Mycological Progress

, Volume 16, Issue 3, pp 191–203 | Cite as

Species clarification of oyster mushrooms in China and their DNA barcoding

Original Article


Species of the Pleurotus ostreatus complex (oyster mushrooms) are one of the main groups of cultivated edible mushrooms in China, with abundant strains and a complex genetic background. Although the cultivation of oyster mushrooms is a very important industry in China, the inconsistent nomenclature of the complex is controversial and has led to much confusion and economic loss in the mushroom industry. In this study, we performed molecular identification of the species complex based on sequences of the internal transcribed spacer region (ITS), translation elongation factor 1-α (TEF1), and two genes that encode subunits of RNA polymerase II (RPB1 and RPB2). A total of 284 samples with different commercial names gathered from different mushroom spawn preservation centers, companies, and field isolations were investigated via phylogenetic analyses inferred from a single locus or multi-loci. Our analyses indicated that 56% of the strains were labeled with improper scientific names and that all of the strains could be divided into seven lineages, representing one morphological species each, namely P. abieticola, P. eryngii, P. cf. floridanus, P. ostreatus, P. placentodes, P. pulmonarius, and P. tuoliensis. Reference sequences for accurate identification of the taxa were developed or suggested. Meanwhile, we evaluated the suitability of the four candidate segments as DNA barcodes. Our analyses indicated that RPB2 was the most promising candidate segment of DNA barcode for the P. ostreatus species complex, taking into consideration the polymorphisms and other aspects of the four markers.


Phylogenetic analyses Species recognition Strain identification DNA sequence markers 



The authors are very grateful to Prof. Dr. Y.C. Zhao (Yunnan Academy of Agricultural Sciences), Mr. F. Nie (Anhui Academy of Agricultural Sciences), Ms. J. Yang (Fujian Academy of Agricultural Sciences), Dr. P. Zhang (Hunan Normal University), Prof. J.W. Shen (Henan Agricultural University), and Dr. X.D. Yu (Shenyang Agricultural University) for providing strains and specimens for this study. The anonymous reviewers are acknowledged for their constructive comments and suggestions. This study is supported by the National Basic Research Program of China (973 Program, No. 2014CB138305) and the Large-scale Scientific Facilities of the Chinese Academy of Sciences (2009-LSFGBOWS-01).

Supplementary material

11557_2016_1266_Fig5_ESM.gif (362 kb)
Suppl. Fig. 1

Phylogenetic tree inferred from ML analysis with branch support obtained by ML and BI analyses based on a four-locus (ITS, TEF1, RPB1, and RPB2) dataset. Branch support values are indicated by numbers above branches (ML-BP/BI-PP). Taxon labels indicate strain number listed in Suppl. Table 1. Location of P. ostreatus (P382, P383, CCMSSC 00323) and P. cf. floridanus (P289, P290) are highlighted in blue and red, respectively (GIF 361 kb)

11557_2016_1266_MOESM1_ESM.tif (2.9 mb)
High Resolution (TIF 3012 kb)
11557_2016_1266_Fig6_ESM.gif (1.2 mb)
Suppl. Fig. 2

Phylogenetic tree inferred from ML analysis with branch support obtained by ML and BI analyses based on ITS, TEF1, RPB1, and RPB2. Branch support values are indicated by numbers above branches (ML-BP/BI-PP). Taxon labels indicate strain number listed in Suppl. Table 1. Strains of P. ostreatus and P. cf. floridanus in single gene phylogeny (ITS, TEF1, and RPB1) are highlighted in purple and red, respectively (GIF 1184 kb)

11557_2016_1266_MOESM2_ESM.tif (31.1 mb)
High Resolution (TIF 31805 kb)
11557_2016_1266_Fig7_ESM.gif (445 kb)
Suppl. Fig. 3

Phylogenetic tree inferred from ML analysis with branch support obtained by ML and BI analyses based on RPB2. Branch support values are indicated by numbers above branches (ML-BP/BI-PP). The strain names labeled on our collections correspond to Suppl. Fig. 2 (GIF 445 kb)

11557_2016_1266_MOESM3_ESM.tif (250.3 mb)
High Resolution (TIF 256266 kb)
11557_2016_1266_Fig8_ESM.gif (130 kb)
Suppl. Fig. 4

