Research Article
Research Article
Three coralloid species of the genus Trechispora (Trechisporales, Basidiomycota) in China: two newly discovered taxa and one reported for the first time
expand article infoPeng-Tao Deng, Jun Yan, Xiang-Fen Liu, Zheng-Mi He, Yuan Lin§, Ming-Xin Lu§, Ping Zhang
‡ Hunan Normal University, Changsha, China
§ Bureau of Forestry,Tongdao Dong Autonomous, Huaihua, China
Open Access


Two new species of Trechispora indigenous to southern China, T. laxa and T. tongdaoensis, are described and illustrated, and the first record of T. khokpasiensis in China is reported. Molecular phylogenetic analyses of the concatenated nuclear rDNA ITS1–5.8S–ITS2 and nuclear large subunit sequences supported the inclusion of the three species within the Trechispora clade, together with species formerly classified in Scytinopogon. The new species are similar in micromorphology to species of Trechispora (as traditionally circumscribed) but are distinguished by having coralloid basidiomata. A key to the known coralloid Trechispora species in China is provided.

Key words

Coral fungi, Phylogenetic analysis, Scytinopogon, Taxonomy


The genus Trechispora P. Karst was established by Karsten (1890) with Trechispora onusta P. Karst as the type species. Trechispora is the largest genus in the order Trechisporales (Larsson 2007), which is highly diverse in morphology: stipitate, clavarioid, or resupinate basidiomata (Meiras-Ottoni et al. 2021; Sommai et al. 2023); smooth, grandinioid, odontoid, hydnoid, or poroid hymenophores; short cylindric basidia, clamped, with 2 or 4 sterigmata on the basidia; and smooth or variously ornamented basidiospores. In addition, ampullate septa is an important character in Trechispora (Bernicchia and Gorjón 2010, Ordynets et al. 2015, Meiras-Ottoni et al. 2021, Liu et al. 2022). Calcium oxalate crystals usually accumulate on the mycelium or subhymenial hyphae, and the crystal morphology can be useful for species identification (Larsson 1994). Currently, approximately 90 species are accepted in Trechispora (Liu et al. 2022; Sommai et al. 2023). Consistent with previous studies (Birkebak et al. 2013), most Trechispora species are distributed in the tropics or subtropics (Chikowski et al. 2020). The placement of Trechispora in the order Trechisporales is supported by phylogenetic analyses of molecular data (Hibbett et al. 2007; Larsson 2007).

Scytinopogon Singer, erected by Singer (1945), has been assigned to several different families in the past: Clavariaceae Chevall (Corner 1970), Thelephoraceae Chevall (Donk 1964), and Gomphaceae (Maas Geesteranus 1962). The genus has also been suggested to be related to the Hydnodontaceae (Jülich 1981). Morphologically, Scytinopogon is characterized by clavarioid basidiomata with flattened and dense branches, which distinguish the species from Trechispora. However, phylogenetic analyses of molecular data indicate that Scytinopogon is nested within Trechispora (Hydnodontaceae), and no clear delimitation exists between the two genera. Because the name Trechispora has nomenclatural priority, Scytinopogon has been synonymized with Trechispora, thus rendering Trechispora a large, monophyletic genus (Meiras-Ottoni et al. 2021). To determine the correct name for Scytinopogon species, sequences and specimens for the type species were needed (Meiras-Ottoni et al. 2021). For this reason, to avoid misinterpretations in species delimitation within Trechispora, not all currently accepted Scytinopogon species have been transferred to Trechispora. Scytinopogon cryptomerioides W.R. Lin and P.H. Wang has recently been described from Taiwan (Lin et al. 2022).

During research on clavarioid fungi indigenous to southern China, two undescribed and one recently described coralloid Trechispora species were collected. Descriptions and illustrations of these three species are provided, and phylogenetic reconstructions based on nuclear rDNA ITS1–5.8S–ITS2 (ITS) and nuclear large subunit (LSU) sequences support the distinction of the new species and their placement in Trechispora.

Materials and methods

Specimen sources

Field work was conducted and specimens gathered by the authors from 2011 to 2022 in Hainan, Hunan, and Guangdong provinces, China. The habitat and morphological characters of fresh specimens were recorded in the field, including their dimensions and color. The fresh fruiting bodies were dried using heat or silica gel. The dried specimens were deposited in the Mycological Herbarium of Hunan Normal University (MHHNU), Changsha, China.

