Mycologia
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Phylogenetic analysis and morphological
characteristics of laccate Ganoderma specimens in
Finland
Marta Cortina-Escribano, Pyry Veteli, Michael John Wingfield, Brenda
Diana Wingfield, Martin Petrus Albertus Coetzee, Henri Vanhanen & Riikka
Linnakoski
To cite this article: Marta Cortina-Escribano, Pyry Veteli, Michael John Wingfield, Brenda
Diana Wingfield, Martin Petrus Albertus Coetzee, Henri Vanhanen & Riikka Linnakoski (2024)
Phylogenetic analysis and morphological characteristics of laccate Ganoderma specimens in
Finland, Mycologia, 116:6, 1046-1062, DOI: 10.1080/00275514.2024.2381424
To link to this article:  https://doi.org/10.1080/00275514.2024.2381424
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Phylogenetic analysis and morphological characteristics of laccate Ganoderma 
specimens in Finland
Marta Cortina-Escribano a,b, Pyry Vetelic, Michael John Wingfieldd, Brenda Diana Wingfieldd, 
Martin Petrus Albertus Coetzee d, Henri Vanhanen a, and Riikka Linnakoski c
aProduction Systems, Natural Resources Institute Finland (LUKE), Joensuu, North Karelia 80100, Finland; bSchool of Forest Sciences, 
University of Eastern Finland, Joensuu, North Karelia 80100, Finland; cNatural Resources, Natural Resources Institute Finland (LUKE), 
Uusimaa, Helsinki 00790, Finland; dDepartment of Biochemistry, Genetics and Microbiology, Forestry and Agricultural Biotechnology 
Institute (FABI), University of Pretoria, Pretoria 0028, South Africa
ABSTRACT
The Ganoderma lucidum complex includes fungi with similar morphologies but which are thought to 
represent different species. The lack of available type material and associated absence of multiple 
locus sequence data has complicated identification of these fungi. The aim of this study was to clarify 
the identity of the laccate Ganoderma species occurring in Finland by inferring a phylogeny using 
DNA sequences from available boreal-temperate material. DNA from Finnish isolates together with 
an older G. lucidum isolate originating from the United Kingdom was sequenced, and the morpho-
logical features of the Finnish specimens were examined. The phylogenetic analysis of the internal 
transcribed spacer region (ITS), the elongation factor 1-α (tef1), RNA polymerase II subunit (rpb2), and 
partial β-tubulin (β-tub) genes revealed that the G. lucidum isolate from the United Kingdom did not 
fall within a well-supported clade with other G. lucidum sequences or related species. The Finnish 
isolates were closely related to the G. tsugae lineage in tef1, rpb2, and β-tub phylogenies. However, 
G. tsugae appears morphologically distinct from the Finnish material. The results suggest that 
G. tsugae, or a species phylogenetically closely related to it, may occur in Finland. But further 
investigation into the relationship between G. tsugae and G. lucidum from Europe will be needed 
to clarify the identity of the laccate Ganoderma species in Finland.
ARTICLE HISTORY 
Received 9 November 2023  
Accepted 5 July 2024 
KEYWORDS 
Basidiomycota; biodiversity; 
Ganodermataceae; fungal 
diversity; Polyporales
INTRODUCTION
Ganoderma P. Karst includes a diversity of wood-decaying 
polyporoid fungal species. Several of these species are 
agents of tree diseases and thus of interest to tree health 
specialists (e.g., Coetzee et al. 2011, 2015; Paterson 2007). 
There are seven Ganoderma species known to occur in 
Europe, namely, G. adspersum (Schulzer) Donk, 
G. applanatum (Pers.) Pat., G. carnosum Pat., G. lucidum 
(Curtis) P. Karst., G. pfeifferi Bres., G. resinaceum Boud, 
and G. valesiacum Boud. Of these, G. adspersum, 
G. applanatum, G. pfeifferi, and G. resinaceum cause 
wood decay in living trees (e.g., Schwarze 2001; Terho 
et al. 2007). The other species are found mostly on dead 
trees and rarely on roots of living trees.
