Mycorrhizal fungal community composition in seven orchid species inhabiting Song Mountain, Beijing, China
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摘要
Mycorrhizal fungi play an important role in the germination and growth of orchids essentially influencing their survival,abundance, and spatial distribution. In this study, we investigated the composition of the mycorrhizal fungal community in seven terrestrial orchid species inhabiting Song Mountain, Beijing, China, using Illumina MiSeq high-throughput sequencing. The mycorrhizal communities in the seven orchids were mainly composed of members of the Ceratobasidiaceae, Sebacinales, and Tulasnellaceae, while a number of ectomycorrhizal fungi belonging to the Russulaceae, Tricholomataceae, Thelephoraceae, and Cortinariaceae were occasionally observed. However, the dominant fungal associates and mycorrhizal community differed significantly among the orchid species as well as subhabitats. These findings confirm the previous observation that sympatric orchid species show different preferences for mycorrhizal fungi, which may drive niche partitioning and contribute to their cooccurrence.
Mycorrhizal fungi play an important role in the germination and growth of orchids essentially influencing their survival,abundance, and spatial distribution. In this study, we investigated the composition of the mycorrhizal fungal community in seven terrestrial orchid species inhabiting Song Mountain, Beijing, China, using Illumina MiSeq high-throughput sequencing. The mycorrhizal communities in the seven orchids were mainly composed of members of the Ceratobasidiaceae, Sebacinales, and Tulasnellaceae, while a number of ectomycorrhizal fungi belonging to the Russulaceae, Tricholomataceae, Thelephoraceae, and Cortinariaceae were occasionally observed. However, the dominant fungal associates and mycorrhizal community differed significantly among the orchid species as well as subhabitats. These findings confirm the previous observation that sympatric orchid species show different preferences for mycorrhizal fungi, which may drive niche partitioning and contribute to their cooccurrence.
Mycorrhizal fungi play an important role in the germination and growth of orchids essentially influencing their survival,abundance, and spatial distribution. In this study, we investigated the composition of the mycorrhizal fungal community in seven terrestrial orchid species inhabiting Song Mountain, Beijing, China, using Illumina MiSeq high-throughput sequencing. The mycorrhizal communities in the seven orchids were mainly composed of members of the Ceratobasidiaceae, Sebacinales, and Tulasnellaceae, while a number of ectomycorrhizal fungi belonging to the Russulaceae, Tricholomataceae, Thelephoraceae, and Cortinariaceae were occasionally observed. However, the dominant fungal associates and mycorrhizal community differed significantly among the orchid species as well as subhabitats. These findings confirm the previous observation that sympatric orchid species show different preferences for mycorrhizal fungi, which may drive niche partitioning and contribute to their cooccurrence.
引文
Altschul,S.F.,Gish,W.,Miller,W.,Myers,E.W.,and Lipman,D.J.(1990).Basic local alignment search tool.J Mol Biol 215,403-410.
Bengtsson-Palme,J.,Ryberg,M.,Hartmann,M.,Branco,S.,Wang,Z.,Godhe,A.,et al.(2013).Improved software detection and extraction of ITS1 and ITS2 from ribosomal ITS sequences of fungi and other eukaryotes for analysis of environmental sequencing data.Methods Ecol Evol 4,914-919.
Bidartondo,M.I.,Burghardt,B.,Gebauer,G.,Bruns,T.D.,and Read,D.J.(2004).Changing partners in the dark:isotopic and molecular evidence of ectomycorrhizal liaisons between forest orchids and trees.Proc RSoc London Ser B-Biol Sci 271,1799-1806.
Cameron,D.D.,Johnson,I.,Read,D.J.,and Leake,J.R.(2008).Giving and receiving:measuring the carbon cost of mycorrhizas in the green orchid,Goodyera repens.New Phytol 180,176-184.
Cribb,P.J.,Kell,S.P.,Dixon,K.W.,and Barrett,R.L.(2003).Orchid conservation:a global perspective.In Dixon KW,Kell SP,Barrett RL,Cribb PJ,eds.(Kota Kinabalu,Sabah,Natural History Publications),pp.1-24.
Darriba,D.,Taboada,G.L.,Doallo,R.,and Posada,D.(2012).jModelTest2:more models,new heuristics and parallel computing.Nat Methods 9,772.
