Research Article
Research Article
Revisiting the phylogeny and taxonomy of the genus Sidera (Hymenochaetales, Basidiomycota) with particular emphasis on S. vulgaris
expand article infoVassiliki Fryssouli, Elias Polemis, Milton A. Typas§, Georgios I. Zervakis
‡ Agricultural University of Athens, Athens, Greece
§ National and Kapodistrian University of Athens, Athens, Greece
Open Access


The genus Sidera (Hymenochaetales, Basidiomycota) comprises white-rot, mono- or dimitic fungi with poroid or hydnoid hymenophore. It has a worldwide distribution albeit with fewer species present in the Southern Hemisphere. Although recent studies revealed the existence of several new Sidera species, there are still taxonomic inconsistencies and obscure phylogenetic relationships amongst certain taxa of the genus. In this work, a large number of Sidera collections were used to obtain an updated phylogeny, based on ITS and 28S rDNA sequences by including new material from Mediterranean Europe. The monophyly of the genus was strongly supported and all species with poroid hymenophore formed a highly-supported lineage with two major subclades. In total, 23 putative species were recognised. Amongst those, five are considered to possibly represent entities new to science, but further work is required since they are represented by single specimens or environmental sequences. Examined collections originally named S. lenis from southern Europe were grouped within S. vulgaris. Similarly, several collections under various names were hereby identified as S. vulgaris, including those of the recently described species S. tibetica. Furthermore, a critical discussion (based on morphoanatomical findings) is made on the key features that could be used to distinguish S. lenis from S. vulgaris.

Key words

Basidiomycetes, biodiversity, fungal phylogeny, Mediterranean Europe, mushroom, white-rot fungi


The genus Sidera Miettinen & K.H. Larss. (Sideraceae, Hymenochaetales, Basidiomycota) was established to harbour four resupinate, wood-inhabiting, white-rot species, in accordance with morphological and phylogenetic evidence: S. lenis (P. Karst.) Miettinen (type species), S. lowei (Rajchenb.) Miettinen, S. lunata (Romell ex Bourdot & Galzin) K.H. Larsson and S. vulgaris (Fr.) Miettinen (Miettinen and Larsson 2011). Amongst them, S. lowei was originally described from Brazil (Rajchenberg 1987), while all the others were described from material collected in Europe. Recently, the diversity of the genus was significantly expanded as a result of phylogenetic studies using nuclear ribosomal markers (ITS and LSU) to describe new taxa from boreal, temperate and tropical habitats (Du et al. 2019, 2020; Liu et al. 2021, 2022, 2023; Xu et al. 2023).

Morphologically, Sidera species are characterised by resupinate, whitish to cream-coloured, yellowish or buff, rarely pinkish or bluish basidiomata, poroid hymenophore with middle-sized to small pores (and, in only one case, hydnoid hymenophore; i.e. S. lunata), monomitic or dimitic hyphal system, generative hyphae with clamps, rather loosely arranged skeletal hyphae, presence of rosette-like crystals in subiculum and/or trama, hymenial cystidia as thin-walled hyphidia (cystidioles) and minute, allantoid to lunate, hyaline, thin-walled, negative in Melzer’s reagent and acyanophilous basidiospores (Miettinen and Larsson 2011).

After re-examining the lectotype of Physisporus lenis (designated by Lowe 1956), Niemelä and Dai (1997) recognised two morphologically distinct species: one that was more common in boreal, mostly pine-inhabiting old-growth forests, i.e. Skeletocutis lenis (P. Karst.) Niemelä and another species that came across more as generalist – in terms of substrate preference – and with a more southern distribution, i.e. Sk. vulgaris (Fr.) Niemelä & Y.C. Dai. In the original description of the genus Sidera, Miettinen and Larsson (2011) used two sequences which were then considered to represent S. vulgaris, both originating from Oceania (Tasmania, Gates FF257; New Zealand, Ryvarden 37198). However, no sequences from European collections were included in their phylogenetic analysis. The sequence from Tasmania nested close to S. lowei, while the sequence from New Zealand appeared as sister to S. lenis, indicating that they possibly correspond to two distinct species. Recent phylogenetic studies in the genus Sidera repeatedly used the aforementioned two specimens (Du et al. 2019, 2020) without including any S. vulgaris material from Europe. In fact, in one of these publications (Du et al. 2020), the specimen from Tasmania was identified as S. minutipora (Rodway & Cleland) Y.C. Dai, F. Wu, G.M. Gates & Rui Du. Moreover, Liu et al. (2021) used the term “Sidera vulgaris sensu lato” to refer to the New Zealand sequence and to some newly-generated sequences from Chinese collections, which were later linked to several new species, i.e. S. americana Z.B. Liu & Yuan Yuan, S. borealis Z.B. Liu & Yuan Yuan, and S. tibetica Z.B. Liu, J. Yu & F. Wu (Liu et al. 2022, 2023). Therefore, the phylogenetic position of S. vulgaris within the genus remains obscure and controversial.

In order to resolve this issue and provide an updated phylogeny of the genus Sidera, several collections from Mediterranean Europe, initially identified as Skeletocutis sp., Sk. lenis and Sk. vulgaris from Mediterranean Europe, were included in this study, together with a large number of publicly available sequences. In addition, answers were sought to the following key questions: (a) Could available specimens confirm the presence of S. lenis in the Mediterranean Region? (b) Is there adequate evidence that S. vulgaris indeed has a cosmopolitan distribution? (c) Which Sidera taxa are accommodated within S. vulgaris sensu lato and are they related to new species correctly introduced in this genus? (d) What are the key morphological features to distinguish S. vulgaris from S. lenis?


