Two new agaricoid species of the family Clavariaceae (Agaricales, Basidiomycota) from China, representing two newly recorded genera to the country
expand article infoMing Zhang, Chao-Qun Wang, Tai-Hui Li
‡ Guangdong Institute of Microbiology, Guangdong Academy of Sciences, Guangzhou, China
Open Access


Two new lamellar species, Camarophyllopsis olivaceogrisea and Hodophilus glaberripes, of the family Clavariaceae were discovered in the subtropical zone of China. Camarophyllopsis olivaceogrisea is morphologically characterized by its hygrophanous basidiomata, greenish gray to dull green pileus, shortly decurrent lamellae, broadly elliptic basidiospores 4–5.5 × 3.5–4.5 μm in size, and cutis-like pileipellis composed of cylindrical cells. Hodophilus glaberripes is mainly characterized by its white to brownish pileus, glabrous stipe, slight yam bean smell, broadly elliptic basidiospores 5–6.5 × 4–5 μm in size, and epithelium-like pileipellis composed of inflated cells. Phylogenetic placement of the two species was determined by the combined analyses of a DNA data matrix containing ITS and LSU, and showed that collections of the two species formed two independent lineages in the Camarophyllopsis and Hodophilus clades respectively. The delimitation of C. olivaceogrisea and H. glaberripes were evaluated using molecular, morphological, and ecological methods. This is the first report of the genera Camarophyllopsis and Hodophilus in China.


Camarophyllopsis, Hodophilus, phylogenetic analysis, subtropical zone, taxonomy


Clavariaceae Chevall. (Agaricales, Basidiomycota) is a genetically monophyletic but morphologically diverse family. Members of this family show variations in the macromorphology of their sporocarps and are pendant-hydnoid, cylindrical, clavate, coralloid, resupinate, pileate, or lamellate-stipitate (Larsson et al. 2004; Dentinger and McLaughlin 2006; Matheny Curtis et al. 2006; Larsson 2007; Birkebak et al. 2013). Generally, Clavariaceae was known as a coral fungal group, including club-like (clavarioid) genera such as Clavaria Vaill. ex L., and Clavulinopsis Overeem, and branch-form (coralloid) genera, such as Ramariopsis (Donk) Corner (Birkebak et al. 2013, 2016). However, some agaricoid genera, such as Camarophyllopsis Herink, Hodophilus R. Heim ex R. Heim and Lamelloclavaria Birkebak & Adamčík, showed phylogenetic affinity within Clavariaceae (Matheny Curtis et al. 2006; Birkebak et al. 2013), thus have been added to the family based on phylogenetic analyses in recent years (Birkebak et al. 2016).

Camarophyllopsis can be easily distinguished from other genera in the family by its small agaricoid basidiomata, hygrophanous pileus, subglobose to broadly ellipsoidal basidiospores, and epithelium pileipellis composed of chains of erect, ascending or repent, subcylindrical to ellipsoid terminal cells (Arnolds 1986; Young 2005). Hodophilus differs from Camarophyllopsis in the hymeniderm pileipellis composed of typically perpendicular, broadly inflated, globose, and obpyriform to sphaero-pendunculate terminal elements (Birkebak et al. 2016). Lamelloclavaria can be distinguished from the above genera by its rimulose non-hygrophanous pileus and cutis pileipellis (Birkebak et al. 2016).

In this study, some agaricoid collections were identified in China. Morphological observation and phylogenetic analyses confirmed that they are two novel taxa in the genera Camarophyllopsis and Hodophilus. This is the first report of these two genera in China.

Materials and methods

Morphological studies

Photographs of basidiomata were taken in type localities when collected. Macro-morphological characteristics were recorded for fresh specimen. Specimens were dried and then deposited in the Guangdong Institute of Microbiology (GDGM). Methods used for morphological descriptions were followed Zhang et al. (2015). Colors were recorded and described in general terms according to the method of Kornerup and Wanscher (1978). Microstructures were observed from rehydrated materials, and the notation “basidiospores (n/m/p)” indicates that the measurements were conducted for n basidiospores from m basidiomata of p collections. Line drawings were prepared by free hand.

DNA extraction, PCR amplification, and sequencing

Total genomic DNA of each voucher specimen was extracted from silica-gel-dried materials using the Sangon Fungus Genomic DNA Extraction kit (Sangon Biotech, Shanghai, China) according to the manufacturer’s instructions. Primer pairs ITS1/ITS4 (White et al. 1990) and LR0R/LR5 (Vilgalys and Hester 1990) were used to amplify the internal transcribed spacer (ITS) region and the large subunit nuclear ribosomal RNA (nrLSU) region, respectively. PCR protocol and sequencing were conducted following the method of Zhang et al. (2015).