Morphological observation

Macroscopic characteristics were mainly derived from record sheets and photographs. The colors cited in the descriptions are based on those of Kornerup and Wanscher (1978) and Ridgway (1912). Dried fruiting body sections were placed in 3% KOH solution containing 1% Congo red solution. Microscopic characters were observed from a small portion of dried hymenial tissue using a light microscope to observe the basidiospores (100×), basidia, and hyphae. Scanning electron microscopy (SEM) was conducted with a TESCAN CLARA Xplore 30 operating at 2 keV. Forty basidiospores of each specimen were randomly selected for measurement. The spore size is expressed in the form (a–) b–c (– d), where a and d are the minimum and maximum dimensions of spores, respectively, and b and c encompass the majority of the spore dimensions. The abbreviation [n/m/p] refers to n spores measured from m basidiomata of p specimens. In addition, the Q value represents the length: width ratio of basidiospores, and the Qm value is the average Q ± standard deviation.

DNA extraction, PCR amplification, and sequencing

Genomic DNA was extracted from dried specimens using the EZup Column Fungal Genomic DNA Extraction Kit (Sangon Biotech, Shanghai, China). A 20 mg sample of a dried specimen was ground to powder in liquid nitrogen in accordance with the manufacturer’s instructions. The primer pairs ITS4/ITS5 and LR5/LR0R were used to amplify the ITS and LSU regions, respectively (Vilgalys and Hester 1990; White et al. 1990; Gardes and Bruns 1993). The PCR amplification reactions were performed on an Eppendorf Mastercycler thermal cycler in a 25 µL volume containing 1 µL DNA, 2 µL primers, 9.5 µL ddH2O, and 12.5 µL 2× Es Taq Master Mix. The amplification procedure consisted of pre-denaturation at 94 °C for 4 min, then 32 cycles comprising denaturation at 94 °C for 40 s, annealing at 55 °C for 40 s, and extension at 72 °C for 1 min, followed by a final extension at 72 °C for 8 min, and held at 4 °C (Liu et al. 2022). The PCR products were separated by electrophoresis on a 1% agarose gel. An ABI 3730 DNA Analyzer (PerkinElmer Inc., USA) was used to sequence the PCR products. The newly generated sequences (seven ITS and seven LSU) were deposited in GenBank (Table 1).

Alignment and phylogenetic analysis

The newly generated sequences were aligned with publicly available ITS and LSU sequences of Scytinopogon and Trechispora species from GenBank (see Table 1 for the accession numbers and sources of previously published sequences). Sequences for the corresponding regions from single accessions of Brevicellicium olivascens K.H. Larss. and Hjortstam and Brevicellicium atlanticum Melo, Tellería, M. Dueñas and M.P. Martín were used as the outgroup and included in the ITS+LSU sequence matrix. The ITS and LSU sequences were aligned using MAFFT v7.471 with default settings of gap openings and extension penalties (Katoh and Standley 2016). The final ITS+LSU dataset comprising 151 sequences and 1663 aligned positions (77 ITS and 74 LSU). It was assembled with SEQUENCEMATRIX v1.7.8 (Vaidya et al. 2011) and used for a multimarker phylogenetic analysis. A maximum likelihood (ML) analysis was conducted with RAxML v7.2.6 (Stamatakis et al. 2005, Stamatakis 2006) using the GTR+Gamma evolutionary model (Stamatakis et al. 2008). ML bootstrapping (BS) was performed with 1000 replicates. Bayesian inference (BI) was performed using MrBayes v3.2.7 (Ronquist and Huelsenbeck 2003); analyses were run for 2,000,000 generations using four Metropolis-coupled Monte Carlo Markov chains to calculate posterior probabilities (PP). FigTree 1.4.2 (Rambaut 2012) was used to visualize the tree files, which were edited using Adobe Photoshop CS6 (Adobe Systems Inc., USA).

Table 1.

Details of the ITS and 28S rDNA sequences used for phylogenetic analyses. The sequences newly generated in this study are highlighted in bold, and all types marked with an asterisk.

