Phylogenomics and systematics of the Gomphaceae with a focus on species diversity in China

Introduction

The basidiomycete order Gomphales Jülich (Agaricomycetes) currently comprises three families (Clavariadelphaceae Corner, Gomphaceae Donk, and Lentariaceae Jülich) and 22 genera (Donk 1961; Jülich 1982; Villegas et al. 2005; Hyde et al. 2024). Among the 22 genera formerly placed in Gomphales, some had been tentatively transferred to other orders, such as Ceratellopsis Konrad & Maubl. and Destuntzia Fogel & Trappe (Fogel & Trappe 1985; Olariaga et al. 2020; Mujic et al. 2024). Additionally, a few genera, including Araeocoryne Corner, Delentaria Corner, Terenodon Maas Geest., Protogautieria A.H. Sm., and Pseudogomphus R. Heim, Schildia Franchi & M. Marchetti, consisted of only one or two species each in Gomphales but have never been re-reported since their initial classification (Corner 1950, 1970; Eriksson 1954; Maas 1971; Kirk et al. 2008; Franchi & Marchetti 2021). A few of the genera had been placed in different families: Beenakia D.A. Reid and Clavariadelphus Donk belonged to Clavariadelphaceae, whereas Gomphocantharellus L. Fan, Y.Y. Xu, Zhu L. Yang & S.P. Jian, Hydnocristella R.H. Petersen, Kavinia Pilát, and Lentaria Corner were placed in Lentariaceae (Borgarino et al. 2005; Chen et al. 2015; Liu et al. 2017; Huang et al. 2020; Sandoval-Leiva et al. 2022).

Gomphaceae is the most morphologically diverse and species-rich family within Gomphales. However, the current classification systems for Gomphaceae remain inconsistent (Pine et al. 1999; Villegas et al. 1999; Humpert et al. 2001;

Hosaka et al. 2006; Giachini et al. 2010; Zhao 2017; He et al. 2019; Fan et al. 2023). Donk (1961) established Gomphaceae, with Gomphus designated as the type genus. Later, he proposed that Gomphaceae included genera Ramaricium, Kavinia, Ramaria, Clavariadelphus, Beenakia, Gomphus, Chloroneuron Murrill (now known as Phaeoclavulina Brinkmann), and Gloeocantharellus (Donk 1964). Recent decades have witnessed considerable advances in the molecular systematics of Gomphales, particularly in resolving intergeneric relationships within Gomphaceae. Early molecular phylogenetic studies based on multi-gene sequences revealed the non-monophyly of some genera in the family, viz., Gomphus Pers., Ramaria Fr. ex Bonord and Gloeocantharellus Singer (Pine et al. 1999; Villegas et al. 1999; Humpert et al. 2001; Hosaka et al. 2006; Giachini et al. 2010). Subsequent phylogenetic analyses employing broader sampling confirmed that Gomphaceae is paraphyletic and comprises several distinct lineages. Several cantharelloid-gomphoid genera such as Gloeocantharellus, Gomphus, Turbinellus Earle, or sequestrate genera like Gautieria Vittad, were found to be closely related to ramarioid genus Ramaria (Zhao 2017; He et al. 2019; Xu et al. 2022; Fan et al. 2023; Peng et al. 2023). Besides, despite being incongruent with nomenclature rules, a few ramarioid genera such as Phaeoclavulina, corticioid Ramaricium J. Erikss., as well as cantharelloid-gomphoid genera such as Luteodorsum Z.J. Peng, and several lignicolous species of Ramaria, have been temporarily assigned to Gomphaceae (Eriksson 1954; Jülich 1982; Pine et al. 1999; Villegas et al. 1999, 2005; Humpert et al. 2001; Giachini 2004a, b; Hosaka et al. 2006; James et al. 2006; Kirk et al. 2008; Bau & Liu 2013; Linhares et al. 2016; Zhao 2017; Ralaiveloarisoa et al. 2021; Li et al. 2022; Xu et al. 2022; Fan et al. 2023; Peng et al. 2023; Vidal et al. 2023; Hyde 2024; Kirk 2024). These findings highlight the need for a renewed and comprehensive taxonomic revision of Gomphaceae.

Ramaria s.l. has the largest number of species within the Gomphaceae, containing more than 300 species described worldwide (Corner 1950; Petersen 1968, 1969; Corner 1970; Petersen 1972, 1974a, b, 1975, 1976; Schild 1978; Petersen 1979, 1981, 1982; Petersen & Zang 1986; Petersen 1987, 1988a, b; Petersen & Scates 1988; Petersen 1989a; Petersen & Watling 1989; Petersen & Zang 1989; Petersen 1990; Schild 1992a, 1998; Petersen & Scates 2000; Zhang et al. 2005; Exeter et al. 2006; Kirk et al. 2008; Christan & Yorou 2009; Petersen et al. 2014; Franchi & Marchetti 2021; Li et al. 2024). The genus, with the type species R. botrytis (Pers.) Bourdot, is commonly classified into four subgenera, namely subgen. Echinoramaria, Laeticolora, Lentoramaria, and Ramaria (Corner 1950, 1970; Marr & Stuntz 1973; Petersen 1972, 1975, 1981, 1990; Thind and Sharda 1985b; Sharda and Thind 1986; Petersen & Zang 1989). In China, 45 taxa in Ramaria have been reported, including seven species in subgen. Echinoramaria, two species in subgen. Lentoramaria, four species in subgen. Ramaria, and 32 species or varieties in subgen. Laeticolora (Petersen & Zang 1986, 1989; Petersen 1989b, 1990; Zhang et al. 2005; Li et al. 2024; Zhou et al. 2024). Christan & Hahn (2005) discovered star-shaped crystals in R. flaccida (Fr.) Bourdot, and established a new subgenus, namely, Asteroramaria Christan & C. Hahn, however, the species was subsequently transferred to Phaeoclavulina, with R. flaccida now recognized as Phaeoclavulina flaccida (Fr.) Giachini (Giachini et al. 2010). Subgen. Lentoramaria mainly includes lignicolous species, whereas the other three subgenera are primarily terrestrial; subgen. Echinoramaria is characterized by echinulate basidiospores; Laeticolora, the largest of the four, is predominantly composed of species with verrucose basidiospores; and subgen. Ramaria is defined by striate basidiospores. Some mycologists classified the four subgenera into different sections or series, for example, Corner (1950, 1970) established several series, while Marchetti and Goya (2001) proposed divisions into various sections. However, these classifications were primarily based on morphological features, lacking molecular support. Molecular phylogenetic studies have demonstrated that Ramaria s.l. is not monophyletic in phylogenetic analyses. For instance, subgen. Echinoramaria and Laeticolora exhibit internal polyphyly or paraphyly. In addition, subgen. Lentoramaria occupies a phylogenetic position that is distinctly distant from other subgenera of Ramaria, suggesting that Ramaria s.l. does not belong to a natural, monophyletic group (Jülich 1982; Hibbett et al. 1997; Pine et al. 1999; Villegas et al. 1999, 2005; Humpert et al. 2001; Giachini 2004; Hosaka et al. 2006; James et al. 2006; Giachini et al. 2010; Gruhn et al. 2017; Zhao 2017; Li et al. 2022; Fan et al. 2023).

In 2010, Giachini et al. reported that Gomphus s.l. and spiny-spored Ramaria s.l. species (i.e., subgen. Echinoramaria) constitute a monophyletic clade, and divided Gomphus s.l. into Gloeocantharellus, Gomphus s.s., Phaeoclavulina, and Turbinellus. Consequently, Ramaria subgen. Echinoramaria was removed from Ramaria, a decision supported by several mycologists (Franchi & Marchetti 2021; Liu et al. 2022). In addition, Ramaria also had a complex taxonomic history. It was once placed in the family Clavariaceae (Persoon 1797a, 1801; Coker 1923; Donk 1933). Based on basidiospores color, Donk (1933) merged Gomphus and Ramaria in the tribe Ramariae within the subfamily Clavarioineae of the family Aphyllophoraceae, in the order Aphyllophorales. Eriksson (1954) proposed that Ramaricium J. Erikss. and Kavinia Pilát, which have corticioid basidiomata and the basidiospores wall surface is stained with Cotton Blue, were phylogenetically related to Ramaria. Corner (1970) disputed Donk’s classification of the genus Ramaria within the family Gomphaceae and established a new family, Ramariaceae, which included the genera Kavinia, Lentaria, Ramaria, and a newly proposed genus, Delentaria. Petersen (1971) disagreed with the establishment of the Ramariaceae family and believed that the above genera should all be retained in Gomphaceae. Subsequently, Petersen (1988b) revised Gomphaceae, with not only the cantharelloid genus Gomphus but also the genera Beenakia, Kavinia, Ramaria, Ramaricium, and Ramariopsis, of which Ramaria and Ramariopsis were the only clavarioid genera within the family. In recent decades, taxonomists have continued to argue over the taxonomic status of Ramaria, the validity of the Ramariaceae, and the genetic composition of the Gomphaceae. Villegas et al. (1999) studied the Gomphaceae s.l. and supported recognition of the Ramariaceae as proposed by Corner and proposed that Gomphales comprises four families: Gomphaceae (including genera Gomphus and Gloeocantharellus), Beenakiaceae Jülich (including genera Beenakia, Kavinia, and Ramaricium), Ramariaceae (including genus Ramaria), and Lentariaceae Jülich (including genus Lentaria). In addition, these authors removed Delentaria, Kavinia, and Lentaria from the Ramariaceae.

As described above, the classification of Gomphaceae has been contentious. Advances in DNA sequencing and genomics have provided new support for its reclassification. The high diversity and complex phylogenetic relationships within fungal groups can be better resolved using phylogenomics analyses, particularly for taxa with uncertain systematic positions. However, phylogenomic analyses targeting Gomphaceae remain scarce, and previous studies have sequenced the genomes of only a few representative species within the order (He et al. 2019). In recent years, several new genera within Gomphales, such as Gomphocantharellus, Luteodorsum and Schildia have been proposed, however, due to the lack of comprehensive phylogenomic data, their exact phylogenetic positions remain unresolved, and they are currently recognized only at the ordinal level (Earle 1909; Giachini 2004, 2010; Franchi & Marchetti 2021; Xu et al. 2022; Fan et al. 2023; Peng et al. 2023).

Based on the above observations, the aims of this study were to: (1) reconstruct the phylogenies of Gomphaceae and Ramaria s.l. using concatenated sequences of single-copy orthologous genes (SOGs) and two-gene sequence data; (2) evaluate the phylogenetic relationships of Ramaria s.l. with the four related genera: Gautieria, Gloeocantharellus, Gomphus, and Turbinellus; and (3) document and characterize both the known and newly discovered Chinese species of Gomphaceae based on morphological and molecular phylogenetic evidence.

Materials and methods

Specimen collection and morphological descriptions

This study included materials from a total of 506 specimens, representing more than 100 species collected in China over the last two decades through field collections, specimen loans, and donations from other collectors. These specimens are housed in the Fungal Herbarium of Hainan Medical University (FHMU), the Mycological Herbarium of Hunan Normal University (MHHNU), the Herbarium of Cryptogams of Kunming Institute of Botany, Chinese Academy of Sciences (HKAS), and the Mycological Herbarium of Kunming Medical University (MHKMU).

The morphological descriptions of new species were based on morphological examinations of fresh and/or dry materials. The color designations followed those of Kornerup & Wanscher (Kornerup & Wanscher 1978). The notation [n/m/p] was used for descriptions of basidiospore dimensions, representing n basidiospores measured from m basidiomata of p collections. The notation (a)b–c(d) was used for ranges, where “b−c” contained a minimum of 90% of the measured values, and extreme values (a and d) were provided in parentheses. The ratio of basidiospore length to width was abbreviated as Q, Q_m was the average Q of all basidiospores measured, and the Q_m value ± sample standard deviation was provided (Bas 1969). L_m and l_m represented the average length and width of all basidiospores measured, respectively.

The generic names applied in this paper are abbreviated as follows: Acutiramaria = “A.”, Brachyramaria = “B.”, Clavariadelphus = “C.”, Gautieria = “Gau.”, Gloeocantharellus = “Glo.”, Gomphus = “Gom.”, Gracimaria = “Gra.”, Lentoramaria = “L.”, Phaeoclavulina = “P.”, Ramaria = “R.”, Rubellaria = “Rub.”, and Turbinellus = “T.”.

DNA extraction, sequencing, and data processing

Whole-genome sequence data generated by next-generation sequencing (NGS) was employed to resolve the phylogeny of Gomphaceae. A total of 70 genomes from Gomphaceae and related genera were included, comprising four retrieved from GenBank and 66 newly generated in this study. Fruiting bodies of 66 selected species were sent to Novogene Co., Ltd. (Beijing) for DNA extraction, library construction, and then sequencing on an Illumina NovaSeq X Plus platform, generating 150 bp paired-end reads for each DNA fragment. Over 5 Gb of sequence data was obtained for each sample. The raw sequence reads were preprocessed using Trimmomatic v0.33 (Bolger et al. 2014), which consisted of trimming adapters and removing low-quality bases. The processed reads were then assembled de novo using SPAdes (Bankevich et al. 2012) with automatic K selection based on read length. Benchmarking Universal Single-Copy Orthologs (BUSCO v5.4.7) was employed to assess the completeness and quality of the genome assemblies (Manni et al. 2021), using the Agaricomycetes OrthoDB v10 database (Zdobnov et al. 2021) as a reference. This database includes 2,898 predefined orthologs (BUSCO genes) derived from 76 samples of Agaricomycetes. These BUSCO genes were used as a query in tBLASTn to search for homologous sequences in our assembled genomes (Gertz et al. 2006). To further confirm the presence of these identified putative BUSCO genes, AUGUSTUS v3.5.0 was run to predict their gene structure (Stanke et al. 2006). HMMER was then used to score these candidate BUSCO genes (Zhang and Wood 2003). Thereafter, BUSCO genes for each sample were retrieved from its corresponding genome. Four genomes that contained fewer than 1900 single-copy BUSCO genes were excluded from the final phylogenomic analyses due to their low quality. In addition, the genome of R. rubella (Schaeff.) R.H. Petersen and three outgroup species, Clathrus columnatus Bosc, Hysterangium stoloniferum Tul. & C. Tul. and Sphaerobolus stellatus Tode, were retrieved from GenBank and included in the phylogenomic analyses.

The two-gene (ITS+nrLSU) dataset: The following primer pairs were used for PCR amplification and sequencing of 257 specimens: ITS5 and ITS4 for the internal transcribed spacers 1 and 2 including the 5.8S rDNA gene (ITS) (Vilgalys & Hester 1990; Gardes & Bruns 1993); LR0R and LR5 (Vilgalys & Hester 1990) for the D1–D3 domains of the nuclear 28S rDNA gene (nrLSU). These sequences were analyzed together with those from 248 strains downloaded from GenBank. Alignments were generated separately for each gene fragment using MAFFT v7.310 (Katoh & Standley 2013). The resulting alignments were optimized and trimmed using BioEdit v7.0.9 (Hall 1999), and subsequently concatenated for maximum likelihood (ML) analysis using OFPT v1 (Zeng et al. 2023). Pterula gracilis (Desm. & Berk.) Corner and Pterula echo D.J. McLaughlin & E.G. McLaughlin in Pterulaceae Corner as outgroup taxa.

Phylogenetic analyses

Two datasets were assembled for this study: Dataset I comprised species in the Gomphaceae and related genera represented by the concatenated SOGs (single-copy orthologous genes), obtained through whole-genome sequencing, while Dataset II was a multi-copy gene dataset encompassing most species of Ramaria s.l. worldwide, incorporating concatenated ITS and nrLSU sequences.

In dataset I (SOGs), 66 specimens representing 66 species of Gomphaceae and related genera were included. The 1090 SOGs shared by more than 95% of samples were aligned using MAFFT v7.505 (Katoh & Standley 2013) under the L-INS-I setting. Alignments with fewer than 100 aligned amino acids were excluded from further phylogenomic analyses. Ambiguous regions in the remaining alignments were then eliminated using Gblocks 0.91b (Talavera & Castresana 2007), with “allowed gap positions” set to “half”, “minimum length of a block: set to “5”, and other default parameters. To investigate the potential effects of sequence biases in the dataset on phylogenomic reconstruction, several related tests were performed using TreSpEx (Struck 2014), which was recently developed to detect possible sequence biases, including long branch attraction (LBA) and saturation. After that, 65 of the 1090 SOGs were excluded from the final analysis. Phylogenomic analyses were then performed based on the remaining 1025 SOGs using both concatenation and coalescence methods. For the concatenation analysis, the SOGs were concatenated using PhyKIT (Steenwyk et al. 2021) and then subjected to the ML analysis using IQ-TREE v.2.0-rc1 with automatic detection for the best-fit model with the “−MFP” option (Nguyen et al. 2015). Branche supports were generated with 1,000 UFBoot (ultrafast bootstrap) replicates (Hoang et al. 2018). For the coalescent analysis, sequences with more than 60% gap characters were removed from each SOG to improve the accuracy of individual gene analyses (Sayyari et al. 2017). Individual gene trees were then generated by RAxML v8.2.12 with the GTR+GAMMA model and 1000 rapid bootstrap replicates (Stamatakis, 2014). We next contracted branches with low bootstrap support values (≤30%) for these gene trees (Zhang et al. 2017, 2018), and then used these partially multifurcating gene trees to infer the coalescent-based species tree with ASTRAL v.5.7.3 (Zhang et al. 2018). The local posterior probabilities (LPP) were used to assess clade support (Sayyari & Mirarab 2016).

Dataset II (ITS/nrLSU) comprised 489 ITS sequences and 406 nrLSU sequences representing a total of 506 specimens belonging to more than 100 species. The total aligned length, including gaps at different positions for different taxa, was 2502 bp, with 1530 bp for ITS and 972 bp for nrLSU. The optimal partition schemes and evolutionary models for dataset II were as follows: ITS (GTR+F+R6) and nrLSU (TIM3e+R5). For taxa described from southwestern China by Petersen & Zang (Petersen & Zang 1986, 1989; Petersen 1990), we were unable to obtain any DNA sequences from their type specimens through PCR amplification and Sanger sequencing. To overcome this problem, next-generation sequencing (NGS) was performed to obtain the target loci. Following the genome skimming approach on a DNBSEQ-T7 platform (BGI, Shenzhen, China), 15 isotype specimens of Ramaria s.l. and one specimen of R. hemirubella were sequenced. The resulting ITS and nrLSU sequences were extracted from the corresponding genomic data. These sequences were selected and added to Dataset II for phylogenetic analysis.

Results

SOGs phylogenomic analysis of Gomphaceae and related genera

In this study, Ramaria sensu lato (Ramaria s.l.) is used to include Ramaria and the formerly recognized subgen. Echinoramaria and Lentoramaria, like those treated in previous studies. On the other hand, the term Ramaria sensu stricto (Ramaria) refers specifically to the newly revised circumscription of the genus in this study.

Dataset I included newly circumscribed Ramaria and eight related genera of Gomphaceae, i.e. Clavariadelphus, Gautieria, Gloeocantharellus, Gomphus, Lentaria, Luteodorsum, Phaeoclavulina, and Turbinellus, along with the outgroup. According to Marr & Stunz (1973), the genus Ramaria was traditionally divided into four subgenera: subgen. Echinoramaria, Laeticolora, Lentoramaria, and Ramaria. In the phylogenomic analyses generated from the 1025 SOGs, the monophyly of Gomphaceae is strongly supported (BS = 100%, PP = 1), with two well-resolved genera (Gloeocantharellus and Gomphus) and a major lineage harboring species of Gautieria, Ramaria and Turbinellus. Gloeocantharellus formed a highly supported monophyletic clade (BS = 100%, PP = 1), which was a sister group to another strongly supported clade comprising species from Gautieria, Gomphus, Turbinellus, R. subgen. Laeticolora, R. subgen. Ramaria. Gomphus clustered with some species from R. subgen. Laeticolora that exhibit purple basidiomata and clamp connections (e.g., R. violacea), forming a distinct, well-supported clade (BS = 100%, PP = 1). Gautieria was closely related to R. subgen. Ramaria with strong support (BS = 100%, PP = 1). Turbinellus is grouped with a few species of R. subgen. Laeticolora. Previously defined Ramaria species did not form a monophyletic group but rather appeared as a paraphyletic assemblage.

Species of Ramaria subgen. Lentoramaria were predominantly segregated into five clades. One clade, including R. concolor (Corner) R.H. Petersen, R. paraconcolor Franchi & M. Marchetti, R. stricta (Pers.) Quél., and an unknown species, formed a strongly supported monophyletic group (BS = 100%, PP = 1). The remaining five clades each contained one or two species: R. comitis Schild and R. rainierensis Marr & D.E. Stuntz grouped together, while R. apiculata (Fr.) Donk, R. gracilis Quél. R. rubella, and R. suecica (Fr.) Donk were each placed in separate clades.

Species within subgen. Echinoramaria formed a strongly supported clade (BS = 100%, PP = 1) that is distinct from Gomphaceae, corroborating the viewpoint of Giachini et al. (2010) and Hyde et al. (2024), to which the name Phaeoclavulina would apply at the generic rank.

Based on SOGs phylogenomic analysis, taxonomic conclusions for Gomphaceae are as follows: (1) Gomphaceae comprises Gloeocantharellus, a redefined Gomphus (including species formerly in Gomphus and some ramarioid species from subgen. Laeticolora), and a redefined Ramaria (including the remaining species of subgen. Laeticolora as well as the genera Gautieria and Turbinellus). (2) The former subgen. Echinoramaria and Lentoramaria are excluded from Gomphaceae. The subgen. Echinoramaria corresponds to the genus Phaeoclavulina, while the subgen. Lentoramaria should be divided into five genera (see taxonomic part).

ITS/nrLSU phylogenetic analysis of gen. Ramaria s.l.

Dataset II (ITS/nrLS) comprises more than 100 species (504 strains) of Gomphaceae and two outgroup species, including 256 newly sequenced strains in this study and 248 strains with sequences downloaded from GenBank. A phylogenetic analysis was conducted using the maximum likelihood (ML) method (Fig. 2).

Findings from phylogenetic analyses (Fig. 2) of Chinese specimens are summarized as follows:

Together, 113 species within Gomphaceae are reported in this study, including 68 new species, and 32 known ramarioid species and 13 new combinations. In addition, 20 cantharelloid-gomphoid species are also documented.

Fig. 1. The concatenation-based phylogenomic analyses of DNA sequences of 1025 single-copy orthologous genes (SOGs) inferred by IQ-TREE (A) and ASTRAL (B). Numbers associated with nodes represent ultrafast bootstrapping support and ASTRAL local posterior probabilities, respectively. Species highlighted in red are those recommended for genus name revision.

Based on SOGs and two-gene clustering results (Fig. 1 and Fig. 2), the genus name “Ramaria” among the published species is distributed across 27 clades. To keep the nomenclatural and taxonomic stability of the ramarioid fungi, we did not replace the genus name Ramaria with the earlier established names Gautieria or Gomphus, nor did we revise the generic status of the cantharelloid-gomphoid genus Turbinellus, which is nested within Ramaria. Instead, Gautieria and Turbinellus are provisionally treated as embedded lineages within Ramaria s.l. Once the legitimacy of Ramaria is resolved, we recommend transferring Gautieria and Turbinellus to Ramaria rather than maintaining them as embedded monophyletic genera.

Taxonomy

Based on SOGs and two-gene molecular, morphological, and ecological data, we present a new classification of Gomphaceae. Additionally, we propose establishing five new genera which previously classified under Ramaria subgen. Lentoramaria. The taxa in these five new genera currently have unresolved phylogenetic positions within the order Gomphales. Furthermore, this study explores species diversity within these five new genera and other Gomphaceae taxa documented in China.

I. Ramaria subgen. Lentoramaria doesn’t belong to Gomphaceae and should be divided into five independent genera.

Acutiramaria P. Zhang & J. Zhong, gen. nov.

MycoBank: MB 852898

Etymology: Acutiramaria, from acut = sharp, signifies that the genus exhibits acute apices.

Diagnosis: Differs from the genus Lentoramaria by its basidiomata that are light pinkish ochraceous or creamy yellow, and apices color that is white or concolorous with branches, sometimes greenish.

Type species: Acutiramaria apiculata (Fr.) P. Zhang & J. Zhong

Description:1) basidiomata small to medium sized, on wood or thick leaf litter; 2) arising from a basal mat of mycelium or easily observable rhizomorphic strands; 3) light pinkish ochraceous or creamy yellow when young, and becoming deeper ochraceous brown, vinaceous-cinnamon or rufescent alutaceous from the base upward at maturity; 4) hyphae thin walled or thick walled, monomitic; 4) hyphae and basidia base with clamps; 5) basidiospores roughened in profile.

Ecology: Gregarious on rotting deciduous and coniferous wood; primarily in summer and autumn.

Distribution: At least in Asia, north Europe, and North America.

Commentary: Based on two-gene phylogenetic analyses, species of subgen. Lentoramaria within the genus Ramaria were predominantly segregated into six independent genera (Fig. 2). Additionally, A. apiculata and its varieties formed a high supported clade distinct from the Gomphaceae. Furthermore, species within the genus sometimes exhibit greenish hues and consistently lack skeletal hyphae, distinguishing them from the genus Lentoramaria. These findings support the proposal of a new genus.

Fig. 2. The maximum likelihood phylogenetic tree of Gomphaceae inferred using the combined ITS+nrLSU sequences to aid species recognition within the family. ML bootstrap values >90% are displayed at relevant nodes. A black star marks a holotype, isotype, epitype, or a specimen representing a known Chinese Ramaria species, including new combinations. A red star indicates the holotype of a newly identified Ramaria species from China. Every sample of a species without a star indicates that the species was not originally described from China. Species names highlighted in red represent either cantharelloid gomphoid taxa within the Gomphaceae or species whose genus or species names require revision.

1. Acutiramaria apiculata (Fr.) P. Zhang & J. Zhong, comb. nov. Fig. 3

MycoBank: MB 853040

Basionym: Clavaria apiculata Fr., Syst. mycol. 1: 470 (1821). [≡ Clavaria apiculata Fr., Observ. mycol. (Havniae) 2: 288 (1818)]

≡ Clavariella apiculata (Fr.) P. Karst., Revue mycol., Toulouse 3 (no. 9): 21 (1881)

= Clavaria densissima Peck, Bull. Torrey bot. Club 30: 98 (1903)

= Clavaria tsugina Peck, Bull. N.Y. St. Mus. 67: 27 (1903)

= Clavaria dendroidea var. compacta Bourdot & Galzin, Hyménomyc. de France (Sceaux): 98 (1928) [1927]

≡ Ramaria apiculata (Fr.) Donk, Biblthca Mycol. 21: 105 (1933)

= Ramaria apiculata var. compacta (Bourdot & Galzin) Corner, Monograph of Clavaria and allied Genera, (Annals of Botany Memoirs No. 1): 556 (1950)

= Ramaria apiculata var. brunnea R.H. Petersen, Am. J. Bot. 59: 1042 (1972)

= Ramaria apiculata f. compacta (Bourdot & Galzin) R.H. Petersen, Biblthca Mycol. 43: 51 (1975)

Etymology: “apiculata” signifies the presence of acute apices.

Diagnosis: Differs from other known species by its light pinkish ochraceous or creamy yellow basidiomata, and apices whitish then paler concolorous, sometimes greenish.

Epitype: Franchi & Marchetti, Riv. Micol. 60(1): 43. 2017; Europe, Italy, Vpiteno, AMB 18207 (GenBank Acc. No.: ITS = MF288927, nrLSU = MF288935).

Description: Basidiomata 3−11 × 3−9 cm stipitate or branched from the base, repeatedly branched, arising from a basal mat and rhizomorphic strands. Rhizomorphic strands white, extensive, ill-defined, often minutely webbed, mealy; basal mat not extensive, off-white, felty to minutely strigose, often mealy. Stipe (when present) and major branches pallid ochraceous tan when very young, maturing to cinnamon brown to cinnamon tan, slowly changing to chocolate brown when bruised. Stipe base, axils and/or apices often pallid green; Branches flattened somewhat, erect to ascending; axils flattened; Apices acute to sub-palmate, usually flattened, tan to beige.

Basidiospores [30/2/1] (7.2–)7.6–9.8(–10.0) × (3.6–)4.0–5.5(–6.3) µm, L_m = 8.68 µm, l_m = 4.57 µm, Q = (1.59–)1.65–2.11(–2.32), Q_m = 1.89 ± 0.18, roughened in profile, ornamentation of ill-defined small, low, cyanophilous warts and small ridges. Hyphae of rhizomorphic stands and basal mat monomitic, of two types: a) generative, slender, conspicuously clamped, and b) very rare skeletonized generative segments 3−4.5 µm diam, equal, arising and ending in clamps, opalescent under phase contrast.

Ecology: Usually gregarious in a forest with Picea or Abies with lignicolous or humicolous habit; primarily in summer and autumn.

Distribution: At least in Asia, north Europe, and North America (Petersen 1975; Wu 1987; Franchi & Marchetti 2021).

Additional specimens examined: CHINA, SICHUAN PROVINCE: Ganzi, Kangding City, Gongga Mountain, in Quercus semecarpifolia and Picea asperata forest, altitude 3900 m, 5 September 2021, P. Zhang 4261 (MHHNU 10761); same city, Pusharong Township, Yuelianghu, in mixed forest, altitude 3600 m, 25 August 2017, P. Zhang 2761 (MHHNU 9261), P. Zhang 2766 (MHHNU 9266). YUNNAN PROVINCE: Shangri-La City, Pudacuo, in the forest with Quercus semecarpifolia, altitude 3400 m, 14 August 2018, P. Zhang 3394 (MHHNU 9894); same location, Shudu Lake, in the forest with Quercus semecarpifolia, altitude 3600 m, 15 August 2018, P. Zhang 3422(MHHNU 9922), same city, Xiaozhongdian Township, Tuanjie Village, Qianhu Mountain, in mixed forest, altitude 3450 m, 2 September 2022, P. Zhang 4870 (MHHNU 11370), P. Zhang 4873 (MHHNU 11373); same city, Deqin County, Baima Snow Mountain Nature Reserve, in Picea forest, altitude 3750 m, 17 August 2018, P. Zhang 3447 (MHHNU 9947), P. Zhang 3450 (MHHNU 9950); same location, 18 August 2018, P. Zhang 3459 (MHHNU 9959), P. Zhang 3460 (MHHNU 9960), P. Zhang 3462 (MHHNU 9962).

Commentary: Petersen (1975) reported frequent encounters of this species in northern North America (and at higher altitudes southward) and northern Europe. However, specimens from northern India and Himalayas exhibit a more pallid coloration and a hint of violet or purplish shades. Fries (1821) observed that the greenish branch tips were not consistently present. Due to historical misinterpretations of the taxon, collections lacking greenish coloration were often misidentified as R. stricta. In our phylogenetic analysis of Database I and Database II, this species shows a distant relation to true R. stricta, and the specimens collected by us lack the greenish tinge at the apices. Additionally, the species shares a close genetic distance with epitype specimens of A. apiculata collected in Italy and exhibits morphological characteristics similar to A. apiculata var. brunnea. The monophyletic clade to which this species belongs is distantly related to the Ramaria s.l. and Lentoramaira, and its habitat differs from the predominantly terricolous species of Ramaria s.l.

Brachyramaria P. Zhang & J. Zhong, gen. nov.

MycoBank: MB 858493

Etymology: Brachyramaria is derived from “brachy-” (short) and Ramaria, indicating that the genus exhibits short, ramarioid branches.

Diagnosis: Brachyramaria is characterized by ivory white to cream basidiomata with short, ramarioid branches.

Type species: Brachyramaria suecica (Fr.) P. Zhang & J. Zhong

Description:1) Basidiomata small, occurring on wood or thick leaf litter. 2) arising from a basal mat of mycelium or easily observable rhizomorphic strands; 3) Basidiomata ivory white to cream, with short, ramarioid branches. 4) hyphae thin walled or thick walled, monomitic; 4) hyphae and basidia base with clamps; 5) basidiospores roughened in profile.

Ecology: Gregarious on rotting coniferous wood; primarily in summer and autumn.

Distribution: Known from Asia and North America.

Commentary: Based on SOGs and two-gene phylogenetic analyses, Brachyramaria was segregated from Gomphaceae, and B. suecica as its sole species. Its distinguishing features—ivory white to cream basidiomata, short, ramarioid branches, and a monomitic hyphal system—clearly differentiate it from other genera in Gomphaceae.

2. Brachyramaria suecica (Fr.) P. Zhang & J. Zhong, comb. nov. Fig. 4

Basionym: Ramaria suecica (Fr.) Donk, Rev. Niederl. Homob. Aphyll. 2: 105 (1933)

Description: Basidiomata 3–6 × 3–5 cm, gregarious or cespitose on rotting wood. Rhizomorphic strands are small, white, and tomentose. Stipe 1–3 × 0.5–1 cm, distinct, single, cream to pale yellow. Major branches several, erect. Branches in 3–5 ranks, erect, short, smooth, strict to lax and open in one form, off-white to cream; internodes gradually shortening upward, V-shaped; Apices erect, short and acute, dichotomous to polychotomous, off-white to cream. Odor weakly fragrant; taste negligible.

Basidiospores [60/2/2] (7.8–)8.3–10.4(–11.3) × (3.0–)3.1–4.6(–5.0) μm, L_m = 9.44 µm, l_m = 4.32 µm, Q = 2.12–3.00, Q_m = 2.48± 0.28, elongate-ellipsoidal, roughened in profile, thick-walled, and ornamented with cyanophilous warts and small ridges. Basidia 47–60 × 9–12 μm, clavate, thin-walled, hyaline, clamped; sterigmata 4, stout, straight, 6–8 μm. Hyphae of rhizomorphic stands and basal mat monomitic, thin-walled, clamps, which are occasionally swollen and tightly arranged.

Fig. 3. Fresh basidiomata and microscopic features of A. apiculate. a–b Fresh basidiomata (a, MHHNU 11373; b, MHHNU 10721); c basidiospores (MHHNU 11373). Scale bars: a–b =2 cm, c =10 µm.

Fig. 4. Fresh basidiomata and microscopic features of B. suecica. a–b Fresh basidiomata (a, MHHNU 9981; b, MHHNU 10389); c basidiospores (MHHNU 10389). Scale bars: a–b =2 cm, c =10 µm.

Ecology: In Quercus or Picea forests or broad-leaved forests. Basidiomata occurs in summer and autumn.

Distribution: Known from China (Henan, Xinjiang, Xizang, Yunnan) and North America.

Materials examined: HENAN PROVINCE: Luoyang City, Muzaling Primeval Forest, in larch forest, altitude 1800 m, 8 August 2020, P. Zhang 3889(MHHNU 10389), P. Zhang 3897(MHHNU 10397). XINJIANG PROVINCE: Ili, Tekes County, Qiongkushitai, in a Picea forest, altitude 2000, 21 August 2019, P. Zhang 3719 (MHHNU 10219), P. Zhang 3724 (MHHNU 10224). XIZANG AUTONOMOUS REGION: Linzhi City, Milin County, Zhaxiraodeng Village, in Picea forest, altitude 3040 m, 24 August 2018, P. Zhang 3481 (MHHNU 9981). YUNNAN PROVINCE: Lijiang City, Yulong County, Laojun Mountain, altitude 3000 m, 14 July 2007, Q. Zhao LJ-100 (HKAS 59224).

Commentary: Brachyramaria suecica is primarily characterized by its off-white to cream basidiomata, with a white mycelial mat at the base. Unlike other species within subgen. Lentoramaria, this species has a monomitic hyphal system with thin-walled hyphae compared to other members of the subgenus.

Gracimaria P. Zhang & J. Zhong, gen. nov.

MycoBank: MB 858496

Etymology: Gracimaria is derived from gracilis (slender) and Ramaria, indicating that the genus exhibits slender, ramarioid branches.

Diagnosis: Gracimaria is characterized by its slender branches, acute apices and dimitic or monomitic hyphae.

Type species: Gracimaria gracilis (Pers.) P. Zhang & J. Zhong

Description:1) basidiomata small to medium sized, on wood or thick leaf litter; 2) arising from a basal mat of mycelium or easily observable rhizomorphic strands; 3) slender and ramarioid branches.; 4) hyphae thin walled or thick walled, dimitic or monomitic; 4) hyphae and basidia base with clamps; 5) basidiospores roughened in profile.

Ecology: Gregarious on rotting deciduous and coniferous wood; primarily in summer and autumn.

Distribution: At least in Asia, Europe, and North America.

Commentary: Based on SOGs and two-gene phylogenetic analyses, Gracimaria was separated from Gomphaceae, consisting only of the two species, Gra. Gracilis and Gra. comitis. It is distinguished by its off-white basidiomata, short and slender branches, and a dimitic hyphal system.

Key to the species of genus Gracimaria from China

1. Fruiting bodies off-white to cream, branches sparse, apices acute, spores L_m < 6 μm Gra. gracilis

1. Fruiting bodies cream to cinnamon, branches dense and parallel, apices acute, spores L_m > 6 μm Gra. Comitis

Fig. 5. Fresh basidiomata of representative species in Gracimaria. a−b Gra. comitis (a, MHHNU 9163, b, MHHNU 10507); c−d Gra. gracilis (c, MHHNU 10493, d, MHHNU 7477). Bar = 2 cm.

3. Gracimaria comitis (Schild) P. Zhang & J. Zhong, comb. nov. Fig. 5 and Fig. 6

Basionym: Ramaria comitis Schild, Z. Mykol. 64(1): 56 (1998)

Holotype: ZT Myc 55622 (new identifier for the material deposited by Schild in ZT con n. 2228), Italy, Poggibonsi, Siena Province, Pínuswald, collected on 7 November 1996 by P. Amedeo.

Original Description (Franchi & Marchetti 2021): Basidiomata 6.5−8.5(−9.5) × 2.5−4.4(−5.0) cm, densely branched. Stipe 0.8−1.8 × 1.0−1.3 cm, irregularly cylindrical or somewhat inflated, white at the base, becoming dirty white to creamy upwards, unchanging upon touch. Branches 2.5–6 mm thick at the base, repeatedly dichotomous or polychotomous upwards, somewhat divergent or parallel, grayish brown, turning dark purple at age, especially at the apices. Flesh dirty white to creamy-white, unchanging color when touch. Odor reminiscent of moss or wood, becoming slightly anise-like with age. Taste slightly acrid or somewhat bitter.

Basidiospores [51/1/1] 6.4−9.4 × 3.7−5.1 μm, L_m = 7.9 μm, l_m = 4.4 μm, Q = 1.6−2.0, Q_m = 1.7, in mass pale ochraceous, verrucose, cyanophilous. Basidia 38–55 × 6.3–8.4 μm, mostly 4-spored (rarely three-spored), clamps. Hyphal system monomitic, composed of generative hyphae, clamps. Rhizomorphic strands are formed by generative hyphae, clamp, slightly thick-walled, smooth.

Ecology: Gregarious on rotting wood in pine forests.

Distribution: Known from Aisa, North America, and northern Europe (Franchi & Marchetti 2021).

Additional materials examined: CHINA, NEI MONGOL AUTONOMOUS REGION: Hulunbuir City, Genhe County, Delpur, altitude 1050 m, 8 August 2013, P. Zhang 1293 (MHHNU 7793). SICHUAN PROVINCE: Ganzi, Kangding City, Waze Township, in a mixed forest of Abies and Picea, altitude of 4000 m, 23 August 2017, P. Zhang 2740 (MHHNU 9240). JILIN PROVINCE: YanbianAntu County, Changbai Mountain, Hongsongwang Scenic Area, in mixed forest, altitude 1100 m, 7 August 2017, P. Zhang 2663 (MHHNU 9163); Baishan City, Fusong County, Lushuihe National Forest Park, 26 August 2020, P. Zhang 4007 (MHHNU 10507).

Commentary: The species could be mistaken for Gra. rainierensis, but it can be immediately distinguished by the monomitic hyphal and smaller basidiospores (8.5 × 5.0 μm, Marr & Stuntz 1973).

Fig. 6. Basidiospores features of Gra. comitis (MHHNU 10507). Bars: a =10 µm.

4. Gracimaria gracilis (Pers.) P. Zhang & J. Zhong, comb. nov. Fig. 5 and Fig. 7

Basionym: Ramaria gracilis (Pers.) Quél., Fl. mycol. France (Paris): 463 (1888)

≡ Merisma gracile (Pers.) Spreng., Syst. veg., Edn 16 4(1): 496 (1827)

≡ Clavaria gracilis Pers., Comm. fung. clav. (Lipsiae): 50 (1797)

= Clavaria fragrantissima G.F. Atk., Annls mycol. 6(1): 57 (1908)

= Clavaria stricta var. alba Cotton & Wakef., Trans. Br. mycol. Soc. 6(2): 174 (1919) [1918]

= Clavariella gracilis (Pers.) P. Karst., Revue mycol., Toulouse 3(no. 9): 21 (1881)

Lectotype: [illustration in] Persoon, Commentatio de Fungis Clavaeformibus (Leipzig): Tab. 4, Fig. 2, 1797 (Persoon 1797a; Franchi & Marchetti 2021 2017).

Epitype: AMB 18210, Italy, Passo San Pellegrino, Soraga (TN), in mixed forest of Picea abies and Pinus sylvestris, altitude 1300 m, collected by L. Lami on 13 August 2005, GenBank number: MF288929 (ITS), MF288937 (LSU) (Franchi & Marchetti 2021 2017).

Description: Basidiomata 1–5 × 3−4 cm, upward off-white to creamy, base yellow brown. Stipe solitary, 0.5–1 × 0.3–0.5 cm, slender, creamy-white becoming darker with age. Branches 5−6 times, slender, white turning pale yellow-brown when mature, dichotomous, straight or curved, dense, smooth in surfaced; internodes V-shaped, long, parallel. Apices acute, off-white, dichotomous, or polychotomous. Rhizomorphic strands white. Flesh leathery. Odor and taste are not recorded.

Basidiospores [40/2/2] 5.0−6.0(−7.0) × 3.0−4.0 μm, L_m = 6.49 μm, l_m = 4.63 μm, Q = (1.25−)1.60−2.00(−2.30), Q_m = 1.81, narrowly ellipsoid, roughened in profile; strong cyanophilous; apiculus prominent. Basidia (24–)30–43(–48) × 7–9 μm, clavate, thin-walled, hyaline, clamps at base, 4-spored, sterigmata 4–5 μm long. Hyphae system dimitic, densely interwoven, conspicuously clamped; Rhizomorphic strands are formed by generative hyphae, clamps, thick-walled.

Ecology: Gregarious on rotting deciduous and coniferous wood.

Distribution: Known from Aisa and Europe (Franchi & Marchetti 2021).

Additional materials examined: CHINA, HUNAN PROVINCE: Yongzhou City, Dong'an County, altitude of 300 m, Z. H. Chen 30703, 16 September 2010 (MHHNU 30703); Changde City, Shimen County, Hupingshan National Nature Reserve, in mixed forest, altitude 500 m, 8 July 2011, P. Zhang 977 (MHHNU 7477); Yongshun County, Xiaoxi National Nature Reserve, in mixed forest, altitude 900 m, 28 August 2014, P. Zhang 1776 (MHHNU 8276); Zhangjiajie City, Wulingyuan National Forest Park, altitude 800 m, 10 July 2020, Z. H. Chen 32678 (MHHNU 32678), 12 July 2020, Z. H. Chen 32704 (MHHNU 32704), Z. H. Chen 32717 (MHHNU 32717). YUNNAN PROVINCE: Chuxiong, Nanhua County, Yulu Township, in broad-leaved forest, altitude 2000 m, 10 August 2016, P. Zhang 2396 (MHHNU 8896); Lijiang, Yulong County, Yulong Snow Mountain, altitude 3700 m, 26 July 2017, P. Zhang 2793 (MHHNU 9293); Wenshan, Malipo County, Donggan Town, Malibao Village, in broad-leaved forest with Fagaceae, altitude 1700 m, 6 August 2018, P. Zhang 3359 (MHHNU 9859); Chuxiong, Zixi Mountain, in broad-leaved forests, altitude 2200 meters, 25 August 2022, P. Zhang 4798 (MHHNU 11298). JILIN PROVINCE: YanbianAntu County, Changbai Mountain, Hongsongwang Scenic Area, in mixed forest, altitude 1100 m, 7 August 2017, P. Zhang 2665 (MHHNU 9165), 25 August 2019, P. Zhang 3821 (MHHNU 10321), 27 August 2019, P. Zhang 3827 (MHHNU 10327), 11 September 2019, P. Zhang 3829 (MHHNU 10329); Erdaobaihe Town, Changbai Mountain, 25 August 2020, P. Zhang 3993 (MHHNU 10493). HUBEI PROVINCE: Enshi, Hefeng County, Mulinzi Nature Reserve, in mixed forest, altitude 1200 m, 23 September 2020, Z. H. Chen 32544 (MHHNU 32544). GUIZHOU PROVINCE: Longli County, Longjiashan National Forest Park, in coniferous-broadleaf mixed forest, altitude 1300 m, 20 June 2023, P. Zhang 4957 (MHHNU 11457); Leishan County, Leigongshan National Forest Park, in coniferous-broadleaf mixed forest, altitude 1600 m, 21 June 2023, P. Zhang 4962 (MHHNU 11462).

Commentary: This species was first described in 1790 by the naturalist and illustrator James Bolton in An History of Fungusses Growing About Halifax (Vol. 3, Tab. 111, Fig. 1). The English author conceived the taxon as a small, unbranched clavarioid species with a dirty white color. Persoon (Persoon 1797a), in his monographic work on simple or branched clavarioid fungi, described Clavaria gracilis but, unlike Bolton, associated the species with a branched species, initially whitish in color, with dimensions similar to those of Bolton’s species. Even Fries (Fries 1821) described Clavaria gracilis, nom., favoring Persoon’s later interpretation. Consequently, with the name, Bolton’s concept of taxon fell into obscurity.

Gra. gracilis can be recognized by its small basidiomata, which range from white to creamy, and especially by its anise-like odor. Microscopically, it is distinguished by its dimitic hyphal structure, present both in the context of the stipe and branches, as well as in the mycelium and rhizomorphic strandse.

From a SODs and two-gene perspective, the epitype of Gra. gracilis is placed within a monophyletic clade derived from other species.

Fig. 7. Basidiospores features of Gra. gracilis (MHHNU 7477). Bars: a =10 µm.

Lentoramaria (Corner) P. Zhang & J. Zhong, comb. & stat. nov.

MycoBank: MB 852899

Basionym: Ramaria subgen. Lentoramaria Corner Beih. Nova Hedwigia 33: 239 (1970)

Etymology: The combination of “lento”, meaning “flexible”, and the genus name “ramaria” signifies that species within this genus exhibit flexible characteristics that differ from those of the genus Ramaria.

Diagnosis: Differs from the genus Acutiramaria by its dimitic hyphal system and paler basidiomata, and from the genus Ramaria s.l. by its lignicolous or humicolous habit, in contrast to the terricolous habit of genus Ramaria s.l.

Type species: Lentoramaria stricta (Pers.) P. Zhang & J. Zhong (MycoBank: MB 853041)

Description: 1) basidiomata small to medium sized, on wood or thick leaf litter; 2) arising from a basal mat of mycelium or easily observable rhizomorphic strands; 3) white to cream when young, brownish at maturity; 4) hyphae thin walled or thick walled, usually dimitic; 4) hyphae and basidia base with clamps; 5) basidiospores roughened in profile.

Ecology: Gregarious on rotting deciduous and coniferous wood; primarily in summer and autumn.

Distribution: Widespread in Asia, Europe, North America, north Africa (Corner 1950; Petersen 1975; Wu 1987; Zhang 2010; Franchi & Marchetti 2021).