Phylogenetic tree inferred from ML analysis with branch support obtained by ML and BI analyses based on TEF1(a) and, RPB1(b). Branch support values are indicated by numbers above branches (ML-BP/BI-PP). The samples of P. ostreatus and P. cf. floridanus were highlighted in different colors (GIF 130 kb)

11557_2016_1266_Fig9_ESM.gif (126 kb)
Suppl. Fig. 4

Phylogenetic tree inferred from ML analysis with branch support obtained by ML and BI analyses based on TEF1(a) and, RPB1(b). Branch support values are indicated by numbers above branches (ML-BP/BI-PP). The samples of P. ostreatus and P. cf. floridanus were highlighted in different colors (GIF 130 kb)

11557_2016_1266_MOESM4_ESM.tif (63.4 mb)
High Resolution (TIF 64912 kb)
11557_2016_1266_MOESM5_ESM.tif (116.3 mb)
High Resolution (TIF 119085 kb)
11557_2016_1266_Fig10_ESM.jpg (1 mb)
Suppl. Fig. 5

Phylogenetic relationships among reference collections and other samples of Pleurotus inferred from ITS sequences using ML analysis. Branch support values are indicated by numbers above branches (ML-BP ≥ 70%). Accession numbers for sequences retrieved from GenBank database are listed in Suppl. Table 1. Specimens in P. ostreatus species complex from China are highlighted in blue. Reference collections in single-locus and multi-locus phylogenetic analyses are labeled with a black rectangle (JPEG 1.03 MB)

11557_2016_1266_MOESM6_ESM.doc (566 kb)
Suppl. Table 1 Collections of Pleurotus species used in this study and their GenBank accession numbers (DOC 565 kb)


