874.x |journal=Ecological Entomology |language=en |volume=32 |issue=4 |pages=349–356 |doi=10.1111/j.1365-2311.2007.00874.x |issn=0307-6946|hdl=10261/47760 |hdl-access=free }}</ref> Dung beetles do not necessarily have to eat or drink anything else, because the dung provides all the necessary [[nutrient]]s.{{Citation needed|date=August 2019}}

Most dung beetles search for dung using their sensitive [[olfaction|sense of smell]]. Some smaller species simply attach themselves to the dung-providers to wait for the dung. After capturing the dung, a dung beetle rolls it, following a straight line despite all obstacles. Sometimes, dung beetles try to steal the dung ball from another beetle, so the dung beetles have to move rapidly away from a dung pile once they have rolled their ball to prevent it from being stolen. Dung beetles can roll up to 10 times their weight. Male ''Onthophagus taurus'' beetles can pull 1,141 times their own body weight: the equivalent of an average person pulling six double-decker buses full of people.<ref>{{cite news| url=https://www.theguardian.com/environment/2010/mar/24/dung-beetles | location=London | work=The Guardian | first=Homa | last=Khaleeli | title=Just how strong is a dung beetle? | date=24 March 2010}}</ref>

A species of dung beetle (the African ''Scarabaeus zambesianus'') [[animal navigation|navigates]] by [[Polarization (waves)|polarization]] patterns in [[moonlight]],{{Sfn|Tills|2013|p=123}}  the first animal known to do so.<ref>{{Cite journal | doi = 10.1038/424033a| pmid = 12840748| title = Animal behaviour: Insect orientation to polarized moonlight| journal = Nature| volume = 424| issue = 6944| page = 33| year = 2003| last1 = Dacke | first1 = M. | last2 = Nilsson | first2 = D. E. | last3 = Scholtz | first3 = C. H. | last4 = Byrne | first4 = M. | last5 = Warrant | first5 = E. J. | bibcode=2003Natur.424...33D| s2cid = 52859195| doi-access = free}}</ref><ref>{{cite journal | last1 = Milius | first1 = Susan | year = 2003 | title = Moonlighting: Beetles navigate by lunar polarity | url = http://www.sciencenews.org/articles/20030705/fob4.asp | journal = Science News | volume = 164 | issue = 1| pages = 4–5 | doi = 10.2307/3981988 | jstor = 3981988 }}</ref><ref>Roach, John (2003). [https://web.archive.org/web/20030707041737/http://news.nationalgeographic.com/news/2003/07/0702_030702_dungbeetle.html "Dung Beetles Navigate by the Moon, Study Says"], ''National Geographic News''. Retrieved on 2007-08-02.</ref><ref>{{Cite journal | doi = 10.2307/3981988| jstor = 3981988| title = Moonlighting| journal = Science News| volume = 164| issue = 1| pages = 4–5| year = 2003| last1 = Milius | first1 = S. }}</ref> Dung beetles can also navigate when only the [[Milky Way]] or clusters of bright stars are visible,<ref name="auto">{{cite journal |doi=10.1016/j.cub.2012.12.034|pmid=23352694|title=Dung Beetles Use the Milky Way for Orientation|journal=Current Biology|volume=23|issue=4|pages=298–300|year=2013|last1=Dacke|first1=Marie|last2=Baird|first2=Emily|last3=Byrne|first3=Marcus|last4=Scholtz|first4=Clarke H.