View source for Epichloë

{{Redirect|Choke disease||Choke (disambiguation){{!}}Choke}}
{{Italic title}}
{{Taxobox
| name = ''Epichloë''
| image = Epichloe_typhina.JPG
| image_caption = "Choke disease": ''[[Epichloë typhina]]'' stroma on [[Poa pratensis|bluegrass]]
| regnum = [[Fungus|Fungi]]
| divisio = [[Ascomycota]]
| classis = [[Ascomycetes]]
| subclassis = [[Sordariomycetes]]
| ordo = [[Hypocreales]]
| familia = [[Clavicipitaceae]]
| tribus = [[Balansiae]]
| genus = '''''Epichloë'''''
| genus_authority = ([[Elias Magnus Fries|Fr.]]) [[Louis René Tulasne|Tul.]] & [[Charles Tulasne|C. Tul]]
| type_species = ''[[Epichloë typhina]]''
| type_species_authority = (Fr.) Tul. & C. Tul.
| diversity = 35 species, see text
}}

'''''Epichloë''''' is a genus of [[Ascomycota|ascomycete]] [[Fungus|fungi]] forming an [[Endophyte|endophytic]] [[symbiosis]] with [[Poaceae|grasses]]. Grass '''choke disease''' is a symptom in grasses induced by some ''Epichloë'' species, which form [[spore (mycology)|spore]]-bearing mats ([[Stroma (tissue)|stromata]]) on [[tiller (botany)|tiller]]s and suppress the development of their [[host plant]]'s [[inflorescence]]. For most of their life cycle however, ''Epichloë'' grow in the intercellular space of stems, leaves, inflorescences, and seeds of the grass plant without incurring symptoms of disease. In fact, they provide several benefits to their host, including the production of different herbivore-deterring [[alkaloids]], increased stress resistance, and growth promotion.

Within the family [[Clavicipitaceae]], ''Epichloë'' is embedded in a group of endophytic and [[plant pathogen]]ic fungi, whose common ancestor probably derived from an animal pathogen. The genus includes both species with a sexually reproducing ([[teleomorph]]ic) stage and [[asexual reproduction|asexual]], [[anamorph]]ic species. The latter were previously placed in the [[form genus]] ''[[Neotyphodium]]'' but included in ''Epichloë'' after [[molecular phylogenetics]] had shown asexual and sexual species to be intermingled in a single [[clade]]. [[Hybrid speciation]] has played an important role in the evolution of the genus.

''Epichloë'' species are ecologically significant through their effects on host plants. Their presence has been shown to alter the composition of [[plant communities]] and [[food web]]s. Grass varieties, especially of [[tall fescue]] and [[ryegrass]], with symbiotic ''Epichloë'' endophyte strains, are commercialised and used for [[pasture]] and [[turf]].

==Taxonomy==

[[Elias Fries]], in 1849, first defined ''Epichloë'' as a [[subgenus]] of ''[[Cordyceps]]''.<ref name="Fries1849"/> As [[type species]], he designated ''[[Epichloë typhina|Cordyceps typhina]]'',<ref name="Fries1849"/> originally described by [[Christiaan Hendrik Persoon]].<ref name="Persoon1798"/> The brothers [[Charles Tulasne|Charles]] and [[Louis René Tulasne]] then raised the subgenus to genus rank in 1865.<ref name="Tulasne1865"/> ''Epichloë typhina'' would remain the only species in the genus until the discovery of fungal grass endophytes causing livestock intoxications in the 1970s and 1980s, which stimulated the description of new species.<ref name="Leuchtmann2014"/> Several species from Africa and Asia that develop stromata on grasses were split off as a separate genus ''[[Parepichloë]]'' in 1998.<ref name="WhiteReddy1998"/>

Many ''Epichloë'' species have forms that reproduce sexually, and several purely asexual species are closely related to them. These [[anamorphs]] were long classified separately: Morgan-Jones and Gams (1982) collected them in a section (''Albo-lanosa'') of genus ''[[Acremonium]]''.<ref name="Morgan-Jones1982"/> In a [[molecular phylogenetic]] study in 1996, Glenn and colleagues found the genus to be [[polyphyletic]] and proposed a new genus ''[[Neotyphodium]]'' for the anamorphic species related to ''Epichloë''.<ref name="Glenn1996"/> A number of species continued to be described in both genera until Leuchtmann and colleagues (2014) included most of the [[form genus]] ''Neotyphodium'' in ''Epichloë''.<ref name="Leuchtmann2014"/> Phylogenetic studies had shown both genera to be intermingled, and the [[International Code of Nomenclature for algae, fungi, and plants|nomenclatural code]] required since 2011 that one single name be used for all stages of development of a fungal species. Only ''Neotyphodium starrii'', of unclear status, and ''[[Neotyphodium chilense|N. chilense]]'', which is unrelated, were excluded from ''Epichloë''.<ref name="Leuchtmann2014"/>

<imagemap>
File:Epichloë tubB phylogeny.jpg|center|750px|alt=Phylogeny of the fungal genus Epichloë from aligned tubB gene sequences.|Phylogeny of the genus *Epichloë*. Clicking on a species in the phylogeny will load the appropriate wikipedia article.

desc bottom-left

rect 844 337  1195 377  [[w:Epichloë amarillans|Epichloë amarillans]]
rect 605 689  977  728  [[w:Epichloë aotearoae|Epichloë aotearoae]]
rect 931 534  1412 578  [[w:Epichloë baconii|Epichloë baconii]]
rect 606 636  1022 682  [[w:Epichloë brachyelytri|Epichloë brachyelytri]]
rect 495 234  1206 277  [[w:Epichloë bromicola|Epichloë bromicola]]
rect 496 282  796  332  [[w:Epichloë elymi|Epichloë elymi]]
rect 902 384  1454 475  [[w:Epichloë festucae|Epichloë festucae]]
rect 432 84   1105 180  [[w:Epichloë gansuensis|Epichloë gansuensis]]
rect 433 184  942  233  [[w:Epichloë glyceriae|Epichloë glyceriae]]
rect 902 485  1170 528  [[w:Epichloë mollis|Epichloë mollis]]
rect 433 32 985 75      [[w:Epichloë sibirica|Epichloë sibirica]]
rect 932 586  1560 632  [[w:Epichloë stromatolonga|Epichloë stromatolonga]]
rect 841 938  1581 1032 [[w:Epichloë sylvatica|Epichloë sylvatica]]
rect 771 736  1720 931  [[w:Epichloë typhina|Epichloë typhina]]
rect 924 1037 1716 1131 [[w:Epichloë typhina|Epichloë typhina]]
</imagemap>

==Species==

As of 2020, there are 35 accepted species in the genus, with 3 [[subspecies]] and 6 [[variety (botany)|varieties]] described.  14 species, 3 subspecies and 5 varieties are haploid.  21 species and 1 variety are hybrids ([[Polyploidy#Allopolyploidy|allopolyploids]]). Several taxa are only known as anamorphic (asexual) forms, most of which have previously been classified in ''Neotyphodium''.<ref name="Leuchtmann2014"/>