  1. Albertó EO, Petersen RH, Karen WH, Bernardo L (2002) Miscellaneous notes on Pleurotus. Persoonia 18:55–69Google Scholar
  2. Anderson NA, Schwandt JW (1973) The Pleurotus ostreatus-sapidus species complex. Mycologia 65:28–35CrossRefGoogle Scholar
  3. Armstrong KF, Ball SL (2005) DNA barcodes for biosecurity: invasive species identification. Philos Trans R Soc B 360:1813–1823CrossRefGoogle Scholar
  4. Avin FA, Bhassu S, Shin TY, Sabaratnam V (2012) Molecular classification and phylogenetic relationships of selected edible Basidiomycetes species. Mol Biol Rep 39:7355–7364PubMedCrossRefGoogle Scholar
  5. Avise JC, Wollenberg K (1997) Phylogenetics and the origin of species. Proc Natl Acad Sci USA 94:7748–7755PubMedPubMedCentralCrossRefGoogle Scholar
  6. Balasundaram SV, Engh IB, Skrede I, Kauserud H (2015) How many DNA markers are needed to reveal cryptic fungal species? Fungal Biol 119:940–945PubMedCrossRefGoogle Scholar
  7. Bao DP, Ishihara H, Mori N (2004) Phylogenetic analysis of oyster mushrooms (Pleurotus spp.) based on restriction fragment length polymorphism of 5′portion of 26s rDNA. J Wood Sci 50:169–176Google Scholar
  8. Bobovčák M, Kuniaková R, Gabriž J, Majtán J (2010) Effect of Pleuran (β-glucan from Pleurotus ostreatus) supplementation on cellular immune response after intensive exercise in elite athletes. Appl Physiol Nutr Metab 35:755–762PubMedCrossRefGoogle Scholar
  9. Buchanan PK (1993) Identification, names and nomenclature of common edible mushrooms. In: Chang ST, Buswell JA, Chiu SW (eds) Mushroom Biology and Mushroom Products. The Chinese University Press, Hong Kong, pp 21–32Google Scholar
  10. Bunyard BA, Chaichuchote S, Nicholson MS, Royse DJ (1996) Ribosomal DNA analysis for resolution of genotypic classes of Pleurotus. Mycol Res 100:143–150CrossRefGoogle Scholar
  11. Choi DB, Ding JL, Cha WS (2007) Homology search of genus Pleurotus using an internal transcribed spacer region. Korean J Chem Eng 24:408–412CrossRefGoogle Scholar
  12. Cox JPL (2001) Barcoding objects with DNA. Analyst 126:545–547PubMedCrossRefGoogle Scholar
  13. Dentinger BTM, Didukh MY, Moncalvo JM (2011) Comparing COI and ITS as DNA barcode markers for mushrooms and allies (Agaricomycotina). PLoS One 6:e25081PubMedPubMedCentralCrossRefGoogle Scholar
  14. Dettman JR, Jacobson DJ, Turner E, Pringle A, Taylor JW (2003) Reproductive isolation and phylogenetic divergence in Neurospora: comparing methods of species recognition in a model eukaryote. Evolution 57:2721–2741PubMedCrossRefGoogle Scholar
  15. Dettman JR, Jacobson DJ, Taylor JW (2006) Multilocus sequence data reveal extensive phylogenetic species diversity within the Neurospora discreta complex. Mycologia 98:436–446PubMedCrossRefGoogle Scholar
  16. Doyle JJ, Doyle JL (1987) A rapid DNA isolation procedure for small quantities of fresh leaf material. Phytochem Bull 19:11–15Google Scholar
  17. Du XH, Zhao Q, Yang ZL, Hansen K, Taskin H, Buyukalaca S, Dewsbury D, Moncalvo JM, Douhan GW, Robert VARG, Crous PW, Rehner SA, Rooney AP, Sink S, O′Donnell K (2012) How well do ITS rDNA sequences differentiate species of true morels (Morchella). Mycologia 104:1351–1368PubMedCrossRefGoogle Scholar
  18. Dupuis JR, Roe AD, Sperling FAH (2012) Multi-locus species delimitation in closely related animals and fungi: one marker is not enough. Mol Ecol 21:4422–4436PubMedCrossRefGoogle Scholar
  19. Eason RG, Pourmand N, Tongprasit W, Herman ZS, Anthony KP, Jejelowo O, Davis R, Stolc V (2004) Characterization of synthetic DNA barcodes in Saccharomyces cerevisiae gene-deletion strains. Proc Natl Acad Sci USA 2:285–288Google Scholar
  20. Edgar RC (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Res 32:1792–1797PubMedPubMedCentralCrossRefGoogle Scholar
  21. Eger G (1976) Rapid method for breeding Pleurotus ostreatus. Mushroom Sci 9:567–576Google Scholar
  22. Eger G, Eden G, Wissig E (1976) Pleurotus ostreatus-breeding potential of a new cultivated mushroom. Theor Appl Genet 47:155–163PubMedCrossRefGoogle Scholar