De Cáceres,M.,Legendre,P.,and Moretti,M.(2010).Improving indicator species analysis by combining groups of sites.Oikos 119,1674-1684.
Dearnaley,J.D.W.(2007).Further advances in orchid mycorrhizal research.Mycorrhiza 17,475-486.
Dearnaley,J.D.W.,Martos,F.,and Selosse,M.A.(2012).Orchid mycorrhizas:molecular ecology,physiology,evolution and conservation aspects.In Fungal Association,2nd ed.B,Hock.(Berlin:SpringerVerlag),pp.207-230.
Edgar,R.C.(2010).Search and clustering orders of magnitude faster than BLAST.Bioinformatics 26,2460-2461.
Edgar,R.C.(2013).UPARSE:highly accurate OTU sequences from microbial amplicon reads.Nat Methods 10,996-998.
Esposito,F.,Jacquemyn,H.,Waud,M.,and Tyteca,D.(2016).Mycorrhizal fungal diversity and community composition in two closely related Platanthera(Orchidaceae)species.PLoS ONE 11,e0164108.
van der Heijden,M.G.A.,Martin,F.M.,Selosse,M.A.,and Sanders,I.R.(2015).Mycorrhizal ecology and evolution:the past,the present,and the future.New Phytol 205,1406-1423.
Ihrmark,K.,B?deker,I.T.M.,Cruz-Martinez,K.,Friberg,H.,Kubartova,A.,Schenck,J.,Strid,Y.,Stenlid,J.,Brandstr?m-Durling,M.,Clemmensen,K.E.,et al.(2012).New primers to amplify the fungal ITS2 region-evaluation by 454-sequencing of artificial and natural communities.FEMS Microbiol Ecol 82,666-677.
Ives,A.R.,and Godfray,H.C.J.(2006).Phylogenetic analysis of trophic associations.Am Natist 168,E1-E14.
Jacquemyn,H.,Brys,R.,Lievens,B.,and Wiegand,T.(2012a).Spatial variation in below-ground seed germination and divergent mycorrhizal associations correlate with spatial segregation of three co-occurring orchid species.J Ecol 100,1328-1337.
Jacquemyn,H.,Brys,R.,Merckx,V.S.F.T.,Waud,M.,Lievens,B.,and Wiegand,T.(2014).Coexisting orchid species have distinct mycorrhizal communities and display strong spatial segregation.New Phytol 202,616-627.
Jacquemyn,H.,Brys,R.,Waud,M.,Busschaert,P.,and Lievens,B.(2015a).Mycorrhizal networks and coexistence in species-rich orchid communities.New Phytol 206,1127-1134.
Jacquemyn,H.,Deja,A.,De hert,K.,Cachapa Bailarote,B.,and Lievens,B.(2012b).Variation in mycorrhizal associations with Tulasnelloid fungi among populations of five Dactylorhiza species.PLoS ONE 7,e42212.
Jacquemyn,H.,Honnay,O.,Cammue,B.P.A.,Brys,R.,and Lievens,B.(2010).Low specificity and nested subset structure characterize mycorrhizal associations in five closely related species of the genus Orchis.Mol Ecol 19,4086-4095.
Jacquemyn,H.,Merckx,V.,Brys,R.,Tyteca,D.,Cammue,B.P.A.,Honnay,O.,and Lievens,B.(2011).Analysis of network architecture reveals phylogenetic constraints on mycorrhizal specificity in the genus Orchis(Orchidaceae).New Phytol 192,518-528.
Jacquemyn,H.,Waud,M.,Brys,R.,Lallemand,F.,Courty,P.E.,Robionek,A.,and Selosse,M.A.(2017).Mycorrhizal associations and trophic modes in coexisting orchids:an ecological continuum between autoand mixotrophy.Front Plant Sci 8,1497.
Jacquemyn,H.,Waud,M.,Lievens,B.,and Brys,R.(2016a).Differences in mycorrhizal communities between Epipactis palustris,E.helleborine and its presumed sister species E.neerlandica.Ann Bot 118,105-114.
Jacquemyn,H.,Waud,M.,Merckx,V.S.F.T.,Lievens,B.,and Brys,R.(2015b).Mycorrhizal diversity,seed germination and long-term changes in population size across nine populations of the terrestrial orchid Neottia ovata.Mol Ecol 24,3269-3280.
