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Journal of Fungi
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12 July 2024

Four New Species of Jelly Fungi from Northeastern China

and
1
College of Mycology, Jilin Agricultural University, Changchun 130118, China
2
Key Laboratory of Edible Fungal Resources and Utilization (North), Ministry of Agriculture and Rural Affairs, Jilin Agricultural University, Changchun 130118, China
*
Author to whom correspondence should be addressed.
J. Fungi2024, 10(7), 480;https://doi.org/10.3390/jof10070480
This article belongs to the Special Issue Diversity, Phylogeny and Ecology of Forest Fungi
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Abstract

Four new species of jelly fungi were described from northeastern China based on morphological and molecular evidence. These new species were classified into the four genera Sirobasidium (Sirobasidium jilinense), Calocera (Calocera velutina), Dacrymyces (Dacrymyces jauensis), and Dacryopinax (Dacryopinax manghanensis). Maximum likelihood and Bayesian analyses were performed using a combined nuc rDNA internal transcribed spacer region (ITS) and nuc 28S rDNA (nrLSU) dataset for the construction of phylogenetic trees. Morphological descriptions, line illustrations, and the ecological habits of these new species are provided.

1. Introduction

Sirobasidium Lagerh. & Pat. was established in 1892, with the type species being Sirobasidium sanguineum Lagerh. & Pat [1]. Due to the presence of hypobasidia that are divided vertically, the genus was first assigned to Tremellaceae Fr. In 1895, Möller proposed the establishment of Sirobasidiaceae Lindau due to the unique chain-like hypobasidia of this species [2]. Two years later, Sirobasidiaceae was officially published by Lindau [3]. Lowy then discovered that the French record of S. cerasi Bourdot & Galzin did not match the requirements of the genus concerning the size of its basidiomata, the quantity of chain-shaped basidia, and the size of its basidiospores. Lowy also provided a key for the recognized species S. brefeldianum Möller, S. magnum Boedijn, S. albidum, and S. sanguineum [4]. It was not until Bandoni examined the basidia and basidiospores of Sirobasidium that the spindle-shaped epiprobasidia shed by species in this genus were recognized as basidiospores [5]. In 2015, Roedel and Putzmann compiled a key for the Sirobasidium in their study of S. albidum, which included eight species [6]. Unfortunately, molecular information was not provided. At present, the genus contains ten species [3], three of which have been identified in China: S. japonicum, S. magnum, and S. sanguineum [7]. The chain-like hypobasidia and exfoliated epiprobasidia are the two primary characteristics of members of this genus. Phylogenetic analyses have shown the genus to be polyphyletic [8,9,10,11,12,13]. It is currently challenging to confirm the taxonomic status of Sirobasidium, however, as there are currently no molecular sequences available for the type species.
Dacrymycetaceae J. Schröt. (1888) belongs to Basidiomycota [3]. This family includes a variety of small-sized, morphologically diverse, and gelatinous species that are hard to find when dry. The majority of the basidiomata are yellow and contain carotene. The majority of basidiospores are often separated transversely, occasionally they can also be separated longitudinally or obliquely, or not septate when immature, septate by thin or thick walls; germination produces conidia or not; basidia and basidiospores contain oil droplets; hyphidia are present or absent; and with or without clamp connection [14].
In the 1960s and 1970s, McNabb conducted a detailed study on the genera of Dacrymycetaceae based on their morphology and compiled a key [15,16,17]. Since the 21st century, with the development of molecular systematics, researchers have conducted more in-depth research on Dacrymycetaceae species. In 2007, Shirouzu et al. found, in their phylogenetic analysis of Dacrymyces Nees based on 28S, that Dacrymyces was polyphyletic. While some species of Dacrymyces nest in Cerinomycetaceae Jülich or form a separate clade within Dacrymycetes, the majority of Dacrymyces species are found nested in Dacrymycetaceae [18]. In their 2009 study of Dacrymycetes in Japan, Shirouzu et al. described five new species and provided line illustrations [14]. Based on morphological observations, Malysheva (2013) identified the species of Calocera (Fr.) Fr. in Russia, described eight species of the genus, and provided line illustrations [19]. Later, Shirouzu et al. published eight new species of Dacrymycetes from New Zealand [20]. Dacryopinax G.W. Martin species from Brazil and Mexico were studied by Renato et al. and Castro-Santiuste et al., respectively [21,22]. The above studies show that most genera in Dacrymycetaceae are polyphyletic, and species are scattered in various branches of Dacrymycetaceae.
“Flora Fungorum Sinicorum Vol. 2 Tremellales et Dacrymycetales” by Liu was the primary focus of early Chinese research on Dacrymycetaceae species [7]. At the time, only morphological research was included due to the restricted conditions. Chinese researchers have recently studied species such as Calocera and Dacrymyces using multi-gene fragments [23,24,25]. However, most of the specimens used in these studies were collected from southern China, while there are relatively few specimens from northeastern China.
To enrich the species resources of jelly fungi in Northeast China. We have discovered four new species through morphological research and phylogenetic analysis. This study provided a detailed description and discussion of these species.

2. Materials and Methods

2.1. Specimen Collection

The specimens for this study were collected from Jilin Province and the Inner Mongolia Autonomous Region in China and deposited in the herbarium Fungarium of Jilin Agricultural University (FJAU).