  23. Estrada AE, Jimenez-Gasco MM, Royse DJ (2010) Pleurotus eryngii species complex: sequence analysis and phylogeny based on partial EF1α and RPB2 genes. Fungal Biol 114:421–428PubMedCrossRefGoogle Scholar
  24. Eugenio CP, Anderson NA (1968) The genetics and cultivation of Pleurotus ostreatus. Mycologia 60:627–634CrossRefGoogle Scholar
  25. Fell JW, Scorzetti G, Statzell-Tallman A, Boundy-Mills K (2007) Molecular diversity and intragenomic variability in the yeast genus Xanthophyllomyces: The origin of Phaffia rhodozyma? FEMS Yeast Res 7:1399–1408PubMedCrossRefGoogle Scholar
  26. Froslev TG, Matheny PB, Hibbett DS (2005) Lower level relationships in the mushroom genus Cortinarius (Basidiomycota, Agaricales): a comparison of RPB1, RPB2, and ITS phylogenies. Mol Phylogenet Evol 37:602–618PubMedCrossRefGoogle Scholar
  27. Gao S, Huang CY, Chen Q, Bian YB, Zhang JX (2008) Phylogenetic relationship of Pleurotus species based on nuclear large subunit ribosomal DNA sequence. J Plant Genet Resour 9:328–334Google Scholar
  28. Garbelotto M, Otrosina WJ, Cobb FW, Bruns TD (1998) The European S and F intersterility groups of Heterobasidion annosum may represent sympatric protospecies. Can J Bot 76:397–409Google Scholar
  29. Gonzalez P, Labarère J (2000) Phylogenetic relationships of Pleurotus species according to the sequence and secondary structure of the mitochondrial small-subunit rRNA V4, V6 and V9 domains. Microbiology 146:209–221PubMedCrossRefGoogle Scholar
  30. Guzmán G, Montoya L, Mata G, Salmones D (1994) Studies in the genus Pleurotus, III: The varieties of P. ostreatus-complex based in interbreeding strains and in the study of basidiomata obtained in culture. Mycotaxon 50:365–378Google Scholar
  31. Guzmán G, Montoya L, Bandala VM, Mata G, Salmones D (1995) Studies in the genus Pleurotus, IV. Observations on the pink forms growing in Mexico based in the interbreeding of two different strains. Mycotaxon 53:247–259Google Scholar
  32. Hadar Y, Kerem Z, Gorodecki B (1993) Biodegradation of lignocellulosic agricultural wastes by Pleurotus ostreatus. J Biotechnol 30:133–139CrossRefGoogle Scholar
  33. Hall TA (1999) BioEdit: A user-friendly biological sequence alignment editor and analysis program for windows 95/98/NT. Nucleic Acids Symp Ser 41:95–98Google Scholar
  34. Hebert PDN, Cywinska A, Ball SL, DeWaard JR (2003) Biological identification through DNA barcodes. Proc R Soc London B 270:313–231CrossRefGoogle Scholar
  35. Hughes KW, Petersen RH (2001) Apparent recombination or gene conversion in the ribosomal ITS region of a Flammulina (Fungi, Agaricales) hybrid. Mol Biol Evol 18:94–96PubMedCrossRefGoogle Scholar
  36. Iracalbal B, Zervakis G, Labarere J (1995) Molecular systematics of the genus Pleurotus: analysis of restriction polymorphisms in ribosomal DNA. Microbiology 141:1479–1490CrossRefGoogle Scholar
  37. Ito Y, Fushimi T, Yanagi SO (1998) Discrimination of species and strains of basidiomycete genus Coprinus by random amplified polymorphic DNA (RAPD) analysis. Mycoscience 39:361–365CrossRefGoogle Scholar
  38. James SA, West C, Davey RP, Dicks J, Robert LN (2016) Prevalence and dynamics of ribosomal DNA micro-heterogeneity are linked to population history in two contrasting yeast species. Sci Rep. doi: 10.1038/srep28555 Google Scholar
  39. Justo A, Vizzini A, Minnis AM, Menolli N, Capelari M, Rodriguez O, Malysheva E, Contu M, Ghignone S, Hibbett DS (2011) Phylogeny of the Pluteaceae (Agaricales, Basidiomycota): taxonomy and character evolution. Fungal Biol 115:1–20PubMedCrossRefGoogle Scholar
  40. Kauserud H, Svegården IB, Decock C, Hallenberg N (2007) Hybridization among cryptic species of the cellar fungus Coniophora puteana (Basidiomycota). Mol Ecol 16:389–399PubMedCrossRefGoogle Scholar
