Research Article
Research Article
The genus Castanediella
expand article infoChuan-Gen Lin§, Darbhe J. Bhat|, Jian-Kui Liu#, Kevin D. Hyde, Yong Wang§
‡ Mae Fah Luang University, Chiang Rai, Thailand
§ Guizhou University, Guiyang, China
| Azad Housing Society, Goa Velha, India
¶ Goa University, Goa, India
# University of Electronic Science and Technology of China, Chengdu, China
Open Access


Two new species, Castanediella brevis and C. monoseptata, are described, illustrated and compared with other Castanediella taxa. Evidence for the new species is provided by morphological comparison and sequence data analyses. Castanediella brevis can be distinguished from other Castanediella species by the short hyaline conidiophores and fusiform, aseptate hyaline conidia, while C. monoseptata differs from other Castanediella species by its unbranched conidiophores and fusiform, curved, 0–1-sepatate, hyaline conidia. Phylogenetic analysis of combined ITS and LSU sequence data was carried out to determine the phylogenetic placement of the species. A synopsis of hitherto described Castanediella species is provided. In addition, Castanediella is also compared with morphologically similar-looking genera such as Idriella, Idriellopsis, Microdochium, Neoidriella, Paraidriella and Selenodriella.


new taxa, Castanediellaceae, hyphomycetes, phylogeny, Sordariomycetes


Hernández-Restrepo et al. (2017) introduced the family Castanediellaceae for the genus Castanediella within Xylariales and it was consolidated in recent study by Wijayawardene et al. (2018). The asexual morphs in Castanediellaceae are hyphomycetous and characterized by macronematous, mononematous or sporodochial, branched, brown to pale brown conidiophores, with monoblastic or polyblastic, sympodial, discrete, cylindrical to lageniform, hyaline to subhyaline conidiogenous cells, that produce unicellular or transversely septate, cylindrical, fusiform or lunate, hyaline conidia (Hernández-Restrepo et al. 2017).

The genus Castanediella was established by Crous et al. (2015) to accommodate C. acaciae, C. cagnizarii and C. ramosa within Xylariales genera incertae sedis. The genus contains twelve species (Costa et al. 2018; Wanasinghe et al. 2018), each characterized by branched, hyaline to pale brown conidiophores, holoblastic, sympodial conidiogenous cells and falcate, cylindrical or fusiform, 0–3-sepate, hyaline conidia (Crous et al. 2015; Costa et al. 2018).

During a survey of hyphomycetes in Thailand, two hyaline-spored hyphomycetes were collected. They were shown to belong to the genus Castanediella based on morphology and phylogeny analyses of ITS and LSU sequence data. The new species C. brevis and C. monoseptata are introduced.

Materials and methods

Collection and isolation of fungi

Dead leaves from a variety of plants in two forests (Lampang province and Chiang Mai province) were collected in 2016 in Thailand. Samples were taken to the laboratory in Zip-lock plastic bags for examination. The specimens were incubated in sterile moist chambers and examined using a Motic SMZ 168 series microscope. Fungi were removed with a needle and placed in a drop of distilled water on a slide for morphological study. Photomicrographs of fungal structures were captured with a Canon 600D digital camera attached to a Nikon ECLIPSE Ni compound microscope. All measurements were made by the Tarosoft (R) Image FrameWork program. Photo-plates were made with Adobe Photoshop CS3 (Adobe Systems, USA). Isolation of the fungi on to potato dextrose agar (PDA) was performed by the single spore isolation method (Chomnunti et al. 2014). Dried material was deposited in the Herbarium of Mae Fah Luang University (MFLU), Chiang Rai, Thailand and herbarium of Kunming Institute of Botany, Chinese Academy of Sciences (HKAS), Kunming, China. Cultures were deposited at Mae Fah Luang University Culture Collection (MFLUCC), Chiang Rai, Thailand and Kunming Institute of Botany, Chinese Academy of Sciences (KUMCC), Kunming, China. FacesofFungi and Index Fungorum numbers were registered (Jayasiri et al. 2015; Index Fungorum 2018).