|last5=Warrant|first5=Eric J.|doi-access=free}}</ref>  making them the only insects known to orient themselves by the Milky Way.<ref name="scienceDaily">{{cite news|last=Wits University |url=https://www.sciencedaily.com/releases/2013/01/130124123203.htm |title=Dung Beetles Follow the Milky Way: Insects Found to Use Stars for Orientation|work=ScienceDaily|date=24 January 2013 |access-date=25 January 2013}}</ref><ref name="auto"/> Research using 1 kg bolus of elephant dung found that a larger number exploit it during the night (13,700) than during the day (3,330).<ref name="j480" />  The eyes of dung beetles are superposition [[compound eye]]s typical of many scarabaeid beetles;<ref>{{cite journal|author=Meyer-Rochow, V.B.|title=Retina and dioptric apparatus of the dung beetle ( Euoniticellus africanus ; Scarabaeidae)|journal=Journal of Insect Physiology|date=1978|volume= 24|issue=2|pages= 165–179|doi=10.1016/0022-1910(78)90114-2}}</ref><ref>{{cite journal| author1=Meyer-Rochow, V.B.| author2= Gokan, N.| title=The eye of the dung beetle Onthophagus posticus (Coleoptera, Scarabaeidae)| date=1990| journal= New Zealand Journal of Entomology| volume=13| pages=7–15| doi= 10.1080/00779962.1990.9722582}}</ref>
The sequence of images shows a sequence of the beetle rolling a dung ball. It does this to navigate.
<gallery mode="packed" heights="120px">
Plum dung beetle (Anachalcos convexus) 1 of 4.jpg|The beetle climbs onto the ball
Plum dung beetle (Anachalcos convexus) 2 of 4.jpg|The beetle starts to turn around
Plum dung beetle (Anachalcos convexus) 3 of 4.jpg|The beetle continues turning around
Plum dung beetle (Anachalcos convexus) 4 of 4.jpg|The beetle rolls the ball with its hind legs
</gallery>
Cambefort and Hanski (1991) classified dung beetles into three functional types based on their feeding and nesting strategies such as – Rollers, Tunnelers and Dwellers. The "rollers" roll and bury a dung ball either for food storage or for making a brooding ball. In the latter case, two beetles, one male and one female, stay around the dung ball during the rolling process. Usually it is the male that rolls the ball, while the female hitch-hikes or simply follows behind. In some cases, the male and the female roll together. When a spot with soft soil is found, they stop and bury the ball, then [[mating|mate]] underground. After the mating, one or both of them prepares the brooding ball. When the ball is finished, the female lays eggs inside it, a form of [[mass provisioning]].