2.2. Morphological Studies

Field records of fresh basidiomata, photographs of the habitat, and records after drying were used for macromorphological descriptions. The basidiomata’s color description was derived from Kornerup and Wanscher [26]. Using the technique of freehand slicing, utilizing water and 5% KOH solution as the floating carrier, and dyeing with 1% Phloxine B solution, the structures of basidia, basidiospores, hyphidia, and hyphae were examined under an optical microscope (Olympus CX33, Olympus, Tokyo, Japan). Twenty mature microstructures were chosen at random, examined under a microscope, and measured. “a–b × c–d” indicates the measurement of basidia, basidiospores, and hyphidia; “e–f” indicates the measurement of hyphae. Moreover, we determined the basidiospore’s Q value (Q = length/width). The extreme values of length, width, and diameter are denoted by the letters “a–b”, “c–d”, and “e–f”, respectively. For a given species, “n” denotes the ratio of the total number of measured basidiospores to the total number of measured specimens. All data were collected within a measurement range of 90%. The width was measured by choosing the area that was the widest. The apiculum was not taken into account when measuring the length of basidiospores. Along with the macroscopic morphology and microstructure features, a description and line charts are provided.

2.3. DNA Extraction, PCR and Sequencing

To extract DNA from dry specimens, the plant genome extraction kit from Kangwei Century (CWBIO, Beijing, China) was used. The ITS1F/ITS4 [27] and LR0R/LR7 [28,29] primers were used to amplify and sequence the ITS and nrLSU fragments, respectively. Sangon Bioengineering (Shanghai, China) Co., Ltd. was entrusted with the sequencing work, and the PCR reaction procedure was based on the method of Wang and Bau [30].

