As a first step of a project aimed at the identification of potential biocontrol agents of Rhizoctonia solani, the rice sheath blight fungus, we surveyed the biodiversity of the genus Trichoderma based on sequence of the internal transcribed spacer (ITS) 1 and 2 of the ribosomal RNA gene cluster in paddy fields in Mazandaran province, Northern Iran. Amongst the six obtained species of Trichoderma, T. harzianum and T. virens proved to be the most frequent species in this habitat. Sequence alignment and phylogenetic analysis revealed that the T. harzianum isolates can be divided into 14 different ITS genotypes clustering in four groups. Our results are in agreement with previous molecular studies, which also revealed that T. harzianum is a complex species comprising more or less different ITS genotypes. T. virens was not as diverse as T. harzianum and three different genotypes were distinguished which constituted only one cluster. All T. atroviride and T. hamatum strains had identical ITS sequences.
Bissett, J. 1984. A revision of the genus Trichoderma. I. Sect. Longibrachiatum sect. nov. Can. J. Bot. 62:924–931.
Bissett J. , 'A revision of the genus Trichoderma. I. Sect. Longibrachiatum sect. nov ' (1984 ) 62 Can. J. Bot. : 924 -931 .
Bissett, J. 1991a. A revision of the genus Trichoderma. II. Infrageneric classification. Can. J. Bot. 69:2357–2372.
Bissett J. , 'A revision of the genus Trichoderma. II. Infrageneric classification ' (1991 ) 69 Can. J. Bot. : 2357 -2372 .
Bissett, J. 1991b. A revision of the genus Trichoderma. III. Sect. Pachybasium. Can. J. Bot. 69:2373–2417.
Bissett J. , 'A revision of the genus Trichoderma. III. Sect. Pachybasium ' (1991 ) 69 Can. J. Bot. : 2373 -2417 .
Bissett, J. 1991c. Arevision of the genus Trichoderma. IV. Additional notes on section Longibrachiatum. Can. J. Bot. 69:2418–2420.
Bissett J. , 'Arevision of the genus Trichoderma. IV. Additional notes on section Longibrachiatum ' (1991 ) 69 Can. J. Bot. : 2418 -2420 .
Chaverri, P., Castlebury, L.A., Samuels, G.J., Geiser, D.M. 2003. Multilocus phylogenetic structure within the Trichoderma harzianum/Hypocrea lixii complex. Mol. Phylogenet. Evol. 23:302–313.
Geiser D.M. , 'Multilocus phylogenetic structure within the Trichoderma harzianum/Hypocrea lixii complex ' (2003 ) 23 Mol. Phylogenet. Evol. : 302 -313 .
Cumagun, C.J.R., Hockenhull, J., Lübeck, M. 2000. Characterization of Trichoderma isolates from Philippine rice fields by UP-PCR and rDNA-ITS1 analysis: Identification of UP-PCR markers. J. Phytopathol. 148:109–115.
Lübeck M. , 'Characterization of Trichoderma isolates from Philippine rice fields by UP-PCR and rDNA-ITS1 analysis: Identification of UP-PCR markers ' (2000 ) 148 J. Phytopathol. : 109 -115 .
Dodd, S.L., Crowhurst, R.N., Rodrigo, A.C., Samuels, G.J., Hill, R.A., Stewart, A. 2000. Examination of Trichoderma phylogenies derived from ribosomal DNA sequence data. Mycol. Res. 104:23–34.
Stewart A. , 'Examination of Trichoderma phylogenies derived from ribosomal DNA sequence data ' (2000 ) 104 Mycol. Res. : 23 -34 .
Druzhinina, I., Kopchinskiy, A., Komoñ, M., Bissett, J., Szakacs, G., Kubicek, C.P. 2005. An oligonucleotide barcode for species identification in Trichoderma and Hypocrea. Fungal Genet. Biol. 42:813–828.
Kubicek C.P. , 'An oligonucleotide barcode for species identification in Trichoderma and Hypocrea ' (2005 ) 42 Fungal Genet. Biol. : 813 -828 .
Druzhinina, I., Kubicek, C.P. 2005. Species concepts and biodiversity in Trichoderma and Hypocrea: from aggregate species to species clusters? J. Zhejiang Univ. Sci. 6:100–112.
