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Research Article
Three new species of Candolleomyces (Agaricomycetes, Agaricales, Psathyrellaceae) from the Yanshan Mountains in China
expand article infoHao Zhou, GuiQiang Cheng, XiMei Sun, RuiYi Cheng§, HongLiang Zhang§, YanMin Dong§, ChengLin Hou
‡ Capital Normal University, Beijing, China
§ Beijing Songshan National Nature Reserve Management Office, Beijing, China
Open Access

Abstract

Three new species, Candolleomyces incanus, C. subcandolleanus and C. yanshanensis, were found and described from Yanshan Mountains in China. The identification is based on morphological observation combined with phylogenetic analysis of ITS-LSU-Tef1α-TUB2. This study enriched the species diversity of Candolleomyces in Yanshan Mountains and provided important data support for the systematic study of Candolleomyces in the future.

Keywords

molecular systematics, new taxon, Psathyrellaceae, taxonomy

Introduction

Candolleomyces Wächter & A. Melzer was established in 2020, belonging to Basidiomycota, Agaricomycetes, Agaricales, Psathyrellaceae (Wächter and Melzer 2020). In a previous study, this genus was subordinate to Psathyrella (Fr.) Quél. (1872) and molecular sequence data have improved understanding of relationships of Psathyrella species (Hopple and Vilgalys 1999; Moncalvo et al. 2002; Matheny et al. 2006). However, the combination analysis of the ITS and LSU regions showed that the delimitation of some species within Psathyrella are still unclear (Larsson and Örstadius 2008). In more recent studies, multi-gene loci (for example, ITS, LSU, Tef1α and TUB2) became the main methods for identification of Psathyrella (Wang and Bau 2014; Yan and Bau 2017, 2018a, 2018b, 2021; Yan 2018; Yan et al. 2019).

In previous studies of Psathyrella, there are approximately 100 taxa lacking pleurocystidia, but this feature has not been used as a key distinguishing feature (Fries 1838; Smith 1972; Kits van Waveren 1985; Örstadius and Kundsen 2012; Battistin et al. 2014). Based on extensive specimen collection, morphological studies and phylogenetic analyses, Candolleomyces has been separated from Psathyrella as a new genus and it differs from Psathyrella s.s. in lacking pleurocystidia. (Wächter and Melzer 2020).

Currently, there are 25 recognised species in Candolleomyces in the Index Fungorum website (http://www.indexfungorum.org, until Jan. 2022) and 10 species were reported in China (Yan 2018; Bau and Yan 2021).

Yanshan Mountains are located in North China and have a warm temperate continental monsoon climate, with higher plant diversity. The dominant plants include Quercus spp., Betula spp., Abies spp. and Pinus tabuliformis Carr. et al. (Wang et al. 2021). There is no information about Candolleomyces as yet. In this study, based on morphological characters and the phylogenetic analyses, three new species of Candolleomyces from Yanshan Mountains in China are described.

Materials and methods

Morphological studies

Collections were obtained and photographed in the field from Yanshan Mountains in China from 2017 to 2020. The collected specimens were dehydrated with a dryer (Dorrex) at 50 °C and the specimens were deposited in the Herbarium of the College of Life Science, Capital Normal University, Beijing, China (BJTC). Macroscopic characters were recorded from specimens. Microscopic characters were observed in thin sections of specimens mounted in 3% potassium hydroxide (KOH) or sterilised water. The shape and size of microscopic structures were observed and noted using a light microscope [Olympus DP71, Tokyo, Japan]. The measurements and Q values are given as (a)b–c(d), in which “a” is the lowest value, “b–c” covers a minimum of 90% of the values and “d” is the highest value. Q stands for the ratio of length and width of a spore (Bau and Yan 2021). Nomenclatural details were submitted to the MycoBank. In this study, the morphological colour comparison was compared to the reference website colorhexa (https://www.colorhexa.com).