Two Ganoderma species, G. lucidum (subgenus 
Ganoderma) and G. applanatum (subgenus Elfvingia), 
have previously been recorded in Finland (Niemelä 
1982; Niemelä and Kotiranta 1986). Both these taxa are 
known to include species having similar morphology but 
that are genetically distinct. They are therefore usually 
referred to as representing species complexes. In general, 
laccate specimens are referred to as G. lucidum and non- 
laccate specimens as G. applanatum.
In 1881, P. A. Karsten described the genus 
Ganoderma P. Karst, with G. lucidum as the only species 
(Karsten 1881). Karsten (1889) collected several speci-
mens growing on oak (Quercus robur), alder (Alnus sp.), 
and spruce (Picea abies) wood stumps located in 
Ruissalo, Merimasku, and Vaasa (southwest and wes-
tern Finland). The taxonomic history of G. lucidum 
dates back to 1781 when Curtis described Boletus luci-
dus Curtis (now the basionym of G. lucidum) using 
a specimen collected from London, UK, in 1780. The 
species has been reported worldwide based on identifi-
cations relying on morphological characteristics, 
although many of these collections represent other spe-
cies. The holotype specimen has been lost and only an 
illustration of the basidiocarp is available (Curtis 1781), 
CONTACT Marta Cortina-Escribano marta.cortina.escribano@luke.fi
Supplemental data for this article can be accessed online at https://doi.org/10.1080/00275514.2024.2381424.
MYCOLOGIA                                                  
2024, VOL. 116, NO. 6, 1046–1062 
https://doi.org/10.1080/00275514.2024.2381424
© 2024 The Author(s). Published with license by Taylor & Francis Group, LLC.  
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, 
distribution, and reproduction in any medium, provided the original work is properly cited. The terms on which this article has been published allow the posting of the Accepted 
Manuscript in a repository by the author(s) or with their consent.
Published online 12 Sep 2024
complicating efforts to stabilize the correct identity of 
G. lucidum. Attempts to locate a neotype specimen from 
the original type locality (Hyde Park, Peckham, 
London) have failed (Steyaert 1972). Consequently, 
this species lacks a type specimen suitable for DNA 
sequence–based studies that would be required to 
resolve this question.
Application of DNA sequence–based techniques in 
fungal taxonomy has improved the differentiation of 
species that share phenotypic characteristics and that 
therefore cannot be delineated based on morphology 
alone. Earlier studies have made use of single-locus 
sequences to differentiate species in Ganoderma and 
other fungi (Kwon et al. 2016; Moncalvo et al. 1995; 
Wang et al. 2009). However, several authors suggest that 
the analysis of a single or low number of genes may not 
resolve taxonomic questions, as a single (or few) gene 
tree do not necessarily reflect the species phylogeny 
(e.g., Rokas et al. 2003; Sun et al. 2022). Therefore, to 
discriminate between Ganoderma species, multilocus 
phylogenetic analyses have been performed by various 
authors to resolve problems that have arisen when only 
a single locus is used (Cabarroi-Hernández et al. 2019; 
Hapuarachchi et al. 2015; Loyd, Barnes et al. 2018a; Sun 
et al. 2022; Wang et al. 2012; Zhou et al. 2015). Also, 
a genealogical concordance phylogenetic species recog-
nition approach that uses the concordance of different 
gene trees has been employed to resolve species in 
Ganoderma (e.g., Tchotet Tchoumi et al. 2019).
DNA sequences included in previous studies on 
Ganoderma were mainly from genes or regions located 
on the nuclear genome, with few studies including 
sequences from the mitochondrial genome. Sequences 
from the nuclear genome were generated from the 
internal transcribed spacer region (ITS) region (Cao 
et al. 2012; Hennicke et al. 2016; Jargalmaa et al. 2017; 
Loyd, Barnes et al. 2018a; Loyd, Richter et al. 2018b; 
Moncalvo et al. 1995; Park, Kwon, Son, Yoon, Han, 
Nam et al. 2012; Park, Kwon, Son, Yoon, Han et al. 