Commentary: In our SODs and two-gene phylogenetic analysis, the species of subgen. Lentoramaria (L. gracilioides, L. paraconcolor, L. concolor, L. stricta, and R. sp) are excluded from the family Gomphaceae. Additionally, their lignicolous or humicolous habit differs from the terricolous habit of genus Ramaria s.l. Therefore, we propose elevating subgen. Lentoramaria to genus level.

Key to the species of genus Lentoramaria from China

1. Branches sparse, not strict parallel L. affinis

1. Branches dense, parallel 2

2. Apices and branches are almost concolorous with each other; branches are medium sized, and contain gloeoplerous hyphae L. concolor

2. Without gloeoplerous hyphae 3

3. Apices and branches are almost concolorous; medium sized branches, lignicolous habit L. paraconcolor

3. Apices and branches are almost concolorous with each other; branches slender L. gracilioides

5. Lentoramaria affinis (L. Fan, Y. Li & N. Mao) P. Zhang & J. Zhong, comb. nov.

Basionym: Ramaria affinis L. Fan, Ying Li & N. Mao, Mycology: 10.1080/21501203.2024.2388910, 10 (Li et al. 2024)

Original description in Li et al. (2024).

6. Lentoramaria concolor (Corner) P. Zhang & J. Zhong, comb. nov. Fig. 8 a−b and Fig. 9

MycoBank: MB 853468

Basionym: Ramaria stricta var. concolor Corner, Monograph of Clavaria and allied Genera, (Annals of Botany Memoirs No. 1): 700 (1950).

≡ Ramaria concolor (Corner) R.H. Petersen, Biblthca Mycol. 43: 54 (1975)

= Ramaria stricta var. orientalis R.H. Petersen, Biblthca Mycol. 43: 54 (1975)

= Ramaria concolor f. marrii R.H. Petersen, Biblthca Mycol. 43: 62 (1975)

= Ramaria stricta var. laxiramosa Marr & D.E. Stuntz, Biblthca Mycol. 43: 140 (1974) [1975]

Diagnosis: Differs from other species of the genus by having apices that are either concolorous with branches or creamy, branches are medium sized and contain gloeoplerous hyphae.

Neotype: The U. S. A., 4 August 1917, illustrated by Coker, 1923.

Fig. 8. Fresh basidiomata of representative species in Lentoramaria. a−b L. concolor (a, MHHNU 9155, b, MHHNU 9152); c−d L. gracilioides (c, MHHNU 9154, d, MHHNU 10506); e−f L. paraconcolor (e, MHHNU 10311, f, MHHNU 7090). Bars = 2 cm.

Fig. 9. Basidiospores of L. concolor (MHHNU 9155). Bars =10 µm.

Description: Basidiomata 14 × 10 cm, solitary, gregarious or cespitose on rotting wood (either coniferous or deciduous), arising from a small, white, tomentose to felty basal mycelial mat, usually quickly becoming involved in a copious tangle of stout, white, ropy rhizomorphic strands permeating the substrate to an indefinite distance. Stipe variable, up to 1.2 cm thick, often branched almost from the base, cinnamon when fresh and young, becoming either somewhat violaceous or ruddy in age or deep chocolate brown. Major branches several, up to 8 mm thick, usually divergent secondarily by the force of the copious branches above, but in one form lax ascending. Branches numerous, erect, strict to lax and open in one form, dull ochraceous to tan, becoming dark in age; axils rounded; internodes up to 1.5 cm long below, gradually shortening; upper branches and apices erect, rounded in cross-section; strict, often somewhat prolonged, dichotomous to polychotomous, some shade of pale tan or ochre-tan, but not yellow or greenish yellow; hymenium usually unilateral, with sterile areas restricted to narrow lines decurrent from axils. Odor strong, of anise or aromatic; taste mildly bitter to bitter, sometimes mildly acrid.

Basidiospores [40/4/2] 7.0−9.0 × 4.0−5.0 μm, L_m = 7.83 μm, l_m = 4.61 μm, Q = 1.40−2.00, Q_m = 1.72 ± 0.18, roughened in profile; moderately cyanophilous; ornamentation of very obscure, low warts or ridges. Hyphae of basal mat and rhizomorphic stands of three types: a) generative, densely interwoven, conspicuously clamped; b) skeletal, thick walled, hyaline, straight, aseptate, arising from a clamp connection but ending blindly; and c) gloeoplerous hyphae 3.5−4.5 μm diam, thin walled.

Ecology: Gregarious on rotting deciduous and coniferous wood, primarily in summer and autumn.

Distribution: At least in Asia, North America, northern and central Europe (Petersen 1975; Wu 1987; Franchi & Marchetti 2021).

Additional materials examined: CHINA, JILIN PROVINCE: YanbianAntu County, Erdaobaihe Town, Changbai Mountain, 25 August 2020, P. Zhang 3986 (MHHNU 10486); same location, 6 August 2017, P. Zhang 2646 (MHHNU 9146), P. Zhang 2647 (MHHNU 947), P. Zhang 2652 (MHHNU 9152); same location, 21 August 2002, P. Zhang 230 (MHHNU 6730); same location, 19 August 2002, P. Zhang 191 (MHHNU 6691), P. Zhang 192 (MHHNU 6692); Baishan City, Fusong County, Lushuihe National Forest Park, 18 August 2002, P. Zhang 184 (MHHNU 6684). HUNAN PROVINCE: Zhangjiajie City, Sangzhi County, Badagongshan National Nature Reserve, 22 October 2014, P. Zhang 1886 (MHHNU 8386).

Commentary: Lentoramaria concolor and L. stricta are almost identical in microscopic characteristics, odor, taste, and habit, leading to Coker’s (1923) mixing up of the two species. L. stricta clearly exhibits yellow to greenish yellow apices both in Europe and North America. However, L. concolor has branch apices without this characteristic distinctive color, exhibiting a paler color as the branches. According to Petersen (Petersen 1975), the true L. stricta produces a somewhat larger fruiting body, but individual specimens cannot be reliably distinguished on this basis alone, especially when dried. Having not observed living material, Corner (1950) proposed Coker’s taxon as a variety of L. stricta and its concolorous counterpart, with the concolorous form also rarely occurring in Europe. Petersen (Petersen 1975) considered the two-color forms distinct and accepted them at the species level, dividing L. concolor into four types.

7. Lentoramaria gracilioides (Franchi & M. Marchetti) P. Zhang & J. Zhong, comb. nov. Fig. 8 c−d and Fig. 10

MycoBank: MB 853466

Basionym: Ramaria gracilioides Franchi & M. Marchetti, Index Fungorum 457: 5 (2020).

Etymology: “gracilioides” refers to the species that resemble Ramaria gracilis.

Diagnosis: Differs from other species of the genus by its apices and branches, which are almost concolorous with each other, and its dense, slender branches.

Holotype: Europe, Italy, Caldes, 8 September 2007, AMB 18534 (GenBank Acc. No.: ITS = MT055925, nrLSU = MT053212).

Description: Basidiomata 4−6 × 2−2.5 cm, pale yellow or whitish. white rhizomorphic strands or felty basal mycelial mat at base. Stipe single, 2−5 × 0.5−0.7 cm, off-white, deeper tones over the time, white or brown tomentose at base, hollow when maturity. Branches numerous, erect, strict, becoming dark in age; axils rounded, U-shaped; internodes up to 1.5 cm long below, gradually shortening; upper branches and apices erect, rounded in cross-section; strict, often somewhat prolonged, dichotomous to polychotomous. Apices acute and short, pale yellow when young, concolor with branches at age. Odor strong, of anise or aromatic; taste mildly bitter, sometimes mildly acrid.

Basidiospores [32/2/1] (6.5–)6.7–8.9(–9.2) × (3.1–)3.5–4.9(–5.1) µm, L_m = 7.69 µm, l_m = 4.03 µm, Q = (1.44–)1.64–2.23(–2.28), Q_m = 1.92 ± 0.21, roughened in profile; moderately cyanophilous; apiculus prominent, eccentric, truncate, ornamentation of very obscure, low warts or ridges. Basidia 40−50 × 6−8 μm, clavate, clamped. Hyphae of basal mat and rhizomorphic stands of three types: a) generative, densely interwoven, conspicuously clamped; b) skeletal, thick walled, hyaline, straight, aseptate, arising from a clamp connection but ending blindly; and c) very rare gloeoplerous hyphae 3.5−4.5 μm diam, thin walled.

Ecology: Gregarious on rotting deciduous and coniferous wood, primarily in summer and autumn.

Distribution: At least in Asia and Europe (Petersen 1975; Wu 1987; Franchi & Marchetti 2021).

Additional materials examined: CHINA, JILIN PROVINCE: YanbianAntu County, Erdaobaihe Town, Changbai Mountain, 6 August 2017, P. Zhang 2654 (MHHNU 9154); Baishan City, Fusong County, Lushuihe National Forest Park, 26 August 2020, P. Zhang 4006 (MHHNU 10506).

Commentary: Lentoramaria gracilioides bears a resemblance in fruiting body coloration to Gra. gracilis, with both species having dimitic rhizomorphic strands. However, the latter possesses relatively smaller basidiospores measuring 5.0−6.0(−7.0) × 3.0−4.0 μm (as measured in this study).

8. Lentoramaria paraconcolor (Franchi & M. Marchetti) P. Zhang & J. Zhong, comb. nov. Fig. 8 e−f and Fig. 11

MycoBank: MB 853465

Basionym: Ramaria paraconcolor Franchi & M. Marchetti, Index Fungorum 457: 6 (2020)

Etymology: “paraconcolor” refers to the species that resembles Ramaria concolor.

Diagnosis: Differs from other species of the genus by its lignicolous habit and short basidiospores.

Holotype: Europe, Italy, Bondo, 14 September 2002, AMB 18546 (GenBank Acc. No.: ITS = MT055928, nrLSU = MT053215).

Description: Basidiomata 6−9 × 2.5−2.5 cm, ochre-yellow color when young, turn to cinnamon to reddish-brown color at maturity, with injured areas turning purplish brown. white rhizomorphic strands or felty basal mycelial mat at base. Stipe single, 2−5 × 0.5−0.7 cm, off-white, deeper tones over the time, white or brown tomentose at base, hollow when maturity. Branches numerous, erect, strict, becoming dark in age; axils rounded, U-shaped; internodes up to 1.5 cm long below, gradually shortening; upper branches and apices erect, rounded in cross-section; strict, often somewhat prolonged, dichotomous to polychotomous. Apices acute and short, pale yellow when young, concolor with branches at age. Odor and taste are not recorded.

Basidiospores [32/2/1] (6.2–)6.3–8.4(–8.8) × 3.7–4.9(–5.1) µm, L_m = 7.30 µm, l_m = 4.29 µm, Q = (1.37–)1.41–1.97 (–2.19), Q_m = 1.71 ± 0.19, roughened in profile; moderately cyanophilous; apiculus prominent, eccentric, truncate, ornamentation of very obscure, low warts or ridges. Basidia 20−40 × 6−8 μm, clavate, clamped. Hyphae of basal mat and rhizomorphic stands of three types: a) generative, densely interwoven, conspicuously clamped; b) skeletal, thick walled, hyaline, straight, aseptate, arising from a clamp connection but ending blindly; and c) very rare gloeoplerous hyphae 3.5−4.5 μm diam, thin walled.

Fig. 10. Basidiospores of L. gracilioides (MHHNU 9154). Scale bars: a =10 µm

Ecology: Gregarious on rotting deciduous and coniferous wood, primarily in summer and autumn.

Distribution: At least in Asia and Europe (Petersen 1975; Wu 1987; Franchi & Marchetti 2021).

Additional material examined: CHINA, JILIN PROVINCE: 23 August 2019, P. Zhang 3811 (MHHNU 10311).

Commentary: In our two-gene phylogenetic analysis, L. paraconcolor is related to L. gracilioedes. In the description of Franchi & Marchetti (Franchi & Marchetti 2021), L. paraconcolor has the narrowest basidiospores width compared to L. concolor and L. gracilioides. However, we believe the basidiospores of these species are within a similar range and cannot be strictly distinguished. This species grows on decaying wood, and the whiteness of the fruiting body is a more obvious feature.

Rubellaria P. Zhang & J. Zhong, gen. nov.

MycoBank: MB 858497

Etymology: Rubellaria is derived from rubella (reddish or wine-red) and Ramaria, indicating that the genus exhibits wine-red basidiomata with ramarioid and sparse branches.

Diagnosis: Rubellaria is characterized by its wine-red basidiomata, ramarioid and sparse branches, and dimitic hyphae.

Type species: Rubellaria rubella (Schaeff.) P. Zhang & J. Zhong

Description: 1) Basidiomata small to medium sized, growing on wood, reddish or wine-red, retaining color upon maturity; 2) Stipe not distinct and branches arise directly from the base.; 3) hyphae thin walled or thick walled, monomitic; 4) hyphae and basidia base with clamps; 5) basidiospores roughened in profile.

Ecology: Gregarious on rotting deciduous and coniferous wood; primarily in summer and autumn.

Distribution: At least in Asia, Europe, and North America.

Commentary: Based on SOGs and two-gene phylogenetic analyses, Rubellaria was separated from Gomphaceae and consisting only one species, Rub. rubella. It is distinguished by its wine-red basidiomata, ramarioid and sparse branches, and dimitic hyphae.

Fig. 11. Basidiospores of L. paraconcolor (MHHNU 10311). Scale bars: a = 10 µm

9. Rubellaria rubella (Schaeff.) P. Zhang & J. Zhong, comb. nov. Fig. 12

Basionym: Clavaria rubella Schaeff., Fung. bavar. palat. nasc. (Ratisbonae) 4: 119 (1774)

= Clavaria acris Peck, Ann. Rep. Reg. N.Y. St. Mus. 54: 155 (1902) [1901]

= Ramaria acris (Peck) Corner, Trans. Br. mycol. Soc. 44(2): 235 (1961)

= Ramaria rubella var. rubella (Schaeff.) R.H. Petersen, Am. J. Bot. 61(7): 746 (1974)

= Ramaria rubella f. blanda R.H. Petersen, Biblthca Mycol. 43: 89 (1975)

= Ramaria rubella var. himalaica K.S. Thind & Sharda, Proc. Indian Acad. Sci., Pl. Sci. 95(1): 60 (1985)

Etymology: The specific epithet refers to the reddish color of the basidiomata.

Lectotype: [illustration in] Schaeffer, Fungorum qui in Bavaria et Palatinatu circa Ratisbonam nascuntur 2: Tab. 177 fig. 1−2, 1763 (designated by Petersen, 1974: “The excellent illustration which must serve as type for the fungus”).

Description: Basidiomata 5−6 × 3−4 cm, wine red color. Stipe 1−2 ×0.5−1 cm, solitary or clustered, light brown, later becoming dark brown; flesh leathery. Main branches arise directly from the ground. Branches 5−6 times, reddish-brown, dichotomous, V-shaped, straight or curved, dense, smooth surface, internodes long, parallel. Apices are acute and short, yellowish brown then reddish-brown, dichotomous, or polychotomous to crest-like structure. White rhizomorphic strands at the base, leathery. Odor and taste are not recorded.

Basidiospores [55/2/2] (5.0−)6.0−7.0(−8.0) × 4.0−5.0 μm, L_m = 6.49 μm, l_m = 4.63 μm, Q = (1.60−)1.75−2.50, Q_m = 1.41 ± 0.19, roughened in profile; strong cyanophilous; apiculus prominent, eccentric, truncate, ornamentation of very obscure, low warts or ridges. Basidia 40−60 × 7−9 μm μm, clavate, clamped. Hyphae of rhizomorphic stands and basal mat monomitic, thin-walled, clamps, which are occasionally swollen and tightly arranged.

Ecology: Gregarious of rotting deciduous and coniferous wood.

Distribution: Known from Aisa, North America, and northern Europe (Petersen 1975; Wu 1987).

Additional materials examined: CHINA, HUNAN PROVINCE: Zhangjiajie City, Sangzhi County, Badagongshan National Nature Reserve, Tianping Mountain, 27 July 2020, Zuo H. Chen 31953 (MHHNU 31953); same location, 28 July 2020, Zuo H. Chen 31973 (MHHNU 31973).

Commentary: The wine-red color of its basidiomata distinguishes it from other lignicolous or humicolous species. Additionally, its basidiospores are short and wide. The skeletal hyphae in rhizomorphic strands differs from those of L. stricta but resembles those of Gra. gracilis.

II. Ramaria subgen. Laeticolora and subgen. Ramaria both belong to Ramaria in Gomphaceae, with notable species diversity observed in China.

Ramaria Fr. ex Bonord. 1851, Handb. Mykol. 166. nom. cons., non Ramaria Holmsk. ex A. Gray, Nat. Arr. Brit. P1. 1: 655, 1821.

= Ramaria Holmsk., Beata ruris Otia fung. Dan. 1: 77, 1790

= Ramaria Holmsk. ex Raf., Anal. Nat. ou Tabi. Univ.: 211, 1815

= Ramaria Holmsk. ex Gray, Nat. arrang. Brit. pi. 1: 655, 1821

= Clavaria tribu Ramaria Fr., Epicr. Syst. Mycol.: 571, 1838

= Ramaria Holmsk. ex Quél., FI. Mycol. Fr.: 462, 1888

Type species: Ramaria botrytis (Pers.) Bourdot, Rev. Sci. Bourb. Centr. Fr. 7: 119-126 (1894)

Fig. 12. Fresh basidiomata and microscopic features of R. rubella. a–b Fresh basidiomata (a, MHHNU 31973; b, MHHNU 31953); c basidiospores (MHHNU 31953). Bars: a–b =2 cm, c =10 µm, d = 20 µm

Description: 1) Fruiting bodies are terrestrial, small to large size, repeatedly branched, displaying various colors such as red, orange, yellow, purple, and brown, contexts fibrous, fleshy, gelatinous, or leathery. 2) Hymenium turning green of applying 10% ferric sulphate. 3) Basidiospores colored inspect deposit, walls smooth in a few species, mostly verrucose, or striations, basidia mostly 4 sterigmata, basally clamped or not, contextual hyphae monomitic, clamps or not. 4) No cystidia are present. The hyphae of the context are monomitic, have clamps or without clamps, with elongated cells and lacking secondary septa.

The key to the series of Ramaria, along with illustrations of 97 species within the genus Ramaria is presented as follows.

Key to the series of genus Ramaria

1. Basidiospores smooth under electron microscope Ser. Leiosporae

1. Basidiospores rough 2

2. Basidiospores almost smooth under biological microscope, but with minute ornamentation under electron microscope 3

2. Basidiospores with obvious ornamentation 4

3. Basidioma pale colored, not changing color when injured Ser. Laeviformosoides

3. Basidioma yellow colored, changing color to brown-yellow or red-brown Ser. Flavae

4. Spore ornamentation with distinct stripes 5

4. Spore ornamentation in verrucose, or short ridges 6

5. Stipes white to light yellow; almost no change in color on bruising; apices flesh-colored to light red Ser. Hemirubellae

5. Stipes are stout; changing color on bruising; apices or young branches deep purple-red, orange-purple, or light brown Ser. Botrytes

6. Basidioma with purple hue 7

6. Basidioma yellow, orange, red, or brown in color, without purple hue 9

7. With clamps Ser. Robustipurpureae

7. Without clamps 8

8. Basidioma overall purple-brown Ser. Spinulosae

8. Basidioma pale yellow to salmon, only apices have pale purple hue Ser. Pallidolilacinae

9. Basidioma are bright orange, red-orange, pink, or red, displaying vivid colors 10

9. Basidioma light yellow to yellow, with brownish tones, without pink, red, or orange hues 17

10. Stipes, branches and apices almost identical in color 11

10. Stipe, branches or apices in different color, stipe paler in color 14

11. Basidioma red-orange to bright orange; flesh white to pale yellow-orange 12

11. Basidioma with pinkish, red-orange color; flesh is dark red-orange 13

12. Basidioma small to medium sized, with sparse branching; stipe slender to moderately thick, bright orange when young, lightens when matures Ser. Brunnescentiaurantiae

12. Basidioma large, with dense branching, yellow to bright orange Ser. Laetae

13. Basidioma yellow, pink or bright orange overall, stipe usually single or slender, basidiospores relatively short (L_m < 11 μm) and not distinct basidiospores ornamentation Ser. Nanisporae

13. Basidioma red hint, or context flesh red to orange, stipes usually falsely fasciculate or stout, with distinct basidiospores ornamentation Ser. Pseudobotrytes

14. Basidioma yellow to salmon, apices yellow to red, changing color to red when injured 15

14. Basidioma pale yellow, not changing color when injured 16

15. Apice yellow to red; basidiospores relatively longer (L_m > 10 μm) Ser. Sanguinipesae

15. Apice yellow; basidiospores relatively shorter (L_m < 10 μm) Ser. Luteae

16. Young branches or apices pink, branches and stipe brown Ser. Roseobrunneae

16. Basidioma yellow to salmon, branches slender and apices pink, context of stipe gelatinous, most species without clamps Ser. Gelatinosae

17. Cespitose habit 18

17. Not cespitose habit 20

18. Basidioma pale yellow to yellow brown, context fleshy Ser. Pallidochraceus

18. Basidioma generally small, light yellow to dark yellow, context leathery, tough 19

19. Basidioma deep yellow, context fleshy Ser. Sparsiaggregatae

19. Basidioma pale yellow, stipe and branches-tinged pink Ser. Conjunctipesae

20. Apices and branches distinctly different colored 21

20. Apices almost concolorous with branches 22

21. Apices off-white, basidioma pale salmon to yellow ochre Ser. Formosae

21. Apices yellow, stipe and branches pale yellow to pale salmon, basidiospores small (L_m < 10 μm) Ser. Pseudocystidiophorae

22. Context gelatinous or subgelatinous Ser. Neoformosae

22. Context fleshy to fibrous 23

23. Basidioma small to medium sized; stipe single to subfasciculate; branches strict, very sparse; flesh of brancher red to orange Ser. Semigelatinosae

23. Flesh of flesh not red to orange color 24

24. Apices acute, branches very slender, small basidiospores (L_m < 10 μm) Ser. Linearflavae

24. Apices rounded 25

25. Obvious change in color when injured 26

25. Not changing or slightly changing color on handling 27

26. Changing color to red-brown, branches slender, stipe stout, basidioma pale yellow to cinnamon, clamped Ser. Flavescentoides

26. Changing color to brown, with dense branches, stipe tapering sharply downward to an acute Ser. Densiapicalisae

27. Basidiospores relatively longer (L_m > 11 μm) Ser. Longisporae

27. Basidiospores relatively shorter (L_m < 11 μm) Ser. Zebrisporae

Key to the species of ser. Botrytes from China

1. Average length of basidiospores L_m > 18 μm R. sinobotrytis

1. Average length of basidiospores L_m < 18 μm. 2

2. Average length of basidiospores L_m < 12 μm 3

2. Average length of basidiospores 12 μm < L_m < 18 μm 4

3. Basidioma light color, creamy, distributed in southwestern China R. luteoliparva

3. Basidioma purple-red, turning wine-red on bruising, distributed in central China R. orientalibotrytis

4. Stipe not changing color on bruising 5

4. Stipe changing color on bruising 6

5. Apices are darker in color, purplish-red with orange tones R. violaceoaurantia

5. Young branches and apices are purplish-red; creamy on maturity R. magnibotrys

6. Stipes changing color to lemon yellow on bruising R. flavopurpura

6. Stipes changing color to purple-red color on bruising R. rubropurpurea

Key to the species of ser. Brunnescentiaurantiae from China

1. Basidioma red-orange; stipe changing color to reddish-brown on bruising R. brunnescentiaurantia

1. Basidioma red-orange; stipe not changing color on bruising 2

2. Basidiospores relatively broader (L_m > 4 µm) R. aurantibrevispora

2. Basidiospores relatively narrower (L_m < 4 µm) R. verrucosiflava

Key to the species of ser. Conjunctipesae from China

1. Apices yellow, stipe and branches salmon to light pink 2

1. Basidioma pale yellow to dark yellow overall 3

2. Average length of basidiospores > 8 µm R. microsporoflava

2. Average length of basidiospores < 8 µm R. sinoconjuntcipes

3. Slightly changing color to brownish yellow on bruising R. flaviaggregata

3. Almost not changing color on bruising R. pallidoflava

Key to the species of ser. Flava from China

1. Basidiospores elongated, with a high length-to-width ratio (Q_m > 3) R. flavolongispora

1. Basidiospores have a slightly smaller length-to-width ratio (Q_m < 3) 2

2. Basidioma deep yellow to light brown 3

2. Basidioma creamy to light yellow when young; color deepens on maturity or injury 4

3. Basidioma yellow, stipe stout stipes R. pallidobrunnea

3. Basidioma light brown, stipe relatively slender; apices bruise brown R. barenthalensis

4. Not changing color on bruising R. verrucula

4. Changing color on bruising 5

5. Basidiospores have a relatively long average length (L_m > 13 μm), dark brown on bruising R. laevibrunnea

5. Basidiospores have a slightly shorter average length (L_m < 13 μm) R. hepatica

Key to the species of ser. Flavescentoides from China

1. Basidioma pale yellow and easily brunnescent, stout stipes, slender branches R. brunneipes

1. Basidioma pale yellow to dark yellow, not changing color 2

2. Basidioma pale yellow, branches short R. flavescentoides

2. Basidioma dark yellow, stipe stout 3

3. Basidioma cauliflower-shaped, branches short and stout, flatted stipe R. compressipes

3. Branches slender and long R. obtusa

Key to the species of ser. Formosae from China

1. Average basidiospores length < 10 μm R. pallidicarnea

1. Average basidiospores length > 10 μm. 2

2. Stipe long, branches sparse R. longicarnea

2. Basidioma cream to ocher-yellow, branches dense to sparse R. formosoides

Key to the species of ser. Gelatinosae from China

1. Clamps is frequent 2

1. Clamps are uncommon in both basidia and tramal hyphae 3

2. Basidioma yellow-orange; basidiospores relatively longer (8.9–11.1 × 3.5–5.1 μm); distributed in southwestern China R. gelatinostipes

2. Basidioma saffron-yellow; basidiospores relatively shorter (7.0–10.0 × 4.0–5.0 μm); distributed in north China R. flavicoralloides

3. Basidioma light to vivid yellow 4

3. Basidioma yellow-orange or apices pink to red 9

4. Average basidiospores length < 10 μm R. brevigelatinosa

4. Average basidiospores length > 10 μm 5

5. Distributed in north China (Shanxi) 6

5. Distributed in southwestern 7

6. Basidiomata saffron-yellow to light brown; basidiospores relatively shorter (9.0–11.0 × 3.0–5.0 μm); distributed in north China (Shanxi) R. apicaliochracea

6. Basidiomata pale yellow to yellow; basidiospores relatively longer (10.0–12.5 × 3.8–5.0 µm); distributed in north China (Shanxi) R. platyrugosa

7. Distributed in central China R. hunanensis

7. Distributed in southwestern China 8

8. Branches slender, basidioma bright yellow R. subthindii

8. Basidioma vivid yellow R. gelatinosivivida

9. Basidioma or apices pink to red color 10

9. Basidioma yellow to orange overall 13

10. Average basidiospores length > 10 μm R. rubrisubtilis

10. Average basidiospores length < 10 μm 11

11. Average basidiospores length < 8 μm, apices pink R. indoyunnaniana

11. Average basidiospores length 8 μm < L_m < 10 μm 12

12. Apices dark red, color does not notably fade after drying R. rubroapicalis

12. Apices relatively light in color, color fades at maturity R. microspore

13. Slender branches, average basidiospores length > 13 μm R. sinensis

13. Average basidiospores length < 13 μm 14

14. Branching dense, no change in color on bruising R. subgelatiniaurantia

14. Surface of stipes and branches wrinkled and change color to brown on bruising R. gelatinifusca

Key to the species of ser. Hemirubellae from China

1. Average basidiospores length < 10 μm R. hemirubella

1. Average basidiospores length > 10 μm 2

2. Spore length (L_m) exceeding 12.5 μm R. lingkongshanensis

2. Spore length between 10 μm < L_m < 12.5 μm 3

3. Apices light pinkish cinnamon to light ochraceous cinnamon R. eryuanensis

3. Apices pink to purple-red colored, distributed in Yunnan Province R. delicorosea

Key to the species of ser. Laetae from China

1. Clamps rare of basidia base and tramal hyphae 2

1. Clamps common 3

2. Basidioma bright orange; apices turned to dark brown when hurt R. aurantiapicalis

2. Basidioma pale yellow; apices concolor with branches R. cadmioaurantiaca

3. Basidioma yellow to yellowish orange R. robustiflava

3. Basidioma flesh pink to bright orange 4

4. Basidioma bright orange R. laeta

4. Basidioma flesh pink R. xizangensis

Key to the species of ser. Laeviformosoides from China

1. Basidiospores almost smooth, basidioma light colored. R. laeviformosoides

1. Basidiospores almost smooth, basidioma yellow colored R. laeviflava

Key to species of ser. Leiosporae from China

This series is monotypic, containing only R. schildii.

Key to species of ser. Lineariflavae from China

This series is monotypic, containing only R. lineariflava.

Key to species of ser. Longisporae from China

This series is monotypic, containing only R. pseudolongispora.

Key to species of ser. Luteae from China

This series is monotypic, containing only R. sanguinixantha.

Key to the species of ser. Nanisporae from China

1. Basidioma red-orange, with a reddish hue; average basidiospores length > 10 µm R. rubrosporula

1. Basidioma red-orange, with a reddish hue; average basidiospores length < 10 µm 2

2. Hyphae and basidia base both have clamps R. nanispora

2. Hyphae almost lack clamps, without clamps at basidia base R. corallina

Key to the species of ser. Neoformosae from China

1. With clamps ………… 2

1. Clamps absent 13

2. Basidioma pale yellow to light orange; average basidiospores length L_m < 10 µm R. luteoaurantiaca

2. Average basidiospores length L_m > 10 µm 3

3. Basidioma pale yellow, clamps common in basidia but rare in hyphae, average basidiospores length L_m = 14.20 µm R. distinctissima

3. Clamps common in basidia and hyphae 4

4. Distributed in north China (Shanxi) R. cyanophila

4. Distributed in southwestern China 5

5. Average basidiospores length L_m > 14 µm 6

5. Average basidiospores length 10 ≤ L_m ≤ 14 µm 7

6. Basidioma orange-red to light brown, average basidiospores length L_m > 15 µm R. sinolargentii

6. Basidioma salmon to orange, average basidiospores length > 16 µm R. longiaurantia

7. Basidioma pale yellow, lemon-yellow on bruising; average basidiospores length 12 < L_m < 14 µm (L_m = 12.52 µm) R. brunnescentiflava

7. Average basidiospores length 10 µm < L_m < 12 µm 8

8. Basidioma reddish-orange 9

8. Basidioma salmon to pale yellow 10

9. Apices yellow-orange, darker than the pale yellow of branches, changing color to brown on bruising, distributed in central and northeastern China R. aureoapicalis

9. Branches salmon, distributed in southwest and eastern China R. hilaris

10. Stipe single 11

10. Stipe falsely fasciculate 12

11. Stipe stout; changing color to brown on bruising R. robustibrunnea

11. Stipe slender; not changing color on bruising R. aureolinearis

12. Basidioma short and stout, stipe surface rapidly fading after collection R. ephemeroderma

12. Basidioma slender, changing color to light brown on bruising R. aurantiformosa

13. Average basidiospores length L_m > 13 µm, width l_m > 13 µm 14

13. Average basidiospores length 12 µm ≤ L_m ≤ 13 µm, width 4 µm l_m ≤ 6 µm 15

14. Basidioma red-orange, branches very crowded, stipe slender R. aurantilinearis

14. Basidioma long and high, stipes long R. hypogaeoides

15. Branches elongate; distributed in central China (Shanxi) R. subcolumnaris

15. Branches relatively dense and short 16

16. Basidioma salmon; branches dispersing, internodes gradually wide; distributed in north China (Shanxi) R. peisicinoflava

16. Basidioma pale orange to orange 17

17. Basidioma broad and short, stipes stout and short R. linearis

17. Basidioma salmon to yellow-orange, stipe falsely fasciculate 18

18. Branches dense, short and stout, apices rounded R. pseudoobtusissima

18. Branches relatively slender, apices acute 19

19. Growing in coniferous forests, distributed in southwestern and northwestern and east China R. conferta

19. Growing in broad-leaved or mixed forests, distributed in southwestern China R. sinosubtilis

Key to species of ser. Pallidochraceus from China

This series is monotypic, containing only R. pallidochracea.

Key to species of ser. Pallidolilacinae from China

This series is monotypic, containing only R. pallidolilacina.

Key to species of ser. Pseudobotrytes from China

This series is monotypic, containing only R. rufoaurea.

Key to species of ser. Pseudocystidiophorae from China

This series is monotypic, containing only R. pseudocystidiophora.

Key to species of ser. Robustipurpureae from China

This series is monotypic, containing only R. robustipurpurea.

Key to the species of ser. Roseobrunneae from China

1. Basidioma pale yellow to pale orange; with clamps R. densiapicalis

1. Basidioma brown, with pink tint; without clamps R. roseobrunnea

Key to the species of ser. Sanguinipesae from China

1. Basidia base or contextual hyphae with clamps R. rubriattenuipes

1. Basidia base and contextual hyphae with clamps 2

2. Basidioma pale yellow, changing color to dark red on bruising R. sanguinipes

2. Basidioma salmon, apices deep red R. rubriapicalis

Key to species of ser. Semigelatinosae from China

This series is monotypic, containing only R. semigelatinosa.

Key to species of ser. Sparsiaggregatae from China

This series is monotypic, containing only R. sparsiaggregata.

Key to the species of ser. Spinulosae from China

1. Basidiospores rounded and broad, length-to-width ratio < 2 R. purpureivariabilis

1. Basidiospores length-to-width ratio > 2 2

2. Basidiospores average dimensions 10.01 × 4.89 µm, distributed in Yunnan Province

R. purpurea

2. Basidiospores average dimensions 11.38 × 5.62 µm, distributed in Sichuan Province R. sinospinulosa

Key to the species of ser. Zebrisporae from China

1. Basidiospores larger than 10 μm R. zebrispora

1. Basidiospores smaller than 10 μm R. linearioides

Fig. 13. Fresh basidiomata of representative species in Ramaria ser. Botrytes. a−b R. flavopurpurea (Holotype, a, MHHNU 6595; b, MHHNU 11634); c−d R. luteoliparva (Holotype, c, MHHNU 10013; d, MHHNU 10025); e−f R. magnibotrys (Holotype, e, MHHN11364; f, MHHN11365); g−h R. orientalibotrytis (Holotype, g, h, MHHNU 7165); i−j R. rubropurpurea (Holotype, i, MHHNU 10846; j, MHHNU 10767); k−l R. sinobotrytis (Holotype, k, l, MHHNU 9268); m−l R. violaceoaurantia (Holotype, m, MHHNU 10074; n, MHHNU 10002; l, MHHNU 10078); Bars =2 cm.

Ramaria ser. Botrytes Corner, Beih. Nova Hedwigia 33: 240 (1970)

Diagnosis: This series includes 7 species collected in China, all of which feature striped basidiospores, stout stipes, with bruising causing a color from light yellow to wine-red. Branching and apices display colors ranging from creamy to purple red.

Type species: R. botrytis (Pers.) Bourdot, Rev. Sci. Bourb. Centr. Fr. 7: 119-126 (1894)

Description: 1) basidiomata are milk white to pale yellow, with apices ranging from pale red, red-orange, dark purple; 2) the stipe is stout and nearly does not change color when injured; 3) basidia base and hyphae have clamps; 4) basidiospores are ornamented with conspicuous striae.

Commentary: This series includes seven species collected in China.

10. Ramaria flavopurpurea P. Zhang & Xue L. Gao, sp. nov. Fig. 13 a−b and Fig. 40

MycoBank: MB 852968

Etymology: The proposed compound epithet “flavopurpurea” refers to the basidiomata being red-purple in color, with the stipe of the species changing to lemon yellow when handled.

Diagnosis: Differs from other species of ser. Botrytes by its stipe bruising to a lemon-yellow color.

Type: CHINA, YUNNAN PROVINCE: Chuxiong, Zixi Mountain, in a forest with Lithocarpus and Pinus yunnanensis, altitude 2400 m, 2 August 2001, P. Zhang 95 (Holotype, MHHNU 7165, GenBank Acc. No.: ITS = PP467112, nrLSU = PP493444, tef1-α = PP480338).

Description: Basidiomata 13 cm high and 11 cm broad. Stipe 6 × 3 cm, single, falsely fasciculate, to compound-connate, with a few abortive branchlets, off-white (1A1) to cream (4A3), with a hint of pink upward; slowly changing color to lemon yellow (4A3) when handle; flesh solid, off-white (1A1), flesh to fibrous, drying soft, easily penetrated. Major branches several, compound at base, ascending. Branches in 2–5 ranks, short and crowded; reddish lilac (14B4); axils narrowly rounded; internodes all short. Apices are sometimes finely divided, acute, double-dichotomous, usually gross, short and crowded, rose (12A4) to grayish magenta (13D5). Odor weakly fragrant; taste negligible.

Fig. 14. Fresh basidiomata of representative species in Ramaria ser. Brunnescentiaurantiae. a−b R. aurantibrevispora (a, MHHNU 11650; b, MHHNU 11642); c−d R. brunnescentiaurantia (Holotype, c, MHHNU 10772; d, MHHNU 10812); e−f R. verrucosiflava (Holotype, e, MHHNU 9908; f, MHHNU 9904). Bars = 2 cm.

Fig. 15. Fresh basidiomata of representative species in Ramaria ser. Conjunctipesae. a−b R. flaviaggregata (Holotype, a, MHHNU 10565; b, MHHNU 6952); c−d R. microsporoflava (Holotype, c, MHHNU 31326; d, MHHNU 11313); e−f R. pallidoflava (Holotype, e, MHHNU 11652; f, MHHNU 33498); g−h R. sinoconjunctipes (g, MHHNU 11645; h, MHHNU 9940). Bars =2 cm.

Fig. 16. Fresh basidiomata of representative species in Ramaria ser. Flavae. a−b R. barenthalensis (a, MHHNU 10091; b, MHHNU 10819); c−d R. flavolongispora (Holotype, c, MHHNU 11653; d, HKAS123587); e−f R. hepatica (Holotype, e, MHHNU 10916; f, MHHNU 10915); g−h R. laevibrunnea (Holotype, g, HKAS 105942; h, MHHNU 11639); i−j R. pallidobrunnea (Holotype, i, MHHNU 9213; j, MHHNU 9985); k−l R. verrucula (Holotype, k, MHHNU 31735; l, MHHNU 31736). Bars =2 cm.

Basidiospores [60/2/2] (13.0–)14.0–18.0 × 5.0–6.0(–6.5) µm, L_m = 15.93 µm, l_m = 5.48 µm, Q = (2.15–) 2.50–3.60, Q_m = 2.92 ± 0.39, long-ellipsoid, conspicuously striae in profile, strongly cyanophilous; Basidia 60–75 × 8–12 µm, clavate, clamped at base; sterigmata mostly 4, 3–8 μm, stout and straight. Subhymenium rudimentary, hyphal. Hymenium unthickening. Tramal hyphae of stipe 6–12 µm wide, hyaline, conspicuously clamped, thin to thick-walled (wall up to 0.5 µm thick), easily broken, interwoven, not agglutinated or adherent; ampulliform clamps up to 15 µm broad, thick-walled (wall up to 1 µm thick), with conspicuous stalactiform ornamentation; gloeoplerous hyphae not observed. Tramal hyphae of upper branches up to slightly narrowly, hyaline, thin walled, conspicuously clamped, not agglutinated or adherent, parallel and hardly branched inward; ampulliform clamps occasional; gloeoplerous hyphae not observed.

Ecology: In broad-leaved or coniferous forest; basidiomata occurs in summer and autumn.

Distribution: Known from southwestern China (Yunnan).

Additional material examined: CHINA, YUNNAN PROVINCE: Chuxiong, Lufeng City, Tuoan Township, 21 July 2022, Xue L. Gao 19718 (MHHNU 11630).

Commentary: Its basidiomata morphology is quite distinctive. Unlike most species in ser. Botrytes, where the reddish-purple coloration is more intense at the apices and bruising appears light brown to wine-red, features of R. flavopurpurea branches that are entirely purple, with the stipe bruising to lemon yellow. This characteristic allows for quick identification in the wild.

Fig. 17. Fresh basidiomata of representative species in Ramaria ser. Flavescentoides. a R. brunneipes (MHHNU 6623); b R. compressipes (Holotype, MHHNU 11644); c−d R. flavescentoides (c, MHHNU 9996; d, MHHNU 9983); e−f R. obtusa (e, MHHNU 10553; f, MHHNU 7764). Bars =2 cm.

11. Ramaria luteoliparva P. Zhang & J. Zhong, sp. nov. Fig. 13 c−d and Fig. 41

MycoBank: MB 852972

Etymology: luteoliparva = luteolus + parva, proposed because this species has cream-colored basidiomata and small basidiospores.

Diagnosis: Differs from other species in the series by having the smallest basidiospores size and its basidiomata apices that lack the reddish-purple hue, appearing light brown instead.

Type: CHINA, XIZANG AUTONOMOUS REGION: Linzhi, Sejila Mountain, Bayi District, Lulang Town, in a subalpine forest dominated by Quercus, altitude 3400 m, 26 August 2018, P. Zhang 3513 (Holotype, MHHNU 10013, GenBank Acc. No.: ITS = PP467120, nrLSU = PP493451, tef1-α = PP480345).

Description: Basidiomata 14−15 × 10−15 cm high and 8−28 cm broad, big-sized. Stipe stout, up to 2 × 1 cm, single, falsely fasciculate, to compound-connate, with a few abortive branchlets, off white (1A1) to cream (4A3), not changing color or slowly and slightly turned to grayish yellow (2B5) where handled; flesh off-white (1A1), flesh to fibrous, drying soft, easily penetrated. Major branches several, indistinct. Branches in 2–4 ranks, short and crowded; cream (4A3) to dull yellow (3B3); axils narrow rounded; internodes all short. Apices short and crowded, cream (4A3) to yellow ochre (5C7). Odor weakly fragrant; taste sweet.

Fig. 18. Fresh basidiomata of representative species in Ramaria ser. Formosae. a−b R. formosoides (a, MHHNU 10760; b, MHHNU 10554); c−d R. longicarnea (Holotype, c, MHHNU 11329; d, MHHNU 11249); e−f R. pallidicarnea (Holotype, e, MHKMU HT469; f, MHHNU 11635). Bars =2 cm.

Basidiospores [60/2/2]9.0–12.7(–14.0) × (3.0–)3.50–4.8(–5.0) µm, L_m = 10.75 µm, l_m = 3.92 µm, Q = (2.00–)2.25–3.33(–3.53), Q_m = 2.74 ± 0.35, subcylindrical, conspicuously striae in profile, strongly cyanophilous; Basidia 45–60 × 7–12 μm, clavate, clamped at base; sterigmata mostly 4, 2–5 µm, stout and straight. Hymenium unthickening; Subhymenium rudimentary, hyphal. Tramal hyphae of stipe 6–12 µm wide, hyaline, conspicuously clamped, thin to thick-walled, interwoven, not agglutinated or adherent; ampulliform clamps up to 15 µm broad, usually not thick-walled; gloeoplerous hyphae not observed. Tramal hyphae of upper branches slightly narrowly, hyaline, thin walled, conspicuously clamped, not agglutinated or adherent, parallel and tightly interwoven; ampulliform clamps occasional, unornamented; gloeoplerous hyphae not observed.

Ecology: In broad-leaved or coniferous forests; basidiomata occurs in summer and autumn.

Distribution: Known from southwestern China (Xizang, Yunnan).

Additional materials examined: CHINA, SICHUAN PROVINCE: Ganzi, Kangding City, Gongga Mountain, in Quercus semecarpifolia and Picea asperata forest, altitude 4000 m, 7 September 2021, P. Zhang 4329 (MHHNU 10829). XIZANG AUTONOMOUS REGION: Linzhi, Sejila Mountain, Bayi District, Lulang Town, in a subalpine forest dominated by Quercus, altitude 3400 m, 26 August 2018, P. Zhang 3525 (MHHNU 10025).

Commentary: The species has the smallest basidiospores within the series. Other main features of R. luteoliparva include: 1) large-sized basidiomata; 2) basidiomata are cream-colored with yellow ochre color apices; 3) color changes to pale brown to dark brown when injured; 4) basidiospores are ornamented with longitudinal striae and are less 11 μm in size; 5) hyphae and basidia base have clamps.

Fig. 19. Fresh basidiomata of representative species in Ramaria ser. Gelatinosae. a−b R. brevigelatinosa (Holotype, a, MHHNU 11283; b, MHHNU 11269); c−d R. hunanensis (Holotype, c, MHHNU 7140; d, MHHNU 9727); e−f R. gelatinifusca (Holotype, e, MHHNU 9936; f, MHHNU 11347); g−h R. gelatinosivivida (Holotype, g, MHHNU 10089; h, MHHNU 10083); i−j R. gelatinostipes (i, MHHNU 10826; j, MHHNU 10762); k−l R. indoyunnaniana (k, MHHNU 9134; l, MHHNU 9839); m−n R microspora (m, MHHNU 9914; n, MHHNU 6619); o−p R. rubrisubtilis (Holotype, o, MHHNU 7160; p, MHHNU 11651); q−r R. rubroapicalis (Holotype, q, MHHNU 11218; r, MHKMU HT277); s−t R. sinensis (s, MHHNU 9177; t, MHHNU 9878); u−v R. subgelatiniaurantia (Holotype, u, MHHNU 10859; v, MHHNU 10858); w−x R. subthindii (Holotype, w, MHHNU 9131; x, HKAS55387). Bars =2 cm.

Fig. 20. Fresh basidiomata of representative species in Ramaria ser. Hemirubellae. a−b R. eryuanensis (a, MHHNU 10837; b, MHHNU 9915); c−d R. delicorosea (Holotype, c, MHHNU 31322; d, MHHNU 8882); e−f R. hemirubella (e, MHHNU 31267 and f, MHHNU 7202); g−h R. lingkongshanensis (g, MHHNU 9873; h, MHHNU 9874). Bars =2 cm.

Fig. 21. Fresh basidiomata of representative species in Ramaria ser. Laetae. a−b R. aurantiapicalis (Holotype, a, MHHNU 10099; b, MHHNU 10100); c−d R. laeta (Holotype, c, HKAS49037; d, MHHNU 33922); e−f R. robustiflava (Holotype, e, MHHNU 10090; f, MHHNU 9931). Bars =2 cm.

12. Ramaria magnibotrys P. Zhang & J. Zhong, sp. nov. Fig. 13 e−f and Fig. 42

MycoBank: MB 852959

Etymology: magnibotrys, from magni = large, and “-botrys” means clusters, is proposed because the basidiomata of this species are large and form clusters.

Diagnosis: Large basidiomata with apices that are wine-colored when young and cream-colored at maturity. Basidiomata do not significantly change color when injured. Basidiospores are ornamented with longitudinal striae. Hyphae and basidia base have clamps.

Type: CHINA. YUNNAN PROVINCE: Shangri-La City, Pudacuo National Park, Tianbaotai, in the mixed forest with Quercus and Picea. altitude 3680 m, 1 September 2022, P. Zhang 4864 (Holotype, MHHNU 11364, GenBank Acc. No.: ITS = PP467116, nrLSU = PP493447, tef1-α = PP480341).

Description: Basidiomata 6−25 cm high and 8−28 cm broad, big-sized. Stipe stout, up to 3−18 × 4−16 cm, single, falsely fasciculate, to compound-connate, with a few abortive branchlets, milk white (1A2) to cream (4A3), not changing color or slowly and slightly turned to grayish yellow (2B5) where handled; flesh off-white (1A1), flesh to fibrous, drying soft, easily penetrated. Major branches several, indistinct. Branches in 2–4 ranks, short and crowded; cream (4A3) to light red (10A5) upward, concolor with stipe below; axils narrow rounded; internodes all short. Apices short and crowded, cream (4A3) to grayish magenta (13D5). Odor weakly fragrant; taste sweet.

