Leucocoprinus gongylophorus
| Leucocoprinus gongylophorus | |
|---|---|
| |
Scientific classification
| |
| Domain: | Eukaryota |
| Kingdom: | Fungi |
| Division: | Basidiomycota |
| Class: | Agaricomycetes |
| Order: | Agaricales |
| Family: | Agaricaceae |
| Genus: | Leucocoprinus |
| Species: | L. gongylophorus
|
| Binomial name | |
| Leucocoprinus gongylophorus (Alfred Möller)Heim(1957)
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| Synonyms | |
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Rozites gongylophorusMöller(1893) | |
Leucocoprinus gongylophorusis afungusin the familyAgaricaceaewhich is cultivated by certainleafcutter ants.[1]Like other species of fungi cultivated by ants,L. gongylophorusproducesgongylidia,nutrient-richhyphalswellings upon which the ants feed.[2]Production ofmushroomsoccurs only once ants abandon the nest.[3]L. gongylophorusis farmed by leaf cutter ant species belonging to the generaAttaandAcromyrmex,amongst others.
Description
[edit]L. gongylophorusis completely dependent on ants for survival with workers feeding it cut plant matter and new queens carrying a piece ofmyceliumin their infrabuccal pocket (a specialised structure within the mouth) in order to found a new colony. For both its source of nutrition and mechanism of spreading it is reliant on the ants. The species has co-evolved with ants so thoroughly that it can no longer rely upon producing mushrooms to emit spores as a viable survival mechanism as it has lost the ability to produce sexual spores.[4]The mutualistic relationship also includes a third species asPseudonocardiabacteria coat the exoskeleton of the ants and produce antibiotics which protect the fungal garden frommycoparasiticmicrofungilike members of theEscovopsisgenus. Without this antibiotic production theL. gongylophorusfungi on which the ants rely could be destroyed and take the whole nest with it. Leafcutter ant species may exhibit specially modified exoskeletons better capable of housing beneficial bacteria with which the fungal garden is inoculated as the ants tend to it. A fourth hypothesized beneficial symbiont exists in the form ofPhialophorablack yeast which grow on the cuticle of the ants and which may fulfill a beneficialsaprophyticrole. Including the parasiticEscovopsismicrofungi ormicromycetesthis means a leafcutter ant colony may in fact be asymbiotic relationshipbetween five species.[5][6]The fungal lineage ofL. gongylophorusmay have been continually propagated for over 23 million years which helps explain why it is now entirely reliant on the ants.[7]
L. gongylophorusproduces a diverse array of enzymes to facilitate lignocellulose degradation in ant gardens. As leafcutter ants can have such a profound effect on their environment and account for about 25% of plant consumption in their forest ecosystems[8]this meansL. gongylophorusmay be the primary driver of plant biomass degradation in this ecosystem.[9]The sheer scale of the ants' farming can see 10-15% of leaves in a colony's foraging range being cut and fed to the fungi[8]which results in significant carbon dioxide emissions fromL. gongylophorusas this plant biomass is broken down. Nest openings can have CO2emissions 100,000 times greater than the surrounding soils with nests and their surrounding soil emitting 15-60% more CO2than soil without nests. Emissions were observable for more than two years after nests were abandoned with nests contributing an estimated total of 0.2-0.7% of the ecosystem's soil emissions.[10]Due to the enhanced rate of breakdown of the plant biomass facilitated by the fungus compared to the slower natural degradation of leaves that fall on the surface, leafcutter ants and their symbiotic fungi partners may play a surprisingly significant role in the ecosystem's ability to sequester carbon.