{| class="wikitable collapsible sortable"
! scope="col" style="width: 200px;" | Haploid Taxa !! Known Distribution !! Sexual Reproduction !! Vertical Transmission !! Known Host Range !! scope="col" style="width: 350px;" | Reference to Species Description
|-
! '''''[[Epichloë amarillans]]'''''
J.F. White
| North America || Observed || Present || ''[[Agrostis]]'' ''hyemalis'', ''[[Agrostis perennans]]'', ''[[Calamagrostis canadensis]]'', ''[[Elymus virginicus]]'', ''[[Sphenopholis]]'' ''nitida'', ''[[Sphenopholis obtusata]]'', ''[[Sphenopholis]]'' ''× pallens'', ''[[Ammophila breviligulata]]'' || {{cite journal|last1=White|first1=James F.|title=Endophyte-host associations in grasses. XX. Structural and reproductive studies of ''Epichloë amarillans'' sp. nov. and comparisons to ''E. typhina''|journal=Mycologia|volume=86|issue=4|year=1994|pages=571–580|issn=0027-5514|doi=10.1080/00275514.1994.12026452}}
|-
! '''''[[Epichloë aotearoae]]'''''
(C.D. Moon, C.O. Miles & Schardl) Leuchtm. & Schardl
| New Zealand, Australia || Not observed || Present || ''[[Echinopogon]]'' ''ovatus'' || {{cite journal|last1=Moon|first1=Christina D.|last2=Miles|first2=Christopher O.|last3=Järlfors|first3=Ulla|last4=Schardl|first4=Christopher L.|title=The evolutionary origins of three new ''Neotyphodium'' endophyte species from grasses indigenous to the Southern Hemisphere|journal=Mycologia|volume=94|issue=4|year=2002|pages=694–711|issn=0027-5514|doi=10.1080/15572536.2003.11833197}}
|-
! '''''[[Epichloë baconii]]'''''
J.F. White
| Europe || Observed || Absent || ''[[Agrostis capillaris]]'', ''[[Agrostis stolonifera]]'', ''[[Calamagrostis villosa]]'', ''[[Calamagrostis varia]]'', ''[[Calamagrostis]]'' ''purpurea'' || {{cite journal|last1=White|first1=James F.|title=Endophyte-host associations in grasses. XIX. A systematic study of some sympatric species of ''Epichloë'' in England|journal=Mycologia|volume=85|issue=3|year=1993|pages=444–455|issn=0027-5514|doi=10.1080/00275514.1993.12026295}}
|-
! '''''[[Epichloë brachyelytri]]'''''
Schardl & Leuchtm.
| North America || Observed || Present || ''[[Brachyelytrum]]'' ''erectum'' || {{cite journal|last1=Schardl|first1=Christopher L.|last2=Leuchtmann|first2=Adrian|title=Three new species of ''Epichloë'' symbiotic with North American grasses|journal=Mycologia|volume=91|issue=1|year=1999|pages=95–107|issn=0027-5514|doi=10.1080/00275514.1999.12060996}}
|-
! '''''[[Epichloë bromicola]]'''''
Leuchtm. & Schardl
| Europe, Asia || Observed on ''[[Bromus erectus]]'', ''[[Elymus repens]]'' and [[Elymus (plant)|''Roegneria'']] ''kamoji'' || Present in ''[[Bromus]]'' ''benekenii'', ''[[Bromus ramosus]]'' and ''[[Hordelymus]]'' ''europaeus'', ''[[Hordeum]]'' ''brevisubulatum'', ''[[Leymus chinensis]]'' and [[Elymus (plant)|''Roegneria'']] ''kamoji''; absent in ''[[Bromus erectus]]'' and ''[[Elymus repens]]'' || Europe: ''[[Bromus]]'' ''benekenii'', ''[[Bromus erectus]]'', ''[[Bromus ramosus]]'', ''[[Elymus repens]]'', ''[[Hordelymus]]'' ''europaeus'', ''[[Hordeum]]'' ''brevisubulatum''. Asia: ''[[Leymus chinensis]]'', [[Elymus (plant)|''Roegneria'']] ''kamoji'' || {{cite journal|last1=Leuchtmann|first1=Adrian|last2=Schardl|first2=Christopher L.|title=Mating compatibility and phylogenetic relationships among two new species of ''Epichloë'' and other congeneric European species|journal=Mycological Research|volume=102|issue=10|year=1998|pages=1169–1182|issn=09537562|doi=10.1017/S0953756298006236}}
|-
! '''''[[Epichloë elymi]]'''''
Schardl & Leuchtm.
| North America || Observed || Present || ''[[Bromus kalmii]]'', [[Elymus (plant)|''Elymus'']] spp. (including [[Elymus hystrix|''Elymus patula'']]) || {{cite journal|last1=Schardl|first1=Christopher L.|last2=Leuchtmann|first2=Adrian|title=Three new species of ''Epichloë'' symbiotic with North American grasses|journal=Mycologia|volume=91|issue=1|year=1999|pages=95–107|issn=0027-5514|doi=10.1080/00275514.1999.12060996}}
|-
! '''''[[Epichloë festucae]]'''''
Leuchtm., Schardl & M.R. Siegel
| Europe, Asia, North America || Observed || Present || ''[[Festuca]]'' spp., ''[[Koeleria]]'' spp., ''Schedonorus'' spp. || {{cite journal|last1=Leuchtmann|first1=Adrian|last2=Schardl|first2=Christopher L.|last3=Siegel|first3=Malcolm R.|title=Sexual compatibility and taxonomy of a new species of ''Epichloë'' symbiotic with fine fescue grasses|journal=Mycologia|volume=86|issue=6|year=1994|pages=802–812|issn=0027-5514|doi=10.1080/00275514.1994.12026487}}
|-
! [[Epichloë festucae|'''''Epichloë festucae''''' '''var.''' '''''lolii''''']]
(Latch, M.J. Chr. & Samuels) C.W. Bacon & Schardl
| Europe, Asia, North Africa, introduced in New Zealand, Australia and elsewhere || Not observed || Present || [[Lolium perenne|''Lolium perenne'' subsp. ''perenne'']] || {{cite journal|last1=Latch|first1=G.C.M.|last2=Christensen|first2=M.J.|last3=Samuels|first3=G.J.|title=Five endophytes of ''Lolium'' and ''Festuca'' in New Zealand|journal=Mycotaxon|volume=20|year=1984|pages=535–550}}
|-
! '''''[[Epichloë gansuensis]]'''''
(C.J. Li & Nan) Schardl
| Asia || Not observed || Present || ''[[Achnatherum]]'' ''inebrians'', ''[[Achnatherum]]'' ''sibiricum'', ''[[Achnatherum pekinense]]'' || {{cite journal|last1=Li|first1=C.J.|last2=Nan|first2=Z.B.|last3=Paul|first3=V.H.|last4=Dapprich|first4=P.D.|last5=Liu|first5=Y.|title=A new ''Neotyphodium'' species symbiotic with drunken horse grass (''Achnatherum inebrians'') in China|journal=Mycotaxon|volume=90|year=2004|pages=141–147}}
|-
! [[Epichloë gansuensis|'''''Epichloë gansuensis''''' '''var.''' '''''inebrians''''']]
(C.D. Moon & Schardl) Schardl
| Asia || Not observed || Present || ''[[Achnatherum]]'' ''inebrians'' || {{cite journal|last1=Moon|first1=Christina D.|last2=Guillaumin|first2=Jean-Jacques|last3=Ravel|first3=Catherine|last4=Li|first4=Chunjie|last5=Craven|first5=Kelly D.|last6=Schardl|first6=Christopher L.|title=New ''Neotyphodium'' endophyte species from the grass tribes Stipeae and Meliceae|journal=Mycologia|volume=99|issue=6|year=2007|pages=895–905|issn=0027-5514|doi=10.1080/15572536.2007.11832521}}
|-
! '''''[[Epichloë glyceriae]]'''''
Schardl & Leuchtm.
| North America || Observed || Absent || ''[[Glyceria]]'' spp. || {{cite journal|last1=Schardl|first1=Christopher L.|last2=Leuchtmann|first2=Adrian|title=Three new species of ''Epichloë'' symbiotic with North American grasses|journal=Mycologia|volume=91|issue=1|year=1999|pages=95–107|issn=0027-5514|doi=10.1080/00275514.1999.12060996}}