Kubicek C.P. , 'Species concepts and biodiversity in Trichoderma and Hypocrea: from aggregate species to species clusters? ' (2005 ) 6 J. Zhejiang Univ. Sci. : 100 -112 .
Druzhinina, I.S., Kopchinskiy, A.G., Kubicek C.P. 2006. The first 100 Trichoderma species characterized by molecular data. Mycoscience 47:55–64.
Kubicek C.P. , 'The first 100 Trichoderma species characterized by molecular data ' (2006 ) 47 Mycoscience : 55 -64 .
Druzhinina, I.S., Kubicek, C.P., Komoñ-Zelazowska, M., Mulaw, T.B., Bissett, J. 2010. The Trichoderma harzianum demon: Complex speciation history resulting in coexistence of hypothetical biological species, recent agamospecies and numerous relict lineages. BMC Evol. Biol. 10:94.
Bissett J. , 'The Trichoderma harzianum demon: Complex speciation history resulting in coexistence of hypothetical biological species, recent agamospecies and numerous relict lineages ' (2010 ) 10 BMC Evol. Biol. : 94 -.
Gherbawy, Y., Druzhinina, I., Shaban, G.M., Wuczkowsky, M., Yaser, M., El-Naghy, M.A., Prillinger, H.J., Kubicek, C.P. 2004. Trichoderma populations from alkaline agricultural soil in the Nile valley, Egypt, consist of only two species. Mycol. Prog. 3:211–218.
Kubicek C.P. , 'Trichoderma populations from alkaline agricultural soil in the Nile valley, Egypt, consist of only two species ' (2004 ) 3 Mycol. Prog. : 211 -218 .
Gomez, I., Chet, I., Herrera-Estrella, A. 1997. Genetic diversity and vegetative compatibility among Trichoderma harzianum isolates. Mol. Gen. Genet. 256:127–135.
Herrera-Estrella A. , 'Genetic diversity and vegetative compatibility among Trichoderma harzianum isolates ' (1997 ) 256 Mol. Gen. Genet. : 127 -135 .
Harman, G.E., Howell, C.R., Viterbo, A., Chet, I., Lorito, M. 2004. Trichoderma species — opportunistic avirulent plant symbionts. Nature Rev. Microbiol. 2:43–56.
Lorito M. , 'Trichoderma species — opportunistic avirulent plant symbionts ' (2004 ) 2 Nature Rev. Microbiol. : 43 -56 .
Hatvani, L., Antal, Z., Manczinger, L., Szekeres, A., Druzhinina, I.S., Kubicek, C.P., Nagy, A., Nagy, E., Vágvölgyi, C., Kredics, L. 2007. Green mould diseases of Agaricus and Pleurotus are caused by related but phylogenetically different Trichoderma species. Phytopathology 97:532–537.
Kredics L. , 'Green mould diseases of Agaricus and Pleurotus are caused by related but phylogenetically different Trichoderma species ' (2007 ) 97 Phytopathology : 532 -537 .
Hermosa, M.R., Grondona, I., Iturriaga, E.A., Diaz-Minguez, J.M., Castro, C., Monte, E., Garcia-Acha, I. 2000. Molecular characterization of biocontrol isolates of Trichoderma. Appl. Environ. Microbiol. 66:1890–1898.
Garcia-Acha I. , 'Molecular characterization of biocontrol isolates of Trichoderma ' (2000 ) 66 Appl. Environ. Microbiol. : 1890 -1898 .
Hjeljord, L., Tronsmo, A. 1998. Trichoderma and Gliocladium in biological control: An overview. In: Harman, G.E., Kubicek, C.P. (eds), Trichoderma and Gliocladium. Enzymes, Biological Control and Commercial Applications, vol. II. Taylor & Francis, London, UK, pp. 131–151.
Tronsmo A. , '', in Trichoderma and Gliocladium. Enzymes, Biological Control and Commercial Applications, vol. II , (1998 ) -.
Howell, C.R. 2003. Mechanisms employed by Trichoderma species in the biological control of plant diseases: The history and evolution of current concepts. Plant Dis. 87:4–10.
Howell C.R. , 'Mechanisms employed by Trichoderma species in the biological control of plant diseases: The history and evolution of current concepts ' (2003 ) 87 Plant Dis. : 4 -10 .
Hoyos-Carvajal, L., Orduz, S., Bissett, J. 2009. Genetic and metabolic biodiversity of Trichoderma from Colombia and adjacent neotropic regions. Fungal Genet. Biol. 46:615–631.