2012; Wang et al. 2009; Zhou et al. 2015; Zhang et al. 
2017), as well as partial sequences from genes coding the 
elongation factor 1-α (tef1), RNA polymerase II subunit 
(rpb2) (Cao et al. 2012; Jargalmaa et al. 2017; Loyd, 
Barnes et al. 2018a; Loyd, Richter et al. 2018b; Sun 
et al. 2022; Zhou et al. 2015), and partial β-tubulin (β- 
tub) (Hennicke et al. 2016; Park, Kwon, Son, Yoon, Han 
et al. 2012). Other than a small number of ITS 
sequences, there is no robust DNA sequence data set 
for Finnish Ganoderma specimens.
The aim of this study was to clarify the identity of the 
laccate Ganoderma species occurring in Finland. 
Phylogenetic and morphological studies on selected 
boreal-temperate collections from Satakunta and 
Uusimaa provinces in Finland were performed. We 
generated sequences from the ITS region and partial 
tef1, rpb2, and β-tub genes. These sequences were then 
compared with available sequence data of G. lucidum 
specimens from Europe and related species from East 
Asia and North America using phylogenetic methods. 
The morphological characteristics of selected Finnish 
specimens, including specimens from Karsten’s collec-
tion (1889), were investigated and compared with those 
of other regions and other described species.
MATERIALS AND METHODS
Fungal isolates.—Laccate polypore specimens with 
a macromorphology similar to G. lucidum growing on 
Picea abies and Betula pubescens wood stumps were 
collected from Satakunta and Uusimaa provinces in 
Finland (TABLE 1). Dikaryotic strains were isolated 
from 25 specimens by cutting a 0.5 × 0.5 mm piece of 
tissue from the surface of the freshly collected wild 
basidiocarps context and transferring this to 2% malt 
extract agar [MEA: 20 g/L malt extract (VWR 
International LLC, USA) and 20 g/L agar bacteriologi-
cal] medium and incubating the cultures at room tem-
perature. Pure cultures were obtained by repeated 
transfers from the emerging colony margins, and these 
were stored on 5% MEA medium at 5 C. The dikaryotic 
state was evident from the presence of clamped septa in 
the cultures. The fungal isolates are preserved in the 
Culture Collection of Natural Resources Institute 
Finland (LUKE), Helsinki, Finland. Isolate CBS 170.30 
labeled as G. lucidum collected in London and deposited 
by Kenneth St. G. Cartwright was obtained from the 
culture collection of the Westerdijk Fungal Biodiversity 
Institute, Utrecht, the Netherlands, and included in this 
study. The isolate was transferred to 2% MEA medium 
and kept at room temperature.
Herbarium specimens.—Two specimens were col-
lected for morphological characterization, one grow-
ing on P. abies and another on B. pubescens. The 
specimens were dried at 35 C and frozen for 2 weeks 
at −20 C to prevent invertebrate damage. The dried 
basidiocarps were deposited in the Botanical 
Museum of the University of Helsinki after examina-
tion. Herbarium specimens of several species of 
Ganoderma were obtained from the University of 
Turku Herbarium (Turku, Finland) and the Finnish 
Museum of National History Botanical Herbarium 
(Helsinki, Finland) (TABLE 2) for morphological 
studies. The Ganoderma specimens from conifers 
originated from Finland, Estonia, Russia (Central 
MYCOLOGIA 1047
Table 1. Strain names, origin and GenBank accession numbers of the sequences used in the phylogenetic analyses.