  41. Kawai G, Babasaki K, Neda H (2008) Taxonomic position of a Chinese Pleurotus “Bai-Ling-Gu”: it belongs to Pleurotus eryngii (DC.: Fr.) Quél. and evolved independently in China. Mycoscience 49:75–87CrossRefGoogle Scholar
  42. Kimura M (1980) A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide-sequences. J Mol Evol 16:111–120PubMedCrossRefGoogle Scholar
  43. Kiss L (2012) Limits of nuclear ribosomal DNA internal transcribed spacer (ITS) sequences as species barcodes for Fungi. Proc Natl Acad Sci USA 109:e1811PubMedPubMedCentralCrossRefGoogle Scholar
  44. Le Gac M, Hood ME, Fournier E, Giraud T (2007) Phylogenetic evidence of host-specific cryptic species in the anther smut fungus. Evolution 61:15–26PubMedCrossRefGoogle Scholar
  45. Le QV, Won HK, Lee TS, Lee CY, Lee HS, Ro HS (2008) Retrotransposon microsatellite amplified polymorphism strains fingerprinting markers applicable to various mushroom species. Mycobiology 36:161–166PubMedPubMedCentralCrossRefGoogle Scholar
  46. Lee HK, Shin CS, Min KB, Choi KS, Kim BG, Yoo YB, Min KH (2000) Molecular systematics of the genus Pleurotus using sequence-specific oligonucleotide probes. Sci Cultivation Edible Fungi 1–2:207–213Google Scholar
  47. Li Y, Jiao L, Yao YJ (2013) Non-concerted ITS evolution in fungi, as revealed from the important medicinal fungus Ophiocordyceps sinensis. Mol Phylogenet Evol 68:373–379PubMedCrossRefGoogle Scholar
  48. Librado P, Rozas J (2009) DnaSP v5: a software for comprehensive analysis of DNA polymorphism data. Bioinformatics 25:1451–1452PubMedCrossRefGoogle Scholar
  49. Lindner DL, Banik MT (2011) Intragenomic variation in the ITS rDNA region obscures phylogenetic relationships and inflates estimates of operational taxonomic units in genus Laetiporus. Mycologia 103:731–740PubMedCrossRefGoogle Scholar
  50. Liu YJ, Hall BD (1999) Phylogenetic relationships among ascomycetes: Evidence from an RNA polymerase II subunit. Mol Biol Evol 16:1799–1808PubMedCrossRefGoogle Scholar
  51. Liu SY, Zhang A, Li Y (2012) Principle and procedure of fungal DNA barcoding. J Fungal Res 10:205–209Google Scholar
  52. Liu XB, Feng B, Li J, Yan C, Yang ZL (2016a) Genetic diversity and breeding history of Winter Mushroom (Flammulina velutipes) in China uncovered by genomic SSR markers. Gene 591:227–235PubMedCrossRefGoogle Scholar
  53. Liu XB, Li J, Horak E, Yang ZL (2016b) Pleurotus placentodes, originally described from Sikkim, rediscovered after 164 years. Phytotaxa 267:137–145CrossRefGoogle Scholar
  54. Mapook A, Boonmee S, Ariyawansa HA, Tibpromma S, Campesori E, Gareth Jones EB, Bahkail AH, Hyde KD (2016) Taxonomic and phylogenetic placement of Nodulosphaeria. Mycol Prog. doi: 10.1007/s11557-016-1176-x Google Scholar
  55. Matheny PB (2005) Improving phylogenetic inference of mushrooms with RPB1 and RPB2 nucleotide sequences (Inocybe, Agaricales). Mol Phylogenet Evol 35:1–20PubMedCrossRefGoogle Scholar
  56. Matheny PB (2006) PCR Primers to Amplify and Sequence rpb2 (RNA polymerase II second largest subunit) in the Basidiomycota (Fungi).
  57. Matheny PB, Liu YJ, Ammirati JF, Hall B (2002) Using RPB1 sequences to improve phylogenetic inference among mushrooms (Inocybe, Agaricales). Am J Bot 89:688–698PubMedCrossRefGoogle Scholar
  58. Meng Y, Jiang CS, Liao WT, Zhang YZ (2003) AFLP fingerprinting map analysis of Pleurotus ostreatus. J Genet Genomics 30:1140–1146Google Scholar
  59. Menolli N Jr, Breternitz BS, Capelari M (2014) The genus Pleurotus in Brazil: a molecular and taxonomic overview. Mycoscience 55:378–389CrossRefGoogle Scholar
  60. Meyer CP, Paulay G (2005) DNA barcoding: Error rates based on comprehensive sampling. PLoS Biol 3:e422PubMedPubMedCentralCrossRefGoogle Scholar
  61. Montcalvo JM, Lutzoni FM, Rehner SA, Johnson J, Vilgalys R (2000) Phylogenetic relationships of agaric fungi based on nuclear large subunit ribosomal DNA sequences. Syst Biol 49:278–305CrossRefGoogle Scholar