Taxon Voucher GenBank No. (ITS) GenBank No. (28S) Geographical origin References
Trechispora araneosa KHL8570 AF347084 AF347084 Sweden Larsson et al. (2004)
T. bambusicola CLZhao3302 MW544021 MW520171 China Zhao et al. (2021)
T. bambusicola He3381 OM523405 OM339227 China Liu et al. (2022)
T. chaibuxiensis He5072 OM523408 OM339230 China Liu et al. (2022)
T. chaibuxiensis LWZ2017081434 OM523409 OM339231 China Liu et al. (2022)
T. copiosa* AMO422 MN701013 MN687971 Brazil Meiras-Ottoni et al. (2021)
T. copiosa AMO427 MN701015 MN687973 Brazil Meiras-Ottoni et al. (2021)
T. copiosa AMO453 MN701018 MN687975 Brazil Meiras-Ottoni et al. (2021)
T. confinis KHL11064 AF347081 AF347081 Sweden Larsson et al. (2004)
T. confinis LWZ2021092023b OM523414 OM339235 China Liu et al. (2022)
T. constricta He5899 OM523417 OM339236 China Liu et al. (2022)
T. constricta LWZ2021092430a OM523418 OM339237 China Liu et al. (2022)
T. caulocystidiata* FLOR56314 MK458772 Brazil Furtado et al. (2021)
T. crystallina LWZ201707292 OM523419 OM339238 China Liu et al. (2022)
T. crystallina LWZ 201710137 OM523420 OM339239 Vietnam Liu et al. (2022)
T. dimitiella Dai21181 OK298493 OK298949 China Liu et al. (2022)
T. dimitiella* Dai 21931 OK298492 OK298948 China Liu et al. (2022)
T. dealbata FLOR56183 MK458777 Brazil Furtado et al. (2021)
T. fimbriata He 4873 OM523424 OM339243 China Liu et al. (2022)
T. fimbriata He 6134 OM523425 OM339244 China Liu et al. (2022)
T. fissurata He6190 OM523427 OM339245 China Liu et al. (2022)
T. fissurata He6322 OM523428 OM339246 China Liu et al. (2022)
T. foetida* FLOR56315 MK458769 Brazil Furtado et al. (2021)
T. farinacea KHL 8451 AF347082 AF347082 Sweden Larsson et al. (2004)
T. farinacea KHL 8454 AF347083 AF347083 Larsson (2001)
T. gelatinosa AMO824 MN701020 MN687977 Brazil Meiras-Ottoni et al. (2021)
T. gelatinosa* AMO1139 MN701021 MN687978 Brazil Meiras-Ottoni et al. (2021)
T. havencampii* SFSUDED8300 NR154418 NG059993 Africa Desjardin and Perry (2015)
T. hymenocystis KHL16444 MT816397 MT816397 Norway Meiras-Ottoni et al. (2021)
T. hymenocystis KHL8795 AF347090 AF347090 Sweden Larsson et al. (2004)
T. khokpasiensis* MMCR00009 MZ687107 MZ683197 Thailand Sommai S et al. (2023)
T. latehypha He5848 OM523446 OM339262 Sri Lanka Liu et al. (2022)
T. latehypha LWZ2017061116 OM523447 OM339263 China Liu et al. (2022)
T. longiramosa HG140168 OM523448 OM339264 China Liu et al. (2022)
T. longiramosa CH 19233 OM523449 China Liu et al. (2022)
T. laxa MHHNU10379 OP959649 OP954660 China This study
T. laxa * MHHNU10714 OP959650 OP954661 China This study
T. malayana Dai17876 OM523452 OM339265 Singapore Liu et al. (2022)
T. malayana He4156 OM523453 OM339266 Thailand Liu et al. (2022)
T. minispora AM170 MK328885 MK328894 Mexico Yuan et al. (2020)
T. minispora AM176 MK328886 MK328895 Mexico Yuan et al. (2020)