Biological material – Morphology

Voucher specimens studied were deposited in the fungaria of the Laboratory of General and Agricultural Microbiology (Agricultural University of Athens, ACAM), the University of Oslo (O, HUBO) and the University of Salamanca (Salamanca, SALA-Fungi). Pore density was studied using a stereomicroscope (Zeiss Stemi 2000‐C) at 10–20× magnification by measuring the number of pores per mm. Microscopic examination was performed with a Zeiss AxioImager A2 microscope under bright field and differential interference contrast (DIC) and microphotographs were taken with the aid of a mounted digital camera (Axiocam). Examination of microscopic features were performed in Cotton Blue, Melzer’s reagent and 5% potassium hydroxide (KOH) mounting media. Measurements were taken in KOH under 1,000× magnification and DIC. For each specimen studied, a minimum number of 25 basidiospores were measured and their size (with standard deviation, SD) is provided as minimum and maximum average (average – SD and average + SD, respectively). In addition, the quotient (Q) for each basidiospore was calculated and is presented together with the respective average values (Qav). Length and width of basidia and hymenial cystidioles are also presented with the same formula: (n = x/y) refers to x measurements (of pores/mm, basidiospores, basidia and cystidioles) from y specimens. Other essential microscopical features which were also examined, including generative and skeletal hyphae from subiculum and hymenophoral trama (tubes) and the presence of stellate crystals and capitate hyphal tips.

DNA extraction, amplification and sequencing

Total genomic DNA was extracted from dried material using the Nucleospin Plant II kit (Macherey and Nagel, USA) according to manufacturer’s protocol with minor modifications (Zervakis et al. 2019). The two regions of the nuclear ribosomal repeat unit – namely the ITS region and a fragment of the ribosomal large subunit gene (28S/LSU) – were amplified through the polymerase chain reaction (PCR) in a MiniAmp Plus Thermal Cycler (Applied Biosystems, CA, USA). The ITS sequences were generated using the forward and backward primers ITS1 and ITS4 (White et al. 1990) to include partial 18S, complete ITS and partial 28S rDNA. The 28S rDNA sequences included the D1/D2 domain by employing the primers LROR and LR5 (Vilgalys and Hester 1990; Stielow et al. 2015). The PCR procedure for ITS was as follows: initial denaturation at 95 °C for 3 min followed by 35 cycles at 94 °C for 40 sec, 52 °C for 45 sec and 72 °C for 1 min and a final extension step of 72 °C for 10 min. The PCR procedure for 28S rDNA was as follows: initial denaturation at 94 °C for 3 min, followed by 35 cycles at 94 °C for 30 sec, 48 °C for 1 min and 72 °C for 1.5 min and a final extension of 72 °C for 10 min. The PCR products were purified with Pure Clean spin columns (Invitrogen, California, USA) following the manufacturer’s instructions.

The PCR products were sequenced using the same forward and backward primers with the amplification procedure in an automated ABI sequencer (Life Technology) at CeMIA Inc. (Larissa, Greece). Trace files obtained from the sequencer were aligned using MEGA 11 (Tamura et al. 2021). Consensus sequences were manually edited to remove or replace all ambiguous characters and were cross-checked against local and public databases, including the international nucleotide sequence database collaboration (INSDC, Arita et al. (2021)) and UNITE (Nilsson et al. 2019). Validated sequences were submitted to GenBank (Sayers et al. 2024) to obtain accession numbers (Table 1).

Table 1.

Biological material used in the phylogenetic analysis of genus Sidera. Information includes final identification of taxa (as derived from the present study; in bold typeface), initial identification of taxa as submitted in public databases or as it appeared on the material examined here for first time (when another name appears in parenthesis, it corresponds to the one subsequently used when this collection served as type material), specimen code, geographic origin, substrate and corresponding GenBank accession numbers for ITS and 28S rDNA. The reference for each entry is also provided; asterisk (*) indicates those not accompanied by a publication. Collections serving as type material are indicated with superscript letters, i.e., H: holotype, L: lectotype, and P: paratype; ‘n.a.’ denotes not available information.