DNA extraction, PCR amplification and sequencing

Genomic DNA was extracted from fungal mycelium grown on PDA or malt extract agar (MEA) at room temperature using the Fungal gDNA Kit (BioMIGA, USA) according to the manufacturer’s instructions. The internal transcribed spacer region of ribosomal DNA (ITS) and large subunit nuclear ribosomal DNA (LSU) genes were amplified via polymerase chain reaction (PCR) using the following primers: ITS5 and ITS4 (White et al. 1990) for ITS, and LR0R and LR5 (Vilgalys and Hester 1990) for LSU. The PCR products were sequenced with the same primers. The PCR amplification was performed in a 25 μL reaction volume containing 12.5 μL of 2 × Power Taq PCR MasterMix (a premix and ready to use solution, including 0.1 Units/μl Taq DNA Polymerase, 500 μM dNTP Mixture each [dATP, dCTP, dGTP, dTTP], 20 mM Tris-HCl pH 8.3, 100 Mm KCl, 3 mM MgCl2, stabilizer and enhancer), 1 μL of each primer (10 μM), 1 μL genomic DNA extract and 9.5 μL deionised water. The PCR thermal cycle program of ITS and LSU were followed as: initially 94 °C for 3 min., followed by 35 cycles of denaturation at 94 °C for 30 s, annealing at 55 °C for 50 s, elongation at 72 °C for 1 min., and final extension at 72 °C for 10 min.

Phylogenetic analyses

Original sequences were checked using BioEdit version (Hall 1999), and most reference sequences were originated from previous publications. The remaining homogenous sequences were obtained by BLAST searches (Altschul et al. 1990) from GenBank. All sequences used in this study are listed in Table 1. Alignments for each locus were done in MAFFT v7.307 online version (Katoh and Standley 2016) and manually verified in MEGA 6.06 (Tamura et al. 2013). After alignment, the concatenation of different genes was done in SequenceMatrix 1.8 (Vaidya et al. 2011). The interleaved NEXUS files for Bayesian inference analyses were formatted with AliView v1.19-beta1k (Larsson 2014). Maximum parsimony (MP), maximum likelihood (ML) and Bayesian inference (BI) were used for phylogenetic analyses.

Table 1.

GenBank accession numbers of isolates included in this study.

Taxa Isolatea ITS LSU
Castanediella acaciae CPC 24869, CBS 139896 NR_137985 KR476763
Castanediella brevis KUMCC 18-0132 MH806361 MH806358
Castanediella cagnizarii MUCL 41095 KC775732 KC775707
Castanediella cagnizarii CBS 101043 KP859051 KP858988
Castanediella cagnizarii CBS 542.96 KP859054 KP858991
Castanediella camelliae CNUFC-DLHBS5-1 MF926620 MF926614
Castanediella camelliae CNUFC-DLHBS5-2 MF926621 MF926615
Castanediella communis CPC 27631 KY173393
Castanediella couratarii CBS 579.71 NR_145250 KP858987
Castanediella eucalypti CPC 24746, CBS 139897 NR_137981 KR476758
Castanediella eucalypticola CPC 26539 NR_145254 KX228317
Castanediella eucalyptigena CBS 143178, CPC 32055 MG386036 MG386089
Castanediella hyalopenicillata CPC 25873 KX306751 KX306780
Castanediella malaysiana CPC 24918 NR_154810 KX306781
Castanediella monoseptata KUMCC 18-0133 MH806360 MH806357
Castanediella ramosa MUCL 39857 KC775736 KC775711
Subsessila turbinata MFLUCC 15-0831 KX762288 KX762289

The best models of evolution for each gene region were determined using Akaike information criterion (AIC) as implemented in MrModeltest v2 (Nylander 2004). The analyses’ results showed that the models GTR+I and GTR+I+G were the best ones for LSU and ITS sequence data, respectively.

MP analyses were performed in PAUP*4.0b10 (Swofford 2002) following Liu et al. (2016).

ML analyses were carried out in raxmlGUI v 1.5b1 (Silvestro and Michalak 2012) with RAxML v8.2.10 (Stamatakis 2014), using the ML + rapid bootstrap setting and the GTRGAMMAI (viz., GTR + GAMMA + I) substitution model with 1000 bootstrap replicates.