T. mollusca Dai 6191 OM523455 OM339269 China Liu et al. (2022)
T. mollusca Dai 11085 OM523457 OM339270 China Liu et al. (2022)
T. nivea LWZ201808043 OM523461 OM339273 China Liu et al. (2022)
T. nivea MAFungi74044 JX392832 JX392833 Telleria et al. (2013)
T. papillosa AMO713 MN701022 MN687979 Brazil Meiras-Ottoni et al. (2021)
T. papillosa AMO714 MN687980 Brazil Meiras-Ottoni et al. (2021)
T. papillosa* AMO795 MN701023 MN687981 Brazil Meiras-Ottoni et al. (2021)
T. pallescens FLOR56184 MK458767 Brazil A.N.M. Furtado et al. (2021)
T. pallescens FLOR56188 MK458774 Brazil A.N.M. Furtado et al. (2021)
T. aff. pallescens RL 115 MK328887 MK328896 Mexico Unpublished
T. aff. pallescens RL 132 MK328889 MK328898 Mexico Unpublished
T. aff. pallescens RL 133 MK328890 MK328899 Mexico Unpublished
T. robusta FLOR 56179 MK458770 Brazil A.N.M. Furtado et al. (2021)
T. sanpapaoensis MMCR00124.1 MZ687109 MZ683200 Thailand Sommai S et al. (2023)
T. saluangensis* MMCR00260 MZ687104 MZ683201 Thailand Sommai S et al. (2023)
T. saluangensis MMCR00261 MZ687105 MZ683202 Thailand Sommai S et al. (2023)
T. scabra FLOR56189 MK458773 Brazil A.N.M. Furtado et al. (2021)
T. sinensis He3714 OM523464 OM339274 China Liu et al. (2022)
T. sinensis He4314 OM523465 OM339275 China Liu et al. (2022)
T. stevensonii MAFungi70645 JX392843 JX392844 Telleria et al. (2013)
T. stevensonii MAFungi70669 JX392841 JX392842 Telleria et al. (2013)
T. subfissurata LWZ2019061348 OM523491 China Liu et al. (2022)
T. subfissurata He3907 OM523490 OM339298 China Liu et al. (2022)
T. khokpasiensis MHHNU07529 ON897819 ON898005 China This study
T. khokpasiensis MHHNU10662 ON897822 ON898008 China This study
T. khokpasiensis MHHNU10670 ON897823 ON898009 China This study
T. thailandica* He4101 OM523499 OM339307 Thailand Liu et al. (2022)
T. thailandica He 4112 OM523500 OM339308 Thailand Liu et al. (2022)
T. thelephora URM85757 MH280001 Brazil Chikowski et al. (2020)
T. thelephora URM85758 MH280002 Brazil Chikowski et al. (2020)
T. torrendii* URM85886 MK515148 MH280004 Brazil Chikowski et al. (2020)
T. torrendii URM85887 MH280005 Brazil Chikowski et al. (2020)
T. tongdaoensis * MHHNU11083 OP959651 OP954662 China This study
T. tongdaoensis MHHNU11086 OP959652 OP954663 China This study
T. termitophila* AMO396 MN701025 MN687983 Brazil Meiras-Ottoni et al. (2021)
T. termitophila AMO893 MN701026 MN687984 Brazil Meiras-Ottoni et al. (2021)
T. termitophila AMO1169 MN701028 MN687986 Brazil Meiras-Ottoni et al. (2021)
T. tropica LWZ2017061314 OM523502 OM339310 China Liu et al. (2022)
T. tropica LWZ2017061316 OM523503 OM339311 China Liu et al. (2022)
Scytinopogon cryptomerioides* 0906RK10-23 OK422242 China Lin et al. (2022)
Brevicellicium atlanticum LISU178566 9065IM HE963773 HE963774 Portugal Telleria et al. (2013)
B. olivascens MAFungi23496 HE963787 HE963788 Spain Telleria et al. (2013)


