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Two new species of Fuscoporia (Hymenochaetales, Basidiomycota) from southern China based on morphological characters and molecular evidence
expand article infoQian Chen, Yu-Cheng Dai
‡ Beijing Forestry University, Beijing, China
Open Access

Abstract

Fuscoporia (Hymenochaetaceae) is characterized by annual to perennial, resupinate to pileate basidiocarps, a dimitic hyphal system, presence of hymenial setae, and hyaline, thin-walled, smooth basidiospores. Phylogenetic analyses based on the nLSU and a combined ITS, nLSU and RPB2 datasets of 18 species of Fuscoporia revealed two new lineages that are equated to two new species; Fuscoporia ramulicola sp. nov. grouped together with F. ferrea, F. punctatiformis, F. subferrea and F. yunnanensis with a strong support; Fuscoporia acutimarginata sp. nov. formed a strongly supported lineage distinct from other species. The individual morphological characters of the new species and their related species are discussed. A key to Chinese species of Fuscoporia is provided.

Keywords

Hymenochaetaceae, phylogeny, taxonomy, wood-rotting fungi

Introduction

Fuscoporia Murrill (1907) was established based on F. ferruginosa (Schrad.) Murrill. However, the genus has been unconsidered for a long time, reduced as a synonym of Phellinus Quél. (e.g., Overholts 1953; Ryvarden and Johansen 1980; Larsen and Cobb-Poulle 1990). Fiasson and Niemelä (1984) firstly used morphological features to segregate some members of Phellinus into distinct taxonomic entities, including Fomitiporia Murrill, Fulvifomes Murrill, Phellinus Murrill, Porodaedalea Murrill and Fuscoporia.

Fiasson and Niemelä (1984) defined Fuscoporia by annual to perennial, resupinate to pileate basidiocarps, a dimitic hyphal system with generative hyphae in the dissepiment edge and the tube trama often encrusted with crystals, presence of hymenial setae and hyaline, thin-walled, smooth basidiospores. Later on, phylogenetic studies based on nuclear large subunit (nLSU) ribosomal RNA-based phylogeny confirmed that Fuscoporia formed a lineage distinct from Phellinus s. s. (Wagner and Fischer 2001, 2002). Previous studies on Fuscoporia were mostly based on morphological characteristics (Groposo et al. 2007; Baltazar et al. 2009; Baltazar and Gibertoni 2010; Raymundo et al. 2013) but, recently, more new taxa were described based on both molecular analyses and morphology (Niemelä et al. 2001; Jang et al. 2012; Pires et al. 2015; Chen and Yuan 2017; Chen et al. 2019).

In our study, phylogenetic analyses were carried out based on the nLSU and combined ITS, nLSU and RPB2 datasets including 99 (60 newly generated) sequences representing 18 species of Fuscoporia. From the analyses, two new species of Fuscoporia were found and described. In addition, a key to Chinese species in the genus was provided.

Materials and methods

Morphological studies

The studied specimens are deposited in the herbarium of the Institute of Microbiology, Beijing Forestry University (BJFC). Macro-morphological descriptions are based on field observations and notes and dry herbarium specimens. The microscopic analyses followed that described by Cui et al. (2019). Sections were studied at ultimate magnification ×1000 applying Nikon Eclipse 80i microscopy and phase contrast illumination. Drawings were made with the aid of a drawing tube. The measurements and drawings were made from slide preparations stained with Cotton Blue. In recording spore size variation, 5% of measurements were excluded from each end of the range and given in parentheses. The following abbreviations are used in the article: KOH = 5% potassium hydroxide, CB = Cotton Blue, CB- = acyanophilous, IKI = Melzer’s reagent, IKI- = neither amyloid nor dextrinoid, L = mean spore length (arithmetic average of all spores), W = mean spore width (arithmetic average of all spores), Q = variation in the L/W ratios between specimens studied, and n (a/b) = number of spores (a) measured from given number of specimens (b). Special color terms are cited from Petersen (1996).