Species Collection code Geographic origin Substrate GenBank accession no. Reference
S. americana
Sidera sp. (S. americana) Dai 12730H USA: CT on rotten stump of Pinus MW198478 n.a. Liu et al. (2021, 2023)
S. malaysiana Dai 19173 Canada on rotten angiosperm wood MW198477 MW192005 Liu et al. (2022, 2023)
Sidera sp. TUF101553 Estonia Pinus sylvestris UDB015767 n.a. Runnel (2010)*
S. vulgaris Alden Dirks: ACD0413 USA: MI n.a. OL756000 OL742443 Dirks (2021)*
S. borealis
Sidera sp. (S. borealis) Cui 11216H China: SN fallen angiosperm trunk MW198485 n.a. Liu et al. (2021, 2023)
S. cf. vulgaris Dai 22822 China: YN on rotten
wood of Picea
OM974254 OM974246 Liu et al. (2022, 2023)
Sidera sp. TUF122801 Estonia Pinus sylvestris UDB023006 UDB023006 Runnel (2013)*
S. inflata
S. inflata Cui 13610H China: HI on rotten angiosperm wood MW198480 n.a. Liu et al. (2021)
S. lenis
S. lenis O. Miettinen 11036.1L Finland n.a. FN907914 FN907914 Miettinen and Larsson (2011)
S. lenis NSK 1017015 Russia n.a. OR364533 n.a. Vlasenko (2023)*
S. lenis Dai 22834 China: YN on rotten wood of Picea OQ134538 n.a. Liu et al. (2023)
S. lenis TUF111091 Sweden Pinus sylvestris UDB032409 n.a. Sell (2015)*
S. lowei
S. lowei Dollinger 922 USA: FL Quercus KY264044 n.a. Dollinger and Vlasak (2016)*
S. lowei Ryvarden 40576 Venezuela n.a. FN907917 FN907917 Miettinen and Larsson (2011)
S. lunata
Athelopsis lunata JS 15063 (1717) Norway n.a. DQ873593 DQ873593 Larsson et al. (2006)
S. lunata S851 Estonia soil UDB0662815 n.a. Tedersoo et al. (2018)*
S. malaysiana
S. malaysiana Dai 18570H Malaysia on rotten angiosperm wood MW198481 MW192007 Liu et al. (2021)
S. minutipora
S. minutipora Cui 16720 Australia: Tasmania on rotten stump of Eucalyptus MN621349 MN621348 Du et al. (2020)
S. vulgaris G. Gates FF257 Australia: Tasmania n.a. FN907922 FN907922 Miettinen and Larsson (2011)
S. minutissima
S. minutissima Dai 19529 H Sri Lanka on rotten angiosperm branch MN621352 MN621350 Du et al. (2020)
S. minutissima Dai 22495 China n.a. OM974248 OM974240 Liu et al. (2022)
Sidera sp. KAS: L1620 Réunion Island n.a. UDB024833 n.a. Ordynets (2015)*
Sidera sp. TUF123971 Seychelles n.a. UDB039740 n.a. Kõljalg (2018)*
S. parallela
S. parallela Cui 10346H China: YN on rotten angiosperm trunk MK346145 n.a. Du et al. (2020)
S. parallela Cui 10361P China: YN on fallen angiosperm trunk MK346144 n.a. Du et al. (2020)
S. parallela Dai 22038 China n.a. MW477793 MW474964 Liu et al. (2022)
S. punctata
S. punctata Dai 22119H China: HI on rotten angiosperm wood MW418438 MW418437 Liu et al. (2021)
unc. fungus L042880-122-060-A02 Ocean air filter sample GQ999131 n.a. Fröhlich-Nowoisky et al. (2012)
unc. fungus L042881-122-061-B08 Taiwan air filter sample GQ999432 n.a. Fröhlich-Nowoisky et al. (2012)
S. roseo-bubalina
S. roseo-bubalina Dai 11277T China: HA under decay Quercus MW198483 n.a. Liu et al. (2021)
S. salmonea
S. salmonea Dai 23354P China: Tibet Abies OM974250 OM974242 Liu et al. (2022)
S. salmonea Dai 23428 China: Tibet Pinus armandii OM974251 OM974243 Liu et al. (2022)
S. srilankensis
S. srilankensis Dai 19654H Sri Lanka on rotten angiosperm wood MN621344 MN621346 Du et al. (2020)
S. srilankensis Dai 19581P Sri Lanka on rotten angiosperm wood MN621345 MN621347 Du et al. (2020)
S. tenuis
S. tenuis Dai 18697H Australia: Tasmania on rotten stump of Eucalyptus MK331865 MK331867 Du et al. (2020)
S. tenuis Dai 18698P Australia on rotten stump of Eucalyptus MK331866 MK331868 Du et al. (2020)
S. tianshanensis
S. tianshanensis Cui 19143H China: XJ on fallen trunk of Picea schrenkiana OP920995
Xu et al. (2023)
S. tianshanensis Cui 19132
China: XJ on stump of Picea schrenkiana OP920994
OP920986 Xu et al. (2023)
S. vesiculosa
S. vesiculosa BJFC025377T Singapore on rotten angiosperm MH636564 MH636566 Du et al. (2019)
S. vesiculosa BJFC025367P Singapore on rotten angiosperm MH636565 MH636567 Du et al. (2019)
Sidera sp. TUE002764 Papua New Guinea soil UDB07018609 n.a. Tedersoo et al. (2020)*
S. vulgaris
S. vulgaris ACAM 2013-0017 Greece Pinus halepensis PP275215 PP275225 present work
S. vulgaris ACAM DD2559 Greece Abies cephalonica PP275216 PP275226 present work
Skeletocutis vulgaris HUBO 7745 Italy Pinus sylvestris PP275217 PP275227 present work
Skeletocutis lenis HUBO 8296 Italy Fagus PP275218 PP275228 present work
Skeletocutis vulgaris HUBO 8465 Italy Pinus nigra ssp. laricio PP275219 PP275229 present work
Skeletocutis lenis SALA-Fungi 3749 Spain Eucalyptus camaldulensis PP275220 n.a. present work
Skeletocutis lenis SALA-Fungi 3752 Spain Pinus pinaster PP275221 n.a. present work
Skeletocutis sp. SALA-Fungi 4105 Spain Pinus pinaster PP275222 n.a. present work
Skeletocutis sp. SALA-Fungi 4111 Spain Acer monspessulatum PP275223 n.a. present work
S. vulgaris TU114503 Estonia Populus tremula UDB034888 n.a. Sell (2017)*
S. vulgaris TU135349 Estonia Picea abies UDB0754207 n.a. Sell (2018)*
Sidera sp. (S. tibetica) Dai 23648H China: Tibet Pinus armandii OM974253 OM974245 Liu et al. (2022)
Sidera sp. (S. tibetica) Dai 23407P China: Tibet n.a. OM974252 OM974244 Liu et al. (2022)
S. tibetica Dai 22151 China n.a. MW477794 MW477794 Liu et al. (2021)
S. tibetica Dai 21057 Belarus on rotten wood of Picea MW198484 MW192009 Liu et al. (2021)
S. tibetica LE F-342597 Russia Pinus brutia var. eldarica OR457651 n.a. Volobuev (2023)
Schizopora sp. 206 Spain Castanea sativa EM root tips MN947225 n.a. Santolamazza-Carbone (2020)*
Schizopora sp. DLL2009-014 USA: MN Populus spp. JQ673191 n.a. Brazee et al. (2012)
Schizopora sp. FH:BHI-F453 USA: MA n.a. MF161274 n.a. Haelewaters et al. (2018)
Sidera sp. UC2022907 USA: CA on litter or well decayed wood in pinaceous forest KP814250 n.a. Rosenthal et al. (2017)
unc. Hyphodontia 1Bart548S USA: NH n.a. HQ022192 n.a. Vineis (2011)
unc. fungus S38 Germany air sample FJ820526 n.a. Fröhlich-Nowoisky et al. (2012)
Sidera sp. 1
S. vulgaris Ryvarden 37198 New Zealand n.a. FN907918 FN907918 Miettinen and Larsson (2011)
Sidera sp. 2
Sidera sp. UC2023008 USA: MS decayed wood in pinaceous forest KP814157 n.a. Rosenthal et al. (2017)
Sidera sp. 3
S. lowei Ryvarden 38817 New Zealand n.a. FN907919 FN907919 Miettinen and Larsson (2011)
Sidera sp. 4
unc. fungus L042886-122-066-F04 Taiwan air filter GQ999509 n.a. Fröhlich-Nowoisky et al. (2012)
unc. fungus L042881-122-061-B09 Taiwan air filter GQ999433 n.a. Fröhlich-Nowoisky et al. (2012)
Sidera sp. 5
Sidera sp. MEL:2382752 Australia: NT n.a. n.a. KP012935 Bonito et al. (2014)*
Alloclavaria purpurea Miettinen 18831 ΗΠΑ: WA old-growth forest with conifers ON188807 ON188807 Viner (2022)*
Rickenella mellea Lamoure 74 n.a. n.a. U66438 U66438 Lutzoni (1997)
Resinicium furfuraceum (Skvortzovia furfuracea) KHL 11738 Finland n.a. DQ873648 DQ873648 Larsson et al. (2006)
Skvortzovia furfurella KHL 10180 Puerto Rico n.a. DQ873649 DQ873649 Larsson et al. (2006)