Phylogenetic analyses

The phylogeny derived from the ML analysis of the concatenated ITS+LSU dataset, with both PP and BS support values, is shown in Fig. 1. The BI phylogeny (not shown) was very similar in topology and branch support to the ML tree. The ML and BI analyses resolved that the three species collected from southern China each formed a monophyletic lineage within the genus Trechispora with high statistical support values. The species of Trechispora and Scytinopogon were intermixed, which was consistent with previous phylogenetic studies (Meiras-Ottoni et al. 2021; Chikowski et al. 2020). Trechispora laxa and T. tongdaoensis were closely related to the species Trechispora havencampii (D.E. Desjardin and B.A. Perry) Meiras-Ottoni and Gibertoni, Trechispora robusta (Rick) S.L. Liu and L.W. Zhou, Trechispora foetida (A.N.M. Furtado and M.A. Neves) S.L. Liu and L.W. Zhou, Trechispora longiramosa S.L. Liu, G. He, L. Chen Shuang and L.W. Zhou, Trechispora sanpapaoensis Pinruan, Sommai and Khamsuntorn, and Trechispora termitophila Meiras-Ottoni and Gibertoni. Trechispora khokpasiensis Pinruan, Sommai and Khamsuntorn, Trechispora copiosa Meiras-Ottoni and Gibertoni, and Scytinopogon cryptomerioides were grouped in a well-supported subclade (PP 0.99, BS 86%). These results strongly supported the phylogenetic distinction of T. laxa and T. tongdaoensis from other species within the Trechispora clade.

Figure 1. 

Phylogenetic relationships of Trechispora species inferred from a concatenated ITS and LSU sequence dataset under the maximum likelihood optimality criterion. Bayesian posterior probabilities (PP) > 0.95 and bootstrap values (BS) >70% are reported at the nodes (PP/BS); “–” indicates that the support value was less than the respective threshold. The two newly described species and one newly recorded species from China are highlighted in bold.


Trechispora khokpasiensis Pinruan, Sommai & Khamsuntorn

Figs 2, 3, 4


Clavarioid, scattered or fascicled, 25–30 mm tall, 15–36 mm broad, chalk white (1A1), slightly yellow (2A2) with age, apices white (1A1), yellowish white (3A4) when dry. Stipe single, short and flattened, 10–15 × 3–4 mm, white. Branches flattened or palmate, palmately branched from flattened stipe, dense, 4–7 mm wide, polychotomous below, dichotomous towards apices, internodes becoming gradually longer, branches 6–8 mm diam, divided 3–5 times, apices cristate or flattened, blunt, axils V-shaped. Flesh white to pale yellow, waxy. Taste and odor unrecorded.

Figure 2. 

Basidiomata of Trechispora khokpasiensis (MHHNU7529). Scale bars: 1cm.


Generative hyphae septate, clamped, interwoven, smooth, thin-walled, hyaline; tramal hyphae parallel arranged, 2–4 μm wide, smooth, thin-walled, hyaline. Subhymenial hyphae branched and wide, 3–8 μm; ampulliform septa present in the hyphae, 5−6 µm wide. Basidia: approximately 22–28 × 5.5–8 µm with four sterigmata 3–4.5 µm long, hyaline, subcylindrical to clavate, slight constriction, clamp connection in base. Cystidia absent. Basidiospores [40/4/3] 5–6 (–6.5) × 3–4 μm [Q = 1.33–1.72(1.83), Qm = 1.57 ± 0.16], ellipsoid, angular, finely verruculose or echinulate, hyaline, thin-walled, spines 0.5–1 μm long, apex slightly blunt; hilar appendage extremely small, obscured by spore ornamentation, inamyloid, contents usually uniguttulate.

Figure 3. 

Microscopic features of Trechispora khokpasiensis (MHHNU7529) a basidiospores b basidia.

Habit and distribution

Solitary to caespitose, grows on humus in broadleaf forest or grows on soil; basidiomata generally occur in summer. Known from Thailand (Sommai et al. 2023), Laos and China.

Figure 4. 

Scanning electron micrograph of basidiospores of Trechispora khokpasiensis (MHHNU7529).


Trechispora khokpasiensis is mainly characterized by chalk-white basidiomata and flattened branches. Trechispora pallescens (Bres.) Singer is easily mistaken for T. khokpasiensis in the field on account of its similar size, shape, and color. However, the two species occur in different habitats: T. khokpasiensis grows in the humus layer on soil without any plant root association. Trechispora chartacea (Pat.) Gibertoni also has flattened, narrowly spathulate branches, grayish white in age, axils U-shaped, arising from scarce white mycelia on the soil. However, T. khokpasiensis differs in that the axils are V-shaped, the basidiomata are pale yellow with age, and it grows on dead branches and leaves. Trechispora caulocystidiata is distinguished from T. khokpasiensis by having subglobose basidiospores and possessing caulocystidia.