DNA extraction PCR amplification and sequencing

DNA extraction was achieved by the M5 Plant Genomic DNA Kit [Mei5 Biotechnology, Co., Ltd, China]. The purified DNA was dissolved in 1 × TE buffer and stored at – 20 °C for later use. The PCR amplifications were performed in Bio-Rad S1000 TM Thermal Cycler [Bio-Rad Laboratories, Inc, USA]. The primer sets ITS1/ITS4 (White et al. 1990) were used to amplify the rDNA ITS region, LR5/LR0R (Vilgalys and Hester 1990) were used to amplify the large subunit nuclear ribosomal DNA (nuLSU rDNA) region and EF983F/EF2218R (Örstadius et al. 2015) were used to amplify the translation elongation factor subunit 1 alpha (Tef1α) region. The primer sets B36f and B12r (Nagy et al. 2011) were used to amplify the β-tubulin gene (TUB2) region. PCRs were performed in a volume of 25 μl consisted of 2 μl of DNA template; 1 μl of (10 μM) per primer; 12.5 μl 2 × Master Mix [Mei5 Biotechnology, Co., Ltd, China]. PCR amplification conditions refer to Bau and Yan (2021). DNA sequences were sequenced by Zhongkexilin Biotechnology, Co., Ltd, Beijing, China.

Molecular data analyses

The generated raw reads of the DNA sequences were used to obtain consensus sequences using SeqMan v.7.1.0 in the DNASTAR Lasergene Core Suite software (DNASTAR Inc., Madison, WI, USA). All sequences were aligned using MAFFT v.6 (Katoh and Toh 2010) and trimmed manually using MEGA 6 (Tamura et al. 2013). For phylogenetic analyses, newly-obtained sequences and additional reference sequences of Candolleomyces species were included in the dataset of combined ITS-LSU-Tef1α-TUB2 muti-locus DNA (Table 1), with Psathyrella multipedata (Peck) A.H. Sm. (LÖ237-04) used as outgroup. Phylogenetic analyses were performed using PAUP v.4.0b10 for Maximum Parsimony (MP) analysis (Swofford 2003) and MrBayes v.3.1.2 for Bayesian Inference (BI) analysis (Ronquist and Huelsenbeck 2003). ML gene-trees were estimated using the software RAxML 7.4.2 Black Box (Stamatakis 2006; Stamatakis et al. 2008; Zhou and Hou 2019; Zhou et al. 2021).

Table 1.

Sequences information used in the phylogenetic analysis in this study.

Taxa Voucher Locality ITS LSU β-Tub tef-1α
Candolleomyces aberdarensis GLM-F116094 Kenya MH880928
C. albipes DED8340 Sao Tome KX017209
C. badhyzensis 79478 (TAA) Type Turkmenistan KC992883 KC992883
C. badiophyllus SZMC-NL-2347 FN430699 FM876268 FN396261 FM897252
C. cacao SFSU DED 8339 Sao Tome NR148106
C. cacao FP1R4 USA KU847452
C. cacao MP2R2 USA KU847436
C. candolleanus LAS73030 Neotype Sweden KM030175 KM030175
C. cladii-marisci CLUF302 Type Italy MK080112
C. efflorescens Pegler2133 (K) Sri Lanka KC992941
C. eurysporus GLM-F126263 Type Germany MT651560 MT651560
C. incanus BJTC Z777 Type China: Beijing ON042759 ON042766 ON098513 ON098508
C. incanus BJTC S173 China: Beijing ON042760 ON042767 ON098514 ON098509
C. leucotephrus LÖ138-01 (UPS) Sweden KC992885 KC992885 KJ664865 KJ732775
C. luteopallidus Sharp20863 (MICH) Type USA KC992884 KC992884
C. luteopallidus HMJAU5148 China: Jilin MG734736 MW301084 MW314056 MW314073
C. secotioides UES2918 Type Mexico KR003281 KR003282 KR003283
C. singeri HMJUA37867 China: Jilin MG734718 MW301088 MW314059 MW314077
C. singeri HMJAU37877 China: Chongqing MW301073 MW301091 MW314062 MW314080
C. subcacao HMJAU37807 Type China: Henan MW301064 MW301092 MW314063 MW314081
C. subcacao HMJAU37808 China: Henan MW301065 MW301093 MW314064 MW314082
C. subcacao HFJAU1014 China: Jiangxi MW559218
C. subcacao HFJAU1274 China: Jiangxi MW559219
C. subcacao HFJAU1488 China:Anhui MW559220
C. subcandolleanus BJTC Z239 Type China: Tianjin ON042755 ON042762 ON098510 ON098505
C. subcandolleanus BJTC Z232 China: Tianjin ON042756 ON042763
C. subminutisporus HMJAU37801 Type China: Hubei MW301066 MW301094 MW314065 MW314083
C. subminutisporus HMJAU37916 China: Henan MW301067 MW301095 MW314066 MW314084
C. subsingeri HMJAU37811 Type China: Jilin MG734715 MW301097 MW314067 MW314085
C. subsingeri HMJAU37913 China: Jilin MG734725 MW301098 MW314068 MW314086
C. sulcatotuberculosus GB:LO55-12 KJ138422 KJ138422
C. sulcatotuberculosus HFJAU1515 China: Fujian MW375696 MW382967 MW382965
C. sulcatotuberculosus Chiarello 07-10-2013 KJ138423
C. trinitatensis TL9035 (C) Ecuador KC992882 KC992882 KJ664863
C. trinitatensis ADK4162 (BR) Togo KC992886 KC992886
C. yanshanensis BJTC Z783 China: Beijing ON042757 ON042764 ON098511 ON098506
C. yanshanensis BJTC Z110 Type China: Beijing ON042758 ON042765 ON098512 ON098507
Candolleomyces sp. BAB-4773 India KP686450
Candolleomyces sp. BAB-5172 India KR349656
Candolleomyces sp. BAB-4748 India KR154977
Candolleomyces sp. BAB-4747 India KR154976
Candolleomyces sp. BAB-5202 India KT188611
Psathyrella multipedata LÖ237-04 Sweden KC992888 KC992888 KJ664867 KJ732777

