Phytotaxa 575 (2): 140–148
Copyright © 2022 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
140 Accepted by Samantha C. Karunarathna: 5 Nov. 2022; published: 8 Dec. 2022
Russula rubiginosus sp. nov. in Russula subsect. Maculatinae from Heilongjiang
Province, Northeast China
RUI-QING JI1,2,5#, MENG-LE XIE1,6#, JI-JIANG ZHOU1,7, LI-PENG MENG3,8, YU LI1,9, ZHONG-HUI ZHANG3* &
1Engineering Research Center of Edible and Medicinal Fungi, Ministry of Education, Jilin Agricultural University, Changchun 130118,
2Key Laboratory of Edible Fungus Resources Utilization in North China, Ministry of Agriculture and Rural Affairs, Changchun 130118,
Jilin Province, P. R. China
3Wood Research Institute, Jilin Forestry Science Institute, Changchun-130032, Jilin Province, P. R. China
4Key Laboratory of Vegetable Germplasm Innovation and Utilization of Hebei, Collaborative Innovation Center of Vegetable Industry in
Hebei, College of Horticulture, Hebei Agricultural University, Baoding 071001, China
#These authors contributed equally to this work
Russula rubiginosus sp. nov. (Subsect. Maculatinae) associated with Quercus mongolica is described and illustrated based
on phylogenetic analyses of internal transcribed spacer (ITS) ribosomal DNA sequences and morphological evidence. This
species is characterized by the rusty spots on the pileus and stipe, an acrid taste, a yellow spore print, smaller basidiospores,
and abundant cystidia in pileipellis and stipitipellis. Furthermore, the morphological characteristics were described in detail
and compared with R. globispora and R. subrubescens.
Keywords: Basidiomycota, new taxon, phylogeny, Russulales, SEM, taxonomy
Russula Pers. is the largest genus of Russulaceae. There are 3080 Russula epithets belonging to more than 1331
accepted species in Index Fungorum (Catalogue of Life et al. 2021, Index Fungorum 2022). Before 2015, more than
170 species of Russula have been reported in China (Song et al. 2007, Li et al. 2015). In the following years, more than
50 species of Russula have been recorded (Li et al. 2020; Wang et al. 2021a, Wang et al. 2021b; Li & Bau 2022; Zhou
et al. 2022). Hence, these researches showed that the diversity of Russula in China is high, however, many species rich
areas have not yet been investigated.
Russula is characterized by the brightly colored pileus and amyloid ornamentation basidiospores (Romagnesi
1967, Sarnari 1998, 2005). Russula subsection Maculatinae Romagnesi (1962) is currently considered as a well-defined
section with striking macro-morphological characteristics. The group was originally proposed as Maculatae by Konard
& Josserand (1934). Romagnesi (1967) redefined this group to Russula section Maculatinae with the characteristic
morphology of the mainly reddish pileus, yellowish basidiospore print and acrid-tasting context. Sarnari (1998)
grouped member of Maculatinae species in subsection Urentes Maire, series Maculata (1910: 122), it is characterized
by a yellow-brownish context discolouration (and a spotted pileus surface), a fairly dark spore print, large spores
and pileocystidia with a few septa. Furthermore, several researchers have been introduced detailed morphological
descriptions for members of R. subsect. Maculatinae and provided strong and precise phylogeny support (Adamčík &
Jančovičová 2012, 2013, Adamčík et al. 2016).
SPECIES OF RUSSULA RUBIGINOSUS SP. NOV. Phytotaxa 575 (2) © 2022 Magnolia Press • 141
In China, Li (2014) and Li et al. (2018) identified and classified four species as the members of sect. Maculatinae
of Russula subgen. Insidosula followed the classification system of Romagnesi (1967). Recently, Kiran et al. (2021)
provided the mostly detailed phylogenetic analyses of subsect. Maculatinae, including two complex groups, R.
globispora lineage and R. maculata lineage, with a total of 13 species, which were consistent with the results of
Adamčík et al. (2016).