Phylogenetic analysis

Phylogenetic analysis of the genus Sidera was performed using Maximum Likelihood (ML) and Bayesian Inference (BI) approaches with separate ITS and 28S rDNA datasets. In addition, a concatenated two-marker dataset was analysed by including specimens with data available for both markers. Besides the newly-generated sequences, additional reference sequences were retrieved from INSDC and UNITE using BLASTn searches. Skvortzovia furfurella (Bres.) Bononi & Hjortstam and Skvortzovia furfuracea (Bres.) G. Gruhn & Hallenberg were used as outgroups (Liu et al. 2022), while two additional Hymenochaetales species, i.e. Alloclavaria purpurea (O.F. Müll.) Dentinger & D.J. McLaughlin and Rickenella mellea (Singer & Clémençon) Lamoure, were also included in the analysis. A detailed list of specimens examined in the present study, along with corresponding information, for example, initial identification, habitat, sequence database accession number and pertinent publication (when available), is provided in Table 1.

Sequences were aligned by the online version of MAFFT v. 7 (Katoh et al. 2019) using the “G-INS-i” and “Q-INS-I” strategies for ITS for 28S rDNA barcodes, respectively. The alignments were visually inspected and manually adjusted for conspicuous errors and gapped sites in Mesquite 3.81 (Maddison and Maddison 2023). The MUMSA tool (Lassmann and Sonnhammer 2006) was used to select the best alignment.

ML analysis was performed using IQ-TREE v. 2.2.7 (Minh et al. 2022) via the CIPRES Science Gateway (Miller et al. 2015) with a random starting tree. All model parameters were estimated by the software. The best Maximum Likelihood tree was retained from all searches and the Maximum Likelihood bootstrap values (ML-BS) were determined using ultrafast bootstrapping algorithm with 10,000 replicates. BI analysis was implemented in MrBayes v.3.2.7a (Ronquist et al. 2012) employing optimal models of evolution determined for ITS and 28S rDNA with jModelTest2 v.2.1.6 (Darriba et al. 2012) under the Bayesian Information Criterion (BIC). Two parallel runs, each one consisting of four incrementally heated Monte Carlo Markov Chains, were initiated from programme-generated random trees. The analysis involved sampling every 1,000th generation until the average standard deviation of split frequency fell below 0.005. The burn-in phase (first 25% of sampled trees) was discarded. The remaining trees were used to generate a 50% majority rule consensus tree and to estimate the Bayesian posterior probabilities (BPPs). Branches with ML-BS and BPPs equal to or above 65% and 0.95, respectively, were considered as significantly supported.

The best topologies from MP analyses are presented and the final alignments and the phylograms are deposited in TreeBASE ( under accession ID: 31153. The sequence identity was calculated using MAFFT, accessed through EMBL-EBI (


Taxonomy – Morphology

Sidera vulgaris (Fr.) Miettinen, Mycological Progress 10 (2): 136 (2011)

Figs 1, 2

Polyporus vulgaris Fr., Systema Mycologicum 1: 381 (1821). Basionym.

Skeletocutis vulgaris (Fr.) Niemelä & Y.C. Dai, Annales Botanici Fennici 34 (2): 135 (1997). Synonyms.


