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Revision of Lignicolous Tubeufiaceae Based on Morphological Reexamination and Phylogenetic Analysis

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Revision of Lignicolous Tubeufiaceae Based on Morphological Reexamination and Phylogenetic Analysis Fungal Diversity DOI 10.1007/s13225-011-0147-4 Revision of lignicolous Tubeufiaceae based on morphological reexamination and phylogenetic analysis Saranyaphat Boonmee & Ying Zhang & Putarak Chomnunti & Ekachai Chukeatirote & Clement K. M. Tsui & Ali H. Bahkali & Kevin D. Hyde Received: 18 October 2011 /Accepted: 23 October 2011 # Kevin D. Hyde 2011 Abstract In this paper we revisit the family Tubeufiaceae fia, which are retained in Tubeufiaceae; however, we were with notes on genera that we have re-examined where unable to locate the types of these genera during the time possible. Generic type specimens of Acanthophiobolus, frame of this study. Allonecte is excluded from the Kamalomyces, Podonectria, Thaxteriella and Thaxteriel- Tubeufiaceae, as the ascospores are fusiform-ellipsoidal, lopsis were re-examined, described and illustrated and grey-brown and 1-septate and the asci are cylindrical, all of shown to belong to Tubeufiaceae. Notes are provided on which are features more typical of Pleosporaceae, where it Acanthostigma, Chaetosphaerulina, Thaxterina and Tubeu- is transferred. Byssocallis has yellow to orange ascomata and clavate ascospores which is atypical of Tubeufiaceae. Thus its taxonomic status needs to be reevaluated. Lentendraeopsis has an endophytic habit, cylindro-clavate Electronic supplementary material The online version of this article asci and two-celled ascospores more typical of Pleospor- (doi:10.1007/s13225-011-0147-4) contains supplementary material, ales, where it is transferred. Taphrophila has small which is available to authorized users. ascomata, a thin peridium, branching setae around the apex : : : : S. Boonmee Y. Zhang P. Chomnunti E. Chukeatirote of the ascomata, clavate to saccate asci and lacks K. D. Hyde pseudoparaphyses. These are features atypical of the Institute of Excellence in Fungal Research, Tubeufiaceae, and Taphrophila should be placed in the Mae Fah Luang University, Chiang Rai 57100, Thailand Dothideomycetes incertae cedis. Twelve new collections of : : : : Tubeufiaceae from Thailand were isolated, and their DNA S. Boonmee Y. Zhang P. Chomnunti E. Chukeatirote was extracted. The sequence data of LSU, SSU and ITS K. D. Hyde rDNA were amplified and analyzed using parsimony and School of Science, Mae Fah Luang University, Chiang Rai 57100, Thailand likelihood methods. The results of phylogenetic analysis : was used to establish the inter-generic relationships in Y. Zhang K. D. Hyde Tubeufiaceae. Thaxteriellopsis lignicola, epitypified in this Mushroom Research Centre, investigation, is a sister taxon in the family Tubeufiaceae 128 Moo3, Bahn Pa Dheng, T. Pa Pae, A. Mae Taeng, Chiang Mai 50150, Thailand based on phylogenetic analysis of rRNA sequence data. Chlamydotubeufia is introduced as a new genus based on C. K. M. Tsui the production of dictyochlamydosporous anamorphs, Department of Forest Sciences, including two new species. Three new species, one each The University of British Columbia, Vancouver, BC V6T 1Z4, Canada in Acanthostigma, Tubeufia and Thaxteriella are also : described and illustrated. The phylogenetic placement of A. H. Bahkali K. D. Hyde (*) these genera is also discussed. College of Science, Botany and Microbiology Department, King Saud University, . Riyadh 1145, Saudi Arabia Keywords Anamorph Aquaphila Dothideomycetes e-mail: [email protected] Helicosporous . Molecular phylogeny. Woody litter fungi Fungal Diversity Introduction In this study, we have examined the generic types or authentic specimens of several tubuefiaceous genera that The family Tubeufiaceae sensu Barr (1979) included the we have been able to locate. We have also surveyed woody type genus Tubeufia and four representative genera litter in forests throughout northern Thailand and obtained (Letendraea, Melioliphila, Podonectria, Thaxteriella)and several new collections of these fungi, which have been was considered to be pleosporaceous. The representatives described, isolated and sequenced. We compare the se- were saprobic, hyperparasitic or hypersaprobic on asco- quence data of these new collections to those from the mycetes and scale insects, and characterized by brightly generic types and to provide insights on the inter-generic pigmented, fleshy, superficial ascomata, bitunicate asci, and species relationships in Tubeufiaceae. One new genus and mostly hyaline to pale brownish, narrowly-elongate, and five new species encountered during the course of our obovoid or oblong septate ascospores. Barr (1980) field work are also described. included ten genera while Rossman (1987)studiedthe family in detail and included 12 genera (Table 1). Several other genera