During the field visits in the past few years, many specimens of Russula have been obtained in Northeast China.
Here, we describe a new species of subsect. Maculatinae found in Quercus mongolica forests. The morphological
concepts of this new species were fully supported by molecular data.
FIGURE 1. Phylogram generated by Maximum Likelihood (RAxML) analysis based on nrDNA ITS sequence data. The ML bootstrap
values (ML) ≥ 70% and Bayesian posterior probabilities (BPP) ≥ 0.90 are given on the branches (ML/BPP). The tree is rooted with R.
cuprea and R. juniperina. New species is marked with bold text.
Materials and methods
The specimens were collected from Quercus mongolica forests in the Northeast of China. Fresh basidiomata were
photographed under daylight in the field. The ecological information was recorded in their habitat, and the macroscopic
characteristics were recorded from fresh specimens. Color terms and codes followed Ridgway (1912). The microscopic
characteristics were studied based on hand-cut sections of dried sporocarps, which were mounted on slides in 3% KOH,
Melzer’s reagent and Congo red. Basidiospores, basidia, pleurocystidia, cheilocystidia, pileocystidia and caulocystidia
measurements were given as length × width. The statistics of basidiospore size were based on measurements from 20 to
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40 basidiospores and the abbreviation rules followed Li et al. (2012) and Li (2014). Basidiospore dimensions exclude
ornamentation. The voucher specimens were deposited in the Herbarium of Mycology of Jilin Agricultural University
(HMJAU) and the Fungarium of the Institute of Microbiology, Chinese Academy of Sciences (HMAS). Scanning
electron microscope (SEM) photographs were taken with a Hitachi SU8010 Cold Field Microscope (Engineering
Research Center of Edible and Medicinal Fungi, Ministry of Education, Jilin Agricultural University).
FIGURE 2. Basidiomata of Russula rubiginosus. A, B (Holotype, HMJAU58933; C, D (Paratype, HMJAU58934); Scale bars: 1 cm.
DNA extraction, PCR amplification and DNA sequencing
Genomic DNA was extracted from dried specimens by a Cetyltrimethylammonium Bromide (CTAB) method (Zhao et
al. 2011). Polymerase Chain Reaction (PCR) used 25 μL reaction system. PCR primers used for the amplification of
the nrDNA internal transcribed spacer (ITS) region were ITS1 and ITS4 (White et al. 1990). The PCR reactions were
run under the following conditions: 95 ℃ for 5 mins, followed by thirty cycles of 95 ℃ for 40 s, 50 ℃ for 50 s, 72 ℃
for 1 min 20 s, and a final extension step at 72 ℃ for 10 mins before storage at 4 ℃. The PCR products were sequenced
by Sangon Biotech, Shanghai, China. The newly obtained sequences were submitted to GenBank (https://www.ncbi.
The newly generated ITS sequences were tested with blastn in NCBI. Bayesian Inference (BI) and Maximum Likelihood
(ML) methods were implemented for the phylogenetic analyses involving in totally 42 ITS sequences. The sequence
data refer to Caboň et al. (2019).
Sequences were aligned at both ends using Bioedit 7.0.5 (Hall 1999) and Clustal X 1.83 (Thompson et al. 1997).
The best substitution model HKY+G was determined using the Akaike Information Criterion (AIC), implemented in
MrModeltest 2.3 (Nylander 2004). Bayesian Inference analysis was operated in MrBayes 3.2.7a (Huelsenbeck 2012).
Markov Chains Monte Carlo (MCMC) chains were run for 500,000 generations, sampling every 100th generation
at which point the average standard deviation of split frequencies was 0.006373. A 25% of trees were discarded as
burn-in before the construction of a 50% majority rule consensus tree. Maximum Likelihood analysis was carried out
in raxmlGUI 1.5b2 by choosing GTRGAMMA as the model of nucleotide evolution with 1000 bootstrap iterations
(Silvestro & Michalak 2012). Phylogenetic topologies were exported into figures by FigTree 1.4.4.