Species Isolate Origin ITS tef1 β-tub rpb2 Reference
G. adspersum SFC20141001-16 Korea KY364251 KY393284 – KY393270 Jargalmaa et al. (2017)
G. adspersum LGAM 401 = 
ACAM DD2486
Greece MG706206 MG837829 – MG837781 Fryssouli et al. (NP)
G. annulare KCTC 16803 Brazil JQ520160 – JQ675613 – Park, Kwon, Son, Yoon, Han, Nam et al. (2012)
G. applanatum IUM 3985 Netherlands JQ520162 – JQ675615 – Park, Kwon, Son, Yoon, Han et al. (2012)
G. applanatum SFC20150930-02 Korea KY364258 KY393288 – KY393274 Jargalmaa et al. (2017)
G. australe CMW47785 South Africa MH571686 MH567276 MH567310 – Tchotet-Tchoumi et al. (2019)
G. australe HUEFS:DHCR411 Australia MF436675 MF436677 – – Costa-Rezende et al. (2017)
G. boninense WD 2085 Japan KJ143906 KJ143925 – KJ143965 Zhou et al. (2015)
G. curtisii CBS 100132 USA JQ781849 KJ143927 JQ675617 KJ143967 Cao et al. (2012); Park, Kwon, Son, Yoon, Han, 
Nam et al. (2012); Zhou et al. (2015)
G. flexipes Wei 5491 China JQ781850 – – KJ143968 Cao et al. 2012; Zhou et al. (2015)
G. flexipes Wei 5494 China JN383978 – – – Cao and Yuan (2013)
G. gibbosum GZ14070501 China MK345432 – – MK371436 Hapuarachchi et al. (2019)
G. gibbosum SFC20150918-08 Korea KY364271 KY393291 – KY393278 Jargalmaa et al. (2017)
G. lingzhi Wu 1006-38 China JQ781858 JX029976 – JX029980 Cao et al. (2012)
G. lingzhi Dai 12374 China JQ781867 – – – Cao et al. (2012)
G. lingzhi Cui 9166 China KJ143907 JX029974 – JX029978 Zhou et al. (2015)
G. lingzhi SFC20150624-06 Korea – KY393279 – KY393267 Jargalmaa et al. (2017)
G. lingzhi MFLU 19-2209 Thailand – MN423165 – MN423132 Luangharn et al. (NP)
G. lucidum Gl-4/CMI-UNIBO 
Glu5039
Armenia JN588572 – – – Iotti et al. (NP)
G. lucidum IUM 4303 Bangladesh JQ520182 – JQ675635 – Park, Kwon, Son, Yoon, Han, Nam et al. (2012)
G. lucidum FBE 10458 Bulgaria MG706224 – – MG837797 Fryssouli et al. (NP)
G. lucidum ATCC 46755 Canada JQ520185 – JQ675638 – Park, Kwon, Son, Yoon, Han, Nam et al. (2012)
G. lucidum HKAS:48969 China KC222322 – – – Yang and Feng (2013)
G. lucidum IUM 4242 China JQ520186 – JQ675639 – Park, Kwon, Son, Yoon, Han, Nam et al. (2012)
G. lucidum Cui 9207 China KJ143910 KJ143928 – KJ143970 Zhou et al. (2015)
G. lucidum Cui 14405 China MG279182 MG367574 MG367520 Xing et al. (2018)
G. lucidum DB China KX589245 – – – Zhang et al. (2017)
G. lucidum G.260125-1 China – – PRJNA71455 – Chen et al. (2012)
G. lucidum MT 26/10 Czech 
Republic
KJ143912 KJ143930 – – Zhou et al. (2015)
G. lucidum Dai 11593 Finland JQ781852 – – – Cao et al. (2012)
G. lucidum MUS1 Finland ON598647 MW685379 MW685347 OM810210 This study
G. lucidum MUS2 Finland ON598648 MW685380 MW685348 OM810211 This study