Bissett J. , 'Genetic and metabolic biodiversity of Trichoderma from Colombia and adjacent neotropic regions ' (2009 ) 46 Fungal Genet. Biol. : 615 -631 .
Kindermann, J., El-Ayouti, Y., Samuels, G.J., Kubicek, C.P. 1998. Phylogeny of the genus Trichoderma based on sequence analysis of the internal transcribed spacer region 1 of the rDNA clade. Fungal Genet. Biol. 24:298–309.
Kubicek C.P. , 'Phylogeny of the genus Trichoderma based on sequence analysis of the internal transcribed spacer region 1 of the rDNA clade ' (1998 ) 24 Fungal Genet. Biol. : 298 -309 .
Klein, D., Eveleigh, E., 1998. Ecology of Trichoderma. In: Kubicek, C.P., Harman, G.E. (eds), Trichoderma and Gliocladium. Basic Biology, Taxonomy and Genetics. Taylor & Francis Ltd., London, UK, pp. 57–74.
Eveleigh E. , '', in Trichoderma and Gliocladium. Basic Biology, Taxonomy and Genetics , (1998 ) -.
Kredics, L., Antal, Z., Dóczi, I., Manczinger, L., Kevei, F., Nagy, E. 2003. Clinical importance of the genus Trichoderma. A review. Acta Microbiol. Immunol. Hung. 50:105–117.
Nagy E. , 'Clinical importance of the genus Trichoderma ' (2003 ) 50 A review. Acta Microbiol. Immunol. Hung. : 105 -117 .
Kubicek, C.P., Bissett, J., Druzhinina, I., Kullnig-Gradinger, C.M., Szakacs, G. 2003. Genetic and metabolic diversity of Trichoderma: A case study on South East Asian isolates. Fungal Genet. Biol. 38:310–319.
Szakacs G. , 'Genetic and metabolic diversity of Trichoderma: A case study on South East Asian isolates ' (2003 ) 38 Fungal Genet. Biol. : 310 -319 .
Kuhls, K., Lieckfeldt, E., Samuels, G.J., Meyer, W., Kubicek, C.P., Börner, T. 1997. Revision of Trichoderma section Longibrachiatum including related teleomorphs based on an analysis of ribosomal DNA internal transcribed spacer sequences. Mycologia 89:442–460.
Börner T. , 'Revision of Trichoderma section Longibrachiatum including related teleomorphs based on an analysis of ribosomal DNA internal transcribed spacer sequences ' (1997 ) 89 Mycologia : 442 -460 .
Kullnig, C., Szakacs, G., Kubicek, C.P. 2000. Molecular identification of Trichoderma species from Russia, Siberia and the Himalaya. Mycol. Res. 104:1117–1125.
Kubicek C.P. , 'Molecular identification of Trichoderma species from Russia, Siberia and the Himalaya ' (2000 ) 104 Mycol. Res. : 1117 -1125 .
Kullnig-Gradinger, C., Szakacs, G., Kubicek, C.P. 2002. Phylogeny and evolution of the genus Trichoderma: A multigene approach. Mycol. Res. 106:757–767.
Kubicek C.P. , 'Phylogeny and evolution of the genus Trichoderma: A multigene approach ' (2002 ) 106 Mycol. Res. : 757 -767 .
Lieckfeld, E., Seifert, K.A. 2000. An evaluation of the use of ITS sequences in the taxonomy of the Hypocreales. Stud. Mycol. 45:35–44.
Seifert K.A. , 'An evaluation of the use of ITS sequences in the taxonomy of the Hypocreales ' (2000 ) 45 Stud. Mycol. : 35 -44 .
Lieckfeldt, E., Kuhls, K., Muthumeenakshi, M. 1998. Molecular taxonomy of Trichoderma and Gliocladium and their teleomorphs. In: Kubicek, C.P., Harman, G.E. (eds), Trichoderma and Gliocladium, Vol. 1: Basic Biology, Taxonomy and Genetics. Taylor & Francis, London, UK, pp. 35–74.
Muthumeenakshi M. , '', in Trichoderma and Gliocladium, Vol. 1: Basic Biology, Taxonomy and Genetics , (1998 ) -.