  62. Morin L, van der Merwe M, Hartley D, Müller P (2009) Putative natural hybrid between Puccinia lagenophorae and an unknown rust fungus on Senecio madagascariensis in KwaZulu-Natal, South Africa. Mycol Res 113:725–736PubMedCrossRefGoogle Scholar
  63. Moriwaki J, Tsukiboshi T, Sato T (2002) Grouping of Colletotrichum species in Japan based on rDNA sequences. J Gen Plant Pathol 68:307–320CrossRefGoogle Scholar
  64. Mou CJ, Cao YQ, Ma JL (1987) A new variety of Pleurotus eryngii and its cultural characters. Mycosystema 6:153–156Google Scholar
  65. Murakami S, Takemaru T (1990) Genetic studies of Pleurotus salmoneostramineus forming albino basidiocarps. Rep Tottori Mycol Inst 28:199–204Google Scholar
  66. Newcombe G, Stirling B, McDonald S, Bradshaw HD (2000) Melampsora × columbiana, a natural hybrid of M. medusa and M. occidentalis. Mycol Res 104:261–274CrossRefGoogle Scholar
  67. Nylander JAA (2004) MrModeltest v2.2 Program distributed by the author. In: Evolutionary Biology Centre, Uppsala UniversityGoogle Scholar
  68. Omarini AB, Plagemann I, Schimanski S, Krings U, Berger RG (2014) Crosses between monokaryons of Pleurotus sapidus or Pleurotus florida show an improved biotransformation of (+)-valencene to (+)-nookatone. Bioresour Technol 171:113–119PubMedCrossRefGoogle Scholar
  69. Pegler DN (1996) Hyphal analysis of basidiomata. Mycol Res 100:129–142CrossRefGoogle Scholar
  70. Ravash R, Shiran B, Alavi AA, Bayat F, Rajaee S, Zervakis GI (2010) Genetic variability and molecular phylogeny of Pleurotus eryngii species-complex isolates from Iran, and notes on the systematics of Asiatic populations. Mycol Prog 9:181–194CrossRefGoogle Scholar
  71. Rehner S (2001) Primers for the elongation factor 1-α (tef1-α). primer. pdf
  72. Reid DA, Eicker A, De Kock A (1998) Pleurotus fuscosquamulosus–a new species of Pleurotus subgenus Coremiopleurotus from South Africa. Mycotaxon 66:137–152Google Scholar
  73. Roger AJ, Sandblom O, Doolittle WF, Philippe H (1999) An evaluation of elongation factor 1 α as a phylogenetic marker for eukaryotes. Mol Biol Evol 16:218–233PubMedCrossRefGoogle Scholar
  74. Ronquist F, Huelsenbeck JP (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19:1572–1574PubMedCrossRefGoogle Scholar
  75. Rooney AP, Ward TJ (2005) Evolution of a large ribosomal RNA multigene family in filamentous fungi: Birth and death of a concerted evolution paradigm. Proc Natl Acad Sci U S A 102:5084–5089PubMedPubMedCentralCrossRefGoogle Scholar
  76. Saitou N, Nei M (1987) The neighbor-joining method: a new method for reconstructing phylogenetic trees. Mol Biol Evol 4:406–425PubMedGoogle Scholar
  77. Salas-Lizana R, Santini NS, Miranda-Perez A, Pinero DI (2012) The Pleistocene glacial cycles shaped the historical demography and phylogeography of a pine fungal endophyte. Mycol Prog 11:569–581CrossRefGoogle Scholar
  78. Samerpitak K, Van der Linde E, Choi HJ, Gerrits van den Ende AHG, Machouart M, Gueidan C, de Hoog GS (2014) Taxonomy of Ochroconis, genus including opportunistic pathogens on humans and animals. Fungal Divers 65:89–126CrossRefGoogle Scholar
  79. Sánchez C (2004) Modern aspects of mushrooms culture technology. Appl Microbiol Biotechnol 64:756–762PubMedCrossRefGoogle Scholar
  80. Schoch CL, Seifert KA, Huhndorf S et al (2012) Nuclear ribosomal internal transcribed spacer (ITS) region as a universal DNA barcode marker for Fungi. Proc Natl Acad Sci USA 109:6241–6246PubMedPubMedCentralCrossRefGoogle Scholar
  81. Segedin BP, Buchnan PK, Wilkie JP (1995) Studies in the Agaricales of New Zealand: new species, new records and renamed species of Pleurotus (Pleurotaceae). Aust Syst Bot 8:453–482CrossRefGoogle Scholar
  82. Seifert KA, Crous PW (2008) The All-Fungi DNA Barcoding Campaign (FunBOL). Persoonia 20:106Google Scholar
  83. Seifert KA, Samson RA, DeWaard JR, Houbraken J, Lévesque CA, Moncalvo JM, Louis-Seize G, Hebert PDN (2007) Prospects for fungus identification using CO1 DNA barcodes, with Penicillium as a test case. Proc Natl Acad Sci USA 104:3901–3906PubMedPubMedCentralCrossRefGoogle Scholar