DNA extraction and sequencing

Extract total genomic DNA was extracted from dried specimens by CTAB rapid plant genome extraction kit (Aidlab Biotechnologies Co., Ltd., Beijing, China) according to the manufacturer’s instructions with some modifications (Chen et al. 2016; Han et al. 2016). To generate PCR amplicons, the following primer pairs were used: ITS4 and ITS5 (White et al. 1990) for the internal transcribed spacer (ITS), LR0R and LR7 (Vilgalys and Hester 1990) for nuclear large submit (nLSU) and bRPB2-6F and bRPB2-7.1R (Matheny 2005) for partial RNA polymerase II, second largest submit (RPB2). The PCR procedures followed Song and Cui (2017) and Zhu et al. (2019). DNA sequencing was performed at Beijing Genomics Institute and the sequences are deposited in GenBank and listed in Table 1.

Taxa and GenBank accession numbers for ITS, nLSU and RPB2 sequences used in the phylogenetic analyses (Fig. 2).

Species Sample no. Locality GenBank accession no.
ITS nLSU RPB2
Fuscoporia acutimarginata Dai 15137 China MH050751 MH050765 MN159384
Dai 16892 China MH050752 MH050766 MH079393
Dai 15115 China MN121764 MN121823 MN159385
F. atlantica SP 445618 Brazil KP058515 KP058517
SP 465829 Brazil KP058514 KP058516
F. callimorpha Dai 17388 Brazil MN121765 MN121824
Dai 17389 Brazil MN121766 MN121825
F. contigua Dai 16025 USA MG008401 MG008454 MH079406
JV 1204/22.6-J USA MG008403 MG008456 MH079407
Dai 13567A Romania MG008402 MG008455 MN159386
F. ferrea MUCL 45984 France KX961112 KY189112
Cui 11801 China KX961101 KY189101 MN159387
JV 1105/3-J USA MH050760 MH050770 MH079392
F. ferruginosa JV 1309/4 Slovakia KX961102 KY189102 MH079405
JV 1507/11-CN Europe MG008400 MG008453 MH079404
F. gilva Cui 11209 China MN121767 MN121826 MN159388
Dai 15681 China MN121768 MN121827 MN159389
F. insolita SP 5251 Russia KJ677113
SP 5208 Russia MN121769 MN121828
F. punctatiformis Doll#872 USA MH050753
Dai 17443 Brazil MH050755 MH050764
F. ramulicola Dai 15723 China MH050749 MH050762 MH079398
Dai 16155 China MH050750 MH050763 MH079399
F. rufitincta JV 1008/25 USA KJ940029 KX058575
JV 0904/142 USA KJ940030 KX058574
F. senex KUC 20110922-13 Korea JX463658 JX463652
MEL:2382630 Australia KP012992 KP012992
F. setifer Dai 15710 China MH050758 MH050767 MN159390
Dai 15706 China MH050759 MH050769 MN159391
F. subferrea Dai 16326 China KX961097 KY053472 MH079400
Dai 16327 China KX961098 KY053473 MH079401
F. torulosa JV 1405/2 Czech KX961106 KY189106 MN159392
JV 1312/19-Kout Spain KX961107 KY189107 MN159393
F. viticola JV 0911/6 Czech KX961110 KY189110
He 2081 USA MN121770 MN121829
F. wahlbergii Dai 15636 China MG008397 MG008450 MH079402
Dai 15659 China MG008398 MG008451 MH079403
F. yunnanensis Cui 8182 China MH050756 MN159394
Dai 15637 China MH050757 MH050768 MN159395
Coniferiporia sulphurascens Cui 10429 China KR350565 KR350555
C. weirii CFS 504 Canada AY829341 AY829345

Phylogenetic analyses

Sixty new sequences (nineteen ITS, seventeen nLSU and twenty-four RPB2) of Fuscoporia species were newly generated (Table 1). All sequences of ITS+nLSU+RPB2 analysis (Fig. 2) were shown in Table 1. Additional sequences of representatives genera of Hymenochaetaceae included in nLSU analysis (Fig. 1) were downloaded from GenBank to explore the phylogenetic relationships of Fuscoporia, which were used in the previous phylogenetic study (Zhou et al. 2016; Chen et al. 2019), thus not shown in Table 1. Oxyporus corticola (Fr.) Ryvarden, Oxyporus populinus (Schumach.) Donk, and Hyphodontia pallidula (Bres.) J. Erikss. were included as outgroups in nLSU analysis based on previous studies (Zhou et al. 2016; Chen et al. 2019). The outgroups selected for ITS+nLSU+RPB2 analysis were Coniferiporia weirii (Murrill) L.W. Zhou & Y.C. Dai and Coniferiporia sulphurascens (Pilát) L.W. Zhou & Y.C. Dai because Coniferiporia resulted as a sister group of Fuscoporia in previous studies (Fig. 1; Zhou et al. 2016; Chen and Yuan 2017; Chen et al. 2019).