Some species do not leave after this stage, but remain to safeguard their offspring. The dung beetle goes through a complete [[metamorphosis (biology)|metamorphosis]]. The larvae live in brood balls made with dung prepared by their parents. During the larval stage, the beetle feeds on the dung surrounding it.

The behavior of the beetles was poorly understood until the studies of [[Jean Henri Fabre]] in the late 19th century. For example, Fabre corrected the myth that a dung beetle would seek aid from other dung beetles when confronted by obstacles. By observation and experiment, he found the seeming helpers were in fact awaiting an opportunity to steal the roller's food source.<ref>{{cite book |author=Fabre, J. Henri |author-link = Henri Fabre |date=1949 |title=The Insect World of J. Henri Fabre |url=https://archive.org/details/insectworldofjhe0000fabr |url-access=registration |others=De Mattos, Alexander Teixeira (translator) |publisher=Dodd, Mead & Company |page=[https://archive.org/details/insectworldofjhe0000fabr/page/99 99] |quote=I ask myself in vain what Proudhon introduced into Scarabaean morality the daring paradox that 'property means plunder', or what diplomatist taught the Dung-beetle the savage maxim that 'might is right'.}}</ref>

They are widely used in ecological research as a good [[bioindicator]] group to examine the impacts of climate disturbances, such as extreme droughts<ref>{{Cite journal |doi = 10.1098/rstb.2019.0116|pmid = 31983328 |title = Climatic and local stressor interactions threaten tropical forests and coral reefs |journal = Philosophical Transactions of the Royal Society B|volume = 375|issue = 1794 |pages = 20190116|year = 2020|last1 = França |first1 = FM|last2 = Benkwitt |first2 = CE|last3 = Peralta |first3 = G |last4 = Robinson |first4 = JPW|last5 = Graham|first5 =  NAJ |last6 = Tylianakis |first6 = JM|last7 = Berenguer |first7 = E |last8 = Lees |first8 = AC|last9 = Ferreira |first9 = J|last10 = Louzada |first10 = J |last11 = Barlow |first11 = J|pmc = 7017775 }}</ref> and associated fires,<ref name="auto1">{{Cite journal|last1=França|first1=Filipe|last2=Ferreira|first2=J| last3=Vaz-de-Mello|first3=FZ| last4=Maia|first4=LF| last5=Berenguer|first5=E| last6=Palmeira|first6=A| last7=Fadini|first7=R| last8=Louzada|first8=J |last9=Braga|first9=R| last10=Oliveira|first10=VH| last11=Barlow|first11=J| date=10 February 2020|title= El Niño impacts on human-modified tropical forests: Consequences for dung beetle diversity and associated ecological processes |journal=Biotropica |volume=52 | issue = 1 |pages=252–262|doi=10.1111/btp.12756|doi-access=free}}</ref> and human activities on tropical biodiversity<ref>{{Cite journal|last1=França|first1=Filipe|last2=Louzada|first2=Julio|last3=Korasaki|first3=Vanesca|last4=Griffiths|first4=Hannah|last5=Silveira|first5=Juliana|last6=Barlow|first6=Jos|title= Do space-for-time assessments underestimate the impacts of logging on tropical biodiversity? An Amazonian case study using dung beetles|journal=Journal of Applied Ecology|year=2016|volume=53 | issue = 4 |pages=1098–1105|doi=10.1111/1365-2664.12657|doi-access=free}}</ref><ref>{{Cite journal|last1=França|first1=Filipe|last2=Barlow|first2=Jos|last3=Araújo|first3=Bárbara|last4=Louzada|first4=Julio|date=4 November 2016|title=Does selective logging stress tropical forest invertebrates? Using fat stores to examine sublethal responses in dung beetles|journal=Ecology and Evolution|volume=6 | issue = 23 |pages=8526–8533|doi=10.1002/ece3.2488|pmid=28031804|pmc=5167030}}</ref> and ecosystem functioning,<ref>{{Cite journal|last1=França|first1=Filipe M.|last2=Frazão|first2=Fábio S.|last3=Korasaki|first3=Vanesca|last4=Louzada|first4=Júlio|last5=Barlow|first5=Jos|date=2017-12-01|title=Identifying thresholds of logging intensity on dung beetle communities to improve the sustainable management of Amazonian tropical forests|journal=Biological Conservation|volume=216|issue=Supplement C|pages=115–122|doi=10.1016/j.biocon.2017.10.014|url=https://eprints.lancs.ac.uk/id/eprint/88405/1/170918_Manuscript.pdf}}</ref> such as seed dispersal, soil bioturbation and [[nutrient cycling]].<ref name="auto1"/>
<gallery mode="packed" heights="120px">
Scarabaeus laticollis.jpg|An earth-boring dung beetle working
DungBeetle.jpg|A dung beetle with two balls of dung
Scarabaeus laticollis 2.jpg|Two dung beetles fighting over a ball of dung
</gallery>