  84. Selvakumar P, Rajasekar S, Babu AG, Periasamy K, Raaman N, Reddy MS (2015) Improving biological efficiency of Pleurotus strain through protoplast fusion between P. ostreatus var. florida and P. djamor var. roseus. Food Sci Biotechnol 24:1741–1748CrossRefGoogle Scholar
  85. Shen JW, Zhao X, Li Y, Guan YY, Wang Z, Qi YC, Zhang JX (2011) Evaluation of germplasm resources with cultivated Pleurotus spp. strains in Henan Province. J Henan Agric Univ 45:297–301Google Scholar
  86. Shen LL, Chen JJ, Wang M, Cui BK (2016) Taxonomy and multi-gene phylogeny of Haploporus (Polyporales, Basidiomycota). Mycol Prog 15:731–742CrossRefGoogle Scholar
  87. Shnyreva AA, Shnyreva AV (2015) Phylogenetic analysis of Pleurotus species. Russ J Genet 51:177–187CrossRefGoogle Scholar
  88. Shnyreva A, Shtaer O (2006) Differentiation of closely related oyster fungi Pleurotus pulmonarius and P. ostreatus by mating and molecular markers. Russ J Genet 42:539–545CrossRefGoogle Scholar
  89. Shnyreva AV, Belokon YS, Belokon MM, Altukhov YP (2004) Interspecific genetic variability of the oyster mushroom Pleurotus ostreatus as revealed by allozyme gene analysis. Russ J Genet 40:871–881CrossRefGoogle Scholar
  90. Singer R (1986) The Agaricales in modern taxonomy. 4th edit. Koeltz Scientific Books, Koenigstein, pp 174–179Google Scholar
  91. Skrede I, Carlsen T, Stensrud Ø, Kauserud H (2012) Genome wide AFLP markers support cryptic species in Coniophora (Boletales). Fungal Biol 116:778–784PubMedCrossRefGoogle Scholar
  92. Song C, Chen Q, Xu JY, Zhang JX, Bian YB, Huang CY (2011) Application of CO1 for rapid identification of Pleurotus species. Mycosystema 30:663–668Google Scholar
  93. Stamatakis A (2006) RAxML-VI-HPC: Maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 22:2688–2690PubMedCrossRefGoogle Scholar
  94. Stamatakis A, Hoover P, Rougemont J (2008) A rapid bootstrap algorithm for the RAxML web servers. Syst Biol 57:758–771PubMedCrossRefGoogle Scholar
  95. Stephens M, Donnelly P (2003) A comparison of Bayesian methods for haplotype reconstruction from population genotype data. Am J Hum Genet 73:1162–1169PubMedPubMedCentralCrossRefGoogle Scholar
  96. Stielow JB, Lévesque CA, Seifert KA et al (2015) One fungus, which genes? Development and assessment of universal primers for potential secondary fungal DNA barcodes. Persoonia 35:242–263PubMedPubMedCentralCrossRefGoogle Scholar
  97. Tamura K, Peterson D, Peterson N, Steker G, Nei M, Kumar S (2011) MEGA5: molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Mol Biol Evol 28:2731–2739PubMedPubMedCentralCrossRefGoogle Scholar
  98. Tanabe Y, Saikawa M, Watanabe MM, Sugiyama J (2004) Molecular phylogeny of Zygomycota based on EF-1α and RPB1 sequences: limitations and utility of alternative markers to rDNA. Mol Phylogenet Evol 30:438–449PubMedCrossRefGoogle Scholar
  99. Taniguchi M, Suzuki H, Watanabe D, Sahai K, Hoshino K, Tanaka T (2005) Evaluation of pretreatment with Pleurotus ostreatus for enzymatic hydrolysis of rice straw. J Biosci Bioeng 100:637–643PubMedCrossRefGoogle Scholar
  100. Taylor JW, Jacobson DJ, Kroken S, Kasuga T, Geiser DM, Hibbett DS, Fisher MC (2000) Phylogenetic species recognition and species concepts in fungi. Fungal Genet Biol 31:21–32PubMedCrossRefGoogle Scholar
  101. Terakawa H (1960) The incompatibility factors in Pleurotus ostreatus. Set Pap Coll Gen Educ, Univ Tokyo 10:65–71Google Scholar
  102. Vilgalys R, Moncalvo JM, Liou SR, Volovsek M (1996) Recent advances in molecular systematics of the genus Pleurotus. Mushroom Biol Mushroom Prod 91–101Google Scholar
  103. Wang DM, Yao YJ (2005) Intrastrain internal transcribed spacer heterogeneity in Ganoderma species. Can J Microbiol 51:113–121PubMedCrossRefGoogle Scholar
  104. Wang WJ, Wang XL, Wang XC, Yu XD, Li Y, Wei TZ, Yao YJ (2009) Application of DNA Barcoding in Fungal Research. Front Sci 3:4–12Google Scholar