|-
! '''''[[Epichloë mollis]]'''''
(Morgan-Jones & W. Gams) Leuchtm. & Schardl
| Europe || Observed || Present || ''[[Holcus mollis]]'' || {{cite journal|last1=Morgan-Jones|first1=G.|last2=Gams|first2=W.|title=Notes on hyphomycetes. XLI. An endophyte of ''Festuca arundinacea'' and the anamorph of ''Epichloë typhina'', new taxa in one of two new sections of ''Acremonium''.|journal=Mycotaxon|volume=15|year=1982|pages=311-318|issn=0093-4666}}
|-
! '''''[[Epichloë sibirica]]'''''
(X. Zhang & Y.B. Gao) Tadych
| Asia || Not observed || Present || ''[[Achnatherum]]'' ''sibiricum'' || {{cite journal|last1=Zhang|first1=Xin|last2=Ren|first2=An-Zhi|last3=Wei|first3=Yu-Kun|last4=Lin|first4=Feng|last5=Li|first5=Chuan|last6=Liu|first6=Zhi-Jian|last7=Gao|first7=Yu-Bao|title=Taxonomy, diversity and origins of symbiotic endophytes of ''Achnatherum sibiricum'' in the Inner Mongolia Steppe of China|journal=FEMS Microbiology Letters|volume=301|issue=1|year=2009|pages=12–20|issn=03781097|doi=10.1111/j.1574-6968.2009.01789.x|doi-access=free}}
|-
! '''''[[Epichloë stromatolonga]]'''''
(Y.L. Ji, L.H. Zhan & Z.W. Wang) Leuchtm.
| Asia || Not observed || Present || ''[[Calamagrostis epigejos]]'' || {{cite journal|last1=Ji|first1=Yan-ling|last2=Zhan|first2=Li-hui|last3=Kang|first3=Yan|last4=Sun|first4=Xiang-hui|last5=Yu|first5=Han-shou|last6=Wang|first6=Zhi-wei|title=A new stromata-producing ''Neotyphodium'' species symbiotic with clonal grass ''Calamagrostis epigeios'' (L.) Roth. grown in China|journal=Mycologia|volume=101|issue=2|year=2009|pages=200–205|issn=0027-5514|doi=10.3852/08-044}}
|-
! '''''[[Epichloë sylvatica]]'''''
Leuchtm. & Schardl
| Europe, Asia || Observed || Present || ''[[Brachypodium sylvaticum]]'', ''[[Hordelymus]]'' ''europaeus'' || {{cite journal|last1=Leuchtmann|first1=Adrian|last2=Schardl|first2=Christopher L.|title=Mating compatibility and phylogenetic relationships among two new species of ''Epichloë'' and other congeneric European species|journal=Mycological Research|volume=102|issue=10|year=1998|pages=1169–1182|issn=09537562|doi=10.1017/S0953756298006236}}
|-
! [[Epichloë sylvatica|'''''Epichloë sylvatica''''' '''subsp.''' '''''pollinensis''''']]
Leuchtm. & M. Oberhofer
| Europe || Observed || Present || ''[[Hordelymus]]'' ''europaeus'' || {{cite journal|last1=Leuchtmann|first1=Adrian|last2=Oberhofer|first2=Martina|title=The ''Epichloë'' endophytes associated with the woodland grass ''Hordelymus europaeus'' including four new taxa|journal=Mycologia|volume=105|issue=5|year=2013|pages=1315–1324|issn=0027-5514|doi=10.3852/12-400}}
|-
! '''''[[Epichloë typhina]]'''''
(Pers.) Tul. & C. Tul.
| Europe, introduced in North America and elsewhere || Observed || Present in ''[[Puccinellia distans]]''; absent in other hosts || ''[[Anthoxanthum odoratum]]'', ''[[Brachypodium]]'' ''phoenicoides'', ''[[Brachypodium pinnatum]]'', ''[[Dactylis glomerata]]'', ''[[Lolium perenne]]'', ''[[Milium effusum]]'', [[Timothy-grass|''Phleum pratense'']], ''[[Poa trivialis]]'', ''[[Poa]]'' ''silvicola'', ''[[Puccinellia distans]]'' || {{cite journal|last1=Tulasne|first1=L.R.|last2=Tulasne|first2=C.|title=Nectriei-Phacidiei-Pezizei|journal=Selecta Fungorum Carpologia|volume=3|year=1865|pages=24|publisher=Imperial: Paris}}
|-
! [[Epichloë typhina|'''''Epichloë typhina''''' '''subsp.''' '''''clarkii''''']]
(J.F. White) Leuchtm. & Schardl
| Europe || Observed || Absent || ''[[Holcus lanatus]]'' || {{cite journal|last1=White|first1=James F.|title=Endophyte-host associations in grasses. XIX. A systematic study of some sympatric species of ''Epichloë'' in England|journal=Mycologia|volume=85|issue=3|year=1993|pages=444–455|issn=0027-5514|doi=10.1080/00275514.1993.12026295}}
|-
! [[Epichloë typhina|'''''Epichloë typhina''''' '''subsp.''' '''''poae''''']]
(Tadych, K.V. Ambrose, F.C. Belanger & J.F. White) Tadych
| Europe, North America || Observed on ''[[Poa nemoralis]]'' and ''[[Poa pratensis]]'' || Present in ''[[Poa nemoralis]]'', [[Poa secunda|''Poa secunda'' subsp. ''juncifolia'']]; absent in ''[[Poa pratensis]]'' || Europe: ''[[Poa nemoralis]]'', ''[[Poa pratensis]]''. North America: [[Poa secunda|''Poa secunda'' subsp. ''juncifolia'']], ''[[Poa]]'' ''sylvestris'' || {{cite journal|last1=Tadych|first1=Mariusz|last2=Ambrose|first2=Karen V.|last3=Bergen|first3=Marshall S.|last4=Belanger|first4=Faith C.|last5=White|first5=James F.|title=Taxonomic placement of ''Epichloë poae'' sp. nov. and horizontal dissemination to seedlings via conidia|journal=Fungal Diversity|volume=54|issue=1|year=2012|pages=117–131|issn=1560-2745|doi=10.1007/s13225-012-0170-0}}
|-
! [[Epichloë typhina|'''''Epichloë typhina''''' '''subsp.''' '''''poae''''' '''var.''' '''''aonikenkana''''']]
Iannone & Schardl
| Argentina (Santa Cruz) || Not observed || Present || ''[[Bromus]]'' ''setifolius'' || {{cite journal|last1=Mc Cargo|first1=Patricia D.|last2=Iannone|first2=Leopoldo J.|last3=Vignale|first3=María Victoria|last4=Schardl|first4=Christopher L.|last5=Rossi|first5=María Susana|title=Species diversity of ''Epichloë'' symbiotic with two grasses from southern Argentinean Patagonia|journal=Mycologia|volume=106|issue=2|year=2017|pages=339–352|issn=0027-5514|doi=10.3852/106.2.339}}
|-
! [[Epichloë typhina|'''''Epichloë typhina''''' '''subsp.''' '''''poae''''' '''var.''' '''''canariensis''''']]
(C.D. Moon, B. Scott, & M.J. Chr.) Leuchtm.
| Canary Islands || Not observed || Present || ''[[Lolium]]'' ''edwardii'' || {{cite journal|last1=Moon|first1=Christina D.|last2=Scott|first2=Barry|last3=Schardl|first3=Christopher L.|last4=Christensen|first4=Michael J.|title=The evolutionary origins of ''Epichloë'' endophytes from annual ryegrasses|journal=Mycologia|volume=92|issue=6|year=2000|pages=1103–1118|issn=0027-5514|doi=10.1080/00275514.2000.12061258}}
|-
! [[Epichloë typhina|'''''Epichloë typhina''''' '''subsp.''' '''''poae''''' '''var.''' '''''huerfana''''']]
(J.F. White, G.T. Cole & Morgan-Jones) Tadych & Leuchtm.
| North America || Not observed || Present || ''[[Festuca arizonica]]'' || {{cite journal|last1=Glenn|first1=Anthony E.|last2=Bacon|first2=Charles W.|last3=Price|first3=Robert|last4=Hanlin|first4=Richard T.|title=Molecular phylogeny of ''Acremonium'' and its taxonomic implications|journal=Mycologia|volume=88|issue=3|year=1996|pages=369–383|issn=0027-5514|doi=10.1080/00275514.1996.12026664}}
|}