Figure 1. 

Phylogenetic positions of Fuscoporia and the new species within the Hymenochaetaceae inferred from the nLSU sequences. Topology is from MP tree and statistical values (MP/BI/ML) are indicated for each node that simultaneously received BS from ML and MP not below 75%, and BPP from BI not below 0.9. Names of new species are in bold.

Figure 2. 

Phylogeny of Fuscoporia inferred from ITS+nLSU+RPB2 sequences. Topology is from MP tree and statistical values (MP/BI/ML) are indicated for each node that simultaneously received BS from ML and MP not below 75%, and BPP from BI not below 0.9. Names of new species are in bold.

Sequences were aligned with BioEdit (Hall 1999) and ClustalX (Thompson et al. 1997). Prior to phylogenetic analysis, ambiguous sequences at the start and the end were deleted and gaps were manually adjusted to optimize the alignment. Sequence alignment was deposited at TreeBase (http://purl.org/phylo/treebase; submission ID 22620). Phylogenetic analysis was carried out according to previous studies (Zhou 2015; Shen et al. 2019; Zhu et al. 2019). Maximum parsimony (MP), bayesian inference (BI) and maximum likelihood (ML) methods were employed to perform phylogenetic analysis of the two aligned datasets. MP analysis were performed using PAUP* 4.0b10 (Swofford 2002); BI was calculated with MrBayes3.1.2 (Ronquist and Huelsenbeck 2003); RAxML v.7.2.6 (Stamatakis 2006) was used for ML analysis. The three phylogenetic methods resulted in similar topologies for each dataset, and, thus, only the topology from the MP tree is presented along with statistical values from the ML/BI/MP algorithms (simultaneous MP/BI not less than 75 % and BPP not less than 0.9) at the nodes.

Results

The nLSU datasets included 23 representatives genera of Hymenochaetaceae and the combined ITS+nLSU+RPB2 datasets included 41 fungal specimens representing 20 species. In addition to sequences of new species, 14 new sequences of three species without published DNA sequences were uploaded – F. punctatiformis (Murrill) Zmitr., Malysheva & Spirin, F. setifer (T. Hatt.) Y.C. Dai and F. yunnanensis Y.C. Dai.

The nLSU dataset had an aligned length of 1386 characters, of which 996 were constant, 96 were variable but parsimony-uninformative, and 294 were parsimony-informative. Maximum Parsimony (MP) analysis yielded four equally most parsimonious trees (TL = 1639, CI = 0.350, RI = 0.733, RC = 0.256, HI = 0.650). Bayesian (BI) resulted in a similar consensus tree to that of the Maximum Parsimony (MP) and Maximum Likelihood (ML) analysis, with 1 million generations and an average standard deviation of split frequencies = 0.009570.

The three-gene dataset had an aligned length of 2950 characters, of which 1990 were constant, 90 were variable but parsimony-uninformative, and 870 were parsimony-informative. Maximum Parsimony (MP) analysis yielded 4 most parsimonious trees with near-identical topologies (TL = 2631, CI = 0.569, RI = 0.807, RC = 0.459, HI = 0.431). Bayesian (BI) resulted in a similar consensus tree to that of the Maximum Parsimony (MP) and Maximum Likelihood (ML) analysis, with 1 million generations and an average standard deviation of split frequencies = 0.005640.

Eighteen species of Fuscoporia formed a well-supported clade (94/1/96 in Fig. 1) within the Hymenochaetaceae. Fuscoporia is a sister genus to Coniferiporia. Two samples from southern China are clustered into a new highly supported lineage (100/1/100 in Fig. 2) and in a clade with F. ferrea (Pers.) G. Cunn., F. punctatiformis, F. subferrea Q. Chen & Y. Yuan, and F. yunnanensis with high support (99/1/98 in Fig. 1; 100/1/100 in Fig. 2); it is described as F. ramulicola sp. nov. Another three specimens formed a distinct lineage with strong support (100/1/100 in Fig. 2) in Fuscoporia. This clade is interpreted as a new species, F. Acutimarginata sp. nov.