For BI analysis, Posterior probabilities (PP) (Rannala and Yang 1996; Zhaxybayeva and Gogarten 2002) were determined by Markov Chain Monte Carlo sampling (BMCMC) in MrBayes v 3.2.6 (Ronquist et al. 2012). For the combined dataset, the models were set to nst = 6 and rates = propinv for LSU and nst = 6 and rates = invgamma for ITS. Two independent analyses of two parallel runs and six simultaneous Markov chains were run for 1,000,000 generations, trees were sampled every 100th generation and the temperature value of the heated chains was set at 0.15. The first 25% sampled trees of each run were discarded as “burn-in”, and the remaining trees were used for calculating posterior probabilities (PP) in the majority rule consensus tree with the sumt command in MrBayes.

Phylogenetic trees were drawn with TreeView 1.6.6 (Page 1996).


Molecular phylogeny

The aligned sequence matrix comprises LSU and ITS sequence data for 16 taxa (ingroup) and one outgroup taxon with a total of 1438 characters after alignment including the gaps, of which 120 were parsimony informative, 77 parsimony-uninformative, and 1241 characters constant. The dataset consists of thirteen species within the genus. The tree was rooted with Subsessila turbinata (MFLUCC 15-0831). Maximum parsimony analysis resulted in two trees with TL = 391, CI = 0.657, RI = 0.642, RC = 0.422, HI = 0.343. For the Bayesian analysis, two parallel runs with six chains were run for 1,000,000 generations and trees were sampled every 100th generation, resulting in 20002 trees from two runs of which 15002 trees were used to calculate the posterior probabilities (each run resulted in 10001 trees of which 7501 trees were sampled).The MP and ML (lnL = -4041.301739) analyses based on combined LSU and ITS sequence data provided similar tree topologies, and the result of MP analysis is shown in Fig. 1.

Figure 1. 

Phylogenetic tree generated from MP analysis based on combined LSU and ITS sequence data for the genus Castanediella. Bootstrap support values for maximum parsimony (MP, first set) and maximum likelihood (ML, second set) greater than 50% are indicated above or below the nodes. Ex-type strains are in bold, the new isolates are in red. The tree is rooted with Subsessila turbinata (MFLUCC 15-0831).

The novelty of the species, Castanediella brevis and C. monoseptata, described in this study are supported by sequence data analyses as belonging to the genus Castanediella, but with low bootstrap support values. Isolates of Castanediella brevis and C. monoseptata formed separate clades in the phylogenetic inference, respectively. Castanediella brevis is sister to C. malaysiana and C. ramosa, while C. monoseptata shows close phylogenetic relationship to C. couratarii and C. malaysiana. Both the new taxa can be recognized as phylogenetically distinct species and are clearly novel based on the recommendations for molecular data (Jeewon and Hyde 2016).

MP, ML and BI were used for phylogenetic analyses in this study. The tree topologies of MP and ML resulted from the combined LSU and ITS sequence data are similar, but most of the nodes are in low bootstrap support (Fig. 1). Polytomy structure was formed in the BI tree generated from the combined LSU and ITS sequence data. More sequence data, especially the protein-coding genes, e.g. TEF1-α, RPB2, β-tubulin, are required in the future study of the genus Castanediella.


Castanediella brevis s C.G. Lin & K.D. Hyde, sp. nov.

MycoBank No: MB828879
Facesoffungi number: FoF04929
Figure 2


THAILAND. Lampang: Amphoe Mueang Pan, Tambon Chae Son, on decaying leaves, 24 September 2016, Chuangen Lin, LCG 10-1 (MFLU 18-1695, holotype; HKAS 102198, isotype), ex-type living cultures KUMCC 18-0132.

GenBank number

ITS: MH806361, LSU:MH806358


In reference to the short conidiophores.

Saprobic on plant host. Asexual morph: Colonies on substrate effuse, white. Mycelium partly superficial, composed of septate, branched, smooth, hyaline to subhyaline hyphae. Conidiophores macronematous, mononematous, solitary, erect, unbranched, straight or flexuous, short, 0–1-septate, hyaline, subcylindrical, ampulliform, smooth, often reduced to conidiogenous cells. Conidiogenous cells holoblastic, polyblastic, sympodial, integrated, terminal, subcylindrical, ampulliform, hyaline, denticulate, with 2–4 tiny protuberant denticles, 3–14 × 1.5–5.5 μm. Conidia solitary, dry, acropleurogenous, smooth, fusiform, curved, aseptate, hyaline, 12.5–21.7 × 1.2–3 μm (av. 16.95 × 2.2 μm, n = 60). Sexual morph: Undetermined.