Basidioma —Annual to biennial, resupinate, soft when fresh and rather tough, soft-corky after drying, confluent and widely effused covering extended under-surface of decaying logs, 0.8–2.0 mm thick at the centre; pore surface white to cream when fresh, becoming yellowish to buff when dry; sterile margin indistinct, cottony, white, thinning-out; pores very small, roundish, (5) 6–8 (10) per mm (n = 273/13); dissepiments thin, entire to slightly lacerate; subiculum very thin, cottony, concolorous with the tube layer; tubes concolorous with the poroid surface, up to 2 mm long.

Figure 1. 

S. vulgaris specimen in situ (ΑCAM 2013-0017). Scale bar: 5 cm (left); 2 cm (right).

Hyphal system dimitic in all parts of the basidioma; generative hyphae smooth, without encrustations, septa with clamp connections; skeletal hyphae not reacting with Cotton Blue, Melzer’s reagent or KOH.

Subiculum —Hyphae interwoven, skeletal hyphae dominating, skeletals (1.7) 2–3.5 (4.0) μm in diameter, rosette-like crystal clusters rare to common.

Tubes —Hyphae subparallel to moderately interwoven. Generative hyphae, thin to slightly thick-walled, poorly branched, 1.7–3.0 μm in diameter. Skeletal hyphae, thick-walled to subsolid, hyaline, rarely branched, flexuous, 1.7–3.5 μm in diameter, with scattered swellings up to 7 μm. Dissepiment edges with both generative and skeletal hyphae that often bear a swollen, capitate apex, generative hyphae sometimes covered by a mucous droplet, rosette-like crystals frequent in mature basidiomata. Cystidioles seldom to abundant, fusoid, thin-walled, hyaline, basally swollen, with hyphoid neck and mostly obtuse or capitate tip, some bearing crystals at apex (asterocystidia), a few modified as halocystidia were also observed, (9.3) 12.4–19.9 (25.0) × (2.2) 2.8–4.0 (5.3) μm (n = 125/15). Basidia barrel-shaped to somewhat short-clavate, with four sterigmata and a basal clamp, (6.2) 6.8–9.9 (14.6) × (3.1) 3.8–4.7 (5.6) μm (n = 185/15); basidioles barrel-shaped, slightly shorter than the basidia.

Figure 2. 

Micromorphological features of S. vulgaris; scale bar 5 μm [except of f and g 10 μm] a basidiospores (all specimens) b basidia (ACAM 2013-0017, ACAM DD2559, HUBO 7745, HUBO 8296, HUBO 8465, SALA-Fungi 3752) c hymenium with basidia and basidioles (HUBO 8465) d branched and unbranched cystidioles bearing crystals at apex (asterocystidia) (ACAM DD2559, SALA-Fungi 3752, SALA-Fungi 4111) e hymenial cystidioles (ACAM DD2559, HUBO 8296, SALA-Fungi 3749) f hyphae of the subiculum with dominating skeletals (ACAM 2013-0017) g dissepiment edges with skeletal and generative hyphal ends (HUBO 7745) h details of the rosette-like crystal clusters from tramal hyphae (HUBO 7745) i capitate ends of generative hyphae from dissepiments and hymenium (with mucous droplets) (HUBO 8296, SALA-Fungi 3749, SALA-Fungi 3752) j skeletal hyphae from dissepiments with swellings (ACAM 2013-0017, HUBO 7745, SALA-Fungi 3752, SALA-Fungi 4111).

Basidiospores —Cylindrical, moderately curved to lunate, thin-walled, hyaline, smooth, negative in Melzer’s reagent, acyanophilous, (3.0) 3.4–3.9 (4.3) × (1.2) 1.4–1.6 (1.8) μm, Average = 3.6 × 1.5 μm, Q = (1.95) 2.24–2.60 (3.08) QAV = 2.41 (n = 399/15).

Distribution and hosts

The species is reported from Mediterranean Europe (e.g. Portugal, Spain, France, Italy, Croatia and Greece), Germany, Slovakia, Poland, Estonia, Sweden, Belarus, Russia, as well as from Armenia, Georgia, Iran, Kazakhstan, China, USA and Canada (Niemelä and Dai 1997; Ghobad-Nejhad 2011; Bernicchia et al. 2020; Liu et al. 2022; this work). It occurs on various broadleaved trees of (Alnus, Eucalyptus, Fagus, Populus, Quercus, Sorbus and Ulmus), as well as on coniferous trees, i.e. Picea, Pinus (P. halepensis, P. nigra ssp. laricio, P. pinaster, P. sylvestris) or Juniperus and on Abies cephalonica (this work).

Specimens examined

Greece: Sterea Ellas, Fthiotida, Gardiki, on trunk of Abies cephalonica, 28 April 2007, ACAM DD2559, coll. D. Dimou. Attica, Mt. Parnitha, on trunk of P. halepensis, 30 May 2013, ACAM 2013-0017. coll. E. Polemis. ITALY: Emilia Romana, Forli, Pian del Pero Cullacea, on Ulmus glabra, 7 October 2002, HUBO 7629, coll. A. Bernicchia (as Sk. vulgaris); ibidem. on Fagus sp. 11 October 2006, HUBO 8296, coll. A. Bernicchia (as Sk. lenis); Ferrara, Bosco della Mesola, on Populus sp. 12 November 2003, HUBO 7701, coll. A. Bernicchia (as Sk. lenis); Bologna, Parko la Martina, on P. sylvestris, 16 July 2003, HUBO 7745, coll. A. Bernicchia (as Sk. vulgaris); Ravena, Pineta San Vitale, on Populus alba, 4 November 2003, HUBO 7811, coll./det. A. Bernicchia (as Sk. vulgaris). Sardinia, Tonara, Isca de sa Mela, on P. nigra ssp. laricio, 14 October 2007, HUBO 8465, coll. L. Arras (as Sk. vulgaris); Sorgono, Isca de sa Mela, on P. nigra ssp. laricio, 18 November 2009, HUBO 8522, coll. A. Bernicchia (as Sk. lenis). SPAIN: Castile-Leon, Garcibuey, on Eucalyptus camaldulensis, 7 November 2005, SALA-Fungi 3749, ibidem. on P. pinaster, 22 November 2006, SALA-Fungi 3747, coll. S.P. Gorjón (as Sk. lenis); Herguijuela de la Sierra, on P. pinaster, 18 November 2007, SALA-Fungi 3752, coll. S.P. Gorjón (as Sk. lenis); Miranda del Castañar, on P. pinaster, 22 November 2006, SALA-Fungi 4105, coll. S.P. Gorjón (as Skeletocutis sp.); San Martín del Castañar, on Acer monspessulatum, 14 October 2007, SALA-Fungi 4111, coll. S.P. Gorjón (as Skeletocutis sp.); Cepeda, on Alnus glutinosa, 29 November 2006, SALA-Fungi 3745, coll. S.P. Gorjón (as Sk. lenis).