Basidiospores [60/2/2]12.0–15.1(–16.5) × (4.5–)5.0–7.0 µm, L_m = 13.31 µm, l_m = 5.66 µm, Q = (1.71–)1.93–2.67(–3.01), Q_m = 2.30 ± 0.23, subcylindrical, conspicuously striae in profile, strongly cyanophilous; Basidia 60–75 × 8–12 μm, clavate, clamped at base; sterigmata mostly 4, 3–7 µm, stout and straight. Hymenium unthickening; Subhymenium rudimentary, hyphal. Tramal hyphae of stipe 6–12 µm wide, hyaline, conspicuously clamped, thin to thick-walled, interwoven, not agglutinated or adherent; ampulliform clamps up to 15 µm broad, usually not thick-walled; gloeoplerous hyphae not observed. Tramal hyphae of upper branches slightly narrowly, hyaline, thin walled, conspicuously clamped, not agglutinated or adherent, parallel and tightly interwoven; ampulliform clamps occasional, unornamented; gloeoplerous hyphae not observed.

Ecology: Found in broad-leaved or coniferous forests; basidiomata occurs in summer and autumn.

Distribution: Known from southwestern China (Yunnan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Shangri-a City, Pudacuo National Park, Tianbaotai, in the mixed forest with Quercus and Picea. altitude 3680 m, 1 September 2022, P. Zhang 4865 (MHHNU 11365); same location, in the mixed forest with Quercus and Picea. altitude 3523 m, 3 August 2022, Jian W. Liu MF94 (MHHNU 11646).

Fig. 22. Fresh basidiomata of representative species in Ramaria ser. Laeviformosoides. a−b R. laeviflava (Holotype, a, MHHNU 9909; b, MHHNU 9907); c−d R. laeviformosoides (c, MHHNU 8599 and d, HKAS 55281). Bars =2 cm.

Fig. 23. Fresh basidiomata of representative species in Ramaria ser. Leiosporae. a–b R. schildii (a, MHHNU 9285; b, MHHNU 9242). Bars =2 cm.

Fig. 24. Fresh basidiomata of representative species in Ramaria ser. Lineariflavae. a–b R. lineariflava (Holotype, a, MHHNU 7162; b, MHHNU 10543). Bars =2 cm.

Fig. 25. Fresh basidiomata of representative species in Ramaria ser. Longisporae. a–b R. pseudolongispora (Holotype, a, MHHNU 10059; b, MHHNU 9888). Bars =2 cm.

Fig. 26. Fresh basidiomata of representative species in Ramaria ser. Luteae. a–b R. sanguinixantha (Holotype, a, MHHNU 10842; b, MHHNU 10863). Bars =2 cm.

Commentary: Ramaria magnibotrys resembles R. rubrievanescens and R. rubropurpurea, as all three species exhibit stout stipes and relatively large basidiomata with creamy to purplish-red apices. However, R. rubropurpurea changes color to dark red, with apices that display a deep purplish-red hue that is slow to fade. Ramaria rubrievanescens, on the other hand, has smaller basidiospores (L_m = 11.8 µm, Marr & Stuntz 1973).

13. Ramaria orientalibotrytis P. Zhang & G.W. Li, sp. nov. Fig. 13 g−h and Fig. 43

MycoBank: MB 852961

Etymology: The proposed compound epithet “orientalibotrytis” refers to an eastern species that is similar to R. botrytis.

Diagnosis: Basidiomata are medium sized to large, with a white stipe and pallid wine-colored apices. Branches are light wine-colored and turn wine-colored when injured. Basidiospores are ornamented with longitudinal striae.

Type: CHINA, HUNAN PROVINCE: Chenzhou City, Yizhang County, Mangshan National Nature Reserve, Datang Keng, in a mixed forest of broad-leaved forest and bamboo, altitude 1000 m, 6 September 2007, P. Zhang 665 (Holotype, MHHNU 7165, GenBank Acc. No.: ITS = PP467123, nrLSU = PP493454).

Description: Basidiomata 6–12 cm high and 5–10 cm broad, cauliflower-shaped in outline. Stipe 4–6 × 3–5 cm, single, falsely fasciculate, to compound-connate, with a few abortive branchlets, off-white (1A1) with a hint of shell pink (8A3) upward, flesh (6B3) in age, slowly easily wine red (11D8) where handled or chaffed; flesh solid, off-white (1A1), in age suffused flesh color, moist but not gelatinous or slippery, drying soft, light in weight, easily penetrated. Major branches 4, terete, ascending. Branches in 5–6 ranks, tapering upward, with shell pink (8A3) longitudinally rugulose, turned to wine red (11D8) when hurt, axils "U" shaped, straight, or slightly curved, crowded, rugulose surface, internodes diminishing gradually at maturity. Apices short and rounded, dark red (11C8) color. Odor weakly fragrant; taste negligible.

Fig. 27. Fresh basidiomata of representative species in Ramaria ser. Nanisporae. a−b R. corallina (Holotype, a, MHHNU 11212; b, MHHNU 10541); c−d R. nanispora (c, HKAS 50198; d, MHHNU 11636); e−f R. rubrosporula (Holotype, e, MHHNU 9921; f, MHHN9918). Bars = 2 cm.

Fig. 28. Fresh basidiomata of representative species in Ramaria ser. Neoformosae. a−b R. aurantiformosa (Holotype, a, MHHNU 9180; b, MHHNU 11361); c−d R. aurantilinearis (Holotype, c, MHHNU 10756; d, MHHNU 10816); e R. aureoapicalis(Holotype, e, MHHNU 9328); f R. aureolinearis (Holotype, f, MHHNU 10533); g R. brunnescentiflava (Holotype, g, MHHNU 9290); h R. distinctissima (h, MHHNU 9980); i R. ephemeroderma (i, MHHNU 6601); j R. hilaris (j, MHKMU Yang409); k R. hypogaeoides (Holotype, k, MHHNU 9211); l, R. linearis (l, MHHNU 10003); m R. longiaurantia (Holotype, m, MHHNU 10806); n R. luteoaurantiaca (n, MHHNU 10235); o−p R. pseudoobtusissima (Holotype, o, MHHNU 10064; p, MHHNU 10071); q−r R. robustibrunnea (Holotype, q, MHHNU 9269; r, MHHNU 9271); s−t R. sinolargentii (Holotype, s, MHHNU 34000; t, MHHNU 34029); u−v R. sinosubtilis (Holotype, u, MHHNU 10060; v, MHHNU 9993); w−x R. conferta (Holotype, w, MHHNU 10088; x, MHHNU 9209). Bars = 2 cm.

Fig. 29. Fresh basidiomata of representative species in Ramaria ser. Pallidochraceus. a–b R. pallidochracea (Holotype, a, MHHNU 6753; b, MHHNU 6752). Bars = 2 cm.

Fig. 30. Fresh basidiomata of representative species in Ramaria ser. Pallidolilacinae. a–b R. pallidolilacina (Holotype, a, HKAS46112; b, MHHNU 10080) (Zhang et al. 2005). Bars = 2 cm.

Fig. 31. Fresh basidiomata of representative species in Ramaria ser. Pseudobotrytes. a–b R. rufoaurea (Holotype, a, MHHNU 10026; b, MHHNU 10007). Bars = 2 cm.

Fig. 32. Fresh basidiomata of representative species in Ramaria ser. Pseudocystidiophorae. a–b R. pseudocystidiophora (Holotype, a, MHHNU 10023; b, MHHNU 10825). Bars = 2 cm.

Basidiospores [60/2/2](9.0–)9.1–11.9(–12.9) × 4.0–6.2(–6.4) μm, Lm = 10.48 µm, lm = 4.90 µm, Q = (–1.64)1.71–2.50(–2.75), Qm = 2.06 ± 0.24, ellipsoid to subcylindrical, conspicuously striae in profile, strongly cyanophilous; Basidia 44–60 × 9–12 μm, clavate, thin-walled, hyaline, clamped at base; weakly cyanophilous; sterigmata 4, 6–8 μm, stout and straight. Subhymenium rudimentary. Hymenium thickening. Tramal hyphae of stipe 6–12 µm wide, hyaline, conspicuously clamped, thin to thick-walled (wall up to 0.5 µm thick), easily broken, interwoven, not agglutinated or adherent; ampulliform clamps up to 15 µm broad, thick-walled (wall up to 1 µm thick), with conspicuous ornamentation; gloeoplerous hyphae not observed. Tramal hyphae of upper branches up to 2–11 µm wide, hyaline, thick-walled, conspicuously clamped, not agglutinated or adherent, parallel and hardly branched inward, with cells cigar to barrel-shaped, outward more interwoven, gnarled, freely branched; ampulliform clamps common, not unusually thick-walled, unornamented; gloeoplerous hyphae not observed.

Ecology: Found in broad-leaved or coniferous forests; basidiomata occurs in summer and autumn.

Distribution: Known from central China (Hunan).

Additional material examined: CHINA, HUNAN PROVINCE: Chenzhou City, Yizhang County, Mangshan National Nature Reserve, Zeziping, in broad-leaved forest with Fagaceae, altitude 1290 m, P. Zhang 350 (MHHNU 6850).

Commentary: Ramaria orientalibotrytis is the only species found in Hunan Province, central China, characterized by basidiospores ornamented with striae. Typically, species within Ramaria are commonly found in temperate regions with mild climates and are rarely reported in subtropical areas. However, R. orientalibotrytis thrives in broad-leaved forests within distinct subtropical areas. It resembles R. botrytis but differs in having smaller basidiospores that turn wine-color when injured (14.2 × 5.2 µm, Franchi & Marchetti 2021). Compared to the basidiospores size of R. rubropurpurea (L_m = 14.15 µm, as measured in this study), R. orientalibotrytis has smaller basidiospores.

Fig. 33. Fresh basidiomata of representative species in Ramaria ser. Robustipurpureae. a–b R. robustipurpurea (Holotype, a, MHHNU 8877; b, MHKMU Muman575). Bars = 2 cm.

Fig. 34. Fresh basidiomata of representative species in Ramaria ser. Roseobrunneae. a–b R. densiapicalis (Holotype, a, MHHNU 10012; b, MHHNU 10549); c–d R. roseobrunnea (Holotype, c, d, MHHNU 31571). Bars = 2 cm.

Fig. 35. Fresh basidiomata of representative species in Ramaria ser. Sanguinipesae. a−b R. rubriapicalis (Holotype, a, MHHNU 11382; b, MHHNU 11268); c−d R. rubriattenuipes (c, MHHNU 6578; d, HKAS 54802); e−f R. sanguinipes (e, MHHNU 11250; f, MHHNU 11253). Bars = 2 cm.

Fig. 36. Fresh basidiomata of representative species in Ramaria ser. Semigelatinosae. a–b R. semigelatinosa (Holotype, a, MHHNU 11310, b, MHHNU 10311). Bars = 2 cm.

Fig. 37. Fresh basidiomata of representative species in Ramaria ser. Sparsiaggregatae. a–b R. sparsiaggregata (Holotype, a, MHHNU 10016; b, MHHNU 10017). Bars = 2 cm.

Fig. 38. Fresh basidiomata of representative species in Ramaria ser. Spinulosae. a−b R. purpurea (Holotype, a, MHHNU 11267; b, MHHNU 9879); c−d R. purpureivariabilis (Holotype, c, d, MHHNU 31737); e−f R. sinospinulosa (Holotype, e, MHHNU 10788; f, MHHNU 10808). Bars = 2 cm.

Fig. 39. Fresh basidiomata of representative species in Ramaria ser. Zebrisporae. a−b R. linearioides (a, MHHNU 10046; b, MHHNU 10082); c−d R. zebrispora (c, MHHNU 11272; d, MHHNU 6621). Bars = 2 cm.

Fig. 40. Microscopic features of R. flavopurpurea (MHHNU 6595). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm

14. Ramaria rubropurpurea P. Zhang & J. Zhong, sp. nov. Fig. 13 i−j and Fig. 44

MycoBank: MB 852944

Etymology: rubropurpurea, from rubro = red color, and purpurea = purple color, proposed because this species has wine-colored basidiomata that turn red when injured.

Diagnosis: The apices are wine to grayish red in color and turn wine-red when bruised. The basidiospores are ornamented with longitudinal striae; hyphae and basidia base with clamps.

Type: CHINA. SICHUAN PROVINCE: Ganzi, Kangding City, Gongga Mountain, in Quercus semecarpifolia and Picea asperata forest, altitude 4000 m, 7 September 2021, P. Zhang 4346 (Holotype, MHHNU 10846, GenBank Acc. No.: ITS = PP467118, nrLSU = PP493449, tef1-α = PP480343).

Fig. 41. Microscopic features of R. luteoliparva (MHHNU 10013). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm

Fig. 42. Microscopic features of R. magnibotrys (MHHN 11364). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm

Fig. 43. Microscopic features of R. orientalibotrytis (MHHNU 7165). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Description: Basidiomata 14 cm high and 8 cm broad, big-sized. Stipe stout, up to 2.5 × 7 cm, single, falsely fasciculate, to compound-connate, smooth, with a few abortive branchlets, off-white (1A1) to cream (4A3), slowly turned to dark red (11C8) to wine red (11D8) where handled; flesh off-white (1A1), flesh to fibrous, drying soft, easily penetrated. Major branches several, indistinct. Branches in 2–4 ranks, short and crowded; dark red (11C8) to wine red (11D8) upward, concolor with stipe below; axils narrow rounded; internodes all short. Apices short and crowded, to grayish magenta to wine red (11D8). Odor weakly fragrant; taste sweet.

Basidiospores [60/2/2](12.0–)13.0–16.0(–17.0) × 5.0–8.0 µm, L_m = 14.15 µm, l_m = 6.25 µm, Q = (1.75–)1.86–2.80(–3.00), Q_m = 2.25 ± 0.25, subcylindrical, conspicuously striae in profile, strongly cyanophilous; Basidia 60–80 × 8–13 μm, clavate, clamped at base; sterigmata 1–4, mostly 4, 3–7 µm, stout and straight. Hymenium unthickening; Subhymenium rudimentary, hyphal. Tramal hyphae of stipe 6–12 µm wide, hyaline, conspicuously clamped, thin to thick-walled, interwoven, not agglutinated or adherent; ampulliform clamps up to 15 µm broad, usually not thick-walled; gloeoplerous hyphae not observed. Tramal hyphae of upper branches slightly narrowly, hyaline, thin walled, conspicuously clamped, not agglutinated or adherent, parallel and tightly interwoven; ampulliform clamps occasional, unornamented; gloeoplerous hyphae not observed.

Fig. 44. Microscopic features of R. rubropurpurea (MHHNU 10864). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm

Ecology: Found in broad-leaved or coniferous forests; basidiomata occurs in summer and autumn.

Distribution: Known from southwestern China (Sichuan, Yunnan, Xizang).

Additional materials examined: CHINA, SICHUAN PROVINCE: Ganzi, Kangding City, Gongga Mountain, in broad-leaved forest with Fagaceae, altitude 3500 m, 1 August 2012, P. Zhang 1211 (MHHNU 7711); same city, Mugecuo, in forest dominated by Abies and Pinaceae, altitude 3800 m, 21 August 2017, P. Zhang 2715 (MHHNU 9215); same city, Sewulong Village, within a Picea forest, altitude 3450 m, 5 September 2018, P. Zhang 3597 (MHHNU 10097), P. Zhang 3598(MHHNU 10097), P. Zhang 3601(MHHNU 10101), P. Zhang 3603(MHHNU 10103), P. Zhang 3604(MHHNU 10104); same location, Gongga Mountain, in the high-altitude Quercu and Picea forests, altitude 3900 m, 5 September 2021, P. Zhang 4267 (MHHNU 10767), P. Zhang 4270 (MHHNU 10770), P. Zhang 4271(MHHNU 10771), P. Zhang 4273(MHHNU 10773), P. Zhang 4274(MHHNU 10774), P. Zhang 4275(MHHNU 10775); same location, 6 September 2021, P. Zhang 4290 (MHHNU 10790), P. Zhang 4301 (MHHNU 10301), P. Zhang 4302 (MHHNU 10302), P. Zhang 4313 (MHHNU 10313), P. Zhang 4318 (MHHNU 10318); same location, 8 September 2021, P. Zhang 4245 (MHHNU 10745), P. Zhang 4250 (MHHNU 10750), P. Zhang 4278 (MHHNU 10778);same location, 9 September 2021, P. Zhang 4269 (MHHNU 10769). YUNNAN PROVINCE: Shangri-La City, Haba Snow Mountain, Mianshaba, altitude 3100 m, 13 August 2008, Li-P. Tang 620 (HKAS 54851); same city, in the mixed forest with Quercus and Picea. altitude 3429 m, 10 August 2020, Jian W. Liu MF368 (MHHNU 11648); Lijiang, Yulong County, Yulong Snow Mountain, altitude 3700 m, 26 July 2017, P. Zhang 2797 (MHHNU 9297).

Commentary: Ramaria magnibotrys P. Zhang & J. Zhong and R. rubrievanescens Marr & D.E. Stuntz closely resemble R. rubropurpurea. These species are characterized by their large basidiomata, stout stipes, reddish-purple apices, and striped basidiospores. Compared to R. rubropurpurea, R. magnibotrys exhibits relatively lighter-colored apices and pale stain color on bruising, with slightly smaller basidiospores. In contrast, R. rubropurpurea has deeper red apices that retain their hue even when dry compared to R. rubrievanescens. Specimens collected from some locations exhibit pale to pinkish apices similar to R. rubrievanescens. This discrepancy may arise from environmental factors such as moisture or low light conditions causing the fading of the reddish-purple pigments. Nonetheless, basidiospores of R. rubrievanescens are smaller (L_m = 11.8 µm).

15. Ramaria sinobotrytis P. Zhang & J. Zhong, sp. nov. Fig. 13 k−l and Fig. 45

MycoBank: MB 852971

Etymology: The proposed compound epithet “sinobotrytis” refers to the species collected in China and being similar to R. botrytis.

Diagnosis: The apices and young branches exhibit a pale red-purple color, while the stipe surface gradually turns slightly lemon yellow upon handled. Basidiospores are ornamented with longitudinal striae and measure over 18 μm, with hyphae and basidia base showing clamps.

Type: CHINA, SICHAUN PROVINCE: Ganzi, Kangding City, Gongga Township, Sewurong Village, altitude 3700 m, 26 August 2017, P. Zhang 2768 (Holotype, MHHNU 9268, GenBank Acc. No.: ITS = PP467122, nrLSU = PP493453, tef1-α = PP480347).

Description: Basidiomata 12 cm high and 15 cm broad. Stipe 3 × 5 cm, single, falsely fasciculate, tapering gradually downward, to compound-connate, with a few abortive branchlets, off-white (1A1) to cream (4A3), with a hint of pink upward; nearly not changing color or slowly and slightly changing color to lemon yellow (4A3) when handle; flesh solid, off-white (1A1), flesh to fibrous, drying soft, easily penetrated. Major branches several, compound at base. Branches in 2–5 ranks, short and crowded; flesh (6B3) to brownish orange (6C3), with a hint of red-purple hue; axils narrowly rounded; internodes all short. Apices rounded, short, crowded, rose (12A4) to grayish orange (6B5). Odor weakly fragrant; taste negligible.

Basidiospores [60/2/2]16.0–21.0(–22.0) × 6.0–9.0 µm, L_m = 18.2 µm, l_m = 7.3 µm, Q = (2.11–)2.13–3.00(–3.14), Q_m = 2.51 ± 0.27, long-ellipsoid, conspicuously striae in profile, strongly cyanophilous; Basidia 50–85 × 9–18 µm, clavate, clamped at base; sterigmata mostly 4, 4–9 μm, stout and straight. Subhymenium rudimentary, hyphal. Hymenium unthickening. Tramal hyphae of stipe 6–14 µm wide, hyaline, conspicuously clamped, thin to thick-walled, interwoven, not agglutinated or adherent; ampulliform clamps up to 18 µm broad, usually not thick-walled; gloeoplerous hyphae not observed. Tramal hyphae of upper branches up to slightly narrowly, hyaline, thin walled, conspicuously clamped, not agglutinated or adherent, parallel and hardly branched inward; ampulliform clamps occasional; gloeoplerous hyphae not observed.

Fig. 45. Microscopic features of R. sinobotrytis (MHHNU 9268). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Fig. 46. Microscopic features R. violaceoaurantia (MHHNU 10074). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Ecology: Found in broad-leaved or coniferous forests; basidiomata occurs in summer and autumn.

Distribution: Known from southwestern China (Sichuan).

Commentary: This species is similar to R. botrytis, whose basidiospores measure 14.2 × 5.2 µm (Franchi & Marchetti 2021). Although only one specimen was collected, it includes two additional young basidiomata. Moreover, it features the largest basidiospores among current species within the genus Ramaria, making it easily distinguishable from others. Our phylogenetic analysis indicates that the species exhibits the closest phylogenetic distance to R. botrytis.

16. Ramaria violaceoaurantia P. Zhang & J. Zhong, sp. nov. Fig. 13 m−o and Fig. 46

MycoBank: MB 852939

Etymology: violaceoaurantia = violacea + aurantia = orange, proposed because this species has purple to orange color on the apices.

Diagnosis: The apices are wine to purple-orange in color, turning brown-yellow when bruised, basidiospores are ornamented with longitudinal striae, and hyphae and basidia base feature clamps.

Type: CHINA. SICHUAN PROVINCE: Ganzi, Kangding City, Waze Township, in Picea forest, altitude 3800 m,1 September 2018, P. Zhang 3574 (Holotype, MHHNU 10074, GenBank Acc. No.: ITS = PP467115, nrLSU = PP493446, tef1-α = PP480340).

Description: Basidiomata 11–12 cm high and 10–16 cm broad, big-sized. Stipe stout, up to 2–4 × 2–3 cm, single, falsely fasciculate, to compound-connate, smooth, with a few abortive branchlets, off-white (1A1) to cream (4A3), slowly turned to lemon yellow (3B8) to brownish yellow (5B3) where handled; flesh off-white (1A1), flesh to fibrous, drying soft, easily penetrated. Major branches several, indistinct. Branches in 2–4 ranks, short and crowded; grayish orange (6B5) to brownish yellow (6C8) upward, concolor with stipe below; axils narrow rounded; internodes all short. Apices short and crowded, to reddish lilac (14B4) to brownish yellow (6C8). Odor weakly fragrant; taste sweet.

Basidiospores [60/2/2]12.0–16.0 × 5.0–7.0(–8.0) µm, L_m = 14.35 µm, l_m = 5.83 µm, Q = (1.71–) 1.92–3.10(–3.20), Q_m = 2.40 ± 0.33, subcylindrical, conspicuously striae in profile, strongly cyanophilous; Basidia 60–80 × 8–13 μm, clavate, clamped at base; sterigmata 1–4, mostly 4, 3–7 µm, stout and straight. Hymenium unthickening; Subhymenium rudimentary, hyphal. Tramal hyphae of stipe 6–12 µm wide, hyaline, conspicuously clamped, thin to thick-walled, interwoven, not agglutinated or adherent; ampulliform clamps up to 15 µm broad, usually not thick-walled; gloeoplerous hyphae not observed. Tramal hyphae of upper branches slightly narrowly, hyaline, thin walled, conspicuously clamped, not agglutinated or adherent, parallel and tightly interwoven; ampulliform clamps occasional, unornamented; gloeoplerous hyphae not observed.

Ecology: Found in broad-leaved or coniferous forests; basidiomata occurs in summer and autumn.

Distribution: Known from southwestern China (Sichuan, Yunnan, Xizang).

Additional materials examined: CHINA, SICHUAN PROVINCE: Ganzi, Kangding City, Waze Township, in Picea forest, altitude 3800 m,1 September 2018, P. Zhang 3578 (MHHNU 10078). YUNNAN PROVINCE: Lijiang City, Baisha Town, Yulong County, Yulong Snow Mountain, in broad-leaved forest, altitude 3760 m, 17 August 2019, Jian W. Liu 1765; same location, altitude 3805 m, 24 August 2019, Jian W. Liu 1797. XIZANG AUTONOMOUS REGION: Linzhi, Sejila Mountain, Bayi District, Lulang Town, in a subalpine forest dominated by Quercus, altitude 3400 m, 26 August 2018, P. Zhang 3499 (MHHNU 9999), P. Zhang 3500 (MHHNU 10000), P. Zhang 3501 (MHHNU 10001), P. Zhang 3502 (MHHNU 10002).

Commentary: Ramaria violaceoaurantia is distinguished by its prominently orange apices. It resembles European species R. parabotrytis Franchi & M. Marchetti, North American R. aurantiiramosa Marr & D.E. Stuntz (≡ R. botrytis var. aurantiiramosa) Marr & D.E. Stuntz, and R. rubripermanens Marr & D.E. Stuntz. Among these similar species, R. aurantiiramosa also exhibits orange apices, but with slightly narrower basidiospores (12–16 × 4–6 µm, L_m = 13.5 µm, l_m = 4.7 µm, Q_m = 2.87; Marr & Stuntz 1973; Franchi & Marchetti 2021, 2018). Ramaria parabotrytis and R. rubripermanens display more distinctly reddish-purple apices in maturity and have slightly smaller basidiospores. Although variability in basidiospores size exists within the R. botrytis complex, our molecular evidence indicates significant genetic divergence among these species, which are also geographically distant from each other.

Ramaria ser. Brunnescentiaurantiae P. Zhang & J. Zhong, ser. nov.

MycoBank: MB 852731

Diagnosis: Basidiomata are pale orange to bright orange, with stipe, branches, and apices concolorous with each other. Bruised results in a purple-brown coloration. Hyphae and basidia base are nearly devoid of clamps.

Type species: R. brunescensaurantius P. Zhang & J. Zhong.

Etymology: brunescensaurantius = brunescens (turned to brown when injured) + aurantius (orange).

Commentary: This series comprises three species collected in China, in addition to other species found elsewhere including R. amyloidea Marr & D.E. Stuntz, R. claviramulata Marr & D.E. Stuntz (synonymous: R. celerivirescens Marr & D.E. Stuntz), R. magnifica Schild, and its variety R. magnifica var. sardiniensis.

17. Ramaria aurantibrevispora P. Zhang & J. Zhong, sp. nov. Fig. 14 a–b and Fig. 47

MycoBank: MB 853372

Etymology: aurantibrevispora = “auranti-” + “brevispora”, referring to the orange-colored basidiomata of the species and its short basidiospores.

Diagnosis: Small-sized basidiomata, yellowish orange to salmon-colored; short branches with yellow apices or concolorous with the branches; bruised turn grayish purple; occasional pseudo clamps in hyphae, with clamps absent at basidia base.

Type: CHINA, YUNNAN PROVINCE: Chuxiong, Nanhua County, Dazhongshan Forestry Farm, in a forest dominated by Fagaceae, altitude 2200m, 10 October 2021, Jian W. Liu 2549 (Holotype, MHHNU 11650, GenBank Acc. No.: ITS = PP599212, nrLSU = PP593674).

Description: Basidiomata 8–9 cm high and 5–5.5 cm broad, obtriangular to fusiform in outline. Stipe 1–4 × 0.5–2 cm, single to falsely fasciculate, tomentose at base, pale orange (5A3) to salmon (6A4) color, turned to gray-lilac (15B2) when hurt. Major branches few, terete, slender, pale orange (5A3) to salmon (6A4); flesh orange white (5A2). Branches in 3−5 ranks, ascending, terete, melon yellow (5A6) when young, pale orange (5A3) to salmon (6A4) at age; flesh more pallid, brittle to brittle-fibrous, gelatinous; internodes diminishing gradually at maturity; axils narrowly rounded throughout. Apices are short and acute, double-dichotomous, concolorous with branches. Odor and taste faintly fabaceous.

Fig. 47. Microscopic features of R. aurantibrevispora (MHHNU 11650). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Basidiospores [62/2/2] (7.2–)7.5–9.8(–10.6) × (3.7–)3.9–4.8(–5.2) µm, L_m = 8.68 µm, l_m = 4.33 µm, Q = (1.65–)1.74–2.29(–2.42), Q_m = 2.01 ± 0.16, subcylindrical to ellipsoid, slightly roughened in profile; hilar not prominent; ornamentation of small, randomly scattered flat warts. Basidia 30−45 × 7−19 μm, clavate, without clamps; sterigmata 4, long, slender, straight. Subhymenium extensive, hyphae. Hymenium unthickening. Tramal hyphae of stipe 5-17 μm wide, hyaline, without clamps, tightly interwoven, not agglutinated or adherent; ampulliform clamps up to 16 μm broad, not unusually thick-walled, with delicate ornamentation; gloeoplerous hyphae not observed. Tramal hyphae of upper branches up to 4-15 μm wide, hyaline, thin walled, without clamped, loosely parallel, more inflated inward, agglutinated to adherent; ampulliform clamps up to 17 μm broad, gloeoplerous hyphae occasionally.

Ecology: Found in broad-leaved forests. Basidiomata occurs in summer and autumn.

Distribution: Known from southwestern China (Yunnan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Chuxiong, Zixi Mountain, in a forest with Lithocarpus and Pinus yunnanensis, altitude 2400 m, 2 August 2001, P. Zhang 97 (MHHNU 6597); same location, Xianren Valley, in broad-leaved forest, altitude 2400 m, 24 August 2023, P. Zhang 5039 (MHHNU 11539); same city, Nanhua County, Dazhongshan Forestry Farm, in mixed broadleaf-coniferous forest, altitude 2314 m, 11 August 2017, Jian W. Liu 864 (MHHNU 11642).

Commentary: In our two-gene phylogenetic analysis, the species most closely related to R. aurantibrevispora, which has its type locality in Italy, are R. magnifica Schild, and it's two variety, R. magnifica var. magnifica and var. sardinian (now called R. sardiniensis). They all share characteristics such as 1) absence clamps in hyphae and basidia base; 2) basidiospores average length less than 10 µm; 3) gray-purple bruised; 4) small to medium sized basidiomata with yellowish orange to salmon colors; 5) axils with narrow acute angles, V-shaped. Ramaria magnifica var. magnifica and R. sardiniensis differ in the former having larger basidiomata, more yellow-orange hues, and slightly larger basidiospores. However, both of these species show molecular sequence differences from R. aurantibrevispora, with morphological distinctions being subtle, particularly in basidiospores size, where R. aurantibrevispora basidiospores have an average length of less than 9 µm.

In Dr. Zhang Ping's postdoctoral thesis (2010), this species is considered a variant of R. cyaneigranosa, a species described by Marr & Stuntz in the United States. Ramaria cyaneigranosa exhibits an intermediate color between red and salmon in its basidiomata, rough basidiospores surfaces, and intensely cyanophilic contents in basidia. Ramaria cyaneigranosa comprises three varieties: var. cyaneigranosa with yellow branch tips, var. persicina with pink or salmon-colored branches and yellow spots on the tips, and var. elongata. Morphologically, the basidiomata color and basidiospores size of R. aurantibrevispora from Zixi Mountain in Yunnan are indeed similar to R. cyaneigranosa, but they show distant molecular phylogenetic relationships.

Another species resembling R. aurantibrevispora is R. brunnescentiaurantia, both being small to medium sized with yellowish orange basidiomata. The distinguishing feature of R. brunnescentiaurantia is its longer basidiospores (10.35 × 5.13 µm, as measured in this study).

18. Ramaria brunnescentiaurantia P. Zhang & J. Zhong, sp. nov. Fig. 14 c−d and Fig. 48

MycoBank: MB 853235

Etymology: brunnescentiaurantia = “brunnescenti” + “aurantia”, referring to the species having red to orange basidiomata that turn brown when injured.

Diagnosis: Basidiomata are salmon to orange, with stipe or main branches elongated and concolorous with branches. Bruised results in a grayish purple coloration. Hyphae and basidia base are without clamps.

Type: CHINA, SICHUANG PROVINCE: Ganzi, Kangding City, Gongga Mountain, in Quercus semecarpifolia and Picea asperata forest, 5 September 2021, P. Zhang 4272 (Holotype, MHHNU 10772, GenBank Acc. No.: ITS = PP467243, nrLSU = PP493552, tef1-α = PP480418).

Fig. 48. Microscopic features of R. brunnescentiaurantia (MHHNU 10772). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Description: Basidiomata 10.5 cm high and 8 cm broad, linear, long fusiform in outline. Stipe 1.4 × 2 cm, fasciculate or with a very small point of union at the base, or solitary, smooth, pale orange (5A3) to salmon (6A4), turned to brownish violet (11F8) when hurt. Major branches 2–4, ascending, terete, pale orange (5A3) to salmon (6A4); flesh white (1A1) to pallid orange (5A2) near hymenium; Branches in 3−5 ranks, ascending, more or less terete, pale orange (5A3) to salmon (6A4); flesh more pallid, brittle to brittle-fibrous, more or less gelatinous; internodes diminishing gradually at maturity; axils narrowly rounded throughout, "V" shaped. Apices broadly rounded, double-dichotomous, concolorous with branches. Odor and taste faintly fabaceous.

Basidiospores [60/2/2] (8.8–)9.1–11.2(–11.9) × (4.3–)4.6–5.9(–6.3) µm, L_m = 10.35 µm, l_m = 5.13 µm, Q = (1.68–)1.81–2.23(–2.42), Q_m = 2.02 ± 0.14, ellipsoid, slightly roughened in profile; hilar prominent; ornamentation of small, randomly scattered flat warts. Basidia 40−55 × 6−10 μm, clavate, without clamps; sterigmata 4, long, slender, curved. Subhymenium extensive, hyphae. Hymenium unthickening. Tramal hyphae of stipe 6-15 μm wide, hyaline, without clamps, loosely parallel, not agglutinated or adherent; ampulliform clamps up to 17 μm broad, not unusually thick-walled, with delicate ornamentation; gloeoplerous hyphae not observed. Tramal hyphae of upper branches up to 5-15 μm wide, hyaline, thin walled, without clamped, loosely parallel, more inflated inward, more or less agglutinated to adherent; ampulliform clamps up to 18 μm broad, gloeoplerous hyphae occasionally.

Ecology: Found in broad-leaved, coniferous, or mixed forests. Basidiomata occurs in summer and autumn.

Distribution: Known from southwestern China (Yunnan, Sichuan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Shangri-La City, Haba Snow Mountain, Mianshaba, altitude 3100 m, 10 August 2008, Li-P. Tang 619 (HKAS 54850). SICHUANG PROVINCE: Ganzi, Kangding City, Gongga Mountain, in Quercus semecarpifolia and Picea asperata forest, altitude 4000 m, 7 September 2021, P. Zhang 4375 (MHHNU 10875); same location, altitude 3900 m, 6 September 2021, P. Zhang 4293 (MHHNU 10293), P. Zhang 4294 (MHHNU 10294), P. Zhang 4309 (MHHNU 10309), P. Zhang 4312 (MHHNU 10312).

Commentary: Ramaria brunnescentiaurantia closely resembles to R. aurantibrevispora (basidiospore 8.68 × 4.33 µm, as measured in this study), as both species have yellowish orange basidiomata, non-clamped hypha and basidia, stain purplish-gray upon injury. However, R. brunnescentiaurantia is distinguished by its longer basidiospores and typical inhabit in high-altitude regions.

Distinguishing R. brunnescentiaurantia from R. magnifica and its variants primarily involves slight differences in basidiospores length. In Zhang's 2010 postdoctoral thesis (Zhang 2010), it is proposed that R. brunnescentiaurantia may be synonymous with R. cyaneigranosa var. elongata. However, this species exhibits only moderate cyanophilous reaction within its basidia.

19. Ramaria verrucosiflava P. Zhang & J. Zhong, sp. nov. Fig. 14 e−f and Fig. 49

MycoBank: MB 853191

Etymology: verrucosiflava = verrucosus + flava, proposed because this species has fine warts basidiospores and orange-yellow basidiomata.

Diagnosis: Basidiomata salmon to orange yellow; basidia and hyphae with clamps; basidiospores small sized and finely warty; apices yellow.

Type: CHINA, YUNNAN PROVINCE: Shangri-La, Pudacuo, Shudu Lake, in the forest with Quercus semecarpifolia, altitude 3600 m, 15 August 2018, P. Zhang 3408 (Holotype, MHHNU 9908, GenBank Acc. No.: ITS = PP467288, nrLSU = PP493590, tef1-α = PP480448).

Description: Basidiomata 10 cm high and 6 cm broad, medium sized. Stipe 3 × 3 cm, single to falsely fasciculate, tapering gradually downward, smooth, off-white (1A1) to melon yellow (5A6), with occasional abortive branchlets; flesh solid, off-white (1A1) to orange white (5A2), moist, but not gelatinous or slippery. Major branches several, stout and short, terete, ascending, concolorous with stipe downward, concolorous with branches upward, not changing color on bruising. Branches in 3−4 ranks, dichotomous, terete, ascending, orange white (5A2) to melon yellow (5A6); flesh pale orange (5A3), soft; internodes length ratio diminishing gradually at maturity; axils mostly acute, both U-shaped and V-shaped. Apices crowded, double dichotomous, vivid yellow (2A8) to yellowish orange (4A8); Odor faintly penetrating; taste negligible.

Basidiospores [22/2/2] (7.4−)8.0−8.9(−9.3) × (2.8−)3.1−3.8(−3.9) μm, L_m = 8.59 µm, l_m = 3.61 µm, Q = 2.16–2.82(–3.16), Q_m = 2.39 ± 0.23, ellipsoid, nearly smooth in profile; hilar appendix prominent; ornamentation finely or or extremely obscurely undulate in profile. Basidia 40−55 × 7−11 µm, clavate, clamped; sterigmata 4, long, spindly, straight. Subhymenium rudimentary, pseudoparenchymatous. Hymenium thickening. Tramal hyphae of stipe 4-12 µm wide, usually inflated, hyaline, thin- to thick walled, loosely parallel, interwoven, not agglutinated or adherent; ampulliform clamps up to 23 μm broad, thick-walled, with coarse ornamentation; gloeoplerous hyphae not observed; Tramal hyphae of upper branches up to 3-14 µm wide, hyaline, thin-to thick-walled, clamped, secondarily septum, loosely parallel, not agglutinated or adherent; ampulliform clamps up to 22 µm broad, thick-walled, unornamented; gloeoplerous hyphae not observed.

Fig. 49. Microscopic features of R. verrucosiflava (MHHNU 9908). a basidiospores; b basidia. Scale bars: a =10 µm, b =20 µm.

Ecology: Found in broad-leaved forests; basidiomata occurs in summer and autumn.

Distribution: Known from southwest China (Yunnan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Shangri-La, Pudacuo, Shudu Lake, in the forest with Quercus semecarpifolia, altitude 3600 m, 15 August 2018, P. Zhang 3404 (MHHNU 9904), P. Zhang 3419 (MHHNU 9919), P. Zhang 3420 (MHHNU 9920); Qujing City, Qilin District, near Longyuan Zhumushan Wind Farm, in broad-leaved forest, altitude 2250 m, 19 August 2022, P. Zhang 4706 (MHHNU 11206); Chuxiong, Zixi Mountain, in mixed forests dominated by Fagaceae and Pinaceae, altitude 2250 meters, 24 August 2022, P. Zhangping 4784 (MHHNU 11284).

Commentary: Ramaria verrucosiflava bears a strong resemblance to R. amyloidea (basidiospore 8.9 × 3.6 µm, Marr & Stuntz 1973). Both species feature similarly sized and nearly smooth basidiospores, along with clamped basidia and hyphae. However, R. amyloidea is characterized by a distinctive flesh band of pale brown hyphae visible in the basal region of a radially sectioned stipe and amyloid flesh. Our two-gene phylogenetic analysis indicates a close relationship between these species, confirming their distinct taxonomic identities despite their similarities.

Ramaria ser. Conjunctipesae P. Zhang & J. Zhong, ser. nov.

MycoBank: MB 852741

Etymology: “conjunc” derives from “joined”, and “-ipes” from “foot”, “conjunctipes” refers to the stipe base being joined together.

Type species: R. conjuncta (Peck) Corner, Monograph of Clavaria and allied Genera, (Annals of Botany Memoirs No. 1): 566 (1950)

Diagnosis: Gregarious habit; basidiomata are small to medium sized, ranging in color from salmon, yellow to pallid orange; stipe flesh is tough; basidiospores are small and broadly (L_m<10 μm), small warty; hyphae and basidia lack clamps; no color change upon bruising.

Commentary: This series includes four species collected in China. Additional species found elsewhere include R. orithamnus (Berk.) R.H. Petersen, R. ignicolor Corner, R. conjunctipes (Coker) Corner and its varieties R. conjunctipes var. tsugensis Marr & D.E. Stuntz, and R. conjunctipes var. sparsiramosa Marr & D.E. Stuntz.

20. Ramaria flaviaggregata P. Zhang & J. Zhong, sp. nov. Fig. 15 a–b and Fig. 50

MycoBank: MB 853299

Etymology: flaviaggregata = flava+ aggregate, referring to the species having pale yellow-colored basidiomata and a fasciculate habit.

Diagnosis: Basidiomata are small to medium sized and pastel yellow in color; apices are dichotomous, ranging from yellow to pale yellow; basidiospores are small and broadly, slightly roughened; flesh of the stipe and branches is freshy-fibrous, turning to brownish yellow when bruised; hyphae and basidia base lack clamps.

Type: CHINA, HUNAN PROVINCE: Chenzhou City, Rucheng County, Daping Town, Jiulongjiang National Forest Park, in broad-leaved forest, altitude 100 m, 2 October 2020, P. Zhang 4065 (Holotype, MHHNU 10565, GenBank Acc. No.: ITS = PP467224, nrLSU = PP493538).

Description: Basidiomata 8−10 cm high and 4−6 cm broad, coralline form, fasciculate habit. Stipe 3 × 1.5 cm, individual stipes cluster together at base, or branching directly from the ground; off-white at base (1A1), surface pale yellow (2A3) to cream (4A3) upward; slightly changing color to brownish yellow (5B3) on bruising; surface tomentose or longitudinally gnarled and rugose; occasionally with abortive branches; context of stipe off-white (1A1) when fresh, freshy-fibrous, occasionally hollow, drying hard, cartilaginous. Branches in 4−5 ranks, major branches 4−several, commonly polychotomous, branches above mostly dichotomous; cream (4A3) to butter yellow (4A5) when young, pale yellow (2A3) to pastel yellow (2A4) at age; axils broadly rounded, mostly U-shaped, mostly bifid near apices; internodes diminishing gradually at maturity, lower internodes up to 5 cm; branches slender, commonly 3–8 mm, terete, subparallel; context of main branches and upper branches freshy-fibrous, off-white (1A1), slightly changing color to brownish yellow (5B3) on bruising. Apices acute, short, claw-like, 1–4 mm long, dichotomous, butter yellow (4A5) when young, turned to pall yellow (2A3) at age, the tips slightly brownish yellow (5B3) or concolorous with branches. Odor and taste faintly fabaceous.

Fig. 50. Microscopic features of R. flaviaggregata (MHHNU 10565). a basidiospores; b basidia, a =10 µm, b = 20 µm

Basidiospores [60/2/2] (6.0–)6.5–8.9(–9.6) × (4.2–)4.5–5.7(–6.1) µm, L_m = 7.55 µm, l_m = 5.05 µm, Q = (1.20–)1.30–1.68 (–1.81), Q_m = 1.50 ± 0.14, ovate to shortly ellipsoidal; finely ornamented with linearly lobed, cyanophilous warts; hilar appendix small but conspicuous. Basidia 30–45 × 5–9 μm, clavate, without clamps at base; sterigmata 2–4, mostly 4, 5–9 µm long, incurved or straight, slightly divergent. Hymenium unthickening, 50 µm. Subhymenium extensive, 10–30 µm thick, hyphae, interwoven, 2–5 µm diam, thin-walled, without clamps. Contextual hyphae: Tramal hyphae of stipe 5–11 μm wide, loosely interwoven, inflating slightly, terminal swollen hyphae wide to 13 µm, with moderately ornamented; tramal hyphae of branches central hyphae slightly lysing, terminal swollen hyphae wide to 12 µm, with smooth and thin wall; ampulliform hyphae rarely; without clamps; gloeoplerous hyphae occasionally.

Ecology: Found in broad-leaved forests. Basidiomata occurs in summer and autumn.

Distribution: Known from southwestern and central China (Yunnan, Hunan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Xishuangbanna City, Menglun Town, near Kino Gurry, in the Castanopsis forest on the ridge, altitude 1000 m, 14 August 2005, P. Zhang 452 (MHHNU 6952), P. Zhang 454 (MHHNU 6954); same city, bought from mushroom market, 11 October 2006, P. Zhang 583 (MHHNU 7083); Kunming City, Near Qiongzhu Temple, altitude 1900 m, 26 July 2011, Nian K. Zeng 929 (FUMU 570).

Commentary: The geographic distribution of R. flaviaggregata is extensive, ranging from the southwest to the central China. It belongs to the R. conjunctipes complex group, but its coloration sets it apart within this group. Unlike the pink to salmon hues observed in R. conjunctipes or mustard yellow of R. lorithamnus, R. flaviaggregata exhibits a distinctive off-white to pale yellow color. Ramaria pallidoflava is similar in color and basidiospores size (7.57 × 5.15 µm, as measured in this study), making differentiation challenging. Subtle distinctions include R. flaviaggregata having lower axils for the stipe, a slender stipe, and a slightly more browning color on bruising.

21. Ramaria microsporoflava P. Zhang & J. Zhong, sp. nov. Fig. 15 c−d and Fig. 51

MycoBank: MB 853296

Etymology: “microsporoflava” refers to the species being yellow colored with small basidiospores.

Fig. 51. Microscopic features of R. microsporoflava (MHHNU 31326). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Diagnosis: Basidiomata are small, ranging from salmon to pallid orange in color; apices are vivid yellow colored; basidiospores are small and broadly, ovate to shortly ellipsoidal, and nearly smooth; The color does not change upon bruising; hyphae and basidia base lack clamps.

Type: CHINA, YUNNAN PROVINCE: Mouding County, Huafo Mountain, altitude 2500 m, 23 August 2018, Zuo H. Chen 31326 (Holotype, MHHNU 31326, GenBank Acc. No.: ITS = PP467216, nrLSU = PP493530).

Description: Basidiomata 4–6 cm high and 0.3–2 2.5 cm broad, usually shrubby with a fasciculate habit. Stipe 0.3–2 × 0.5 cm, rarely single, usually fasciculate, tapering gradually downward; off-white (1A1) at base, pale yellow (2A3) to pale orange (5A3) above; not changing color on bruising; covered with a white tomentum where buried in the substratum; without abortive branches; context of stipe off-white (1A1), freshy-fibrous, more or less gelatinous when fresh, occasionally hollow, drying brittle and with distinct chalky-friable properties. Branches in 3−6 ranks, major branches commonly polychotomous, and branches above dichotomous, elongated, slightly divergent; cream (4A3) when young, salmon to butter yellow (6A4–4A5) at age; axils acute to V-shaped, mostly bifid near apices; internodes diminishing gradually at maturity, lower internodes up to 2 cm; branches slender, commonly 3-9 mm, seldom over to 2 cm, terete to flattened, compact and subparallel; context of main branches freshy-fibrous, off-white (1A1), context of upper branches freshy-fibrous, more or less gelatinous, pall orange (5A3), bruising or staining reactions slight or entirely absent. Apices acute, crowded, dichotomous, vivid yellow (2A8). Odor and taste faintly fabaceous.