SPECIES OF RUSSULA RUBIGINOSUS SP. NOV. Phytotaxa 575 (2) © 2022 Magnolia Press • 143
The new ITS sequences were submitted to GenBank with the accession number OL828348 (Holotype) and OL828349.
The results of ML and BI analyses did entirely agree with their topology. The obtained phylogenetic tree showed that
all species were grouped in two well-supported subclades: R. maculata lineage and R. globispora lineage. The R.
maculata lineage (95%/1) contains three well supported terminal clades that include R. maculata, R. mansehraensis
and R. nympharum. The R. globispora lineage (100%/1) was in a large part polytomy showing only good support at
some terminal nodes, which correspond to the nine species: R. abbottabadensis, R. globispora, R. heilongiangensis,
R. mansehraensis, R. mattiroloana, R. mediterraneensis, R. quercus-floribundae, R. rubiginosus and R. subrubescens.
Russula rubiginosus (100%/1) formed a well-supported clade in R. globispora lineage.
Russula rubiginosus R. Q. Ji, M.L. Xie & J. J. Zhou, sp. nov., Figures 3
FIGURE 3. Microscopic structures of Russula rubiginosus (HMJAU58933 holotype). (A) Basidiospores in Melzer’s reagent. (B)
Cheilocystidia. (C) Basidia. (D) Hyphal terminations of the pileus. (E) Pleurocystidia. (F) Caulocystidia. (G) Pileocystidia. Bars: A = 2
µm, B–G = 5 µm.
Diagnosis:—Differs from Russula subrubescens by small-sized basidiospores and large-sized cystidia.
Etymology:—Refers to the rusty spots on the cap and stipe of basidiomata.
Holotype:—CHINA. Heilongjiang Province: Wudalianchi City, ASL 300 m, 126°11’E, 48°37’N, 13 August 2018,
Peng-Jie Xing & Yang Xu (HMJAU58933, GenBank OL828349).
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TABLE 1. Taxa, voucher and GenBank numbers of ITS sequences used in phylogenetic tree. New species is marked with
Taxa Voucher Locality GenBank number
Russula abbottabadensis LAH310099 Pakistan MG386702
R. abbottabadensis (Type) FH00304589 Pakistan MG386704
R. cuprea 2010 BT168 (GENT) Germany MG386711
R. dryadicola AT2004140 (UPS) Sweden KU928145
R. dryadicola TURA152390 Finland MG386712
R. dryadicola TURA151632 Finland KU928146
R. globispora HK12021 (SAV) Sweden KU886595
R. globispora FH-2007-BT121 (GENT) Germany KU886594
R. globispora FH-2007-BT111 (GENT) Germany KU928144
R. globispora FH-2007-BT98 (GENT) Germany MG386717
R. heilongiangensis (Type) HMAS255142 China MG719932
R. heilongiangensis HMAS279587 China MG719933
R. juniperina F-4998 (SAV) Italy MG386714
R. maculate F-933 (SAV) Slovakia KU928150
R. maculata F-2130 (SAV) Estonia KU928151
R. maculata FH-2009-BT05 (GENT) Germany KU928155
R. maculata FH-2010-BT184 (GENT) Germany KU928154