G. lucidum MUS3 Finland ON598649 MW685381 MW685349 OM810212 This study
G. lucidum MUS4 Finland ON598650 MW685382 MW685350 OM810213 This study
G. lucidum MUS5 Finland ON598651 MW685383 MW685351 OM810214 This study
G. lucidum MUS6 Finland MT334583 MW685384 – OM810215 This study
G. lucidum MUS7 Finland ON598652 – MW685352 OM810216 This study
G. lucidum MUS8 Finland ON598653 MW685385 MW685353 OM810217 This study
G. lucidum MUS9 Finland MT334585 MW685386 MW685354 OM810218 This study
G. lucidum MUS10 Finland ON598654 – MW685355 OM810219 This study
G. lucidum MUS12 Finland MT334586 MW685387 MW685356 OM810220 This study
G. lucidum MUS13 Finland ON598655 MW685388 MW685357 OM810221 This study
G. lucidum MUS14 Finland ON598656 MW685389 MW685358 – This study
G. lucidum MUS16 Finland ON598657 MW685390 MW685359 OM810222 This study
G. lucidum MUS17 Finland ON598658 – – – This study
G. lucidum MUS18 Finland ON598659 MW685391 MW685360 OM810223 This study
G. lucidum MUS19 Finland MT334587 MW685392 – – This study
G. lucidum MUS20 Finland ON598660 MW685393 MW685361 OM810224 This study
G. lucidum MUS21 Finland ON598661 MW685394 MW685362 OM810225 This study
G. lucidum MUS23 Finland ON598662 MW685395 MW685363 OM810226 This study
G. lucidum MUS67 Finland – MW685396 MW685364 OM810227 This study
G. lucidum MUS68 Finland – MW685397 MW685365 OM810228 This study
G. lucidum MUS75 Finland MT334584 – MW685366 OM810229 This study
G. lucidum MUS126 Finland – MW685398 – – This study
G. lucidum MUS192 Finland MT334582 – – – Cortina-Escribano et al. (2020)
G. lucidum M9720 France KU310900 – KU310902 – Hennicke et al. (2016)
G. lucidum Rivoire 4195 France KJ143909 – – KJ143969 Zhou et al. (2015)
G. lucidum MUCL 31549 France MG706230 MG837845 – MG837804 Fryssouli et al. (NP)
G. lucidum LGAM 484 = 
ACAM 2013-0022
Greece MG706227 MG837842 – MG837801 Fryssouli et al. (NP)
G. lucidum BCRC36123/ 
ATCC 32471
India EU021459 – – – Wang et al. (2009)
G. lucidum HA2012-001 Iran KX765192 – – – Aghajani et al. (NP)
G. lucidum G1T099 Italy AM269773 – – – Guglielmo et al. (2007)
G. lucidum GlCN04 Italy AM906058 – – – Guglielmo et al. (2008)
G. lucidum WD565 Japan AB462322 – – AB368126 Sotome et al. (2008)
G. lucidum WD2038 Japan EU021456 – – – Wang et al. (2009)
(Continued)
1048 CORTINA-ESCRIBANO ET AL.: PHYLOGENETICS AND MORPHOLOGY OF LACCATE GANODERMA IN FINLAND
and Eastern Siberia), Slovakia, China, Canada, and 
USA, and those from deciduous trees were from 
Finland, Latvia, Sweden, Romania, and China.
DNA extraction, PCR, and sequencing.—Fungal iso-
lates were grown on 50-mm Petri dishes containing 
modified orange serum 2% [MOS: 15 g/L orange 
Table 1. (Continued).