Phylogenetic analyses

Newly generated sequences, related sequences used in previous studies (Adamčík et al. 2018) and a few sequences retrieved from GenBank by a Blast search were used to reconstruct phylogenetic trees. Detailed information on the newly sequenced samples, including the taxon names, voucher numbers, localities and GenBank accession numbers, is shown in Table 1.

Information on newly generated DNA sequences used in this study.

Taxon Voucher Country ITS LSU
Camarophyllopsis olivaceogrisea GDGM44497 China MK894563 MK894551
GDGM44519 China MK894564 MK894552
Camarophyllopsis sp. GDGM44501 China MK894565 MK894553
Hodophilus glaberripes GDGM45940 China MK894566 MK894554
GDGM52374 China MK894567 MK894555
GDGM52530 China MK894568
GDGM52545 China MK894569 MK894556
GDGM52583 China MK894570 MK894557
GDGM55689 China MK894571 MK894558
GDGM70329 China MK894572 MK894559
GDGM70331 China MK894573 MK894560
GDGM72434 China MK894574 MK894561
GDGM72518 China MK894575 MK894562

ITS and LSU sequences were respectively aligned using Clustal X v1.81 (Thompson et al. 1997) and manually modified where necessary in Bioedit v7.0.9 (Hall 1999), and a combined matrix of ITS and LSU sequences was obtained. The combined dataset was then analyzed using RAxML v7.2.6 (Stamatakis 2006) and MrBayes v3.1.2 software (Ronquist and Huelsenbeck 2003) for maximum likelihood (ML) and Bayesian inference (BI) analyses, respectively. For both BI and ML analyses, the substitution model for the two gene partitions was individually determined using the Akaike Information Criterion (AIC) complemented in MrModeltest v2.3 (Nylander 2004). For ML analysis, all parameters were kept at default values, except for choosing the GTRGAMMAI model, and statistical support was obtained using rapid nonparametric bootstrapping with 1000 replicates. BI analysis using selected models and 4 chains was conducted by setting the number of generations to 3 million and stoprul command with the stopval value set to 0.01. Trees were sampled every 100 generations. The first 25% of generations were discarded as burn-ins and posterior probabilities (PP) were calculated from the posterior distribution of the retained Bayesian trees.


Molecular phylogenetic results

For phylogenetic analyses, 25 sequences (13 ITS and 12 LSU) were newly generated from 13 collections, 219 related sequences (123 ITS and 96 LSU) were retrieved from GenBank, and Ramariopsis corniculata (Schaeff.) R.H. Petersen selected as an outgroup based on previous studies (Birkebak et al. 2016; Adamčík et al. 2016, 2018). The combined matrix of 137 samples with 1614 nucleotide sites was constructed for phylogenetic analyses and the final alignment was submitted to TreeBASE (Submission ID 24440). SYM+I and SYM+G were chosen as the best substitution models for ITS and LSU, respectively. The ML and BI analyses generated nearly identical tree topologies with minimal variation in statistical support values; thus, a ML tree was selected for display (Figure 1).

Figure 1. 

Phylogenetic placements of Camarophyllopsis olivaceogrisea and Hodophilus glaberripes inferred from the combined ITS and LSU dataset using RAxML. Ramariopsis corniculata was selected as an outgroup. The lineages with new species were shown in bold. BS ≥ 50% and PP ≥ 0.90 were indicated around the branches.

The tree topologies generated in this study are similar to those obtained by Adamčík et al. (2018) and are therefore not described in detail here, except for the results relevant to the two new species. The two monophyletic genera Camarophyllopsis and Hodophilus were highly supported (Figure 1). Two collections (GDGM44497 and GDGM44519) formed an independent lineage with strong statistical support (BS = 100, PP = 1), located within the Camarophyllopsis clade, and presented as a sister group to a collection numbered as GDGM44501 from China (with low statistical support). Ten collections (GDGM45940, GDGM52374, GDGM52530, GDGM52545, GDGM52583, GDGM55689, GDGM70329, GDGM70331, GDGM72434 and GDGM72518) were grouped together with strong statistical supports (BS = 100, PP = 1) and formed an independent lineage in the Hodophilus clade, and were revealed as a sister group to H. indicus K.N.A. Raj, K.P.D. Latha & Manim. with significant statistical support (BS = 100, PP = 1).