In the present phylogenetic analyses, Scytinopogon cryptomerioides was close to T. khokpasiensis, but the two species differ in that T. khokpasiensis has relatively smaller basidia (22–28 × 6–8 μm vs. 35–42 × 5.5–6 μm in S. cryptomerioides). Trechispora copiosa has similar branches to T. khokpasiensis, but in T. copiosa the branches are moderately open and the basidia are primarily 2–4-spored.

Trechispora laxa P. Zhang & P.T. Deng, sp. nov.

MycoBank No: MycoBank No: 846842
Figs 5, 6, 7


Differs from Trechispora havencampii by the loose branches and 4-spored basidia.


China, Hainan Province, Baoting County, Qixianling Hot Springs National Forest Park, 18°70′24″N, 109°69′35″E, 300 m asl, 31 July 2021, leg. P. Zhang (holotype MHHNU10714).


laxus (Latin), loose, referring to the loose branching.


Solitary or scattered, fleshy consistency, 45–55 mm tall, 30–35 mm broad, fresh color (8B6–7), apices white when young but turning grayish purple (14B6) with age, drying pale grayish beige (4C3). Stipe single, white (1A1), 10–15 mm tall. Branches polychotomous from the stipe, dichotomous towards apices, not flattened, loose, divided 3–5 times, terminal branches relatively short and with color transitions to lilac, apices pale purple or white, acute, axils U-shaped. Taste and odor not recorded.

Figure 5. 

Basidiomata of Trechispora laxa (MHHNU10714). Scale bars: 1 cm.


Context with parallel arranged hyphae, 2–4 μm wide; generative hyphae clamped, smooth, thin-walled, hyaline, no calcium oxalate crystals. Subhymenial hyphae branched and wide, 3–8 μm; ampullate septa present at the base of the stipe, up to 6–8 μm wide. Basidia 20–26 × 7–9 µm with four sterigmata 4–5 µm long and a basal clamp connection, hyaline, subclavate, barrel-shaped. Cystidia absent. Basidiospores [40/3/2] 5–6 × 3–4 μm [Q = 1.25–1.71(1.83), Qm = 1.46 ± 0.16], ellipsoid, slightly irregular, inner side slightly concave, aculeate or finely verrucose, spines 1–1.5 μm long, apex not sharp but blunt; hilar appendage obscured by spore ornamentation; usually uniguttulate; hyaline, thin-walled, inamyloid.

Figure 6. 

Microscopic features of Trechispora laxa (MHHNU10714) a basidiospores b basidia.

Habit and distribution

Solitary or scattered, grows in soil in broadleaf forest; basidiomata generally occur in summer. Known only from the type locality in China.

Figure 7. 

Scanning electron micrograph of basidiospores of Trechispora laxa (MHHNU10714).


The branches of T. laxa are scattered, not dense, and the apices are white or lilac gray with age. In the field, T. laxa and Trechispora havencampii are similar because of their pale grayish brown coloration, but T. havencampii has dense branches, tips white and axils V-shaped, and 2-spored basidia with long sterigmata (5–9.5µm). Trechispora longiramosa differs from T. laxa by having long terminal branches, densely branched and white to honey-yellow tips. Trechispora sanpapaoensis has smaller basidia (11–26 × 5.5–11.0 μm) and a grayish brown stipe. Trechispora termitophila develops abundant basidiomata in active termite nests, but T. laxa generally grows in the soil of broadleaf forest. Trechispora foetida has a reddish brown to deep brown, flattened stipe, and branching in one plane. In comparison, T. laxa is flesh colored, turning grayish purple in the terminal branches, the branches are not flattened, the branches are not white, and the stipe is non-flat. An additional pigmented species, Trechispora robusta described from Brazil, is pale grayish with internodes irregular, branches flattened to subcylindrical, and inflated hyphae.

Trechispora tongdaoensis P. Zhang & P.T. Deng, sp. nov.

MycoBank No: MycoBank No: 846849
Figs 8, 9, 10


Differs from Trechispora termitophila by the white fruiting body and 4-spored basidia.