Maximum Parsimony analysis was performed by a heuristic search option of 1000 random-addition sequences with a tree bisection and reconnection (TBR) algorithm. Maxtrees were set to 1000, branches of zero length were collapsed and all equally parsimonious trees were saved. Other calculated parsimony scores were tree length (TL), consistency index (CI), retention index (RI) and rescaled consistency (RC) (Zhou and Hou 2019).

Maximum Likelihood analysis was performed with a GTR site substitution model (Guindon et al. 2010). Branch support was calculated with a bootstrapping (BS) method of 1000 replicates (Hillis and Bull 1993). Bayesian Inference (BI) analysis, using a Markov Chain Monte Carlo (MCMC) algorithm, was performed (Rannala and Yang 1996). MrModeltest v. 2.3 was used to estimate the best model. Two MCMC chains were run from random trees for 10,000,000 generations and stopped when the average standard deviation of split frequencies fell below 0.01. Trees were saved for each 1000 generations. The first 25% of trees were discarded as the burn-in phase of each analysis. Branches with significant Bayesian posterior probabilities (BPP) were estimated in the remaining trees (Posada and Crandall 1998).

The combined alignment and phylogenetic tree were submitted on TreeBASE (www.treebase.org, study 29579).

Result

Phylogenetic analyses

For the ITS-LSU- Tef1α-TUB2 sequence dataset, a total of 3459 characters including gaps (694 for ITS, 1316 for LSU, 1023 for Tef1α, and 426 for TUB2) were included in the phylogenetic analysis. Using RAxML, MrBayes and PAUP to construct ML, Bayesian and MP phylogenetic trees, the results show that the topology and branching order were similar and the Bayesian tree is shown in this paper (Fig. 1). The Maximum likelihood analysed was performed with a GTR model. For the Bayesian analyses, the GTR + I + G models were recommended by MrModeltest. The heuristic search using Maximum Parsimony (MP) generated 1000 parsimonious trees (TL = 1168, CI = 0.768, RI = 0.815, RC = 0.232) and branches of zero length were collapsed and all multiple parsimonious trees were saved.

Figure 1. 

Multi-gene phylogenetic tree obtained from the Bayesian analysis. Numbers above branches are Bayesian posterior probability (pp) values, Maximum Likelihood bootstrap (MLB) and Maximum parsimony bootstrap (MP) values. Asterisks (*) denote branches with pp = 1.00, MLb = 100% and MPb = 100%. Numbers above branches represent strongly and moderately support (pp ≥ 0.95, MLb ≥ 50% and MPb ≥ 50%). The red font indicates the position of the new species.