Culture characteristics: Conidia germinating on PDA within 24 h. Colonies on PDA effuse, greyish white to dark from above and below, reaching a diam. of 5–7 cm in 30 days at 25 °C.


Based on a megablast search of the NCBI nucleotide database using the ITS sequence of the ex-type culture, the highest similarities found were with Castanediella malaysiana (GenBank NR_154810; identities = 526/537(98%), gaps = 1/537(0%)) and C. couratarii (GenBank KX960789; identities = 521/538(97%), gaps = 3/538(0%)). Castanediella brevis differs from these two species by its conidiophore morphology. Castanediella couratarii has pale brown conidiophores and longer conidiogenous cells (10.5–37 × 2–3.5 μm) whereas C. malysiana has pale brown and longer conidiophores (76–157 × 2.5–3 μm).

Among the species that produce more or less falcate and aseptate conidia, Castanediella communis, C. eucalypti, C. eucalypticola and C. eucalyptigena are most similar to C. brevis. However, Castanediella brevis differs from these species by its short, unbranched and 0–1-septate conidiophores.

Figure 2. 

Castanediella brevis (MFLU 18-1695, holotype) a host material b conidiophores on the host surface c–g conidiophores, conidiogenous cells with conidia h conidia. Scale bars: 10 μm (c–g), 5 μm (h).

Castanediella monoseptata C.G. Lin & K.D. Hyde, sp. nov.

MycoBank No: MB828881
Facesoffungi number: FoF04930
Figure 3


THAILAND. Chiang Mai: on decaying leaves, 24 August 2016, Chuangen Lin, MRC 3-1 (MFLU 18-1696, holotype; HKAS 102199, isotype), ex-type living cultures KUMCC 18-0133.

GenBank number

ITS: MH806360, LSU: MH806357


In reference to the 0–1-septate conidia

Saprobic on plant host. Asexual morph: Colonies on substrate effuse, white. Mycelium partly superficial, composed of septate, branched, hyaline to subhyaline, smooth hyphae. Conidiophores macronematous, mononematous, solitary, erect, unbranched, straight or flexuous, septate, hyaline, subcylindrical, smooth, 8–29 × 2–4 μm. Conidiogenous cells polyblastic, integrated, sympodial, subcylindrical, hyaline, with several scars. Conidia solitary, dry, acropleurogenous, smooth, fusiform, curved, 0–1-sepatate, hyaline, 15.4–25.8 × 1.5–2.3 μm (av. 23.03 × 1.98 μm, n = 45). Sexual morph: Undetermined.

Culture characteristics: Conidia germinating on PDA within 24 h. Colonies on PDA effuse, grayish white to dark from above and below, reaching a diam. of 5–7 cm in 30 days at 25 °C.


A megablast search of the NCBI nucleotide database using the ITS sequence of the ex-type culture showed the highest similarities with uncultured Sordariales fungi (GenBank GQ268569; identities = 518/539(96%), gaps = 3/539(0%)) and Castanediella couratarii (GenBank KX960789; identities = 516/540(96%), gaps = 4/540(0%)).

Five Castanediella species, C. cagnizarii, C. diversispora, C. hyalopenicillata, C. malaysiana and C. ramosa, were reported to produce 1-septate conidia. Castanediella monoseptata can be distinguished from these species by its unbranched conidiophores and falcate and 15.4–25.8 × 1.5–2.3 μm conidia. Castanediella monoseptata is phylogenetically closely related to C. couratarii and C. ramosa, but differs from both species by its conidial morphology. Castanediella couratarii has shorter conidia (9.5–19 × 2–3 μm) are aseptate and C. ramosa has larger conidia (26–44 × 2–3 μm) that are 0–3-septate.

Figure 3. 

Castanediella monoseptata (MFLU 18-1696, holotype) a host material b conidiophores on the host surface c–f conidiophores, conidiogenous cells with conidia g–l conidia. Scale bars: 10 μm (c, d), 5 μm (e–l).


In this study, two new Castanediella species, C. brevis and C. monoseptata, were identified from decaying leaves in Thailand and a synopsis of hitherto described Castanediella species is provided (Table 2).

Table 2.

Synopsis of Castanediella species.