2.4. Molecular Phylogenetic Analyses

Sequencher 5.4.5 was used to observe the peak maps, further screen and sort the self-test sequences, and perform blast alignment [31]. ITS and nrLSU sequences were downloaded from GenBank based on the alignment outcomes and morphological similarity [32] (Table 1). The ITS and nrLSU sequences were aligned using the GINS-i algorithm with two iterative cycles only, using the online Mafft tool version 7 [33]. The alignment was manually adjusted and trimmed using MEGA 7.0.
The concatenated alignment of ITS (1–646) and nrLSU (647–1248) of Sirobasidium comprised 64 sequences. The concatenated alignment of ITS (1–682) and nrLSU (683–1517) of Dacrymycetaceae comprised 197 sequences. The best-fit model (edge-unlinked) was selected using the BIC criterion with ModelFinder [34]. The best-fit model of Sirobasidium and Dacrymycetaceae, according to BIC, was both SYM + I + G4. With the use of IQ-TREE [35] and the Shimodaira–Hasegawa-like approximate likelihood ratio test [36], maximum likelihood phylogenies were inferred under the edge–unlinked partition model for 10,000 ultrafast [37] bootstraps. The best-fit partition model (edge-unlinked) was again selected using the BIC criterion with ModelFinder. The best-fit model of Sirobasidium according to BIC was SYM + I + G4 (ITS) and K2P + I + G4 (nrLSU). The best-fit model of Dacrymycetaceae according to BIC was SYM + I + G4 (ITS) and GTR + F + I + G4 (nrLSU). Under the partition model, in which the first 25% of sampled data were discarded as burn-in, Bayesian inference phylogenies were inferred using MrBayes 3.2.6 [38]. Fig Tree v1.4.3 [39] and Photoshop 2021 (Adobe, Sam Jose, CA, USA) were then used to visualize the phylogenetic tree.
Table 1. Sequence information from phylogenetic trees.
Table 1. Sequence information from phylogenetic trees.
SpeciesLocalitySample No.GenBank No.References
ITSnrLSU
Bullera albaUSACBS 501TAF444368AF075500[40]
Bullera hannaeUSACBS 8286TAF444486AF363661[40]
Bullera penniseticolaUSACBS 8623TAF444471AF363649[40]
Bullera unicaUSACBS 8290TAF444441AF075524[40]
Calocera bambusicolaChinaWu9910 12FJ195751-[23]
Calocera corneaUSAAFTOL ID 438AY789083AY701526[41]
Calocera corneaSwedenUPS F 940774MN595626MN595626[41]
Calocera corneaGermanyMW 55-AF291302[42]
Calocera corneaCanadaCBS 124 84AB712437AB472738[43]
Calocera furcataRussiaH Spirin 10949MW191975MW159088[43]
Calocera fuscaNew ZealandPDD 107972LC131406LC131365[20]
Calocera glossoidesUkraineCWU6247MW191968MW159084[44]
Calocera luteaNew ZealandPDD 107841LC131413LC131372[20]
Calocera palmataJapanCBS 127 51MH856777MH868295[32]
Calocera pedicellataNew ZealandPDD 107830LC131415LC131374[20]
Calocera sinensisChinaFJAU68949PP749224-This study
Calocera sinensisChinaDai22645OL518939OL518948[32]
Calocera tibeticaChinaDai20171MW549777MW750403
Calocera velutinaChinaFJAU68950TPP776476PP776564This study
Calocera velutinaChinaFJAU68951PP776543PP776565This study
Calocera viscosaJapanTUFC12873AB712439-[43]
Calocera viscosaChinaFJAU68917-PP749235This study
Calocera viscosaJapan175-AB299048[18]
Calocera viscosaGermanyAFTOL ID 1679TDQ520102DQ520102[41]
Calocera viscosaSwedenUPS F 940773MN595628MN595628[41]
Cerinomyces cokeriCanadaTU135089MW191983MW191983[44]
Cerinomyces concretusColombiaO F 919450TMW191933-[44]
Cerinomyces fugaxUSAHHB 8856TMW191905MW159051[44]
Cerinomyces hesperidisUSANY01782362TMW191921MW159065[44]
Cerinomyces inermisNew ZealandPDD87816TMW191887-[44]
Cerinomyces lipoferusNetherlandsENZ20001MZ147626MZ147626[44]
Cerinomyces nepalensisNepalO F 904088MW191896-[44]
Cerinomyces neuhoffiiSwedenUPS F 941020TMN595625MN595625[41]
Cerinomyces pallidusUSAARIZ AN09245MZ147624-[44]
Cerinomyces paulistanusBrazilO Ryvarden 24759TMW191934-[44]
Cerinomyces pinguisNepalO F 904085TMW191907MW159043[44]
Cerinomyces tortusSwedenUPS F 946515MW191999MW191999[44]
Cerinomyces volaticusRussiaLE295748MW191901MW159047[44]
Dacrymyces adpressusJapan554-AB472729[14]
Dacrymyces ancyleusJapanMAFF 241177TAB712448AB472713[43]
Dacrymyces capitatusChinaFJAU68952PP749225PP749236This study
Dacrymyces cerebriformisChinaDai 19832TOM955202OM955197[25]
Dacrymyces chiangraiensisThailandMFLU 16 0572KY498587-[45]
Dacrymyces chrysocomusJapanCBS 280 84AB712451AB712427[43]
Dacrymyces chrysospermusChinaFJAU68956PP749226PP749237This study
Dacrymyces citrinusNew ZealandPDD 107915TLC131417LC131376[20]
Dacrymyces corticioidesUSANY02686162MW191944MW159068[44]
Dacrymyces cylindricusNew ZealandPDD 105052TLC131419LC131378[20]
Dacrymyces cyrtosporusNew ZealandPDD 107980TLC131422LC131381[20]
Dacrymyces dictyosporusJapanHHB 8618AB712454AB712429[43]
Dacrymyces fennicusFinlandH Miettinen 21174MW191957MW159071[44]
Dacrymyces flabelliformisNew ZealandHHB 18308TAB712455AB712430[43]
Dacrymyces grandinioidesKenyaK M 237139TMW191980MW191980[44]
Dacrymyces intermediusNew ZealandPDD 107851-LC131384[20]
Dacrymyces invisibilisChile14597MDTMH230100MH230102[46]
Dacrymyces jauensisChinaFJAU68959PP776532PP776577This study
Dacrymyces jauensisChinaFJAU68960PP776533PP776578This study
Dacrymyces jauensisChinaFJAU68961TPP776534PP776579This study
Dacrymyces lacrymalisJapanTUFC13327AB712456AB299069[43]