==Benefits and uses==
[[File:Caution Dung Beetles, South Africa.jpg|thumb|upright|Caution sign showing the importance of dung beetles in [[South Africa]]]]
Dung beetles play a role in [[agriculture]] and [[tropical forest]]s. By burying and consuming dung, they improve nutrient recycling and soil structure.<ref>{{Cite journal | doi = 10.1016/j.apsoil.2010.05.010| title = Dung beetles (Coleoptera: Scarabaeidae) can improve soil hydrological properties| journal = Applied Soil Ecology| volume = 46| pages = 9–16| year = 2010| last1 = Brown | first1 = J. | last2 = Scholtz | first2 = C. H. | last3 = Janeau | first3 = J. L. | last4 = Grellier | first4 = S. | last5 = Podwojewski | first5 = P. | hdl = 2263/14419| url = http://repository.up.ac.za/bitstream/2263/14419/1/Brown_Dung%282010%29.pdf | hdl-access = free }}</ref><ref>{{Cite journal|last1=Nichols|first1=E. |last2=Spector|first2=S. |last3=Louzada|first3=J. |last4=Larsen|first4=T. |last5=Amezquita|first5=S. |last6=Favila|first6=M.E. |title=Ecological functions and ecosystem services provided by Scarabaeinae dung beetles|journal=Biological Conservation|volume=141|issue=6|pages=1461–1474 |doi=10.1016/j.biocon.2008.04.011|year=2008}}</ref> Dung beetles have been further shown to improve soil conditions and plant growth on rehabilitated coal mines in South Africa.<ref>{{Cite journal | doi = 10.1016/j.apsoil.2018.08.011| title = Dung beetle activity improves herbaceous plant growth and soil properties on confinements simulating reclaimed mined land in South Africa| journal = Applied Soil Ecology| volume = 132| pages = 53–59| year = 2018| last1 = Badenhorst | first1 = J. | last2 = Dabrowski | first2 = J. | last3 = Scholtz | first3 = C. H. | last4 = Truter | first4 = W. F. | hdl = 2263/67268 | s2cid = 91863191| hdl-access = free }}</ref> They are also important for the dispersal of seeds present in animals' dung,<ref>{{Cite journal|last1= França|first1=Filipe |last2=Louzada|first2=Julio |last3=Barlow|first3=Jos |date=2018-02-15|title= Selective logging effects on 'brown world' faecal-detritus pathway in tropical forests: A case study from Amazonia using dung beetles|journal= Forest Ecology and Management|language=en|volume=410|pages=136–143 |doi=10.1016/j.foreco.2017.12.027|issn=0378-1127 |url=https://eprints.lancs.ac.uk/id/eprint/89648/1/171215_Franca_et_al_Brown_world.pdf}}</ref> influencing seed burial and seedling recruitment in tropical forests.<ref>{{Cite journal|last1=Griffiths|first1=Hannah M. |last2=Bardgett|first2=Richard D.|last3=Louzada|first3=Julio |last4=Barlow|first4=Jos |date=2016-12-14|title=The value of trophic interactions for ecosystem function: dung beetle communities influence seed burial and seedling recruitment in tropical forests|journal=Proc. R. Soc. B|language=en |volume=283|issue=1844|pages=20161634 |doi=10.1098/rspb.2016.1634|issn=0962-8452 |pmc=5204146|pmid=27928036}}</ref> They can protect livestock, such as [[cattle]], by removing the dung which, if left, could provide habitat for [[pest (animal)|pests]] such as [[fly|flies]]. Therefore, many countries have [[Biological pest control|introduced]] the creatures for the benefit of [[animal husbandry]]. The [[American Institute of Biological Sciences]] reports that dung beetles save the [[United States of America|United States]] cattle industry an estimated [[United States dollar|US$]]380 million annually through burying above-ground livestock feces.<ref>{{Cite journal| doi = 10.1641/0006-3568(2006)56[311:TEVOES]2.0.CO;2| title = The Economic Value of Ecological Services Provided by Insects| journal = BioScience| volume = 56| issue = 4| pages = 311–23| year = 2006| last1 = Losey| first1 = J. E.| last2 = Vaughan| first2 = M.| doi-access = free}}</ref>

In Australia, the [[Commonwealth Scientific and Industrial Research Organisation]] (CSIRO) commissioned the [[Australian Dung Beetle Project]] (1965–1985) which, led by [[George Bornemissza]], sought to introduce species of dung beetles from South Africa and Europe. The successful introduction of 23 species was made, most notably ''[[Digitonthophagus gazella]]'' and ''[[Euoniticellus intermedius]]'', which has resulted in improvement of the quality and fertility of Australian cattle pastures, along with a reduction in the population of pestilent [[Musca vetustissima|Australian bush flies]] by around 90%.<ref name="adbp">{{cite journal | last1 = Bornemissza | first1 = G. F. | year = 1976 | title = The Australian dung beetle project 1965–1975 | journal = Australian Meat Research Committee Review | volume = 30 | pages = 1–30 }}</ref><!---Missylisa153, August 2009---><ref>{{cite web |last1=Ward |first1=Colin |title=Dung beetle program |url=https://csiropedia.csiro.au/Dung-beetle-program/ |website=CSIROpedia |archive-url= https://web.archive.org/web/20201124144551/https://csiropedia.csiro.au/Dung-beetle-program/ |archive-date=24 November 2020 |language=en-AU |date=21 February 2011 |url-status=live}}</ref> In 1995 it was reported that dung beetles were being trialled in the Sydney beach suburb of [[Curl Curl]] to deal with dog droppings.<ref>{{cite news|url=http://edition.cnn.com/EARTH/9511/dung_beetle/ |title=CNN - Dung beetles devour 'poo-lution' Down Under|date=4 November 1995 |website=CNN|author=Brady Halls}}</ref>

