Editing Domestication (section)

== Domestication by insects ==

At least three groups of insects, namely ambrosia beetles, leafcutter ants, and fungus-growing termites, have domesticated species of [[Fungus|fungi]].<ref name="Purugganan 2022"/><ref name="Mueller Gerardo 2005"/>

=== Ambrosia beetles ===

{{further|Ambrosia beetle}}

Ambrosia beetles in the [[weevil]] subfamilies [[Scolytinae]] and [[Platypodinae]] excavate tunnels in dead or stressed trees into which they introduce fungal gardens, their sole source of nutrition. After landing on a suitable tree, an ambrosia beetle excavates a tunnel in which it releases [[Ambrosia fungi|its fungal symbiont]]. The fungus penetrates the plant's [[xylem]] tissue, extracts nutrients from it, and concentrates the nutrients on and near the surface of the beetle gallery. Ambrosia fungi are typically poor wood degraders and instead utilize less demanding nutrients.<ref>{{cite journal |last1=Kasson |first1=Matthew T. |last2=Wickert |first2=Kristen L. |last3=Stauder |first3=Cameron M. |last4=Macias |first4=Angie M. |last5=Berger |first5=Matthew C. |last6=Simmons |first6=D. Rabern |last7=Short |first7=Dylan P. G. |last8=DeVallance |first8=David B. |last9=Hulcr |first9=Jiri |title=Mutualism with aggressive wood-degrading Flavodon ambrosius (Polyporales) facilitates niche expansion and communal social structure in Ambrosiophilus ambrosia beetles |journal=Fungal Ecology |date=October 2016 |volume=23 |pages=86–96 |doi=10.1016/j.funeco.2016.07.002 |doi-access=free |bibcode=2016FunE...23...86K }}</ref> Symbiotic fungi produce and detoxify ethanol, which is an attractant for ambrosia beetles and likely prevents the growth of antagonistic pathogens and selects for other beneficial symbionts.<ref>{{cite journal |last1=Ranger |first1=Christopher M. |last2=Biedermann |first2=Peter H. W. |last3=Phuntumart |first3=Vipaporn |last4=Beligala |first4=Gayathri U. |last5=Ghosh |first5=Satyaki |last6=Palmquist |first6=Debra E. |last7=Mueller |first7=Robert |last8=Barnett |first8=Jenny |last9=Schultz |first9=Peter B. |last10=Reding |first10=Michael E. |last11=Benz |first11=J. Philipp |title=Symbiont selection via alcohol benefits fungus farming by ambrosia beetles |journal=Proceedings of the National Academy of Sciences |date=24 April 2018 |volume=115 |issue=17 |pages=4447–4452 |doi=10.1073/pnas.1716852115 |pmc=5924889 |pmid=29632193 |bibcode=2018PNAS..115.4447R |doi-access=free }}</ref> Ambrosia beetles mainly colonise wood of recently dead trees.<ref name= Hulcr2017>{{cite journal |last1=Hulcr |first1=Jiri |last2=Stelinski |first2=Lukasz L. |title=The Ambrosia Symbiosis: From Evolutionary Ecology to Practical Management |journal=Annual Review of Entomology |date=31 January 2017 |volume=62 |pages=285–303 |doi=10.1146/annurev-ento-031616-035105 |pmid=27860522 |doi-access=free }}</ref>