Species Isolate Origin ITS tef1 β-tub rpb2 Reference
G. lucidum RDA-Yeongji-1/ 
ASI 7004
Korea JQ520167 – JQ675620 – Park, Kwon, Son, Yoon, Han, Nam et al. (2012)
G. lucidum IUM 0047 Korea JQ520174 – JQ675627 – Park, Kwon, Son, Yoon, Han et al. (2012)
G. lucidum KACC 42232 Korea KT717954 – – – Kwon et al. (2016)
G. lucidum RYV 33217 Norway Z37096/ 
Z37073
– – – Moncalvo et al. (1995)
G. lucidum FCL191 Poland JQ627589 – – – Siwulski et al. (NP)
G. lucidum FCL188 Poland JN008869 – – – Siwulski et al. (NP)
G. lucidum ZBS1 Russia MF419230 – – – Kokaeva et al. (NP)
G. lucidum GL Slovakia MK415269 – – – Gasparcova et al. (NP)
G. lucidum GL81 Slovenia KC311369 – – – Tang and Zhang (NP)
G. lucidum GLS-1 Spain KT805317 – – – Ozcariz Fermoselle (NP)
G. lucidum Dai 2272 Sweden JQ781851 – – – Cao et al. (2012)
G. lucidum BCRC37033 Taiwan EU021462 – – – Wang et al. (2009)
G. lucidum ATCC 64251 Taiwan JQ520187 – JQ675640 – Park, Kwon, Son, Yoon, Han, Nam et al. (2012)
G. lucidum KCTC 16802 Thailand JQ520188 – JQ675641 – Park, Kwon, Son, Yoon, Han et al. (2012)
G. lucidum K 175217 UK KJ143911 KJ143929 – KJ143971 Zhou et al. (2015)
G. lucidum HMAS86597 UK AY884176 – – JF915436 Wang et al. (2012)
G. lucidum CBS 176.30 UK ON600478 OM810241 OM810242 OM810243 This study
G. lucidum SP26 USA AM269772 – – – Guglielmo et al. (2007)
G. lucidum UMNCA6 USA MG654067 MG754724 – – Loyd, Barnes et al. (2018a)
G. lucidum UMNUT7 USA – MG754726 – – Loyd, Richter et al. (2018)
G. lucidum UMNUT1 USA – MG754725 – – Loyd, Barnes et al. (2018)
G. lucidum UMNUT2 USA MG654068 – – – Loyd, Richter et al. (2018)
G. lucidum GLVN02 Vietnam MN636776 – – – Bui and Nguyen (NP)
G. mirabile CBS 218.36 Philippines – – JQ675645 – Park, Kwon, Son, Yoon, Han, Nam et al. (2012)
G. multipileum CWN 04670 China KJ143913 KJ143931 – KJ143972 Zhou et al. (2015)
G. oregonense ASI 7067 USA JQ520197 – JQ675650 – Park et al. (2012)
G. oregonense CBS 265.88 USA JQ781875 KJ143933 – KJ143974 Cao et al. (2012); Zhou et al. (2015)
G. oregonense UMNAK1 USA – MG754740 – – Loyd, Barnes et al. (2018)
G. resinaceum IUM 3651 Czech 
Republic
JQ520204 – JQ675657 – Park, Kwon, Son, Yoon, Han, Nam et al. (2012)
G. resinaceum LGAM 344 = 
ACAM DD2380
Greece MG706245 MG837853 – MG837816 Fryssouli et al. (NP)
G. resinaceum CBS 194.76/ 
BCRC36147
Netherlands KJ143916 KJ143934 – – Zhou et al. (2015)
G. resinaceum CBS 152.27 UK JQ520200 – JQ675653 – Park, Kwon, Son, Yoon, Han, Nam et al. (2012)
G. resinaceum HMAS86599 UK AY884177 JF915435 Wang et al. (2012)
G. sichuanense HMAS42798 China JQ781877 – – – Cao et al. (2012)
G. sichuanense HMAS252081 China KC662402 Yao et al. (2013)
G. tsugae CBS 223.48 Canada Z37054/ 
Z37079
– – – Moncalvo et al. (1995)
G. tsugae KCTC6457/ATCC 
64795
Canada JQ520215 – JQ675668 – Park, Kwon, Son, Yoon, Han, Nam et al. (2012)
G. tsugae Dai 3937 China JQ781853 – – – Cao et al. (2012)