An application made by [[Landcare Research]] to import up to 11 species of dung beetle into [[New Zealand]]<ref>{{cite web |url=http://www.ermanz.govt.nz/BertDocs/ERMA200599_Application%20summary%20ERMA200599%20FINAL.pdf |title=Application summary – ERMA200599 |publisher=ERMA |access-date=12 December 2010 }}{{dead link|date=September 2017 |bot=InternetArchiveBot |fix-attempted=yes }}</ref> was approved in 2011.<ref>{{Cite web |date=2 February 2011 |title=HSNO Application |url=https://www.epa.govt.nz/database-search/hsno-application-register/view/ERMA200599 |access-date=21 October 2024 |website=www.epa.govt.nz}}</ref> As well as improving pasture soils the Dung Beetle Release Strategy Group said that it would result in a reduction in emissions of [[nitrous oxide]] (a [[greenhouse gas]]) from agriculture.<ref>{{cite news |url= http://www.nzherald.co.nz/nz/news/article.cfm?c_id=1&objectid=10675713 |title=Let's roll... dung beetle to combat global warming|last=Ihaka|first=James|date=24 September 2010|work=[[The New Zealand Herald]]|access-date=12 December 2010}}</ref> There was, however, strong opposition from some at the [[University of Auckland]], and a few others, based on the risks of the dung beetles acting as vectors of disease.<ref>{{cite web |author=17 Aug 2014, 01:09 am |url=http://www.fwplus.co.nz/article/experts-dump-on-dung-beetle?p=6 |title=Experts dump on dung beetle |publisher=Fwplus.co.nz |date=2012-05-07 |access-date=2014-08-16 |url-status=dead |archive-url=https://archive.today/20130414001314/http://www.fwplus.co.nz/article/experts-dump-on-dung-beetle?p=6 |archive-date=2013-04-14 }}</ref><ref>{{cite news|url=http://www.nzherald.co.nz/environment/news/article.cfm?c_id=39&objectid=10866149 |title=Grant Guilford: Dung beetle plan a risky gamble |publisher=Nzherald.co.nz |date= 2013-02-18|access-date=2014-08-16|newspaper=NZ Herald |last1=Guilford |first1=Grant }}</ref> There were public health researchers at the University of Auckland who agreed with the [[Environmental Protection Authority (New Zealand)|Environmenal Protection Authority]]'s risk assessment.<ref>{{Cite journal|author=drj9 |url=http://janinepaynter.wordpress.com/2013/03/17/fickle-thy-name-is-vox-populi-puzzle-over-dung-beetles-and-science-reporting/ |title='Fickle thy name is' vox populi : puzzle over dung beetles and science reporting   |journal =Journal of Outrageous Speculation |date=2013-03-17 |access-date=2014-04-22}}</ref>  Several Landcare programmes in Australia involved schoolchildren collecting dung beetles.<ref>{{cite web|url=http://dungbeetle.org.nz/release-programme/ |title=Release Programme &#124; Dung Beetles in New Zealand |publisher=Dungbeetle.org.nz |date=2013-09-26 |access-date=2014-04-22}}</ref>

The African dung beetle (''D. gazella'') was introduced in several locations in North and [[South America]] and has been spreading its distribution to other regions by natural dispersal and accidental transportation, and is now probably naturalized in most countries between [[México]] and [[Argentina]]. The exotic species might be useful for controlling diseases of [[livestock]] in commercial areas, and might displace native species in modified landscapes; however, data is not conclusive about its effect on native species in natural environments and further monitoring is required.<ref name=DgazellaVenezuela>{{cite journal|last1=Ferrer-Paris|first1=José R.|title=El escarabajo estercolero africano, ''Digitonthophagus gazella'', (Coleoptera: Scarabaeidae) en la región Neotropical, ¿beneficioso o perjudicial?|journal=Boletín de la Red Latinoamericana Para el Estudio de Especies Invasoras|date=2014 |volume=4|issue=1|pages=41–48 |url=https://www.researchgate.net/publication/268510737}}</ref>

Like many other insects, (dried) dung beetle, called ''qiāngláng'' (蜣蜋) in Chinese, is used in [[Chinese materia medica|Chinese herbal medicine]]. It is recorded in the "Insect section" (蟲部) of the ''[[Bencao Gangmu|Compendium of Materia Medica]]'', where it is recommended for the cure of 10 diseases.{{citation needed|date=December 2019}}

In [[Isan]], Northeastern Thailand, the local people eat many different kinds of insects, including the dung beetle. There is an Isan song กุดจี่หายไปใหน "Where Did the Dung Beetle Go", which relates the replacement of [[water buffalo]] with the [[automobile|"metal" buffalo]], which does not provide the dung needed for the dung beetle and has led to the increasing rarity of the dung beetle in the agricultural region.{{citation needed|date= April 2020}}