{| class="wikitable collapsible sortable"
! scope="col" style="width: 200px;" | Hybrid Taxa !! Progenitor Species !! Known Distribution !! Sexual Reproduction !! Vertical Transmission !! Known Host Range !! scope="col" style="width: 350px;" | Reference to Species Description
|-
! '''''[[Epichloë alsodes]]'''''
T. Shymanovich, C.A. Young, N.D. Charlton & S.H. Faeth
| ''[[Epichloë amarillans]]'' × [[Epichloë typhina|''Epichloë typhina'' subsp. ''poae'']] || North America || Not observed || Present || ''[[Poa]]'' ''alsodes'' || {{cite journal|last1=Shymanovich|first1=Tatsiana|last2=Charlton|first2=Nikki D.|last3=Musso|first3=Ashleigh M.|last4=Scheerer|first4=Jonathan|last5=Cech|first5=Nadja B.|last6=Faeth|first6=Stanley H.|last7=Young|first7=Carolyn A.|title=Interspecific and intraspecific hybrid ''Epichloë'' species symbiotic with the North American native grass ''Poa alsodes''|journal=Mycologia|volume=109|issue=3|year=2017|pages=459–474|issn=0027-5514|doi=10.1080/00275514.2017.1340779}}
|-
! '''''[[Epichloë australiensis]]'''''
(C.D. Moon & Schardl) Leuchtm. & Schardl
| ''[[Epichloë festucae]]'' × ''[[Epichloë typhina]]'' complex (from ''[[Poa pratensis]]'') || Australia || Not observed || Present || ''[[Echinopogon]]'' ''ovatus'' || {{cite journal|last1=Moon|first1=Christina D.|last2=Miles|first2=Christopher O.|last3=Järlfors|first3=Ulla|last4=Schardl|first4=Christopher L.|title=The evolutionary origins of three new ''Neotyphodium'' endophyte species from grasses indigenous to the Southern Hemisphere|journal=Mycologia|volume=94|issue=4|year=2017|pages=694–711|issn=0027-5514|doi=10.1080/15572536.2003.11833197}}
|-
! '''''[[Epichloë cabralii]]'''''
Iannone, M.S. Rossi & Schardl
| ''[[Epichloë amarillans]]'' × ''[[Epichloë typhina]]'' complex (from ''[[Poa nemoralis]]'') || Argentina (Santa Cruz, Tierra del Fuego) || Not observed || Present || ''[[Phleum alpinum]]'' || {{cite journal|last1=Mc Cargo|first1=Patricia D.|last2=Iannone|first2=Leopoldo J.|last3=Vignale|first3=María Victoria|last4=Schardl|first4=Christopher L.|last5=Rossi|first5=María Susana|title=Species diversity of ''Epichloë'' symbiotic with two grasses from southern Argentinean Patagonia|journal=Mycologia|volume=106|issue=2|year=2017|pages=339–352|issn=0027-5514|doi=10.3852/106.2.339}}
|-
! '''''[[Epichloë canadensis]]'''''
N.D. Charlton & C.A. Young
| ''[[Epichloë amarillans]]'' × ''[[Epichloë elymi]]'' || North America || Not observed || Present || ''[[Elymus canadensis]]'' || {{cite journal|last1=Charlton|first1=N. D.|last2=Craven|first2=K. D.|last3=Mittal|first3=S.|last4=Hopkins|first4=A. A.|last5=Young|first5=C. A.|title=''Epichloë canadensis'', a new interspecific epichloid hybrid symbiotic with Canada wildrye (''Elymus canadensis'')|journal=Mycologia|volume=104|issue=5|year=2012|pages=1187–1199|issn=0027-5514|doi=10.3852/11-403}}
|-
! '''''[[Epichloë chisosa]]'''''
(J.F. White & Morgan-Jones) Schardl
| ''[[Epichloë amarillans]]'' × ''[[Epichloë bromicola]]'' × ''[[Epichloë typhina]]'' complex (from ''[[Poa pratensis]]'') || North America || Not observed || Present || ''[[Achnatherum]]'' ''eminens'' || {{cite journal|last1=Glenn|first1=Anthony E.|last2=Bacon|first2=Charles W.|last3=Price|first3=Robert|last4=Hanlin|first4=Richard T.|title=Molecular phylogeny of ''Acremonium'' and its taxonomic implications|journal=Mycologia|volume=88|issue=3|year=2018|pages=369–383|issn=0027-5514|doi=10.1080/00275514.1996.12026664}}
|-
! '''''[[Epichloë coenophiala]]'''''
(Morgan-Jones & W. Gams) C.W. Bacon & Schardl
| ''[[Epichloë baconii]]'' (''[[Lolium]]'' associated clade) × ''[[Epichloë festucae]]'' × [[Epichloë typhina|''Epichloë typhina'' complex]] (from ''[[Poa nemoralis]]'') || Europe, North Africa, introduced in North America and elsewhere || Not observed || Present || [[Festuca arundinacea|''Schedonorus arundinaceus'']] [synonyms: ''Festuca arundinacea'', ''Lolium arundinaceum''] || {{cite journal|last1=Morgan-Jones|first1=G.|last2=Gams|first2=W.|title=Notes on hyphomycetes. XLI. An endophyte of ''Festuca arundinacea'' and the anamorph of ''Epichloë typhina'', new taxa in one of two new sections of ''Acremonium''.|journal=Mycotaxon|volume=15|year=1982|pages=311-318|issn=0093-4666}}
|-
! '''''[[Epichloë danica]]'''''
Leuchtm. & M. Oberhofer
| ''[[Epichloë bromicola]]'' × ''[[Epichloë sylvatica]]'' || Europe || Not observed || Present || ''[[Hordelymus]]'' ''europaeus'' || {{cite journal|last1=Leuchtmann|first1=Adrian|last2=Oberhofer|first2=Martina|title=The ''Epichloë'' endophytes associated with the woodland grass ''Hordelymus europaeus'' including four new taxa|journal=Mycologia|volume=105|issue=5|year=2017|pages=1315–1324|issn=0027-5514|doi=10.3852/12-400}}
|-
! '''''[[Epichloë disjuncta]]'''''
Leuchtm. & M. Oberhofer
| [[Epichloë typhina|'Epichloë typhina'']] complex × second unknown ancestor || Europe || Not observed || Present || ''[[Hordelymus]]'' ''europaeus'' || {{cite journal|last1=Leuchtmann|first1=Adrian|last2=Oberhofer|first2=Martina|title=The ''Epichloë'' endophytes associated with the woodland grass ''Hordelymus europaeus'' including four new taxa|journal=Mycologia|volume=105|issue=5|year=2017|pages=1315–1324|issn=0027-5514|doi=10.3852/12-400}}
|-