Dacrymyces longistipitatusNew ZealandPDD 107997TLC131426LC131387[20]
Dacrymyces microsporusChinaWu561AB712458OL546779[32]
Dacrymyces minorJapanTUFC12836AB712458-[43]
Dacrymyces minutusJapan648AB712459AB472733[43]
Dacrymyces novae-zelandiaeNew ZealandPDD 107953LC131428LC131391[20]
Dacrymyces ovisporusNorwayH Spirin 11145MW191960MW159073[44]
Dacrymyces pachysporusNew ZealandPDD 105004TLC131429LC131392[20]
Dacrymyces parastenosporusNew ZealandPDD 104963TLC131432LC131395[20]
Dacrymyces pezizoidesJapanTNS F 54909LC386890LC386894[47]
Dacrymyces pinacearumJapanUPS F 593533MN595637MN595637[41]
Dacrymyces puniceusChinaFJAU68955PP749227PP749238This study
Dacrymyces san-augustiniiChinaWu544OL587813OL546781[32]
Dacrymyces sinostenosporusChinaDai20003TMW540888MW540890[25]
Dacrymyces stenosporusNew ZealandPDD 105018TLC131433LC131396[20]
Dacrymyces stillatusSwedenUPS F 939814 1MN595677MN595677[41]
Dacrymyces subalpinusJapan228-AB299060[18]
Dacrymyces subantarcticensisNew ZealandPDD 107948LC131435LC131399[20]
Dacrymyces subarcticusJapanHNo 544AB712467-[43]
Dacrymyces variisporusJapanTUFC14203AB712470-[43]
Dacrymyces venustusEthiopiaO Adane 150MW191949MW159075[44]
Dacryomitra pusillaSwedenUPS F 176774MN595639MN595639[41]
Dacryopinax elegansUSAHHB 18731AB712471AB712433[43]
Dacryopinax indacocheaeVenezuelaCRM 72AB712472AB712434[43]
Dacryopinax lowyiMexico27618MN733712MN733723[22]
Dacryopinax manghanensisChinaFJAU68940PP749230PP749241This study
Dacryopinax manghanensisChinaFJAU68938PP749228PP749239This study
Dacryopinax manghanensisChinaFJAU68941PP749231PP749242This study
Dacryopinax manghanensisChinaFJAU68943TPP749233PP749244This study
Dacryopinax manghanensisChinaFJAU68942PP749232PP749243This study
Dacryopinax manghanensisChinaFJAU68939PP749229PP749240This study
Dacryopinax martiniiMexico682MN733715MN733726[22]
Dacryopinax maxidoriiBrazilRLMA306OK257531OK257536[48]
Dacryopinax primogenitusCosta RicaMIN 862738KT251038KT251039[49]
Dacryopinax sp.KenyaH7008759MW191959MW159091[44]
Dacryopinax spathulariaChinaFJAU68908PP749234PP749245This study
Dacryopinax spathulariaUSAH Miettinen 16740MW191973MW159085[44]
Dacryopinax spathulariaIndonesiaH Miettinen 20559MW191976MW159092[44]
Dacryopinax spathulariaChinnaWu 331OL614833OL616184[32]
Dacryopinax sphenocarpaJapan534-AB472726[14]
Dendrodacrys brasilienseBrazilINPA 241458TAB744230AB723514[50]
Dendrodacrys ciprenseCyprusUPS F 946590TOM519385OM519385[51]
Dendrodacrys concrescensSwedenUPS F 946602TOM519390OM519390[51]
Dendrodacrys dendrocalamiJapanTUFC13914AB712453AB712428[43]
Dendrodacrys ellipsosporumSpainUPS F 946604TOM519392OM519392[51]
Dendrodacrys kennedyaePanamaTAAM192134TOP529836OP529830[51]
Dendrodacrys laetumKenyaH7008757TOP529841OP529841[51]
Dendrodacrys oblongisporumNorwayUPS F 946599OM519399OM519399[51]
Dendrodacrys rigoratumJapanTUFC12845TAB712447-[14]
Ditiola haasiiCzech RepublicPRM 944647MW557322MW557322[52]
Ditiola peziziformisFinlandH Haikonen 2426MW191972MW159070[18]
Fellomyces horovitziaeUSACBS 7515T-AF189856[53]
Femsjonia monosporaThailandMFLU 16 0608KY498588-[45]
Femsjonia uniseptataJapanTNS F 54019TNR_164095NG_060435[20]
Fibulobasidium inconspicuumUSACBS 8237TAF444318AF363641[40]
Fibulobasidium murrhardtenseChinaCBS 9109TGU327540AF416648[54]
Fibulobasidium sirobasidioidesGermanyRJB 12787-AF416644[55]
Fonsecazyma mujuensisChinaCBS 10308TKF036595DQ333884[12]
Genolevuria amylolyticaChinaCBS 10048TKF036585AY562134[12]
Genolevuria armeniacaChinaCBS 10050KF036587AY562140[12]
Genolevuria bromeliarumBrazilBI20EU386359DQ784566[56]
Genolevuria tibetensisChinaAS 2 2653TEF363146EF363143[57]
Guepiniopsis alpinaUSAHAY F 002724OR767855-[32]
Guepiniopsis buccinaUSAAFTOL ID 888DQ206986AY745711[41]
Guepiniopsis estonicaSwedenUPS F 940137MN595632MN595632[41]
Heterotextus miltinusNew ZealandPDD 107924LC131438LC131402[20]
Kockovaella imperataeUSACBS 7554TAB054091AF189862[53]
Pseudotremella allantoinivoransUSACBS 9604TAY315664AY315662[58]
Pseudotremella moriformisUSACBS 7810AF444331AF075493[40]
Sirobasidium apiculatumJapanMY62 1LC203425LC203426[13]
Sirobasidium apiculatumJapanMY62 4LC203428LC203429[13]
Sirobasidium brefeldianumSpainAM71JN053472JN043578[9]
Sirobasidium brefeldianumUSACBS 7805AF444330AF075492[40]
Sirobasidium japonicumJapanMY111 05LC203420LC016573[13]
Sirobasidium japonicumJapanMY111 09LC203422LC203423[13]
Sirobasidium jilinenseChinaFJAU68667PP749214-This study
Sirobasidium jilinenseChinaFJAU68670TPP749217PP749222This study
Sirobasidium jilinenseChinaFJAU68668PP749215PP749220This study
Sirobasidium jilinenseChinaFJAU68669PP749216PP749221This study
Sirobasidium jilinenseChinaFJAU68672PP749219-This study
Sirobasidium jilinenseChinaFJAU68671PP749218PP749223This study
Sirobasidium magnumSpainAM70JN053497JN043603[9]
Sirobasidium magnumNetherlandsCBS 6803AF444314KY109657[40]
Sirobasidium magnumNetherlandsCBS 6964KY105421KY109658[59]
Sirobasidium magnumChinaFJAU68996PP741731PP741733This study
Sirobasidium magnumChinaFJAU68997PP741732PP741734This study
Tremella giraffaGermanyCCJ 1553AF042453AF042271[60]
Note: “-” denotes the absence of pertinent genetic data. Sequences newly generated in this study are indicated in bold. The type specimens are those designated with a “T”.