The Mediterranean dung beetle (''Bubas bison'') has been used in conjunction with [[Biochar#Stock fodder|biochar stock fodder]] to reduce emissions of [[Nitrous oxide#Greenhouse effect|nitrous oxide]] and [[Carbon dioxide#In Earth's atmosphere|carbon dioxide]], which are both greenhouse gases. The beetles work the biochar-enriched dung into the soil without the use of machines.<ref name="abc.net.au-2019-10-18-beetles-charcoal">{{cite news
 | url =https://www.abc.net.au/news/rural/2019-10-18/wa-farmer-uses-beetles-and-charcoal-to-combat-climate-change/11613846
 | title =Poo-eating beetles and charcoal used by WA farmer to combat climate change
 | last =Daly
 | first =Jon
 | date =18 October 2019
 | newspaper =ABC News
 | publisher =Australian Broadcasting Corporation
 | access-date =18 October 2019
 | quote =Mr Pow said his innovative farming system could help livestock producers become more profitable while helping to address the impact of climate change. }}</ref>

Scientists in [[Canberra]] in 1965 discovered that Dung beetles (Scarabaeids), specifically [[Onthophagus|''Onthophagus australis'' Guérin-Méneville]], improve plant yields using their dung. [[Japanese millet]] was studied and data on nutrient uptake. These plants were placed in pots lacking [[nitrogen]], [[phosphorus]], and [[sulfur]]. [[Cow dung|Cow-dung]] was then added in treatment groups with or without ''O. australis.'' Some treatment groups even had two out of the three nutrients supplemented in the pots. Comparisons of the treatment and control groups were made to show that top growth and roots significantly increased when the dung was mixed well into the soil in the pots. Results showed that dung beetle activity greatly improved plant life. The dung has little impact alone, but in combination with the dung beetle, the nutritional value for the plants increases greatly. This suggests that dung beetles have many positive implications for the environment, including a beneficial role with plant life.<ref>{{Cite journal |last1=BORNEMISSZA |first1=G.F. |last2=WILLIAMS |first2=C.H. |date=January 1970 |title=An effect of dung beetle activity on plant yield |journal=Pedobiologia |volume=10 |issue=1 |pages=1–6 |doi=10.1016/s0031-4056(23)00394-3 |issn=0031-4056|doi-access=free }}</ref> 

==In culture==
Some dung beetles are used as food in South East Asia and a variety of dung beetle species have been used [[therapeutically]] (and are still being used in traditionally living societies) in potions and folk medicines to treat a number of illnesses and disorders.<ref>{{cite journal |last=Meyer-Rochow | first=V.B. |title=Therapeutic arthropods and other, largely terrestrial, folk-medicinally important invertebrates: a comparative survey and review. |journal=Journal of Ethnobiology and Ethnomedicine| date=January 2017| volume=13 |issue=9 |pages=9| doi=10.1186/s13002-017-0136-0| pmid=28173820 | pmc=5296966 | doi-access=free }}</ref>

===Ancient Egypt===
{{Main|Scarab (artifact)}}
[[File:Karnak temple scarab beetle A.jpg|thumb|right|A scarab statue at the [[Karnak]] temple complex]]

Several species of the dung beetle, most notably the species ''[[Scarabaeus sacer]]'' (often referred to as ''the sacred scarab''), enjoyed a sacred status among the [[ancient Egypt]]ians.
{{hiero|ḫpr|<hiero>xpr</hiero>|align=right|era=egypt}}Egyptian hieroglyphic script uses the image of the beetle to represent a [[Egyptian language|triliteral phonetic]] that Egyptologists transliterate as ''xpr'' or ''ḫpr'' and translate as "to come into being", "to become" or "to transform". The derivative term ''xprw'' or ''ḫpr(w)'' is variously translated as "form", "transformation", "happening", "mode of being" or "what has come into being", depending on the context. It may have existential, fictional, or ontologic significance.