  105. Wasser SP, Sokolov D, Reshetnikov SV, Timor-Tismenetsky M (2000) Dietary Supplements from medicinal mushrooms: diversity of types and variety of regulations. Int J Med Mushrooms 2:1–19CrossRefGoogle Scholar
  106. White TJ, Burns T, Lee S, Taylor J (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In: Innis MA, Gelfand DH, Sninsky JJ, White TJ (eds) PCR protocols: a guide to methods and applications. Academic Press, San Diego, pp 315–322Google Scholar
  107. Wu SR, Zhao CY, Hou B, Tai LM, Gui MY (2013) Analysis on Chinese edible fungus production area layout of nearly five years. Edible Fungi China 32:51–53Google Scholar
  108. Xu K, Kanno M, Yu H, Li Q, Kijima A (2011) Complete mitochondrial DNA sequence and phylogenetic analysis of Zhikong scallop Chlamys farreri (Bivalvia: Pectinidae). Mol Biol Rep 38:3067–3074PubMedCrossRefGoogle Scholar
  109. Yang ZL (2011) Molecular techniques revolutionize knowledge of basidiomycete evolution. Fungal Divers 50:47–58CrossRefGoogle Scholar
  110. Zeng ZQ, Zhao P, Lou J, Zhuang WY, Yu ZH (2012) Selection of a DNA barcode for Nectriaceae from fungal whole genomes. Sci China Life Sci 55:80–88PubMedCrossRefGoogle Scholar
  111. Zervakis GI (1998) Mating competence and biological species within the subgenus Coremiopleurotus. Mycologia 90:1063–1074CrossRefGoogle Scholar
  112. Zervakis GI, Balis C (1996) A pluralist approach in the study of Pleurotus species with emphasis on compatibility and physiology of the European morphotaxa. Mycol Res 100:717–731CrossRefGoogle Scholar
  113. Zervakis G, Sourdis J, Balis C (1994) Genetic variability and systematics of eleven Pleurotus ostreatus species based on isozyme analysis. Mycol Res 98:329–341CrossRefGoogle Scholar
  114. Zervakis G, Venturella G, Papadopoulou K (2001) Genetic polymorphism and taxonomic relationships of the Pleurotus eryngii species-complex as resolved through the analysis of random amplified DNA patterns, isozyme profiles and ecomorphological characters. Microbiology 147:3183–3194PubMedCrossRefGoogle Scholar
  115. Zervakis GI, Ntougias S, Gargano ML, Besi MI, Polemis E, Typas MA, Venturella G (2014) A reappraisal of the Pleurotus eryngii complex–New species and taxonomic combinations based on the application of a polyphasic approach, and an identification key to Pleurotus taxa associated with Apiaceae plants. Fungal Biol 118:814–834PubMedCrossRefGoogle Scholar
  116. Zhang JX, Huang CY, Zhang RY, Guan GP (2004) RAPD and IGS analysis of Pleurotus nebrodensis cultivars in China. Mycosystema 23:514–519Google Scholar
  117. Zhang JX, Huang CY, Ng TB, Wang HX (2006) Genetic polymorphism of ferula mushroom growing on Ferula sinkiangensis. Appl Microbiol Biotechnol 71:304–309PubMedCrossRefGoogle Scholar
  118. Zhao P, Luo J, Zhuang WY, Liu XZ, Wu B (2011) DNA barcoding of the fungal genus Neonectria and the discovery of two new species. Sci China Life Sci 54:664–674PubMedCrossRefGoogle Scholar
  119. Zhao MR, Zhang JX, Chen Q, Wu XL, Gao W, Deng WQ, Huang CY (2016) The famous cultivated mushroom Bailinggu is a separate species of the Pleurotus eryngii species complex. Sci Rep 6:33066PubMedPubMedCentralCrossRefGoogle Scholar
  120. Zheng SY, Zhang JX, Wang HX, Huang CY (2003) Polyphasic taxonomy of cultivated oyster mushroom in China. Edible Fungi China 22:3–6Google Scholar
  121. Zheng HB, Ma ZG, Lv ZZ, Yu ZH (2006) Identification and evaluation of main cultivated species of Pleurotus in China based on ITS sequence analysis. Mycosystema 25:398–407Google Scholar

Copyright information

© German Mycological Society and Springer-Verlag Berlin Heidelberg 2016

Authors and Affiliations

  1. 1.Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of BotanyChinese Academy of SciencesKunmingChina
  2. 2.University of Chinese Academy of SciencesBeijingChina
  3. 3.State Key Laboratory of Conservation and Utilization for Bioresources in YunnanYunnan UniversityKunmingChina

Personalised recommendations