Migheli, Q., Balmas, V., Komoñ-Zelazowska, M., Scherm, B., Fiori, S., Kopchinskiy, A.G., Kubicek, C.P., Druzhinina, I.S. 2009. Soils of a Mediterranean hot spot of biodiversity and endemism (Sardinia, Tyrrhenian Islands) are inhabited by pan-European, invasive species of Hypocrea/Trichoderma. Environ. Microbiol. 11:35–46.
Druzhinina I.S. , 'Soils of a Mediterranean hot spot of biodiversity and endemism (Sardinia, Tyrrhenian Islands) are inhabited by pan-European, invasive species of Hypocrea/Trichoderma ' (2009 ) 11 Environ. Microbiol. : 35 -46 .
Mostafa Kamal, M.D., Shahjahan, A.K.M. 1995. Trichoderma in rice field soils and their effect on Rhizoctonia solani. Bangladesh J. Bot. 24:75–79.
Shahjahan A.K.M. , 'Trichoderma in rice field soils and their effect on Rhizoctonia solani ' (1995 ) 24 Bangladesh J. Bot. : 75 -79 .
Mulaw, T.G., Kubicek, C.P., Druzhinina, I.S. 2010. The rhizosphere of Coffea arabica in its native highland forests of Ethiopia provides a niche for a distinguished diversity of Trichoderma. Diversity 2:527–549.
Druzhinina I.S. , 'The rhizosphere of Coffea arabica in its native highland forests of Ethiopia provides a niche for a distinguished diversity of Trichoderma ' (2010 ) 2 Diversity : 527 -549 .
Naeimi, S., Okhovvat, S.M., Javan-Nikkhah, M., Kredics, L., Khosravi, V. 2010. Biological control of Rhizoctonia solani AG1-1A, the causal agent of rice sheath blight with Trichoderma strains. Phytopathol. Mediterr. 49: 287–300.
Khosravi V. , 'Biological control of Rhizoctonia solani AG1-1A, the causal agent of rice sheath blight with Trichoderma strains ' (2010 ) 49 Phytopathol. Mediterr. : 287 -300 .
Nagamani, A., Mew, T.W. 1987. Trichoderma in Philippine rice field soils. Int. Rice Res. Newslett. 12:25.
Mew T.W. , 'Trichoderma in Philippine rice field soils ' (1987 ) 12 Int. Rice Res. Newslett. : 25 -.
Ospina-Giraldo, M.D., Royse, D.J., Chen, X., Romaine, C.P. 1999. Molecular phylogenetic analyses of biological control strains of Trichoderma harzianum and other biotypes of Trichoderma spp. associated with mushroom green mold. Phytopathology 89:308–313.
Romaine C.P. , 'Molecular phylogenetic analyses of biological control strains of Trichoderma harzianum and other biotypes of Trichoderma spp. associated with mushroom green mold ' (1999 ) 89 Phytopathology : 308 -313 .
Rifai, M.A. 1969. A revision of the genus Trichoderma. Mycol. Pap. 116:1–116.
Rifai M.A. , 'A revision of the genus Trichoderma ' (1969 ) 116 Mycol. Pap. : 1 -116 .
Sharma, K., Mishra, A.K., Misra, R.S. 2009. Morphological, biochemical and molecular characterization of Trichoderma harzianum isolates for their efficacy as biocontrol agents. J. Phytopathol. 157:51–56.
Misra R.S. , 'Morphological, biochemical and molecular characterization of Trichoderma harzianum isolates for their efficacy as biocontrol agents ' (2009 ) 157 J. Phytopathol. : 51 -56 .
White, T.J., Bruns, T., Lee, S., Taylor, J. 1990. Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In: Innes, M.H., Gelfand, D.H., Sninsky, J.J., White, T.J. (eds), PCR protocols. Academic Press, San Diego, CA, USA, pp. 315–322.
Taylor J. , '', in PCR protocols , (1990 ) -.
Wuczkowski, M., Druzhinina, I., Gherbawy, Y., Klug, B., Prillinger, H.J., Kubicek, C.P. 2003. Species pattern and genetic diversity of Trichoderma in a mid-European, primeval floodplain-forest. Microbiol. Res. 158:125–133.
Kubicek C.P. , 'Species pattern and genetic diversity of Trichoderma in a mid-European, primeval floodplain-forest ' (2003 ) 158 Microbiol. Res. : 125 -133 .