have since been added and the family now Material and methods encompasses 23 genera (Lumbsch and Huhndorf 2010)or22 genera (Kirk et al. 2001). Most taxa of Tubeufiaceae are Sample collection and specimen examination commonly found on woody litter, although some species can also be found on leaf litter or even decaying cloth and some Decaying wood was randomly collected from forests at are associated with other fungi or scale insects (Barr 1980; various sampling sites in northern Thailand (i.e. Chiang Rossman 1987; Kodsueb et al. 2006; Promputtha and Miller Mai, Chiang Rai, Lam Pang and Pha Yao). Material was 2010; Sánchez and Bianchinotti 2010). The anamorphs of examined under a Motic SMZ 168 Series microscope. Tubeufiaceae have been well-studied and are mostly related Micromorphological structures were photographed using a to helicosporous taxa such as Helicoma, Helicomyces and Nikon ECLIPSE 80i compound microscope with a Canon Helicosporium (Tsui et al. 2006, 2007;Hydeetal.2011). 450D digital camera, and measurements were made using Table 1 Various treatments of Tubeufiaceae Barr, 1979 Barr 1980 Rossman 1987 Kirk et al. 2001 Lumbsch and Huhndorf 2010 This paper Letendraea Allonectria Allonectria Acanthophiobolus Acanthostigma Acanthostigma Melioliphila Boerlagiomyces Boerlagiomyces Acanthostigmella Acanthophiobolus Acanthophiobolus Podonectria Byssocallis Byssocallis Allonecte Acanthostigmella ?Acanthostigmella Rebentischia Letendraea Letendraea Amphinectria Allonecte ?Amphinectria Thaxteriella Melioliphila Malacaria Boerlagiomyces ?Amphinectria Aquaphila Tubeufia Paranectriella Melioliphila Borinquenia Boerlagiomyces ?Boerlagiomyces Podonectria Paranectriella Byssocallis Byssocallis ?Chaetocrea Puttemansia Podonectria Chaetocrea Chaetocrea Chaetosphaerulina Rebentischia Puttemansia Glaxoa Chaetosphaerulina ?Glaxoa Tubeufia Rebentischia Letendraea Glaxoa Kamalomyces Tubeufia Letendraeopsis Letendraeopsis ?Malacaria Uredinophila Malacaria Malacaria ?Melioliphila Melioliphila Melioliphila ?Paranectriella Paranectriella Paranectriella Podonectria Podonectria Podonectria ?Puttemansia Puttemansia Puttemansia Rebentischia Rebentischia Rebentischia Thaxteriella Taphrophila Taphrophila Thaxteriellopsis Thaxterina Thaxteriella Tubeufia Tubeufia ?Thaxteriellopsis ?Uredinophila Uredinophila Thaxterina Tubeufia Uredinophila Fungal Diversity the Tarosoft (R) Image Frame Work program. Images used analyzed using PAUP* v. 4.0b10 (Swofford 2002). Am- for figures were processed with Adobe Photoshop CS3 biguous regions in the alignments were excluded from the Extended version 10.0 software (Adobe Systems Inc., The phylogenetic analyses. Maximum Parsimony (MP) was United States). The type herbarium material is deposited in performed with stepwise additions of sequences using the herbarium of Mae Fah Luang University (MFLU), 1,000 random replicates and tree-bisection-reconnection Chiang Rai, Thailand, and cultures are deposited in the Mae (TBR) branch-swapping algorithm, with MAXTREES Fah Luang University Culture Collection (MFLUCC), setting at 1000. The parsimony tree scores including tree BIOTEC Culture Collection (BCC) and IFRD culture length (TL), consistency index (CI), retention index (RI), collections, International Fungal Research & Development rescaled consistency index (RC) and homoplasy index (HI) Centre, Kunming, China, the latter under material transfer were also calculated. Bootstrap support for the branches agreement No. 4/2010 (MTA). was estimated based on 500 MP replicates with a single sequence addition replicates in each bootstrap replicate. Isolation of fungi Models of nucleotide substitution for each gene were determined using MrModeltest 2.3 (Nylander 2004). GTR Single spore isolates were cultured on water agar (WA) or + I + G evolutional model were selected for likelihood malt extract agar (MEA, Difco Laboratories, Detroit, analysis using MrBayes v. 3.1.2 (Ronquist and Huelsenbeck Michigan, USA). Germinating spores were transferred to 2003). The Markov Chain Monte Carlo (MCMC; Rannala MEA media and incubated at room temperature for 7 days. and Yang 1996) algorithm was used to estimate posterior Isolates were grown for 1–2 months when morphological probabilities (PP). Six MCMC chains were run from a characters in culture, such as the extent of the mycelium, random starting tree for 1,000,000 generations and trees colour, shape and texture were recorded. These cultures sampled every 100 generations. The first 2,000 trees were were also used for DNA extraction and sequencing. discarded as burn-in prior to convergence of the chains. The remaining 10,001 trees based on 1,000 replicates were used DNA extraction, PCR amplification and sequencing to construct 50% majority rule consensus tree and to calculate posterior probabilities were determined more than Fungal mycelium was scraped from colonies growing on 95% PP given each on branches.
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