Phylogenetic analysis

To estimate the phylogeny of the genus Sidera, datasets of ITS and 28S rDNA sequences were compiled, including sequences from collections with a Mediterranean distribution, as well as from pertinent specimens and environmental samples deposited in INSDC and UNITE in order to cover as much as possible the diversity and distribution of the genus. The total dataset consisted of 69 collections represented by 68 ITS and 36 28S sequences (Table 1). The material examined for the first time in the present study included nine collections from Mediterranean Europe, from which nine ITS and five 28S sequences were obtained. Additional information on the phylogenetic analyses performed for each dataset is provided in Suppl. material 1. Both applied phylogenetic strategies, ML and BI, produced phylograms characterised by a consistent topology, devoid of any supported conflicts.

The phylogenetic reconstruction, based on the ITS sequences (Fig. 3), recovered Sidera as a strongly-supported monophyletic clade (ML-BS 100%, BPP 1.00), which is further segregated into three well-supported main clades, A through to C. In total, 22 highly-supported terminal clades were recovered including those corresponding to the 18 formally described taxa; amongst them, 14 are represented by type sequences (plus one representing S. tibetica, which, however, should be considered as synonym of S. vulgaris as explained below). No sequences from type specimens were available for S. lunata, S. vulgaris, S. lowei or S. minutipora. In addition, four terminal clades do not correspond to the already known taxa and they could represent undescribed species. They are provisionally referred to as ‘Sidera sp. 1, 2, 3 and 4’.

Figure 3. 

Phylogenetic relationships within the genus Sidera inferred by using ML analysis on the ITS sequence dataset. ML BS ≥ 65% and BPP ≥ 0.95 are appended to nodes; asterisk denotes 100% ML BS and/or 1.00 BBP. Specimens studied are followed by their voucher code and geographic origin. Sequences determined in the present study appear in bold, while those representing type material are underlined. The phylogram is rooted with Skvortzovia furfuracea and Skvortzovia furfurella. The scale bar indicates 0.1 expected change per site per branch.

Clade A (100%, 1.00) includes only S. lunata, which is distantly related to the rest of the Sidera spp. and it has a hydnoid hymenophore. All other taxa have poroid hymenophores and are grouped with significant support (95%, 0.98). They are further subdivided into clades B (100%, 1.00) and C (99%, 1.00) consisting of eight and 13 species, respectively. Clade B includes S. lenis – the type species of the genus – represented by collections from Sweden, Finland, Russia and China, as well as a cluster composed of S. borealis and two closely-related taxonomic entities. The first of them is hereby designated ‘Sidera sp. 1’ (UNITE DOI: SH1110196.09FU); it corresponds to the specimen Ryvarden 37198 from New Zealand, initially identified as S. vulgaris, but apparently not related to the real S. vulgaris, which is grouped in Clade C and includes material from the Northern Hemisphere. The second is represented by the specimen Sidera sp. UC2023008 from USA. Although closely positioned to S. borealis, it is considered as distinct from the latter species since it shows a rather low ITS sequence identity (96.5–98.4%) and distant geographic occurrence (S. borealis is reported from Europe and China). We provisionally call it ‘Sidera sp. 2’ (UNITE DOI: SH1110192.09FU). Moreover, clade B includes two pairs of sister species (100%, 1.00), i.e. S. tianshanensis B.K. Cui & T.M. Xu and S. salmonea Z.B. Liu, Jian Yu & F. Wu (both from Asia), as well as S. parallela Y.C. Dai, F. Wu, G.M. Gates & Rui Du and S. americana (the former originates from Asia, while the latter from North America and north Europe).

Clade C comprises the main diversity of the genus by accommodating 11 species and two entities possibly corresponding to new taxa. S. srilankensis Y.C. Dai, F. Wu, G.M. Gates & Rui Du and S. malaysiana Z.B. Liu & Y.C. Dai form a robustly-supported clade (100%, 1.00) consisting of Asian specimens. Similarly, sequences deriving from the Neotropics correspond to S. lowei. However, another collection (Ryvarden 38817) – initially identified as S. lowei from New Zealand – is phylogenetically separated from the previous species and seems to represent a distinct taxon (ITS sequence identity: 83.0–83.8%), herein called ‘Sidera sp. 3’ (UNITE DOI: SH1110192.09FU). Furthermore, S. roseobubalina Z.B. Liu & Y.C. Dai is represented only by the holotype, originating from China. It is related to two sequences derived from environmental samples (air filters, Taiwan; ITS sequence identity to S. roseobubalina: 93.5–93.6%); hence, the latter could possibly correspond to an undescribed taxon which is provisionally named ‘Sidera sp. 4’ (UNITE DOI: SH1111516.09FU). The aforementioned taxa are strongly linked (96%, 1.00) with a group consisting of S. punctata Z.B. Liu & Y.C. Dai and S. vesiculosa Rui Du & M. Zhou; these four species are represented by sequences from material of Asian origin. Finally, a well-supported cluster (97%, 1.00) is composed by S. minutissima Y.C. Dai, F. Wu, G.M. Gates & Rui Du (including specimens from islands of the Indian Ocean and China), S. inflata Z.B. Liu & Y.C. Dai from China (sequence data available only from the type collection), the sister species S. minutipora (Rodway & Cleland) Y.C. Dai, F. Wu, G.M. Gates & Rui Du and S. tenuis Y.C. Dai, F. Wu, G.M. Gates & Rui Du (consisting of material from Australia) and S. vulgaris.