The scarab was linked to [[Khepri]] ("he who has come into being"), the god of the rising [[sun]]. The ancients believed that the dung beetle was only male-sexed, and reproduced by depositing [[semen]] into a dung ball. The supposed self-creation of the beetle resembles that of Khepri, who creates himself out of nothing. Moreover, the dung ball rolled by a dung beetle resembles the sun. [[Plutarch]] wrote: {{blockquote|The race of beetles has no female, but all the males eject their sperm into a round pellet of material which they roll up by pushing it from the opposite side, just as the sun seems to turn the heavens in the direction opposite to its own course, which is from west to east.<ref>[https://penelope.uchicago.edu/Thayer/E/Roman/Texts/Plutarch/Moralia/Isis_and_Osiris*/E.html#ref373 "Isis and Osiris"], ''Moralia'', in volume V of the [[Loeb Classical Library]] edition, 1936, now in the public domain. Retrieved on 2007-08-02.</ref>}}

The ancient Egyptians believed that Khepri renewed the sun every day before rolling it above the horizon, then carried it through the other world after sunset, only to renew it, again, the next day. Some [[New Kingdom of Egypt|New Kingdom]] royal tombs exhibit a threefold image of the sun god, with the beetle as symbol of the morning sun. The astronomical ceiling in the tomb of [[Ramses VI]] portrays the nightly "death" and "rebirth" of the sun as being swallowed by [[Nut (goddess)|Nut]], goddess of the sky, and re-emerging from her womb as Khepri.

The image of the scarab, conveying ideas of transformation, renewal, and resurrection, is ubiquitous in ancient Egyptian religious and [[funerary art]].

Excavations of ancient Egyptian sites have yielded images of the scarab in [[bone]], [[ivory]], stone, [[Egyptian faience]], and precious metals, dating from the Sixth Dynasty and up to the period of Roman rule. They are generally small, bored to allow stringing on a necklace, and the base bears a brief inscription or [[cartouche]]. Some have been used as [[Impression seal|seals]]. [[Pharaoh]]s sometimes commissioned the manufacture of larger images with lengthy inscriptions, such as the [[The lion hunts of Amenhotep III during the first ten years of his reign|commemorative scarab]] of Queen [[Tiye]]. Massive sculptures of scarabs can be seen at [[Luxor Temple]], at the Serapeum in Alexandria (see [[Serapis]]) and elsewhere in Egypt.

[[File:Egypt.KV6.04.jpg|thumb|left|A scarab, depicted on the walls of [[KV6|Tomb KV6]] in the [[Valley of the Kings]]]]

The scarab was of prime significance in the funerary cult of ancient Egypt. Scarabs, generally, though not always, were cut from green stone, and placed on the chest of the deceased. Perhaps the most famous example of such "heart scarabs" is the yellow-green [[pectoral (Ancient Egypt)|pectoral]] scarab found among the entombed provisions of [[Tutankhamen]]. It was carved from a large piece of [[Libyan desert glass]]. The purpose of the "heart scarab" was to ensure that the heart would not bear witness against the deceased at judgement in the Afterlife. Other possibilities are suggested by the "transformation spells" of the ''Coffin Texts'', which affirm that the soul of the deceased may transform (''xpr'') into a human being, a god, or a bird and reappear in the world of the living.

One scholar comments on other traits of the scarab connected with the theme of death and rebirth:
{{blockquote|It may not have gone unnoticed that the pupa, whose wings and legs are encased at this stage of development, is very mummy-like. It has even been pointed out that the egg-bearing ball of dung is created in an underground chamber which is reached by a vertical shaft and horizontal passage curiously reminiscent of Old Kingdom mastaba tombs."<ref>Andrews, Carol (1994). ''Amulets of Ancient Egypt''. Texas: University of Texas Press. {{ISBN|0-292-70464-X}}. p. 51.</ref>}}

In contrast to funerary contexts, some of ancient Egypt's neighbors adopted the scarab motif for [[scaraboid seal|seals]] of varying types. The best-known of these being Judean [[LMLK seal]]s (8 of 21 designs contained scarab beetles), which were used exclusively to stamp impressions on storage jars during the reign of [[Hezekiah]].

The scarab remains an item of popular interest thanks to modern fascination with the art and beliefs of ancient Egypt. Scarab beads in semiprecious stones or glazed ceramics can be purchased at most bead shops, while at Luxor Temple a massive ancient scarab has been roped off to discourage visitors from rubbing the base of the statue "for luck".