Basidiospores [60/2/2] (6.6–)7.4–9.2(–9.5) × (4.4–)4.8–6.2(–6.5) µm, L_m = 8.25 µm, l_m = 5.43 µm, Q = (1.28–)1.35–1.70(–1.82), Q_m = 1.52 ± 0.11, ovate to shortly ellipsoidal; finely ornamented with linearly lobed, cyanophilous warts; hilar appendix small but conspicuous. Basidia 35–50 × 7–11 μm, clavate, without clamps; sterigmata 1-4, mostly 4, irregular, 5–10 µm long, incurved or straight, slightly divergent, occasionally two sterigmata connate at base. Hymenium unthickening. Subhymenium extensive, 25–30 µm thick, hyphae, interwoven, 3–5 µm diam, thin-walled, without clamps. Contextual hyphae: parallel to subparallel in both base and branches, not-inflating or inflating slightly, up to 4–12 μm wide; outer hyphae of the stipe collapsing and forming a densely stratified parallel tissue, somewhat cornenous; the central context of both stipe and branches lysing, hyphae walls frequently undulate, mostly smooth and slightly gelatinized, thin to moderately thick-walled, strongly cyanophilous; ampulliform hyphae rarely, 8–13 µm, nearly smooth; without clamps; gloeoplerous hyphae uncommon, mostly of narrow diam, 2–3 µm.

Ecology: Found in broad-leaved and coniferous mixed forests. Basidiomata occurs in summer and autumn.

Distribution: Known from southwestern China (Yunnan).

Additional material examined: CHINA, YUNNAN PROVINCE: Binchuan County, Jizu Mountain, with broad-leaved, coniferous, or mixed forests, altitude 2300 m, 29 August 2022, P. Zhang 4813 (MHHNU 11313).

Commentary: The species most similar to R. microsporopflava is R. ignicolor Corner. Corner described R. ignicolor as having non-inflated hyphae without clamps and firmly gelatinous walls. Franchi notes that R. ignicolor is easily distinguished among yellow to orange Ramaria species due to its unique macroscopic and microscopic characteristics: the branches are usually hollow, the hyphae lack clamps, and they possess gelatinous hyphae.

The basidiomata of R. microsporoflava (4–6 × 0.6–2.5 cm) is smaller than those of R. ignicolor described by Franchi (13 × 4–7 cm) (2021) and more akin to the dimensions reported by Corner from Bresadola's records (height 4 cm) (1950). Although the average basidiospores length (L_m = 8.25 µm) is slightly longer than that described by Franchi (L_m = 7.9 µm), the basidiospores width is similar (l_m = 5.43 µm), being slightly larger than the dimensions provided by Corner (7–8.5 × 4–5 µm). Despite minimal differences in basidiospores size, significant differences exist in basidiomata coloration. The Latin term "ignicolor" denotes a fiery red color. Although Corner did not collect this species, observations of dried specimens indicate a light orange-ochraceous color, suggesting that fresh basidiomata may also be darker. In contrast, R. microsporoflava tends to exhibit yellow to salmon colors when fresh, transitioning to a light leather yellow when dried specimens. Franchi's color illustrations confirm that the Italian-described R. ignicolor indeed possesses a more vibrant orange hue, with a higher degree of gelatinization in both the stipe and branch flesh.

22. Ramaria pallidoflava P. Zhang & J. Zhong, sp. nov. Fig. 15 e−f and Fig. 52

MycoBank: MB 853297

Etymology: pallidoflava = pallidus + flava, referring to the species having pall yellow-colored basidiomata.

Fig. 52. Microscopic features of R. pallidoflava (MHHNU 11652). a basidiospores; b basidia. Scale bars: a = 10 µm, b = 20 µm.

Diagnosis: Basidiomata are small to medium sized and pallid yellow in color; apices are dichotomous, ranging from yellow to pallid yellow; basidiospores are small and broadly, slightly roughened; flesh of stipe and branches is freshy-fibrous, turning brownish yellow when bruised; hyphae and basidia base lack clamps.

Type: CHINA, YUNNAN PROVINCE: Nanhua County, Dazhongshan Forestry Farm, in a forest dominated by Fagaceae, altitude 2400 m, 11 October 2021, Jian W. Liu 2576 (Holotype, MHHNU 11652, GenBank Acc. No.: ITS = PP467223, nrLSU = PP493537).

Description: Basidiomata 14 cm high and 10 cm broad, usually single to fasciculate habit. Stipe 2 × 5 cm, terete when young, flatten due to hollow flesh at maturity, clustered together at base; off-white at base (1A1), surface pale yellow (2A3) to cream (4A3) upward; slightly changing color to brownish yellow (5B3) on bruising; surface tomentose or longitudinally gnarled and rugose; occasionally with abortive branches; context of stipe off-white (1A1) when fresh, freshy-fibrous, occasionally hollow, drying hard, cartilaginous. Branches in 3−5 ranks, major branches commonly polychotomous, cluster below, branches above mostly dichotomous; cream (4A3) when young, pale yellow (2A3) to pastel yellow (2A4) at age; axils broadly rounded, mostly U-shaped, mostly bifid near apices; internodes diminishing gradually at maturity, lower internodes up to 4 cm; branches slender, commonly 3–10 mm, terete or flattened, crowded, subparallel; context of main branches and upper branches freshy-fibrous, off-white (1A1), bruising or staining reactions entirely absent. Apices acute, short, 1–4 mm long, dichotomous, concolorous with branches. Odor and taste faintly fabaceous.

Basidiospores [60/2/2] (6.7–)6.9–8.6(–8.8) × (4.4–)4.5–6.3(–6.6) µm, L_m = 7.57 µm, l_m = 5.15 µm, Q = (1.25–)1.32–1.66(–1.78), Q_m = 1.47 ± 0.11, ovate to shortly ellipsoidal; finely ornamented with linearly lobed, cyanophilous warts; hilar appendix small but conspicuous. Basidia 35–45 × 6–11 μm, clavate, without clamps at base; sterigmata 2-4, mostly 4, 3–6 µm long, incurved or straight, slightly divergent. Hymenium unthickening, 50 µm. Subhymenium extensive, 10–20 µm thick, hyphae, interwoven, 2–5 µm diam, thin-walled, without clamps. Contextual hyphae: Tramal hyphae of stipe 6–13 μm wide, loosely interwoven, inflating slightly, terminal swollen hyphae wide to 14 µm, with moderately ornamented; tramal hyphae of branches central hyphae slightly lysing, terminal swollen hyphae wide to 13 µm, with smooth and thin wall; ampulliform hyphae rarely; without clamps; gloeoplerous hyphae occasionally.

Ecology: Found in a broad-leaved forest. Basidiomata occurs in summer and autumn.

Distribution: Known from southwestern and central China (Yunnan, Hunan, Hubei).

Additional materials examined: CHINA, HUNAN PROVINCE: Zhangjiajie City, Sangzhi County, Badagong Mountain National Nature Reserve, Tianping Mountain, in a broad-leaved forest, altitude 1400 m, 27 July 2022, Zuo H. Chen 33498 (MHHNU 33498). HUBEI PROVINCE: Enshi, Hefeng County, Wuyang Township, Hewan Village, altitude 1000 m, 24 September 2019, Zuo H. Chen 31726 (MHHNU 31726).

Commentary: Ramaria pallidoflava belongs to the R. conjunctipes complex. Distinguished by its coloration, which resembles that of R. flaviaggregata, it leans towards a pale-yellow hue. Within the R. conjunctipes complex, apart from R. lorithamnus, most species exhibit ranging from pink to salmon. Ramaria pallidoflava and R. flaviaggregata (basidiospore 7.55× 5.05 µm, as measured in this study) share similar coloration and basidiospores size, making differentiation challenging, with subtle distinctions such as the higher axils of the stipe in R. pallidoflava and a stouter stipe.

23. Ramaria sinoconjunctipes R.H. Petersen & M. Zang, Acta bot. Yunn.12(1): 50, 1990. Fig. 15 g−h and Fig. 53

Description: Basidiomata 6–8 cm high and 2–3 cm broad, usually shrubby with a fasciculate habit. Stipe 1–2 × 2–5 cm, rarely single, clustered together at base, flatten due to hollow flesh, or branching directly from the ground; off-white at base, pale yellow to pale orange above; not changing color on bruising; surface nearly smooth; without abortive branches; context of stipe off-white, freshy-fibrous, more or less gelatinous when fresh, occasionally hollow, drying cartilaginous. Branches in 3−6 ranks, major branches commonly polychotomous, and branches above dichotomous; cream when young, salmon to butter yellow at age; axils acute to U-shaped, mostly bifid near apices; internodes diminishing gradually at maturity, lower internodes up to 4 cm; branches slender, commonly 3–6 mm, seldom over to 1 cm, terete, crowded, subparallel; context of main branches freshy-fibrous, off-white, context of upper branches freshy-fibrous, more or less gelatinous, pall orange, bruising or staining reactions slight or entirely absent. Apices acute, 1–3 mm long, crowded, dichotomous, vivid yellow. Odor and taste faintly fabaceous.

Fig. 53. Basidiospore features of R. sinoconjunctipes (MHHNU 11645). Scale bar: a = 10 µm

Basidiospores [60/2/2] (6.6–)7.0–9.0(–9.7) × (4.1–)4.3–6.1(–6.3) µm, L_m = 7.13 µm, l_m = 5.17 µm, Q = (1.27–)1.32–1.74(–1.86), Q_m = 1.51 ± 0.31, ovate to shortly ellipsoidal; finely ornamented with linearly lobed, cyanophilous warts; hilar appendix small but conspicuous. Basidia 40–50 × 7–12 μm, clavate, without clamps at base; sterigmata 2-4, mostly 4, 3–6 µm long, incurved or straight, slightly divergent, occasionally two sterigmata connate at base. Hymenium unthickening, 55 µm. Subhymenium extensive, 25–30 µm thick, hyphae, interwoven, 2–5 µm diam, thin-walled, without clamps. Contextual hyphae: Tramal hyphae of stipe 4–9 μm wide, hyaline, loosely interwoven, inflating slightly, terminal swollen hyphae wide to 13 µm, with moderately ornamented; tramal hyphae of branches up to 2–6 μm wide, central hyphae slightly lysing, terminal swollen hyphae wide to 12 µm, with smooth and thin wall; ampulliform hyphae rarely; without clamps; gloeoplerous hyphae occasionally.

Ecology: Found in a coniferous mixed forest. Basidiomata occurs in summer and autumn.

Distribution: Known from southwestern China (Yunnan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Lijiang, Black-White Water, 7 September 1983, Petersen 47274 (isotype, HKAS 20064); same city, Yulong County, Yulong Snow Mountain, altitude 3700 m, 25 July 2017, P. Zhang 2795 (MHHNU 9295); same location, 2019; same city, Baisha Township, Lijiang Alpine Botanical Garden, altitude 3569 m, 29 September 2019, Jian W. Liu 2029 (MHHNU 11645); Chuxiong, Nanhua County, Dazhongshan Forestry Farm, in a forest dominated by Fagaceae, altitude 2356 m, 10 October 2021, Jian W. Liu 2425 (MHHNU 11649); Deqin County, Baima Snow Mountain Nature Reserve, in Picea forest, altitude 3750 m, 17 August 2018, P. Zhang 3440 (MHHNU 9940), P. Zhang 3444 (MHHNU 9944), P. Zhang 3454 (MHHNU 9954).

Commentary: The isotype specimens HKAS17873 (R. distinctissima) and HKAS17855 (R. rubriattenuipes), both identified by Petersen in 1989, were incorrectly assigned. Only the isotype specimen HKAS20064 of R. sinoconjunctipes was correctly identified. This taxonomic complex is characterized by basidiomata with fasciculate, attenuate stipes, hyphae without clamps, short and wide basidiospores. The fruiting bodies of most included taxa exhibit pink to salmon branches and yellow apices, but R. lorithamnus has basidiomata that are yellow ochre all over. Among those with a salmon to yellow color pattern, only R. ignicolor is found in Europe.

Marr & Stuntz's (1973) concept of R. conjunctipes was probably not contain with typical material from eastern North American. They described branches as hollow and flesh as watery-gelatinous, characteristics not typical of eastern North American specimens. While this might seem like a minor distinction, the flesh in R. sinoconjunctipes is not gelatinous (microscopically hyphae are not agglutinated or adherent), but rather watery-hydrophanous. It is this quality, not gelatinization, that distinguishes it. Petersen also did not recall seeing gelatinous flesh in those taxa, although hollow branches are commonly observed.

Sharda (1983) and Thind (1985b) described their concept of R. conjunctipes from the eastern Himalayas. They noted fruitbody colors as “light salmon to salmon yellow colored, tips light yellow” and provided basidiospores dimensions of 7–8.5(–9.5) × 4–5(–5.5) µm. Their concept likely corresponds to R. sinoconjunctipes, with the only differing feature being the presence of sclerotic basidia in the Himalayan specimens, which was not observed in Petersen's collections.

Also considered part of this taxonomic complex is R. petersenii K.S. Thind & Sharda (1985b). The fruiting bodies are characterized by very sparse branching, with a long, single, slender stipe that is not fasciculate. Basidiospores are wide (7–11.5 × 5–6.5 µm), and septa lack clamps. The color of the fruiting bodies was described as “orange, white to light orange in the central part, with pale orange apices.”

Wu has described R. hypogaea Q.X. Wu (Wu 1987), whose basidiomata display fasciculate single stipes and a stature similar to R. sinoconjunctips. However, the fruitbody coloration of R. sinoconjunctipes, becoming increasingly orange towards the apices. Despite these differences, R. hypogaea certainly belongs to the same taxonomic complex as R. sinoconjunctips.

Ramaria ser. Flavae (Franchi & M. Marchetti) P. Zhang & J. Zhong, stat. nov.

Fungi Non Delineati, Raro vel Haud Perspecte et Explorate Descripti aut Definite Picti 16: 26 (2001)

Synonym: Ramaria sect. Flavae Franchi & M. Marchetti

Type species: Ramaria flava (Schaeff.) Quél., Fl. mycol. France (Paris): 466 (1888).

Diagnosis: Basidiomata range from pale yellow to dark yellow; frequently brunnescent or rubribrunnescent; basidiospores are often smooth or finely warty; 4) hyphae may be clamped or clampless.

Commentary: This series includes six species collected in China, as well as additional species such as R. obtusissima (Peck) Corner, R. pseudoflava, R. flava, R. barenthalensis, R. edwinii, R. magnipes, R. rasilispora, and its variety R. rasilispora var. rasilispora, as well as R sanguinea. Here are the characteristics of some species within this series:

(1) R. highlandensis Petersen: The basidiomata primarily yellow to yellow-brown colors, with bright yellow apices; browning on bruising; Relatively pronounced aromatic odor, especially when the flesh is exposed. Basidiospores measure 11.5−13.7× 4.0−4.7 μm (Q = 2.67−3.27, Q_m = 2.95, L_m = 12.43 μm), and the type specimen from the United States.

(2) R. magnipes Marr & D.E. Stuntz and its varieties (R. magnipes var. magnipes Marr & D.E and R. magnipes var. albidior R.H. Petersen): This species emerges in spring and features cauliflower-like basidiomata with a yellowish-green hue. The stipes are stout, and the basidiospores measure 11.8 × 3.7 μm. Clamps are present.

(3) R. rasilispora Marr & Stuntz and its varieties (R. rasilispora var. rasilispora Marr & D.E. Stuntz, and R. rasilispora var. scatesiana Marr & D.E): This species appears in spring, with basidiomata ranging from light yellow to ochre-yellow. The basidiospores are smaller than 11 μm, and clamps are present.

(4) R. rasilisporoides Exeter: This species is relatively large cauliflower-like basidiomata, up to 28 × 20 cm. They are light yellow to pale orange in color. The stipe's flesh has a strong amyloid reaction. Basidiospores measure 9.1 × 3.6 μm, and clamps are present. The type specimen of originates from North America.

Additionally, Corner and others considered R. obtusissima (Peck) Corner to be a species with smooth basidiospores. Its basidiomata ranges from white to buff-yellow or ochraceous, and clamps are present. The type specimen of originates from North America. When Corner reorganized the species in 1950, it was considered a mixed group with smooth basidiospores. Petersen reexamining the type specimen in 1967, also noted the basidiospores surface to be nearly smooth. However, in 1992, Schild conducted further studies on the type specimens and observed basidiospores using electron microscopy. It was discovered that the basidiospores of the type specimen of R. obtusissima are not smooth but have verrucose ornamentation. Consequently, Schild designated those species with truly smooth basidiospores as a new species, named R. schildii.

24. Ramaria barenthalensis Franchi & M. Marchetti, Riv. Micol. 61: 199 (2019). Fig. 16 a−b and Fig. 54

Description: Basidiomata 14 × 13 cm, medium sized, usually obtriangular to broadly fusiform in outline. Stipe 2 × 3 cm, single to falsely fasciculate, tapering gradually downward, smooth, without abortive branches, off-white where stipe protected below, cream to pale brown above, weakly brunnescent, sometimes with pale brown to dark brown spots; flesh off-white, mottled hydrophanous, not gelatinous or slippery. Major branches 2−4, stout, terete, ascending, cream to yellow when young, mellowing to salmon or cream-ochre in age; flesh white, solid, not hydrophanous. Branches in 2−5 ranks, stout, ascending, concolorous with upper major branches below; internodes elongating through development, diminishing gradually at maturity; axils broad rounded throughout (both U-shaped and V-shaped). Apices dichotomous, stout, stiffly digitate, short and crowded, yellow to dark ochraceous, sometimes dark brown. Odor negligible or faintly fragrant; taste negligible or fabaceous.

Fig. 54. Basidiospore features of R. barenthalensis (MHHNU 10091). Scale bar: a = 10 µm.

Basidiospores [30/2/2] (10.9–)11.2–13.6 × 4.4–5.5(–5.7) µm, L_m = 12.23 µm, l_m = 4.93 µm, Q = (2.17–)2.21–2.79(–2.92), Q_m = 2.48 ± 0.19, sub-cylindrical, slightly roughened in profile; hilar appendix prominent, ornamentation of small warts and short ridges randomly placed. Basidia 50-85 × 7-14 μm, elongate-clavate, clamped at base; sterigmata 4, long, stout, slightly curved. Subhymenium rudimentary. Hymenium thickening. Tramal hyphae of stipe 5−16 μm wide, hyaline, clamped common, thick-walled (wall up to 1 μm thick), interwoven, not agglutinated or adherent; ampulliform clamps up to 18 μm broad, asymmetrical, not unusually thick-walked, with coarse ornamentation; gloeoplerous hyphae not observed. Tramal hyphae of upper branches 4−18 μm wide, inflated especially in medullary regions, hyaline, conspicuously clamped, thick-walled (wall up to 1 μm thick), loosely parallel; ampulliform inflations rare, gloeoplerous hyphae not observed.

Ecology: In the forests with Fagaceae, Picea, Quercus semecarpifolia, or in other mixed forests.

Distribution: Known from southwest, central, and northeast China and at high altitude location (Yunnan, Sichuan, Xizang, Hubei, Jinlin).

Additional materials examined: CHINA, YUNNAN PROVINCE: Deqin County, Baima Snow Mountain Nature Reserve, altitude 3750 m, 17 August 2018, P. Zhang 3446 (MHHNU 9946); Qujing City, Qilin District, near Longyuan Zhumushan Wind Farm, altitude 2250 m, 19 August 2022, P. Zhang 4719 (MHHNU 11219). SICHUAN PROVINCE: Ganzi, Kangding City, Gongga Mountain, in Quercus semecarpifolia and Picea asperata forest, altitude 3900 m, 6 September 2021, P. Zhang 4310 (MHHNU 10810), P. Zhang 4319 (MHHNU 10819); same location, 5 September 2021, P. Zhang 4268 (MHHNU 10768); same location, in the Abies forest, altitude 3500 m, 2 September 2021, P. Zhang 3586 (MHHNU 10086), P. Zhang 3591(MHHNU 10091); same city, Gongga Township, in a mixed forest, altitude 3700 m, 24 August 2017, P. Zhang 2757 (MHHNU 9257), P. Zhang 2754 (MHHNU 9254); same city, Mugecuo, in broad-leaved forest with Fagaceae, altitude 3500 m, 1 August 2012, Ping Zhang 1208 (MHHNU 7708). XIZNAG AUTONOMOUS REGION: Qamdo City, Changdu County, Ranwu Lake, in Picea forest, altitude 3,900 meters, 22 August 2018, P. Zhang 3479 (MHHNU 9979). HUBEI PROVINCE: Enshi, Hefeng County, Xiaping Township, Aimin Village, altitude 1100 m, 23 September 2019, Zuo H. Chen 31718 (MHHNU 31718). JILIN PROVINCE: Tonghua County, Baijifeng National Forest Park, in broadleaf forest, altitude 800 m, 6 August 2022, P. Zhang 4698 (MHHNU 11198); Jilin City, Jiaohe City, Shasong Mountain, Kangdalazi, altitude 1000 m, 9 August 2017, P. Zhang 2810 (MHHNU 9310). Liaodong Peninsula, in broad-leaved forest, 30 August 2018, P. Zhang 3609 (MHHNU 10109).

25. Ramaria flavolongispora P. Zhang & J. Zhong, sp. nov. Fig. 16 c−d and Fig. 55

MycoBank: MB 853370

Etymology: flavolongispora= flava+ longa + spora, referring to the species having yellow-colored basidiomata and long basidiospores.

Diagnosis: Salmon-colored basidiomata, massive stipe, long basidiospores, crowded apices.

Type: CHINA, YUNNAN PROVINCE: Chuxiong, Nanhua County, Dazhongshan Forestry Farm, in a forest dominated by Fagaceae, altitude 2400 m, 11 October 2021, Jian W. Liu 2607 (Holotype, MHHNU 11653, GenBank Acc. No.: ITS = PP467278).

Description: Basidiomata 11 cm high and 10 cm broad, medium sized, usually obovate to obpyriform in outline. Stipe 4 × 5 cm, single to falsely fasciculate, stout, tapering gradually downward, longitudinally rugose, often with abortive branchlets, off-white (1A1) to cream (4A3), weakly brunnescent when injury; flesh solid, off-white (1A1), brittle. Major branches 2−4, stout, terete, ascending, milk white (1A2) to cream (4A3); flesh off-white (1A1), solid, not hydrophanous. Branches in 2−5 ranks, ascending, milk white (1A2) to cream (4A3), weakly brunnescent; internodes all short when young, elongating below through development, diminishing gradually at maturity; axils narrowly rounded throughout, usually V-shaped. Apices double-dichotomous, crowded, narrowly rounded, digitate at maturity, concolorous with branches. Odor negligible or faintly fragrant; taste negligible or fabaceous.

Basidiospores [61/1/1] (11.1–)11.4–14.2(–15.4) × (3.1–)3.4–4.7(–4.9) µm, L_m = 12.60 µm, l_m = 4.05 µm, Q = (2.50–)2.63–3.79(–4.11), Q_m = 3.15 ± 0.35, cylindrical, roughened in profile, strongly cyanophilous; hilar appendix prominent, ornamentation of warts and short ridges randomly placed. Basidia 48-70 × 5-12 µm μm, elongate-clavate, clamped at base; sterigmata 4, stout and long. Subhymenium rudimentary. Hymenium thickening. Tramal hyphae of stipe 5−13 μm wide, hyaline, conspicuously clamped, tightly interwoven, not agglutinated or adherent; ampulliform clamps up to 20 μm broad, with extensive, coarse, stalactitiform ornamentation; gloeoplerous hyphae rare. Tramal hyphae of upper branches 5−13 μm wide, inflated especially in medullary regions, hyaline, conspicuously clamped, tightly interwoven, ampulliform clamps up to 19 μm broad, gloeoplerous hyphae occasional.

Fig. 55. Microscopic features of R. flavolongispora (MHHNU 11653). a basidiospores; b basidia. Scale bars: a =10 µm, b =20 µm.

Ecology: Solitary or gregarious on soil in broad-leaved forests; basidiomata occurs in summer and autumn.

Distribution: Known from southwest China (Yunnan).

Additional material examined: CHINA, YUNNAN PROVINCE: Wenshan, Malipo County, Mengdong Yao Township, Ailao Mountain range, in broad-leaved forest, altitude 1250 m, 29 July 2017, G. Wu 2273 (HKAS123587).

Commentary: Ramaria flavolongispora is closely related to R. magnipes and R. rasilispora. However, R. flavolongispora possesses the largest basidiospores among the three species (R. magnipes and R. rasilispora having basidiospores measuring 11.8 × 3.7 µm and 9.5 × 3.4 µm respectively). Additionally, the apices and branches of R. flavolongispora share the same color, whereas the apices of the other species are more or less brown in color.

26. Ramaria hepatica P. Zhang & G.W. Li, sp. nov. Fig. 16 e−f and Fig. 56

MycoBank: MB 853367

Etymology: “hepatica” refers to the species turning brown when injured or damaged.

Diagnosis: Basidiomata are cauliflower-like in appearance, start pale yellow when young and turns red brown as it matures, stipe is massive and turns dark brown with a yellowish tinge when bruised, basidiospores are cylindrical.

Type: CHINA, HUNAN PROVINCE: Zhangjiajie City, Yongding District, Tianmenshan National Forest Park, in a broadleaf forest, altitute 800 m, 9 September 2021, P. P. Zhang 4416 (Holotype, MHHNU 10916, GenBank Acc. No.: ITS = PP467277, nrLSU = PP493583, tef1-α = PP482597).

Description: Basidiomata 8–10 cm high and 5–9 cm broad, small to medium sized, usually obovate to obpyriform in outline. Stipe 3–4 × 2–3 cm, single to falsely fasciculate, rounded at base, smooth upward, often with abortive branchlets, off-white (1A1) to pale yellow (2A3) when young, grayish orange (6B5) in age, slowly brunnescent when injury. flesh solid, off-white (1A1), slowly brunnescent on handing, brittle, not gelatinous or slippery. Major branches 4−8, stout, terete, ascending, pale yellow (2A3) to salmon (6A4) when young or flesh (6B3), easily mellowing to light brown (6D6) to brown (6E8) through development; flesh off-white (1A1), solid, not hydrophanous. Branches in 3−6 ranks, ascending, concolorous with upper major branches, internodes all short when young, elongating below through development, diminishing gradually at maturity; axils narrowly rounded throughout, usually (both U-shaped and V-shaped). Apices are double-dichotomous, crowded when young, acute, short, digitate at maturity, finally concolorous with branches, light brown (6D6) to brown (6E8) in age. Odor negligible or faintly fragrant; taste negligible or fabaceous.

Basidiospores [40/2/2] 11.0–13.0 × 4.0–5.0 µm, L_m = 12.00 µm, l_m = 4.68 µm, Q = (2.17–)2.20–3.00(–3.25), Q_m = 2.59 ± 0.29, ellipsoid to cylindrical, roughened in profile, strongly cyanophilous; hilar appendix prominent, yellow to brown, 6-8 μm long, ornamentation of warts and short ridges randomly placed. Basidia 42–65 × 9–12 μm, elongate-clavate, clamped at base; sterigmata 4, stout and long. Subhymenium rudimentary. Hymenium thickening. Tramal hyphae of stipe 5−14 μm wide, hyaline, conspicuously clamped, tightly interwoven, not agglutinated or adherent; ampulliform clamps up to 16 μm broad, with extensive, coarse, stalactitiform ornamentation; gloeoplerous hyphae rare. Tramal hyphae of upper branches 5−16 μm wide, inflated especially in medium sized regions, hyaline, conspicuously clamped, tightly interwoven, ampulliform clamps up to 16 μm broad, gloeoplerous hyphae occasional.

Ecology: Solitary or gregarious on soil in broad-leaved forests; basidiomata occurs in summer and autumn.

Distribution: Known from central China (Hunan).

Additional materials examined: CHINA, HUNAN PROVINCE: Zhangjiajie City, Yongding District, Tianmenshan National Forest Park, in a broadleaf forest, altitute 800 m, 9 September 2021, P. Zhang 4415 (MHHNU 10915); Changde City, Shimen County, Hupingshan Town, Hupingshan National Nature Reserve, in a broad-leaved forest, altitude 1000 m, 3 October 2010, Zuo H. Chen 30754 (MHHNU 30754).

Commentary: Ramairia brunneicontusa R. H. Petersen, R. pseudoflava Franchi & M. Marchetti, and R. flava greatly resemble species in Ramaria, characterized by yellow basidiomata and clamps. Ramaria hepatica and R. flava share a similar basidiospores size, but the former exhibits a pale yellow basidiomata when young, turning red brown with age, whereas the latter’s basidiomata remains consistently yellow. Ramaria pseudoflava has smaller basidiospores (10.7 × 4.0 µm; Franchi & Marchetti 2021) and its basidiomata are gold yellow at maturity without changing color. Ramaria hepatica and R. brunneicontusa both change color when bruised or handled, but R. brunneicontusa has longer basidiospores measuring (13.0–16.6 × 4.7–6.1 μm; Petersen 1989).

Fig. 56. Microscopic features of R. hepatica (MHHNU 10916). a basidiospores; b basidia. Scale bars: a =10 µm, b =20 µm.

27. Ramaria laevibrunnea P. Zhang & Zai W. Ge, sp. nov. Fig. 16 g−h and Fig. 57

MycoBank: MB 853371

Etymology: laevibrunnea = “laevi-” + “brunnea”, referring to the brown-colored basidiomata of the species and its nearly smooth basidiospores.

Diagnosis: Yellow to salmon basidiomata; clamped basidia and hyphae; basidiospores are nearly smooth; turns dark brown when bruised or handled.

Type: CHINA, YUNNAN PROVINCE: Dali, Yongping County, Beidou Township, Yuntai Mountain, in a mixed forest by Keteleeria and Pinus, altitude 2400 m 22 August 2017, Zai W. Ge 4028 (Holotype, HKAS 105942 GenBank Acc. No.: ITS = PP467280, nrLSU = PP493584).

Description: Basidiomata 9 cm high and 8 cm broad, medium sized, usually subcircular to commashaped in outline. Stipe 3 × 2 cm, falsely fasciculate, nearly smooth, tapering gradually downward, without abortive branchlets at maturity; off-white (1A1) to pastel yellow (2A4) downward, ivory (4B3) to cinnamon brown (6D6) upward, weakly brunnescent when hurt or handle; flesh off-white (1A1), solid, pale yellow (2A3), not gelatinous, or slippery. Major branches several, ascending, terete, concolorous with stipe downward, concolorous with branches upward, weakly brunnescent when hurt or handle. Branches in 2−4 ranks, terete, cream (4A3) to sunshine yellow (2A5) colored, maturity to cinnamon brown (6D6); flesh paler than hymenium, soft; internodes all short when young, elongating from below, diminishing gradually upward at maturity; axils broadly rounded throughout (U shaped). Apices rounded and pale yellow (2A3) when young, acute and cinnamon brown (6D6) in age; Odor faintly penetrating; taste negligible.

Basidiospores [31/1/1] (11.8–)12.3–14.9(–16.9) × (4.1–)4.2–5.3(–5.8) µm, L_m = 13.70 µm, l_m = 4.82 µm, Q = (2.19–)2.46–3.36(–3.54), Q_m = 2.86 ± 0.28, cylindrical to ellipsoid, smooth or extremely obscurely undulate in profile; wall up to 0.2 µm thick; hilar appendix small, not prominent, truncate; ornamentation none. Basidia 30-42 × 6–12 μm, clavate, short, clamped; sterigmata (2–) 4, short, spindly, straight. Subhymenium rudimentary, pseudoparenchymatous. Hymenium thickening. Tram-al hyphae of stipe 4-12 µm wide, usually inflated, hyaline, thick walled (wall up to 1.5 µm thick locally), tightly interwoven, not agglutinated or adherent; ampulliform clamps up to 16 μm broad, thick-walled (wall up to 1.5 µm thick), with coarse ornamentation; gloeoplerous hyphae not observed; Tramal hyphae of upper branches up to 3–10 µm wide, hyaline, thin-to thick-walled (wall up to 1 µm thick locally), clamped, secondarily septum, interwoven, not agglutinated or adherent; ampulliform clamps up to 14 µm broad, thick-walled (wall up to 1 µm thick), coarse ornamentation; gloeoplerous hyphae not observed.

Ecology: Found in Picea and broad-leaved mixed forests; basidiomata occurs in summer and autumn.

Distribution: Known from southwest China (Yunnan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Dali, Dali City, Xianyangfeng, in a mixed forest, altitude 2800 m, 12 August 2018, P. Zhang 3375 (MHHNU 9875); Chuxiong, Nanhua County, Dazhongshan Forestry Farm, in mixed broadleaf-coniferous forest, altitude 2218 m, 9 August 2017, Jian W. Liu 781 (MHHNU 11639).

Commentary: Ramaria laevibrunnea is distinguished as the first smooth-spores Ramaria with young salmon coloration that darkens to brown with age. The species closely resemble R. laeviformosoides, both sharing nearly smooth basidiospores, clamps, and similar salmon basidiomata. However, R. laevibrunnea noticeably turns dark brown, while R. laeviformosoides only exhibits slight browning.

Fig. 57. Microscopic features of R. laevibrunnea (MHHNU 11639). a basidiospores; b basidia. Scale bars: a =10 µm, b =20 µm.

28. Ramaria pallidobrunnea P. Zhang & J. Zhong, sp. nov. Fig. 16 i−j and Fig. 58

MycoBank: MB 853369

Etymology: pallidobrunnea = pallida + brunnea, refers to the basidiomata of the species being light brown colored.

Diagnosis: Basidiomata are yellow, easily and extensively turning brunnescent; basidia are clamped and long; basidiospores are smooth, virtually unornamented.

Type: CHINA, SICHUAN PROVINCE: Ganzi, Kangding County, Mugecuo, in forest dominated by Abies and Pinaceae, altitude 3800 m, 21 August 2017, P. Zhang 2713 (Holotype, MHHNU 9213, GenBank Acc. No.: ITS = PP467270, nrLSU = PP493576, tef1-α = PP480439).

Description: Basidiomata 20 cm high and 13 cm broad, massive, circular to broadly obovate in outline. Stipe 4 × 4 cm, small to large, rounded to lappering sharply to acute or rooting base, smooth, with a few abortive stumps high on stipe or low on lower branches, off-white (1A1) to yellowish white (2A2), very easily brunnescent; flesh off-white (1A1), slowly brunnescent when cut, solid, mottled-hydrophanous, but not gelatinous or slippery, often with a yellowish band suffused near base, dry flesh firm but easily penetrated. Major branches 2−4, massive, hardly terete, ascending, pale yellow (2A3) to flesh (6B3) when young, easily mellowing to pastel yellow (2A4) through development; flesh off-white (1A1), solid, not hydrophanous. Branches in 3−6 ranks, stout, ascending, concolorous with upper major branches, internodes all short when young, elongating below through development, diminishing gradually at maturity; axils rounded throughout. Apices are sometimes finely divided, acute, double-dichotomous, usually gross, molar-like, short-digitate at maturity, pastel yellow (2A4) to chrome yellow (3A8), off-white (1A1) where protected. Odor negligible or faintly fragrant; taste negligible or fabaceous.

Basidiospores [50/2/2](12.0–)13.0–15.2(–15.5) × 4.2–5.6(–5.8) µm, L_m = 14.05 µm, l_m = 4.81 µm, Q = (2.60–)2.67–3.33, Q_m = 2.92 ± 0.21, boletoid to cylindrical, flattened adaxially or with a suprahilar swelling, smooth in profile; wall up to 0.2 μm thick, hilar appendix not prominent, ornamentation usually none, occasionally a suggestion of ill-defined, moderately cyanophilous markings of delicate ridges and small patches, in more or less longitudinal orientation. Basidia 95−115 × 10−11 μm, elongate-clavate, clamped at base; sterigmata 4, long, slender, straight. Subhymenium rudimentary. Hymenium thickening. Tramal hyphae of stipe 5−16 μm wide, hyaline, clamped, thick-walled (wall up to 1.5 μm thick), interwoven, not agglutinated or adherent; ampulliform clamps up to 18 μm broad, asymmetrical, not unusually thick-walked, with extensive, coarse, stalactitiform ornamentation; gloeoplerous hyphae rare, 3-5 μm, yellow- refringent. Tramal hyphae of upper branches 4−28 μm wide, inflated especially in medullary regions, hyaline, conspicuously clamped, thin to thick-walled (wall up to 1 μm thick locally), with common false clamps, loosely parallel, ampulliform inflations not observed, gloeoplerous hyphae occasional, 2.5-5 μm diam, yellow, refringent, with abrupt swellings.

Ecology: Found on soil in a broad-leaved and coniferous mixed forest; basidiomata occurs in summer and autumn.

Distribution: Known from southwest China (Yunnan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Shangri-La, Bitahai, in a subalpine forest, altitude 3800 m, 26 August 2020, Zuo H. Chen 32322 (MHHNU 32322); same city, Zhongdian County, Zhongdian Snow Mountain, 24 August 2000, Zhu L. Yang 3015 (HKAS 36561). SICHUAN PROVINCE: Ganzi, Xiangcheng County, Reda Township, altitude 3400 m, Zai W. Ge 123 (HKAS 45890). XIZANG AUTONOMOUS REGION: Linzhi City, Milin County, Zhaxiraodeng Village, in Picea forest, altitude 3040 m, 24 August 2018, Ping Zhang 3485 (MHHNU 9985); Leiwuqi, in a subalpine forest, altitude 3900 m, 24 August 2004, Zhu L. Yang 4355 (HKAS 45734); Jiangda County, altitude 3800 m, Zai W. Ge 229 (HKAS 46009).

Commentary: A taxonomic complex of nearly smooth-spored taxa within Ramaria includes North American species such as R. obtusissima var. obtusissima (Corner 1950) and R. highlandensis (Petersen 1989), which occur in summer, along with one western North American taxon and its varieties (R. magnipes var. magnipes, Marr & Stuntz 1973; and var. Albidior, Petersen & Scates 1988), known as vernal fruiters. Additionally, there are two European taxa (R. obtusissima var. Scandinavica, Petersen 1989; and an undescribed taxon), and one North American taxon exhibiting weakly amyloid stipe flesh but otherwise showing macro-chemical inertness.

The common features among R. pallidobrunnea include basidiomata displaying yellow or ochre-yellow tones, basidiospores with subtle roughness, and slight browning upon minor injury. Due to their morphological similarities, accurate differentiation of these species requires integrating molecular biological evidence and geographical factors.

Fig. 58. Microscopic features of R. pallidobrunnea (MHHNU 9213). a basidiospores; b basidia. Scale bars: a =10 µm, b =20 µm.

29. Ramaria verrucula P. Zhang & J. Zhong, sp. nov. Fig. 16 k−l and Fig. 59

MycoBank: MB 853360

Etymology: “verrucula” refers to the finely warty basidiospores of this species.

Diagnosis: Basidiomata are light yellow in color, with apices and stipe brunnescent reluctantly; stipe is massive; basidia are slightly short and conspicuously clamped; basidiospores’ ornamentation is usually fine.

Type: CHINA, HUBEI PROVINCE: Enshi, Hefeng County, Wuyang Township, Hewan Village, altitude 1000 m, 24 September 2019, Zuo H. Chen 31735 (Holotype, MHHNU 31735, GenBank Acc. No.: ITS = PP467272, nrLSU = PP493578, tef1-α = PP480441).

Description: Basidiomata 10 cm high and 7 cm broad, small to medium sized, usually obovate to obpyriform in outline. Stipe 3 × 6 cm, single to falsely fasciculate, large to stout, tapering gradually to the acute, pruinose at base, sometimes with very few abortive branchlets, off-white (1A1) to pale yellow (2A3); flesh solid, off-white (1A1), not gelatinous or slippery, reluctantly brunnescent on handing, not gelatinous or slippery. Major branches 2−4, massive, terete, ascending, pale yellow (2A3) when young or flesh (6B3), easily mellowing to grayish orange (6B5) through development; flesh off-white (1A1), solid, not hydrophanous. Branches in 3−6 ranks, ascending, concolorous with upper major branches, internodes all short when young, elongating below through development, diminishing gradually at maturity; axils narrowly rounded throughout. Apices cuspidate or double-dichotomous, crowded when young, sunshine yellow (2A5) to grayish orange (6B5), brownish in age. Odor negligible or faintly fragrant; taste negligible or fabaceous.

Basidiospores [58/2/2] (10.9–)11.8–13.5(–13.9) × (3.6–)3.9–5.3(–5.6) µm, L_m = 12.11 µm, l_m = 4.64 µm, Q = (2.19–)2.28–3.07(–3.37), Q_m = 2.64 ± 0.26, cylindrical; hilar appendix prominent, ornamentation usually fine. Basidia 35-70 × 5-11 μm, elongate-clavate, clamped at base; sterigmata 4, stout and long. Subhymenium rudimentary. Hymenium thickening. Tramal hyphae of stipe 5−14 μm wide, hyaline, conspicuously clamped, thick-walled (wall up to 1 μm thick), interwoven, not agglutinated or adherent; ampulliform clamps up to 18 μm broad, asymmetrical, with extensive, coarse, stalactitiform ornamentation; gloeoplerous hyphae rare. Tramal hyphae of upper branches 5−16 μm wide, inflated especially in medium sized regions, hyaline, conspicuously clamped, thin to thick-walled (wall up to 1 μm thick locally), loosely parallel, ampulliform clamps up to 16 μm broad, gloeoplerous hyphae occasional.

Ecology: Found in a broad-leaved and coniferous mixed forest; basidiomata occurs in autumn.

Distribution: Known from central China (Hubei).

Commentary: In the genus Ramaria, there are several species characterized by minutely warted basidiospores and yellow basidiomata. The species most closely related to this group is R. hepatica, which shares similarities in basidiospores size (12.00 × 4.68 µm, as measured in this study) and ornamentation. However, the basidiomata of R. hepatica are cauliflower-like in appearance, initially pale yellow when young, and turning reddish brown as they mature.

Fig. 59. Microscopic features of R. verrucula (MHHNU 31735). a basidiospores; b basidia. Scale bars: a =10 µm, b =20 µm.

Ramaria ser. Flavescentoides P. Zhang & J. Zhong, ser. nov.

MycoBank: MB 852775

Etymology: Species features are very similar to R. flavescens.

Type species: R. flavescentoides Hanif & Khalid, Mycotaxon 134(2): 402 (2019).

Diagnosis: Basidiomata are pale yellow to pale orange in color; apices are broadly rounded and concolorous with branches; basidiospores are long and conspicuously roughened in profile; hyphae and basidia base have clamps.

Commentary: This series includes four species collected in China, along with additional species such as R. rubricarnata Marr & D.E. Stuntz, R. thiersii R.H. Petersen & Scates, R. flavescens (Schaeff.) R.H. Petersen.

30. Ramaria brunneipes R.H. Petersen & M. Zang, Acta bot. Yunn. 11(4): 374, 1989. Fig. 17 a and Fig. 60

Fig. 60. Basidiospore features of R. brunneipes (MHHNU 6623). Scale bar: a =10 µm.

Description: Basidiomata 12 cm high and 7 cm broad, medium sized to large, usually single to fasciculate habit. Stipe 3 × 4 cm, single to compound, fleshy, large to robust, tapering gradually downward, rounded at base; off-white at base, off-white to butter yellow upward; not changing color or slightly changing color to brownish yellow to brown on bruising; surface smooth; occasionally with abortive branches; context of stipe off-white, freshy, drying light in weight, easily penetrated, but not friable. Branches in 3−6 ranks, major branches and branches above commonly polychotomous; inner branches and young branches pale yellow to pale orange when young, cream to butter yellow at age; axils narrowly rounded, mostly V-shaped; internodes all short, lower internodes up to 1.5 cm; branches slender and crowded, commonly 2–5 mm, seldom over 1 cm diam, terete, subparallel; context of main branches and upper branches freshy-gelatinous, off-white. Apices cuspidate, short and crowded, 1–3 mm long, polychotomous, concolorous with branches. Odor and taste faintly fabaceous.

Basidiospores [60/2/2] (9.2–)9.6–12.7(–13.0) × (3.3–)3.9–6.0 µm, L_m = 11.06 µm, l_m = 4.71 µm, Q = (1.67–)2.00–2.78(–3.34), Q_m = 2.38 ± 0.29, subcylindrical; ornamented with conspicuously warts and short meandering ridges randomly placed, strongly cyanophilous warts; hilar conspicuous, slightly curved. Basidia 50–65 × 9–13 μm, clavate, with basal clamps; sterigmata 4, 4–6 µm long, incurved or straight, slightly divergent. Hymenium unthickening, 55–70 µm. Subhymenium extensive, 15–30 µm thick, hyphae, interwoven, 3–6 µm diam, thin-walled, with clamps. Contextual hyphae: Tramal hyphae of stipe 3–15 μm wide, hyaline, loosely interwoven, inflating slightly, terminal swollen hyphae wide to 19 µm, with moderately ornamented; tramal hyphae of branches central hyphae slightly lysing, terminal swollen hyphae wide to 18 µm, with smooth and thin wall; ampulliform hyphae rarely; with clamps; gloeoplerous hyphae common, 7−10 μm diam.

Ecology: Found in broad-leaved, coniferous, or mixed forests. Basidiomata occurs in summer and autumn.

Distribution: Known from southwestern (Yunnan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Lijiang, bought from mushroom market, 9 September 1986, Petersen 47303 (isotype, HKAS 20048); Chuxiong, Zixi Mountain, in a forest with Lithocarpus and Pinus yunnanensis, altitude 2400 m, 3 August 2001, P. Zhang 123 (MHHNU 6623).

Commentary: Several yellowish basidiomata species share similarities with R. brunneipes, including R. flavescentoides, R. flavescens and its variants, R. formosa and its variants, R. pseudoflava, and R. barenthalensis. However, these species differ in various aspects such as basidiospores size, color, and apices morphology. A distinguishing feature for R. brunneipes is their stout stipes and extremely slender, crowded and subparallel branches. Notably, most of the other yellow species have primarily been reported in North America and Europe, whereas R. brunneipes is confirmed through our two-gene molecular phylogenetic analysis to be distinct from other yellow Ramaria species. The species most closely related to R. obtusa is R. flavescens. These two species share similar coloration, basidiospores sizes, and exhibit comparable features of a stout stipe and crowded branches. However, a key distinction lies in the branching pattern: R. flavescens has shorter internodes, and the branches are more dispersed, whereas R. brunneipes is has longer internodes and branches that are sub-parallel.

31. Ramaria compressipes P. Zhang & J. Zhong, sp. nov. Fig. 17 b and Fig. 61

MycoBank: MB 853293

Etymology: “compressipes” denotes the characteristic of the species having flatted stipes.

Diagnosis: Basidiomata are medium sized to large, cauliflower-like, cream to salmon colored; branches and apices are crowded, with apices concolorous with branches or turning brownish in maturity; slightly turn brownish when injured; basidiospores are long and narrowly, roughened in profile; hyphae and basidia base have clamps.

Type: CHINA, YUNNAN PROVINCE: Lijiang City, Yulong County, Baisha Township, Lijiang Alpine Botanical Garden, altitude 3569 m, 29 September 2019, Jian W. Liu 1809 (Holotype, MHHNU 11644, GenBank Acc. No.: ITS = PP467208).

Description: Basidiomata 8 cm high and 9 cm broad, single to fasciculate habit, medium sized, cauliflower-like in outline. Stipe 5 × 4 cm, single to compound, fleshy, large to robust, tapering gradually downward to narrowly rounded at base; off-white (1A1) at base, cream (4A3) to butter yellow (4A5) upward; slightly changing color to brownish yellow (5B3) on bruising; surface smooth; without abortive branches; context of stipe off-white (1A1), fibrous to spongy, drying light in weight, easily penetrated. Branches in 3−5 ranks, major branches and branches above commonly polychotomous; young branches pale yellow (2A43) to butter yellow (4A5), cream (4A3) to grayish orange (6B5) at age; axils narrowly rounded, mostly V-shaped; internodes all short, lower internodes up to 1 cm; branches slender and crowded, commonly 1–3 mm, terete, ascending; context of main branches and upper branches fleshy to gelatinous, pale yellow (2A3). Apices short and crowed, digitate to molar-like when young, coarsely digitate by maturity, 1–2 mm long, polychotomous, axils near apices 2–8 times. Odor and taste faintly fabaceous.