Camarophyllopsis olivaceogrisea Ming Zhang, C.Q. Wang & T.H. Li, sp. nov.

MycoBank No: MB831122
Figs 2a–b, 3


The epithet “olivaceogrisea” refers to the olive-gray pileus color.


This new species is morphologically distinguished from other taxa in the genus by its smaller basidiomata, greenish gray to dull green pileus, white and short decurrent lamellae, and broadly elliptic basidiospores.


CHINA. Guizhou Province: Leishan County, Leigongshan National Nature Reserve, alt. 1260 m, 22 July 2014, M. Zhang (holotype: GDGM44519!).


Basidiomata small-sized. Pileus 7–12 mm broad, hemispherical, convex to plano-convex at first, then gradually applanate, becoming depressed at disc when mature, non-striate to weakly striate; margin slightly inflexed at first, soon straight, slightly crenate or lacerate when mature; surface matt, velvety, hygrophanous, greenish gray (or olive gray) to dull green (27D 2–30D2, 27D3–30D3, 27E2–30E2, 27E3–30E3), often paler at margin. Flesh 1–3 mm thick in the stipe, white to grayish white, unchanging when exposed. Lamellae 1–2 mm deep, L = 20–34, l = 1–3, short to moderately decurrent, white to weakly grayish white (1A1–1B1) at first, white to weakly greenish white (27A2–30A2) when mature, unchanging when bruised; edge entire, concolorous with the sides. Stipe 13–25 × 1.5–2.5 mm, central, cylindrical and becoming narrower downwards; glabrous and shiny, hardly tomentulose or pruinose, hygrophanous, white to greenish white at first (28A1–28A2, 29A1–29A2), becoming greenish white to light greenish gray (28A2–28B2, 29A2–29B2) when mature and in dry condition. Odor none. Taste mild.

Basidiospores [60/2/2] 4–5.5(–6) × 3.5–4.5(–5) μm, av. 4.6 × 3.8 μm, Q = (1.12)1.14–1.28(1.43), av. Q = 1.21 ± 0.08, broadly ellipsoid, hyaline, smooth, inamyloid, thin-walled. Basidia 4-spored, occasionally 2-spored, (10–)15–26(–30) × 5–7 μm, av. 24.5 × 5.8 μm, hyaline, narrowly clavate, attenuated and flexuous toward base, sterigmata up to 4 μm long. Basidioles cylindrical to narrowly clavate, often flexuous, obtuse, (18–)20–37.5(–40) × 6–8 μm, av. 23 × 6.8 μm. Pleurocystidia absent. Marginal cells on the lamellar edges not well differentiated, similar to basidioles on lamellar sides. Lamellar trama composed of sub-parallel or occasionally interwoven and irregularly inflated hyphae (23–)35–50(–104) × 4–8(–10) μm, av. 46.5 × 7 μm. Pileipellis a cutis of numerous repent branched hyphal 4–8 μm wide, with terminal chains of ellipsoid or cylindrical cells. Pileus trama composed of cylindrical and occasionally branched hyphae (23–)35–50(–70) × (4–)6–10 μm, av. 45.5 × 7.6 μm. Stipitipellis formed of parallel, thin-walled and narrow hyphae 3–8 μm diam. Caulocystidia not observed. Clamp connections absent in all tissues.

Figure 2. 

a, b Basidiomata of Camarophyllopsis olivaceogrisea (GDGM44519, holotype) c–f basidiomata of Hodophilus glaberripes (e. GDGM72518, holotype). Scale bars: 20 mm (a, b); 50 mm (c–f).

Figure 3. 

Camarophyllopsis olivaceogrisea. a Basidiospores b basidioles c Basidia d hyphal terminations in pileipellis e stipitipellis. Scale bars: 10 μm (a–c); 20 μm (d, e).

Figure 4. 

Hodophilus glaberripes. a Basidiospores b basidia c basidioles d hyphal terminations in pileipellis e hyphal terminations in stipitipellis. Scale bars: 10 μm (a–c); 20 μm (d, e).

Habit, ecology and distribution

Solitary, scattered on soil in mixed forests; currently only known from the Guizhou Province of China.

Additional specimens examined

CHINA. Guizhou Province: Leishan County, Leigongshan National Nature Reserve, alt. 1120 m, 22 July 2014, J. Xu (GDGM44497).

Hodophilus glaberripes Ming Zhang, C.Q. Wang & T.H. Li, sp. nov.

MycoBank No: MB831125
Figs 2c–f, 4


The epithet refers to the glabrous stipe.