Taxonomy

Fuscoporia acutimarginata Y.C. Dai & Q. Chen, sp. nov.

MycoBank No: 824732
Figs 3A, 4

Type

China. Yunnan Province: Kunming, Wild Duck Park, 2 August 2016, on fallen angiosperm branch, Dai 16892 (holotypes: BJFC 022998).

Etymology

Acutimarginata” (Latin): referring to the species with a sharp margin of fruiting body.

Description

Basidiocarps annual, effused-reflexed to pileate, broadly attached, without taste or odor and soft corky when fresh. Pilei conchate, laterally fused, convex towards margin, projecting up to 1.5 cm, 7 cm wide and 6 mm thick at base. Pileal surface yellowish brown to dark brown, velutinate, concentrically sulcate with zoned; margin acute, yellowish brown. Pore surface yellowish brown when dry, glancing; margin distinct, yellowish, up to 2 mm wide; pores circular to angular, 5–7 per mm; dissepiments thin, entire. Context yellowish brown to dull brown, corky, up to 3 mm thick. Tubes yellowish brown, paler than context, corky, up to 3 mm long.

Hyphal structure

Hyphal system dimitic; generative hyphae simple septate; tissue darkening but otherwise unchanged in KOH.

Subiculum

Generative hyphae rare, hyaline to pale yellowish, thin- to slightly thick-walled, occasionally branched, 2–3.5 μm in diam; skeletal hyphae dominant, yellowish brown, thick-walled with a wide lumen, unbranched, aseptate, interwoven, 2–4.3 μm in diam.

Tubes

Generative hyphae hyaline, thin-walled, occasionally branched, 2–3 μm in diam, occasionally encrusted at dissepiment edges; skeletal hyphae dominant, yellowish brown, thick-walled with a wide lumen, unbranched, aseptate, straight, subparallel along the tubes, 2–4 μm in diam. Irregular crystals occasionally present among trama and hymenia.

Hymenium

Hymenial setae rare, mostly originating from tramal hyphae, subulate, dark brown, thick-walled, 20–40 × 3–7 μm; cystidioles frequent, fusoid, sometimes covered with crystals, hyaline, thin-walled, 16.5–26 × 4–6.5 μm; basidia broadly clavate, with four sterigmata and a simple septum at the base, 14–17 × 4.8–6.5 μm; basidioles similar in shape to basidia, but slightly smaller. Basidiospores cylindrical, hyaline, thin-walled, smooth, IKI–, CB–, (7–)7.5–9(–9.8) × (2.2–)2.5–3.2 μm, L = 8.12 μm, W = 2.87μm, Q = 2.73–2.95 (n = 60/2).

Additional specimens examined (paratypes)

China. Hunan Province: Yizhang County, Mangshan Nature Reserve, Guizizhai, 16 Aug 2014, on fallen angiosperm trunk, Dai 15115 (BJFC 018227), Dai 15137 (BJFC 018253).

Figure 3. 

Basidiocarps of Fuscoporia. A F. acutimarginata (Dai 15137) B F. ramulicola (Dai 16155). Scale bars: 10 mm.

Figure 4. 

Microscopic structures of Fuscoporia acutimarginata (Holotype, Dai 16892) A basidiospores B basidia and basidioles C cystidioles D hymenial setae E generative hyphae at dissepiment edge F hyphae from trama G hyphae from subiculum.

Fuscoporia ramulicola Y.C. Dai & Q. Chen, sp. nov.

MycoBank No: 824734
Figs 3B, 5

Type

China. Yunnan Province: Binchuan County, Jizushan Park, 30 August 2015, on fallen angiosperm branch, Dai 15723 (holotypes: BJFC 019827).

Etymology

Ramulicola” (Latin) referring to the species growing on branches.