=== Leafcutter ants ===

{{further|Leafcutter ant}}

The leafcutter ants are any of some 47 species of leaf-chewing ants in the genera ''[[Acromyrmex]]'' and ''[[Atta (genus)|Atta]]''. The ants carry the discs of leaves that they have cut back to their nest, where they feed the leaf material to the fungi that they tend. Some of these fungi are not fully domesticated: the fungi farmed by ''[[Mycocepurus smithii]]'' constantly produce spores that are not useful to the ants, which eat fungal [[hypha]]e instead. The process of domestication by ''Atta'' ants, on the other hand, is complete; it took 30 million years.<ref name="Shik Gomez Kooij 2016">{{cite journal |last1=Shik |first1=Jonathan Z. |last2=Gomez |first2=Ernesto B. |last3=Kooij |first3=Pepijn W. |last4=Santos |first4=Juan C. |last5=Wcislo |first5=William T. |last6=Boomsma |first6=Jacobus J. |title=Nutrition mediates the expression of cultivar–farmer conflict in a fungus-growing ant |journal=Proceedings of the National Academy of Sciences |volume=113 |issue=36 |date=6 September 2016 |pmid=27551065 |pmc=5018747 |doi=10.1073/pnas.1606128113 |pages=10121–10126 |doi-access=free |bibcode=2016PNAS..11310121S }}</ref>

=== Fungus-growing termites ===

{{further|Macrotermitinae|Termitomyces}}

Some 330 fungus-growing termite species of the subfamily [[Macrotermitinae]] cultivate ''[[Termitomyces]]'' fungi to eat; domestication occurred exactly once, 25–40 [[Million years ago|mya]].<ref name="Purugganan 2022"/><ref name="Mueller Gerardo 2005">{{cite journal |last1=Mueller |first1=Ulrich G. |last2=Gerardo |first2=Nicole M. |last3=Aanen |first3=Duur K. |last4=Six |first4=Diana L. |last5=Schultz |first5=Ted R. |title=The Evolution of Agriculture in Insects |journal=Annual Review of Ecology, Evolution, and Systematics |volume=36 |issue=1 |date=1 December 2005 |doi=10.1146/annurev.ecolsys.36.102003.152626 |pages=563–595 |url=https://www.academia.edu/download/51001206/The_Evolution_of_Agriculture_in_Insects20161221-18052-s5ycda.pdf<!--not redundant to DOI-->}}</ref> The fungi, described by [[Roger Heim]] in 1942, grow on 'combs' formed from the termites' excreta, dominated by tough woody fragments.<ref name="Heim 1942">{{cite journal |last=Heim |first=Roger |author-link=Roger Heim |title=Nouvelles études descriptives sur les agarics termitophiles d'Afrique tropicale |journal=Archives du Muséum National d'Histoire Naturelle |language=French |trans-title=New Descriptive Studies on the Termitophile Mushrooms of Tropical Africa |year=1942 |volume=18 |issue=6 |pages=107–166}}</ref> The termites and the fungi are both obligate [[symbiosis|symbionts]] in the relationship.<ref name="NobreAanen2010">{{cite journal |last1=Nobre |first1=T. |last2=Aanen |first2=D. K. |journal=Communicative & Integrative Biology |title=Dispersion and colonisation by fungus-growing termites |volume=3 |issue=3 |pages=248–250 |date=1 May 2010 |url=https://doi.org/10.4161/cib.3.3.11415 |doi=10.4161/cib.3.3.11415 |pmid=20714406 |pmc=2918769 }}</ref>

<gallery class=center mode=nolines widths=180 heights=180 caption="Domestication by insects">
File:Xylosandrus crassiusculus galleryR.jpg|Gallery of the ambrosia beetle ''[[Xylosandrus crassiusculus]]'' split open, with pupae and black fungus. The fungus decomposes materials in the wood, providing food for the beetles.
File:leaf cutter ants arp.jpg|Leafcutter ants ''[[Atta cephalotes]]'' carrying discs of leaf material back to their nest to feed to their domesticated fungus
Ancistrotermes latinotus.jpg|Inside the nest of the fungus-cultivating termite ''[[Ancistrotermes]]''
File:Termitomyces heimii.jpg|''[[Termitomyces heimii]]'' growing on 'comb' inside a termite mound
File:Termitomyces reticulatus 37340.jpg|''[[Termitomyces]]'' fungi are mutually dependent on [[Macrotermitinae]] termites for their survival.
</gallery>