! '''''[[Epichloë funkii]]'''''
(K.D. Craven & Schardl) J.F. White
| ''[[Epichloë elymi]]'' × ''[[Epichloë festucae]]'' || North America || Not observed || Present || ''[[Achnatherum robustum]]'' || {{cite journal|last1=Moon|first1=Christina D.|last2=Guillaumin|first2=Jean-Jacques|last3=Ravel|first3=Catherine|last4=Li|first4=Chunjie|last5=Craven|first5=Kelly D.|last6=Schardl|first6=Christopher L.|title=New ''Neotyphodium'' endophyte species from the grass tribes Stipeae and Meliceae|journal=Mycologia|volume=99|issue=6|year=2017|pages=895–905|issn=0027-5514|doi=10.1080/15572536.2007.11832521}}
|-
! '''''[[Epichloë guerinii]]'''''
(Guillaumin, Ravel & C.D. Moon) Leuchtm. & Schardl
| ''[[Epichloë gansuensis]]'' × ''[[Epichloë typhina]]'' complex || Europe || Not observed || Present || ''[[Melica ciliata]]'', ''[[Melica transsilvanica]]'' || {{cite journal|last1=Moon|first1=Christina D.|last2=Guillaumin|first2=Jean-Jacques|last3=Ravel|first3=Catherine|last4=Li|first4=Chunjie|last5=Craven|first5=Kelly D.|last6=Schardl|first6=Christopher L.|title=New ''Neotyphodium'' endophyte species from the grass tribes Stipeae and Meliceae|journal=Mycologia|volume=99|issue=6|year=2017|pages=895–905|issn=0027-5514|doi=10.1080/15572536.2007.11832521}}
|-
! '''''[[Epichloë hordelymi]]'''''
Leuchtm. & M. Oberhofer
| ''[[Epichloë bromicola]]'' × ''[[Epichloë typhina]]'' complex || Europe || Not observed || Present || ''[[Hordelymus]]'' ''europaeus'' || {{cite journal|last1=Leuchtmann|first1=Adrian|last2=Oberhofer|first2=Martina|title=The ''Epichloë'' endophytes associated with the woodland grass ''Hordelymus europaeus'' including four new taxa|journal=Mycologia|volume=105|issue=5|year=2017|pages=1315–1324|issn=0027-5514|doi=10.3852/12-400}}
|-
! '''''[[Epichloë hybrida]]'''''
M.P. Cox & M.A. Campell
| [[Epichloë festucae|''Epichloë festucae'' var. ''lolii'']] × ''[[Epichloë typhina]]'' || Europe || Not observed || Present || ''[[Lolium perenne]]'' || {{cite journal|last1=Campbell|first1=Matthew A.|last2=Tapper|first2=Brian A.|last3=Simpson|first3=Wayne R.|last4=Johnson|first4=Richard D.|last5=Mace|first5=Wade|last6=Ram|first6=Arvina|last7=Lukito|first7=Yonathan|last8=Dupont|first8=Pierre-Yves|last9=Johnson|first9=Linda J.|last10=Scott|first10=D. Barry|last11=Ganley|first11=Austen R. D.|last12=Cox|first12=Murray P.|title=''Epichloë hybrida'', sp. nov., an emerging model system for investigating fungal allopolyploidy|journal=Mycologia|volume=109|year=2017|pages=1–15|issn=0027-5514|doi=10.1080/00275514.2017.1406174|doi-access=free}}
|-
! '''''[[Epichloë liyangensis]]'''''
Z.W. Wang, Y. Kang & H. Miao
| ''[[Epichloë bromicola]]'' × ''[[Epichloë typhina]]'' complex (from ''[[Poa nemoralis]]'') || Asia || Observed || Present || [[Poa pratensis|''Poa pratensis'' subsp. ''pratensis'']] || {{cite journal|last1=Yan|first1=Kang|last2=Yanling|first2=Ji|last3=Kunran|first3=Zhu|last4=Hui|first4=Wang|last5=Huimin|first5=Miao|last6=Zhiwei|first6=Wang|title=A new ''Epichloë'' species with interspecific hybrid origins from ''Poa pratensis'' ssp. ''pratensis'' in Liyang, China|journal=Mycologia|volume=103|issue=6|year=2017|pages=1341–1350|issn=0027-5514|doi=10.3852/10-352}}
|-
! '''''[[Epichloë melicicola]]'''''
(C.D. Moon & Schardl) Schardl
| ''[[Epichloë aotearoae]]'' × ''[[Epichloë festucae]]'' || South Africa || Not observed || Present || ''[[Melica racemosa]]'', ''Melica decumbens'' || {{cite journal|last1=Moon|first1=Christina D.|last2=Miles|first2=Christopher O.|last3=Järlfors|first3=Ulla|last4=Schardl|first4=Christopher L.|title=The evolutionary origins of three new ''Neotyphodium'' endophyte species from grasses indigenous to the Southern Hemisphere|journal=Mycologia|volume=94|issue=4|year=2017|pages=694–711|issn=0027-5514|doi=10.1080/15572536.2003.11833197}}
|-
! '''''[[Epichloë novae-zelandiae]]'''''
Leuchtm. & A.V. Stewart
| ''[[Epichloë amarillans]]'' × ''[[Epichloë bromicola]]'' × [[Epichloë typhina|''Epichloë typhina'' subsp. ''poae'']] || New Zealand || Not observed || Present || ''[[Poa]]'' ''matthewsii'' || {{cite journal|last1=Leuchtmann|first1=Adrian|last2=Young|first2=Carolyn A.|last3=Stewart|first3=Alan V.|last4=Simpson|first4=Wayne R.|last5=Hume|first5=David E.|last6=Scott|first6=Barry|title=''Epichloë novae-zelandiae'', a new endophyte from the endemic New Zealand grass ''Poa matthewsii''|journal=New Zealand Journal of Botany|volume=57|issue=4|year=2019|pages=271–288|issn=0028-825X|doi=10.1080/0028825X.2019.1651344}}
|-
! '''''[[Epichloë occultans]]'''''
(C.D. Moon, B. Scott & M.J. Chr.) Schardl
| ''[[Epichloë baconii]]'' (''[[Lolium]]'' associated clade) × ''[[Epichloë bromicola]]'' || Europe, North Africa, introduced in New Zealand and elsewhere || Not observed || Present || [[Festuca perennis|''Lolium multiflorum'']], ''[[Lolium rigidum]]'' u.a. || {{cite journal|last1=Moon|first1=Christina D.|last2=Scott|first2=Barry|last3=Schardl|first3=Christopher L.|last4=Christensen|first4=Michael J.|title=The evolutionary origins of ''Epichloë'' endophytes from annual ryegrasses|journal=Mycologia|volume=92|issue=6|year=2019|pages=1103–1118|issn=0027-5514|doi=10.1080/00275514.2000.12061258}}
|-
! '''''[[Epichloë pampeana]]'''''
(Iannone & Cabral) Iannone & Schardl