Description

Basidiocarps annual, resupinate, effused, inseparable, without taste or odor and corky when fresh, light-weight and hard corky when dry, up to 10 cm long, 2.2 cm wide and 1 mm thick at center. Pore surface grayish brown, fawn, cracked with age; sterile margin yellowish brown to olivaceous buff, distinctly paler than tubes, up to 1 mm wide; pores more or less angular, 6–7 per mm; dissepiments thin, sometimes irregular to slightly lacerate; abundant setae seen in tube cavities (under lens). Subiculum reddish brown, corky, very thin, about 0.1 mm thick. Tubes olivaceous buff, paler contrasting with pores and subiculum, hard corky, up to 0.9 mm long.

Hyphal structure

Hyphal system dimitic; generative hyphae simple septate; tissue darkening but otherwise unchanged in KOH.

Subiculum

Generative hyphae rare, hyaline, thin-walled, occasionally branched and simple septate, 2.5–3 μm in diam; skeletal hyphae dominant, rust-brown, thick-walled with a wide lumen, unbranched, aseptate, flexuous, loosely interwoven, 3–3.8 μm in diam.

Tubes

Generative hyphae rare, mostly present at dissepiment edges and subhymenium, hyaline, thin-walled, occasionally branched and frequently simple septate, 1.8–2.8 μm in diam, some of them at dissepiment edges and in the hymenium encrusted with small crystals; skeletal hyphae dominant, yellowish brown, thick-walled with a wide lumen, unbranched, aseptate, flexuous to more or less straight, subparallel along the tubes, 2.5–3.8 μm in diam. Irregular crystals usually present among trama and hymenia.

Hymenium

Hymenial setae frequent, mostly originating from hymenium, subulate, dark brown, thick-walled, 35–60 × 4.5–7 μm; cystidioles fusoid, sometimes covered with crystals, hyaline and thin-walled, 15–22 × 3–5 μm; basidia barrel-shaped, with four sterigmata and a simple septum at the base, 9–11 × 4.5–5.5 μm; basidioles frequently in hymenium, similar in shape to the basidia, but slightly smaller. Basidiospores cylindrical, hyaline, thin-walled, smooth, usually glued in tetrads, IKI–, CB–, with some small guttules, (5.2–)5.8–7(–7.2) × (1.8–)2–2.5(–2.8) μm, L = 6.37 μm, W = 2.28 μm, Q = 2.57–2.88 (n = 60/2).

Additional specimen examined (paratype)

China. Hainan Province: Wuzhishan County, Wuzhishan Nature Reserve, 14 Nov 2015, on fallen angiosperm branch, Dai 16155 (BJFC 020252).

Figure 5. 

Microscopic structures of Fuscoporia ramulicola (Holotype, Dai 15723) A basidiospores B basidia and basidioles C cystidioles D hymenial setae E generative hyphae at dissepiment edge F hyphae from trama G hyphae from subiculum.

Discussion

In the study, sixteen previously accepted species of Fuscoporia were referred to morphological examination and phylogenetic analyses. Two species of Fuscoporia from China, F. acutimarginata and F. ramulicola, are described as new on the basis of molecular data and morphology. F. acutimarginata is characterized by annual, effused-reflexed to pileate basidiocarps, small pores (5–7 per mm), a dimitic hyphal structure, hymenial setae rarely present, originating from tramal hyphae, the presence of cystidioles, and cylindrical basidiospores measuring 7.5–9 × 2.5–3.2 μm. Phylogenetically, samples of F. acutimarginata formed a well-supported monophyletic lineage distinct from other Fuscoporia species (Fig. 2). F. acutimarginata is very similar to F. setifer in having annual, effused-reflexed basidiocarps and the presence of cystidioles, but the latter has bigger pores (3–4 per mm) and smaller basidiospores (5.8–7 × 2–2.5 μm, Dai 2010). F. acutimarginata has a similar macromorphology to F. gilva (Schwein.) T. Wagner & M. Fisch., but the latter has frequently septate skeletal hyphae and ellipsoid to ovoid basidiospores (4–5 × 3–3.5 μm, Ryvarden 2004).