R. mansehraensis HUP-SU33 Pakistan MG948635
R. mansehraensis (Type) HUP-SUR180 Pakistan MG948636
R. mansehraensis HUP-SU803 Pakistan MG948637
R. mansehraensis MAM-0077 (LAH) Pakistan KU886598
R. mattiroloana JMV800644 (BCN) Greece MK105653
R. mattiroloana JMV800713 (BCN) Poland MK105656
R. mediterraneensis (Type) GK6710 Greece MK105660
R. mediterraneensis MG630 Italy MK105661
R. mediterraneensis 29086 (MCVE) Italy MK105662
R. nympharum HJB10019 Belgium DQ422015
R. nympharum (Type) RUS11121505(GENT) Spain KU928157
R. quercus-floribundae (Type) LAH36219 Pakistan MN053395
R. quercus-floribundae LAH36220 Pakistan MN053391
R. rubiginosus (Type) HMJAU58933 China OL828349
R. rubiginosus HMJAU58934 China OL828348
R. rubiginosus QL041 China HM105533
R. rubiginosus HMAS267826 China KX441111
R. rubiginosus HMAS267741 China KX441095
R. rubricolor LAH35070 Pakistan MK142795
R. rubricolor (Type) LAH35071 Pakistan MK142798
R. rubricolor LAH35072 Pakistan MK142796
R. subrubescens (Type) F18339 (TENN) USA NR153232
R. tengii (Type) HMAS262728 China NR169926
R. tengii HMAS281318 China MK966689
R. tengii HMAS290030 China MK966688
SPECIES OF RUSSULA RUBIGINOSUS SP. NOV. Phytotaxa 575 (2) © 2022 Magnolia Press • 145
FIGURE 4. Basidiospores (SEM photo) of Russula rubiginosus (Holotype, HMJAU49121). Bars=10 μm
Description:—Basidiomata medium-sized. Pileus 40‒72 mm diam., fleshy, firm, first hemispherical, then turned
applanate with depressed center after maturation, margin becoming somewhat sulcate with age, glabrous, corinthain
pink (xxvii 3ʹʹ d) to livid pink (xxvii 3ʹʹ f) when juvenile, corinthain red (xxvii 3ʹʹ) to corinthain pink (xxvii 3ʹʹd) with
age, center orient pink (ii 11 f) to orange pink (ii 9 f) when juvenile, smiona orange (ii 11 b) to orange pink (ii 9 f)
with age; mostly more or less covered with small ochraceous tawny (xv 15ʹ i) to mikado brown (xxix 13ʹʹ i) spots.
Lamellae adnate, crowded, up to 6 mm wide, more or less furcate towards the stipe, slightly anastomosing, fragile,
light ochraceous salmon (xv 13ʹ d) first, primuline yellow (xvi 19ʹ) when mature or dry, often with rust spots at
lamellae. Stipe cylindrical to slightly clavate with rounded base, enlarged towards the base, firm, 40‒60 × 8‒13 mm,
surface minutely wrinkled, white (liii) when juvenile, becoming baryta yellow (iv 21f) towards the base when old or
dry, appearing ochraceous tawny (xv 15ʹ i) to mikado brown (xxix 13ʹʹ i) patches or spots when handled or with age,
especially bottom. Context firm, white (liii) when juvenile, naples yellow (xvi 19ʹ d) with age, distinctly yellowing
when dry or bruised. Taste acrid. Odor indistinct.