S. vulgaris forms a highly-supported terminal clade (100%, 1.00) composed of 22 sequences labelled with various names, for example, S. vulgaris, S. lenis, S. tibetica, Sidera sp., Skeletocutis sp. and Schizopora sp. All samples originated from the Northern Hemisphere (Europe, Asia and North America). In particular, the clade includes all material studied for the first time in the framework the present study (collected from various substrates in Spain, Italy and Greece), as well as sequences from Germany, Estonia, Belarus, Russia, China (incl. Tibet) and the USA (UNITE DOI: SH1262165.09FU).

Although represented by fewer sequences, the phylogenetic reconstructions that were based on 28S or on the concatenated ITS and 28S sequences (Suppl. material 2 and Fig. 4, respectively) provided similar topologies as the ITS tree by maintaining the same phylogenetic positions of S. lunata and of species within Clades B and C (100%, 1.00). It is also interesting that the unnamed taxa ‘Sidera sp. 1’ and ‘Sidera sp. 3’ seem to be well-separated from the already known species, as indicated above (Fig. 3). In addition, the specimen MEL:2382752 (Australia), originally identified as Sidera sp., appears to be distinct from the two most closely-related taxa, i.e. S. srilankensis and S. malaysiana; therefore, it is assigned with the provisional name ‘Sidera sp. 5’ (Suppl. material 2).

Figure 4. 

Phylogenetic relationships within the genus Sidera inferred by using ML analysis on the concatenated ITS and 28S rDNA sequence dataset. ML BS ≥ 65% and BPP ≥ 0.95 are appended to nodes; asterisk denotes 100% ML BS and/or 1.00 BBP. Specimens studied are followed by their voucher code and geographic origin. Sequences determined in the present study appear in bold, while those representing type material are underlined. The phylogram is rooted with Skvortzovia furfuracea and Skvortzovia furfurella. The scale bar indicates 0.1 expected change per site per branch.


This study mainly deals with the taxonomic uncertainty associated with collections under the names S. vulgaris and S. lenis. A great deal of confusion stems from the erroneous initial identifications of such specimens and, as explained below, further obstacles were raised by the description of the (allegedly) new species S. tibetica (Liu et al. 2022).

Miettinen and Larsson (2011), who introduced Sidera as a new genus to accommodate – amongst others – the dimitic polypores Sk. vulgaris and Sk. lenis, studied the former by examining three collections: one European (Poland, Niemelä 5981) and two from the Southern Hemisphere. However, sequence data were available only from the latter two. As our results show, the collection from New Zealand (Ryvarden 37198), that was considered to represent S. vulgaris and was included as such in several phylogenetic studies, was assigned to clade B in the present study (as ‘Sidera sp. 1’). Furthermore, the collection from Tasmania (G. Gates FF257) was grouped in clade C and was identified as S. minutipora (Du et al. 2020). Therefore, it is clear that these two collections are not related to S. vulgaris. Moreover, several specimens included either as S. vulgaris or S. vulgaris sensu lato in the previous publications (Liu et al. 2021, 2022) were later linked to other species of this genus by the same group of authors (Liu et al. 2023), for example, to S. americana (Dai 12730 from USA and Dai 19173 from Canada) and S. borealis (Cui 11216 and Dai 22822 from China).

Most importantly, another new Sidera species was recently introduced under the name S. tibetica (Liu et al. 2022). It was described on the basis of two sequenced specimens from Tibet, but without being examined versus any real/authentic S. vulgaris collections. In fact, the three “S. vulgaris” collections included in the aforementioned study were Ryvarden 37198 (New Zealand), Dai 19173 (Canada) and Dai 22822 (China). However, the first of these corresponds to Sidera sp. 1 (as the present work demonstrated), the second to S. americana and the third to S. borealis. In addition, all S. tibetica sequences from the work of Liu et al. (2022) [and from other recent publications where this name was also erroneously used (e.g. Volobuev (2023))] were grouped (100%, 1.00) together with a large number of S. vulgaris specimens originating from Europe (#15), Asia (#3) and USA (#4). It should additionally be noted that Liu et al. (2021) had indications that the allegedly new species occurred also in Europe (since they have examined the specimen Dai 21057, initially identified as S. vulgaris sensu lato from Belarus). However, they did not include it in the description of S. tibetica (Liu et al. 2022), whereas it was used in their more recent study under this name (Liu et al. 2023). Therefore, it is evident that S. tibetica was erroneously introduced as a new species since the respective examined material corresponds to S. vulgaris, i.e. the already existing species, previously described from specimens originating from Sweden (Niemelä and Dai 1997). Apparently, the lack of sequence data from the type material of S. vulgaris and not including correctly identified collections of the appropriate geographic origin (e.g. Europe) were the main reasons for this major issue detected in the publication of Liu et al. (2022). Hence, S. tibetica should be considered as a synonym of S. vulgaris.