Basidiospores [33/1/1] (9.6–)10.2–13.0(–13.4) × (3.3–)3.8–4.9(–5.4) µm, L_m = 11.34 µm, l_m =4.25 µm, Q = (2.26–)2.39–3.09(–3.19), Q_m = 2.68 ± 0.23, subcylindrical to narrowly ellipsoid; ornamented with conspicuously warts and short meandering ridges randomly placed, strongly cyanophilous warts; hilar conspicuous, slightly curved. Basidia 50–65 × 8–12 μm, clavate, with basal clamps; sterigmata 4, 3–6 µm long, incurved or straight, slightly divergent. Hymenium unthickening, 55–70 µm. Subhymenium extensive, 15–30 µm thick, hyphae, interwoven, 3–6 µm diam, thin-walled, with clamps. Contextual hyphae: Tramal hyphae of stipe 3–15 μm wide, hyaline, loosely interwoven, inflating slightly, terminal swollen hyphae wide to 19 µm, with moderately ornamented; tramal hyphae of branches central hyphae slightly lysing, terminal swollen hyphae wide to 18 µm, with smooth and thin wall; ampulliform hyphae rarely; with clamps; gloeoplerous hyphae common, 7−10 μm diam.

Fig. 61. Microscopic features of R. compressipes (MHHNU 11644). a basidiospores; b basidia. Scale bars: a = 10 µm, b = 20 µm.

Ecology: Found in broad-leaved forests. Basidiomata occurs in summer and autumn.

Distribution: Known from southwestern (Yunnan).

Additional material examined: CHINA, YUNNAN PROVINCE: Deqin County, Feilaisi Forest Park, altitude 3400 m, 19 August 2018, P. Zhang 3417 (MHHNU 9917).

Commentary: Ramaria compressipes shares similarities with the R. thiersii R.H. Petersen & Scates in North American. Both exhibit cauliflower-like structures in basidiomata, obpyramidal and stout stipe, similar yellow to salmon-colored basidiomata, and are exclusively found in high-altitude regions (above 3000 meters) on gritty soil under the humus layer. The distinguishing feature lies in the smaller basidiospores of R. compressipes.

Furthermore, Ramaria compressipes exhibits similarities with R. rubricarnata Marr & D.E. Stuntz and its variants in the North American, displaying a similar botrytis-like morphology. The main differences lie in the latter's orange coloration with slightly smaller basidiospores. Notably, the hymenium of R. rubricarnata young specimens are beautiful rich maize yellow colored, and context is salmon colored, as opposed to the light orange to pinkish color in the lower branch’s context.

Additionally, R. compressipes shares morphological similarities with R. brunneipes. However, R. compressipes is distinguished by its short, digitate to molar-like apices, whereas R. brunneipes has slender apices.

32. Ramaria flavescentoides Hanif & Khalid, Mycotaxon 134(2): 402 (2019). Fig. 17 c−d and Fig. 62

Description: Basidiomata 6–9 cm high and 3–5 cm broad, repeatedly branched, usually obovate to obconic in outline. Stipe 1–2 × 1–3 cm, compound to fasciculate in groups of 2–5, tapering gradually downward into a basal mycelial mat or white tomentum covering the stipe base; off-white to pale orange at base, pale yellow to pale orange upward; not changing color on bruising; surface smooth; without abortive branches; context of stipe off-white, solid, dried flesh firm but easily penetrated, gelatinous or slippery. Branches in 4−7 ranks, major branches 4–6, 1.5–2 cm diam, stout, more or less terete, ascending, pale yellow to pale orange below, upward paler; axils narrowly rounded, mostly U-shaped; internodes diminishing gradually at maturity, lower internodes up to 3 cm; branches above commonly polychotomous, broader upward, commonly 5–20 mm diam, terete or flattened, crowded, ascending, slightly divergent; context of main branches and upper branches gelatinous, off-white to pale orange. Apices broadly rounded, short, 2–4 mm long, polychotomous, concolorous with branches or sometimes slightly changing color to brownish yellow on bruising. Odor and taste faintly fabaceous.

Basidiospores [60/2/2] (10.2–)11.1–14.9(–15.6) × (4.0–)4.5–6.5(–7.0) µm, L_m = 12.40 µm, l_m = 5.27 µm, Q = (1.82–)2.01–2.88(–3.27), Q_m = 2.38 ± 0.26, oblong to ellipsoid; ornamented with low coarse warts and delicate meandering ridges covering significant areas of wall, strongly cyanophilous warts; hilar conspicuous, slightly curved at one end. Basidia 55–65 × 9–13 μm, clavate, with basal clamps; sterigmata 2–4, mostly 4, 5–8 µm long, incurved or straight, slightly divergent. Hymenium unthickening, 75–80 µm. Subhymenium rudimentary, 15–30 µm thick, hyphae, interwoven, 3–6 µm diam, thick-walled, with clamps. Contextual hyphae: Tramal hyphae of stipe 6–12 μm wide, hyaline, parallel, inflating slightly, terminal swollen hyphae wide to 15 µm, with moderately ornamented; tramal hyphae of branches central hyphae slightly lysing, terminal swollen hyphae wide to 15 µm, with smooth and thin wall; ampulliform hyphae occasional; with frequently clamps; gloeoplerous frequently, 7−10 μm diam.

Ecology: Found in a mixed forest with Quercus and Picea. Basidiomata occurs in summer and autumn.

Distribution: Known from northeast China and southwest China (Xizang), Pakistan.

Additional materials examined: CHINA, XIZANG AUTONOMOUS REGION: Linzhi City, Milin County, Zhaxiraodeng Village, in Picea forest, altitude 3040 m, 24 August 2018, P. Zhang 3483 (MHHNU 9983), P. Zhang 3489 (MHHNU 9989); same city, 25 August 2018, P. Zhang 3496 (MHHNU 9996).

Fig. 62. Basidiospore features of R. flavescentoides (MHHNU 9996). Scale bar: a = 10 µm.

Commentary: The main features of R. flavescentoides include: 1) basidiomata medium sized, obovate to obconic basidiomata, pale yellow to pale orange colored; 2) apices broadly rounded, concolorous with branches or slightly changing color to brown when matured; 3) basidiospores long, slightly curved, conspicuously roughened in profile; 4) hyphae and basidia base with clamps.

R. flavescentoides resembles R. flavescens (Schaeff.) R.H. Petersen in shape, branching pattern, and basidiomata color, also sharing a similar basidiospores shape and ornamentation and the presence of basidial clamps. However, R. flavescens has smaller basidiomata and smaller basidiospores. Hanif & Khalid noted that it’s have stellate crystalline material in the hyphae of upper branches, which similar crystals have been reported by Zhang & al (Zhang et al. 2005) in the stipe and upper branches of R. luteoaeruginea P. Zhang & Zhu L. Yang in subgen. Echinoramaria (Phaeoclavulina).

R. flavescentoides also resembles R. rasilisporoides, however, R. rasilisporoides differs from R. flavescentoides in its pale yellow basidiomata, amyloid stipe tissues, deeply rooting stipe with numerous abortive branchlets, and smaller basidiospores (Exeter et al. 2006).

R. flavescentoides and R. mairei Donk were the only two species of old subgen. Laeticolora previously reported from Pakistan (Hanif et al. 2019), the latter is distinguished by larger (18 cm) basidiomata, 8 cm high stipe, numerous pallid or deep lemon to pale ochraceous and rather lax or crowded branches, and slightly longer (10–15 μm) oblong to ellipsoid roughened pale ochraceous basidiospores.

33. Ramaria obtusa L. Fan, Ying Li & N. Mao, in Li, Mao, Zhang, Fu & Fan, Mycology: 10.1080/21501203.2024.2388910, 23 (2024) Fig. 17 e−f and Fig. 63

Description: Basidiomata 12 cm high and 7 cm broad, medium sized to large, usually single to fasciculate habit. Stipe 3 × 4 cm, single to compound, fleshy, large to robust, tapering gradually downward, rounded at base; off-white at base, off-white to butter yellow upward; not changing color or slightly changing color to brownish yellow to brown on bruising; surface smooth; occasionally with abortive branches; context of stipe off-white, freshy, drying light in weight, easily penetrated, but not friable. Branches in 3−6 ranks, major branches and branches above commonly polychotomous; inner branches and young branches pale yellow to pale orange when young, cream to butter yellow at age; axils narrowly rounded, mostly V-shaped; internodes all short, lower internodes up to 1.5 cm; branches slender and crowded, commonly 2–5 mm, seldom over 1 cm diam, terete, subparallel; context of main branches and upper branches freshy-gelatinous, off-white. Apices cuspidate, short and crowded, 1–3 mm long, polychotomous, concolorous with branches. Odor and taste faintly fabaceous.

Basidiospores [60/2/2] (9.2–)9.6–12.7(–13.0) × (3.3–)3.9–6.0 µm, L_m = 11.06 µm, l_m = 4071 µm, Q = (1.67–)2.00–2.78(–3.34), Q_m = 2.38 ± 0.29, subcylindrical; ornamented with conspicuously warts and short meandering ridges randomly placed, strongly cyanophilous warts; hilar conspicuous, slightly curved. Basidia 50–65 × 9–13 μm, clavate, with basal clamps; sterigmata 4, 4–6 µm long, incurved or straight, slightly divergent. Hymenium unthickening, 55–70 µm. Subhymenium extensive, 15–30 µm thick, hyphae, interwoven, 3–6 µm diam, thin-walled, with clamps. Contextual hyphae: Tramal hyphae of stipe 3–15 μm wide, hyaline, loosely interwoven, inflating slightly, terminal swollen hyphae wide to 19 µm, with moderately ornamented; tramal hyphae of branches central hyphae slightly lysing, terminal swollen hyphae wide to 18 µm, with smooth and thin wall; ampulliform hyphae rarely; with clamps; gloeoplerous hyphae common, 7−10 μm diam.

Ecology: Found in broad-leaved, coniferous, or mixed forests. Basidiomata occurs in summer and autumn.

Distribution: Known from central, north and northeastern of China (Hunan, Jilin, Shanxi).

Additional materials examined: CHINA, HUNAN PROVINCE: Changde City, Shimen County, Hupingshan National Nature Reserve, Huier Bay, in broadleaf forest, altitude 1600 m, 10 September 2012, P. Zhang 1264 (MHHNU 7764). JILIN PROVINCE: Tonghua City, Baijifeng Forest Park, in mixed forests dominated by Acer, Quercus and Pinus, altitude 800 m, 28 Aug. 2020, P. Zhang 4043 (MHHNU 10553); 28 Aug. 2020, P. Zhang 4053 (MHHNU 10091).

Commentary: Several yellowish basidiomata species share similarities with R. obtusa, including R. flavescentoides, R. flavescens and its variants, R. formosa and its variants, R. pseudoflava, and R. barenthalensis. However, these species differ in various aspects such as basidiospores size, color, and apices morphology. A distinguishing feature for R. obtusa is its stout stipes and extremely slender, crowded and subparallel branches. Notably, most of the other yellow species have primarily been reported in North America and Europe, whereas R. obtusa is confirmed through our two-gene molecular phylogenetic analysis to be distinct from other yellow Ramaria species. The species most closely related to R. obtusa is R. flavescens. These two species share similar coloration, basidiospores sizes, and exhibit comparable features of a stout stipe and crowded branches. However, a key distinction lies in the branching pattern: R. flavescens has shorter internodes, and the branches are more dispersed, whereas R. obtusa has longer internodes and branches that are subparallel.

Ramaria ser. Formosae Corner, Beih. Nova Hedwigia 33: 240 (1970)

Synonym: Ramaria sect. Formosae (Corner) Franchi & M. Marchetti.

Type species: R. formosa (Pers.) Quél., Fl. mycol. France (Paris): 466 (1888)

Diagnosis: Basidiomata are pale yellow to dark yellow in tone; branches are subparallel, with off-white apices; basidia base and hyphae have clamps; basidiospores are conspicuously rough.

Commentary: This series includes three species collected in China, but the type species R. formosa is not founded in China. This series is characterized by white apices and basidiomata ranging from yellow to ocher-yellow color.

Fig. 63. Basidiospore features of R. obtuse (MHHNU 10553). Scale bar: a = 10 µm.

34. Ramaria formosoides L. Fan, Ying Li & N. Mao, in Li, Mao, Zhang, Fu & Fan, Mycology: 10.1080/21501203.2024.2388910, 20 (2024) Fig. 18 a–b and Fig. 64

Description: Basidiomata 15 cm high and 9 cm broad. Stipe 1–3.2 × 5.5 cm, off-white at base, cream to salmon; not changing color or slightly changing color to brownish yellow to yellow ochre on bruising; surface tomentose, gnarled or rugose; without abortive branches; context of stipe off-white to cream, freshy-fibrous, drying chalky-friable, brittle. Major branches several, cream to salmon. Branches in 3−6 ranks, butter yellow to salmon when young, apricot yellow at age; axils broadly rounded, mostly U-shaped; internodes diminishing gradually at maturity. Apices acute to rounded, crowded, polychotomous, off-white when young, darker at age. Odor and taste faintly fabaceous.

Basidiospores [30/2/1] 9.0–12.0(–13.0) × 4.5–5.2 (–5.5) µm, L_m = 10.64 µm, l_m = 4.84 µm, Q = (1.8–)1.9–2.5(–2.6), Q_m = 2.20 ± 0.19, subcylindrical; ornamented with conspicuously warts, cyanophilous warts. Basidia 45–65 × 8–12 μm, clavate, with basal clamps; sterigmata mostly 4, 3–7 µm long, incurved or straight, slightly divergent. Subhymenium rudimentary, hyphae, interwoven, thin walled, with clamps. Hymenium unthickening. Contextual hyphae: Tramal hyphae of stipe 4–11 μm wide, hyaline, loosely interwoven, inflating slightly; tramal hyphae of branches central hyphae slightly lysing; ampulliform hyphae rarely; with clamps; gloeoplerous hyphae occasionally.

Ecology: Found in coniferous and broad-leaved mixed forests. Basidiomata occurs in summer and autumn.

Distribution: Known from southwest, north and northeast of China (Yunnan, Sichuan, Jilin, Shanxi).

Additional materials examined: CHINA, YUNNAN PROVINCE: Shangri-La City, Haba Snow Mountain, Mianshaba, altitude 3100 m, 12 August 2008, Li P. Tang 588 (HKAS 54819); same city, Deqin County, Baima Snow Mountain Nature Reserve, altitude 3750 m, 18 August 2018, P. Zhang 3467(MHHNU 9967). SICHUAN PROVINCE: Ganzi, Kangding City, Gongga Mountain, in Quercus semecarpifolia and Picea asperata forest, altitude 3900 m, 5 September 2021, P. Zhang 4259 (MHHNU 107659), P. Zhang 4260 (MHHNU 10760), P. Zhang 4276 (MHHNU 10776), P. Zhang 4277 (MHHNU 107677); same location, 6 September 2021, P. Zhang 4287 (MHHNU 107687), P. Zhang 4291(MHHNU 10791), P. Zhang 4300 (MHHNU 10300), P. Zhang 4305 (MHHNU 107305); same location, 7 September 2021, P. Zhang 4367 (MHHNU 107867); same location, 8 September 2021, P. Zhang 4379 (MHHNU 10879). JILIN PROVINCE: Tonghua County, Baijifeng National Forest Park, in broadleaf forest, altitude 800 m, 6 August 2022, P. Zhang 4701 (MHHNU 11201).

Commentary: Dr. Zhang identified this species as R. obtusissima (Peck) Corner (Zhang 2010). In Corner's 1950 description (Corner 1950), R. obtusissima was considered a complex group, including five different types: rough spored form, pink form, vinescent form, rich yellow, and typical (Corner 1950). In 1967, Petersen, redescribed the type specimen and restricted R. obtusissim. However, in 1992, Schild reexamining the type specimen, found that the basidiospores of R. obtusissima were not smooth as previously described, but had significant warty ornamentation (Schild 1992b). He assigned those fungi with smooth basidiospores, previously considered R. obtusissima, to a new species named R. schildii. Compared to the fine rough basidiospores of the type species R. obtusissima, the ornamentation of this species is significantly rougher. Moreover, this species has a wider distribution in China, although some basidiomata have rounded apices similar to R. obtusissima, basidiomata with acute apices are more common.

The white to creamy apices of R. formosoides are compared to R. formosa, with the latter having distinctly yellow apices. Compared to the R. pallidicarnea, it has slightly larger basidiospores (9.85 × 4.06 µm, as measured in this study). Compared to R. longicarnea, the basidiospores are slightly smaller (11.45 × 6.10 µm, as measured in this study).

Fig. 64. Basidiospore features of R. formosoides (MHHNU 10760). Scale bar: a = 10 µm.

35. Ramaria longicarnea P. Zhang & J. Zhong, sp. nov. Fig. 18 c−d and Fig. 65

MycoBank: MB 853141

Etymology: “longicarnea” = “long-” + “carnea”, proposed because this species has long stipes and pink to salmon color of basidiomata.

Diagnosis: Basidiomata are salmon to fleshy in color; basidia and hyphae have clamps; stipe is long with white rugulose hyphae; basidiospores are broad, strongly cyanophilous, and rough in outline.

Type: CHINA, YUNNAN PROVINCE: Dali, Binchuan County, Jizu Mountain, with broad-leaved, coniferous, or mixed forests, altitude 2300 m, 29 August 2022, P. Zhang 4829 (Holotype, MHHNU 11329, GenBank Acc. No.: ITS = PP467291, nrLSU = PP493593, tef1-α = PP480451).

Description: Basidiomata 13 × 5.5 cm. Stipe 7 × 1.5 cm, single, off-white (1A1) at base, salmon (6A4) upward, tapering gradually downward, with abortive branchlets; flesh solid, off-white (1A1), moist, but not gelatinous or slippery. Major branches 3–5, stout, crowded, small branching angle, nearly terete. Branches in 3−6 ranks, ascending, small branching angle, crowded, cream (4A3) to salmon (6A4), fleshy, drying soft, easily penetrated; internodes length ratio diminishing gradually at maturity; axils mostly acute, V-shaped. Apices crowded and acute, cuspidate, off-white (1A1) when young, finally concolorous with branches. Odor and taste faintly fabaceous.

Basidiospores [64/2/2] (9.2–)9.7–13.2(–15.6) × (4.7–)5.1–7.3(–7.9) µm, L_m = 11.45 µm, l_m = 6.10 µm, Q = 1.59–2.32, Q_m = 1.92 ± 0.16, ellipsoid, roughened in profile; wall up to 0.3 μm thick, strongly cyanophilous; hilar appendix prominent; ornamentation of prominent, discrete, low warts and short ridges randomly placed. Basidia 38-45 × 5-9 μm, clavate, clamped at base, strong cyanophilous, 4-spored; sterigmata 4−6 μm long. Subhymenium rudimentary, hyphal. Hymenium unthickening. Tramal hyphae of stipe 4−10 μm wide, hyaline, thin-walled to thick-walled, loosely parallel, clamped, interwoven; ampulliform clamps up to 13 μm broad, asymmetrical, thick-walled, with coarse ornamentation; gloeoplerous hyphae not observed. Tramal hyphae of upper branches 3−10 μm wide, hyaline, clamped, thin-walled to thick-walled, loosely parallel; ampulliform clamps up to 13 μm broad, asymmetrical, thick-walled, with coarse ornamentation; gloeoplerous hyphae not observed.

Ecology: Solitary on soil in mixed forests dominated by Fagaceae and Pinaceae.

Distribution: known from southwestern China (Yunnan).

Additional materials examined: CHINA YUNNAN PROVINCE: Chuxiong, Mouding County, Huafo Mountain, in a broad-leaved forest, altitude 2350 m, 21 August 2022, P. Zhang 4731 (MHHNU 11231), P. Zhang 4749 (MHHNU 11249); same location, Mouding County, Huafo Mountain, 27 July 2021, Z. H. Chen 32918 (MHHNU 32918), Z. H. Chen 32921 (MHHNU 32921); Kunming, bought from mushroom market, 23 July 2017, P. Zhang 2600 (MHHNU 9100).

Commentary: Compared with R. pallidicarnea (basidiospores 9.85 × 4.06 µm, as measured in this study) and R. formosoides (10.64 × 4.84 µm, as measured in this study), it has slightly larger basidiospores and a longer stipes. Ramaria atkinsonii (Petersen 1984) and R. formosa resembles R. longicarnea in appearance, they share salmon to yellow basidiomata and clamped basiama. Longer stipe can separate R. formosa and R. longicarnea, and the later have off-white apice when young. Ramaria atkinsonii and R. longicarneahave the same sized basidiospores, however, the former has near smooth to finely rough spore.

Fig. 65. Microscopic features of R. longicarnea (MHHNU 11329). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

36. Ramaria pallidicarnea P. Zhang & Xue L. Gao, sp. nov. Fig. 18 e−f and Fig. 66

MycoBank: MB 853138

Etymology: pallidicarnea = pallida+ carnea, referring to its off-white apices and salmon to pink basidiomata.

Diagnosis: Apices are off-white to cream; basidiomata pink to salmon; hyphae and basidia base have clamps; drying results in a brittle and chalky-friable texture; does not change color or slightly change color to brownish yellow.

Type: CHINA, YUNNAN PROVINCE: Lijiang City, Shuhe Ancient Town, in a mixed coniferous and broad-leaved forest, altitude of 3172 m, 22 August 2020, Li P. Tang HT469 (Holotype, MHKMU HT469, GenBank Acc. No.: ITS = PP467293).

Description: Basidiomata 12 cm high and 7 cm broad. Stipe 2 × 2 cm, lobed or muscled; off-white (1A1) at base, cream (4A3) to salmon (6A4); not changing color or slightly changing color to brownish yellow (5B3) to yellow ochre (5C7) on bruising; surface tomentose, gnarled or rugose; without abortive branches; context of stipe off-white (1A1) to cream (4A3), freshy-fibrous, drying chalky-friable, brittle. Major branches several, cream (4A3) to salmon (6A4). Branches in 3−6 ranks, butter yellow (4A5) to salmon (6A4) when young, apricot yellow (5B6) at age; axils broadly rounded, mostly U-shaped; internodes diminishing gradually at maturity. Apices acute to rounded, crowded, polychotomous, off-white (1A1) when young, darker at age. Odor and taste faintly fabaceous.

Basidiospores [30/2/1] (8.3–)8.7–11.0(–11.7) × (3.1-)3.4–4.9(5.0) µm, L_m = 9.85 µm, l_m = 4.06 µm, Q = 2.10–2.87(–3.00), Q_m = 2.45 ± 0.26, subcylindrical; ornamented with conspicuously warts, cyanophilous warts. Basidia 45–65 × 7–12 μm, clavate, with basal clamps; sterigmata mostly 4, 3–7 µm long, incurved or straight, slightly divergent. Subhymenium rudimentary, hyphae, interwoven, thin walled, with clamps. Hymenium unthickening. Contextual hyphae: Tramal hyphae of stipe 4–11 μm wide, hyaline, loosely interwoven, inflating slightly; tramal hyphae of branches central hyphae slightly lysing; ampulliform hyphae rarely; with clamps; gloeoplerous hyphae occasionally.

Ecology: Found in coniferous and broad-leaved mixed forests. Basidiomata occurs in summer and autumn.

Distribution: Known from southwest China (Yunnan).

Additional material examined: CHINA, YUNNAN PROVINCE: Chuxiong, Lufeng City, Tuoan Township, 6 September 2022, Xue L. Gao 20586 (MHHNU 11635).

Commentary: The isotype specimen HKAS 17744, which Petersen (1989) classified as R. laeviformosoides, cannot be attributed to that species because it exhibits distinctly warty basidiospores, whereas R. laeviformosoides is characterized by nearly smooth basidiospores. Consequently, the original isotype HKAS 17744 is described here as a new species, R. pallidicarnea. Ramaria pallidicarnea is distinguished from other species in ser. formosae by its average basidiospores length being less than 10 µm.

Ramaria ser. Gelatinosae P. Zhang & J. Zhong, ser. nov.

MycoBank: MB 852719

Etymology: “gelatinose” means the stipe somewhat gelatinous.

Type species: R. indoyunnaniana R.H. Petersen & M. Zang, Acta bot. Yunn. 8(3): 287, 1986.

Diagnosis: Basidiomata are pale yellow to salmon, with apices yellow to pink; stipe is somewhat gelatinous or slippery; branches are slender; most species lack clamps.

Commentary: This series includes fifteen species collected in China, including R. pallidosaponaria R.H. Petersen 1989, R. flavigelatinosa Marr & D.E. Stuntz, R. flavoides Schild and its varities R. flavoides var. betonensis Franchi & M. Marchetti (= R. abetonensis Franchi & M. Marchetti), R. verlotensis Marr & D.E. Stuntz, R. ossolana Franchi & M. Marchetti, R. primulina R.H. Petersen, R. gelatiniaurantia Marr & D.E. Stuntz and its varities R. gelatiniaurantia var violeitingens Marr & D.E. Stuntz, R. gelatiniaurantia var gelatiniaurantia Marr & D.E. Stuntz.

Fig. 66. Microscopic features of R. pallidicarnea (MHHNU 11635). a basidiospores; b basidia. Scale bars: a =10 µm, b =20 µm.

37. Ramaria apicaliochracea L. Fan, Ying Li & N. Mao, in Li, Mao, Zhang, Fu & Fan, Mycology: 10.1080/21501203.2024.2388910, 14 (2024)

Original description in Li et al. (2024).

38. Ramaria brevigelatinosa P. Zhang & J. Zhong, sp. nov. Fig. 19 a–b and Fig. 67

MycoBank: MB 853036

Etymology: brevigelatinosa = brevi- + gelatinosa, where "brevi-" refers to the stipe being short, and “gelatinosa” means the species has a gelatinous context.

Diagnosis: Basidiomata are small to medium sized and pale yellow; stipe is short, stout, gelatinous, and smooth; hyphae and basidia base lack clamps; average basidiospores length is smaller than 10 µm.

Type: CHINA, YUNNAN PROVINCE: Chuxiong, Zixi Mountain, in mixed forests dominated by Fagaceae and Pinaceae, altitude 2100 meters, 24 August 2022, P. Zhang 4783 (Holotype, MHHNU 11283, GenBank Acc. No.: ITS = PP467184, nrLSU = PP493505).

Description: Basidiomata 3–6 × 1.5–3.5 cm, fasciculate habit, usually obpyriform to broadly fusiform in outline. Stipes up to 0.5 × 1.5 cm, single to compound, tapering gradually downward; milk white (1A2) at base, milk white (1A2) to pale yellow (2A3) upward; surface smooth; with abortive branches; some region changing color to brownish gray (6E2) on bruising or purple spots caused by soil particles; context of stipe off-white (1A1) to jelly-like pastel yellow (2A4) when fresh, drying light in weight, easily penetrated, gelatinous, hard. Major branches several, ascending, terete. Branches in 3−5 ranks, major branches and lower branches commonly dichotomous; branches pale yellow (2A3) to chicken yellow (2A6); axils broadly rounded, mostly U-shaped; internodes all short; branches terete, sparsely, subparallel upward; context of main branches and upper branches gelatinous, viscid, brittle, pastel yellow (2A4). Apices broadly rounded, molar-like when young, terraced, short-digitate at maturity, concolorous with branches or slightly pale yellow (2A3) to chicken yellow (2A6). Odor weakly fragrant, aromatic; taste negligible to faintly astringent.

Basidiospores [50/2/2] (8.5–)9.0–11.0(–11.5) × 4.5–5.8(–6.0) µm, L_m = 9.92 µm, l_m = 5.21µm, Q = (–1.61)1.77–2.29(–2.33), Q_m = 2.01 ± 0.15, ellipsoid, noticeably rough; ornamented with warts and short meandering ridges randomly placed, strongly cyanophilous warts; hilar conspicuous, slightly curved. Basidia 45–65 × 6–8 μm, clavate, without basal clamps; sterigmata 4, 3–6 µm long, short and straight, somewhat scattered. Hymenium unthickening, 50–70 µm. Subhymenium extensive, hyphae, interwoven, 1–3 µm diam, adherent, without clamps. Contextual hyphae: Tramal hyphae of stipe 3–10 μm wide, hyaline, tightly interwoven, heavily agglutinated, ampulliform inflations occasionally; tramal hyphae of branches central hyphae slightly narrow, 2–10 µm, hyaline, thin-walled, agglutinated, strictly parallel, tightly packed, ampulliform inflations rare; gloeoplerous hyphae commonly, 4−8 μm diam.

Ecology: Solitary or gregarious on soil in mixed forests dominated by Fagaceae and Pinaceae. Basidiomata occurs in summer and autumn.

Distribution: Known from southwest China (Yunnan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Chuxiong, Huafo Mountain, in the mixed forests, altitude 2200 m, 23 August 2022, P. Zhang 4769 (MHHNU 11269); same locaiton, altitude 2300 m, 21 August 2022, P. Zhang 4748 (MHHNU 11248); Chuxiong, Shuangbaisi, 22 August 2018, Zuo H. Chen 31306 (MHHNU 31306); Kunming, Panlong District, A Ziying, 2 August 2001, P. Zhang 128 (MHHNU 6628); Wenshan, Malipo County, Xinzhai Township, in broad-leaved forest, altitude 1566 m, 27 July 2017, G. Wu 2207 (HKAS123447), G. Wu 2208 (HKAS123448).

Commentary: Species with a closer affinity to R. brevigelatinosa include R. thindii, and R. hunanensis. These species share common features such as yellow basidiomata, gelatinous context, and absence of clamps. Molecular phylogenetic analysis confirms that they are not the same species. Microscopically, only R. brevigelatinosa and R. hunanensis (basidiospore 11.51 × 5.70 µm, as measured in this study) exhibit a slightly smaller length-to-width ratio of basidiospores compared to the others, with the former having even smaller basidiospores dimensions. Additionally, R. brevigelatinosa is characterized by shorter and stouter stipes, and relatively fat branches.

Fig. 67. Microscopic features of R. brevigelatinosa (MHHNU 11283). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

39. Ramaria hunanensis P. Zhang & J. Zhong, sp. nov. Fig. 19 c−d and Fig. 68

MycoBank: MB 853374

Etymology: hunanensis, indicating that the species is collected in the Hunan Province of China and is characterized by its yellow basidiomata and gelatinous flesh features.

Diagnosis: Basidiomata are overall light-yellow in color; stipe is slender, gelatinous, and smooth; changes color to fawn brown on bruising; hyphae and basidia base lack clamps; basidia are elongated, and basidiospores relatively large.

Type: CHINA, HUNAN PROVINCE: Chenzhou City, Yizhang County, Mangshan National Nature Reserve, in mixed forests dominated by Fagaceae and Pinaceae. altitude 1500 m, 2 September 2007, P. Zhang 640 (Holotype, MHHNU 7140, GenBank Acc. No.: ITS = PP467189, nrLSU = PP493509, tef1-α = PP480391).

Description: Basidiomata 2–8 × 1.5–7 cm, fasciculate habit. Stipes up to 2–4 × 1–2 cm, single to compound, slender, tapering gradually downward; milk white (1A2) to chicken yellow (2A6); changing color to fawn brown (7E4) on bruising; surface smooth; without abortive branches; context of stipe off-white (1A1) to jelly-like pastel yellow (2A4) when fresh, drying light in weight, easily penetrated, gelatinous, brittle. Major branches several, ascending, terete. Branches in 2−5 ranks, major branches and lower branches commonly dichotomous; branches milk white (1A2) to chicken yellow (2A6); axils broadly rounded, mostly U-shaped; internodes long, diminishing gradually at maturity; branches terete, strict, sparsely, subparallel upward; context of main branches and upper branches gelatinous, viscid, brittle, pale yellow (2A3). Apices broadly rounded, 1–3 mm long, fragile, dichotomous, milk white (1A2) to pastel yellow (2A4). Odor weakly fragrant, aromatic; taste negligible to faintly astringent.

Basidiospores [60/2/2] 10.0–13.0(–14.0) × 4.0–7.0 µm, L_m = 11.51 µm, l_m = 5.70 µm, Q = (1.67–)1.71–2.75, Q_m = 2.05 ± 0.26, subcylindrical to long ellipsoid, noticeably rough; ornamented with moderate warts and short meandering ridges randomly placed, strongly cyanophilous warts; hilar conspicuous, slightly curved. Basidia 53–80 × 8–13 μm, clavate, without basal clamps; sterigmata 4, 6–9 µm long, long and straight, somewhat scattered. Hymenium unthickening, 60–90 µm. Subhymenium extensive, hyphae, interwoven, 1.5–3 µm diam, adherent, without clamps. Contextual hyphae: Tramal hyphae of stipe 2–14 μm wide, hyaline, tightly interwoven, heavily agglutinated, ampulliform inflations up to 17 µm; tramal hyphae of branches central hyphae slightly narrow, 2–12 µm, hyaline, thin-walled, agglutinated, strictly parallel, tightly packed, ampulliform inflations associated with septa, up to 17 µm broad, thick-walled (wall up to 1 µm thick), unornamented; gloeoplerous hyphae commonly, 4−8 μm diam.

Ecology: Solitary or gregarious on soil in mixed forests dominated by Fagaceae and Pinaceae. Basidiomata occurs in summer and autumn.

Distribution: Known from central China (Hunan).

Additional materials examined: CHINA, HUNAN PROVINCE: Chenzhou City, Yizhang County, Mangshan National Nature Reserve, in mixed forests dominated by Fagaceae and Pinaceae. 26 July 2018, P. Zhang 3227 (MHHNU 9727); same location, Guizizhai, in broad-leaved forest with Fagaceae, 14 September 2003, P. Zhang 375 (MHHNU 6875), P. Zhang 376 (MHHNU 6876), P. Zhang 377 (MHHNU 6877).

Commentary: Within the group of taxa with light yellow to yellow gelatinous features and no clamps, there are species such as R. verlotensis Marr & D. E. Stuntz, R. flavigelatinosa, and its variants. Ramaria verlotensis has cauliflower-like, pale yellowish pink basidiomata, and its basidiospores are slightly smaller (9.0–11.0 × 4.5–6.0 µm, L_m = 10.1 µm, l_m = 4.9 µm, Q_m = 2.06; Marr & Stuntz 1973). Ramaria flavigelatinosa has a white base of the fruiting body, branches ranging from light yellow to maize yellow, and apices concolorous with branches or slightly brighter, resembling sunflower yellow. However, its basidiospores are shorter than 10 µm (9.6 × 4.1 µm; Marr & Stuntz 1973). Ramaria flavigelatinosa var. megalospora, has basidiospores sizes similar to R. hunanensis, but its fruiting body tends towards melon yellow, slightly more orange yellow, unlike the yellowish-white color of R. hunanensis. Importantly, R. hunanensis shows purple-brown bruising where it contacts soil particles.

Fig. 68. Microscopic features of R. hunanensis (MHHNU 7140). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

40. Ramaria flavicoralloides L. Fan, Ying Li & N. Mao, in Li, Mao, Zhang, Fu & Fan, Mycology: 10.1080/21501203.2024.2388910, 19 (2024)

Original description in Li et al. (2024).

41. Ramaria gelatinifusca P. Zhang & J. Zhong, sp. nov. Fig. 19 e−f and Fig. 69

MycoBank: MB 853376

Etymology: gelatinifusca = gelatinosa + fusca, indicating that the species has a gelatinous stipe and changes color to brown when bruised.

Diagnosis: Basidiomata are salmon color; apices are broadly rounded, yellow to salmon; stipe is gelatinous, gnarled or rugose; hyphae and basidia base are nearly without clamps; basidiospores are broad.

Type: CHINA, YUNNAN PROVINCE: Deqin County, Baima Snow Mountain Nature Reserve, 17 August 2018, P. Zhang 3436 (Holotype, MHHNU 9936, GenBank Acc. No.: ITS = PP467193, nrLSU = PP493513).

Fig. 69. Microscopic features of R. gelatinifusca (MHHNU 9936). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Description: Basidiomata 9–15 × 8–12 cm, fasciculate habit. Stipes up to 4 × 5 cm, single to compound, tapering gradually downward; off-white (1A1) at base, light orange(5A3) to yellowish orange (4A8); changing color to brownish gray (6E2) on bruising; longitudinally gnarled or rugose; occasionally with abortive branches; context of stipe off-white (1A1) to jelly-like yellowish orange (4A8) when fresh, drying light in weight, easily penetrated, gelatinous, firm. Major branches several, ascending, hardly terete. Branches in 3−5 ranks, major branches and lower branches commonly polychotomous; branches salmon (6A4) to yellowish orange (4A8); axils narrowly rounded, mostly V-shaped; internodes long, diminishing gradually at maturity; branches terete, sparsely, subparallel; context of main branches and upper branches semi-gelatinous, viscid, brittle, yellowish orange (4A8). Apices broadly rounded, 1–3 mm long, fragile, polychotomous, pale orange (5A3) to chicken yellow (2A6). Odor weakly fragrant, aromatic; taste negligible to faintly astringent.

Basidiospores [30/2/1] (9.0–)10.5–13.7(–14.2) × (4.2–)4.8–6.6(–7.2) µm, L_m = 11.71 µm, l_m = 5.68 µm, Q = (1.73–)1.83–2.36(–2.43), Q_m = 2.07 ± 0.18, subcylindrical to ellipsoid, moderately rough; ornamented with moderate warts and short meandering ridges randomly placed, moderately cyanophilous warts; hilar conspicuous, slightly curved. Basidia 55–75 × 7–15 μm, clavate, without basal clamps; sterigmata 4, 3–6 µm long, slender and straight, somewhat scattered. Hymenium unthickening, 60–80 µm. Subhymenium extensive, hyphae, interwoven, 1.5–3 µm diam, adherent, without clamps. Contextual hyphae: Tramal hyphae of stipe 2–14 μm wide, hyaline, tightly interwoven, heavily agglutinated, ampulliform inflations up to 15 µm; tramal hyphae of branches central hyphae slightly narrow, 2–12 µm, hyaline, thin-walled, agglutinated, strictly parallel, tightly packed, ampulliform inflations associated with septa, up to 15 µm broad, thick-walled (wall up to 1 µm thick), unornamented; gloeoplerous hyphae commonly, 4−8 μm diam.

Ecology: In the mixed forests dominated by Picea. Basidiomata occurs in summer and autumn.

Distribution: Known from southwest China (Yunnan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Shangri-La City, Pudacuo National Park, Bitahai, in the Picea forest, altitude 3500 m, 31 August 2022, P. Zhang 4847 (MHHNU 11347); Deqin County, Baima Snow Mountain Nature Reserve, in the Picea forest, altitude 3750 m, 18 August 2018, P. Zhang 3468 (MHHNU 9968); same location, 17 August 2018, P. Zhang 3441 (MHHNU 9941).

Commentary: The species with salmon-colored basidiomata, gelatinous stipe, and lacking clamps are quite numerous. The most similar species is R. linearis, and the wider basidiospores (13.56 × 5.14 µm, as measured in this study) and bruising to grayish brown are the distinguishing features of R. gelatinifusca, making it easily distinguishable from other species. This species is collected in relatively humid high-altitude areas with Picea forests, and it is prone to mold, producing gray-brown spots. The abundant gelatinous substance imparts a jelly-like semi-translucent quality to the stipe flesh, and the branching tissue exhibits a red-orange color.

42. Ramaria gelatinosivivida P. Zhang & J. Zhong, sp. nov. Fig. 19 g−h and Fig. 70

MycoBank: MB 853377

Etymology: gelatinosivivida = gelatinosa + vivida, referring to the species having a vivid yellow color and gelatinous stipe and branches.

Diagnosis: Basidiomata are vivid yellow to salmon colored; apices are broadly rounded, concolorous with branches, and slightly brownish by maturity; stipe is gelatinous, gnarled or rugose; hyphae and basidia base are nearly without clamps; basidiospores are ornamented with moderate warts, long and narrow.

Type: CHINA, SICHUAN PROVINCE: Ganzi, Kangding City, Gongga Mountain, in Quercus semecarpifolia and Picea asperata forest, altitude 3500 m, 2 September 2018, P. Zhang 3589 (Holotype, MHHNU 10089, GenBank Acc. No.: ITS = PP467195, nrLSU = PP493515).

Fig. 70. Microscopic features of R. gelatinosivivida (MHHNU 10089). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Description: Basidiomata 8–10 × 15 cm, single to fasciculate habit. Stipes up to 3–5 × 3–6 cm, single to compound, stout, tapering gradually downward; off-white (1A1) at base, off-white (1A1) to vivid yellow (2A8); not changing color on bruising; longitudinally gnarled or rugose; with abortive branches; context of stipe off-white (1A1) to pale yellow (62A3) when fresh, drying light in weight, easily penetrated, gelatinous, firm. Major branches several, ascending, hardly terete. Branches in 3−5 ranks, major branches and lower branches commonly polychotomous; branches pale yellow (2A3) to vivid yellow (2A8), sometimes cream (4A3) to salmon (6A4); axils narrowly rounded, mostly V-shaped; internodes all short, branches terete, ascending; context of main branches and upper branches semi-gelatinous, viscid, brittle, pastel yellow (2A4). Apices broadly rounded, 1–3 mm long, fragile, polychotomous, concolorous with branches, with slightly grayish orange (6B5) at maturity. Odor weakly fragrant, aromatic; taste negligible to faintly astringent.

Basidiospores [90/3/3] (9.4–)10.5–12.9(–14.2) × (4.1–)4.3–5.5(–5.9) µm, L_m = 11.64 µm, l_m = 4.80 µm, Q = (1.94–)2.06–2.77(–2.90), Q_m = 2.43 ± 0.23, subcylindrical to narrowly ellipsoid; ornamented with moderate warts and short meandering ridges randomly placed, strongly cyanophilous warts; hilar conspicuous, slightly curved. Basidia 45–70 × 7–12 μm, clavate, without basal clamps; sterigmata 4, 3–6 µm long, short and straight. Hymenium unthickening, 55–80 µm. Subhymenium extensive, hyphae, interwoven, 1.5–3 µm diam, adherent, occasionally with clamps. Contextual hyphae: Tramal hyphae of stipe 2–14 μm wide, hyaline, tightly interwoven, heavily agglutinated, ampulliform inflations up to 15 µm; tramal hyphae of branches central hyphae slightly narrow, 2–12 µm, hyaline, thin-walled, agglutinated, strictly parallel, tightly packed, ampulliform inflations associated with septa, up to 15 µm broad, thick-walled (wall up to 1 µm thick), unornamented; gloeoplerous hyphae commonly, 4−8 μm diam.

Ecology: In the mixed forests dominated by Quercus and Picea. Basidiomata occurs in summer and autumn.

Distribution: Known from southwest China (Sichuan).

Additional materials examined: CHINA, SICHUAN PROVINCE: Ganzi, Kangding City, Gongga Mountain, in Quercus semecarpifolia and Picea asperata forest, altitude 4000 m, 7 September 2021, P. Zhang 4380 (MHHNU 10880); same location, altitude 3500 m, 2 September 2018, P. Zhang 3581 (MHHNU 10081), P. Zhang 3583 (MHHNU 10083); same city, Shade Town, 24 August 2017, P. Zhang 2752 (MHHNU 9252).

Commentary: The basidiomata color of R. gelatinosivivida is similar to R. ephemeroderma, both of which have gelatinous stipes, lack clamps, and exhibit yellow to salmon-colored basidiomata. However, it differs from the latter in having larger basidiospores dimensions and a higher length-to-width ratio (11.91 × 4.47 µm, Q_m = 2.67; as measured in this study). Additionally, R. gelatinosivivida can be confused with some other yellow to salmon-colored species, but the wrinkled and gelatinous stipe, long and narrow basidiospores, and the vivid yellow color when fresh are crucial characteristics for identification.

43. Ramaria gelatinostipes P. Zhang & J. Zhong, sp. nov. Fig. 19 i−j and Fig. 71

MycoBank: 853035

Etymology: gelatinostipes = gelatinosa + stipes, indicating that the species has a gelatinous stipe.

Diagnosis: Basidiomata are cream color; stipe is short with abortive branches, context is hard and gelatinous, heavily agglutinated; apices are broadly rounded and concolorous with branches; hyphae and basidia base exhibit clamps.

Type: CHINA, SICHUAN PROVINCE: Ganzi, Kangding City, Gongga Mountain, in Quercus semecarpifolia and Picea asperata forest, altitude 4000 m, 7 September 2021, P. Zhang 4326 (Holotype, MHHNU 10826, GenBank Acc. No.: ITS = PP467197, nrLSU = PP493517, tef1-α = PP480394).

Fig. 71. Microscopic features of R. gelatinostipes (MHHNU 10826). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Description: Basidiomata 8–11 cm high and 5–8 cm broad, single to fasciculate habit. Stipe 2–4 × 2.–4 cm, single to compound, stout; off-white (1A1) at base, off-white (1A1) to pale yellow (2A3); not changing color on bruising; surface smooth; with abortive branches; context of stipe off-white (1A1) when fresh, drying light in weight, easily penetrated, gelatinous, firm. Major branches 2−4, curved-ascending, hardly terete. Branches in 3−5 ranks, major branches and lower branches commonly polychotomous; branches pale yellow (2A3) to cream (4A3); axils narrowly rounded, mostly V-shaped; internodes diminishing gradually at maturity, lower internodes to 3 cm; branches commonly 2–15 mm, terete, ascending; context of main branches and upper branches semi-gelatinous, viscid, brittle, pastel yellow (2A4). Apices broadly rounded, 1–3 mm long, fragile, polychotomous, concolorous with branches. Odor weakly fragrant, aromatic; taste negligible to faintly astringent.

Basidiospores [63/2/2] (–8.7)8.9–11.1(–11.6) × (3.1–)3.5–5.1(–5.4) µm, L_m = 9.79 µm, l_m =4.37 µm, Q = (–1.90)1.98–2.73(–3.30), Q_m = 2.29 ± 0.17, subcylindrical to narrowly ellipsoid; ornamented with conspicuously warts and short meandering ridges randomly placed, strongly cyanophilous warts; hilar conspicuous, slightly curved. Basidia 40–55 × 8–10 μm, clavate, with basal clamps; sterigmata 4, 3–6 µm long, short and straight. Hymenium unthickening, 55–65 µm. Subhymenium extensive, hyphae, interwoven, 1.5–3 µm diam, adherent, with clamps. Contextual hyphae: Tramal hyphae of stipe 2–14 μm wide, hyaline, tightly interwoven, heavily agglutinated, ampulliform inflations not observed; tramal hyphae of branches central hyphae slightly narrow, 2–12 µm, hyaline, thin-walled, agglutinated, strictly parallel, tightly packed, ampulliform inflations associated with septa, up to 15 µm broad, thick-walled (wall up to 1 µm thick), unornamented; gloeoplerous hyphae commonly, 4−8 μm diam.

Ecology: In the mixed forests dominated by Quercus and Picea. Basidiomata occurs in summer and autumn.

Distribution: Known from southwest China (Sichuan).

Additional materials examined: CHINA, SICHUAN PROVINCE: Ganzi, Kangding City, Gongga Mountain, in Quercus semecarpifolia and Picea asperata forest, altitude 4000 m, 5 September 2021, P. Zhang 4355 (MHHNU 10855), P. Zhang 4362 (MHHNU 10862).

Commentary: The species most similar to R. gelatinostipes is R. ossolana Franchi & M. Marchetti and R. flavoides Schild and its original variety. These species all have basidiomata with a light yellow to cream color, gelatinous flesh, and the presence of clamps. The differences lie in the fact that R. ossolana has a spicy taste, an unpleasant odor, and slightly smaller basidiospores (9.3 × 4.3 µm; Franchi & Marchetti 2021). The basidiomata of R. flavoides and its original variety tend to be bright yellow color.