3. Results

3.1. Phylogenetic Analyses

This study generated a total of 46 new sequences, including 24 ITS sequences and 22 nrLSU sequences. These new sequences have been uploaded to GenBank. The multi-locus dataset (ITS + nrLSU) of Sirobasidium had an aligned length of 1248 total characters), including gaps. The multi-locus dataset (ITS + nrLSU) of Dacrymycetaceae had an aligned length of 1517 total characters, including gaps. Only the topological structures of Bayesian inference are displayed, as the topological structures of ML and BI are very similar. Bootstrap support (BS) values ≥ 75% and Bayesian posterior probability (PP) values ≥ 0.75 are indicated on branches (BS/PP) (Figure 1 and Figure 2).
Figure 1. Phylogeny of Sirobasidium and related species by Bayesian inference based on the ITS + nrLSU dataset. New species are indicated in bold.
Figure 2. Phylogeny of Dacrymycetaceae by Bayesian inference based on the ITS + nrLSU dataset. New species are indicated in bold.
From the phylogenetic tree of Sirobasidium and its related species constructed with Fellomyces horovitziae Spaaij, G. Weber & Oberw. and Kockovaella imperatae Nakase, I. Banno & Y. Yamada as outgroups (highlighted in blue), it can be seen that Sirobasidium is polyphyletic (Figure 1). High support rates (PP = 1.00, MLbs = 100%) were attained by the six specimens of the new species Sirobasidium jilinense T. Bau et X. Wang sp. nov., which were grouped in an independent branch. S. jilinense and S. magnum share a sister relationship and received high support (PP = 1.00, MLbs = 100%). The novel species could also be easily distinguished from other species, thanks to its distinct placement in the phylogenetic tree (Figure 1).
From the phylogenetic tree of Dacrymycetaceae constructed with Cerinomycetaceae as an outgroup (highlighted in blue), it can be seen that Dacrymyces, Dacryopinax, and Calocera are polyphyletic. The species belonging to each genus are dispersed across different branches of Dacrymycetaceae (Figure 2).
The two specimens of the recently discovered species Calocera velutina T. Bau et X. Wang sp. nov. were grouped into a separate branch and had a high support rate (PP = 1.00, MLbs = 100%). C. velutina forms a sister clade with C. cornea (Batsch) Fr. and C. furcata (Fr.) Fr., which is highly supported (PP = 1.00, MLbs = 100%). At the same time, the new species can be distinguished from other species in the genus in the phylogenetic tree (Figure 2).
A high support rate (PP = 1.00, MLbs = 100%) was obtained for the three specimens of the new species, Dacrymyces jauensis T. Bau et X. Wang sp. nov. They were grouped into an independent branch. D. jauensis forms a sister relationship with the clade consisting of D. chiangraiensis Ekanayaka, Karun., Q. Zhao & KD Hyde, and D. san-augustinian Kobayasi and gained a lower support rate (PP = 0.75, MLbs = 78%). This suggests that other species among them have not yet been identified. The new species and other species could also be well distinguished in the phylogenetic tree (Figure 2).
The six specimens of the new species Dacryopinax manghanensis T. Bau et X. Wang sp. nov. were clustered into an independent branch and obtained a high support rate (PP = 1.00, MLbs = 100%). It is a sister clade to Dacryopinax sp. (H7008759), with a high support rate (PP = 1.00, MLbs = 100%), and which is well distinguished from other species.