The molecular evidence provided by the phylogenetic analyses (Fig. 3, Fig. 4 and Suppl. material 2) shows that S. vulgaris is well discriminated from S. lenis since the respective terminal clades are properly defined and clearly separated. On the other hand, distinguishing S. vulgaris from S. lenis morphologically remains a difficult task; this resulted in incorrect identifications of the two species in Europe and elsewhere. Until very recently, the distribution of S. vulgaris in south Europe was considered to be unknown because, as stated, “… it was easily confused with S. lenis” (Bernicchia et al. 2020). It is now clear that S. vulgaris is the only species of the genus which is also present in Mediterranean Europe.

Our morphological studies, in conjunction with the verified identity of specimens from DNA sequencing, revealed that the most stable and reliable character to distinguish these two species is the pore size, which is clearly smaller in S. vulgaris, on average, more than six pores per mm, as opposed to less than six pores per mm in S. lenis (Table 2). The length of the basidia, furthermore, seems to be important in this regard, as they hardly exceed 10 μm in S. vulgaris, while they are always longer in S. lenis. Regarding the size and shape of basidiospores, our measurements indicate a much wider deviation, particularly with regards to the width, which largely affects the quotient (Q, length/width). Our work also suggests that S. vulgaris basidiospores may exceed 1.5 μm in width; this contradicts pertinent generic keys which placed a clear-cut value of 1.5 μm between these species (i.e. Ryvarden and Melo (2017); Bernicchia et al. (2020)). Apparently, this tiny value – and variations thereof – are very difficult to detect with accuracy. In contrast, the average spore length seems to be a more reliable character, since it does not exceed 4 μm in S. vulgaris, in contrast to S. lenis whose spores are usually longer. In our opinion, the presence of the stellate crystal agglomerations and the mucous deposit on the capitate generative hyphal tips are unstable characters, most likely affected by the age of basidiomata and the microscopy techniques used; thus, they are of questionable taxonomic value. In addition, the most important taxonomic features mentioned in the description of S. tibetica (Liu et al. 2022) are similar to (or not differing considerably from) those of S. vulgaris (Table 2). The deviations observed in spore size (especially) and pore density are small and they cannot be considered as significant since only three specimens of S. tibetica were used for its original description.

Table 2.

Comparison of key morphological features from collections of Sidera vulgaris (this work; Niemelä and Dai 1997), S. tibetica (Liu et al. 2022) and S. lenis (Niemelä and Dai 1997).

S. vulgaris (this work) S. vulgaris (Niemelä and Dai 1997) S. tibetica (Liu et al. 2022) S. lenis (Niemelä and Dai 1997)
Pores (per mm) 6–8 6–8 7–8 4–6
Spores 3.2–4.0×1.3–1.7 μm, av. 3.64×1.51 μm Q = 2.24–2.60 2.9–3.6×0.9–1.4 μm, av. 3.14×1.08 μm Q = 2.44–3.11 2.9–3.1×1.0–1.1 μm, av. 3.01×1.05 μm Q = 2.78–2.91 3.9–4.9×1.5–2 μm, av. 4.35×1.76 μm Q = 2.29–2.74
Basidia length 6.8–11 μm 6.5–8.5 μm 8–9.5 μm 10–13.5 μm
Skeletals in KOH 1.7–3.5 μm 2.7–3.5 μm 2.0–4.0 μm 3.5–4.8 μm
Stellate crystals frequent very rare frequent frequent


In this work, a large number of Sidera sequences (ITS and LSU rDNA) were analysed which included new material from Mediterranean Europe, as well as publicly available sequences. The monophyletic nature of the genus was strongly supported in the generated trees. Sidera lunata (characterised by a hydnoid hymenophore) was identified as the sister group to the remainder of the genus in the derived phylogeny, while species with poroid hymenophore formed a robustly supported lineage that was subdivided into two major clades. Amongst 23 species in total, five are possibly new to science, but since they are mostly represented by single collections, further work is needed before any definite conclusions could be drawn. The presence of S. lenis was assessed in north Europe, Russia and China, while examined collections from south Europe under this name were recovered within S. vulgaris. The latter species exhibits a Holarctic distribution. It occurs on dead wood of angiosperms and gymnosperms, including the regions of Eurasia where it was erroneously reported as S. tibetica. As we demonstrate, the description of this allegedly new species was based on collections that are hereby identified as S. vulgaris. This observation also emphasises the need to proceed with the epitypification of S. vulgaris since the type material maintained in Herbarium UPS may be too old for successful sequencing.


Curators of O (University of Oslo, Norway) and SALA (University of Salamanca, Spain) are thanked for loans of specimens. Our gratitude is also extended to A. Bernicchia and S.P. Gorjon for kindly providing access to herbarium collections.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.


No funding was reported.

Author contributions

All authors have contributed equally.

Author ORCIDs

Vassiliki Fryssouli

Elias Polemis

Georgios I. Zervakis

Data availability

All of the data that support the findings of this study are available in the main text or Supplementary Information.


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Supplementary materials

Supplementary material 1 

Detailed characteristics of the phylogenetic analysis performed for each sequence dataset used for the study of Sidera collections

Vassiliki Fryssouli, Elias Polemis, Milton A. Typas, Georgios I. Zervakis

Data type: docx

This dataset is made available under the Open Database License ( The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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Supplementary material 2 

Phylogenetic relationships within the genus Sidera inferred by using ML analysis on the 28S rDNA sequence dataset

Vassiliki Fryssouli, Elias Polemis, Milton A. Typas, Georgios I. Zervakis

Data type: pdf

This dataset is made available under the Open Database License ( The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (316.41 kb)
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