44. Ramaria indoyunnaniana R.H. Petersen & M. Zang, Acta bot. Yunn. 8(3): 287, 1986. Fig. 19 k−l and Fig. 72

Description: Basidiomata 8 cm high and 4 cm broad, obovate in outline. Stipe discrete, short to long, fleshy, solid, without abortive branches, smooth, ivory-colored; flesh off-white, very slightly soapy when sliced and rubbed. Major branches are few, ascending but not strict, shell pink when adolescent; small soil particles causing weak wine-red spots; axils rounded. Apices more or less acute, probably terraced when young, rich muted rose when adolescent, perhaps somewhat pallid when juvenile and probably fading to pallid ochraceous in age; Odor weakly fabaceous; taste negligible; apparently edible.

Fig. 72. Basidiospore features of R. indoyunnaniana (MHHNU 9134). Scale bar: a = 10 µm.

Basidiospores [40/2/2] (5.7–)6.2–8.8(–9.5) × (3.1–)3.3–4.8(–4.9) µm, L_m = 7.18 µm, l_m = 4.16 µm, Q = (1.50–)1.51–2.03(–2.04), Q_m = 1.75 ± 0.15, cylindrical, roughened in profile; wall up to 0.3 µm thick; hilar appendix eccentric, rarely discrete; ornamentation of obscure, hardly cyanophilous patches up to 0.2 µm high. Basidia 35−45 × 6−7.5 μm, clavate, without clamps; sterigmata 4, spindly, straight. Subhymenium extensive, hyphal. Hymenium thickening. Tramal hyphae of stipe 10 µm wide, hyaline, without clamps, probably adherent; Tramal hyphae of upper branches up to 8 µm wide, hyaline, thin walled, without clamps, adherent, tightly packed, strictly parallel.

Ecology: Solitary or gregarious on soil in broad-leaved forests. Basidiomata occurs in summer and autumn.

Distribution: Known in southwestern, central China and east China (Yunnan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Pu’er City, 20 September 1983, Petersen45681 (isotype, HKAS 10422); Pu’er City, Menglang Town, bought from mushroom market, 30 July 2017, P. Zhang 2634 (MHHNU 9134); Wenshan, Malipo County, Xinhe Village, in broad-leaved forest with Fagaceae, altitude 1200 m, 4 August 2018, P. Zhang 3339 (MHHNU 9839).

Commentary: The species is what (Sharda 1983) referred to as R. araiospora var. rubella Marr & Stuntz. The fruiting bodies exhibit a bright pinkish red coloration, not the distinctly magenta red of R. araiospora, and produce somewhat smaller basidiospores compared with R. araiospora (9.9 × 3.7 µm; Marr & Stuntz 1973). Additionally, R. araiospora and other similar species R. cyaneigranosa Marr & Stuntz (9.6 × 4.7 µm; Marr & Stuntz 1973) typically have larger basidiomata compared to those reported by Sharda. Instead, the Asian taxon appears most similar to R. rosella Petersen from the southern Applachian Mountans of eastern North America (Petersen 1985). Ramaria rosella shares almost identical coloration and stature but produces somewhat larger basidiospores (9.87 × 4.16 µm; Petersen 1985).

45. Ramaria microspora (R.H. Petersen & M. Zang) P. Zhang & J. Zhong, stat nov. et comb. nov. Fig. 19 m−n and Fig. 73

≡ Ramaria botrytoides var. microspora R.H. Petersen & M. Zang, Acta bot. Yunn. 11(4): 372 (1989)

MycoBank: MB 853380

Basionym: Ramaria botrytoides var. microspora R.H. Petersen & M. Zang, Acta bot. Yunn. 11(4): 372, 1989.

Etymology: “microspora” refers to the small basidiospores of the species, thus elevating “R. botrytoides var. microspora” to the rank of species “R. microspora”.

Isotype: CHINA, YUNNAN PROVINCE: Lijiang City, bought from mushroom market, under Pinus armandi, P. yunnanensis, and Quercus rehderiana mixed forest, 30 August 1986, HKAS 20072.

Fig. 73. Basidiospore features of R. microspore (MHHNU 9914). Scale bar: a = 10 µm.

Description: Basidiomata 8 cm high and 10.5 cm broad, single to cespitose, obtriangular in outline. Stipes up to 3 × 2 cm, single, tapering sharply downward to an acute or gnarled base, involving significant substrate when picked; white or off-white at base, off-white to pale yellow upward; not changing color on bruising; pruinose at stipe base, smooth upward; with abortive branchlets, abortive branchlets bright pink rose pink where protected; flesh off-white when flesh, appearing hydrophanous, rubbery or firm-gelatinous, drying hard, cartilaginous. Major branches 2-4, curved ascending, hardly terete. Branches in 3-6 ranks, major branches and lower branches commonly polychotomous, orange-white when young, deepens slightly by maturity; axils narrowly rounded, mostly U-shaped; internodes diminishing gradually at maturity, lower internodes to 3 cm; branches commonly 2–10 mm, hardly terete, ascending; context of main branches and upper branches semi-gelatinous, viscid, brittle, off-white. Apice finely divided, 1–3 mm, delicate, double-dichotomous, rose-pink when protected or young, fading rapidly where exposed carrot red, finally concolorous with branches in age. Odor faintly fabaceous; taste negligible.

Basidiospores [22/2/2] (7.0–)7.2–9.5(–10.0) × 4.0–5.0 µm, L_m = 8.49 µm, l_m = 4.32 µm, Q = (1.67–)1.68–2.38, Q_m = 1.97 ± 0.23, short-cylindrical to broadly ellipsoid, flattened adaxially, occasionally pip-shaped, conspicuously roughened in profile; ornamentation of plate-like warts and short ridges randomly placed, strongly cyanophilous warts; hilar appendix prominent, slightly curved. Basidia 57–65 × 8–9 µm, clavate, without clamps at base; sterigmata 4, 4–7 µm long, slender, curved. Hymenium thickening, 60–85 µm. Subhymenium extensive, pseudo-parouchymatous, 1.5–2.5 µm, adherent, without clamps. Contextual hyphae: Tramal hyphae of stipe 4–12 μm wide, hyaline, thin-walled, unclamped, tightly interwoven, heavily agglutinated, ampulliform inflations not observed; tramal hyphae of branches central hyphae slightly narrow, 3–11 µm, hyaline, thin walled, agglutinating substance easily liberated into 2% KOH mounts as amorphous debris), strictly parallel, tightly packed, ampulliform inflations associated with septa, up to 13 µm broad, thick-walled (wall up to 1 µm thick), unornamented; without clamps; gloeoplerous hyphae commonly, 4−8 μm diam.

Ecology: Found in mixed forests dominated by Quercus or in mixed forests. Basidiomata occurs in summer and autumn.

Distribution: Known from southwest China (Yunnan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Shangri-La City, Pudacuo National Park, Bitahai, in the mixed forest with Quercus and Picea. altitude 3500 m, 31 August 2022, P. Zhang 4842 (MHHNU 11342); same location, Shuduhu, in the Quercus forests, altitude 3600 m, 15 August 2018, P. Zhang 3412 (MHHNU 9912), P. Zhang 3414 (MHHNU 9914); Qujing City, Qilin District, near Longyuan Zhumushan Wind Farm, in the broad-leaved forest, altitude 2100 m, 19 August 2022, P. Zhang 4707 (MHHNU 11207); Chuxiong, Zixi Mountain, in a forest with Lithocarpus and Pinus yunnanensis, bought from mushroom market, 3 August 2001, P. Zhang 119 (MHHNU 6619), P. Zhang 120 (MHHNU 6620).

Commentary: The main features of R. microspora include cream-colored branches with bright rose-pink clustered apices, which retain their pink hue well when dry but may fade after maturation or due to environmental factors. The stipe is short with firmly gelatinous flesh characterized by heavily agglutinated hyphae, often forming abortive clusters with rosy-pink apices. Hyphae and basidia base lack clamps, and basidiospores are small sized with a thickened hymenium.

Ramaria botrytoides and their original variety are distinctly different both in the field and microscopically. Widely distributed in North America and reported in China, R. botrytoides differs from R. microspora primarily in having slender basidiospores (7.0–9.5 × 3.0–3.7 µm, Corner 1950) with less pronounced ornamentation.

Despite similarities between two species collected in China and North America—such as gelatinous stipes, pink branch apices, and small basidiospores —there are notable differences. Firstly, basidiospores from the Chinese collection are smaller than the smallest basidiospores found in North American specimens. Secondly, basidiospores ornamentation in the Chinese collection is significantly more prominent compared to North American specimens. Thirdly, Chinese basidiomata react with several macrochemical reagents, whereas North American specimens are relatively inert.

46. Ramaria platyrugosa L. Fan, Ying Li & N. Mao, in Li, Mao, Zhang, Fu & Fan, Mycology: 10.1080/21501203.2024.2388910, 26 (2024)

Original description in Li et al. (2024).

47. Ramaria rubrisubtilis P. Zhang & J. Zhong, sp. nov. Fig. 19 o−p and Fig. 74

MycoBank: MB 853379

Etymology: rubrisubtilis = “rubr” + “subtilis”, referring to the species having slightly red apices.

Diagnosis: Apices slightly pink colored, retaining color when dry but may lighten with mature or environment factors; stipe slender, basidiomata obpyramidal in outline; bruising turns slightly brownish when injured; hyphae and basidia base lack clamps.

Type: CHINA, HUNAN PROVINCE: Chenzhou City, Yizhang County, Mangshan National Nature Reserve, 5 September 2007, P. Zhang 660 (Holotype, MHHNU 7160, GenBank Acc. No.: ITS = PP467202, nrLSU = PP493521).

Description: Basidiomata 6.5–15 cm high and 5–12 cm broad, single to fasciculate habit, usually obovate in outline. Stipe 2–3 × 1–2.5 cm, single, fleshy, slender, tapering downward evenly to narrowly rounded base; off-white (1A1) at base, off-white (1A1) to pale yellow (2A3); slightly changing color to brownish yellow (5B3) on bruising; surface smooth; with abortive branches; context of stipe off-white (1A1) when fresh, drying light in weight, easily penetrated, cartilaginous, brittle. Branches in 3−5 ranks, major branches and lower branches commonly polychotomous, upward branches dichotomous; branches ivory (4B3) to cream (4A3) when young, surface smooth, flesh color (6A3) and rugose at age; axils narrowly rounded, mostly V-shaped; internodes diminishing gradually at maturity, lower internodes long, up to 1.5 cm; branches commonly 1–3 mm, terete, sub-parallel; context of main branches and upper branches fleshy to gelatinous, off-white (1A1). Apices broadly rounded, 1–2 mm long, dichotomous, shell pink (8A3) to pink (10A3) and the color remains unchanged after collecting when young, the color lighter when old or influenced by environmental factors. Odor weakly fragrant, aromatic; taste negligible to faintly astringent.

Fig. 74. Microscopic features of R. rubrisubtilis (MHHNU 7160). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Basidiospores [60/2/2] 10.0–12.0 × 4.0–6.0(–7.0) µm, L_m = 10.76 µm, l_m = 5.24 µm, Q = 1.67–2.50(–2.75), Q_m = 2.07 ± 0.24, subcylindrical to long ellipsoid; ornamented with conspicuously warts and short meandering ridges randomly placed, strongly cyanophilous warts; hilar conspicuous, slightly curved. Basidia 46–72 × 7–12 μm, clavate, without basal clamps; sterigmata 4, 4–7 µm long, incurved or straight, slightly divergent. Hymenium unthickening, 45–75 µm. Subhymenium extensive, 10–15 µm thick, hyphae, interwoven, 3–6 µm diam, thin-walled, without clamps. Contextual hyphae: Tramal hyphae of stipe 3–12 μm wide, hyaline, loosely interwoven, inflating slightly, terminal swollen hyphae wide to 14 µm, with moderately ornamented; tramal hyphae of branches central hyphae slightly lysing, terminal swollen hyphae wide to 15 µm, with smooth and thin wall; ampulliform hyphae rarely; without clamps; gloeoplerous hyphae occasionally, 5−8 μm diam.

Ecology: Found in broad-leaved forests or mixed forests. Basidiomata occurs in summer and autumn.

Distribution: Known from southwestern and central China (Yunnan, Hunan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Chuxiong, Nanhua County, Dazhongshan Forestry Farm, in a forest dominated by Fagaceae, altitude 2400 m, 11 October 2021, Jian W. Liu 2574 (MHHNU 11651). HUNAN PROVINCE: Zhangjiajie City, Sangzhi County, Badagongshan National Nature Reserve, in the mixed forest, altitude 1400 m, 15 September 2020, Zuo H. Chen 32514 (MHHNU 32514).

Commentary: Ramaria rubrisubtilis, with its slightly pink branch apices, is visually similar to R. rubroapicalis, R. indoyunnaniana, R. botrytoides and its variants, all of which have pink to red branch apices. They share a close phylogenetic relationship and lack clamps. The main distinguishing feature of R. rubrisubtilis is that its pink color is only present at the top 1–2 mm of the apices, and the pink color can easily fade with maturity or environmental influence, finally appearing ivory to pale yellow color. Additionally, R. rubroapicalis and R. indoyunnaniana have relatively smaller basidiospores sizes (9.76 × 4.63 µm; 7.18 × 4.16 µm; as measured in this study). As R. botrytoides is a broadly defined species with its type locality in North America, and R. rubrisubtilis has only been found in the central region of China, with basidiospores sizes significantly larger than those documented in the literature, it can be confirmed that the two are not the same species.

48. Ramaria rubroapicalis P. Zhang & J. Zhong, sp. nov. Fig. 19 q−r and Fig. 75

MycoBank: MB 853378

Etymology: rubroapicalis = “rubr” + “apicalis”, referring to the species having red apices.

Fig. 75. Microscopic features of R. rubroapicalis (MHHNU 11218). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Diagnosis: Apices are pink and retain color without fading easily; stipe is single, with basidiomata obpyramidal in outline; slightly turns to brownish when injured; basidiospores are less than 10 µm in average length, roughened in profile; hyphae and basidia base lack clamps.

Type: CHINA, YUNNAN PROVINCE: Qujing City, Qilin District, near Longyuan Zhumushan Wind Farm, altitude 2250 m, 19 August 2022, P. Zhang 4718 (Holotype, MHHNU 11218, GenBank Acc. No.: ITS = PP467205, nrLSU = PP493523).

Description: Basidiomata 8.5 cm high and 8 cm broad, single to fasciculate habit, medium-sized, obpyramidal in outline. Stipe 3 × 2 cm, single, fleshy, moderately to large, tapering gradually downward, rounded at base; off-white (1A1) at base, cream (4A3) to pale yellow (2A3) with slightly shell pink (8A3) upward; slightly changing color to brownish yellow (5B3) on bruising; surface smooth; without abortive branches; context of stipe off-white (1A1) when fresh, fibrous to spongy, drying light in weight, easily penetrated. Branches in 3−7 ranks, major branches and branches above commonly polychotomous; branches off-white (1A1) to pale yellow (2A3); axils narrowly rounded, mostly V-shaped; internodes diminishing gradually at maturity, lower internodes up to 3 cm; branches commonly 3–8 mm, terete, ascending; context of main branches and upper branches fleshy to gelatinous, off-white (1A1). Apices cusped to minutely double-dichotomous, 1–2 mm long, axils near apices 2–8 times, pink (10A3) to light red (10A5). Odor and taste faintly fabaceous.

Basidiospores [30/2/1] (8.3–)8.9–11.0 × (4.0–)4.1–5.2(–5.3) µm, L_m = 9.76 µm, l_m = 4.63 µm, Q = (1.80–)1.90–2.34(–2.38), Q_m = 2.11 ± 0.15, subcylindrical to narrowly ellipsoid; ornamented with conspicuously warts and short meandering ridges randomly placed, strongly cyanophilous warts; hilar conspicuous, slightly curved. Basidia 35–55 × 8–12 μm, clavate, without basal clamps; sterigmata 4, 2–8 µm long, incurved or straight, slightly divergent. Hymenium unthickening, 55–70 µm. Subhymenium extensive, 15–20 µm thick, hyphae, interwoven, 3–6 µm diam, thin-walled, without clamps. Contextual hyphae: Tramal hyphae of stipe 3–12 μm wide, hyaline, loosely interwoven, inflating slightly, terminal swollen hyphae wide to 14 µm, with moderately ornamented; tramal hyphae of branches central hyphae slightly lysing, terminal swollen hyphae wide to 18 µm, with smooth and thin wall; ampulliform hyphae rarely; without clamps; gloeoplerous hyphae occasionally, 7−10 μm diam.

Ecology: In broad-leaved forests. Basidiomata occurs in summer and autumn.

Distribution: Known from southwestern and east China (Yunnan, Hunan, Fujian, Zhejiang).

Additional materials examined: CHINA, YUNNAN PROVINCE: Qujing City, Qilin District, near Longyuan Zhumushan Wind Farm, altitude 2160 m, 28 August 2020, Li P. Tang HT277 (MHKMU HT277). Xishuangbanna, Jinghong City, Menglun Town, near the Kino Gurry, in the broad-leaved forest, altitude 900 m, 14 August 2005, P. Zhang 451 (MHHNU 6951); same city, altitude 900 m, 9 July 2006, Jun F. Liang 351 (HKAS 50067), Jun F. Liang 353 (HKAS 50069); HUNAN PROVINCE: Chenzhou City, Rucheng County, Daping Town, Jiulongjiang National Forest Park, in broad-leaved forest, altitude 100 m, 2 October 2020,P. Zhang 4069 (MHHNU 10569); same location, 22 June 2020, P. Zhang 3582 (MHHNU 10352). FUJIAN PROVINCE: Sanming City, Jiangle County, Longqi Mountain National Nature Reserve, 21 August 2023, Nian K. Zeng 8161 (MHHNU 11626). ZHEJIANG PROVINCE, Lishui City, Qingyuan County, 11 October 2023, P. Zhang 5100 (MHHNU 11600), P. Zhang 5101 (MHHNU 11601).

Commentary: Ramaria rubroapicalis have pink apices and white to creamy-colored basidiomata, it resembles to R. indoyunnaniana R.H. Petersen & M. Zang found in the southwestern of China. The key difference lies in the smaller basidiospores size of R. indoyunnaniana (7.18 × 4.16 µm, as measured in this study).

Ramaria rubroapicalis shares morphological similarities with the globally distributed species R. botrytoides (Peck) Corner and its variants. However, the definition of R. botrytoides is unclear, described by Corner as a species with short basidiospores (7.0–9.5 × 3.0–3.7 µm, Corner 1950), pinkish apices, gelatinous flesh, common in North America, ranging from salmon-rose colors, and exhibits similar habits to R. conjunctipes, basidiospores walls finely rough to nearly smooth, and hyphae with clamps. In Zang Mu and Petersen's articles (Petersen & Zang 1989), R. botrytoides is lacks clamps in subgen. Laeticora, with abortive branches in stipe, and apices bright yellow.

Considering the comprehensive information, especially the basidiospores size, it becomes evident that R rubroapicalis does not align with the literature's depiction of R. botrytoides.

49. Ramaria sinensis (R.H. Petersen & M. Zang) P. Zhang & J. Zhong, stat nov. et comb. nov. Fig. 19 s−t and Fig. 76

≡ Ramaria neoformosa var. sinensis R.H. Petersen & M. Zang, Acta bot. Yunn. 11(4): 391 (1989)

MycoBank: MB 855602

Etymology: “sinensis” means collected in China.

Diagnosis: Basidiomata with a linear stature, featuring small but single stipes that superficially appear fasciculate; lower branches are golden yellow, while upper branches exhibit salmon or orange hues with golden yellow apices; basidia lack clamps; basidiospores are prominently roughened.

Isotype: CHINA, YUNNAN PROVINCE: Dali, Eryuan County, Sanying Town, 27 August 1986, Petersen 47305 (HKAS 20043).

Description: Basidiomata 14 cm high and 6 cm broad, linear, elongate obpyriform in outline. Stipes up to 1 × 1.5 cm, small, tapered to an acute base, white, smooth, often with abortive branchlets, reluctantly brunnescent on handing; flesh white, solid, moist but not slippery, often mottled hydrophanous; abortive branchlets yellow to pastel orange; surface of stipe and lower branches becoming grey upon drying. Major branches several, strictly ascending, terete, clear yellow. Branches in 4–7 ranks, terete, strict, orange, pastel orange to pastel salmon, fading in age to "ochraceous buff"; internodes somewhat long throughout, diminishing gradually upward at maturity; axils narrowly rounded below, rounded above; flesh solid, sometimes slippery when cut, orange. Apices fine, delicate, knobby-digitate by maturity, somewhat divaricate, golden yellow to pastel orange. Odor negligible; taste negligible or faintly fungoid.

Basidiospores [50/2/2] (11.2–)11.8–15.0 × (4.8–)5.0–5.8(–6.0) µm, L_m = 13.56 µm, l_m = 5.14 µm, Q = (2.18–)2.24–2.88(–3.13), Q_m = 2.63 ± 0.23, narrowly ellipsoid, conspicuously roughened in profile; wall up to 0.2 µm thick; hilar appendix gradual, prominent, almost perpendicular to basidiospores axis; ornamentation of prominent elongate warts and anastomosing, meandering ridges. Basidia 57–65 × 9–11 µm, clavate, without clamps; moderately cyanophilous; sterigmata 4, short, spindly, straight to curved. Hymenium thickening. Subhymenium rudimentary; hyphae 1.5–2.5 µm, tightly packed, interwoven, without clamps. Tramal hyphae of stipe 4–12 µm wide, hyaline, thick-walled (wall up to 0.5 µm thick), without clamps, tightly interwoven, not agglutinated or adherent; ampulliform swelling rare, not associated with septa, not unusually thick-walled, with coarse, extensive stalactitiform ornamentation; gloeoplerous hyphae not observed. Tramal hyphae of upper branches up to 4–11 µm wide, hyaline, without clamps, thin-walled, parallel, tightly packed, free to locally adherent; ampulliform swellings not observed; gloeoplerous hyphae sometimes common.

Fig. 76. Basidiospore features of R. sinensis (MHHNU 9177). Scale bar: a = 10 µm.

Ecology: In mixed forests dominated by Quercus and Picea. Basidiomata occurs in summer and autumn.

Distribution: Known in southwestern China (Yunnan, Sichuan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Lijiang, Yulong County, Laojun Mountain, in a subalpine forest dominated by Abies, altitude 3800 m, 14 August 2000, Zhu L. Yang 2920 (HKAS 36620); same city, Tiejia Mountain, in a forest with Pinus yunnanensis, altitude 2800 m, 31 July 2001, P. Zhang 79 (MHHNU 6679), P. Zhang 80 (MHHNU 6680); Kunming City, Yiliang County, Zhongdian, Hong Mountain, in Quercus and Pinus mixed forest, altitude 3700 m, 8 August 2005, Zhu L. Yang 4568 (HKAS 48732); Dali, Dali City, Xianyangfeng, in a mixed forest, altitude 2800 m, 12 August 2018, P. Zhang 3378 (MHHNU 9878). SICHUAN PROVINCE: Ganzi, Kangding City, Pengta Township, altitude 3100 m, 19 August 2017, P. Zhang 2677 (MHHNU 9177).

Commentary: Petersen brought a single fruiting body of this species from the market and identified it as a variant of R. neoformosa. Within the species, two varieties have been described, basidiospores of R. neoformosa var. neoformosa are (10.7–11.8 × 4.8–5.6 µm, L_m = 10.9 µm, l_m = 2.09 µm; Petersen 1976), R. neoformosa var. macrospora Pet. known only from eastern Canada and with basidiospores (11.2–14.8 × 3.6–4.3 µm, L_m = 12.57 µm, l_m = 3.01 µm; Petersen 1986). Our basidiospores measurements of R. sinensis slightly exceeded those reported by Petersen, with slightly larger and wider basidiospores. However, the bright pink flesh of branches and our phylogenetic analyses significant its identification as R. neoformosa var. sinensis (≡ R. sinensis).

50. Ramaria subgelatiniaurantia P. Zhang & J. Zhong, sp. nov. Fig. 19 u−v and Fig. 77

MycoBank: 853033

Etymology: The prefix “sub-” indicates “similarity”. The species epithet “subgelatiniaurantia” suggests that this species is similar to R. gelatiniaurantia, sharing features such as gelatinous flesh and an orange color of the basidiomata.

Diagnosis: Basidiomata are orange-yellow in color, with apices concolorous with branches; stipe is short, stout, gelatinous, with abortive branches; does not change color when injured; hyphae and basidia base lack clamps.

Fig. 77. Microscopic features of R. subgelatiniaurantia (MHHNU 10859). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Type: CHINA, SICHUAN PROVINCE: Ganzi, Kangding City, Gongga Mountain, in Quercus semecarpifolia and Picea asperata forest, altitude 4000 m, 7 September 2021, P. Zhang 4359 (Holotype, MHHNU 10859, GenBank Acc. No.: ITS = PP467201, nrLSU = PP493520).

Description: Basidiomata 9 cm high and 9 cm broad, single, usually obtriangular in outline. Stipe 2.5 – 2 cm, single, stout and short, tapering downward evenly to rounded base; off-white (1A1) to pastel yellow (2A4); not changing color on bruising; surface smooth; with abortive branches; context of stipe off-white (1A1) when fresh, drying light in weight, easily penetrated, gelatinous, firm. Branches in 3−5 ranks, major branches and lower branches commonly polychotomous; branches pale yellow (2A3) to yellowish orange (4A8); axils narrowly rounded, mostly V-shaped; internodes diminishing gradually at maturity, lower internodes to 4 cm; branches commonly 2–25 mm, terete, ascending; context of main branches and upper branches semi-gelatinous, off-white (1A1). Apices broadly rounded, 2–6 mm long, polychotomous, concolorous with branches. Odor weakly fragrant, aromatic; taste negligible to faintly astringent.

Basidiospores [60/2/2] (9.0–)9.4–12.0(–12.4) × (3.4–)3.7–5.0(–5.4) µm, L_m = 10.67 µm, l_m =4.47 µm, Q = (1.99–)2.10–2.98(–3.18), Q_m = 2.51 ± 0.26, subcylindrical to narrowly ellipsoid; ornamented with conspicuously warts and short meandering ridges randomly placed, strongly cyanophilous warts; hilar conspicuous, slightly curved. Basidia 46–72 × 7–12 μm, clavate, without basal clamps; sterigmata 4, 4–6 µm long, incurved or straight, slightly divergent. Hymenium unthickening, 60–75 µm. Subhymenium extensive, 20–40 µm thick, hyphae, interwoven, 3–6 µm diam, thin-walled, without clamps. Contextual hyphae: Tramal hyphae of stipe 3–12 μm wide, hyaline, loosely interwoven, inflating slightly, terminal swollen hyphae wide to 14 µm, with moderately ornamented; tramal hyphae of branches central hyphae slightly lysing, terminal swollen hyphae wide to 15 µm, with smooth and thin wall; ampulliform hyphae rarely; without clamps; gloeoplerous hyphae commonly, 4−8 μm diam.

Ecology: Found in mixed forests dominated by Quercus and Picea. Basidiomata occurs in summer and autumn.

Distribution: Known from southwest China (Sichuan).

Additional material examined: CHINA, SICHUAN PROVINCE: Ganzi, Kangding City, Gongga Mountain, in Quercus semecarpifolia and Picea asperata forest, altitude 4000 m, 7 September 2021, P. Zhang 4358 (MHHNU 10858).

Commentary: Ramaria subgelatiniaurantia has only been collected in the southwestern of China until now. In fact, characterized by gelatinous stipes, orange-yellow colored basidiomata, and lack of clamps is a widespread complex group species, found in North America, Asia, and Europe. In Franche's work, he identified a species closely related to R. subgelatiniaurantia as R. primulina.

Ramaria primulina, reported by Petersen in 1986 with its type locality in Canada, has also been observed by Franche in Europe, particularly in Spain, Italy, Finland, and Norway. Franche (2021) described the European specimens as having pale yellow color, U-shaped axils, gelatinous context hyphae, and basidiospores measuring 8.2–11.2 × 3.5–5.2 µm, occasionally with clamps, growing in coniferous forests. However, Petersen described R. primulina with fibrous stipes and have clamps, leading to the suggestion that Franche's identification of R. primulina might not be the same species Petersen described. Franche's observations showing slightly smaller basidiospores on the type specimen compared to Petersen's description. Differences in basidiomata color and basidiospores size, combined with the results of our two-gene phylogenetic analysis, confirm that R. subgelatiniaurantia is not the same as Franche's, but aligns with the complex reported by Marr in 1973, including R. gelatiniaurantia and its variants.

The complex of R. gelatiniaurantia and its variants includes R. gelatiniaurantia, R. hilaris var. olympiana, R. flavigelatinosa, and R. verlotensis. In this complex, R. flavigelatinosa and R. hilaris var. olympiana have yellow branch flesh, while R. verlotensis has a pale orange basidiomata, wide cauliflower-shaped basidiomata with an overall yellow-pink hue. These characteristics confirm that R. subgelatiniaurantia does not belong to the complex described above.

51. Ramaria subthindii P. Zhang & J. Zhong, sp. nov. Fig. 19 w−x and Fig. 78

MycoBank: MB 853373

Etymology: The specific epithet "subthindii" means that the species is similar to R. thindii.

Diagnosis: Basidiomata are small to medium sized and pale yellow; stipe is short, stout, gelatinous, and smooth; hyphae and basidia base lack clamps; average basidiospores length exceeds 10 µm.

Fig. 78. Microscopic features of R. subthindii (MHHNU 9131). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Type: CHINA YUNNAN PROVINCE: Pu’er City, Lancang County, Menglang Town, altitude 980 m, brought in mushroom market, 30 July 2017, P. Zhang 2631 (Holotype, MHHNU 9131, GenBank Acc. No.: ITS = PP467187, nrLSU = PP493507, tef1-α = PP480389).

Description: Basidiomata 6–8 × 6–7 cm, fasciculate habit, usually obpyriforme to broadly fusiform in outline. Stipes up to 1.5–3 × 2–3 cm, single to compound, tapering gradually downward; milk white (1A2) at base, milk white (1A2) to pale yellow (2A3) upward; surface smooth; with abortive branches; some region changing color to brownish gray (6E2) on bruising or purple spots caused by soil particles; context of stipe off-white (1A1) to jelly-like pastel yellow (2A4) when fresh, drying light in weight, easily penetrated, gelatinous, hard. Major branches several, ascending, terete. Branches in 3−5 ranks, major branches and lower branches commonly dichotomous; branches pale yellow (2A3) to chicken yellow (2A6); axils broadly rounded, mostly U-shaped; internodes all short; branches terete, sparsely, subparallel upward; context of main branches and upper branches gelatinous, viscid, brittle, pastel yellow (2A4). Apices broadly rounded, molar-like when young, terraced, short-digitate at maturity, concolorous with branches or slightly pale yellow (2A3) to chicken yellow (2A6). Odor weakly fragrant, aromatic; taste negligible to faintly astringent.

Basidiospores [50/2/2] (8.5–)9.0–12.6(–13.9) × 4.0–5.4(–5.6) µm, L_m = 10.87 µm, l_m = 4.91 µm, Q = (–1.99)2.00–2.71(–2.84), Q_m = 2.34 ± 0.23, ellipsoid, noticeably rough; ornamented with warts and short meandering ridges randomly placed, strongly cyanophilous warts; hilar conspicuous, slightly curved. Basidia 45–65 × 8–11 μm, clavate, without basal clamps; sterigmata 4, 3–6 µm long, short and straight, somewhat scattered. Hymenium unthickening, 50–70 µm. Subhymenium extensive, hyphae, interwoven, 1–3 µm diam, adherent, without clamps. Contextual hyphae: Tramal hyphae of stipe 3–10 μm wide, hyaline, tightly interwoven, heavily agglutinated, ampulliform inflations occasionally; tramal hyphae of branches central hyphae slightly narrow, 2–10 µm, hyaline, thin-walled, agglutinated, strictly parallel, tightly packed, ampulliform inflations rare; gloeoplerous hyphae commonly, 4−8 μm diam.

Ecology: Solitary or gregarious on soil in mixed forests dominated by Fagaceae and Pinaceae. Basidiomata occurs in summer and autumn.

Distribution: Known from southwest China (Yunnan).

Additional material examined: CHINA YUNNAN PROVINCE: Pu’er City, Ning'er Hani and Yi Autonomous County, 1 September 2008, B. Feng 276 (HKAS 55387).

Commentary: The Chinese species R. subthindii shares a close morphological affinity with the type collection of Ramaria thindii from India, both characterized by yellow basidiomata, gelatinous flesh, and absence of clamps. However, R. subthindii has a nearly 1:1 ratio of length to width in its basidiomata, a short and stout stipe, internodasl all short, and an overall yellowish-white coloration without orange hues in the basidiomata. In contrast, R. thindii has more linear basidiomata, slenderer stipe, and mature basidiomata with orange or pink hues, along with slightly smaller basidiospores.

Moreover, R. subthindii is easily confused with R. hunanensis from Hunan, China. Macroscopically, they are indistinguishable in the field, and under the microscope, their basidiospores sizes are quite similar. However, R. hunanensis is currently known only from Hunan, while R. subthindii is distributed in the southwestern China. Our two-gene phylogenetic analysis also indicates a close phylogenetic relationship between the two, confirming them as distinct but closely related species within the Ramaria. Ramadia verlotenses Marr & D.E. Stuntz (1974) was recovered as phylogenetically closely related to R. subthindii. The two species share similar basidiospore dimensions (with R. verlotenses measuring 10.1 × 4.9 µm) but differ in basidiomata coloration: R. subthindii is pale yellow to chicken yellow, whereas R. verlotenses exhibits pallid salmon to salmon-colored basidiomata.

Ramaria ser. Hemirubellae P. Zhang & J. Zhong, ser. nov.

MycoBank: MB 852712

Etymology: “hemi” means “half” or “semi”, “rubella” means “reddish” or “red”, is proposed because the collections of the series are pale reddish.

Type species: R. hemirubella R.H. Petersen & M. Zang, Acta bot. Yunn. 8(3): 285, 1986.

Diagnosis: basidiomata are milk white to pale yellow, with apices concolorous with branches or pale red; Darkens to brunnescent or rubric-brunnescent when injured; basidia base and hyphae exhibit clamps; basidiospores are ornamented with conspicuous longitudinal striae.

Commentary: This series has collected four species in China. Additionally, another taxon within this series is characterized by a yellow basidiomata, known as R. flavissima Schild (= R. pallidissima Schild & G. Ricci).

52. Ramaria eryuanensis R.H. Petersen & M. Zang, Acta bot. Yunn. 11(4): 364, 1989. Fig. 20 a−b and Fig. 79

Fig. 79. Basidiospores of R. eryuanensis (MHHNU 10837). Scale bar: a =10 µm.

Description: Basidiomata 12 cm high and 8 cm broad. Stipe 7 × 5 cm, massive, rounded at base, sometimes rooting, smooth, somewhat flattened, often with a few abortive stumps high on stipe, somewhat flattened laterally, white at base, sometime suffused pinkish at base, off-white above, easily brunnescent on all surfaces where handled or rubbed, flesh white, solid, not gelatinous or slippery, sometimes slowly sordid grey where cut. Major branches 2–4, stout, short, more or less terete, off-white (pale ochraceous buff, pale pinkish cinnamon). Branches in 3–5 ranks, short, elongating reluctantly, crowded, more or less terete, pallid rose-beige (pinkish buff), easily brunnescent; flesh off-white, fibrous; internodes short, hardly diminishing above; axils narrowly rounded throughout. Apices digitate to awl-shaped, long, crowded, pastel rose-tan (light pinkish cinnamon, light ochraceous cinnamon), easily brunnescent. Odor faintly; taste negligible to faintly fabaceous.

Basidiospores [132/5/5](9.5–)10.7–13.1(–14.2) × (4.0–)4.2–6.0(–7.0) µm, L_m = 11.74 µm, l_m = 4.94 µm, Q = (1.57–)1.98–2.80(–3.07), Q_m = 2.39 ± 0.29, subboletoid to narrowly ellipsoid, smooth in profile; wall up to 0.2 µm thick; hilar appendix prominent, with a throat; ornamentation of coarse striae, arranged in abaxial-distal to adaxial-proximal orientation. Subhymenium extensive, hyphal. Hymenium thickening. Tramal hyphae of stipe 3–11 µm wide, hyaline, clamped, thin to thick-walled (wall up to 1 µm thick), freely branched, interwoven, not agglutinated or adherent; ampulliform inflations as clamps or asymmetrical, not unusually thick-walled, with extensive stalactitiform ornamentation; gloeoplerous hyphae not observed. Tramal hyphae of upper branches 2–8 µm diam, hyaline, thick-walled (wall up to 1 µm thick), straight, occasionally branched, parallel, not agglutinated or adherent; ampulliform clamps up to 15 µm broad, not unusually thick-walled, asymmetrical, unornamented; gloeoplerous hyphae not observed.

Ecology: In broad-leaved or coniferous forest; basidiomata occurs in summer and autumn.

Distribution: Known from southwestern China (Yunnan, Sichuan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Shangri-La, Pudacuo, Shudu Lake, in the forest with Quercus semecarpifolia, altitude 3600 m, 15 August 2018, P. Zhang 3415 (MHHNU 9915); Lijiang City, brought from Lijiang mushroom market, 2 September 1986, Petersen 47306 (isotype, HKAS 20044); same city, Yulong County, Yulong Snow Mountain, altitude 3700 m, 27 July 2017, P. Zhang 2800 (MHHNU 9300); same city, Tuanshan Reservoir, altitude 2500 m, Li P. Tang 361 (HKAS54592); Kunming City, bought from mushroom market, 23 July 2017, P. Zhang 2601 (MHHNU 9101); Chuxiong, Zixi Mountain, in a forest with Lithocarpus and Pinus yunnanensis, altitude 2400 m, 2 August 2001, P. Zhang 94 (MHHNU 6594); same city, Nanhua County, Dazhongshan Forestry Farm, in mixed broadleaf-coniferous forest, altitude 2218 m, 9 August 2017, Jian W. Liu 787(MHHNU 11640). SICHUAN PROVINCE: Ganzi, Kangding City, Gongga Mountain, in Quercus semecarpifolia and Picea asperata forest, altitude 4000 m, 7 September 2021, P. Zhang 4336 (MHHNU 10836), P. Zhang 4338 (MHHNU 10838); same city, Sewulong Village, within a Quercus forests, altitude 3700 m, 3 September 2018, P. Zhang 3595 (MHHNU 10095); same location, 26 August 2017, P. Zhang 2776 (MHHNU 9276).

Commentary: Ramaria eryuanensis exhibits light pinkish cinnamon to light ochraceous cinnamon apices. According to the literature (Petersen, & Zang 1989), Kunming Institute of Botany holds two isotype specimens of R. eryuanensis: HKAS 20042 and HKAS 20044, with HKAS 20042 being designated as the isotype for R. eryuanensis. Upon examination of these specimens, we discovered that HKAS 20042 exhibits basidiospores dimensions of 14.5–17.0 × 4.5–6.5 µm, which are notably inconsistent with the literature. Consequently, it may be unsuitable for HKAS 20042 to continue as the isotype for R. eryuanensis. HKAS 20044, on the other hand, aligns with the characteristics detailed in the literature. However, this specimen is also as the isotype for R. linearioides, a species that owns verruculose basidiospores. Thus, it cannot be considered the isotype for R. linearioides.

53. Ramaria delicorosea P. Zhang & J. Zhong, sp. nov. Fig. 20 c−d and Fig. 80

MycoBank: MB 853006

Etymology: delicorosea = delico + rosea, proposed because this species has rose-pink colored apices of basidiomata and is considered delicious.

Diagnosis: basidiomata are off-white to cream color, with apices and young branches ranging from pink to red purple; color change upon handling is minimal, with a slow and slight transition to pale brown; basidiospores are ornamented with longitudinal stria; hyphae and basidia base exhibit clamps.

Type: CHINA, YUNNAN PROVINCE: Chuxiong, Mouding County, Huafo Mountain, altitude 2400 m, 23 August 2018, Zuo H. Chen 31322 (Holotype, MHHNU 31322, GenBank Acc. No.: ITS = PP467107, nrLSU = PP493439).

Fig. 80. Microscopic features of R. delicorosea (MHHNU 31322). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Description: Basidiomata 10 cm high and 10 cm broad, medium sized. Stipe 2 × 1 cm, single, falsely fasciculate, compound-connate, abortive branchlets occasional, off-white (1A1) to cream (4A3); nearly not changing color or slowly and slightly changing color to grayish yellow (2B5) when handle; flesh solid, off-white (1A1), flesh to fibrous, drying soft, easily penetrated. Major branches several, compound at base. Branches in 2–5 ranks, short and crowded; cream (4A3) to purplish red (14A8); axils narrowly rounded; internodes diminishing gradually at maturity. Apices acute, crowded, rose (12A4) to purplish red (14A8). Odor weakly fragrant; taste negligible.

Basidiospores [60/2/2]10.0–12.0(–13.0) × 4.0–6.0(–7.0) µm, L_m = 11.14 µm, l_m = 5.43 µm, Q = (1.57–)1.83–2.75(–2.88), Q_m = 2.29 ± 0.23, long-ellipsoid, conspicuously striae in profile, strongly cyanophilous; Basidia 45–65 × 7–13 µm, clavate, clamped at base; sterigmata mostly 4, 3–6 μm, stout and straight. Subhymenium rudimentary, hyphal. Hymenium unthickening. Tramal hyphae of stipe 6–14 µm wide, hyaline, conspicuously clamped, thin to thick-walled, interwoven, not agglutinated or adherent; ampulliform clamps up to 18 µm broad, usually not thick-walled; gloeoplerous hyphae not observed. Tramal hyphae of upper branches up to slightly narrowly, hyaline, thin walled, conspicuously clamped, not agglutinated or adherent, parallel and hardly branched inward; ampulliform clamps occasional; gloeoplerous hyphae not observed.

Ecology: Found in broad-leaved or forests or coniferous forests; basidiomata occurs in summer and autumn.

Distribution: Known from southwestern China (Yunnan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Chuxiong, Shuangbai County, 22 August 2018, Zuo H. Chen 31297 (MHHNU 31297); Nanhua County, bought from mushroom market, 9 August 2016, P. Zhang 2379 (MHHNU 8879), P. Zhang 2380 (MHHNU 8880), P. Zhang 2381 (MHHNU 8881), P. Zhang 2382 (MHHNU 8882), P. Zhang 2383 (MHHNU 8883), P. Zhang 2385 (MHHNU 8885).

Commentary: This species was only collected in Chuxiong, Yunnan, and is very similar to R. linkongshanensis L. Fan, Ying Li & N. Mao [11–13(–15) × 4.5–5 μm, Q = (2.2–)2.4–2.7, Q_m = 2.53 ± 0.16, Li et al. 2024; 12.97 × 4.73 μm, as measured in this study]. A subtle difference is that the basidiospores of this species are slightly smaller, and the apices of the branches exhibit a pink coloration that does not easily fade.

54. Ramaria hemirubella R.H. Petersen & M. Zang, Acta bot. Yunn. 8(3): 285, 1986. Fig. 20 e−f and Fig. 81

Description: Basidiomata 14 cm high and 8 cm broad, broadly fusiform to obpyriforme in outline. Stipe 5 × 3 cm, single, falsely fasciculate, to compound-connate, smooth, with a few abortive branchlets, off-white with a hint of pink upward, in age fleshly buff, slowly easily brown to red-brown where handled or chaffed, sometimes staining lemon yellow where handled; flesh solid, off-white, in age suffused flesh color, moist but not gelatinous or slippery, drying soft, light in weight, easily penetrated. Major branches several, more or less terete, ascending. Branches in 3-7 ranks, linear ascending, of two types: 1) typical, terete, pallid ochre, sometimes with hint of flesh upward; internodes diminishing gradually upward; Apices short digitate, suffused rose-pink in apical centimeter, deep red apically tan in age; and 2) grossly inflated, longitudinally rugulose, ruddy. Odor weakly fragrant; taste negligible.

Fig. 81. Basidiospores of R. hemirubella (MHHNU 31267). Scale bar: a = 10 µm.

Basidiospores [50/2/2] (8.0–)8.5–11.0 × (3.0–)4.0–5.5 µm, L_m = 9.47 µm, l_m = 4.94 µm, Q = (1.78–)1.80–2.26(–3.33), Q_m = 2.07 ± 0.29, ellipsoid, smooth to very obscurely undulate in profile; wall up to 0.2 µm thick; hilar appendix prominent, with pronounced throat; ornamentation of conspicuous to broken striae. Basidia 68−75 × 8−9 μm, clavate, clamped; weakly cyanophilous; sterigmata (2−) 4, slender, long, straight. Subhymenium rudimentary. Hymenium thickening. Tramal hyphae of stipe 4−12 µm wide, hyaline, conspicuously clamped, thin to thick-walled (wall up to 0.5 µm thick), easily broken, interwoven, not agglutinated or adherent; ampulliform clamps up to 15 µm broad, thick-walled (wall up to 1 µm thick), with conspicuous stalactiform ornamentation; gloeoplerous hyphae not observed. Tramal hyphae of upper branches up to 4-18 µm wide, hyaline, thick-walled (wall up to 1 µm locally), conspicuously clamped, not agglutinated or adherent, parallel and hardly branched inward, with cells cigar to barrel-shaped, outward more interwoven, gnarled, freely branched; ampulliform clamps common, not unusually thick-walled, unornamented; gloeoplerous hyphae not observed.

Ecology: Found in broad-leaved or forests or coniferous forests; basidiomata occurs in summer and autumn.

Distribution: Known from southwest China (Yunnan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Pu'er City, Simao District, 20 September 1983, Petersen 45673 (HKAS 10419); same place, 10 August 2001, altitude 1300 m, Zhu L. Yang 3181 (HKAS 38358); same location, 11 August 2001, altitude 1600 m, Zhu L. Yang 3183 (HKAS 38360); same city, Menglang Town, in a forest dominated by Pinaceae, altitude 980 m, 30 July 2017, P. Zhang 2630 (MHHNU 9130); Kunming, bought from Ciba mushroom Market, 2 August 2008, P. Zhang 702 (MHHNU 7202); Chuxiong, Nanhua County, Yulu Township, Sapiwu Village, with broad-leaved, coniferous or mixed forests, altitude 2000 m, 22 August 2018, Zuo H. Chen 31267 (MHHNU 31267).

Commentary: The isotype of this species (HKAS 10420) was found to have mold growth. Petersen first described R. hemirubella from a single specimen obtained at a mushroom market located in south-central Yunnan (Petersen & Zang 1986, 1989). Based on these additional collections, we can provide color notes and confirm that the apices are naturally deep red, not strained or secondarily suffused. Moreover, the branches constituted only a portion of one basidiomata, while the rest exhibited typical branches and apices.

Wu (1987) may classify this species as R. botrytis var. holorubella (Atk.) Corner. Petersen examined the type specimens of Clavaria holorubella and found the basidiospores to measure 12−15.6 × 3.5−5.0 µm. Wu reported measurements of Chinese material as 9.2−13 × 3−5.0 µm, which are closer to those of R. hemirubella. One collection (no. 47285) produced basidiospores substantially longer than those of others (10.8−13.7 × 3.6−4.7 µm, Q = 2.50−3.17, Q_m = 2.82; L_m =12.26 µm). Basidia of these basidiomata were predominantly 2-sterigmate, which may account for the larger basidiospores size. Although longer than those of other collections of R. hemirubella, they were still considered shorter than those of R. holorubella, and Petersen found little reason to emphasize the collection further.