3.2. Taxonomy

Sirobasidium jilinense T. Bau et X. Wang, sp. nov. (Figure 3 and Figure 4)
Figure 3. Basidiomata of Sirobasidium jilinense: (a) fresh (FJAU68670); (b) fresh (FJAU68668); (c,d) dry (FJAU68667). Scale bars: (a,b) = 1 cm; (c,d) = 0.5 cm.
Figure 4. Sirobasidium jilinense T. Bau et X. Wang, sp. nov. (a) Basidiomata; (b) Epibasidia; (c) Basidiospores; (d) Hyphae; (e) Hypobasidia; (f) Probasidia. (Scale bars: (a) = 1 cm; (b,e,f) = 10 μm; (c,d) = 5 μm).
MycoBank number: 853841
Diagnosis: Sirobasidium jilinense is characterized by cerebriform basidiomata, which are reddish brown to dark reddish brown when fresh and dark brown when dry. The hypobasidia consist of three to nine chains and are divided vertically or obliquely into four cells, whereas the basidiospores are subglobose to broadly ellipsoid, and the hyphae are thick-walled.
Etymology: “jilinense” refers to the discovery of a type specimen in Jilin Province, China.
Type: CHINA. Jilin Province, Jilin, Jiaohe, Shansong Ridge, on rotten wood of Quercus, elev. 577.2 m, 43°75′ N, 127°33′ E, 26 August 2023, T. Bau and Xia Wang, (FJAU68870, holotypus!). Same location, 29 July 2023, Xia Wang, (FJAU68668, paratypus!).
Basidiomata are soft gelatinous when fresh, easily rotting, cerebriform, reddish brown (8E5–E8) to dark reddish brown (8F5–F8), usually remaining coalescing, occasionally separate, 1–3 cm long and 1 cm thick. The hymenium surface is smooth, with full edges, folded, and relatively blunt. Shrinkage when dry, dark brown (8F1–F5), fragile.
Cross section without medulla, composed of the hymenium and intertwined jelly hyphae. Hymenium peripheral, pale yellow, composed of probasidia and hypobasidia. Probasidia 12–20 × 11–16 μm, globose to ellipsoid, light yellow, containing oil droplet-like substances, with clamp connection at the base; hypobasidia, 14–22 × 10–15 μm, ellipsoid to ovoid, the vertical or diagonal partitions are 4-cells, 3–9 chain-like structures, wrinkled at maturity, light yellow, containing oil droplet-like substances, with clamp connection at the base; epibasidia, 13–18 × 5–8 μm, spindle-shaped, detached, containing oil droplet-like substances. Basidiospores 6.7–9.4 × 6.7–7.2 μm, Q = 1.04–1.37 (n = 100/5), subglobose to broadly ellipsoid, apiculate, colorless, smooth, with oil droplets, germinating to produce regenerated spores or germination tubes. Hyphae with rich clamp connection, thick-walled, 2.3–3.6 µm in diameter.
Habitat: In summer and autumn, it grows on fallen trees or branches such as Acer and Quercus in broad-leaved forests.
Distribution: Currently, only known in the Jilin Province, China, Asia.
Additional specimens examined: China. Jilin Province: Jilin, Jiaohe, Shansong Ridge, on fallen branches of Acer, July 24, 2022, Liyang Zhu, FJAU68667 (Z22072428). Same location, Shien Wang, FJAU68669 (E2307209); Jilin, Huandian, Hongshi National Forest Park, on fallen branches of Acer, August 27, 2023, T. Bau & Xia Wang FJAU68671 (W23082706), T. Bau & Mu Liu, FJAU68672 (lm230890).
Calocera velutina T. Bau et X. Wang, sp. nov. (Figure 5 and Figure 6)
Figure 5. Basidiomata of Calocera velutina: (a) fresh (FJAU68950); (b,c) dry (FJAU68951, FJAU68950). Scale bars: (a–c) = 5 mm.
Figure 6. Calocera velutina T. Bau et X. Wang, sp. nov. (a) Basidiomata; (b) Basidiospores; (c) Basidia and hyphidia; (d) Central hyphae. (Scale bars: (a) = 1 cm; (b) = 5 μm; (c,d) = 10 μm).
MycoBank number: 854036
Diagnosis: Calocera velutina is characterized by stipitate and pileate when young, needle-like when mature, light yellow, with a white disc-shaped villus at the base, simple or dichotomously branched. Hyphae thick-walled, without clamp connection. In summer, they grow on decaying trees in coniferous forests.
Etymology: “velutina” refers to the base of the basidiomata with villus.
Type: CHINA. Jilin Province, Yanbian Korean Autonomous Prefecture, Antu County, Erdaobaihe Town, Forest Industry Cultural Park, on rotten wood of Pinus L., elev. 708 m, 42°43′ N, 128°12′ E, 31 July 2022, T. Bau and Xia Wang, (FJAU68950, holotypus!). Jilin Province, Huadian City, Hongshi National Forest Park, 27 August 2023, T. Bau and Mu Liu, (FJAU68951, paratypus!).
Basidiomata are gelatinous when fresh, stipitate and pileate when young, needle-like when mature, light yellow (4A4–A5), slightly lighter at the tip, with white (4A1) disc-shaped villus at the base, simple or dichotomously branched, up to 15 mm high. The hymenium is located in the upper middle part, and the base is sterile. When dry, it is greyish yellow (4B3–B6, 4C5), and the surface under the body mirror is fine and granular.
The transverse section of the apex of the basidiomata is composed of the hymenium and interwoven hyphae. The transverse section of the basidiomata cylinder presents three annular zones. The central hyphae are densely arranged vertically, surrounded by loosely interlaced hyphae, and the periphery is the hymenium. Hymenium amphigenous is composed of probasidia, basidia, and hyphidia. Probasidia 18–25 × 2.9–4.4 μm, sub-clavate, forked when mature, containing oil droplet-like substances, with septate at the base; basidia 21–33 × 3.5–4.8 μm, clavate, containing oil droplet-like substances, with septate at the base; sterigmata 10–17 × 2.3–3.7 μm, cylindrical, the top gradually becomes pointed and gradually weakens when mature, containing oil droplet-like substances. Hyphidia 19–28 × 1.8–2.7 μm, narrowly cylindrical, simple or slightly curved, with septum at the base. Basidiospores 9.1–10.4 × 4.2–5.3 μm, Q = 1.90–2.27 (n = 40/2), curved cylindrical, narrow top, smooth, thin-walled, mature 1 septate, septate thin-walled, contains oil droplet-like substances. No germination was observed. Central hyphae 4.9–6.8 μm in diameter, septate, slightly rough, thick-walled. Interlaced hyphae, 3.4–5.3 μm in diameter, septate, thick-walled, basal hyphae 2.5–3.8 μm in diameter. All structures without clamp connection.
Habitat: In summer, it grows on decaying trees in coniferous forests.
Distribution: Currently, only known in the Jilin Province, China, Asia.
Dacrymyces jauensis T. Bau et X. Wang, sp. nov. (Figure 7 and Figure 8)