China, Hunan Province, Tongdao County, WanFo Mountain Nature Reserve, 26°32′54″N, 109°86′95″E, 523 m asl., 6 July 2022, leg. P. Zhang (holotype MHHNU11083).


tongdaoensis (Latin), referring to the currently known distribution of the species in Tongdao County, Hunan Province, China.


Clavarioid, gregarious to caespitose clusters, 60–90 mm tall, 30–45 mm broad, white (1A1), with pale yellow (1A3). Stipe single, 20–40 mm long, white (1A1), no change in color when dried. Branches dense, branching from the base, repeatedly dichotomous towards apices, divided 3–5 times, branches slender, 2–3 mm wide, internodes becoming gradually longer, terminal branches long and not flat, sometimes split at the tips, acute, axils V-shaped, terminal branches short, tips acute. Context pale yellow. Taste and odor not recorded.

Figure 8. 

Basidiomata of Trechispora tongdaoensis (MHHNU11083). Scale bars: 1 cm.


Context hyphae compact, 3–5.5 μm wide, subparallel arranged, cylindric; generative hyphae with clamp connections but not at every septum, thin-walled, smooth, hyaline, no calcium oxalate crystals; ampullate septa present in the hyphae of the stipe, 7–8 µm wide. Basidia: approximately 18–28 × 6–8 µm with four sterigmata 3–5 µm long, hyaline, subclavate to cylindrical, with clamp connection in base. Cystidia absent. Basidiospores [40/3/2] 4–6(6.5) × 3–5.5 μm [Q = 1.25–1.57(1.83), Qm = 1.50 ± 0.11] ellipsoid, slightly angular, tuberculate or coarsely echinulate, spines 0.5–1 μm long, blunt; hilar appendage ambiguous by spore ornamentation, sometimes contents uniguttulate, inamyloid.

Figure 9. 

Microscopic features of Trechispora tongdaoensis (MHHNU11083) a Basidiospores b Basidia.

Habit and distribution

Caespitose or gregarious on the soil of broadleaf forests; basidiomata generally occur in summer. Known only from the type locality in China.

Figure 10. 

Scanning electron micrograph of basidiospores of Trechispora tongdaoensis (MHHNU11083).


The fruiting body of T. tongdaoensis has a long stalk, 20–40 × 4–6 mm, the terminal branches are long, bifurcate, the tips are white, and the branches are not flattened in a plane. Trechispora foetida differs from T. tongdaoensis by having reddish brown to deep brown basidiomata, and flattened branches and stipe. Trechispora caulocystidiata has cystidia in the stipitipellis as caulocystidial hairs. This feature is obvious under a microscope, but we failed to observe this structure in T. tongdaoensis. In addition, the T. caulocystidiata stipe is relatively shorter (10–20 × 3–5 mm vs. 20–40 × 4–6 mm in T. tongdaoensis). Trechispora copiosa differs from T. tongdaoensis in having acute or flattened branch tips and T. tongdaoensis has relatively smaller spores (4–6 (–6.5) × 3–5.5 μm vs. (5–) 5.5–6.5 (–7) × (3–) 3.5–4 (–4.5) μm in T. copiosa). Trechispora dealbata was classified in Ramariopsis (Donk) Corner based on the gelatinous context by Petersen (1984, 1988), whereas T. tongdaoensis lacks a gelatinous context and differs in having relatively smaller spores (4.0–4.5 × 2.5–3.5 µm vs. 4–6(6.5) × 3–5.5 μm in T. tongdaoensis). In the field, T. gelatinosa has a fleshy or gelatinous consistency, translucent when fresh, small basidia (12–26 × 5–6.5 μm), and small basidiospores (3–) 3.2–4.5 (–5) × (2–) 2.5–3.5 (–4) μm, which distinguishes the species from T. tongdaoensis. In the present phylogenetic analyses, T. tongdaoensis clustered with pigmented species.