Taxa Conidiophores Conidiogenous cells Conidia
Shape Size (μm) Septa Colour
C. acaciae Subcylindrical, medium brown, 40–80 × 2–3 μm. Polyblastic, ampulliform, pale brown, 10–15 × 2–3 μm. Falcate with subobtuse ends (8–)10–11(–12) × 1.5(–2) 0 Hyaline
C. brevis Subcylindrical, ampulliform, hyaline, often reduced to conidiogenous cells Polyblastic, cylindrical, hyaline, 3–14 × 1.5–5.5 μm Fusiform, curved 12.5–21.7 × 1.2–3.0 0 Hyaline
C. cagnizarii Cylindrical, brown at the base, subhyaline towards the apex, up to 45 μm long. Polyblastic, sympodial, subhyaline, 5–22 × 3–4 μm. Cylindrical to fusiform, curved at the ends Two sizes, 10–15 × 2 or 20–26 × 2 Hyaline
C. camelliae Conidiophores reduced to conidiogenous cell. Cylindrical, ampulliform, globose to subglobose, or irregularly-shaped, 5.5–20.5 × 2–4.5 μm. Straight to slightly curved, sometimes swollen in the middle part 18.5–51.5 × 1.6–2.5 Septum indistinct Hyaline
C. communis Subcylindrical, medium brown, 20–60 × 3–4 μm. Polyblastic, subcylindrical to ampulliform, pale brown, 10–35 × 2–4 μm. Falcate with subobtuse ends (13–)17–20(–22) × (2–)2.5(–3) 0 Hyaline
C. couratarii Pale brown Lageniform to cylindrical, hyaline to pale brown, 10.5–37 × 2–3.5 μm Lunate 9.5–19 × 2–3 0 Hyaline
C. diversispora Pale brown to brown Polyblastic, sympodial, pale brown to brown, 4–9 × 2–3.5 μm. Type i) cylindrical, slightly uncinate at the ends, straight Type i) 11.5–16 × 2 Type i) 1-septate Hyaline
Type ii) cylindrical to slightly subacerose, slightly uncinate at the apex, abruptly attenuated at the base, straight Type ii) 19.5–25 × 1.5–2 Type ii) 1-septate
Type iii) long filiform, obtuse or rounded at the apex attenuated at the base, straight or curved Type iii) 28.5–47 × 1 Type iii) 1–3-septate
C. eucalypti Subcylindrical, medium brown, 10–30 × 3–4 μm. Polyblastic, subcylindrical to ampulliform, pale brown, 8–25 × 2.5–4 μm. Falcate, slightly curved, widest in middle with subobtuse ends (15–)18–21(–23) × 2–3 0 Hyaline
C. eucalypticola Subcylindrical, medium brown, 5–30 × 3–5 μm. Polyblastic, subcylindrical to ampulliform or lanceolate, pale brown, 5–20 × 3–3.5 μm. Falcate, straight to curved, widest in the middle, apex subobtusely rounded, base truncate, 0.5 μm diam (15–)20–26(–30) × (2.5–)3 0 Hyaline
C. eucalyptigena Subcylindrical, hyaline, frequently reduced to conidiogenous loci on hyphae, up to 15 μm tall, 3–5 μm diam. Polyblastic, hyaline, ampulliform or subcylindrical, 2–10 × 2–5 μm Falcate, tapering to acute ends that are subobtusely rounded (13–)18–24(–30) × 2(–2.5) 0 Hyaline
C. hyalopenicillata Cylindrical, penicillate, mono-, bi-, and terverticillate, hyaline, 24–69 × 1.5–3 μm. Mono- and polyblastic, short cylindrical, ampulliform, hyaline, 6.5–14 × 2–4 μm Fusiform, base pointed, apex obtuse 14–24 × 2–3 0–1 Hyaline
C. malaysiana Cylindrical, biverticillate, pale brown, 76–157 × 2.5–3 μm. Polyblastic, cylindrical, subcylindrical, hyaline, 19–28 × 2.5–3.5 μm. Fusiform, curved, apex acuminate, and base acuminate or slightly flattened 18–30 × 2–3 0–1 Hyaline
C. monoseptata Subcylindrical, unbranched, hyaline, 8–29 × 2–4 μm Polyblastic, cylindrical, hyaline Fusiform, curved 15.4–25.8 × 1.5–2.3 0–1 Hyaline
C. ramosa Cylindrical, penicillate, brown at the base, subhyaline at the apex, up to 70 μm long Polyblastic, subhyaline, 10–20 x 2.5–3.5 μm Falcate 26–44 × 2.2–3 (0–) 1 (–3) Hyaline