Figure 7. Basidiomata of Dacrymyces jauensis: (a) fresh (FJAU68961); (b) dry (FJAU68961). Scale bars: (a) = 1 cm, (b) = 0.5 cm.
Figure 8. Dacrymyces jauensis T. Bau et X. Wang, sp. nov. (a) Basidiomata; (b) Probasidia; (c) Basidiospores; (d) Conidia; (e) Basidia. (Scale bars: (a) = 1 cm; (b–d) = 10 μm; (e) = 20 μm).
MycoBank number: 854037
Diagnosis: Dacrymyces jauensis is characterized by basidiomata flat to cushion-shaped when young, cinnamon, golden yellow to brownish yellow; basidiospores sausage-shaped, separated horizontally when mature, usually divided into 7 septate; germination produces rod-shaped conidia; hyphidia absent.
Etymology: “jauensis” refers to the discovery of a type specimen at Jilin Agricultural University.
Type: CHINA. Jilin Province, Changchun city, wild plantation garden of the campus of Jilin Agricultural University, grows on highly decayed wood of broad-leaved trees in broad-leaved forests, 43°81′ N, 125°40′ E, 235 m, 16 July 2022, Xia Wang & Tolgor Bau, (FJAU68961, 22071605W, holotypus!). Same location, 4 July 2022, Xia Wang & Tolgor Bau, (FJAU68959, T22070437W; FJAU68960, T22070438W, paratypus!).
Basidiomata are soft gelatinous when fresh, flat to cushion-shaped when young, cinnamon, golden yellow (4C6), up to 0.5 cm, usually remaining coalescing when mature, brownish yellow (5D6, 5E5), sessile, up to 2.5 cm. Hymenium surface with wrinkles and grooves, edge curved, folded, wavy, blunt. When dry, it shrinks and collapses in a brown (6E6), making it difficult to observe.
The transverse section is composed of hymenium and context hyphae. Hymenium amphigenous. Probasidia 32–47 × 3.7–6.3 μm, cylindrical, subclavate to broadly clavate, thin-walled, transparent, forked when mature, containing oil-dripping substance, basally septate; sterigmata 11–33 × 3.0–4.4 μm, cylindrical, apically acuminate when mature, gradually weakening. Hyphidia absent. Basidiospores 15.9–20.9 × 5.1–6.3 μm, Q = 2.80–3.49 (n = 90/3), sausage-shaped, smooth, thin-walled, transparent, separated horizontally when mature, usually divided into 7 septate, occasionally 0–3 septate, thin-walled, containing oil drop-like substance, and conidia germinate at the septate. Conidia 4.8–12.3 × 1.4–1.8 μm, long cylindrical to rod-shaped. Context hyphae 1.6–3.2 μm in diameter, thin-walled, septate, septate not swollen. All structures without clamp connection.
Habitat: In summer, it grows on highly decayed wood of broad-leaved trees in broad-leaved forests.
Distribution: Currently, only known in the Jilin Province, China, Asia.
Dacryopinax manghanensis T. Bau et X. Wang, sp. nov. (Figure 9 and Figure 10)
Figure 9. Basidiomata of Dacryopinax manghanensis: (a) fresh (FJAU68943); (b) fresh (FJAU68940); (c) fresh (FJAU68941); (d) dry (FJAU68938). Scale bars: ((a–c) = 0.5 cm; (d) = 0.2 cm).
Figure 10. Dacryopinax manghanensis T. Bau et X. Wang. (a) Basidiomata; (b) Probasidia and hyphidia; (c) Basidiospores; (d) Basidia; (e) Marginal hyphae. ((a) = 0.5 cm; (b) = 5 μm; (c) = 2 μm; (d,e) = 10 μm).
MycoBank number: 853843
Diagnosis: Dacryopinax manghanensis is characterized by small basidiomata, 1–5 mm wide, 1–8 mm high, spathulate and stipitate to substipitate, light yellow, cross section without medulla, the short and fine villi on the sterile surface. Basidiospores curved cylindrical to navicular, with 0–1 septate at maturity, thin-walled septate, and no germination observed; cortical hyphae curved, thick-walled, swollen, and branched.
Etymology: “manghanensis” is a Mongolian word for “sandy land”, which means that type specimens are collected from rotten wood in sandy land.
Type: CHINA. Inner Mongolia Autonomous Region, Tongliao, in the later stage of the left wing of Horqin, Wudantara Forest Farm, fallen branches of broad-leaved trees, 43°01′ N, 122°44′ E, 363 m, 16 July 2022, Weinan Hou & Tolgor Bau, (FJAU68943, H2207159, holotypus!). Same location, 15 July 2023, Hong Cheng & Tolgor Bau, (FJAU68940, C2371524, paratypus!).
When the basidiomata are fresh, soft gelatinous, spathulate, and stipitate to substipitate when young, the stipitate is light yellow (4A4–A5) and slightly hyaline, petaloid when mature, surface with longitudinal ridges, 1–5 mm wide, and 1-8 mm high. Hymenial surface smooth, with entire edges, blunt. The short and fine villi on the sterile surface are visible under a stereomicroscope. When dry, it shrinks, with a white (4A1) base, a dark yellow (4C8) surface on the hymenial, and obvious longitudinal edges on the stipitate.
Cross section without medulla, composed of the hymenium, context hyphae, and cortical hyphae. Hymenium unilateral, occasionally bilateral, composed of basidia and hyphidia. Probasidia 18–27 × 2.5–3.5 μm, cylindrical to clavate, thin-walled, hyaline, forked when mature, containing oil droplet-like substances, with septate at the base; sterigmata 7–13 × 1.8–2.9 μm, the top gradually becomes pointed and gradually weakens when mature. Hyphidia 15–27 × 1.4–2.2 μm, narrow cylindrical, thin-walled, hyaline, simple or slightly curved, with 1 septum at the base. Basidiospores 7.9–9.0 × 3.7–4.9 μm, Q = 1.73–2.07 (n = 80/4), curved cylindrical to navicular, apiculate, smooth, thin-walled, hyaline, mature 0–1 septate, septate thin-walled, contains oil droplet-like substances. No germination was observed. Cortical hyphae 4.5 μm in diameter, cylindrical branching at the end, curved, septum, constriction at intervals, thick-walled. All structures without clamp connection.
Habitat: In summer and autumn, it decays on fallen branches in broad-leaved forests or mixed coniferous and broad-leaved forests.
Distribution: Currently, it is only distributed in China, Inner Mongolia Autonomous Region, and Jilin Province.
Additional specimens examined: CHINA. Jilin Province, Tonghua, Yuhuangshan Park, broad-leaved forest rotting wood, 14 August 2022, Xia Wang, FJAU68941 (2281405W); Changchun, Jilin Agricultural University Campus Back Mountain, 16 September 2022, Xia Wang, FJAU68942 (2291601W); Tonghua, Yushan Park in Ji’an City, 6 July 2023, Zhengqing Chen, FJAU68938 (Q237603); Changchun, Jingyue Pool National Forest Park, 12 July 2023, Xia Wang, FJAU68939 (W23071206).

