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Download Full Article in PDF Format Cryptogamie, Mycologie, 2017, 38 (2): 191-203 © 2017 Adac. Tous droits réservés Russula dinghuensis sp. nov. and R. subpallidirosea sp. nov., two new species from southern China supported by morphological and molecular evidence Jianbin ZHANG, Jingwei LI, Fang Li &Lihong QIU* State Key Lab of Biocontrol, School of Life Science, Sun Yat-sen University,Guangzhou 510275, China Abstract –Two new taxa of Russula from the Dinghu Mountain, Guangdong Province, China were described and illustrated based on both morphological data and phylogenetic analysis of the internal transcribed spacer sequences. Russula dinghuensis is characterized by the olive green pileus, acute and incurved margin, white and rarely forked lamellae, white spore print, globose to ellipsoid basidiospores with stocky and isolated warts, thick metachromatic pileipellis, and slender,furcated and septated terminal elements of pileipellis. Russula subpallidirosea is recognized by the pale pink to pale grayish-pink pileus, white and forked lamellae, white spore print, subglobose to ellipsoid basidiospores with the isolated, subcylindrical to conical warts, the metachromatic pileipellis, and the short, furcated and septate terminal elements of pileipellis. Both molecular and morphological analyses consistently confirm that these two new taxa are placed into Russula subg. Heterophyllidia subsection Cyanoxanthinae. The morphological differences among these two novel species and the closely related taxa are discussed. Cyanoxanthinae /Dinghu Mountain /ITS /phylogeny /taxonomy INTRODUCTION Russula Pers. (Russulaceae, Basidiomycota), erected by Persoon in 1796, is considered one of the most abundant and widely distributed ectomycorrhizal agaric genera in the world (Buyck et al. 2008, 2015; Buyck &Horak 1999). Species of Russula are recognized by the combination of the following characteristics: mostly conspicuous fruit bodies with colorful pileus, amyloid spore ornamentation, presence of gloeocystidia, brittle context with abundant sphaerocytes, absence of latex, and hyphae lacking clamp connections (Romagnesi 1967; Sarnari 1998; Singer 1986). The genus Russula consists of more than 750 species and ca. 160 of these have been reported from China (Kirk et al. 2008; Song et al. 2007). Compared with the long and rich taxonomic history of Russula in Europe (Romagnesi 1967; Sarnari 1998), new Russula taxa from Asian countries are rarely reported. Up to now,only 22 species and 3varieties were originallydescribed from China (Bi &Li1986; Chiu 1945; Li et al. 2012; Li et al. 2011; G.J. Li et al. 2015; Li et al. 2013; Li et al. 2013; Y.K. Li et al. 2015; Singer 1935; Song et al. 2007; Wang et al. 2009; Wen& *Corresponding author: [email protected] doi/10.7872/crym/v38.iss2.2017.191 192 J. Zhang et al. Ying 2001; Ying 1983; 1989; Zang &Yuan1999; Zhao et al. 2015). Because of the lack of systematic studies on Russula in China, it is believed that many Chinese Russula species are misplaced under European or American species names, which probably led to underestimates of the real diversity of Russula in China (Li et al. 2012; Li et al. 2011). The Dinghu Mountain, located in Zhaoqing city,China, is considered a region highly diverse in macrofungi (Bi et al. 1994; Zheng et al. 1985). More than 51 species of Russula have been recorded from this subtropical region. During recent surveys of Russula in the area, several specimens representing two Russula taxa were collected. Morphological study together with molecular analyses (ITS) showed they represent two novel species of Russula subg. Heterophyllidia (Romagnesi) Sarnari 1998. MATERIALS AND METHODS Sampling and morphological studies Random collection was conducted in September 2013 and May 2015 in Dinghushan Biosphere Reserve, Guangdong Province, China (112°33′ E, 23°10′ N). Specimens were dried at 50-60°C and deposited in the Herbarium of Guangdong Institute of Microbiology,China. Fresh basidiomes were photographed using a Canon IXUS 220 hs digital camera, and macroscopic characteristics of the intact fresh fruit bodies were recorded under daylight conditions in the field. HTML Color Codes (http://www.htmlcolorcode.org/) were used to describe the color of the specimens. For the microscopic characters, tissue sections were immersed in 5% KOH and then stained with 1% aqueous Congo red solution. All tissues were also examined in Cresyl blue to verify presence of ortho- or metachromatic reactions as explained in Buyck (1989). Sulfovanillin (SV) was used to test for reactions of cystidia. Micromorphological features including the pileipellis, basidia, basidiospores, cystidia, and stipitipellis, were observed and photographed using aNIKON E200 microscope equipped with an NIKON E4500 camera. Basidiospores were observed