Based on the results, six specimens were assigned to three branches and were described as three new species. The three new species (Candolleomyces yanshanensis, C. subcandolleanus, C. incanus) and a known species (Candolleomyces badiophyllus (Romagn.) D. Wächt. & A. Melzer etc.) clustered together in the phylogenetic tree. The three new species clustered into together (pp = 0.99, MLbs = 82%,MPbs = 74%), but three new species separately formed three subclades with high support value. Candolleomyces yanshanensis, C. subcandolleanus and C. incanus can be distinguished by the phylogenetic tree, sequence base differences and morphological characteristics.

Taxonomy

Candolleomyces yanshanensis C. L. Hou & H. Zhou, sp. nov.

MycoBank No: 843464
Fig. 2

Etymology

yanshanensis referred to the locality where the type specimen was collected.

Type

China, Beijing, Changping District, Beitaizi Village, 40.272906°N, 116.4203°E, alt. 149 m, 14 Aug 2019, coll. X.Y. Shen, H Zhou and R.T. Zhang, BJTC Z110.

Figure 2. 

Basidiomata and microscopic features of Candolleomyces yanshanensis (BJTC Z110) A, B basidiomata C basidia D pileipellis E basidiospores F cheilocystidia. Scale bars: 20 mm (A, B); 10 μm (C); 20 μm (D); 5 μm (E); 20 μm (F).

Diagnosis

Candolleomyces yanshanensis, pileus 20–60 mm, flabellate, flattening with age, hygrophanous. Basidiospores 5.8–8.2 × 3.3–5.4 μm, often with germ pore. Subglobose cell, irregular oval, (18) 20–27μm broad.

Description

Pileus 20–60 mm, flabellate, flattening with age, hygrophanous, slightly dirty white (#e3dac9) to pale brown (#deb887). Veil white (#ffffff), fibrils in young, evanescent. Context 1.0–2.0 mm broad at centre, same colour as pileus. Lamellae sparsely to moderately, adnate, slightly dirty white (#e3dac9) to champagne (#fad6a5), edge white (#ffffff) as spores mature. Stipes 50–130 × 3–6 mm, smooth, fibrils on the base, cornsilk (#f0ead6) to white (#ffffff).

Basidiospores 5.8–8.2 × 3.3–5.4 μm, Q = 1.4–2.0, ellipsoid to long ellipsoid, ovoid to ellipsoid, partly triangular at base, dark brown (#b8860b) to brown (#b06500) in water, smooth, abundant, multi-guttules, often with germ pore. Basidia 17–31 × 5.8–7.5 μm, short clavate, hyaline, 4-spored. Cheilocystidia 22–35 (40) × 8–11 (15) μm, irregular utriform or claviform, apex obtuse or broadly obtuse or often subcapitate, rarely with deposits. Pileipellis consists of 2–3 cells deep layer of irregular subglobose cell, irregular oval, (18) 20–27μm broad.

Habit and habitat

. Clumped on the ground with rich humus in broad-leaved forests or broad-leaved shrubs.

Additional specimen examined

China, Beijing, Changping District, Tailing, 40.327397°N, 116.21916°E, alt. 172 m, 17 Aug 2020, coll. X.Y. Shen, H Zhou and X.B. Huang, BJTC Z783.

Candolleomyces subcandolleanus C. L. Hou & H. Zhou, sp. nov.

MycoBank No: 843466
Fig. 3

Etymology

subcandolleanus referred to its morphological similarity to Candolleomyces candolleanus (Fr.) D. Wächt. & A. Melzer.

Figure 3. 

Basidiomata and microscopic features of Candolleomyces subcandolleanus. (BJTC Z239) A, B basidiomata C basidia D pileipellis E basidiospores F cheilocystidia. Scale bars: 10 mm (A, B); 10 μm (C); 20 μm (D); 5 μm (E); 20 μm (F).

Type

China, Tianjin, Jizhou District, Sanjiebei, 40.227984°N, 117.43354°E, alt. 235 m, 17 Aug 2019, coll. X.Y. Shen, H. Zhou and R.T. Zhang, BJTC Z239.

Diagnosis

Candolleomyces subcandolleanus, pileus 5–20 mm. Basidiospores 5.5–6.7 × 3.2–4.5 μm, germ pore absent. Cheilocystidia 21–28 (30) × 8–12 (15) μm. Subglobose cell, irregular oval or long oval, (13) 16–25 μm broad.