Trechispora has until recently been considered to encompass a variety of morphological characteristics and broad diversity in hymenophore structure ranging from smooth, grandinioid to odontioid, hydnoid, or poroid, but always resupinate basidiomata. Scytinopogon is characterized by having clavarioid basidiomata and flattened branches, and the basidiomata are mostly white in color with a tough texture. Over time, additional pigmented species of Scytinopogon have been described and some species have rounded instead of flattened branches. A relationship between the flattened coralloid Scytinopogon and the resupinate Trechispora was first suggested by Jülich (1981), who placed the genera in separate families of the order Trechisporales, Hydnodontales. Subsequently, Meiras-Ottoni et al. (2021) analyzed a number of sequences from specimens of each genus, including the type species of each genus, and concluded that Scytinopogon is a synonym of Trechispora. In phylogenetic reconstructions, several transitions between fruiting body types are indicated to have occurred (Meiras-Ottoni et al. 2021). In addition, the color of Trechispora species has changed many times over the course of evolution, from white to pigmented, but the evolutionary trend for the basidiomata is not understood. The transition from resupinate basidiomata to clavarioid basidiomata seems to represent an evolutionary trend. The effect of this transformation is presumably an increase to expand the surface area of the hymenophore. The driving force behind morphological differences in fruiting bodies seems likely to be associated with selection for efficient spore dispersal (Hibbett and Binder 2002). In micromorphology, variation in basidiospores (smooth or ornamented) and basidia (2 or 4 sterigmata) represents different evolutionary directions, but the effects of these differences remain unknown. Most species of Trechispora are distributed in the tropics or subtropics, including coralloid species. Indeed, the two new species described in this study were collected in the subtropics of southern China. However, the current distribution of stipitate and clavarioid species does not include temperate regions. To comprehend the diverse factors contributing to the evolution in Trechispora, a number of specific aspects should be considered: substrate, nutritional mode, and environmental conditions. The ITS region is highly variable among Trechispora species and cannot be aligned reliably within the genus (Meiras-Ottoni et al. 2021), such that other genetic markers (e.g., LSU, SSU, ef1, rpb1, and rpb2) should be used to for phylogenetic analyses in Trechispora.

In China, previous studies have reported one new coralloid species of Trechsipora (T. longiramosa; Liu et al. 2022) and three species of Scytinopogon (S. cryptomerioides, S. echinosporus (Berk. and Broome) Corner and S. pallescens; Zhang and Yang 2003, Lin et al. 2022). The present phylogenetic analysis confirmed that S. cryptomerioides was firmly nested within Trechispora. Together with the previous studies mentioned here, this finding highlights that further exploration for coralloid species of Trechispora is needed. The present study expands our understanding of clavarioid species of Trechispora by providing descriptions and illustrations for two new species. The findings enrich our knowledge of the distribution of coralloid Trechispora species in China and the overall diversity of Trechispora.

Key to coralloid Trechispora species in China

1 Basidiomata pure white to pale yellow 2
Basidiomata grayish brown to pale purple 4
2 Basidiomata with no flattened branches T . tongdaoensis
Basidiomata with flattened branches 3
3 Basidiomata only grow in soil T. pallescens
Basidiomata grow in the humus layer on soil T . khokpasiensis
4 Basidiomata with dense branches and long terminal branches T. longiramosa
Basidiomata with loose branches T . laxa


We thank Robert McKenzie, PhD, from Liwen Bianji (Edanz) ( for editing the English text of a draft of this manuscript.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.


This study was financially supported by the Key Research and Development Program of Hunan Province (No. 2020SK2103)

Author contributions

Conceptualization: Ping Zhang; methodology: Peng-Tao Deng and Jun Yan; performing the experiment: Peng-Tao Deng, Xiang-Fen Liu; resources: Ping Zhang, Peng-Tao Deng, Jun Yan, Ming-Xin Lu and Yuan Lin; writing – original draft preparation: Peng-Tao Deng; writing – review and editing: Ping Zhang and Zheng-Mi He; supervision: Ping Zhang; project administration: Ping Zhang; funding acquisition: Ping Zhang. All authors have read and agreed to the published version of the manuscript.

Author ORCIDs

Peng-Tao Deng

Jun Yan

Zheng-Mi He

Ping Zhang

Data availability

The sequence data generated in this study are deposited in NCBI GenBank.


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Supplementary material

Supplementary material 1 

Multiple sequence alignment

Peng-Tao Deng, Jun Yan, Xiang-Fen Liu, Zheng-Mi He, Yuan Lin, Ming-Xin Lu, Ping Zhang

Data type: fasta file

This dataset is made available under the Open Database License ( The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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