This species is easily distinguished from other species in the genus Hodophilus by its larger basidiomata, white, brownish orange to brown pileus, glabrous stipe, slightly yam bean smell and broadly elliptic basidiospores.


CHINA. Guangdong Province: Shaoguan City, Danxiashan National Nature Reserve, alt. 240 m, 10 May 2018, M. Zhang (holotype: GDGM72518!).


Basidiomata small to medium-sized. Pileus 15–50 mm broad, hemispherical, convex to plano-convex at first, then becoming broadly convex or plano-convex but hardly fully expanded to plane, often depressed at disc when old; white to yellowish white at first, then gradually becoming orange white, pale yellow, pale orange, brownish orange, light brown, brown to reddish brown (5A2, 3A3–5A3, 5C4–7C4, 5D5–9D5) when mature and dry, hygrophanous; margin first slightly inflexed, soon straight, slightly crenate when mature, non-striate or indistinctly translucently striate up to one third when wet; surface matt, velvety and later with fine and darker granules or pruina, at first even, but becoming rugose or rough towards the center when mature, often concentrically cracked in dry conditions. Flesh 0.5–2 mm thick in half radius of the pileus, white, pinkish white to pale beige; Lamellae 3–5 mm deep, distant to subdistant, L = 21–32, l = 1–3, short decurrent, notched, orange white to pinkish white (5A2–10A2) when young, brownish orange, light brown, reddish brown to brownish red (5C4–7C4, 5D6–10D6) when mature, unchanging when bruised; edge entire, concolorous or slightly paler than lamella sides. Stipe (50) 80–100 × 3–5 mm, cental, usually flexuous, cylindrical and slightly narrower downwards; glabrous smooth and shiny, hygrophanous, white to yellowish white at first, becoming pale yellow to pale orange when mature and in dry condition. Odor none or slight yam bean smell, taste mild.

Basidiospores [210/9/9] (4.5)5–6.5(7) × 4–5(5.5) μm, av. 5.9 × 4.7 μm, Q = (1.0)1.11–1.37(1.4), av. Q = 1.20 ± 0.11, broadly ellipsoid to subglobose, hyaline, smooth, inamyloid, thin-walled. Basidia 4-spored, occasionally 2-spored, (32–) 36–46(–66) × (4–)4.5–6(–7) μm, av. 39.5 × 5.9 μm, tenuated and flexuous towards base, with sterigmata up to 7 μm long. Basidioles cylindrical to narrowly clavate, obtuse, often flexuous, (31–)34–42(–60) × (4–)6–8(–10) μm, av. 40.5 × 6.9 μm. Pleurocystidia absent. Marginal cells on the lamellar edges usually not well differentiated, similar to basidioles on lamellar sides. Lamellar trama composed of sub-parallel or occasionally interwoven and irregularly inflated branched hyphae with elongate cells (38–)52–98(–160) × (4–)6–14(–20) μm, av. 88 × 9.5 μm. Subhymenium poorly developed. Pileipellis a transition from hymeniderm to epithelium, with hyphal elements 3–10 μm wide, thin-walled, hyaline, terminations usually composed of 1–3 inflated cells; terminal cells obpyriform, subglobous or ellipsoid, rarely sphaero-pedunculate or broadly clavate, (15–)19–46(–50)× (7–)12–22(–30) μm, av. 38.5 × 18 μm. Pileus trama composed of subparallel hyphae (34–)46–89(–130) × (4–)5.5–10 μm, av. 74 × 7.6 μm. Stipitipellis formed of parallel, thin-walled and narrow hyphae 3–6 μm diam. Caulocystidia usually in dense fascicles or patches, thin-walled, repent or ascending; with terminal cells mainly clavate, occasionally subcapitate or obpyriform, obtuse, often pedicellate and flexuous, measuring (18–)22–53(–60)× (4–)5.5–13 μm, av. 43 × 7.5 μm. Clamp connections absent in all tissues.

Habit, ecology and distribution

Solitary, scattered on soil in broadleaf forests and mixed forests; currently only known from China.

Additional specimens examined

CHINA. Guangdong Province: Huizhou City, Xiangtoushan National Nature Reserve, alt. 640 m, 18 May 2016, H. Huang (GDGM45940); Shaoguan City, Nanling National Nature Reserve, 800 m, 29 July 2017, M. Zhang (GDGM70329 and GDGM70331); Shaoguan City, Danxiashan National Nature Reserve, 200 m, 27 April 2019, X.R. Zhong (GDGM76367 and GDGM76337), J.P. Li (GDGM76300); Jiangxi Province: Jinggangshan Botanical Garden, 884 m, 20 June 2016, H. Huang (GDGM52374), Z.P. Song (GDGM52545); same location, 21 June 2016, Z.P. Song (GDGM52530 and GDGM52583); Hunan Province: Chenzhou City, Jiulongjiang National Forest Park, alt. 230 m, 14 May 2018 X. R. Zhong (GDGM55689).