Jacquemyn,H.,Waud,M.,Merckx,V.S.F.T.,Brys,R.,Tyteca,D.,Hedrén,M.,and Lievens,B.(2016b).Habitat-driven variation in mycorrhizal communities in the terrestrial orchid genus Dactylorhiza.Sci Rep 6,37182.
Kembel,S.W.,Cowan,P.D.,Helmus,M.R.,Cornwell,W.K.,Morlon,H.,Ackerly,D.D.,Blomberg,S.P.,and Webb,C.O.(2010).Picante:R tools for integrating phylogenies and ecology.Bioinformatics 26,1463-1464.
Larkin,M.A.,Blackshields,G.,Brown,N.P.,Chenna,R.,McGettigan,P.A.,McWilliam,H.,Valentin,F.,Wallace,I.M.,Wilm,A.,Lopez,R.,et al.(2007).Clustal W and Clustal X version 2.0.Bioinformatics 23,2947-2948.
Li,N.Q.,Wu,J.G.,Jiang,W.J.,Mu,X.Y.,and Cheng,J.(2013).Biodiversity and conservation of orchids in Songshan National Nature Reserve,Beijing.Plant Sci J 31,510.
Martos,F.,Munoz,F.,Pailler,T.,Kottke,I.,Gonneau,C.,and Selosse,M.A.(2012).The role of epiphytism in architecture and evolutionary constraint within mycorrhizal networks of tropical orchids.Mol Ecol21,5098-5109.
McCormick,M.K.,and Jacquemyn,H.(2014).What constrains the distribution of orchid populations?New Phytol 202,392-400.
McCormick,M.K.,Lee Taylor,D.,Juhaszova,K.,Burnett Jr,R.K.,Whigham,D.F.,and O’neill,J.P.(2012).Limitations on orchid recruitment:not a simple picture.Mol Ecol 21,1511-1523.
McKendrick,S.L.,Leake,J.R.,Taylor,D.L.,and Read,D.J.(2002).Symbiotic germination and development of the myco-heterotrophic orchid Neottia nidus-avis in nature and its requirement for locally distributed Sebacina spp..New Phytol 154,233-247.
Mujica,M.I.,Saez,N.,Cisternas,M.,Manzano,M.,Armesto,J.J.,and Pérez,F.(2016).Relationship between soil nutrients and mycorrhizal associations of two Bipinnula species(Orchidaceae)from central Chile.Ann Bot 118,149-158.
Ogura-Tsujita,Y.,Yokoyama,J.,Miyoshi,K.,and Yukawa,T.(2012).Shifts in mycorrhizal fungi during the evolution of autotrophy to mycoheterotrophy in Cymbidium(Orchidaceae).Am J Bot 99,1158-1176.
Oja,J.,Vahtra,J.,Bahram,M.,Kohout,P.,Kull,T.,Rannap,R.,K?ljalg,U.,and Tedersoo,L.(2017).Local-scale spatial structure and community composition of orchid mycorrhizal fungi in semi-natural grasslands.Mycorrhiza 27,355-367.
Oksanen,J.,Blanchet,F.G.,Kindt,R.,Legendre,P.,Minchin,P.R.,O’Hara,R.B.,Simpson,G.L.,Solymos,P.,Stevens,M.H.H.,and Wagner,H.(2015).Vegan:Community Ecology Package Version 2.3-2.http://cran.r-project.org/.
Rasmussen,H.N.,and Rasmussen,F.N.(2009).Orchid mycorrhiza:implications of a mycophagous life style.Oikos 118,334-345.
Schloss,P.D.,Westcott,S.L.,Ryabin,T.,Hall,J.R.,Hartmann,M.,Hollister,E.B.,Lesniewski,R.A.,Oakley,B.B.,Parks,D.H.,Robinson,C.J.,et al.(2009).Introducing mothur:open-source,platformindependent,community-supported software for describing and comparing microbial communities.Appl Environ Microbiol 75,7537-7541.
Shefferson,R.P.,Weiss,M.,Kull,T.,and Taylor,D.L.(2005).High specificity generally characterizes mycorrhizal association in rare lady’s slipper orchids,genus Cypripedium.Mol Ecol 14,613-626.