in Melzer’s reagent and measured in side view,excluding ornamentation and apiculus which were observed by SEM. The abbreviation [x/y/z]indicates x basidiospores measured from y fruit bodies of z specimens. In the basidiospore dimension notation “(a-) b-m-c (-d)”, b-c is the range including 95% of the measured values for length or width, with a and d corresponding to the extremes of all measurements, and “m” mean value. Qindicates length/width ratio of basidiospores, with Qm the average Q of all basidiospores ± standard deviation. DNA extraction, PCR and sequencing DNA was extracted from afresh fruitbody using the method described by Xu et al. (2010). For PCR, sequences of the internal transcribed spacer (ITS) region of nuclear ribosomal DNA were amplified with the primer pair ITS1F/ITS4 (Gardes &Bruns 1993; White et al. 1990). The protocols for PCR amplification consisted of a5min activation at 94°C, followed by 32 cycles of 30 sat94°C, 30 sat52°C and 1min at 72°C, and a final 12 min extension at 72°C. Direct sequencing of PCR Russula dinghuensis sp. nov.and R. subpallidirosea sp. nov.193 products was performed using the PCR primers as sequencing primers. Sanger dideoxy sequencing were performed with ABI 3730 DNA analyzer (IGE, Guangzhou, China). If direct sequencing of PCR products failed, it was subcloned with the PMD18-T vector (Takara Bio, Japan) before sequencing. The generatedsequences were then submitted in GenBank. PHYLOGENETIC ANALYSES BLAST query in GenBank indicated that sequences of the two novel species are close to R. cyanoxantha (Schaeff.) Fr., aspecies in Russula subgenus Heterophyllidia subsection Cyanoxanthinae.Toestablish their taxonomic position within subgenus Heterophyllidia,28sequences were added to the ITS dataset representing 19 Russula from five subsections of subg. Heterophyllidia (Table 1), according to the classification of Sarnari (1998). Based on the molecular phylogenetic results of Miller &Buyck (2002), Albatrellus flettii Morse ex Pouzar and Gloeocystidiellum aculeatum Sheng H. Wu were chosen as outgroups. Sequences were alignedwith Clustal Xand manually modulated when necessary.Some ambiguously aligned terminal regions were excluded. The final aligned result was submitted to TreeBASE (ID 18975). For phylogenetic analyses, both Neighbor-joining (NJ) and Bayesian inference (BI) algorithms were employed. NJ analysis of the phylogenetic relationships among the taxa was performed using MEGA 5.05 with the Kimura-2- parameter model and gaps in alignment were treated as missing data. Bootstrap analysis was conducted with 1000 replicates. Bootstrap value (BV) exceeding 70% was considered as significantly supported. Bayesian inference analyses were performed with MrBayes v3.2.5 using the Markov chain Monte Carlo method under the GTR +I+Gmodel. Analyses were run with 4chains of 1,000,000 generations, and trees were sampled every 100th generation. Bayesian posterior probabilities(PP) values were obtained from the 50% majority-rule consensus trees. RESULTS Phylogenetic analysis Atotal of 41 ITS sequences (average length 682 bp, see Table 1) representing 23 taxa including two outgroups, were analyzed by NJ and BI analysis. In the aligned ITS matrix, 263 characters were constant, 390 characters were variable, and 295 characters were parsimony-informative. Both the NJ and BI analyses produced similar tree topologies and only the tree inferred from NJ analysis is shown (Fig. 1). The molecular phylogenetic analysis showed that Russula subg. Heterophyllidia was astrongly supported (BV 87%, PP 0.68), monophyletic group. And five subsections of this subgenus also formed well-supported, distinct clades. The overall structure of tree was in line with the result of Dutta et al. (2015) and Zhao et al. (2015). The two new taxa formed astrongly supported clades (BV 100%, PP 1.00) that nested well within subsection Cyanoxanthinae and were distinct from each other 194 J. Zhang et al. Table 1. The rDNA ITS sequences of Russula subg. Heterophyllidia used in this study Taxon Voucher Locality GenBank Accession No. Albatrellus flettii S. MillerWF3 USA AY061738 Gloeocystidiellum aculeatum GB-2647 China AY061739 Russula aeruginea DG88 UK JQ888195 R. aeruginea Nl1292 Germany UDB000341 R. alboareolata SUT-1 Thailand AF345247 R. anatina 13216 Italy JF908698 R. atroaeruginea HKAS53626 China JX391967 R. aurata S. Miller 6001 Europe AY061659 R. crustosa BB2004-208, PC Europe EU598194 R. cyanoxantha SM/BB 5, PC Europe AY061669 R. cyanoxantha HKAS 78376 China KF002766 R. cyanoxantha HKAS 78385 China KF002775 R. dinghuensis GDGM 45244 China KU863579 R. dinghuensis GDGM45243 China KU863580 R. dinghuensis K15052704-3 China KU863581 R. grisea Watling 27098, EEurope AY061679 R. heterophylla Buyck 99.803, PC Europe AY061681 R. ilicis Sarnari
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