55. Ramaria lingkongshanensis L. Fan, Ying Li & N. Mao, in Li, Mao, Zhang, Fu & Fan, Mycology: 10.1080/21501203.2024.2388910, 23 (2024) Fig. 20 g−h and Fig. 82

Description: Basidiomata 10 cm high and 10 cm broad. Stipe 4 × 3 cm, single, falsely fasciculate, compound-connate, abortive branchlets occasional, off-white to cream; nearly not changing color or slowly and slightly changing color to grayish yellow when handle; flesh solid, off-white, flesh to fibrous, drying soft, easily penetrated. Major branches several, compound at base. Branches in 2–5 ranks, short and crowded; cream to rose; axils narrowly rounded; internodes diminishing gradually at maturity. Apices acute, crowded, rose to pink. Odor weakly fragrant; taste negligible.

Fig. 82. Basidiospore features of R. lingkongshanensis (MHHNU 9873). Scale bar: a = 10 µm.

Basidiospores [60/2/2](11.4–)11.9–14.0(–14.3) × (3.9–)4.0–5.0(–5.8) µm, L_m = 12.97 µm, l_m = 4.73 µm, Q = (2.34–)2.40–3.42(–3.50), Q_m = 2.83 ± 0.31, long-ellipsoid, conspicuously striae in profile, strongly cyanophilous; Basidia 35–60 × 6–10 µm, clavate, clamped at base; sterigmata mostly 4, 3–6 μm, stout and straight. Subhymenium rudimentary, hyphal. Hymenium unthickening. Tramal hyphae of stipe 6–14 µm wide, hyaline, conspicuously clamped, thin to thick-walled, interwoven, not agglutinated or adherent; ampulliform clamps up to 18 µm broad, usually not thick-walled; gloeoplerous hyphae not observed. Tramal hyphae of upper branches up to slightly narrowly, hyaline, thin walled, conspicuously clamped, not agglutinated or adherent, parallel and hardly branched inward; ampulliform clamps occasional; gloeoplerous hyphae not observed.

Ecology: Found in broad-leaved or forests or coniferous forests; basidiomata occurs in summer and autumn.

Distribution: Known from southwestern and north China (Yunnan, Shanxi).

Additional materials examined: CHINA, YUNNAN PROVINCE: Chuxiong, Mouding County, Huafo Mountain, in a broad-leaved forest, altitude 2350 m, 21 August 2022, P. Zhang 4737 (MHHNU 11237); same location, altitude 2400 m, 23 August 2023, P. Zhang 5030 (MHHNU 11530); same location, 27 July 2021, Zuo H. Chen 32922 (MHHNU 32933); Dali, Dali City, Xianyangfeng, in a mixed forest, altitude 2800 m, 12 August 2018, P. Zhang 3373 (MHHU 9873), P. Zhang 3374 (MHHNU 9874), P. Zhang 3383 (MHHNU 9883); Kunming, bought from mushroom market, 29 September 2006, P. Zhang 574 (MHHNU 7074).

Commentary: Most species of Ramaria in ser. Hemirubellae are characterized by pink apices. However, R. lingkongshanensis is particularly notable within this series, not only for its pink apices but also for possessing the largest basidiospore, making it a distinctive species among them.

Ramaria ser. Laetae P. Zhang & J. Zhong, ser. nov.

MycoBank: MB 852734

Diagnosis: Differs from other series by its large basidiomata, dense branching, and yellow to bright orange coloration.

Type species: R. laeta P. Zhang & J. Zhong.

Description: 1) basidiomata pale orange to bright orange, big sized; 2) apices, branches and stipe concolorous with each other; 3) changing color to purple-brown on bruising or hurt; 4) branches and apices crowded; 5) basidia base and hyphae clamps or without clamps.

Commentary: This series includes five species collected exclusively in China. The basidiomata of these species range from yellow to orange, with or without clamps.

56. Ramaria aurantiapicalis P. Zhang & J. Zhong, sp. nov. Fig. 21 a–b and Fig. 83

MycoBank: MB 853249

Etymology: aurantiapicalis = aurantia + apicalis, referring to the yellow to orange color of the basidiomata with apices that turn brown when injured.

Diagnosis: Basidiomata exhibits a salmon to orange tone, with apices are yellow to orange and crowded, turning dark brown when injured. Basidia base and hyphae lack clamps.

Type: CHINA, SICHUAN PROVINCE: Ganzi, Kangding City, Sewulong Village, within a Picea forest, altitude 3450 m, 5 September 2018, P. Zhang 3599 (Holotype, MHHNU 10099, GenBank Acc. No.: ITS = PP467237).

Fig. 83. Microscopic features of R. aurantiapicalis (MHHNU 10099). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Description: Basidiomata 9 cm high and 11 cm broad, obtriangular to subcircular in outline. Stipe 4 × 3 cm, single to falsely fasciculate, tapering to narrowly rounded base, pruinose at base, longitudinally gnarled or rugose, with abortive branchlets, brunnescent when injury. flesh off-white (1A1), moist but not gelatinous or slippery. Major branches several, terete, off-white (1A1) at base, cream (4A3) to salmon (6A4) upward. Branches in 3−6 ranks, ascending, terete, salmon (6A4) to apricot yellow (5B6); flesh more pallid, brittle to brittle-fibrous; internodes diminishing gradually at maturity; axils narrowly rounded throughout. Apices short and crowded, double-dichotomous, yellowish orange (4A8) color, turned brown (6E8) when hurt. Odor and taste faintly fabaceous.

Basidiospores [60/2/2] (9.1–)9.4–11.6(–12.4) × (3.5–)3.7–5.1(–5.4) µm, L_m = 10.26 µm, l_m = 4.28 µm, Q = (1.97–)2.10–2.83(–2.99), Q_m = 2.45 ± 0.23, subcylindrical to ellipsoid, slightly roughened in profile; hilar not prominent; ornamentation of small, randomly scattered flat warts. Basidia 40−60 × 6−9 μm, clavate, without clamps; sterigmata 4, very slender, straight. Subhymenium extensive, pseudo-parenchymatous. Hymenium thickening. Tramal hyphae of stipe 4-13 μm wide, hyaline, without clamped, tightly interwoven, not agglutinated or adherent; ampulliform clamps up to 12 μm broad, not unusually thick-walled, with delicate ornamentation; gloeoplerous hyphae not observed. Tramal hyphae of upper branches up to 5-11 μm wide, hyaline, thin walled, without clamps, strictly parallel, more inflated inward, tightly packed, not agglutinated to adherent; ampulliform clamps up to 13 μm broad, gloeoplerous hyphae not observed.

Ecology: In Picea forests or broad-leaved forests. Basidiomata occurs in summer and autumn.

Distribution: Known from southwestern China (Yunnan, Sichuan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Deqin County, Baima Snow Mountain Nature Reserve, in Picea forest, altitude 3750 m, 17 August 2018, P. Zhang 3443 (MHHNU 9943). SICHUAN PROVINCE: Ganzi, Kangding City, Sewulong Village, within a Picea forest, altitude 3450 m, 5 September 2018, P. Zhang 3600 (MHHNU 10100); same city, Gongga Mountain, in Quercus semecarpifolia and Picea asperata forest, altitude 4000 m, 7 September 2021, P. Zhang 4366 (MHHNU 10866).

Commentary: Ramaria aurantiapicalis is similar to R. robustiflava and R. laeta in that they share salmon or yellow basidiomata and have similar-sized basidiospores. However, R. aurantiapicalis differs in that it develops dark brown apices when injured and lacks clamps, whereas the latter two species possess clamps.

57. Ramaria cadmioaurantiaca L. Fan, Ying Li & N. Mao, in Li, Mao, Zhang, Fu & Fan, Mycology: 10.1080/21501203.2024.2388910, 16 (2024)

Original description in Li et al. (2024).

58. Ramaria laeta (R.H. Petersen & M. Zang) P. Zhang & J. Zhong, stat nov. et comb. nov. Fig. 21 c−d and Fig. 84

≡ Ramaria rubricarnata var. laeta R.H. Petersen, in Petersen & Zang, Acta bot. Yunn. 11(4): 394 (1989)

MycoBank: MB 853237

Etymology: “laeta”, referring to the bright orange color of the basidiomata.

Diagnosis: Basidiomata are salmon to orange in tone and large-sized; apices are broadly acute and concolorous with branches; color changes to vinescent upon bruising or injury; branches and apices are crowded; basidia base and hyphae have clamp connection.

Isotype: CHINA, YUNNAN PROVINCE: Dali, Eryuan County, Sanying Township, brought from mushroom market, 27 August 1986, Petersen 47292.

Description: Basidiomata 14 cm high and 10 cm broad, obovate in outline. Stipe 6 × 4 cm, single to falsely fasciculate, stout, with abortive branchlets, minutely pruinose between substrate particles and between stipes, off-white, somewhat rugulose, brunnescent at base; flesh off-white, moist but not gelatinous or slippery. Major branches 3-5, flattened or terete, crowded, brunnescent where handled, concolorous with stipe downward, salmon to orange upward. Branches in 3−5 ranks, ascending, terete, salmon to salmon-orange; flesh more pallid, brittle to brittle-fibrous; internodes diminishing gradually at maturity; axils rounded throughout. Apices short and narrowly rounded, double-dichotomous, crowded, yellowish, minutely short digitate when mature, concolorous with branches. Odor and taste faintly fabaceous.

Fig. 84. Basidiospore features of R. laeta (MHHNU 33922). Scale bar: a = 10 µm.

Basidiospores [50/2/2] (8.8–)9.3–12.6(–13.0) × (3.8–)3.9–5.0(–5.2) µm, L_m = 10.66 µm, l_m = 4.33 µm, Q = (1.92–)2.05–2.75(–2.89), Q_m = 2.46 ± 0.22, ellipsoid to subcylindrical, flattened adaxially, roughened in profile; wall up to 0.2 μm thick; hilar prominent; ornamentation of plat-like wart and small warts randomly scattered. Basidia 40−60 × 6−9 μm, clavate, clamped; sterigmata 4, very slender, straight. Subhymenium extensive, pseudo-parenchymatous. Hymenium thickening. Tramal hyphae of stipe 4-12 μm wide, hyaline, conspicuously clamped, parallel, not agglutinated or adherent; ampulliform clamps up to 14 μm broad, not unusually thick-walled, with delicate ornamentation; gloeoplerous hyphae not observed. Tramal hyphae of upper branches up to 5-10 μm wide, hyaline, conspicuously clamped, thin-walled, parallel, not agglutinated to adherent; ampulliform clamps and gloeoplerous hyphae not observed.

Ecology: In the broad-leaved forests. Basidiomata occurs in summer and autumn.

Distribution: Known from southwestern China (Yunnan, Sichuan).

Additional materials examined: CHINA, SICHUAN PROVINCE: Ganzi, Seda County, in Picea and Abies forest, altitude 4000 m, 8 August 2023, Zuo H. Chen 33922 (MHHNU 33922); Kangding City, Mugecuo, in broad-leaved forest with Fagaceae, altitude 3500 m, 1 August 2012, Ping Zhang 1205 (MHHNU 7705); same city, Jiulong County, Tanggu Township, 18 July 2005, Zai W. 542 (HKAS49037).

Commentary: Ramaria laeta closely resembles R. robustiflava, as both species exhibit yellowish orange basidiomata, stout stipes, clamped hyphae and basidia, and a tendency to turn brown when bruised. The key distinction lies in their coloration: R. laeta leans towards an orange hue, while R. robustiflava tends towards a yellowish tint. Besides, R. laeta closely resembles R. xizangenesis (basidiospores 10.69 × 4.29 µm; Zhou et al. 2024), but the former exhibits yellowish-orange basidiomata, whereas the latter has pinkish-toned basidiomata.

59. Ramaria robustiflava P. Zhang & J. Zhong, sp. nov. Fig. 21 e−f and Fig. 85

MycoBank: MB 853246

Etymology: robustiflava = robust + flava, referring to the species having a stout stipe and yellow-colored basidiomata.

Diagnosis: Basidiomata are yellow with stout stipes, and the basidiospores measure 11.08 × 4.57 µm with warts.

Type: CHINA, SICHUAN PROVINCE: Ganzi, Kangding City, Muge Lake, in the Abies forest, altitude 3500 m, 2 September 2021, P. Zhang 3590 (Holotype, MHHNU 10890, GenBank Acc. No.: ITS = PP467234, nrLSU = PP493546, tef1-α = PP480413).

Fig. 85. Microscopic features of R. robustiflava (MHHNU 10090). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Description: Basidiomata 11 cm high and 10 cm broad, obovate in outline. Stipe 3 × 2 cm, single to falsely fasciculate, stout, with abortive branchlets, minutely pruinose between substrate particles and between stipes, off-white (1A1), brunnescent at base, weakly watery-brunnescent upward; flesh off-white (1A1), moist but not gelatinous or slippery. Major branches 3–5, flattened or terete, crowded, brunnescent or turned to lemon yellow (3B8) where handled, concolorous with stipe downward, yellowish orange (4A8) upward. Branches in 3–5 ranks, ascending, more or less terete, salmon (6A4) to yellowish orange (4A8); flesh more pallid, brittle to brittle-fibrous; internodes diminishing gradually at maturity; axils narrow rounded throughout. Apices short and broadly rounded when young, digitate when mature, double-dichotomous, concolorous with branches. Odor and taste faintly fabaceous.

Basidiospores [30/2/2] (10.0–)10.1–12.0(–12.9) × (3.8–)4.1–5.3(–5.4) µm, L_m = 11.08 µm, l_m = 4.57 µm, Q = (2.01–)2.10–2.83(–2.99), Q_m = 2.45 ± 0.23, ellipsoid, flattened adaxially, roughened in profile; wall up to 0.2 μm thick; hilar prominent; ornamentation of small and flat warts randomly scattered. Basidia 45–65 × 7–10 μm, clavate, clamped; sterigmata 4, stout, straight. Subhymenium extensive, double-dichotomous. Hymenium thickening. Tramal hyphae of stipe 3–14 μm wide, hyaline, conspicuously clamped, thick–walled (wall up to 1 μm thick), tightly interwoven, not agglutinated or adherent; ampulliform clamps up to 16 μm broad, not unusually thick-walled, with delicate ornamentation; gloeoplerous hyphae not observed. Tramal hyphae of upper branches up to 4-15 μm wide, hyaline, conspicuously clamped, thin-walled, parallel, not agglutinated to adherent; ampulliform clamps and gloeoplerous hyphae not observed.

Ecology: Solitary or gregarious on soil in forests dominated by Picea. Basidiomata occurs in summer and autumn.

Distribution: Known from southwestern China (Yunnan, Sichuan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Kunming, bought from mushroom market, 23 July 2017, P. Zhang 2598 (MHHNU 9098); Shangri-La, 8 September 2007, Feng Bang 113 (HKAS 52834);same city, Deqin County, Baima Snow Mountain Nature Reserve, altitude 3750 m, 17 August 2018, P. Zhang 3431 (MHHNU 9931), P. Zhang 3432 (MHHNU 9932), P. Zhang 3448 (MHHNU 9948); same location, altitude 3750 m, 18 August 2018, P. Zhang 3469 (MHHNU 9969).

Commentary: Dr. Zhang (2010) classified this species as R. rubricarnata var. laeta R.H. Petersen. Initially described by Marr & D.E. Stuntz, R. rubricarnata has been collected in North America, exhibiting four variants. Ramaria rubricarnata var. verna R.H. Petersen & Scates and R. rubricarnata var. pallida R.H. Petersen & Scates are both spring-fruiting species found in western North America. Ramaria rubricarnata Marr & D.E. Stuntz and R. rubricarnata var. rubricarnata were collected by Marr & D.E. Stuntz in October to November, characterized by basidiospores less than 10 µm in length, pallid fleshy-yellow branches, yellow apices, and salmon colored flesh.

Petersen (1989) described the variant R. rubricarnata var. laeta in Yunnan, China, noting its shorter basidiospores (L_m = 10.17 µm), salmon-orange branches, pastel gold to bright orange-yellow apices, brunnescent when bruised, and absence of abortive branchlets, collected in Pines yunnanensis forests.

Further observations and molecular identifications of additional specimens have revealed the presence of abortive branchlets, and slightly longer basidiospores compared to R. rubricarnata var. laeta (L_m = 10.17 µm; Petersen & Watling 1989). These differences, including collections from Picea forests with salmon branches and bright orange-yellow apices, warrant the classification of R. robustiflava as a distinct taxon.

60. Ramaria xizangensis C.L. Zhao & H.M. Zhou, in Zhou, Zhang, Li, Wu & Zhao, MycoKeys 106: 216 (2024)

Original description in Zhao et al. (2024).

Ramaria ser. Laeviformosoides P. Zhang & J. Zhong, ser. nov.

MycoBank: MB 852710

Etymology: “laevi” refers to smooth basidiospores, while “formosoides” denotes similarity to R. formosa.

Type species: R. laeviformosoides R.H. Petersen & M. Zang, Acta bot. Yunn. 11(4): 384, 1989.

Diagnosis: Basidiomata displays weak salmon and yellow coloration; basidia are clamped and non-cyanophilous; basidiospores are smooth and weakly amyloid stipe flesh.

Commentary: A potential basonym, Ramaria sect. Gypseae Franchi & M. Marchetti, shares characteristics such as smooth basidiospores and predominantly white basidiomata rather than yellow. However, the type species of this section, R. gypseae, collected by Schild in 1982, has not been rediscovered since.

In China, two species were collected in this series. Ramaria mediterranea Schild & Franchi, which also features white basidiomata and purple apices in this series (Schild 1998; Franchi & Marchetti 2021). Ramaria gypsea Schild (1982) may also belong to this Series, characterized by white basidiomata and basidiospores measuring 10.8−14 × 3.7−4.5 μm with clamps. The type specimen originated from Italy but has not been rediscovered since Schild’s collection in 1982.

61. Ramaria laeviflava P. Zhang & J. Zhong, sp. nov. Fig. 22 a–b and Fig. 86

MycoBank: MB 853227

Etymology: laeviflava = laevi + flava, referring to the species having nearly smooth basidiospores and yellow basidiomata.

Diagnosis: Basidiomata are pallid salmon to yellow in color; basidia and hyphae exhibit clamps; basidiospores are nearly smooth; darkens to dark brown when bruised or handled.

Type: CHINA, YUNNAN PROVINCE: Shangri-La, Pudacuo, Shudu Lake, in the forest with Quercus semecarpifolia, altitude 3600 m, 15 August 2018, P. Zhang 3409 (Holotype, MHHNU 9909, GenBank Acc. No.: ITS = PP467285, nrLSU = PP493587).

Description: Basidiomata 10 cm high and 11 cm broad, medium sized, obtriangular in outline. Stipe 4 × 1.5 cm, single to falsely fasciculate, tapering gradually downward, smooth, off-white (1A1) to pastel yellow (2A4), without abortive branchlets; flesh solid, off-white (1A1) to yellowish white (2A2), moist, but not gelatinous or slippery. Major branches several, stout and short, terete to flattened, ascending, concolorous with stipe downward, concolorous with branches upward, weakly brunnescent. Branches in 3−4 ranks, dichotomous, terete, ascending, yellowish white (2A2) to cream (4A3); flesh paler than hymenium, soft; internodes length ratio diminishing gradually at maturity; axils mostly acute, both U-shaped and V-shaped. Apices crowded, cuspidate, the ends of the major branches are almost in the same plane, pastel yellow (2A4) to yellowish orange (4A8), concolorous with the branches in age; Odor faintly penetrating; taste negligible.

Fig. 86. Microscopic features of R. laeviflava (MHHNU 9909). a basidiospores; b basidia. Scale bars: a =10 µm, b =20 µm.

Basidiospores [62/2/2] (7.0–)7.5–11.8(–12.3) × (3.1–)3.3–5.1(–5.4) µm, L_m = 9.56 µm, l_m = 4.11 µm, Q = (1.67–)1.91–2.71(–2.94), Q_m = 2.33 ± 0.26, ellipsoid, nearly smooth in profile; hilar appendix prominent; ornamentation none or finely and small warts randomly placed. Basidia 35-50 × 5-11 µm, clavate, clamped; sterigmata 4, long, spindly, straight. Subhymenium rudimentary, pseudoparenchymatous. Hymenium unthickening. Tramal hyphae of stipe 4-15 µm wide, usually inflated, hyaline, thin- to thick walled, loosely parallel, interwoven, not agglutinated or adherent; ampulliform clamps up to 15 μm broad, thick-walled, with coarse ornamentation; gloeoplerous hyphae not observed; Tramal hyphae of upper branches up to 3-14 µm wide, hyaline, thin-to thick-walled, clamped, secondarily septum, loosely parallel, not agglutinated or adherent; ampulliform clamps up to 15 µm broad, thick-walled, unornamented; gloeoplerous hyphae not observed.

Ecology: Found in the forests with Quercus semecarpifolia; basidiomata occurs in summer and autumn.

Distribution: Known from southwest China (Yunnan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Shangri-La, Pudacuo, Shudu Lake, in the forest with Quercus semecarpifolia, altitude 3600 m, 15 August 2018, P. Zhang 3406 (MHHNU 9906), P. Zhang 3407 (MHHNU 9907), P. Zhang 3410 (MHHNU 9910), P. Zhang 3411 (MHHNU 9911).

Commentary: Ramaria laeviflava bears a striking resemblance to R. laeviformosoides. Both species share similar-sized and nearly smooth basidiospores, clamped basidia and hyphae, and a comparable pale salmon to yellow basidiomata that darkens when bruised or handled. However, R. laeviflava is typically found in Picea forests, whereas R. laeviformosoides occurs in Quercus forests. In addition, R. laeviformosoides tends to have a relatively paler color (cream with a hint of salmon), while R. laeviflavahas exhibits a more yellowish hue. Ramaria laeviflava is phylogenetically close to R. mediterranea, but the latter possesses smaller basidiospores (7.7 × 4.2 µm; Franchi & Marchetti 2021).

62. Ramaria laeviformosoides R.H. Petersen & M. Zang, Acta bot. Yunn. 11(4): 384, 1989. Fig. 22 c–d and Fig. 87

Description: Basidiomata 11 cm high and 7 cm broad, small-sized, circular to broadly obovate in outline. Stipe 6 × 5 cm, single to falsely fasciculate, stout, tapering gradually downward, smooth upward, tomentose at base, with abortive branchlets, ridged to appear fasciculate, pruinose at base, smooth upward, with no abortive branchlets; white downward, weakly brunnescent; flesh white, solid, moist, but not gelatinous or slippery, soft, drying firm but easily penetrated, not friable, yellow downward. Major branches several, ascending, terete, concolorous with stipe downward, concolorous with branches upward, weakly brunnescent. Branches in 3−6 ranks, terete, strict, pallid flesh to pink colored, fading by maturity to cream; flesh paler than hymenium, soft; internodes all short when young, elongating from below, diminishing gradually upward at maturity; axils rounded throughout. Apices coarse, acute, divaricate when young, stiffly digitate at maturity, light ochraceous color when young, concolorous with the branches in age; Odor faintly penetrating; taste negligible.

Fig. 87. Basidiospore features of R. laeviformosoides (MHHNU 8599). Scale bar: a = 10 µm.

Basidiospores [60/2/2]7.0−9.5 ×3.5−4.2(−4.5) μm, L_m = 8.56 µm, l_m = 3.89 µm, Q = (1.75−)1.84–2.57, Q_m = 2.19 ± 0.23, cylindrical, smooth, or extremely obscurely undulate in profile; wall up to 0.2 µm thick; hilar appendix small, not prominent, truncate; ornamentation none. Basidia 60−70 × 9−10 μm, clavate, clamped; sterigmata (2−) 4, short, spindly, straight. Subhymenium rudimentary, pseudoparenchymatous. Hymenium thickening. Tramal hyphae of stipe 3−23 µm wide, usually inflated, hyaline, thin- to occasionally thick walled (wall up to 1 µm thick locally), interwoven, tightly packed, not agglutinated or adherent; ampulliform clamps up to 16 μm broad, thick-walled (wall up to 1.5 µm thick), with coarse ornamentation; gloeoplerous hyphae rare, as clavate cell tips, undelimited at narrow end, yellow-refringent; Tramal hyphae of upper branches up to 3−12 µm wide, hyaline, thin-to thick-walled (wall up to 0.5 µm thick locally), clamped, secondarily septum, strictly parallel, not agglutinated or adherent; ampulliform clamps up to 14 µm broad, thick-walled (wall up to 1 µm thick), unornamented; gloeoplerous hyphae occasional, 2−3.5 μm, refringent.

Ecology: Found in Picea forests; basidiomata occurs in summer and autumn.

Distribution: Known from southwest China (Yunnan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Lijiang, Lijiang mushroom market, 26 August 2008, Qi Zhao 8319 (HKAS 55281); Chuxiong, 17 August 2015, P. Zhang 2099 (MHHNU 8599).

Commentary: Diagnostic characters for the species include the following: 1) pale salmon and yellow coloration; 2) clamped basidia; 3) smooth basidiospores; and 4) weakly amyloid stipe flesh.

The isotype designation (HKAS 17744) in Petersen's publication is erroneous, as the species exhibits warty basidiospores, which contradicts the smooth basidiospores that were described (Petersen & Zang 1989). Zhang's perspective is accurate because R. laeviformosoides stands out as the only smooth-spored Ramaria species lacking predominant yellow or orange coloration (Zhang 2010). It bears a resemblance to R. formosa in stature, browning tendency, and overall coloration. Notably, both species share characteristics such as soft stipe flesh when fresh and dry, and clamped basidia. However, they differ significantly in basidiospores morphology—R. laeviformosoides basidiospores are smooth without ornamentation, whereas R. formosa basidiospores are conspicuously roughened and coarsely ornamented. Additionally, the basidiomata of R. laeviformosoides exhibit more muted colors compared to R. formosa. Ramaria laeviformosoides is phylogenetically close to R. mediterranea, but the latter possesses smaller basidiospores (7.7 × 4.2 µm; Franchi & Marchetti 2021).

Ramaria ser. Leiosporae P. Zhang & J. Zhong, ser. nov.

MycoBank: MB 852709

Etymology: Leiosporae = "Leio" + "sporae", "leios" (λεῖος), meaning "smooth".

Type species: Ramaria schildii R.H. Petersen, Mycologia 80(2): 229 (1988).

Diagnosis: Basidiomata displays weak salmon and yellow coloration; basidia are clamped; basidiospores are smooth with electron microscope.

Commentary: In China, only one species has been collected in this series.

63. Ramaria schildii R. H. Petersen, Mycologia 80(2): 229 (1988). Fig. 23 a–b and Fig. 88

Description: Basidiomata 10−18 cm high, up to 5−10 cm broad, superficially obconical to trumpet-shaped in outline, but old basidiomata becoming lax and assuming semi-grotesque position. Stipe 2 × 3−5 cm, rooting to some extent, fasciculate or with a very small point of union at the base, or solitary, minutely tomentose and off-white below substrate, creamy above, becoming darker to tannish-ochre in age; surface obviously weakly brunnescent where handled; flesh white, fibrous-fleshy, moist but not slippery or gelatinous, slowly weakly brunnescent. Major branches few per fascicle, often terete or flattened, light yellow (with a hint of ochre) when young, fading to pale fleshy tan in age; flesh yellowish near hymenium; surface vinescent around dirt specks and flesh slowly so when ripped; axils narrowly rounded to acute when young, often lunate in age; internodes diminishing gradually upward, giving an open, lax appearance. Apices short and straight, lemon yellow when young, becoming divaricate and lobed in age, and then fading through more ochre to finally concolorous with branches. Taste and odor not recorded.

Fig. 88. Basidiospores of R. schildii (MHHNU 9285). Scale bar: a = 10 µm.

Basidiospores [60/2/2](8.8−)9.3−14.0(−14.9) × (3.3−) 3.4−5.1(−5.8) μm, L_m = 12.09 μm, l_m = 4.16 μm, Q = (2.2−)2.4−3.4(−3.5), Q_m = 2.85 ± 0.29, cylindrical, smooth in profile; wall up to 0.2 μm thick; hilar appendix truncates, somewhat prominent, abrupt. Basidia 40−65 × 9−12 μm, clavate, clamped; sterigmata 4, long, stout, slightly divergent. Hymenium thickening. Tramal hyphae of stipe hyaline, thin walled, clamped, weakly cyanophilous, of two types: 1) 3−13 μm wide, parallel, somewhat inflated; and 2) 1.6−2.0 μm wide, tortuous, often with abundant side branches arising at almost right angles and usually ending as abortive pegs or short branches, infrequently clamped, locally densely interwoven with more inflated hyphae. Stipe surface hyphae identical to predominant flesh hyphae. Tramal hyphae of upper branches 3−10 μm wide, thin-walled, parallel, clamped, hyaline, free.

Ecology: Solitary or gregarious in forests with Picea or Abies.

Distribution: Known from southwestern China (Sichuan); Italy and Sweden in Europe (Petersen 1988a).

Additional materials examined: CHINA, SICHUAN PROVINCE: Ganzi, Kangding City, Mugecuo, in a Picea asperata forest, altitude 3500 m, 21 August 2017, P. Zhang 2702 (MHHNU 9202); same city, Tagong Township, in a mixed forest of Abies and Picea, altitude of 3900m, 22 August 2017, P. Zhang 2727 (MHHNU 9227) and P. Zhang 2730 (MHHNU 9230); same city, Waze Township, in a mixed forest of Abies and Picea, altitude of 4000 m, 23 August 2017, P. Zhang 2742 (MHHNU 9242); same city, Gongga Township, in mixed forests dominated by Picea asperata and Betula platyphylla, altitute 3700 m, 27 August 2017, P. Zhang 2785 (MHHNU 9285). XIZANG AUTONOMOUS REGION: Changdu City, Leiwuqi County, August 2004, Zhu L. Yang 4378 (HKAS 45735).

Commentary: The species is the only one in Ramaria of China with smooth basidiospores.

Ramaria ser. Lineariflavae P. Zhang & J. Zhong, ser. nov.

MycoBank: MB 852771

Etymology: linearflava = linear + flava, indicating that this species has slender branches and yellowish fruitbodies.

Type: R. linearflava P. Zhang & J. Zhong.

Diagnosis: Basidiomata displays yellow coloration; branches slender; basidia are clamped; basidiospores are small and warty.

Commentary: This series was collected in China and comprises one species, the other species is R. flavosalmonicolor Schild.

64. Ramaria lineariflava P. Zhang & J. Zhong, sp. nov. Fig. 24 a–b and Fig. 89

MycoBank: MB 853206

Etymology: lineariflava = linear + flava, indicating that this species has slender branches and yellowish basidiomata.

Diagnosis: Basidiomata are pale orange; stipe ranges from single to fasciculate; branches are slender; basidia are clamped; basidiospores are coarsely ornamented.

Type: CHINA, HUNAN PROVINCE: Chenzhou City, Yizhang County, Mangshan National Nature Reserve, altitude 1400 m. 5 September 2007, P. Zhang 662 (Holotype, MHHNU 7162, GenBank Acc. No.: ITS = PP467263, nrLSU = PP493569, tef1-α = PP480433).

Fig. 89. Microscopic features of R. lineariflava (MHHNU 7162). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Description: Basidiomata 14 cm high and 7 cm broad, repeatedly branched, usually obpyriforme in outline. Stipe 2 × 1.5 cm, single to falsely fasciculate, slender, tapering gradually downward, tomentose at base, smooth upward, without abortive branches, off-white (1A1) to cream (2A2), not changing color on bruising; flesh white, solid, moist but not gelatinous or slippery. Major branches 2−6, terete, up to 0.5 cm thick, ascending, concolorous to branches above. Branches in 4−6 ranks, ascending, polychotomous, slender, crowded; pale yellow (2A2–3), cream (2A2), pale orange (5A2–3); internodes diminishing gradually upwards at maturity; axils mostly acute (U-shaped). Apices narrowly rounded, concolorous with the branches. Odor negligible or faintly fresh; taste negligible.

Basidiospores [85/3/3] (6.0–)7.0–7.5(–9.0) × 3.0–4.0 µm, L_m = 7.56 µm, l_m = 3.40 µm, Q = 1.75–2.67, Q_m = 2.18 ± 0.27, thick-walled, cylindrical to ellipsoid, more or less flattened adaxially, conspicuously roughened in profile; wall up to 0.5 μm thick, cyanophilous; hilar appendix prominent; ornamentation of prominent, discrete, low warts and short ridges randomly placed. Basidia 32−54 × 7−9 μm, clavate, thin-walled, hyaline, clamped at base, 4-spored; sterigmata 6−7 μm long. Subhymenium rudimentary. Hymenium unthickening. Tramal hyphae of stipe 3−15 μm wide, hyaline, thin-walled, clamped, interwoven; ampulliform clamps up to 14 μm broad, gloeoplerous hyphae not observed. Tramal hyphae of upper branches 3−10 μm wide, hyaline, thin-walled, interwoven, loosely parallel, clamped; gloeoplerous hyphae not observed.

Ecology: Solitary or gregarious on soil in broad-leaved forests with Fagaceae or mixed forests dominated by Fagaceae, Oak and Pinaceae; basidiomata occurs in summer and autumn.

Distribution: Known from southwest, northeast, and central China (Yunnan, Jilin, Hunan, Hubei).

Additional materials examined: CHINA, JILIN PROVINCE: Tonghua County, Baijifeng Forest Park, in a temperate forest, altitude 800 m, 28 Aug. 2020, P. Zhang 4043 (MHHNU 10543). HUNAN PROVINCE: Chenzhou City, Yizhang County, Mangshan National Nature Reserve, altitude 1400 m. 3 September 2007, P. Zhang 645 (MHHNU 7145); same location, 27 July 2018, P. Zhang 3247 (MHHNU 9747); HUBEI PROVINCE: Yichang County, Shennongjia, in a mixed forests of broad-leaved and coniferous, altitude 1100 m, 29 July 2006, P. Zhang 554 (MHHNU 7054). YUNNAN PROVINCE: Lijiang City, Yulong Snow Mountain, 7 September 1986, Petersen 47307 (HKAS 20046).

Commentary: The isotype species sequences of HKAS 20046, which Petersen identified as R. hilaris (Petersen & Zang 1989), actually cluster with R. lineariflava in our phylogenetic analyses. Examination of the HKAS 20046 specimen revealed an average basidiospore size of 7.3 × 3.4 µm and without clamps, which is inconsistent with Petersen’s description of R. hilaris. This confirms that HKAS 20046 represents R. lineariflava rather than R. hilaris. Moreover, R. lineariflava is characterized by smaller basidiospores than those of true R. hilaris (9.7 × 4.75 µm, Petersen & Zang 1989) and exhibits clamps, which are absent in R. hilaris.

R. sandaracina (hototype from U.S.A.) could be mistaken for R. lineariflava. All have orange branches with clamped basidia, but R. lineariflava have stouter stipe and R. sandaracina have more darker orange color (Marr & Stuntz 1973).

Ramaria ser. Longisporae P. Zhang & J. Zhong, ser. nov.

MycoBank: MB 852773

Etymology: “longispora” refers to long basidiospores.

Lectotype: R. longispora Marr & D.E. Stuntz, Biblthca Mycol. 38: 101.

Diagnosis: 1) flesh fleshy to fibrous; 2) Basidiomata are yellow to bright orange, without brown or red-brown patches; 3) long basidiospores; 4) long basidia and without clamps.

Commentary: One species of this series was collected in China. The other species, R. longispora Marr & D.E. Stuntz are found in North American, R. atractospora Franchi & M. Marchetti and R. rielii Boudhowever are found in Europe. All species within this series possess bright yellow color fruiting bodies and exhibit long basidiospores.

65. Ramaria pseudolongispora P. Zhang & J. Zhong, sp. nov. Fig. 25 a–b and Fig. 90

MycoBank: MB 853222

Etymology: “pseudolongispora” refers to the species having long basidiospores and being similar to R. longispora.

Diagnosis: Stipe context ranges from fleshy to fibrous; basidiomata are yellow to bright orange, lacking brown to red-brown stains; basidiospores are long; basidia are long and feature clamps.

Type: CHINA, YUNNAN PROVINCE: Shangri-La, Pudacuo, in the forest with Quercus semecarpifolia, altitude 3400 m, 14 August 2018, P. Zhang 3388 (Holotype, MHHNU 9888, GenBank Acc. No.: ITS = PP467183, nrLSU = PP493504, tef1-α = PP480388).

Fig. 90. Microscopic features of R. pseudolongispora (MHHNU 10059). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Description: Basidiomata 11–12 cm high and 4–6 cm broad, small to medium sized. Stipes up to 0.5–1 × 3–5 cm, single, or more commonly sub-compound, consisting of 2–6 primary axes arising from a primary root-like structure; off-white (1A1) at base, surface pale yellow (2A3) to sunflower yellow (4A7); not changing color on bruising; surface smooth or rugose, hardly erect; without abortive branches; context of stipe pale orange (5A3) when fresh, fleshy to fibrous, not slippery or soapy to the touch. Major branches 2–6, polychotomous, erect, more or less terete; Branches in 3–6 ranks, commonly dichotomous; pale orange (5A3) to melon yellow (5A6) by maturity; axils narrowly rounded, mostly V-shaped; internodes length ratio diminishing gradually at maturity, distinctly elongated at maturity; context of main branches and upper branches fleshy to fibrous, pale orange (5A3), not slippery; Apices generally subacute, sometimes swollen, dichotomous, vivid yellow (2A8) to yellowish orange (4A8), concolorous with branches at maturity. Odor negligible; taste weak, pleasant.

Basidiospores [60/2/2] (12.1–)13.1–16.9(–18.1) × (4.2–)4.5–6.1(–7.3) µm, L_m = 14.95 µm, l_m = 5.19 µm, Q = (2.26–)2.46–3.27(–3.94), Q_m = 2.89 ± 0.29, cylindrical to long ellipsoid; roughened in profile; ornamentation of numerous, distinct, cyanophilous warts. hilar appendix moderate; Basidia 60–80 × 8–12 μm, long, clavate, without basal clamps; sterigmata 4, 4–7µm long, straight. Hymenium unthickening, 75–90 µm; Subhymenium rudimentary, 10–25 µm. Contextual hyphae: Tramal hyphae of stipe 3–12 μm wide, loosely parallel, not adherent; inflated clamps up to 15 µm broad, with coarse ornamentation; tramal hyphae of branches central hyphae slightly narrow, inflated clamps up to 15 µm broad, hyaline, thin-walled, unornamented; without clamps; gloeoplerous hyphae not observed.

Ecology: Solitary or gregarious on soil in broad-leaved or mixed forests. Basidiomata occurs in summer and autumn.

Distribution: Known from southwest China (Xizang, Yunnan, Sichuan).

Additional materials examined: CHINA, XIZANG AUTONOMOUS REGION: Linzhi, Sejila Mountain, in a subalpine forest dominated by Picea, altitude 3300 m, 27 August 2018, P. Zhang 3527 (MHHNU 10027), P. Zhang 3534 (MHHNU 10034), P. Zhang 3537 (MHHNU 10037), P. Zhang 3549 (MHHNU 10049), P. Zhang 3550 (MHHNU 10050), P. Zhang 3553 (MHHNU 10053), P. Zhang 3554 (MHHNU 10054), P. Zhang 3554 (MHHNU 10059). YUNNAN PROVINCE: Shangri-La, Pudacuo, in the forest with Quercus semecarpifolia, altitude 3400 m, 14 August 2018, P. Zhang 3389 (MHHNU 9889). SICHUAN PROVINCE: Ganzi, Kangding City, Waze Township, in a mixed forest of Abies and Picea, altitude of 4000 m, 23 August 2017, P. Zhang 2741 (MHHNU 9241).

Commentary: In the field, this species is often mistaken for R. largentii, R. sandaracina, R. gelatiniaurantia, R. aurantiisiccescens due to their similar coloration. However, the first two species can be distinguished by their clamped hyphae, while the latter two differ in various aspects, including basidiospores dimensions.

Additionally, species like R. longispora from North America, R. rielii Boud and R. atractospora Franchi & M. Marchetti from Italy share a similar yellowish basidiomata and elongated basidiospores. Distinguishing factor lies in the basidiospores morphology: R. rielii has slightly narrower and longer (13.5–18.0 × 3.5–5.0 µm, L_m = 14.9 µm, l_m = 4.4 µm, Q = 2.7–4.1, Q_m = 3.4, Franchi & Marchetti 2021), whereas R. longispora and R. atractospora have slightly smaller basidiospores in size (Marr & Stuntz 1973; Franchi & Marchetti 2021).

Ramaria ser. Luteae (Franchi & M. Marchetti) P. Zhang & J. Zhong stat. nov.

Synonym: Ramaria sect. Luteae Franchi & M. Marchetti, Fungi Non Delineati, Raro vel Haud Perspecte et Explorate Descripti aut Definite Picti 16: 32 (2001)

Diagnosis: Basidiomata terrestrial, cream to yellow, with more brightly colored apices or branches that are concolorous; clamps absent, basidiospores pale ochraceous, more or less verrucose, longitudinally striated or smooth.

Type species: Ramaria lutea (Vitt.) Schild, Persoonia Vol. 9 (3): 409, 1977.

Description: This series includes one species collected in China.

66. Ramaria sanguinixantha P. Zhang & J. Zhong, sp. nov. Fig. 26 a−b and Fig. 91

MycoBank: MB 853234

Etymology: sanguinixantha = "sanguini-" + "-xantha", referring to the yellow basidiomata that turn blood-red when injured.

Diagnosis: Basidiomata exhibits a salmon to yellow hue with apices that are yellow or concolorous with the branches, turning to blood-red when injured, and the base of basidia and hyphae lack clamps.

Type: CHINA, SICHUAN PROVINCE: Ganzi, Kangding City, Gongga Mountain, in Quercus semecarpifolia and Picea asperata forest, altitude 4000 m, 7 September 2021, P. Zhang 4342 (Holotype, MHHNU 10842, GenBank Acc. No.: ITS = PP467240, nrLSU = PP493550, tef1-α = PP480416).

Fig. 91. Microscopic features of R. sanguinixantha (MHHNU 10842). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Description: Basidiomata 11 cm high and 15 cm broad, obtriangular to usually subcircular in outline. Stipe 4 × 4 cm, single to falsely fasciculate, tapering to narrowly abounded base, smooth, with occasional abortive branchlets, brunnescent when injury; flesh off-white (1A1), moist but not gelatinous or slippery. Major branches several, terete, pale yellow (2A3) to salmon (6A4). Branches in 2−6 ranks, ascending, terete, pale yellow (2A3) to salmon (6A4); flesh more pallid, brittle to brittle-fibrous, gelatinous; internodes diminishing gradually at maturity; axils narrowly rounded throughout. Apices short and crowded, double-dichotomous, pale yellow (2A3), turned to wine red (11D8) when injury. Odor and taste faintly fabaceous.

Basidiospores [60/2/2] (8.1–)8.7–11.2(–11.9) × (3.4–)4.0–5.2(–5.9) µm, L_m = 9.97 µm, l_m = 4.50 µm, Q = (1.82–)2.00–2.56(–2.87), Q_m = 2.23 ± 0.19, subcylindrical to ellipsoid, slightly roughened in profile; hilar not prominent; ornamentation of small, randomly scattered flat warts. Basidia 30−60 × 5−10 μm, clavate, without clamps; sterigmata 4, stout and straight. Subhymenium extensive, hyphae. Hymenium unthickening. Tramal hyphae of stipe 5-13 μm wide, hyaline, without clamps, tightly interwoven, not agglutinated or adherent; ampulliform clamps up to 16 μm broad, not unusually thick-walled, with delicate ornamentation; gloeoplerous hyphae not observed. Tramal hyphae of upper branches up to 4-15 μm wide, hyaline, thin walled, without clamped, loosely parallel, more inflated inward, agglutinated to adherent; ampulliform clamps up to 16 μm broad, gloeoplerous hyphae occasionally.

Ecology: In Quercus or Picea forests or broad-leaved forests. Basidiomata occurs in summer and autumn.

Distribution: Known from southwestern China (Sichuan, Xizang, Yunnan).

Additional materials examined: CHINA, SICHUAN PROVINCE: Ganzi, Kangding City, Gongga Mountain, in Quercus semecarpifolia and Picea asperata forest, altitude 4000 m, 7 September 2021, P. Zhang 4363 (MHHNU 10863); same location, altitude 3900 m, 6 September 2021, P. Zhang 4317 (MHHNU 10817); same city, Sewulong Village, within a Quercus forest, altitude 3700 m, 3 September 2018, P. Zhang 3594 (MHHNU 10094); same city, Gongga Township, Sewurong II Village, altitude 3700 m, 26 August 2017, P. Zhang 2772 (MHHNU 9272), P. Zhang 2773 (MHHNU 9273); same city, Pengta Township, altitude 3100 m, 19 August 2017, P. Zhang 2678 (MHHNU 9178). XIZANG AUTONOMOUS REGION: Linzhi, Sejila Mountain, in a subalpine forest dominated by Picea, altitude 3300 m, 27 August 2018, P. Zhang 3548 (MHHNU 10048), 3561 (MHHNU 10061); same location, altitude 3400 m, 26 August 2018, P. Zhang 3524 (MHHNU 10024). YUNNAN PROVINCE: Shangri-La, Pudacuo, Shudu Lake, in the mixed forest with Quercus and Picea. altitude 3429 m, 10 August 2020, Jian W. Liu MF367 (MHHNU 11647); same location, in the forest with Quercus semecarpifolia, altitude 3600 m, 15 August 2018, P. Zhang 3416 (MHHNU 9916) and P. Zhang 3424 (MHHNU 9924).

Commentary: Reddish-brown staining species are distributed worldwide, with a total of 11 species reported. Among these, five species possess clamps (R. cystidiophora var. maculans, R. maculatipes, R. vinosimaculans, R. eosanguinea, R. sanguinipes), and six species without clamps (R. rubribrunnescentiflava, R. synaptopoda, R. rubiginosa, R. sanguinea, R. rubriattenuipes, R. xanthosperma). The main characteristics of each species are as follows.

Among the species without clamps, R. rubribrunnescentiflava, R. rubriattenuipes and R. xanthosperma have basidiospores sizes greater than 10 µm, while R. sanguinea and other two species have basidiospores sizes smaller than 10 µm.

Compared to the other two species with similar basidiospores sizes, R. synaptopoda is distinguished by its a pronounced fasciculate habit and small basidiomata, resembling R. conjunctipes.

Marr & Stuntz questioned the application of R. sanguinea (Pers.) Quél. Coker identified his specimens based on a colored illustration and Persoon’s original description, and according to Corner, “the exact identity of Clavaria sanguinea Per. is debatable”. In 1968, Marr considered the collection now known as R. rubiginosa, to be R. sanguinea as described by Coker (1923). Since then, Marr examined the collections that formed the basis of Coker’s description and found differences: Marr's specimens were more robust with broader stipes, and larger, more finely ornamented basidiospores, In 1966, Corner reclassified R. sanguinea (formerly C. sanguinea) as a variety of R. flava (Schaeff.) Quél., designating Coker No. 2656 (NCU) as the type specimen. To clarify whether R. sanguinea should be understood in Persoon’s sense, a type specimen from Persoon’s collections needs identification or a neotype must be designated from the type locality. Based on color notes from Doty (1944) and examination of two collections he cited, Marr concluded that C. sanguinea sensu differs from R. rubiginosa.

When comparing the eastern R. xanthosperma (Peck) Corner with the western R. rubiginosa and R. sanguinea, Petersen (1976) observed that R. rubiginosa “shows a generally brighter yellow coloration in the lower branches, stouter fruit bodies, positive reaction of branch trama with iron salts, and slightly smaller basidiospores.” He further commented that if this combination of characteristics did not sufficiently distinguish the taxon at the species level, “R. rubiginosa would appear as a synonym under R. sanguinea.”

For comparisons between R. rubiginosa and R. sanguinixantha, the former collected in the western Hemlock Forest in Washington, USA, while the latter was collected in Quercus, Picea, or broad-leaved forests in the highlands of China.