Description

Pileus 5–20 mm, campanulate to conical, smooth, fibrils in young, evanescent, brown (#b06500) to golden brown (#996515). Veil white (#ffffff), fibrils in young, evanescent. Context 0.2–0.5 mm broad at centre, same colour as pileus. Lamellae moderately to normally, adnate, slightly dirty white (#e3dac9) to white (#ffffff), edge white (#ffffff) as spores mature. Stipes 20–60 × 1–3 mm, smooth, fibrils on the base, cornsilk (#f0ead6) to white (#ffffff).

Basidiospores 5.5–6.7 × 3.2–4.5 μm, Q = 1.4–2.0, ellipsoid to ovoid, pale cream (#fffff0) to pale lemon (#fffacd) in water, smooth, multi-guttules, germ pore absent. Basidia 18–27 × 5–10 μm, short clavate, hyaline, 4-spored. Cheilocystidia 21–28 (30) × 8–12 (15) μm, utriform or claviform, apex obtuse or broadly obtuse or often subcapitate, rarely with deposits. Trama of gills irregular. Pileipellis consists of irregular subglobose cell, irregular oval or long oval, (13) 16–25 μm broad.

Habit and habitat

Clumped on the ground with rich humus in broad-leaved forests or broad-leaved shrubs.

Additional specimen examined

China, Tianjin, Jizhou District, Huangyaguan Great Wall, 40.245615°N, 117.44047°E, alt. 235 m, 17 Aug 2019, coll. X.Y. Shen, H. Zhou and R.T. Zhang, BJTC Z232.

Candolleomyces incanus C. L. Hou & H. Zhou, sp. nov.

MycoBank No: 843465
Fig. 4

Etymology

incanus referred to the basidiomata appears incanus.

Type

China, Beijing, Changping District, Sidaohe Village, 40.246374°N, 116.4406°E, alt. 114 m, 16 Aug 2020, coll. X.Y. Shen, H Zhou and X.B. Huang, BJTC Z777.

Figure 4. 

Basidiomata and microscopic features of Candolleomyces incanus (BJTC Z777) A, B basidiomata C basidia D pileipellis E cheilocystidia F basidiospores. Scale bars: 20 mm (A, B); 10 μm (C); 20 μm (D, E, F).

Diagnosis

Candolleomyces incanus, pileus 5–25 mm, hemispherical to conical. Basidiospores 6.0–7.0 × 3.2–4.5 μm. Stipe 40–70×4–6 mm, smooth, germ pore absent. Subglobose cell, irregular oval or long oval, (22) 25–32 μm broad.

Description

Pileus 5–25 mm, hemispherical to conical, hygrophanous, incanus (#f2f3f4) to nude (#fdf5e6). Veil white (#ffffff), fibrils in young, evanescent. Context 0.5–1.0 mm broad at centre, same colour as pileus. Lamellae moderately to normally, adnate, off-white (#f2f3f4) to white (#ffffff), edge white (#ffffff) as spores mature. Stipes 40–70 × 4–6 mm, smooth, hygrophanous, cornsilk (#f0ead6) to white (#ffffff).

Basidiospores 6.0–7.0 × 3.2–4.5 μm, Q = 1.4–1.9, ellipsoid, floral white (#fffaf0) to dark yellow (#eedc82) in water, smooth, abundant, multi-guttules, germ pore absent. Basidia 15–20 × 5–8 μm, short clavate, hyaline, 4-spored. Cheilocystidia 17–27 (31) × 7–11 (13) μm, utriform, apex obtuse or broadly obtuse or often subcapitate, rarely with deposits. Trama of gills irregular. Pileipellis consisted of irregular subglobose cell, irregular oval or long oval, (22) 25–32 μm broad.

Habit and habitat

Clumped on the ground with rich humus in deciduous broad-leaved or deciduous coniferous forests.

Additional specimen examined

China, Beijing, Yanqing District, Yudu Mountain, 40.54399°N, 115.893984°E, alt. 860 m, 12 Sep 2018, coll. C.L. Hou, H Zhou and J.Q. Li, BJTC 646.