Fuscoporia ramulicola is distributed in southern China and characterized by annual and resupinate basidiocarps, small pores (6–7 per mm), a dimitic hyphal system, subulate hymenial setae, the presence of cystidioles, and cylindrical basidiospores measuring 5.8–7 × 2–2.5 μm. F. ferrea, F. subferrea, F. yunnanensis and F. ramulicola have overlapping distribution in China and clustered together with F. punctatiformis in a clade with strong support (99/1/98 in Fig. 1, 100/1/100 in Fig. 2). They are distinguishable according to their morphology and molecular data: F. ferrea is distinguished from F. ramulicola by its perennial basidiocarps and widely distributed in northern China, Europe and North America (Ryvarden and Gilbertson 1994, Lowe 1966, Chen et al. 2019); F. subferrea has smaller pores (7–10 per mm) and shorter basidiospores (4.2–6.2 × 2–2.6 μm, Chen and Yuan 2017); F. yunnanensis has larger pores (2–4 per mm) and wider basidiospores (6–8.3 × 2.4–3 μm, Dai 2010); F. punctatiformis has shorter hymenial setae (18–25 μm vs. 35–60 μm), subcylindrical basidiospores with a pointed apex (4–6 × 1.5–2 μm, Ryvarden and Johansen 1980) and is reported from America, Brazil and USA (Ryvarden 2004; Groposo et al. 2007). Fuscoporia ramulicola is similar to F. contigua and F. ferruginosa in having resupinate basidiocarps; however, F. contigua and F. ferruginosa have mycelial, extra-hymenial setae (Chen et al. 2019); in addition, the three species are distantly related in our phylogenies (Figs 1, 2). Fuscoporia montana Y.C. Dai & Niemelä and F. chrysea (Lév.) Baltazar & Gibertoni are also similar to F. ramulicola in sharing resupinate basidiocarps, abundant subulate hymenial setae, absence of mycelial setae and distributed in southern China. Fuscoporia montana differs from F. ramulicola in having ovoid basidiospores 6.5–8.2 × 3.2–4.2 μm (Niemelä et al. 2001) whereas F. chrysea is different from F. ramulicola by its smaller pores (9–10 per mm) and shorter basidiospores (3–4 × 2–2.5 μm, Dai 2010).

Key to Fuscoporia species in China

1 Basidiocarps usually laterally stipitate; hymenial setae absent F. discipes
Basidiocarps sessile; hymenial setae present 2
2 Basidiocarps completely resupinate 3
Basidiocarps effuse-reflexed to pileate 11
3 Mycelial setae present in the decayed wood and margin of basidiocarps (by hand lens) 4
Mycelial setae absent from the decayed wood and margin of basidiocarps (by hand lens) 5
4 Pores 7–8 per mm F. ferruginosa
Pores 2–3 per mm F. contigua
5 Basidiocarps perennial 6
Basidiocarps annual 9
6 Pores 5–7 per mm 7
Pores 7–10 per mm 8
7 Basidiospores cylindrical, 6–7.8 × 2–2.5 μm F. ferrea
Basidiospores subcylindrical, 4–6 × 1.5–2 μm F. punctatiformis
8 Pores 9–10 per mm; basidiospores ellipsoid, < 5 μm long F. chrysea
Pores 7–8 per mm; basidiospores narrowly ovoid, > 5 μm long F. montana
9 Pores 2–4 per mm F. yunnanensis
Pores 6–10 per mm 10
10 Pores 7–10 per mm; basidiospores 4.2–6.2 µm long F. subferrea
Pores 6–7 per mm; basidiospores 6–7 µm long F. ramulicola
11 Hymenial setae usually hooked F. wahlbergii
Hymenial setae straight 12
12 Pores 3–4 per mm F. setifera
Pores 5–11 per mm 13
13 Basidiocarps annual, margin acute 14
Basidiocarps perennial, margin obtuse 15
14 Skeletal hyphae septate, spores ellipsoid, 3.3–4.2 × 2.2–3 μm F. gilva
Skeletal hyphae aseptate, spores cylindrical, 7.5–9 × 2.5–3.2 μm F. acutimarginata
15 Basidiocarps subungulate; contextual skeletal hyphae aseptate F. torulosa
Basidiocarps usually applanate; contextual skeletal hyphae septate 16
16 Basidiospores 3.3–4.1 × 2.1–2.4 μm, skeletal hyphae unchanged in KOH F. rhabarbarina
Basidiospores 4–4.8 × 3.6–3.9 μm, skeletal hyphae swelling in KOH F. senex

Acknowledgements

We are grateful to Dr. Shuang-Hui He (BJFC, China) for his companionship on field trips. The research is financed by the National Natural Science Foundation of China (Project Nos. U1802231 & 31530002).

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