Stamatakis,A.,Hoover,P.,and Rougemont,J.(2008).A rapid bootstrap algorithm for the RAxML web servers.Systatic Biol 57,758-771.
Stark,C.,Babik,W.,and Durka,W.(2009).Fungi from the roots of the common terrestrial orchid Gymnadenia conopsea.Mycol Res 113,952-959.
Swarts,N.D.,and Dixon,K.W.(2009).Terrestrial orchid conservation in the age of extinction.Ann Bot 104,543-556.
Swarts,N.D.,Sinclair,E.A.,Francis,A.,and Dixon,K.W.(2010).Ecological specialization in mycorrhizal symbiosis leads to rarity in an endangered orchid.Mol Ecol 19,3226-3242.
Taylor,D.L.,and McCormick,M.K.(2008).Internal transcribed spacer primers and sequences for improved characterization of basidiomycetous orchid mycorrhizas.New Phytol 177,1020-1033.
Tedersoo,L.,Bahram,M.,P?lme,S.,K?ljalg,U.,Yorou,N.S.,Wijesundera,R.,Villarreal Ruiz,L.,Vasco-Palacios,A.M.,Thu,P.Q.,Suija,A.,et al.(2014).Global diversity and geography of soil fungi.Science 346,1256688.
Tě?itelová,T.,Jersáková,J.,Roy,M.,Kubátová,B.,Tě?itel,J.,Urfus,T.,Trávní?ek,P.,and Suda,J.(2013).Ploidy-specific symbiotic interactions:divergence of mycorrhizal fungi between cytotypes of the Gymnadenia conopsea group(Orchidaceae).New Phytol 199,1022-1033.
Tě?itelová,T.,Kotilínek,M.,Jersáková,J.,Joly,F.X.,Ko?nar,J.,Tatarenko,I.,and Selosse,M.A.(2015).Two widespread green Neottia species(Orchidaceae)show mycorrhizal preference for Sebacinales in various habitats and ontogenetic stages.Mol Ecol 24,1122-1134.
Waud,M.,Busschaert,P.,Lievens,B.,and Jacquemyn,H.(2016a).Specificity and localised distribution of mycorrhizal fungi in the soil may contribute to co-existence of orchid species.Fungal Ecol 20,155-165.
Waud,M.,Busschaert,P.,Ruyters,S.,Jacquemyn,H.,and Lievens,B.(2014).Impact of primer choice on characterization of orchid mycorrhizal communities using 454 pyrosequencing.Mol Ecol Resour 14,679-699.
Waud,M.,Wiegand,T.,Brys,R.,Lievens,B.,and Jacquemyn,H.(2016b).Nonrandom seedling establishment corresponds with distancedependent decline in mycorrhizal abundance in two terrestrial orchids.New Phytol 211,255-264.
Weiss,M.,Sykorová,Z.,Garnica,S.,Riess,K.,Martos,F.,Krause,C.,Oberwinkler,F.,Bauer,R.,and Redecker,D.(2011).Sebacinales everywhere:Previously overlooked ubiquitous fungal endophytes.PLoS ONE 6,e16793.
Xing,X.,Ma,X.,Deng,Z.,Chen,J.,Wu,F.,and Guo,S.(2013).Specificity and preference of mycorrhizal associations in two species of the genus Dendrobium(Orchidaceae).Mycorrhiza 23,317-324.
Xing,X.,Ma,X.,Men,J.,Chen,Y.,and Guo,S.(2017).Phylogenetic constrains on mycorrhizal specificity in eight Dendrobium(Orchidaceae)species.Sci China Life Sci 60,536-544.
Yagame,T.,Ogura-Tsujita,Y.,Kinoshita,A.,Iwase,K.,and Yukawa,T.(2016).Fungal partner shifts during the evolution of mycoheterotrophy in Neottia.Am J Bot 103,1630-1641.
Yuan,L.,Yang,Z.L.,Li,S.Y.,Hu,H.,and Huang,J.L.(2010).Mycorrhizal specificity,preference,and plasticity of six slipper orchids from South Western China.Mycorrhiza 20,559-568.
Bengtsson-Palme,J.,Ryberg,M.,Hartmann,M.,Branco,S.,Wang,Z.,Godhe,A.,et al.(2013).Improved software detection and extraction of ITS1 and ITS2 from ribosomal ITS sequences of fungi and other eukaryotes for analysis of environmental sequencing data.Methods Ecol Evol 4,914-919.