4. Discussion

Sirobasidium jilinense is easily confused with the type species S. sanguineum in the wild; however, the hypobasidia of the latter are solitary or in chains of 2–4, without a clamp connection at the base, and the clamp connection of hyphae is enlarged (5.5–8 μm). S. jilinense and S. rubrofuscum (Berk.) P. Roberts are similar; the basidiomata are all cerebriform and dark reddish brown, but they can be distinguished by the number of hypobasidia in the chains and the size of the basidiospores. Other species in Sirobasidium are white, off-white, or yellowish and are easily distinguished from S. jilinense (Table 2). Due to its brain-like, reddish-brown basidiomata, S. jilinense is easily confused with individual species of Tremella Pers. and Exidia Fr. in the wild, but it is easily distinguished from the latter under a microscope according to the basidium morphology.
Table 2. Morphological comparison of Sirobasidium species.
Molecular phylogeny analysis showed that members of Fibulobasidium Bandoni and Sirobasidium jilinense, S. magnum, S. apiculatum, and S. japonicum form a separate branch (PP = 0.99, MLbs = 91%), while S. brefeldianum is located in another branch. This suggests that Sirobasidium is polyphyletic, which is consistent with the results of recent studies [9,10,11,13] (Figure 1). The new species S. jilinense and S. magnum have a sister relationship and obtained a high support rate (PP = 1, MLbs = 100%), which is consistent with the results of morphological studies.
Calocera velutina is characterized by basidiomata that are stipitate and pileate when young, needle-like when mature, yellow with a white disc-shaped villus at the base, and simple or dichotomously branched. The hyphae are thick-walled, without clamp connection. In summer, they grow on decaying trees in coniferous forests. In the wild, C. velutina is easy to be confused with C. cornea (Batsch) Fr., C. sinensis McNabb, and C. tibetica F. Wu, L.F. Fan & Y.C. Dai. They all have fresh yellow basidiomata with sharp, unbranched, or even-branched cylindrical upward-narrowing tops; however, the base of the latter three does not have a white disc-shaped villus [7,23,24]. The basidiomata of C. sinensis are small (1–5 mm), and the hyphae have clamp connections [23]. The basidiospores of C. tibetica are divided into three transverse septates when mature [24]. In the phylogenetic tree, they were also found to have a distant relationship.
Dacrymyces jauensis is characterized by basidiomata flat to cushion-shaped when young, cinnamon, golden yellow to brownish yellow; basidiospores sausage-shaped, separated horizontally when mature, usually divided into seven septate; germination produces rod-shaped conidia; hyphidia absent. Dacrymyces jauensis, Dacrymyces san-augustinii Kobayasi, and Dacrymyces chiangraiensis Ekanayaka, Karun., Q. Zhao & K.D. Hyde present multiple transverse septates when basidiospores mature. In the phylogenetic tree, a close relationship can be observed (Figure 2). However, the basidiomata of Dacrymyces san-augustinii are yellow, and the basidia (38–58 × 5.5–7 µm) and basidiospores (16–27.5 × 6–10 µm) are larger, with hyphidia [14]. The basidiomata of Dacrymyces chiangraiensis are yellow to orange, and the basidia (32–53 × 7–10 µm) and basidiospores (19–23 × 6.5–8 µm) are also large, with hyphidia [45].
There are two kinds of basidiomata of Dacryopinax manghanensis, most of which are clustered and spoon-shaped in the sand of the Inner Mongolia Autonomous Region but petal-shaped in Jilin Province (Figure 9). However, their microstructure is the same, as they are clustered in the same branch in the phylogenetic tree, and the shape of the basidiomata may be related to the environment. There are seven species of Dacryopinax distributed in China. The mature basidiospores of Dacryopinax aurantiaca and Dacryopinax spathularia are 0–1 septate, and they are easily confused with Dacryopinax manghanensis in the wild. However, the basidiospores of Dacryopinax aurantiaca are longer (10.4–13 × 3.5–5 μm) [7]. Basidiospores germinate as conidia or germ tubes and are mainly distributed in South China. Dacryopinax spathularia had obvious villi (visible to the naked eye when dry), with medulla on the transverse section and a bulbous hyphae septum [7]. In the phylogenetic tree, they are not closely related (Figure 2). In addition, the basidiospores of Dacryopinax indacocheae Lowy, Dacryopinax primogenitus, and Dacryopinax sphenocarpa Shirouzu & Tokum. are also divided into 0–1 septate at maturity. However, the basidiomata of Dacryopinax indacocheae are yellowish brown and larger (30 mm wide, 20 mm high). The cortical hyphae of the stipe have chain-like enlarged cells, the basidiospores are longer (8–11.5 × 3–3.5 μm), and the hymenium has abundant conidia [15]. Dacryopinax primogenitus has longer basidia (23–49 × 2.5–4 μm) and longer hyphidia (45.5–48.5 × 2.5–3 μm) and germinates as conidia or germ tubes [49]. The basidiomata of Dacryopinax sphenocarpa are yellow-white or light amber, the hyphae have clamp connections, the base of the probasidia has clamp connections, and the basidiospores are larger (10–16 × 4.5–8.5 μm) [14]. Dacryopinax is polyphyletic, with a total of eight species currently supported by molecular data, located in five distinct clades, consistent with recent findings [6,13,49]. In addition, in the phylogenetic tree constructed in this study, the branch composed of Dacryopinax manghanensis and Dacryopinax sp. (H7008759) was closely related to the branch composed of Calocera species (Figure 2); however, the species of Dacryopinax manghanensis and Calocera were different in morphology, so, it was placed in Dacryopinax.
Regarding jelly fungi, due to their gelatinous texture, researchers tend to focus more on damp environments when collecting specimens. In addition to adding three new species from the humid environment of Jilin Province—S. jilinense, C. velutina, and Dacrymyces jauensis—this study also added a new species from the Horqin Sandy Land—Dacryopinax manghanensis. This shows that jelly fungi have a wide geographical distribution and are well-adapted to different conditions and environments. As such, this type of fungus is worthy of further exploration.

Author Contributions

Conceptualization, T.B. and X.W.; methodology, X.W.; software, X.W.; validation, X.W. and T.B.; formal analysis, X.W.; investigation, X.W. and T.B.; resources, X.W. and T.B.; data curation, X.W. and T.B.; writing—original draft preparation, X.W.; writing—review and editing, X.W. and T.B.; visualization, X.W. and T.B.; supervision, T.B.; project administration, T.B.; funding acquisition, T.B. All authors have read and agreed to the published version of the manuscript.

Funding

This study was supported by the Ministry of Education Innovation Team (IRT-15R25).

Institutional Review Board Statement

Not applicable.

Data Availability Statement

All the sequences have been deposited in GenBank (https://www.ncbi.nlm.nih.gov, accessed on 1 May 2024) and MycoBank (https://www.mycobank.org, accessed on 17 May 2024). The data presented in this study are deposited in the Zenodo repository, accession number doi: 10.5281/zenodo.11206684.

Acknowledgments

We appreciate the help of our team members during the field collection process. We thank the reviewers and responsible editors for their corrections and suggestions.

Conflicts of Interest

The authors declare no conflicts of interest.

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