| ''[[Epichloë festucae]]'' × ''[[Epichloë typhina]]'' complex (from ''[[Poa nemoralis]]'') || South America || Not observed || Present || ''[[Bromus]]'' ''auleticus'' || {{cite journal|last1=Iannone|first1=Leopoldo Javier|last2=Cabral|first2=Daniel|last3=Schardl|first3=Christopher Lewis|last4=Rossi|first4=María Susana|title=Phylogenetic divergence, morphological and physiological differences distinguish a new ''Neotyphodium'' endophyte species in the grass ''Bromus auleticus'' from South America|journal=Mycologia|volume=101|issue=3|year=2017|pages=340–351|issn=0027-5514|doi=10.3852/08-156}}
|-
! '''''[[Epichloë schardlii]]'''''
(Ghimire, Rudgers & K.D. Craven) Leuchtm.
| [[Epichloë typhina|''Epichloë typhina'' complex]] (subsp. ''poae'' × subsp. ''poae'') || North America || Not observed || Present || [[Cinna (plant)|''Cinna arundinacea'']] || {{cite journal|last1=Ghimire|first1=Sita R.|last2=Rudgers|first2=Jennifer A.|last3=Charlton|first3=Nikki D.|last4=Young|first4=Carolyn|last5=Craven|first5=Kelly D.|title=Prevalence of an intraspecific ''Neotyphodium'' hybrid in natural populations of stout wood reed (''Cinna arundinacea'' L.) from eastern North America|journal=Mycologia|volume=103|issue=1|year=2017|pages=75–84|issn=0027-5514|doi=10.3852/10-154}}
|-
! [[Epichloë schardlii|'''''Epichloë schardlii''''' '''var.''' '''''pennsylvanica''''']]
T. Shymanovich, C.A. Young, N.D. Charlton & S.H. Faeth
| ''[[Epichloë typhina]]'' complex (subsp. ''poae'' × subsp. ''poae'') || North America || Not observed || Present || ''[[Poa]]'' ''alsodes'' || {{cite journal|last1=Shymanovich|first1=Tatsiana|last2=Charlton|first2=Nikki D.|last3=Musso|first3=Ashleigh M.|last4=Scheerer|first4=Jonathan|last5=Cech|first5=Nadja B.|last6=Faeth|first6=Stanley H.|last7=Young|first7=Carolyn A.|title=Interspecific and intraspecific hybrid ''Epichloë'' species symbiotic with the North American native grass ''Poa alsodes''|journal=Mycologia|volume=109|issue=3|year=2017|pages=459–474|issn=0027-5514|doi=10.1080/00275514.2017.1340779}}
|-
! '''''[[Epichloë siegelii]]'''''
(K.D. Craven, Leuchtm. & Schardl) Leuchtm. & Schardl
| ''[[Epichloë bromicola]]'' × ''[[Epichloë festucae]]'' || Europe || Not observed || Present || [[Festuca pratensis|''Schedonorus pratensis'']] (synonyms: ''Festuca pratensis'', ''Lolium pratense'') || {{cite journal|last1=Craven|first1=K.D.|last2=Blankenship|first2=J.D.|last3=Leuchtmann|first3=A.|last4=Hinight|first4=K.|last5=Schardl|first5=C.L.|title=Hybrid fungal endophytes symbiotic with the grass ''Lolium pratense''|journal=Sydowia|volume=53|year=2001|pages=44–73}}
|-
! '''''[[Epichloë sinica]]'''''
(Z.W. Wang, Y.L. Ji & Y. Kang) Leuchtm.
| ''[[Epichloë bromicola]]'' × ''[[Epichloë typhina]]'' complex || Asia || Not observed || Present || [[Elymus (plant)|''Roegneria'']] spp. || {{cite journal|last1=Yan|first1=Kang|last2=Yanling|first2=Ji|last3=Xianghui|first3=Sun|last4=Lihui|first4=Zhan|last5=Wei|first5=Li|last6=Hanshou|first6=Yu|last7=Zhiwei|first7=Wang|title=Taxonomy of ''Neotyphodium'' endophytes of Chinese native ''Roegneria'' plants|journal=Mycologia|volume=101|issue=2|year=2017|pages=211–219|issn=0027-5514|doi=10.3852/08-018}}
|-
! '''''[[Epichloë sinofestucae]]'''''
(Y.G. Chen, Y.L. Ji & Z.W. Wang) Leuchtm.
| ''[[Epichloë bromicola]]'' × [[Epichloë typhina|''Epichloë typhina'' complex]] || Asia || Not observed || Present || ''[[Festuca parvigluma]]'' || {{cite journal|last1=Chen|first1=Yong-gan|last2=Ji|first2=Yan-ling|last3=Yu|first3=Han-shou|last4=Wang|first4=Zhi-wei|title=A new ''Neotyphodium'' species from ''Festuca parvigluma'' Steud. grown in China|journal=Mycologia|volume=101|issue=5|year=2017|pages=681–685|issn=0027-5514|doi=10.3852/08-181}}
|-
! '''''[[Epichloë tembladerae]]'''''
(Cabral & J.F. White) Iannone & Schardl 
| ''[[Epichloë festucae]]'' × ''[[Epichloë typhina]]'' complex (from ''[[Poa nemoralis]]'') || North America || Not observed || Present || North America: ''[[Festuca arizonica]]''. South America: ''[[Bromus]]'' ''auleticus'', ''[[Bromus]]'' ''setifolius'', ''[[Festuca]]'' ''argentina'', ''[[Festuca]]'' ''hieronymi'', ''[[Festuca]]'' ''magellanica'', ''[[Festuca]]'' ''superba'', ''[[Melica stuckertii]]'', ''[[Phleum alpinum]]'', ''[[Phleum]]'' ''commutatum'', ''[[Poa]]'' ''huecu'', ''[[Poa]]'' ''rigidifolia'' || {{cite journal|last1=Cabral|first1=Daniel|last2=Cafaro|first2=Matías J.|last3=Saidman|first3=B.|last4=Lugo|first4=M.|last5=Reddy|first5=Ponaka V.|last6=White|first6=James F.|title=Evidence supporting the occurrence of a new species of endophyte in some South American grasses|journal=Mycologia|volume=91|issue=2|year=2019|pages=315–325|issn=0027-5514|doi=10.1080/00275514.1999.12061021}}
|-
! '''''[[Epichloë uncinata]]'''''
(W. Gams, Petrini & D. Schmidt) Leuchtm. & Schardl 
| ''[[Epichloë bromicola]]'' × ''[[Epichloë typhina]]'' complex || Europe || Not observed || Present || [[Festuca pratensis|''Schedonorus pratensis'']] (synonyms: ''Festuca pratensis'', ''Lolium pratense'') || {{cite journal|last1=Gams|first1=W.|last2=Petrini|first2=O. J.|last3=Schmidt|first3=D.|title=''Acremonium uncinatum'', a new endophyte in ''Festuca pratensis''|journal=Mycotaxon|volume=37|year=1990|pages=67–71}}A1:G25
|}