In comparing R. sanguinea descripted by Franchi & Marchetti with R. sanguinixantha, our two-gene phylogenetic analysis indicates that are similar but not identical species. Morphologically, the species found in Italy displays a bright yellow color, with bruising brownish, and is collected in Fagus sylvatica and Abies alba forests. In contrast, R. sanguinixantha in China shows deeper wine-red bruising and tends to have salmon-colored basidiomata.

Ramaria ser. Nanisporae P. Zhang & J. Zhong, ser. nov.

MycoBank: MB 852717

Etymology: “nanis” meaning “small or short”, is proposing because the short spore.

Type species: R. nanispora R.H. Petersen & M. Zang, Acta bot. Yunn. 11(4): 389 (1989).

Diagnosis: 1) Basidiomata with salmon to yellow-orange branches; 2) Bright orange flesh; 3) Stipe flesh weakly amyloid; 4) Basidiospores short and small.

Commentary: This series includes three species collected in China. Additionally, other species such as R. araiospora Marr & D.E. Stuntz from North America, and its variants: R. araiospora var. araiospora Marr & D.E. Stuntz, R. araiospora var. rubella Marr & D.E. Stuntz, R. fagetorum Maas Geest. ex Schild, and R. subbotrytis (Coker) Corner are also part of this group.

67. Ramaria corallina P. Zhang & J. Zhong, sp. nov. Fig. 27 a−b and Fig. 92

MycoBank: MB 853184

Etymology: “corallina” refers to the coral red color of the basidiomata.

Diagnosis: This species is distinguished within its series by its coral-red basidiomata and basidiospores measuring less than 9 µm in length.

Type: CHINA, YUNNAN PROVINCE: Qujing City, Qilin District, near Longyuan Zhumushan Wind Farm, altitude 2250 m, 19 August 2022, P. Zhang 4712 (Holotype, MHHNU 11212, GenBank Acc. No.: ITS = PP467305, nrLSU = PP493603).

Fig. 92. Microscopic features of R. corallina (MHHNU 11212). a basidiospores; b basidia. Scale bars: a =10 µm, b =20 µm.

Description: Basidiomata 9 cm high and 5 cm broad, elongate obpyriform to obovate in outline. Stipe 3 × 2 cm, tapering gradually downward, single to falsely fasciculate, without abortive branches, not changing color on bruising, off-white (1A1) below, upward light red (10A5) to coral red (9B7), concolorous to branches above; flesh off-white (1A1) to shell pink (8A3), solid, not slippery, Major branches 2−6, ascending, terete, light red (10A5) to coral red(9B7), concolorous to branches. Branches in 4−6 ranks, light red (10A5) to coral red (9B7), sometimes with a light orange (5A5) tinge; internodes diminishing gradually upwards at maturity; axils narrowly rounded when young, broadly rounded at age. Apices broadly rounded, dichotomous, stiffly digitate, finally concolorous with branches. Odor faintly fragrant to middle; taste negligible.

Basidiospores [30/1/1] (7.6−)8.0−9.8(−10.1) × (2.9−)3.0−4.3 μm, L_m = 8.67 µm, l_m = 3.64 µm, Q = (1.90–)2.04–2.90(–2.98), Q_m = 2.40 ± 0.23, ellipsoid to subcylindrical, roughened in profile; hilar appendix broad; ornamentation of prominent, discrete, low warts and short meandering ridges. Basidia 75−92 × 11−13 μm, clavate, without clamps; sterigmata 4, stout, straight. Subhymenium rudimentary, hyphal. Hymenium unthickening. Tramal hyphae of stipe 5−16 μm wide, hyaline, virtually unclamped, thin to thick walled, interwoven, not agglutinated or adherent; ampulliform clamps up to 15 μm broad, not unusually thick-walled, with extensive, coarse, stalactitiform ornamentation; gloeoplerous hyphae not observed. Tramal hyphae of upper branches 5−12 μm wide, hyaline, usually without clamps, usually inflated, parallel, not agglutinated or adherent, loosely parallel; ampulliform clamps up to 12 μm broad; gloeoplerous hyphae not observed.

Ecology: Gregarious on soil in mixed forests dominated by Acer, Quercus and Pinus; basidiomata occurs in autumn.

Distribution: Known from northeast China and southwest China (Yunnan, Jilin, Liaoning).

Additional materials examined: CHINA, JILIN PROVINCE: Tonghua County, Baijifeng Forest Park, in mixed forests dominated by Acer, Quercus and Pinus, altitude 800 m, 28 Aug. 2020, P. Zhang 4040 (MHHNU 10540), P. Zhang 4041 (MHHNU 10541), P. Zhang 4047 (MHHNU 10547). LIAONING PROVINCE: Liaodong Peninsula, in broad-leaved forest, 30 August 2018, P. Zhang 3610 (MHHNU 10110).

Commentary: Ramaria corallina shares similarities in appearance with R. rubrosporula and R. nanispora supported by our two-gene phylogenetic analysis indicating close genetic relationships among them. However, these species can be distinguished as follows: R. rubrosporula typically has a stouter stipe often longitudinally rugulose, while R. nanispora exhibits clamps at the base of basidia and hyphae. Other morphologically similar species include R. cyaneigranosa (basidiospores 9.6 × 4.7 µm; Marr & Stuntz, 1973) and R. stuntzii, however, R. cyaneigranosa possesses longer basidiospores, while R. stuntzii exhibits more reddish basidiomata.

68. Ramaria nanispora R.H. Petersen & M. Zang, Acta bot. Yunn. 11(4): 389, 1989. Fig. 27 c−d and Fig. 93

Description: Basidiomata 10 cm high and 7 cm broad. Stipe 3 × 1.5 cm, modest, tapering to narrowly rounded base, smooth, without abortive branchlet, off-white, weakly watery brunnescent where handled; flesh white, solid, fibrous, not gelatinous or slippery. Major branches several, ascending, flattened, congested upward, concolorous to branches. Branches in 3−7 ranks, terete, salmon colored, retaining orange coloration when dried; flesh brittle, snappy, bright salmon orange; internodes diminishing gradually at maturity; axils rounded throughout. Apices narrowly rounded double-dichotomous, curved-digitate, pale cream color to fleshy cream colored. Odor and taste weakly fabaceous.

Fig. 93. Basidiospores of R. nanispora (MHHNU 11636). Scale bar: a = 10 µm.

Basidiospores [40/2/2] (7.0−)8.0−10.5(−12.0) × 3.5−4.0 μm, L_m = 9.14 µm, l_m = 3.71 µm, Q = (2.00–)2.24–3.00, Q_m = 2.47 ± 0.24, cylindrical, minutely rough in profile; wall up to 0.2 μm thick; hilar appendix gradual, not prominent; ornamentation of small, irregularly shaped plates randomly oriented. Basidia 48−55 × 8−9 μm, clavate, clamped, congested; contents usually homogeneous distally, granular proximally, the granules refringent; sterigmata 4, straight slender. Subhymenium rudimentary. Hymenium unthickening. Tramal hyphae of stipe 3−10 μm wide, hyaline, thick-walled (wall up to 1 μm thick), very tightly interwoven, clamped, not agglutinated or adherent; ampulliform clamps common, up to 14 μm broad, symmetrical, blunt, thick-walled (wall up to 1.5 μm thick), with coarse ornamentation; gloeoplerous hyphae not observed. Tramal hyphae of upper branches 3−11 μm wide, hyaline, thin-walled, inconspicuously clamped or often unclamped, secondarily septate, short celled, strictly parallel, adherent to free; ampulliform clamps and gloeoplerous hyphae not observed.

Ecology: Solitary or gregarious on soil in forests dominated by Pinus Yunnanensis; basidiomata occur in autumn.

Distribution: Known from southwestern China (Yunnan) (Petersen & Zang 1989; Zhang et al. 2005).

Additional materials examined: CHINA, YUNNAN PROVINCE: Dali, Er Yuan County, San Ying Town market, 27 August 1986, No.47293 (Holotype, TENN; isotype, HKAS); Kunming, Qiongzhusi, in a mixed forest with Fagaceae and Pinaceae, altitude 2000 m, 6 August 2006, Jun-Feng Liang 471 (HKAS 50198); Chuxiong, Nanhua County, Majie Town, Dazhongshan Nature Reserve, in broad-leaved forest dominated by Fagaceae, altitude 458 m, 24 August 2015, Jian W. Liu 368 (MHHNU 11636).

Commentary: The diagnostic features of R. nanispora are distinctive within Ramaria.s.l. Its basidiomata displays a salmon-yellow color pattern, accompanied by bright orange branch flesh. The presence of clamped, short basidia and weakly amyloid stipe flesh further characterize this species. Additionally, it is notable for its small, cylindrical basidiospores.

The bright salmon-orange branch flesh of R. nanispora resembles that of R. ephemeroderma, however, R. ephemeroderma lacks clamps and exhibits gelatinous flesh. Similarly, R. nanispora shares similarities with R. leptoformosa in terms of salmon branches and cream-colored apices (Marr & Stuntz 1973), but R. leptoformosa differs significantly with its linear basidiomata structure and non-amyloid stipe flesh. Another comparable species, R. rubricarnata, shares similar colors and amyloid stipe flesh characteristics, but it differs in basidiomata stature and larger basidiospores size (average length of 9.8 μm across all varieties).

Sharda (1983) previously classified a taxon under R. sandaracina Marr & Stuntz or proposed it as a new variety of that species. In Sharda’s concept, R. sandaracina included orange branches and apices, with basidiospores averaging 8.5 μm in length. A variant labeled as “R. sandaracina var. nov.” described apices as “deep yellow” white flesh (likely referring to stipe flesh), and basidiospores measuring 5.6−7 (−7.5) × 3−3.5 μm.

69. Ramaria rubrosporula P. Zhang & J. Zhong, sp. nov. Fig. 27 e−f and Fig. 94

MycoBank: MB 853194

Etymology: rubrosporula = “rubro-” + “sporosia”, proposed because the species has small basidiospores and red to orange-color basidiomata.

Diagnosis: Basidiomata are red to orange colored; apices are yellow or paler than upper branches; basidia are non-clamped and basidiospores are coarsely ornamented, averaging 10.43 × 3.78 μm.

Type: CHINA, YUNNAN PROVINCE: Shangri-La, Pudacuo, Shudu Lake, in the forest with Quercus semecarpifolia, altitude 3600 m, 15 August 2018, P. Zhang 3421 (Holotype, MHHNU 9921, GenBank Acc. No.: ITS = PP467257, nrLSU = PP493563, tef1-α = PP480429).

Description: Basidiomata 9 cm high and 7 cm broad. Stipe 2 × 2 cm, single to falsely fasciculate, stout, tapering gradually downward or rounded at base, somewhat rugulose, with no abortive branchlets, off-white (1A1) to salmon (6A4) below, yellowish orange (4A8) to carrot red (6B7) above, not changing color on bruising. flesh off-white (1A1), solid, not gelatinous or slippery. Major branches 1−2, stout, terete to flattened, concolorous to branches. Branches in 2−4 ranks, yellowish orange (4A8) when young, in age tend to carrot red (6B7) with a light red (10A5) color; internodes diminishing gradually upwards at maturity; axils narrowly rounded (V shaped). Apices double-dichotomous, narrowly rounded to broadly rounded, crowded, yellowish orange (4A8) to carrot red (6B7). Odor faintly fragrant to middle fresh, taste negligible.

Fig. 94. Microscopic features of R. rubrosporula (MHHNU 9921). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Basidiospores [30/2/1] (9.3−)9.6−11.7(−12.9) × (3.1−)3.3−4.6(−4.7) μm, L_m = 10.38 µm, l_m = 3.78 µm, Q = (2.30−)2.39–3.20(–3.36), Q_m = 2.77 ± 0.24, subcylindrical, conspicuously roughened in profile, flattened adaxially; hilar appendix prominent; ornamentation of meandering and anastomosing ridges and warts. Basidia 35−75 × 8−12 μm, clavate, without clamp at base; sterigmata 4, stout and short. Subhymenium rudimentary, pseudoparenchymatous. Hymenium unthickening. Tramal hyphae of stipe 6−16 μm wide, hyaline, without clamp, thin to thick walled (wall up to 1 μm thick), loosely parallel, not agglutinated or adherent; ampulliform clamps up to 14 μm broad, not unusually thick-walled; gloeoplerous hyphae not observed. Tramal hyphae of upper branches 4−10 μm wide, hyaline, without clamps, thick-walled (wall up to 1 μm thick), usually inflated, parallel, not agglutinated or adherent, tightly packed; ampulliform swellings not observed; gloeoplerous hyphae hardly discernable.

Ecology: Solitary or gregarious on soil in high-altitude forests with Picea and Betula platyphylla; basidiomata occurs in autumn.

Distribution: Known from southwestern China (Yunnan, Sichuan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Shangri-La, Pudacuo, Shudu Lake, in the forest with Quercus semecarpifolia, altitude 3600 m, 15 August 2018, P. Zhang 3418 (MHHNU 9918). Ganzi, Kangding City, Gongga Township, Sewulong 2 village, in mixed forests dominated by Picea and Betula platyphylla, altitute 3700 m, 27 August 2017, P. Zhang 2788 (MHHNU 9288).

Commentary: Ramaria rubrosporula shares significant morphological similarities with R. corallina and R. nanispora (basidiospore 8.67 × 3.64 µm and 9.14 × 3.71 µm, as measured in this study). However, the distinguishing feature lies in the length of the basidiospores, with R. rubrosporula possessing longer basidiospores. Additionally, R. rubrosporula bears resemblance to R. araiospora and R. stuntzii, although these species have not been documented in China. Ramaria araiospora is characterized by a rounded base, while R. stuntzii typically exhibits a tendency towards red-colored basidiomata. Notably, R. rubrosporula has only been observed in high altitude locations in southwestern China.

Ramaria ser. Neoformosae (Franchi & M. Marchetti) P. Zhang & J. Zhong stat. nov.

Synonym: Ramaria sect. Neoformosae Franchi & M. Marchetti, Fungi Non Delineati, Raro vel Haud Perspecte et Explorate Descripti aut Definite Picti 16: 34 (2001).

Type species: R. flavicingula R.H. Petersen, Persoonia 14(1): 28 (1989).

Diagnosis: Basidiomata are yellow to salmon colored; flesh is gelatinous; color may change upon injury; basidia may or may not clamps; branches are slender.

Commentary: This series encompasses a diversity of species, with twenty species collected in China and over fourteen species from other countries. Among them are notable species such as R. cistophila P.P. Daniëls, R. admiratia R.H. Petersen, R. acrisiccescens Marr & D.E. Stuntz, R. praecox Schild, R. parecox var. luteovenalis (Franchi, M. Marchetti & Bottoni) Franchi, M. Marchetti & Bottoni, R. calvodistalis R.H. Petersen, R. aurantiisiccescens Marr & D.E. Stuntz, R. flavicingula R.H. Petersen, R. luteoaurantiaca Franchi & M. Marchetti, R. subtilis (Pers.) Quél., R. albidoflava Schild, R. dendrophora G.C. González, R. largentii Marr & D.E. Stuntz, R. aurea (Schaeff.) Quél.

70. Ramaria aurantiformosa P. Zhang & J. Zhong, sp. nov. Fig. 28 a−b and Fig. 95

MycoBank: MB 853354

Etymology: aurantiformosa = aurantius + formosa, referring to the species beautiful and orange-colored basidiomata.

Diagnosis: Basidiomata are cream to salmon colored; apices are vivid yellow when young, becoming concolorous with branches at maturity; turn brunnescent where handled; absence of clamps.

Type: CHINA, SICHUAN PROVINCE: Ganzi, Kangding City, Pengta Township, altitude 3100 m, 19 August 2017, P. Zhang 2680 (Holotype, MHHNU 9180, GenBank Acc. No.: ITS = PP467149, nrLSU = PP493474).

Description: Basidiomata 8–10 cm high and 2–4 cm broad, obovate in outline. Stipe 1 × 2 cm, single, substantial, pruinose between substrate particles, smooth above, somewhat rugulose, off-white (1A1), brunnescent where handled; flesh off-white (1A1), soft, punky, not gelatinous or slippery, drying firm, easily penetrated, but not friable. Major branches 2–3, stout, short, terete. Branches in 3–6 ranks, terete, ascending, light orange (5A5) to salmon (6A4); flesh soft, beautiful peach (7A4) colored; internodes diminishing gradually upward at maturity; axils narrowly rounded below, rounded above. Apices finely divided, crowded, cristate, light orange (5A5) colored when young, short-digitate by maturity, retaining salmon (6A4) color. Odor negligible; taste negligible when fresh, bitter when dry.

Fig. 95. Microscopic features of R. aurantiformosa (MHHNU 9180). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Basidiospores [50/2/2] (9.5–)10.0–13.0 × 4.0(4.2–)–5.4(–5.5) µm, L_m = 11.48 µm, l_m =4.79 µm, Q = (2.10–)2.17–2.88(–2.89), Q_m = 2.41 ± 0.19, ellipsoid, flattened adaxially, conspicuously roughened in profile; wall up to 0.3 µm thick; hilar appendix not prominent; ornamentation of very complex, narrow, meandering and anastomosing ridges. Basidia 55–70 × 9–10 µm, clavate, without clamps; sterigmata (1–2–3–) 4. Subhymenium rudimentary; hyphae 1.5–2 µm, without clamps, tightly interwoven. Tramal hyphae of stipe 3–13 µm wide, hyaline, gnarled, stiff, freely branched, easily shattered in mounts, thick-walled (wall up to 2 µm thick), interwoven, not agglutinated or adherent; ampulliform septa up to 14 µm broad, thick-walled (wall up to 3 µm thick), with delicate ornamentation; gloeoplerous hyphae not observed. Tramal hyphae of upper branches up to 4–12 µm wide, hyaline, thin- to thick-walled (wall up to 0.5 µm thick), parallel, without clamps, tightly packed, not agglutinated or adherent; ampulliform inflations not always associated with septa, thick-walled (wall up to 1 µm thick), with delicate, extensive ornamentations; gloeoplerous hyphae not observed.

Ecology: Solitary or gregarious on soil in mixed forests or in broad-leaved forests. Basidiomata occurs in summer and autumn.

Distribution: In southwestern China (Yunnan, Sichuan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Shangri-La City, Pudacuo National Park, Bitahai, in the mixed forest with Quercus and Picea. altitude 3500 m, 31 August 2022, P. Zhang 4841 (MHHNU 11341); same location, Tianbaotai, in the mixed forest with Quercus and Picea. altitude 3700 m, 1 September 2022, P. Zhang 4860 (MHHNU 11360), P. Zhang 4861 (MHHNU 11361); Dali, Dali City, Xianyangfeng, in a mixed forest, altitude 2800 m, 12 August 2018, P. Zhang 3375 (MHHNU 9875); Lijiang City, Yulong County, 16 July 2004, Qi Zhao Lj-97 (HKAS 59221). SICHUAN PROVINCE: Ganzi, Kangding City, Pengta Township, altitude 3100 m, 19 August 2017, P. Zhang 2674 (MHHNU 9174), P. Zhang 2679 (MHHNU 9179).

Commentary: In Zhang (2010), this species was identified as R. neoformosa var. sinensis R.H. Petersen & M. Zang. However, further collections revealed that its stipes are falsely fasciculate rather than singly stout. Within the R. neoformosa species complex, there are two other varieties: var. neoformosa and var. macrospora. The latter, found in eastern Canada, features larger basidiospores. Ramaria aurantiformosa shares many similarities with R. formosa in terms of color and basidiomata morphology. However, the presence or absence of clamps easily distinguishes the latter two (Quélet 1888; Corner 1950; Franchi & Marchetti 2021).

Moreover, the species most similar to R. aurantiformosa are R. hilaris and R. aureoapicalis, all of which are salmon to orange colored Ramaria found in broad-leaved or mixed forests. They lack clamps, but the basidiospores of the latter two are slightly smaller. While other features are challenging to differentiate, genetic distances in our two-gene molecular phylogenetic analysis (see Fig. 2) confirm them as three distinct species.

71. Ramaria aurantilinearis P. Zhang & J. Zhong, sp. nov. Fig. 28 c−d and Fig. 96

MycoBank: MB 853355

Etymology: “aurantilinearis” = aurantius + linearis, referring to the basidiomata of the species being orange in color and having slender branches.

Diagnosis: Basidiomata are large-sized, dark orange to salmon; stipe slender, fleshy to fibrous; apices are apricot yellow when young, becoming concolorous with branches at maturity; basidia base and hyphae lack clamps; basidiospores are roughened in profile, long and narrow.

Type: CHINA, SICHUAN PROVINCE: Ganzi, Kangding City, Gongga Mountain, in Quercus and Picea forest, altitude 3900 m, 6 September 2021, P. Zhang 4256 (Holotype, MHHNU 10756, GenBank Acc. No.: ITS = PP467172, nrLSU = PP493495, tef1-α = PP480380).

Description: Basidiomata 12 × 10 cm, medium sized to large. Stipes up to 5 × 5 cm, single, slender; off-white (1A1) below substrate level, surface pale orange (5A3) to yellowish orange (4A8) upward; surface smooth; nearly without abortive branches; not changing color or weakly brownish orange (6C3) where handled; context of stipe off-white (1A1) when fresh, fleshy to fibrous, drying light in weight, easily penetrated, cartilaginous. Major branches two to several, slender, ascending, terete. Branches in 3−7 ranks, slender, crowded, major branches and upper branches commonly polychotomous; branches salmon (6A4) to melon yellow (5A6); axils mixed narrowly rounded and broadly rounded, mixed U-shaped and U-shaped; internodes all short, lower internodes slightly long; context of main branches and upper branches fleshy to fibrous, not gelatinous or slippery, off-white (1A1) to pale orange(5A3). Apices slender and acute, short and crowded, polychotomous, pale orange (5A3) to apricot yellow (5B6) when young, concolorous with branches when maturity. Odor weakly fragrant, aromatic; taste negligible to faintly astringent.

Fig. 96. Microscopic features of R. aurantilinearis (MHHNU 10756). a basidiospores; b basidia. Scale bars: a =10 µm, d = 20 µm.

Basidiospores [60/2/2] (12.6–)13.2–16.5(–19.0) × (4.1–)4.9–6.4(–6.6) µm, L_m = 14.95 µm, l_m = 5.54 µm, Q = (2.18–)2.29–3.17(–3.81), Q_m = 2.73 ± 0.32, subcylindrical to long-ellipsoid; roughened in profile; ornamented with warts and short meandering ridges randomly placed, strongly cyanophilous warts; hilar not conspicuous. Basidia 50–70 × 8–15 µm, clavate, without basal clamps; sterigmata 4, 5–8 µm long, stout, somewhat scattered. Hymenium unthickening, 60–80 µm. Subhymenium extensive, hyphae, interwoven, 2–5 µm diam, thin-walled, without clamps. Contextual hyphae: Tramal hyphae of stipe 5–18 μm wide, loosely parallel, ampulliform inflations up to 18 μm broad, with coarse ornamentation; tramal hyphae of branches central hyphae slightly narrow, hyaline, thin-walled, strictly parallel, ampulliform inflations up to 18 μm broad; without clamps; gloeoplereous hyphae not observed.

Ecology: Solitary or gregarious on soil in mixed forests of Quercus, Picea, and Abies. Basidiomata occurs in summer and autumn.

Distribution: In southwest China (Sichuan).

Additional materials examined: CHINA, SICHUAN PROVINCE: Ganzi, Kangding City, Gongga Mountain, in Quercus and Picea forest, altitude 3900 m, 6 September 2021, P. Zhang 4286 (MHHNU 10786), P. Zhang 4295 (MHHNU 10795), P. Zhang 4298 (MHHNU 10798), P. Zhang 4315 (MHHNU 10815), P. Zhang 4316 (MHHNU 10816).

Commentary: In the wild, the large basidiomata and slender branches of R. aurantilinearis are crucial characteristics for its identification. Currently, this species has only been collected in the high-altitude regions (3000–4000 m above sea level) of Gongga Mountain in China. The small branches are fragile and prone to breaking. Unlike other Ramaria species with large basidiomata that typically have stout stipes, R. aurantilinearis features densely packed and slender branches. As these basidiomata mature, the internodes elongate, and the number of branches ranks increase, resulting in increasingly massive basidiomata. Under microscopic examination, this species lacks clamps, and its basidiospores are relatively narrow and elongated. The combining of these macroscopic and microscopic features makes R. aurantilinearis easily distinguishable among all the yellow/orange Ramaria species.

72. Ramaria aureoapicalis P. Zhang & J. Zhong, sp. nov. Fig. 28 e and Fig. 97

MycoBank: MB 853353

Etymology: "aureoapicalis" conveys the characteristic of having golden-yellow apices.

Diagnosis: Basidiomata are salmon colored; apices are vivid yellow when young, becoming concolorous with branches at maturity; turn brunnescent when handled; and absence of clamps.

Type: CHINA, HUNAN PROVINCE: Zhangjiajie City, Sangzhi County, Badagongshan National Nature Reserve, in broad-leaved forest, altitude 1500 m, 14 September 2017, P. Zhang 2828 (Holotype, MHHNU 9328, GenBank Acc. No.: ITS = PP467146, nrLSU = PP493473, tef1-α = PP480363).

Description: Basidiomata 6–13× 3–10 cm, solitary to gregarious. Stipes 2.3–5.7 × 0.8–2 cm, single to falsely fasciculate, tapering gradually downward, acute at base; off-white (1A1) below, surface milk white (1A2) to pale yellow (2A3); surface smooth; with abortive branches; changing color to brownish orange (6C3) where handled; context of stipe off-white (1A1) to pale yellow (2A3) when fresh, paler than hymenium, fleshy to fibrous, drying light in weight, easily penetrated, cartilaginous. Major branches 2–several, ascending, terete. Branches in 3−5 ranks, major branches and upper branches mostly dichotomous; branches whitish orange (5A2) to cream (4A3); axils narrowly rounded, U-shaped or V-shaped; internodes diminishing gradually at maturity, lower internodes long; context of main branches and upper branches fleshy to fibrous, not gelatinous or slippery, pastel yellow (2A4) to pale orange (5A3). Apices acute and slender, dichotomous, vivid yellow (2A4) when young, concolorous with branches by maturity. Odor weakly fragrant, aromatic; taste negligible to faintly astringent.

Fig. 97. Microscopic features of R. aureoapicalis (MHHNU 9328). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Basidiospores [60/2/2] (8.0–)9.0–13.0(–14.0) × 4.0–6.0(–7.0) µm, L_m = 10.23 µm, l_m = 4.75 µm, Q = 1.67–2.75(–3.00), Q_m = 2.17 ± 0.34, subcylindrical to long-ellipsoid; conspicuously roughened in profile; ornamented with warts and short meandering ridges randomly placed, strongly cyanophilous warts; hilar not conspicuous. Basidia 40–73 × 8–12 µm, clavate, without basal clamps; sterigmata 4, 4–5 µm long, stout and straight, somewhat scattered. Hymenium unthickening, 60–80 µm. Subhymenium extensive, hyphae, interwoven, 2–5 µm diam, thin-walled, without clamps. Contextual hyphae: Tramal hyphae of stipe 5–18 μm wide, loosely parallel, ampulliform inflations up to 20 μm broad, with coarse ornamentation; tramal hyphae of branches central hyphae slightly narrow, hyaline, thin-walled, strictly parallel, ampulliform inflations up to 20 μm broad; without clamps; gloeoplereous hyphae not observed.

Ecology: Solitary or gregarious on soil in broad-leaved forests. Basidiomata occurs in summer and autumn.

Distribution: In central China (Hunan).

Additional materials examined: CHINA, HUNAN PROVINCE: Chenzhou City, Rucheng County, Daping Town, Jiulongjiang National Forest Park, in broad-leaved forest, altitude 100 m, 2 October 2020, P. Zhang 4066 (MHHNU 10566); Zhangjiajie City, Sangzhi County, Badagongshan National Nature Reserve, in the mixed forest, altitude 1400 m, 15 September 2020, Zuo H. Chen 32483 (MHHNU 32483); same location, in broad-leaved forest, altitude 1000 m, 17 September 2019, P. Zhang 3765 (MHHNU 10265).

Commentary: Ramaria aureoapicalis bears a striking resemblance to both R. hilaris and R. aurantiformosa. All three species exhibit salmon to orange-yellow basidiomata with bright yellow apices when young, lack clamps, and show only slight difference in basidiospores size. Our two-gene phylogenetic analysis also indicates a very close genetic relationship among these species. While they all thrive in broad-leaved forests or mixed coniferous-broadleaved forests, R. aurantiformosais is typically found in the southwestern region of China, R. hilaris in eastern China, and R. aureoapicalis in central China.

73. Ramaria aureolinearis P. Zhang & J. Zhong, sp. nov. Fig. 28 f and Fig. 98

MycoBank: MB 853362

Etymology: “aureolinearis” indicates that the species is golden-yellow in color and has slender branches.

Diagnosis: Basidiomata are gold yellow to pale orange; absence of clamps; branches are slender.

Type: CHINA, JILIN PROVINCE: Tonghua City, Baijifeng Forest Park, in mixed forests dominated by Acer, Quercus and Pinus, altitude 800 m, 28 Aug. 2020, P. Zhang 4033 (Holotype, MHHNU 10533, GenBank Acc. No.: ITS = PP467166, nrLSU = PP493489, tef1-α = PP480375).

Description: Basidiomata 14 × 10 cm, medium sized to large. Stipes up to 4 × 2 cm, stingle to falsely fasciculate, tapering gradually downward; off-white (1A1) below substrate level, surface pale yellow (2A3) to cream (4A3) upward; surface smooth; with abortive branches; changing color to weakly lemon yellow (3B8) to brownish orange (6C3) where handled; context of stipe off-white (1A1) when fresh, fleshy to fibrous, drying light in weight, easily penetrated, cartilaginous. Major branches several, ascending, hardly terete. Branches in 3−5 ranks, slender, major branches commonly polychotomous and upper branches mostly dichotomous; branches salmon (6A4) to melon yellow (5A6); axils mixed narrowly rounded and broadly rounded, mixed U-shaped and U-shaped; internodes diminishing gradually at maturity, lower internodes long; context of main branches and upper branches fleshy to fibrous, not gelatinous or slippery, off-white (1A1) to pale orange (5A5). Apices are slender and acute, crowded, polychotomous, pale orange (5A3) to vivid yellow (2A8), or concolorous with branches. Odor weakly fragrant, aromatic; taste negligible to faintly astringent.

Fig. 98. Microscopic features of R. aureolinearis (MHHNU 10533). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Basidiospores [60/2/2] (9.7–)11.4–13.2(–13.7) × (3.3–)3.5–4.9(–5.1) µm, L_m = 11.68 µm, l_m = 4.13 µm, Q = (2.37–)2.44–3.34(–3.41), Q_m = 2.84 ± 0.28, subcylindrical to long-ellipsoid; conspicuously roughened in profile; ornamented with warts and short meandering ridges randomly placed, strongly cyanophilous warts; hilar not conspicuous. Basidia 45–55 × 7–12 µm, clavate, without basal clamps; sterigmata 4, 2–5 µm long, stout, short and straight, somewhat scattered. Hymenium unthickening, 50–60 µm. Subhymenium extensive, hyphae, interwoven, 2–5 µm diam, thin-walled, without clamps. Contextual hyphae: Tramal hyphae of stipe 5–18 μm wide, loosely parallel, ampulliform inflations up to 20 μm broad, with coarse ornamentation; tramal hyphae of branches central hyphae slightly narrow, hyaline, thin-walled, strictly parallel, ampulliform inflations up to 20 μm broad; without clamps; gloeoplereous hyphae not observed.

Ecology: Solitary or gregarious on soil in broad-leaved and coniferous mixed forests. Basidiomata occurs in summer and autumn.

Distribution: In northeast China (Jilin).

Additional materials examined: CHINA, JILIN PROVINCE: Tonghua City, Baijifeng Forest Park, in mixed forests dominated by Acer, Quercus and Pinus, altitude 800 m, 28 Aug. 2020, P. Zhang 4032 (MHHNU 10532).

Commentary: Ramaria aureolinearis bears resemblance to R. aurea and R. albidoflava Schild. Compared to R. aurea (basidiospore 10.9 × 4.7 µm; Franchi & Marchetti 2021), both species have orange color in their basidiomata, but R. aureolinearis has slightly larger basidiospores and more slender branches. Compared to R. albidoflava (= R. subtilis var. crassispora), the latter has slightly larger basidiospores (12.7 × 5.4 µm; Franchi & Marchetti 2021). However, basidiospores sizes in the genus Ramaria can vary during different stages of basidiomata development, so it cannot be used as the sole distinguishing feature. Additionally, while R. aureolinearis has orange to salmon-colored basidiomata, R. albidoflava exhibits a light-yellow color. Our two-gene molecular phylogenetic analysis supports these three distinct species.

74. Ramaria brunnescentiflava P. Zhang & J. Zhong, sp. nov. Fig. 28 g and Fig. 99

MycoBank: MB 853352

Etymology: brunnescentiflava = brunnescens + flava, indicating that the species exhibits a brown coloration in mature basidiomata.

Fig. 99. Microscopic features of R. brunnescentiflava (MHHNU 9290). a basidiospores; b basidia. Scale bars: a =10 µm, b = 20 µm.

Diagnosis: Basidiomata are yellow when young, becoming salmon with brownish tones at maturity; stipe is fasciculate or with a very small point of union at the base, acute at the base, and gelatinous; apices are vivid yellow when young, becoming concolorous with branches at maturity; basidia base and hyphae have clamps; changes color to lemon yellow when injured.

Type: CHINA, YUNNAN PROVINCE: Lijiang, Yulong County, Yulong Snow Mountain, altitude 3700 m, 27 July 2017, P. Zhang 2790 (Holotype, MHHNU 9290, GenBank Acc. No.: ITS = PP467177, nrLSU = PP493499, tef1-α = PP480384).

Description: Basidiomata 11 × 8–10 cm, medium sized. Stipes up to 1 × 2 cm, fasciculate or with a very small point of union at the base, tapering sharply downward to an acute base; off-white (1A1) below substrate level, surface pale yellow (2A3) to salmon (6A4) upward; surface smooth; without abortive branches; changing color to lemon yellow (3C8) where handled; context of stipe ivory white (4B3) to grayish orange (6B5) when fresh, gelatinous, difficult to penetrated, cartilaginous. Major branches 2–4, slender, ascending, terete. Branches in 3−5 ranks, crowded, major branches and upper branches commonly polychotomous; branches salmon (6A4) to melon yellow (5A6); axils mixed broadly rounded, U-shaped; internodes length ratio diminishing gradually at maturity when young, all short by maturity; context of main branches and upper branches gelatinous, ivory white (4B3) to butter yellow (4A5). Apices slender and acute when young, broadly rounded when maturity, short digitate, vivid yellow (2A8) when young, concolorous with branches when maturity. Odor weakly fragrant, aromatic; taste negligible to faintly astringent.

Basidiospores [60/2/2] (10.0–)10.8–14.8(–15.7) × 3.9–5.2(–5.3) µm, L_m = 12.52 µm, l_m = 4.54 µm, Q = (2.23–)2.38–3.17(–3.48), Q_m = 2.77 ± 0.27, subcylindrical to long-ellipsoid; roughened in profile; ornamented with warts and short meandering ridges randomly placed, strongly cyanophilous warts; hilar not conspicuous. Basidia 45–65 × 7–10 µm, clavate, with basal clamps; sterigmata 4, 2–5 µm long, stout and short. Hymenium unthickening, 60–80 µm. Subhymenium extensive, hyphae, interwoven, 2–5 µm diam, thin-walled, with clamps. Contextual hyphae: Tramal hyphae of stipe 5–10 μm wide, tightly interwoven, ampulliform inflations not observed; tramal hyphae of branches central hyphae slightly narrow, hyaline, thin-walled, tightly interwoven, ampulliform inflations not observed; with clamps; gloeoplereous hyphae not observed.

Ecology: Solitary or gregarious on soil in Picea and Abies forests, or in mixed forests. Basidiomata occurs in summer and autumn.

Distribution: In southwest China (Yunnan, Sichuan).

Additional materials examined: CHINA, YUNNAN PROVINCE: Lijiang, Yulong County, Yulong Snow Mountain, altitude 3700 m, 27 July 2017, P. Zhang 2784 (MHHNU 9284), P. Zhang 2786 (MHHNU 9286). SICHUAN PROVINCE: Ganzi, Kangding City, Gongga Township, Sewurong II Village, in a forest with Quercus, altitude 3700 m, 21 August 2017, P. Zhang 2791 (MHHNU 9291).

Commentary: Ramaria brunnescentiflava occupies a unique position in the two-gene phylogenetic analysis, distinguishing it as a distinct species. Mature basidiomata of R. brunnescentiflava are salmon-colored with brownish tones, and their stipes are compound-connate at the base. This species exhibits a characteristic of turning lemon yellow upon injury and is identified by the presence of prominent clamps. It shares a close affinity with a specimen labeled MF80513 collected in the southern United States, both displaying yellow orange basidiomata and gelatinous, falsely fasciculate stipes. Additionally, R. brunnescentiflava is closely related to R. cistophila P.P. Daniëls & M.P. Martín (10.9 × 4.7 µm, Daniëls PP et al. 2017) collected in Spain under Cistus ladanifer, which also has salmon-colored basidiomata but lacks clamps, furthermore, the basidiospores of R. brunnescentiflava are larger.

75. Ramaria conferta L. Fan, Ying Li & N. Mao, in Li, Mao, Zhang, Fu & Fan, Mycology: 10.1080/21501203.2024.2388910, 17 (2024) Fig. 28 w–x and Fig. 100

Description: Basidiomata 10 × 7 cm, medium sized. Stipes up to 3 × 3.5 cm, single, terete, rounded at base; off-white below substrate level, surface pale orange to yellowish orange upward; surface smooth; nearly without abortive branches; not changing color or weakly brownish orange where handled; context of stipe off-white when fresh, fleshy to fibrous, drying light in weight, easily penetrated, cartilaginous. Major branches two to several, slender, ascending, terete. Branches in 3−7 ranks, slender, crowded, major branches and upper branches commonly polychotomous; branches salmon to melon yellow; axils mixed narrowly rounded and broadly rounded, mixed U-shaped and U-shaped; internodes all short, lower internodes slightly long; context of main branches and upper branches fleshy to fibrous, not gelatinous or slippery, off-white to pale orange. Apices slender and acute, short and crowded, polychotomous, concolorous with branches when young, sometimes darker to brown when maturity. Odor weakly fragrant, aromatic; taste negligible to faintly astringent.

Fig. 100. Basidiospores of R. conferta (MHHNU 10088). Scale bar: a = 10 µm.

Basidiospores [60/2/2] (10.1–)10.7–14.3(–14.8) × (4.0–)4.3–6.2(–6.5) µm, L_m = 12.31 µm, l_m = 5.24 µm, Q = (1.99–)2.03–2.72(–3.06), Q_m = 2.37 ± 0.23, subcylindrical to long-ellipsoid; roughened in profile; ornamented with warts and short meandering ridges randomly placed, strongly cyanophilous warts; hilar not conspicuous. Basidia 45–65 × 7–12 µm, clavate, without basal clamps; sterigmata 4, 5–8 µm long, stout, somewhat scattered. Hymenium unthickening, 60–80 µm. Subhymenium extensive, hyphae, interwoven, 2–5 µm diam, thin-walled, without clamps. Contextual hyphae: Tramal hyphae of stipe 5–18 μm wide, loosely parallel, ampulliform inflations up to 18 μm broad, with coarse ornamentation; tramal hyphae of branches central hyphae slightly narrow, hyaline, thin-walled, strictly parallel, ampulliform inflations up to 18 μm broad; without clamps; gloeoplereous hyphae not observed.

Ecology: Solitary or gregarious on soil in Picea and Abies forests. Basidiomata occurs in summer and autumn.

Distribution: In southwest, north and northwest China (Sichuan, Xinjiang, Shanxi).

Additional materials examined: CHINA, SICHUAN PROVINCE: Ganzi, Kangding City, Gongga Mountain, in the Abies forest, altitude 3500 m, 2 September 2021, P. Zhang 3588 (MHHNU 10088); same city, Waze Township, in Picea forest, altitude 3800 m,1 September 2018, P. Zhang 3567 (MHHNU 10067), P. Zhang 3568 (MHHNU 10068), P. Zhang 3570 (MHHNU 10070); same location, 23 August 2017, P. Zhang 2732 (MHHNU 9232); same city, Mugecuo, in a Abies forest, altitude 3500 m, 21 August 2017, P. Zhang 2707 (MHHNU 9207). XINJIANG UYGUR AUTONOMOUS REGION: Urumqi City, Urumqi County, Nan Mountai, in Picea and Abies forest., altitude 2000 m, 24 August 2013, P. Zhang 1335 (MHHNU 7835). GANSU PROVINCE: Lanzhou City, Yuzhong County, Xinglong Mountain Nature Reserve, Dong Mountain, in a coniferous and broad-leaved mixed forest, altitude 2300 m, 7 August 2019, P. Zhang 3705 (MHHNU 10205).

Commentary: The distribution of R. conferta is relatively extensive, ranging from the southwestern to the northwestern of China. It prefers to grow under Picea trees and shares a close phylogenetic relationship with the R. robustibrunnea and R. aurantilinearis. These three species exhibit similar-colored basidiomata and lack clamps. They are commonly found in forests with Picea trees. However, compared to the R. robustibrunnea (11.01 × 4.69 µm), R. conferta has a slender stipe, and its basidiospores are larger. In comparison to R. aurantilinearis (14.95 × 5.54 µm), both the basidiospores and fruiting body sizes of R. conferta are smaller.

76. Ramaria cyanophila L. Fan, Ying Li & N. Mao, in Li, Mao, Zhang, Fu & Fan, Mycology: 10.1080/21501203.2024.2388910, 19 (2024)

Original description in Li et al. (2024).

77. Ramaria distinctissima R.H. Petersen & M. Zang, Acta bot. Yunn. 11(4): 376, 1989. Fig. 28 h and Fig. 101

Description: Basidiomata 5–7 cm high and 3–7 cm broad, obpyriforme in outline. Stipes up to 2 × 1–2 cm, tapering gradually to the acute, often rooting at base, tomentose at base, smooth upward, without abortive branches, off-white where protected, easily brunnescent around substrate bruises but not on handling; flesh white, solid, moist, drying brittle, penetrated with difficulty but not to impenetrable. Major branches 2–4, terete, ascending, concolorous to stipe below, concolorous to branches above. Branches in 3–6 ranks, terete, ascending, pastel yellow orange or yellow below, upward paster orange; internodes diminishing gradually upwards at maturity; axils narrowly rounder below, rounded above; flesh somewhat yellow than hymenium, white inward, drying brittle, friable. Apices double-dichotomous when young, elongating to short-digitate by maturity, rich golden orange to yellowish orange. Odor negligible or faintly fresh; taste negligible.

Fig. 101. Basidiospore of R. distinctissima (MHHNU 9980). Scale bars: a = 10 µm.

Basidiospores [40/2/2] (12.0)12.7–15.6(–16.1) × (4.4–)4.8–6.0(–6.2) µm, L_m = 14.20 µm, l_m =5.22 µm, Q = (2.16–)2.36–3.04(–3.29), Q_m = 2.74 ± 0.22, long-ellipsoid, conspicuously roughened in profile; wall up to 0.3 µm thick; hilar appendix gradual, not prominent, with apparent struma; ornamentation of prominent, discrete, low warts and short meandering ridges. Basidia 85–95 × 10–12 µm, clavate, clamped, in tight cymes; weakly cyanophilous; sterigmata 4, up to 6 µm long, spindly, straight. Subhymenium rudimentary; pseudo-parenchymatous; hyphae 1.5–3 µm, conspicuously clamped. Hymenium thickening; Tramal hyphae of stipe 4–15 µm wide, hyaline, thin to thick-walled (wall up to 0.3 µm thick), interwoven, hyaline, thin to thick-walled (wall up to 0.5 µm thick), without clamps, interwoven, not agglutinated or adherent, thick-walled (wall up to 1.5 µm thick); usually without clamps, tightly packed; ampulliform swelling up to 15 µm broad, often not associated with septa, not unusually thick-walled, with extensive ornamentation; gloeoplerous hyphae not observed. Tramal hyphae of upper branches up to 4-18 µm wide, inflated especially inward, hyaline, thin- to thick-walled (wall up to 0.5 µm thick), almost always without clamps, parallel, not agglutinated or adherent; cells cigar-shaped; ampulliform swelling and gloeoplerous not observed.

Ecology: Solitary or gregarious on soil in Picea and Abies forests. Basidiomata occurs in summer and autumn.

Distribution: In southwest China (Yunnan, Sichuan, Xizang).

Additional materials examined: CHINA, YUNNAN PROVINCE: Lijiang City, Yulong County, 16 July 2004, Qi Zhao Lj-97 (HKAS 59221). SICHUAN PROVINCE: Ganzi, Jiulong County, northern Tanggu Township, Wuxuhai, in Picea and Abies forest, altitude 4100 m, 19 August 2023, Zuo H. Chen 34086 (MHHNU 34086); Shangri-La City, Pudacuo National Park, Tianbaotai, in the mixed forest with Quercus and Picea. altitude 3700 m, 1 September 2022, P. Zhang 4866 (MHHNU 11366). XIZANG AUTONOMOUS REGION: Changdu City, Leiwuqi County, altitude 3900 m, 10 August 2004, Zhu L. Yang 4356 (HKAS 45735); same city, Basu County, Ranwu Lake, in Picea and Abies forest, altitude 3,900 m, 22 August 2018, P. Zhang 3480 (MHHNU 9980).

Commentary: The main features of R. distinctissima include: 1) salmon to orange-colored branches of the basidiomata; 2) stipe tapering gradually to an acute point, fleshy to fibrous, and concolorous with the branches; 3) absence of clamps at the base of basidia and in hyphae; 4) preference for growing in Picea and Abies forests.

The basidiospores size of R. distinctissima is slightly smaller than that of R. sinolargentii. The golden-yellow coloration of the upper stipe and lower major branches may result from exposed hymenium due to injury. This species shares similarities with R. flavicingula (12.8 × 4.8 µm, Franchi & Marchetti 2021), R. largentii (13.4 × 4.5 µm, Marr & Stuntz 1973), and R. luteoaurantiaca (13.0 × 4.6 µm, Franchi & Marchetti 2021). However, its basidiospores are larger than those of the three species.

The virtually indistinguishable R. distinctissima var. americana (Petersen & Scates 1988) in North America differs from the typical variety in the following characters: 1) the stipe of basidiomata tapers more sharply into an obconic base; 2) tramal hyphae of the upper branches are strictly thin walled; 3) basidia are somewhat wider than those of R. distinctissima var. distinctissima.

78. Ramaria ephemeroderma R.H. Petersen & M. Zang, Acta bot. Yunn. 11(4): 378, 1989. Fig. 28 i and Fig. 102

Description: Basidiomata 13 cm high and 6 cm broad, circular to broadly fusiform in outline. Stipe 3 × 2 cm, single, falsely fasciculate, lobed or muscled, tapering to a blunt base, spongy when pressed, pruinose between major branches, otherwise smooth, with no abortive branchlets, off-white, not brunnescent; flesh firmly gelatinous, slippery, off-white below, paster orange at junction with major branches. Major branches 3–several, curved-ascending, hardly terete, pallid salmon-pink with some areas rich yellow. Branches terete, in 3–7 ranks, ascending, salmon color; branch surface (hymenium) quickly fading after picking to pale pink or pale salmon pink; branch flesh watery gelatinous, stuffed to hollow, brittle, bright orange remaining so during drying, drying horny, cartilaginous; internodes diminishing gradually upward at maturity; axils lunate to narrowly rounded. Apices fine, delicate, double-dichotomous when young, delicately digitate by maturity, bright yellow. Odor fainty fresh or penetrating; taste negligible.