Discussion

In this study, three new species were identified by morphology and phylogeny. It is very interesting that the three new species C. yanshanensis, C. subcandolleanus and C. incanus formed a stronger supported clade and they clustered with Candolleomyces badiophyllus (Romagn.) D. Wächt. & A. Melze, Candolleomyces candolleanus, Candolleomyces badhyzensis (Kalamees) D. Wächt. & A. Melzer, Candolleomyces trinitatensis (R.E.D. Baker & W.T. Dale) D. Wächt. & A. Melzer and Candolleomyces cladii-marisci (Sicoli, N.G. Passal., De Giuseppe, Palermo & Pellegrino) J.Q. Yan together in the phylogenetic tree. In addition, three new species were weakly sister to the known species C. badiophyllus in the phylogenetic tree.

Candolleomyces yanshanensis and C. subcandolleanus are different in macroscopic morphology of basidiomata. Candolleomyces yanshanensis is lighter in pileus colour and C. yanshanensis has larger spores (5.8–8.2 × 3.3–5.4 vs. 5.5–6.7×3.2–4.5 μm) and longer cheilocystidia (22–35 × 8–11 vs. 21–28 × 8–12 μm) than those of C. subcandolleanus. Moreover, C. yanshanensis spores often have a germ pore. Candolleomyces subcandolleanus is very easily confused with C. candolleanus in the field because of their similar macroscopic characteristics. In particular, two species in these sections possess the combined characteristics of small basidiomata. C. candolleanus is the type species of Candolleomyces, with early studies on this species being based on the number of pleats and other characteristics, but this also led to confusion in the identification of this species. Candolleomyces subcandolleanus can be distinguished from C. candolleanus by the smaller spores (5.5–6.7 × 3.2–4.5 vs. 7–9 × 4–5 μm) (Kits van Waveren 1980; Breitenbach and Kränzlin 1995; Mifsud 2017).

Candolleomyces incanum, C. badiophyllus, C. candolleanus and C. badhyzensis are close to each other in the phylogenetic tree. However, the four species show significant differences in morphology. These species can be distinguished as follows: C. incanus has smaller and narrower spores (6.0–7.0 × 3.2–4.5 μm), whereas C. candolleanus, C. badhyzensis and C. badiophyllus have larger spores (Spores of C. candolleanus were 7.0–9.0 × 4.0–5.0 μm, spores of C. badhyzensis were 10.2–11.5 × 5.5–6.5 μm, spores of C. badiophyllus were 10–14 × 5–6 μm). In addition, C. incanus has smaller cheilocystidia (17–27 × 7–11 vs. 34–51 × 10–15 μm) than those of C. badhyzensis (Kalamees 1981; Kasik et al. 2004; Wächter and Melzer 2020).

Except for morphological differences, the three new species in this study can also be distinguished by sequence similarity. Candolleomyces yanshanensis (BJTC Z110) can be distinguished, based on nucleotide differences in ITS, LSU, Tef1α and TUB2 loci from C. subcandolleanus (BJTC Z239) (sequence base similarity 93% in ITS, 100% in LSU, 99% in Tef1α and 98% in TUB2); C. yanshanensis (BJTC Z110) can be distinguished, based on nucleotide differences from C. incanus (BJTC Z777) (sequence base similarity 80% in ITS, 99% in LSU, 99% in Tef1α and 96% in TUB2); C. subcandolleanus (BJTC Z239) can be distinguished, based on nucleotide differences from C. incanus (BJTC Z777) (sequence base similarity 81% in ITS, 99% in LSU, 99% in Tef1α and 98% in TUB2). It can also be found that the ITS loci have a greater degree of differentiation for the species in Candolleomyces, Nevertheless, LSU and Tef1α were more conservative for the genus.

According to the research of Wächter and Melzer (2020), the species of Candolleomyces may be more abundant than previously thought and better delimitation of species boundaries is required. While the boundaries of some species are disputed, the number of new taxa is steadily increasing (Sicoli et al. 2019; Büttner et al. 2020; Bau and Yan 2021). However, the continued discovery of clear boundaries in new taxa like this study enhances our comprehension of species in this genus.

It is considered that the natural growth of Candolleomyces may be related to precipitation. However, the investigation and specimen collection in this study were carried out in the rainy season in July to August, with no collection in other periods. Therefore, more species of Candolleomyces might be expected in Yanshan Mountains.

Acknowledgements

This study was supported by the Biodiversity Survey and Assessment Project of the Ministry of Ecology and Environment, China. (2019HJ2096001006) and National Natural Science Foundation of China (No. 31870629).

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