Bidartondo,M.I.,Burghardt,B.,Gebauer,G.,Bruns,T.D.,and Read,D.J.(2004).Changing partners in the dark:isotopic and molecular evidence of ectomycorrhizal liaisons between forest orchids and trees.Proc RSoc London Ser B-Biol Sci 271,1799-1806.
Cameron,D.D.,Johnson,I.,Read,D.J.,and Leake,J.R.(2008).Giving and receiving:measuring the carbon cost of mycorrhizas in the green orchid,Goodyera repens.New Phytol 180,176-184.
Cribb,P.J.,Kell,S.P.,Dixon,K.W.,and Barrett,R.L.(2003).Orchid conservation:a global perspective.In Dixon KW,Kell SP,Barrett RL,Cribb PJ,eds.(Kota Kinabalu,Sabah,Natural History Publications),pp.1-24.
Darriba,D.,Taboada,G.L.,Doallo,R.,and Posada,D.(2012).jModelTest2:more models,new heuristics and parallel computing.Nat Methods 9,772.
De Cáceres,M.,Legendre,P.,and Moretti,M.(2010).Improving indicator species analysis by combining groups of sites.Oikos 119,1674-1684.
Dearnaley,J.D.W.(2007).Further advances in orchid mycorrhizal research.Mycorrhiza 17,475-486.
Dearnaley,J.D.W.,Martos,F.,and Selosse,M.A.(2012).Orchid mycorrhizas:molecular ecology,physiology,evolution and conservation aspects.In Fungal Association,2nd ed.B,Hock.(Berlin:SpringerVerlag),pp.207-230.
Edgar,R.C.(2010).Search and clustering orders of magnitude faster than BLAST.Bioinformatics 26,2460-2461.
Edgar,R.C.(2013).UPARSE:highly accurate OTU sequences from microbial amplicon reads.Nat Methods 10,996-998.
Esposito,F.,Jacquemyn,H.,Waud,M.,and Tyteca,D.(2016).Mycorrhizal fungal diversity and community composition in two closely related Platanthera(Orchidaceae)species.PLoS ONE 11,e0164108.
van der Heijden,M.G.A.,Martin,F.M.,Selosse,M.A.,and Sanders,I.R.(2015).Mycorrhizal ecology and evolution:the past,the present,and the future.New Phytol 205,1406-1423.
Ihrmark,K.,B?deker,I.T.M.,Cruz-Martinez,K.,Friberg,H.,Kubartova,A.,Schenck,J.,Strid,Y.,Stenlid,J.,Brandstr?m-Durling,M.,Clemmensen,K.E.,et al.(2012).New primers to amplify the fungal ITS2 region-evaluation by 454-sequencing of artificial and natural communities.FEMS Microbiol Ecol 82,666-677.
Ives,A.R.,and Godfray,H.C.J.(2006).Phylogenetic analysis of trophic associations.Am Natist 168,E1-E14.
Jacquemyn,H.,Brys,R.,Lievens,B.,and Wiegand,T.(2012a).Spatial variation in below-ground seed germination and divergent mycorrhizal associations correlate with spatial segregation of three co-occurring orchid species.J Ecol 100,1328-1337.
Jacquemyn,H.,Brys,R.,Merckx,V.S.F.T.,Waud,M.,Lievens,B.,and Wiegand,T.(2014).Coexisting orchid species have distinct mycorrhizal communities and display strong spatial segregation.New Phytol 202,616-627.
Jacquemyn,H.,Brys,R.,Waud,M.,Busschaert,P.,and Lievens,B.(2015a).Mycorrhizal networks and coexistence in species-rich orchid communities.New Phytol 206,1127-1134.
Jacquemyn,H.,Deja,A.,De hert,K.,Cachapa Bailarote,B.,and Lievens,B.(2012b).Variation in mycorrhizal associations with Tulasnelloid fungi among populations of five Dactylorhiza species.PLoS ONE 7,e42212.
Jacquemyn,H.,Honnay,O.,Cammue,B.P.A.,Brys,R.,and Lievens,B.(2010).Low specificity and nested subset structure characterize mycorrhizal associations in five closely related species of the genus Orchis.Mol Ecol 19,4086-4095.