This species has become so dependent on the ants to survive that the production of mature fruiting bodies is not often recorded. In the wild it is heavily suppressed by the ants due to their harvesting of the immature mushrooms. InMøller's 1893 study he described the mushrooms ofRozitesgongylophorusgrowing from a nest removed from the wild as follows:
Cap:Up to 16cm in diameter.Stem and cap:Up to 24cm in height.Stem:Up to 4cm diameter at the base and 2cm at the annulus.Spore print:Tinted with a clear ochre colour (possibly due to discolouration with age sinceLeucoagaricusspecies typically have white spores).[11]
In the 1977 publicationStudies on Fungi Cultivated by Antsattempts were made to culture 37 fungi samples obtained from the nests of various fungus farming ants. This study succeeded in producing mushrooms on an oatmeal medium from samples acquired from the nests ofMyrmicocrypta buenzlii,Mycetophylax conformis,Cyphomyrmex costatusandApterostigma auriculatumwhich were described as appearing similar to Møller's observation.[11]However whilst these lower attine ants do farm otherLeucocoprinusspecies, they do not farm the highly specialisedL. gongylophorus.[12]The cultures taken fromAttaandAcromyrmexnests either producedgongylidiaor only mycelium.[11]
Later studies have observed mushrooms fromAttaandAcromyrmexnests growing in the lab or in wild nests during excavations.[13]
Taxonomy
[edit]Originally classified asRozites gongylophorusby the German botanist and mycologistFriedrich Alfred Gustav Jobst Møllerin 1893 who described it as an ant fungus associated withAtta discigera(since reclassified asAcromyrmex disciger)leafcutter ants in Brazil.[11]
It was reclassified asLeucocoprinus gongylophorusby French mycologistRoger Heimin 1957. Heim obtained samples of sterile ant fungi from French Guiana and Panama fromAttaant species which were compared with illustrations and descriptions of an ant fungus obtained fromCyphomyrmex costatusants in Trinidad. Based on this study the fungus was transferred to theLeucocoprinusgenus. It was later transferred toLeucoagaricusby the German mycologistRolf Singerin 1986. More recent studies based upon DNA sequence data fail to differentiate the generaLeucoagaricusandLeucocoprinus.[14][15]Consequently, the type ofLeucoagaricuswas transferred toLeucocoprinusandLeucoagaricusbecomes a synonym ofLeucocoprinus,the older of the two generic names. As a result the nameLeucocoprinus gongylophorusbecomes the correct name.
It was also classified asPholiota gongylophoraby Italian botanist and mycologistPier Andrea Saccardoin 1895 andAttamyces bromatificusby German mycologistHanns Kreiselin 1972 however these species were merged withL. gongylophorus. Attamyceswas amonotypicgenus described by Kreisel after he found the fungus fruiting on the nest of the fungus-growing antsAtta insularisin Cuba. Thespecific epithetbromatificusrefers to bromatia, which are swollen tips on the hyphae that the ants use as food.[16]Bromatia are also calledgongylidiawith 'bromatia' seldom used in modern descriptions so this word could now be considered archaic.
Etymology
[edit]Leucocoprinusgets its name from the GreekLeucomeaning white or bright and the Latin or GreekCoprinusrefers to its look-a-like genusCoprinus.[17]Gongylophorusderives from the Greekgongylomeaning rounded and Greekphorusmeaning bearing or carrying.[18]
Similar species
[edit]Termitomycesis a genus of mushrooms also belonging to the orderAgaricaleswhich is cultivated by various species belonging to the subfamily of termites,macrotermitinae.Whilst not immediately related to species cultivated by leafcutter ants they are noteworthy as another insect-fungus partnership.
Myrmecopterulais a genus of coral fungi belonging to the orderAgaricaleswhich is cultivated byApterostigmaants.
See also
[edit]References
[edit]- ^Aylward FO, Burnum-Johnson KE, Tringe SG, Teiling C, Tremmel DM, Moeller JA, et al. (June 2013)."Leucoagaricus gongylophorus produces diverse enzymes for the degradation of recalcitrant plant polymers in leaf-cutter ant fungus gardens".Applied and Environmental Microbiology.79(12): 3770–8.doi:10.1128/AEM.03833-12.PMC3675943.PMID23584789.
- ^De Fine Licht HH, Boomsma JJ, Tunlid A (December 2014)."Symbiotic adaptations in the fungal cultivar of leaf-cutting ants".Nature Communications.5:5675.Bibcode:2014NatCo...5.5675D.doi:10.1038/ncomms6675.PMID25435021.
- ^McCoy, Peter (2016).Radical Mycology: A Treatise on Seeing & Working With Fungi.Chthaeus Press. p. 215.ISBN978-0986399602.
- ^Espinoza, César; Zavala Izquierdo, Inés; Couttolenc, Alan; Landa-Cadena, Gandhi; Valenzuela, Jorge; Trigos, Ángel; Espinoza, César; Zavala Izquierdo, Inés; Couttolenc, Alan; Landa-Cadena, Gandhi; Valenzuela, Jorge (2017)."In vitro isolation and identification of Leucoagaricus gongylophorus from Atta mexicana (Hymenoptera:Formicidae) fungal garden".Revista mexicana de micología.46:3–8.ISSN0187-3180.
- ^"Fungal Farming in Leafcutter Ants".reed.edu.Retrieved2022-07-07.
- ^Little, Ainslie E.F; Currie, Cameron R (2007-10-22)."Symbiotic complexity: discovery of a fifth symbiont in the attine ant–microbe symbiosis".Biology Letters.3(5): 501–504.doi:10.1098/rsbl.2007.0253.ISSN1744-9561.PMC2396185.PMID17686758.