Key to the species of Camarophyllopsis and Hodophilus

1 Basidiomata agaricoid; pileipellis an epithelium composed of chains of subcylindrical to ellipsoid terminal elements 2 Camarophyllopsis
Basidiomata agaricoid; pileipellis a hymeniderm composed of broadly inflated, globose, and obpyriform to sphaero-pendunculate terminal elements 12 Hodophilus
2 Pileus diameter usually < 30 mm 3
Pileus diameter usually ≥ 30 mm 11
3 Pileus hygrophanous or subhygrophanous 4
Pileus not hygrophanous 5
4 Pileus greenish gray to dull green; lamellae white, decurrent; basidiospores 4–5.5 × 3.5–4.5 μm C. olivaceogrisea
Pileus rugulose, buffy brown to dark hazel; lamellae decurrent, pinkish to hazel; stipe olive brown or grayer; basidiospores globose av. 4 × 5 μm C. rugulosoides
5 Lamellae always with pink tinct, pale pink to pink, decurrent; pileus pale pink to whitish, matt; stipe pale pink to whitish; basidiospores av. 7 × 4.5 μm C. roseola
Lamellae without pink tinct 6
6 Lamellae white, unchanging when mature 7
Lamellae whitish to grayish or brownish 9
7 Basidiospores < 6 μm; pileus brownish to dark brown, subtomentosus, depress in central when mature; stipe concolorous with pileus or slightly faded C. atrovelutina
Basidiospores usually ≥ 6 μm 8
8 Pileus gray, manifestly cleaving at margin when dry; stipe white; basidiospores 5.5–8.5 × 4–5.5 μm C. leucopus
Pileus gray, sub-velvety; stipe pale gray; basidiospores 7–8 × 5.5–6.5 μm C. tetraspora
9 Basidiospores ≥ 7 μm; pileus fuliginous brown, subfibrillose to fibrillose, infundibuliform; lamellae deep decurrent; stipe brown C. araguensis
Basidospores < 7 μm 10
10 Pileus dark brown, subtomentosus; lamellae light grayish, decurrent; stipe white, smooth, basidiospores 5–6.5 × 4–5 μm C. albipes
Pileus yellowish cinnamon to brownish cinnamon or chocolate gray, silky-tomentose or velvet; lamellae decurrent, whitish to grayish or brownish; stipe concolorous with pileus; basidiospores 4–5 × 4–4.5 μm C. schulzeri
11 Pileus brownish gray to grayish brown; lamellae adnate to subdecurrent, white; stipe smooth, white; basidiospores 5–7 × 3.5–4.5 μm C. pedicellata
Pileus pearl gray to drab, often rivulose-cracking; lamellae light gray; stipe light gray, slightly longitudinally fibrillose-striped; basidiospores 5–7 × 3.5–5.2 μm C. dennisiana
12 Basidiomata with a napthalene or an unpleasant odor see Adamčík et al. (2016, 2017)
Basidiomata without napthalene odor 13
13 Stipe with dark dots on surface 14
Stipe never with dark dots on surface 16
14 Lamellae white to grayish white, pileus pale brown; basidiospores 4.5–6 × 4.0–5 μm, pileipellis an epithelium composed of globose to pyriform elements C. kearneyi
Lamellae with orange, brown or light brown tinct 15
15 Pileus yellowish gray to brown when fresh, light brown, orange gray to beige when mature; lamellae beige or orange gray at first, changing to brownish orange to yellowish brown when mature, basidiospores 4.6–5.4 × 3.5–4.2 μm H. atropunctus
Pileus dark brown to pale brown; lamellae light brown to grayish brown when young, changing brown to dark brown when mature; basidiospores 4.8–5.4 × 3.9–4.5 μm H. variabilipes
16 Stipe with yellow tinct see Adamčík et al. (2018)
Stipe without yellow tinct 17
17 Species has south hemisphere distribution, pileus creamy buff-brown or pinkish fawn; lamellae decurrent, pale pinkish; basidiosproes 4.5–6.0 × 4.5–5.5 μm; pileipellis an epithelium of globose or pyriform elements; known from Australia C. darminensis
Species has north hemisphere distribution 18
18 Pileus yellowish white, brownish orange to reddish brown; lamellae orange white to brownish red; basidiospores 5–6.5 × 4–5 μm; known from China H. glaberripes
Pileus grayish brown to brownish orange; lamellae subdecurrent, pale orange; stipe grayish orange, glabrous; basidiospores 4–5 × 3–5 µm; known from India H. indicus