==Life cycle and growth==

[[File:Neotyphodium coenophialum.jpg|thumb|alt=Blue-stained large plant cells with smaller hyphae visible between them|''[[Epichloë coenophiala]]'' hyphae between [[tall fescue]] leaf cells]]

''Epichloë'' species are specialized to form and maintain systemic, constitutive (long-term) [[symbiosis|symbioses]] with plants, often with limited or no disease incurred on the host.<ref name="Spatafora2007"/> The best-studied of these symbionts are associated with the [[grass]]es and [[Cyperaceae|sedges]], in which they infect the leaves and other aerial tissues by growing between the plant cells ([[endophyte|endophytic]] growth) or on the surface above or beneath the cuticle ([[epiphyte|epiphytic]] growth). An individual infected plant will generally bear only a single genetic individual clavicipitaceous symbiont, so the plant-fungus system constitutes a genetic unit called a symbiotum (pl. symbiota).

Symptoms and signs of the fungal infection, if manifested at all, only occur on a specific tissue or site of the host [[Tiller (botany)|tiller]], where the fungal stroma or sclerotium emerges. The stroma (pl. stromata) is a mycelial cushion that gives rise first to asexual spores ([[conidium|conidia]]), then to the sexual fruiting bodies ([[ascocarp]]s; perithecia). [[Sclerotium|Sclerotia]] are hard resting structures that later (after incubation on the ground) germinate to form stipate stromata. Depending on the fungus species, the host tissues on which stromata or sclerotia are produced may be young inflorescences and surrounding leaves, individual florets, nodes, or small segments of the leaves. Young stromata are hyaline (colorless), and as they mature they turn dark gray, black, or yellow-orange. Mature stromata eject meiotically derived spores ([[ascospore]]s), which are ejected into the atmosphere and initiate new plant infections ([[horizontal transmission]]). In some cases no stroma or sclerotium is produced, but the fungus infects seeds produced by the infected plant, and is thereby [[vertical transmission|transmitted vertically]] to the next host generation. Most ''Epichloë'' species, and all asexual species, can vertically transmit.

The taxonomic dichotomy is especially interesting in this group of symbionts, because vegetative propagation of fungal [[mycelium]] occurs by vertical transmission, i.e., fungal growth into newly developing host tillers (=individual grass plants). Importantly, many ''Epichloë'' species infect new grass plants solely by growing into the seeds of their grass hosts, and infecting the growing seedling.<ref name="Schardl2004"/><ref name="Freeman1904"/> Manifestation of the sexual state — which only occurs in ''Epichloë'' species — causes "choke disease", a condition in which grass [[inflorescence]]s are engulfed by rapid fungal outgrowth forming a stroma. The fungal stroma suppresses host seed production and culminates in the ejection of meiospores ([[ascospore]]s) that mediate horizontal (contagious) transmission of the fungus to new plants.<ref name="Schardl2004"/> So, the two transmission modes exclude each other, although in many grass-''Epichloë'' symbiota the fungus actually displays both transmission modes simultaneously, by choking some tillers and transmitting in seeds produced by unchoked tillers.

While being [[obligate symbiont]]s in nature, most epichloae are readily culturable in the [[laboratory]] on [[culture media]] such as [[potato dextrose agar]] or a minimal salts broth supplemented with thiamine, sugars or sugar alcohols, and organic nitrogen or ammonium.<ref name="Blankenship2001"/>

''Epichloë'' species are commonly spread by [[fly|flies]] of the genus ''[[Botanophila]]''. The flies lay their eggs in the growing fungal tissues and the larvae feed on them.<ref name="Goryzynska2010"/>

[[File:Stromata Botanophila Flies.jpg|thumb|A stroma of ''[[Epichloë baconii]]'' formed on the grass ''[[Agrostis stolonifera]]'', showing eggs, brood chambers and larval feeding tracks of ''[[Botanophila]]'' flies.]]

==Evolution==

The epichloae display a number of central features that suggest a very strong and ancient association with their grass hosts. The symbiosis appears to have existed already during the early grass [[evolution]] that has spawned today's [[Pooideae|pooid]] grasses. This is suggested by [[Phylogeny|phylogenetic]] studies indicating a preponderance of codivergence of ''Epichloë'' species with the grass hosts they inhabit.<ref name="Schardl2008"/> Growth of the fungal symbiont is very tightly regulated within its grass host, indicated by a largely unbranched [[mycelium|mycelial]] morphology and remarkable synchrony of grass leaf and [[hypha]]l extension of the fungus;<ref name="Tan2001"/><ref name="Christensen2002"/> the latter seems to occur via a mechanism that involves stretch-induced or ''intercalary'' elongation of the endophyte's hyphae, a process so far not found in any other fungal species, indicating specialized adaptation of the fungus to the dynamic growth environment inside its host.<ref name="Christensen2008"/> A complex [[NADPH oxidase]] enzyme-based [[Reactive oxygen species|ROS]]-generating system in ''Epichloë'' species is indispensable for maintenance of this growth synchrony. Thus, it has been demonstrated that [[Gene deletion|deletion]] of genes encoding these enzymes in ''Epichloë festucae'' causes severely disordered fungal growth in grass tissues and even death of the grass plant.<ref name="Tanaka2006"/><ref name="Takemoto2006"/>

Molecular phylogenetic evidence demonstrates that asexual ''Epichloë'' species are derived either from sexual ''Epichloë'' species, or more commonly, are hybrids of two or more progenitor ''Epichloë'' species.<ref name="Tsai1994"/><ref name="Moon2004"/>

==Bioactive compounds==

[[File:N-formylloline.svg|thumb|125px|''N''-formylloline, an [[insecticide|insecticidal]] alkaloid produced in several ''Epichloë''–grass symbiota.]]