G. tsugae Dai 15529 China MG279197 MG367590 – MG367536 Xing et al. 2018
G. tsugae JV 0307/4 J China – MG367562 – MG367505 Xing et al. (2018)
G. tsugae Cui 14112 China MG279196 MG367587 – MG367534 Xing et al. (2018)
G. tsugae S90 China – PRJNA445345 PRJNA445345 – NA
G. tsugae KL20 India FJ655478 – – – Arulpandi and Kalaichelvan (NP)
G. tsugae AFTOL-ID 771 USA? DQ206985 – DQ408116 Matheny et al. (2007)
G. tsugae ASI 7064/KU- 
4018
USA JQ520216 – JQ675669 – Park, Kwon, Son, Yoon, Han, Nam et al. (2012)
G. tsugae KCTC6290/ATCC 
64794
USA – – JQ675673 – Park, Kwon, Son, Yoon, Han, Nam et al. (2012)
G. tsugae Dai 12760 USA KJ143920 KJ143940 – KJ143978 Zhou et al. (2015)
G. tsugae UMNAZ9 USA MG654321 MG754763 – – Loyd, Barnes et al. (2018)
G. tsugae UMNMI30 USA – MH025362 – MG754871 Loyd, Richter et al. (2018)
G. tsugae UMNNC4 USA MG654329 MG754765 – MG754872 Loyd, Barnes et al. (2018)
G. tsugae UMNMN7 USA MG654327 – – – Loyd, Richter et al. (2018)
G. tsugae UMNWI22 USA MG654344 – – – Loyd, Barnes et al. (2018)
T. colossus CGMCC5.763 Philippines JQ081068 – – JQ081070 Wang et al. (2012)
T. colossus TC-02 Vietnam KJ143923 KJ143943 – – Zhou et al. (2015)
T. suaveolens CBS 446.61 Austria – – FJ410378 – Lesage-Meessen et al. (2011)
Note. NP = Not published.
MYCOLOGIA 1049
serum agar (Berner Oy, Maharashtra, India), 20 g/L Bacto 
malt extract (Berner), 8 g/L Berner dextrose (Berner), 9 g/L 
Bacto agar (Berner)] medium or 2% MEA medium and 
a layer of sterilized cellophane. After 2–3 weeks of growth, 
DNA was extracted from the isolates using PrepMan Ultra 
Sample following the manufacturer’s protocol (Applied 
Biosystems, Fosters City, California). DNA concentration 
was determined using NanoDrop ND-1000 spectrophot-
ometer (NanoDrop Technologies, Madison, Wisconsin). 
The DNA was diluted to a working concentration of 50 ng/ 
µL with SABAX water (Adcock Ingram, Midrand, South 
Africa). Four gene regions were amplified for the isolates. 
These included the ITS and regions of the tef1, rpb2, and β- 
tub genes. The primers used to amplify the ITS region were 
ITS1-F (5′-CTT GGT CAT TTA GAG GAA GTA A-3′) 
forward primer (Gardes and Bruns 1993) and ITS4 (5′- 
TCC TCC GCT TAT TGA TAT GC-3′) reverse primer 
(White et al. 1990). The primers used to amplify the gene 
for the tef1 region were EF595F (5′-CGT GAC TTC ATC 
AAG AAC ATG-3′) forward primer and EF1160R (5′- 
CCG ATC TTG TAG ACG TCC TG-3′) reverse primer 
(Kauserud and Schumacher 2001). The β-tubulin gene 
region was amplified using β-tubF (5′-CCG GTG CAG 
GCA TGG GTA CC-3′) forward primer and β-tubR (5′- 
TGA AGA CGG GGG AAG GGA AC-3′) reverse primer 
(Park, Kwon, Son, Yoon, Han et al. 2012). The primers 
used to amplify the rpb2 gene region were G-RPB2-F1 (5′- 
CAT CGA GTT CTT GGA GGA GTG G-3′) forward 
primer and G-RPB2-R1 (5′-CGG AAT GAT GCT GGC 
ACA GAC A-3′) reverse primer (Cao et al. 2012).