Phylogenetic relationships of the genera within Clavariaceae have been investigated in several studies, and Camarophyllopsis and Hodophilus were well supported as two independent groups at generic level (Birkebak et al. 2013, 2016; Adamčík et al. 2017, 2018). In the present study, phylogenetic analyses based on ITS and LSU showed that the two clades Camarophyllopsis and Hodophilus were well supported with high phylogenetic values (BS/BPP = 100/1), and collections from China formed two strongly supported terminal branches in the two clades. The sequences generated in this study did not match any previously described sequences, validating with strong support the recognition of C. olivaceogrisea and H. glaberripes as two distinct species based on their phenotypic features.

According to the phylogram (Figure 1), C. olivaceogrisea nested well into the Camarophyllopsis clade and formed a sister group of an unidentified Chinese collection (GDGM44501) with low statistical support. Because of the low number of specimens, GDGM44501 was not described here, but it can be easily separated from C. olivaceogrisea by branch distance. The other three species in the phylogenetic tree, C. atrovelutina, C. deceptive and C. schulzeri, also can be easily separated from C. olivaceogrisea. The closest relatives of the new species remain unresolved in this study because of the lack of significant statistical supports and the few available sequences of Camarophyllopsis used in phylogenetic analysis.

In the Hodophilus clade, H. glaberripes is closely related to H. indicus K.N.A. Raj, K.P.D. Latha & Manim., and together formed a well-supported branch, which is a sister clade to the yellow stipe clade (or H. micaceus superclade) as defined by Adamčík et al. (2017, 2018), but with limited statistical support. However, the recently described Indian species, H. indicus, show smaller and brownish orange basidiomata, subdecurrent and pale orange lamellae, and slightly smaller basidiospores (4–5 × 3–5 μm) (Crous et al. 2017).

Morphologically, the most distinctive features of C. olivaceogrisea are the small basidiomata with a greenish gray to dull green pileus, white and decurrent lamellae, broadly ellipsoid basidiospores, narrowly clavate basidia, and a cutis pileipellis composed of chains of cylindrical cells. Camarophyllopsis microspora (A.H. Sm. & Hesler) Bon is similar to C. olivaceogrisea to some extent. However, C. microspora, originally reported in Michigan, differs on account of its fuscous pileus and stipe, dark grayish context and smaller basidiospores (4–4.5 × 2.5–3 μm) (Hesler and Smith 1963). Considering species with a pileus diameter of 10–30 mm, C. olivaceogrisea is similar to C. albipes and C. leucopus (Singer) Boertm. However, C. albipes mainly differs on account of its brown and subtomentose pileus, grayish lamellae with slight veins at the margin, robust stipe, and slightly larger basidiospores (5–6.5 × 4–5 μm) (Singer 1973); C. leucopus mainly differs on account of its gray and sulcate pileus and larger basidiospores (5–8.5 × 4–5.5 μm) (Singer 1973).

Hodophilus glaberripes is characterized by its larger basidiomata, hygrophanous pileus with white, brownish orange to brown color, glabrous stipe, larger and broadly elliptic basidiospores, epithelium-like pileipellis with obpyriform or subglobous terminal cells, and slightly yam bean smell. The combination of these characteristics makes H. glaberripes easily distinguishable from other members of the genus. Hodophilus glaberripes is somewhat similar to H. albofloccipes (Kovalenko, E.F. Malysheva & O.V. Morozova) Looney and Adamčík, H. anatinus Dima, Adamčík & Jančovičová, H. subfoetens Adamčík, Jančovičová & Looney, and H. pallidus Adamčík, Jančovičová & Looney in morphology. However, H. albofloccipes mainly differs by its smaller baisdiomata, ochre or ochre yellow to pale olives pileus, yellow to brownish stipe covered with white pruina or squamula, smaller basidiospores (4–5.7 × 3.5–5 μm), and naphthalene-like odor (Kovalenko et al. 2012); H. anatinus differs by its smaller basidiomata, grayish brown pileus, grayish yellow to brown stipe, and smaller basidiospores (4.8–5.5 × 3.8–4.4 μm) (Adamčík et al. 2018); H. subfoetens differs by its smaller and grayish brown to brownish black basidiomata, with a naphthalene odor, and smaller basidiospores (5–5.7 × 3.9–4.5 μm) (Adamčík et al. 2017a); H. pallidus differs by its smaller basidiomata with a strong naphthalene odor, orange-gray to grayish orange pileus, orange gray to orange brown stipe, and smaller basidiospores (5.1–5.7 × 3.9–4.6 μm) (Adamčík et al. 2017a).