Jacquemyn,H.,Merckx,V.,Brys,R.,Tyteca,D.,Cammue,B.P.A.,Honnay,O.,and Lievens,B.(2011).Analysis of network architecture reveals phylogenetic constraints on mycorrhizal specificity in the genus Orchis(Orchidaceae).New Phytol 192,518-528.
Jacquemyn,H.,Waud,M.,Brys,R.,Lallemand,F.,Courty,P.E.,Robionek,A.,and Selosse,M.A.(2017).Mycorrhizal associations and trophic modes in coexisting orchids:an ecological continuum between autoand mixotrophy.Front Plant Sci 8,1497.
Jacquemyn,H.,Waud,M.,Lievens,B.,and Brys,R.(2016a).Differences in mycorrhizal communities between Epipactis palustris,E.helleborine and its presumed sister species E.neerlandica.Ann Bot 118,105-114.
Jacquemyn,H.,Waud,M.,Merckx,V.S.F.T.,Lievens,B.,and Brys,R.(2015b).Mycorrhizal diversity,seed germination and long-term changes in population size across nine populations of the terrestrial orchid Neottia ovata.Mol Ecol 24,3269-3280.
Jacquemyn,H.,Waud,M.,Merckx,V.S.F.T.,Brys,R.,Tyteca,D.,Hedrén,M.,and Lievens,B.(2016b).Habitat-driven variation in mycorrhizal communities in the terrestrial orchid genus Dactylorhiza.Sci Rep 6,37182.
Kembel,S.W.,Cowan,P.D.,Helmus,M.R.,Cornwell,W.K.,Morlon,H.,Ackerly,D.D.,Blomberg,S.P.,and Webb,C.O.(2010).Picante:R tools for integrating phylogenies and ecology.Bioinformatics 26,1463-1464.
Larkin,M.A.,Blackshields,G.,Brown,N.P.,Chenna,R.,McGettigan,P.A.,McWilliam,H.,Valentin,F.,Wallace,I.M.,Wilm,A.,Lopez,R.,et al.(2007).Clustal W and Clustal X version 2.0.Bioinformatics 23,2947-2948.
Li,N.Q.,Wu,J.G.,Jiang,W.J.,Mu,X.Y.,and Cheng,J.(2013).Biodiversity and conservation of orchids in Songshan National Nature Reserve,Beijing.Plant Sci J 31,510.
Martos,F.,Munoz,F.,Pailler,T.,Kottke,I.,Gonneau,C.,and Selosse,M.A.(2012).The role of epiphytism in architecture and evolutionary constraint within mycorrhizal networks of tropical orchids.Mol Ecol21,5098-5109.
McCormick,M.K.,and Jacquemyn,H.(2014).What constrains the distribution of orchid populations?New Phytol 202,392-400.
McCormick,M.K.,Lee Taylor,D.,Juhaszova,K.,Burnett Jr,R.K.,Whigham,D.F.,and O’neill,J.P.(2012).Limitations on orchid recruitment:not a simple picture.Mol Ecol 21,1511-1523.
McKendrick,S.L.,Leake,J.R.,Taylor,D.L.,and Read,D.J.(2002).Symbiotic germination and development of the myco-heterotrophic orchid Neottia nidus-avis in nature and its requirement for locally distributed Sebacina spp..New Phytol 154,233-247.
Mujica,M.I.,Saez,N.,Cisternas,M.,Manzano,M.,Armesto,J.J.,and Pérez,F.(2016).Relationship between soil nutrients and mycorrhizal associations of two Bipinnula species(Orchidaceae)from central Chile.Ann Bot 118,149-158.
Ogura-Tsujita,Y.,Yokoyama,J.,Miyoshi,K.,and Yukawa,T.(2012).Shifts in mycorrhizal fungi during the evolution of autotrophy to mycoheterotrophy in Cymbidium(Orchidaceae).Am J Bot 99,1158-1176.
Oja,J.,Vahtra,J.,Bahram,M.,Kohout,P.,Kull,T.,Rannap,R.,K?ljalg,U.,and Tedersoo,L.(2017).Local-scale spatial structure and community composition of orchid mycorrhizal fungi in semi-natural grasslands.Mycorrhiza 27,355-367.