- ^Chapela, Ignacio H.; Rehner, Stephen A.; Schultz, Ted R.; Mueller, Ulrich G. (1994-12-09)."Evolutionary History of the Symbiosis Between Fungus-Growing Ants and Their Fungi".Science.266(5191): 1691–1694.Bibcode:1994Sci...266.1691C.doi:10.1126/science.266.5191.1691.ISSN0036-8075.PMID17775630.S2CID22831839.
- ^abSwanson, Amanda C.;Schwendenmann, Luitgard;Allen, Michael F.; Aronson, Emma L.; Artavia-León, Allan; Dierick, Diego; Fernandez-Bou, Angel S.; Harmon, Thomas C.; Murillo-Cruz, Catalina; Oberbauer, Steven F.; Pinto-Tomás, Adrián A. (August 2019). Bennett, Alison (ed.)."Welcome to the Atta world: A framework for understanding the effects of leaf-cutter ants on ecosystem functions".Functional Ecology.33(8): 1386–1399.doi:10.1111/1365-2435.13319.ISSN0269-8463.S2CID92669434.
- ^Aylward, Frank O.; Burnum-Johnson, Kristin E.; Tringe, Susannah G.; Teiling, Clotilde; Tremmel, Daniel M.; Moeller, Joseph A.; Scott, Jarrod J.; Barry, Kerrie W.; Piehowski, Paul D.; Nicora, Carrie D.; Malfatti, Stephanie A. (2013-06-15)."Leucoagaricus gongylophorus Produces Diverse Enzymes for the Degradation of Recalcitrant Plant Polymers in Leaf-Cutter Ant Fungus Gardens".Applied and Environmental Microbiology.79(12): 3770–3778.doi:10.1128/AEM.03833-12.ISSN0099-2240.PMC3675943.PMID23584789.
- ^Fernandez-Bou, A. S.; Dierick, D.; Swanson, A. C.; Allen, M. F.; Alvarado, A. G. F.; Artavia-León, A.; Carrasquillo-Quintana, O.; Lachman, D. A.; Oberbauer, S.; Pinto-Tomás, A. A.; Rodríguez-Reyes, Y. (February 2019)."The Role of the Ecosystem Engineer, the Leaf-Cutter Ant Atta cephalotes, on Soil CO 2 Dynamics in a Wet Tropical Rainforest".Journal of Geophysical Research: Biogeosciences.124(2): 260–273.doi:10.1029/2018JG004723.hdl:10669/79895.ISSN2169-8953.S2CID134458614.
- ^abcdHervey, Annette; Rogerson, Clark T.; Leong, Ina (1977-04-01)."Studies on fungi cultivated by ants".Brittonia.29(2): 226–236.doi:10.2307/2805856.ISSN1938-436X.JSTOR2805856.S2CID41247049.
- ^Mueller, Ulrich G.; Kardish, Melissa R.; Ishak, Heather D.; Wright, April M.; Solomon, Scott E.; Bruschi, Sofia M.; Carlson, Alexis L.; Bacci, Maurício (2018)."Phylogenetic patterns of ant-fungus associations indicate that farming strategies, not only a superior fungal cultivar, explain the ecological success of leafcutter ants".Molecular Ecology.27(10): 2414–2434.doi:10.1111/mec.14588.ISSN1365-294X.PMID29740906.S2CID13702174.
- ^Mueller, Ulrich G. (2002)."Ant versus Fungus versus Mutualism: Ant-Cultivar Conflict and the Deconstruction of the Attine Ant-Fungus Symbiosis".The American Naturalist.160(S4): S67–S98.doi:10.1086/342084.ISSN0003-0147.JSTOR10.1086/342084.PMID18707454.S2CID21576198.
- ^Vellinga EC (2004). "Genera in the family Agaricaceae: evidence from nrITS and nrLSU sequences".Mycol. Res.108(Pt 4): 354–377.doi:10.1017/S0953756204009700.PMID15209277.
- ^Vellinga EC, Sysouphanthong P, Hyde KD (2011). "The family Agaricaceae: phylogenies and two new white-spored genera".Mycologia.103(3): 494–509.doi:10.3852/10-204.PMID21193599.
- ^Kreisel, H. (1972)."Pilze aus Pilzgärten von Atta insularis in Kuba".Zeitschrift für allgemeine Mikrobiologie.12(8): 643–654.doi:10.1002/jobm.19720120805.
- ^"Fungi naming, author citation and biographies".first-nature.Retrieved2022-07-07.
- ^"Botanical Latin (L) & Greek (G)"(PDF).