Ecologically, very little is known about the ecology of Camarophyllopsis species, as for most species only a few verified collections are known and little molecular data is available. Camarophyllopsis species are widely distributed in the southern and northern hemispheres, from tropical zones to cool temperate zones, and in monsoon forest, bushy forest, and grassland habitats, and some species have been shown to be saprotrophic (Young 1999, 2005; Boertmann 2002; Kovalenko et al. 2012). Camarophyllopsis olivaceogrisea was collected from subtropical regions in southwest China at altitudes of over 1000 m, and is typically found in wet areas with moss under mixed forest, which is mainly dominated by broadleaf plants (Castanopsis spp., Fagus spp. and Schima spp.) with few conifer (Pinus massoniana Lamb. and Pinus spp.). This study expands the geographic distribution of the genus Camarophyllopsis to China.

Hodophilus taxa were mainly reported in temperate to boreal zones of the northern hemisphere and can be found in forest, bushy forest margin, grassland and bare soil habitats (Adamčík et al. 2016, 2017a, 2017b, 2018), and a recent study reported a new tropical distribution in India (Crous et al. 2017). In this study, collections of H. glaberripes were distributed from 23°N to 26°N in the subtropical zone of southern China and at altitudes of 200–800 m, mostly occurred in the margin of broadleaf forest (mainly dominated by Fagaceae, Hamamelidaceae and Theaceae plants) and mixed forest (dominated by broadleaf tree mixed with few conifer as Pinus massoniana Lamb., Pinus spp. and Cunninghamia spp.), commonly along the sides of cement road in the forest and preferentially in heavy clay soil to humus. This study revealed an expanded geographic distribution of Hodophilus species to subtropical regions.


Sincere acknowledgements are expressed to Dr. J. Xu, Mr. H. Huang, J.P. Li, Z.P. Song, X.R. Zhong and S.H. Zhou (Guangdong Institute of Microbiology, China) for their kind help during the field trips. This study was supported by the National Natural Science Foundation of China (Nos. 31700021, 31770014, 31670029), the Science and Technology Project of Guangdong Province (2017A030303050, 2018B030324001) and the GDAS’ Special Project of Science and Technology Development (2019GDASYL-0104009).