===In literature===
In [[Aesop's fable]] "[[The Eagle and the Beetle]]", the eagle kills a hare that has asked for sanctuary with a beetle. The beetle then takes revenge by twice destroying the eagle's eggs. The eagle, in despair, flies up to [[Mount Olympus|Olympus]] and places her latest eggs in [[Zeus]]'s lap, beseeching the god to protect them. When the beetle finds out what the eagle has done, it stuffs itself with dung, goes straight up to Zeus and flies right into his face. Zeus is startled at the sight of the unpleasant creature, jumping to his feet so that the eggs are broken. Learning of the origin of their feud, Zeus attempts to mediate and, when his efforts to mediate fail, he changes the breeding season of the eagle to a time when the beetles are not above ground.

[[Aristophanes]] alluded to Aesop's fable several times in his plays. In ''[[Peace (play)|Peace]]'', the hero rides up to Olympus to free the goddess Peace from her prison. His steed is an enormous dung beetle which has been fed so much dung that it has grown to monstrous size.

[[Hans Christian Andersen]]'s "The Dung Beetle" tells the story of a dung beetle who lives in the stable of the king's horses in an imaginary kingdom. When he demands golden shoes like those the king's horse wears and is refused, he flies away and has a series of adventures, which are often precipitated by his [[Superiority complex|feeling of superiority]] to other animals. He finally returns to the stable having decided (against all logic) that it is for him that the king's horse wears golden shoes.<ref>{{cite book|last1=Andersen|first1=Hans Christian|last2=James|first2=M. R. (trans.)|work=Hans Andersen Forty-Two Stories | title=The Beetle |date=1930 |publisher=Faber and Faber |url=https://fadedpage.com/showbook.php?pid=20100119}}</ref>

In [[Franz Kafka]]'s ''[[The Metamorphosis]]'', the transformed character of Gregor Samsa is called an "old dung beetle" (''alter Mistkäfer'') by a [[charwoman]].

==See also==
{{portal|Insects}}
* ''[[Catharsius]]'', an important dung beetle genus in African and Asian environments
* [[Addo Elephant National Park]], site of the largest remaining population of the endangered flightless dung beetle (''Circellium bacchus'').
* [[List of dung beetle and chafer (Scarabaeoidea) species recorded in Britain]]
* [[Rotating locomotion in living systems]]

==References==
{{reflist|2}}

== Bibliography ==
{{refbegin}}
* {{citation|title=Science Year|first=Tony |last=Tills|publisher=World Book, Inc.|year=2013|isbn=9780716605676}}

{{refend}}

==External links==
{{wikispecies|Scarabaeinae}}
{{wikispecies|Aphodiinae}}
{{Commons category|Dung beetles}}
* [https://web.archive.org/web/20121105033930/http://www.cosmosmagazine.com/features/online/5803/on-a-roll Feature: 'What to do with too much poo'] – The success story behind the introduction of dung beetles in Australia at cosmosmagazine.com
* [https://web.archive.org/web/20161018110844/http://www.insects.org/ced1/beetles_rel_sym.html Beetles as religious symbols] at insects.org
* [http://www.scarabnet.org/ Scarabaeinae Research Network] {{Webarchive|url=https://web.archive.org/web/20190122100429/http://www.scarabnet.org/ |date=2019-01-22 }}
* [http://www.amonline.net.au/factsheets/dung-beetles.htm Dung beetles] at the Australia Museum
* [http://www.catharsius.fr/ Catharsius], an international group working on taxonomy and ecology of Western African dung beetles
* Tomas Libich, [http://www.goliathus.com/en/en-dung-beetle.html Congo dung beetle sp1] and [http://www.goliathus.com/en/en-dung-beetle-2.html Congo dung beetle sp2] photos
* [https://archive.today/20130416051320/http://www.wild.org/video/i-brake-for-dung-beetles/ Dung Beetles in action (video)] by The WILD Foundation/Boyd Norton
* [http://www.dungbeetle.org.nz/ Dung Beetles in New Zealand] (proposed release of dung beetles, with background research)
* [http://www.ted.com/talks/marcus_byrne_the_dance_of_the_dung_beetle.html Marcus Byrne The dance of the dung beetle] Ted conference about dung beetle behavior.
* [http://www.dungbeetles.com.au Dung Beetle Ecosystem Engineers] Dung Beetle Ecosystem Engineers – expanding the range of dung beetles in Australia and analysing their performance for livestock producers.

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[[Category:Coprophagous insects]]
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