Presently, the genus Castanediella contains 14 species, and is shown to be diverse in its habitats. Most of Castanediella species have been collected from plant leaves. Castanediella acaciae, C. camelliae, C. communis, C. eucalypti, and C. eucalypticola were isolated from disease symptoms on different host plant leaves (Crous et al. 2015, 2016a, b; Wanasinghe et al. 2018) whereas C. cagnizarii is the only species found on decaying leaves submerged in a stream (Castañeda Ruiz et al. 2005). Some Castanediella species were reported from decaying leaves, such as C. brevis, C. cagnizarii, C. diversispora, C. hyalopenicillata and C. monoseptata (Castañeda Ruiz et al. 2005; Hernández-Restrepo et al. 2016b; Costa et al. 2018). Castanediella couratarii was reported from dead wood (Hernández-Restrepo et al. 2016a).

The genus Castanediella is morphologically similar to Idriella, Idriellopsis, Microdochium, Neoidriella, Paraidriella, Selenodriella (Seifert et al. 2011; Crous et al. 2015; Hernández-Restrepo et al. 2016a). However, these genera can be distinguished by the branching pattern of their conidiophores and conidial shape and septation (Hernández-Restrepo et al. 2016a). Castanediella differs from these genera by its branched conidiophores, ampulliform conidiogenous cells with scars instead of denticles, and filiform, 0–1-septate, straight to curved conidia (Crous et al. 2015). These similar-looking genera are phylogenetically distinct (Crous et al. 2015; Hernández-Restrepo et al. 2016a). A comparative synopsis of these genera is provided (Table 3).

Table 3.

Synopsis of Castanediella-like genera.

Genera Conidiophores Conidiogenous cells Conidia Chlamydospores
Castanediella Branched, pale brown to brown at the base and subhyaline at the apex. Sympodial, small denticles or scars, subhyaline. 0–1-sepate, falcate, lunate, cylindrical or fusiform, hyaline Not observed.
Idriella Brown, mostly reduced to conidiogenous cells Denticulate, sympodial Aseptate, lunate, curved, hyaline Brown, uni- or pluricellular.
Idriellopsis Unbranched, brown at the base, almost hyaline at the apex, mostly reduced to conidiogenous cells Terminal, denticulate 0–1-septate, falcate, curved, hyaline Not observed
Microdochium More or less verticillate, reduced to conidiogenous cells, hyaline Hyaline, sympodial or percurrent, sometimes denticulate Aseptate or multiseptate, lunate, falcate, fusiform, filiform, obovoid or subpyriform, straight or curved, hyaline Terminal or intercalary, solitary, in chains or grouped in clusters, brown.
Neoidriella Mostly unbranched, pale brown, mostly reduced to conidiogenous cells Sympodial, denticulate, terminal. Aseptate, cylindrical to obovoid, hyaline Intercalary or terminal, pale brown.
Paraidriella Unbranched, pale brown, mostly reduced to conidiogenous cells. Sympodial, denticulate, terminal. Aseptate, cylindrical to oblong, hyaline Not observed.
Selenodriella Unbranched or verticillate, brown. Sympodial, denticulate, terminal and intercalary. Aseptate, falcate, hyaline Not observed


We would like to thank Dr. Shaun Pennycook (Landcare Research Manaaki Whenua, New Zealand) for advising on the fungal names. The research is supported by the Thailand Research grants entitled The future of specialist fungi in a changing climate: baseline data for generalist and specialist fungi associated with ants, Rhododendron species and Dracaena species (grant no: DBG6080013), Impact of climate change on fungal diversity and biogeography in the Greater Mekong Subregion (grant no: RDG6130001). Y. Wang would like to thank the projects of the National Natural Science Foundation of China (No. 31560489, 31500451), Talent project of Guizhou science and technology cooperation platform ([2017]5788-5), Bijie Science and Technology Project ([2016]19) and Guizhou science and technology department international cooperation base project ([2018]5806). J.K. Liu would like to thank the National Natural Science Foundation of China (NSFC 31600032) and the Science and Technology Foundation of Guizhou Province (LH [2015]7061).


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