  • Adamčík S, Dima B, Adamčíková K, Harries D, Læssøe T, Moreau PA, Jančovičová S (2018) European Hodophilus (Clavariaceae, Agaricales) species with yellow stipe. Mycological Progress 17: 1097–1111.
  • Adamčík S, Jančovičová S, Looney BP, Adamčíková K, Birkebak JM, Moreau PA, Vizzini A, Matheny PB (2017a) Circumscription of species in the Hodophilus foetens complex (Clavariaceae, Agaricales) in Europe. Mycological Progress 16: 47–62.
  • Adamčík S, Jančovičová S, Looney BP, Adamčíková K, Griffith GW, Læssøe T, Moreau PA, Vizzini A, Matheny PB (2017b) Hodophilus (Clavariaceae, Agaricales) species with dark dots on the stipe: more than one species in Europe. Mycological Progress 16: 811–821.
  • Adamčík S, Looney BP, Birkebak JM, Jančovičová S, Adamčíková K, Marhold K, Matheny PB (2016) Circumscription of species of Hodophilus (Clavariaceae, Agaricales) in North America with naphthalene odours. Botany 94: 941–956.
  • Arnolds E (1986) Notes on Hygrophoraceae – VII. On the taxonomy and nomenclature of some species of Hygrophorus. Persoonia 13: 69–76.
  • Birkebak JM, Adamčík S, Matheny PB (2016) Multilocus phylogenetic reconstruction of the Clavariaceae (Agaricales) reveals polyphyly of the agaricoid members. Mycologia 108: 860–868.
  • Birkebak JM, Mayor JR, Ryberg M, Matheny PB (2013) A systematic, morphological, and ecological overview of the Clavariaceae (Agaricales). Mycologia 105: 896–911.
  • Boertmann D (2002) Index Hygrocybearum. A catalogue to names and potential names in tribus Hygrocybeae Kühner (Tricholomatales, Fungi). Bibliotheca Mycologica 192: 1–168.
  • Crous PW, Wingfield MJ, Burgess TI, Hardy GEStJ, Barber PA, Alvarado P, Barnes CW, Buchanan PK, Heykoop M, Moreno G, Thangavel R, van der Spuy S, Barili A, Barrett S, Cacciola SO, Cano-Lira JF, Crane C, Decock C, Gibertoni TB, Guarro J, Guevara-Suarez M, Hubka V, Kolařík M, Lira CRS, Ordoñez ME, Padamsee M, Ryvarden L, Soares AM, Stchigel AM, Sutton DA, Vizzini A, Weir BS, Acharya K, Aloi F, Baseia IG, Blanchette RA, Bordallo JJ, Bratek Z, Butler T, Cano-Canals J, Carlavilla JR, Chande J, Cheewangkoon R, Cruz RHSF, da Silva M, Dutta AK, Ercole E, Escobio V, Esteve-Raventós F, Flores JA, Gené J, Góis JS, Haines L, Held BW, Horta Jung M, Hosaka K, Jung T, Jurjević Ž, Kautman V, Kautmanova I, Kiyashko AA, Kozanek M, Kubátová A, Lafourcade M, La Spada F, Latha KPD, Madrid H, Malysheva EF, Manimohan P, Manjón JL, Martín MP, Mata M, Merényi Z, Morte A, Nagy I, Normand AC, Paloi S, Pattison N, Pawłowska J, Pereira OL, Petterson ME, Picillo B, Raj KNA, Roberts A, Rodríguez A, Rodríguez-Campo FJ, Romański M, Ruszkiewicz-Michalska M, Scanu B, Schena L, Semelbauer M, Sharma R, Shouche YS, Silva V, Staniaszek-Kik M, Stielow JB, Tapia C, Taylor PWJ, Toome-Heller M, Vabeikhokhei JMC, van Diepeningen AD, Van Hoa N, Van Tri M, Wiederhold NP, Wrzosek M, Zothanzama J, Groenewald JZ (2017) Fungal Planet description sheets: 558–624. Persoonia 38: 240–384.
  • Kornerup A, Wanscher JH (1978) Methuen Handbook of Colour. Eyre Methuen: London, 1–252.
  • Kovalenko AE, Malysheva EF, Morozova OV (2012) The genus Camarophyllopsis in Russia: new records and new species C. albofloccipes. Mikologiya I Fitopatologiya 43(1): 54–66. [in Russian]
  • Matheny Curtis JM, Hofstetter V, Aime MC, Moncalvo JM, Ge ZW, Yang ZL, Slot JC, Ammirati JF, Baroni TJ, Bougher NL, Hughes KW, Lodge DJ, Kerrigan RW, Seidl MT, Aanen DK, DeNitis M, Daniele G, Desjardin DE, Kropp BR, Norvell LL, Parker A, Vellinga EC, Vilgalys R, Hibbett DS (2006) Major clades of Agaricales: a multilocus phylogenetic overview. Mycologia 98: 982–995.
  • Nylander JAA (2004) MrModeltest v2. Program distributed by the author. Evolutionary Biology Centre. Uppsala University, Uppsala.
  • Singer R (1973) Diagnoses fungorum novorum Agaricalium III. Beihefte zur Sydowia. 7: 1–106.
  • Tamura K, Peterson D, Peterson N, Stecher G, Nei M, Kumar S (2011) MEGA5: molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Molecular biology and Evolution 28: 2731–2739.
  • Thomson JD, Gibson TJ, Plewnlak F, Jianmougin F, Higgins DG (1997) The Clustal X windows interfaces: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research 24: 4876–4882.
  • Vilgalys R, Hester M (1990) Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. Journal of Bacteriology 172: 4238–4246.
  • White TJ, Bruns T, Lee S, Taylor J (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenies. In: Innis MA, Gelfand DH, Sninsky JJ, White TJ (Eds) PCR protocols, a guide to methods and applications. Academic Press, San Diego, 315–322.
  • Young AM (1997) Prelimnary observations on the limitations of the Australian Hygrophoraceae (Fungi, Agaricales). Muelleria 10: 131–138.
  • Young AM (1999) The Hygrocybeae (Fungi, Basidiomycota, Agaricales, Hygrophoraceae) of the Lane Cove Bushland Park, New South Wales. Austrobaileya 5: 535–564.
  • Young AM (2005) Fungi of Australia, Hygrophoraceae. Australian Biological Resources Study, Canberra; CSIRO Publishing, Melboune, 1–179.