Oksanen,J.,Blanchet,F.G.,Kindt,R.,Legendre,P.,Minchin,P.R.,O’Hara,R.B.,Simpson,G.L.,Solymos,P.,Stevens,M.H.H.,and Wagner,H.(2015).Vegan:Community Ecology Package Version 2.3-2.http://cran.r-project.org/.
Rasmussen,H.N.,and Rasmussen,F.N.(2009).Orchid mycorrhiza:implications of a mycophagous life style.Oikos 118,334-345.
Schloss,P.D.,Westcott,S.L.,Ryabin,T.,Hall,J.R.,Hartmann,M.,Hollister,E.B.,Lesniewski,R.A.,Oakley,B.B.,Parks,D.H.,Robinson,C.J.,et al.(2009).Introducing mothur:open-source,platformindependent,community-supported software for describing and comparing microbial communities.Appl Environ Microbiol 75,7537-7541.
Shefferson,R.P.,Weiss,M.,Kull,T.,and Taylor,D.L.(2005).High specificity generally characterizes mycorrhizal association in rare lady’s slipper orchids,genus Cypripedium.Mol Ecol 14,613-626.
Stamatakis,A.,Hoover,P.,and Rougemont,J.(2008).A rapid bootstrap algorithm for the RAxML web servers.Systatic Biol 57,758-771.
Stark,C.,Babik,W.,and Durka,W.(2009).Fungi from the roots of the common terrestrial orchid Gymnadenia conopsea.Mycol Res 113,952-959.
Swarts,N.D.,and Dixon,K.W.(2009).Terrestrial orchid conservation in the age of extinction.Ann Bot 104,543-556.
Swarts,N.D.,Sinclair,E.A.,Francis,A.,and Dixon,K.W.(2010).Ecological specialization in mycorrhizal symbiosis leads to rarity in an endangered orchid.Mol Ecol 19,3226-3242.
Taylor,D.L.,and McCormick,M.K.(2008).Internal transcribed spacer primers and sequences for improved characterization of basidiomycetous orchid mycorrhizas.New Phytol 177,1020-1033.
Tedersoo,L.,Bahram,M.,P?lme,S.,K?ljalg,U.,Yorou,N.S.,Wijesundera,R.,Villarreal Ruiz,L.,Vasco-Palacios,A.M.,Thu,P.Q.,Suija,A.,et al.(2014).Global diversity and geography of soil fungi.Science 346,1256688.
Tě?itelová,T.,Jersáková,J.,Roy,M.,Kubátová,B.,Tě?itel,J.,Urfus,T.,Trávní?ek,P.,and Suda,J.(2013).Ploidy-specific symbiotic interactions:divergence of mycorrhizal fungi between cytotypes of the Gymnadenia conopsea group(Orchidaceae).New Phytol 199,1022-1033.
Tě?itelová,T.,Kotilínek,M.,Jersáková,J.,Joly,F.X.,Ko?nar,J.,Tatarenko,I.,and Selosse,M.A.(2015).Two widespread green Neottia species(Orchidaceae)show mycorrhizal preference for Sebacinales in various habitats and ontogenetic stages.Mol Ecol 24,1122-1134.
Waud,M.,Busschaert,P.,Lievens,B.,and Jacquemyn,H.(2016a).Specificity and localised distribution of mycorrhizal fungi in the soil may contribute to co-existence of orchid species.Fungal Ecol 20,155-165.
Waud,M.,Busschaert,P.,Ruyters,S.,Jacquemyn,H.,and Lievens,B.(2014).Impact of primer choice on characterization of orchid mycorrhizal communities using 454 pyrosequencing.Mol Ecol Resour 14,679-699.
Waud,M.,Wiegand,T.,Brys,R.,Lievens,B.,and Jacquemyn,H.(2016b).Nonrandom seedling establishment corresponds with distancedependent decline in mycorrhizal abundance in two terrestrial orchids.New Phytol 211,255-264.
Weiss,M.,Sykorová,Z.,Garnica,S.,Riess,K.,Martos,F.,Krause,C.,Oberwinkler,F.,Bauer,R.,and Redecker,D.(2011).Sebacinales everywhere:Previously overlooked ubiquitous fungal endophytes.PLoS ONE 6,e16793.
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