Basidiospores (40/2/2) 7–10 (12–) × 6–9 μm, Q = (1.2–)1.3–1.25(–1.31), Qav. = 1.19 ± 0.05, broadly ellipsoid,
ornamentation of moderately large and distant: 5–7 (–10) amyloid spines in the circle of diameter of 3 μm on
basidiospore surface, spines 1.0–2.0 μm high, mostly isolated, occasionally fused in short chains or crests (0–2 fusions
in the circle), line connections very rare and short (0–1 line connection in the circle), not forming a mesh. Basidia
(30–) 40–53.7 × (13–) 15–17 (–20) μm, clavate to subclavate, thin-walled, oil granules present when viewed with
KOH, 4‒spored, sometimes 2‒spored; sterigmata 6‒8 × 2‒3 µm. Subhymenium pseudoparenchymatic. Lamellar trama
mainly composed of large sphaerocytes. Pleurocystidia 66–92 × 9–13 μm, fusiform, clavate to subclavate, often with
a subacute tip, thin-walled, sometimes with a small appendage and dense crystal inclusions. Cheilocystidia 60–90 ×
8–13 µm, mostly similar to pleurocystidia. Pileipellis is composed of suprapellis and subpellis; suprapellis strongly
gelatinized, 140–250 μm thick, composed of dense, ascending or erect hyphal ends and protruding and near surface
repent pileocystidia, terminal cells 15–45 × 2–3 μm, cylindrical, often constricted towards the apex; subpellis less
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gelatinized, 40–100 μm thick, hyphae 2–6 µm wide, dense, strongly intricate, often branched. Pileocystidia numerous
and often large sized, narrowly to broadly clavate or fusiform, mainly two or three celled, occasionally one-celled,
rarely with more cells, thin-walled, obtuse to subacute, usually inflated near apical part, with terminal cells measuring
30–50 × 6–8 µm, contents in Congo red heteromorphous, usually granulose, but sometimes partly crystalline or banded.
Stipitipellis cutis composed of thin-walled, septate, cylindrical, hyaline hyphae 2–4 μm wide, caulocystidia present,
dispersed, rare, mainly two or three celled, occasionally one celled, rarely with more cells, thin-walled, obtuse to
subacute, usually inflated near apical part, with terminal cells measuring 33–65 × 5–7 μm. Clamp connections absent
in all tissues.
Habitat and distribution:—Ectomycorrhizal fungi symbiotic with Q. mongolica. Solitary in July to August under
Q. mongolica forests. Known from Northeast China.
Additional specimens examined:—CHINA. Heilongjiang Province: Wudalianchi City, ASL 300 m, 126°11’E,
48°37’N, 13 August 2018, Peng-Jie Xing & Yang Xu (HMJAU58934, GenBank OL828348). Inner Mongolia
Autonomous Region: Zhalantun County, Xiushui Scenic Area, ASL 370 m, 27 July 2013, Sai-Fei Li, Dong Zhao
& Guo-Jie Li (HMAS267826, GenBank KX441111); ibid., 28 July 2013, Sai-Fei Li, Dong Zhao & Guo-Jie Li
(HMAS267741, GenBank KX441095).
Russula rubiginosus is characterized by corinthain pink pileus and white stipe with ochraceous-tawny patches or
spots, acrid taste, basidiospore of isolated spines, and the relatively larger cystidia, which showed affinities with
Russula subgen. Insidosula sect. Maculatinae. This result is consistent with the phylogenetic analyses, which the ITS
sequences of R. rubiginosus clustered in R. globispora lineage of subsect. Maculatinae.
Russula globispora shared similar basidiospores ornamentation (isolated amyloid spines on spores) with R.
rubiginosus. However, R. globispora differs R. rubiginosus by having a considerably large-sized pileus (up to 90 mm
diam.) of colored crimson towards the center and carmine towards the margin, a longer stipe (13 × 1.5 cm), large-
sized basidiospores (8–12 × 8.2–11 μm), and the caulocystidia absent (Blum 1952, Caboň et al. 2019, Tian et al.
2015). Russula dryadicala shared the basidiospores of similar size and ornamentation with R. rubiginosus, but differs
by having large-sized pileus (up to 130 mm diam.), a long pink-tinged stipe (35 × 0.8 cm), larger basidia (48–59 ×
13–15 μm) and pleurocystidia (76–97 × 11–12 μm) (Adamčík et al. 2016). Phylogenetically, all five sequences of R.
rubiginosus were identical and formed a well-supported clade in R. globispora lineage, which aided the novelty of this
This work was supported the by Chinese National Natural Science Foundation of China (Nos. 31600020, 31500013,
30770013); and the Key Project on R & D of the Ministry of Science and Technology (No. 2018YFE0107800). We
express our gratitude to the Language Editing Services of MDPI.
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