Many ''Epichloë'' endophytes produce a diverse range of [[natural product]] compounds with biological activities against a broad range of herbivores.<ref name="Bush1997"/><ref name="Scott2001"/> The purpose of these compounds is as a [[toxicity]] or feeding deterrence against insect and mammalian [[Herbivory|herbivores]].<ref name="Roberts2005"/> [[Ergoline]] alkaloids (which are ergot alkaloids, named after the [[ergot]] fungus, ''Claviceps purpurea'', a close relative of the epichloae) are characterized by a ring system derived from 4-[[prenyl]] [[tryptophan]].<ref name="Schardl2006"/> Among the most abundant ergot alkaloids in epichloë-symbiotic grasses is ergovaline, comprising an ergoline [[Moiety (chemistry)|moiety]] attached to a bicyclic [[tripeptide]] containing the amino acids <small>L</small>-[[proline]], <small>L</small>-[[alanine]], and <small>L</small>-[[valine]]. Key genes and enzymes for ergot alkaloid biosynthesis have been identified in epichloae and include ''dmaW'', encoding [[Tryptophan dimethylallyltransferase|dimethylallyl-tryptophan synthase]] and ''lpsA'', a [[Nonribosomal peptide|non-ribosomal peptide synthetase]].<ref name="Schardl2006"/>

Another group of epichloë alkaloids are the [[indole]]-[[diterpenoid]]s, such as lolitrem B, which are produced from the activity of several enzymes, including [[prenyltransferase]]s and various [[monooxygenase]]s.<ref name="Young2006"/> Both the ergoline and indole-diterpenoid alkaloids have [[biological activity]] against mammalian herbivores, and also activity against some insects.<ref name="Bush1997"/> Peramine is a pyrrolopyrazine alkaloid thought to be biosynthesized from the [[guanidine|guanidinium]]-group-containing amino acid <small>L</small>-[[arginine]], and pyrrolidine-5-[[carboxylate]], a precursor of <small>L</small>-[[proline]],<ref name="Tanaka2005"/> and is an insect-feeding deterrent. The [[loline alkaloids]]<ref name="Schardl2007"/> are 1-aminopyrrolizidines with an oxygen atom linking bridgehead carbons 2 and 7, and are biosynthesized from the amino acids <small>L</small>-proline and <small>L</small>-[[homoserine]].<ref name="Blankenship2005"/> The lolines have [[insecticide|insecticidal]] and insect-deterrent activities comparable to [[nicotine]].<ref name="Schardl2007"/> Loline accumulation is strongly induced in young growing tissues<ref name="Zhang2009"/> or by damage to the plant-fungus symbiotum.<ref name="Gonthier2008"/> Many, but not all, epichloae produce up to three classes of these alkaloids in various combinations and amounts.<ref name="Bush1997"/> Recently it has been shown that Epichloë uncinata infection and loline content afford Festulolium grasses protection from black beetle ([[Heteronychus arator]]).<ref name="Barker2014"/>

Many species in ''Epichloë'' produce biologically active alkaloids, such as [[ergot alkaloid]]s, indole-diterpenoids (e.g., [[lolitrem B]]), [[loline alkaloids]], and the unusual [[guanidinium]] alkaloid, peramine.<ref name="Bush1997"/>

==Ecology==

===Effects on the grass plant===

It has been proposed that vertically transmitted [[symbiont]]s should evolve to be [[Mutualism (biology)|mutualists]] since their reproductive [[fitness (biology)|fitness]] is intimately tied to that of their hosts.<ref name="Ewald1987"/> In fact, some positive effects of epichloae on their host plants include increased growth, drought tolerance, and [[herbivore]] and [[pathogen]] resistance.<ref name="Schardl2004"/><ref name="MalinowskiBelesky2000"/> Resistance against herbivores has been attributed to [[alkaloid]]s produced by the symbiotic epichloae.<ref name="Bush1997"/> Although grass-epichloë symbioses have been widely recognized to be [[Mutualism (biology)|mutualistic]] in many wild and cultivated grasses, the interactions can be highly variable and sometimes antagonistic, especially under nutrient-poor conditions in the soil.<ref name="Saikkonen2002"/>

===Ecosystem dynamics===

Due to the relatively large number of grass species harboring epichloae and the variety of environments in which they occur, the mechanisms underlying beneficial or antagonistic outcomes of epichloë-grass symbioses are difficult to delineate in natural and also agricultural environments.<ref name="Schardl2004"/><ref name="Saikkonen2006"/> Some studies suggest a relationship between grazing by herbivores and increased epichloë infestation of the grasses on which they feed,<ref name="Clay2005"/><ref name="Kohn2007"/> whereas others indicate a complex interplay between plant species and fungal symbionts in response to herbivory or environmental conditions.<ref name="Granath2007"/> The strong anti-herbivore activities of several bioactive compounds produced by the epichloae <ref name="Bush1997"/><ref name="Tanaka2005"/> and relatively modest direct effects of the epichloae on plant growth and physiology<ref name="Hahn2008"/><ref name="Hunt2005"/> suggest that these compounds play a major role in the persistence of the symbiosis.

==References==
{{Reflist|30em|refs=

<ref name="Barker2014">{{cite journal|author1=Barker GM |author2=Patchett BJ |author3=Cameron NE |year=2014|title=Epichloë uncinata infection and loline content afford Festulolium grasses protection from black beetle (Heteronychus arator). Landcare Research, Hamilton, New Zealand & Cropmark Seeds Ltd, Christchurch, New Zealand 20 Dec 2014.|journal=New Zealand Journal of Agricultural Research|doi=10.1080/00288233.2014.978480|volume=58|pages=35–56}}</ref>

<ref name="Blankenship2001">{{cite journal|vauthors=Blankenship JD, Spiering MJ, Wilkinson HH, Fannin FF, Bush LP, Schardl CL |year=2001|title=Production of loline alkaloids by the grass endophyte, ''Neotyphodium uncinatum'', in defined media|journal =Phytochemistry|volume=58|pages=395–401|pmid=11557071|doi=10.1016/S0031-9422(01)00272-2|issue=3}}</ref>

<ref name="Blankenship2005">{{cite journal|vauthors=Blankenship JD, Houseknecht JB, Pal S, Bush LP, Grossman RB, Schardl CL |year=2005|title=Biosynthetic precursors of fungal pyrrolizidines, the loline alkaloids|journal=ChemBioChem|volume=6|pages=1016–1022|pmid=15861432|doi=10.1002/cbic.200400327|issue=6}}</ref>

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}}

{{Taxonbar|from=Q5382570}}

{{DEFAULTSORT:Epichloe}}
[[Category:Epichloë| ]]
[[Category:Ascomycota genera]]
[[Category:Clavicipitaceae]]