Polymerase chain reaction (PCR) mixtures included 
2.5 μL of 10× KAPA Taq BUffer A (Bioline, London, 
United Kingdom), 0.5 μL of 25 mM MgCl2 (Bioline), 
0.5 μL of 10 mM dNTP Mix (Bioline), 1 μL of 10 μM 
forward primer, 1 μL of 10 μM reverse primer, 0.1 μL of 
5 U/μL KAPA Taq DNA polymerase (Bioline), and 
0.5 μL of template DNA (1 µL for tef1 and rbp2 gene 
regions). The volume of the mixture was adjusted to 
25 μL with SABAX water. PCRs used the following 
thermal cycling protocol: initial denaturation (95 C for 
3 min, 1 cycle), denaturation, annealing, extension (95 
C for 30 , 52 C for 30 , 72 C for 1 min, respectively; 35 
cycles), final extension (72 C, 10 min, 1 cycle), and hold 
at 4 C. After PCR amplification, amplicons were stained 
with GelRed nucleic acid gel stain (Biotium, Hayward, 
California) and separated using electrophoresis in a 1% 
agarose gel with 10× Tris-borate-EDTA (TBE) buffer. 
PCR products were observed under ultraviolet (UV) 
light and their sizes compared with a GeneRuler 100 
bp Plus DNA ladder (Thermo Scientific, Vilnius, 
Lithuania) using Image Lab software (Bio-Rad, 
Hercules, California).
PCR products were purified with 8 μL of ExoSAP 
reagent [5 µL of Exo (exonuclease I; Thermo Fisher 
Scientific, Vilnus, Lithuania), 100 µL of SAP 1 U/µL 
(shrimp alkaline phosphatase; Roche, Mannheim, 
Germany), and 895 µL H2O] to the remaining 20 µL of 
post-PCR product. The mixture was incubated at 37 
C for 15 min to degrade remaining primers and nucleo-
tides. A second incubation period at 80 C for 15 min was 
applied to inactivate ExoSAP reagent. The sequencing 
reactions of the purified PCR products were performed 
in a 12-µL reaction mixture [0.5 µL of BigDye 
Terminator v3.1 Ready Reaction Mix (Perkin-Elmer 
Applied Biosystems, Warrington, UK), 2.1 µL sequen-
cing buffer, 1 µL of 10 mM primer, and 2 µL purified 
Table 2. Herbarium specimens examined.
Species Specimen Collector/collection Date Country Region Host Host genus
G. carnosum H7050121 G. Silaghi 16 September 1955 Romania Transilvania Deciduous Quercus
H7055041 P. Vampola exsiccati 139 30 September 1994 Slovakia Badin Conifer Abies
G. lucidum H6049741 P.A. Karsten 1858 Finland Turku Deciduous Quercus
TFU.186924 P. Kunttu 5950 19 November 2009 Finland Kaarina Conifer Picea
H6135371 P. Veteli 410 15 August 2017 Finland Porvoo Conifer Picea
TFU.186580 P. Kunttu 5752 7 October 2009 Finland Kaarina Conifer Picea
TFU.110754 M. Kyröläinen 239 30 August 1993 Finland Vaasa Conifer Picea
TFU.205608 P. Kunttu 7691 10 August 2011 Finland Kotka Deciduous Prunus
H6135369 E. Kalska 2017 Finland Ylöjärvi Conifer Larix
H6135370 P. Veteli 764 1 August 2018 Finland Helsinki Deciduous Betula
TFU.34356 U. Kalamees & K. Kalamees 30 April 1958 Estonia Viimsi-Krillimäe Conifer Picea
TFU.34357 J. Smarods 14 April 1936 Latvia Vidzene Deciduous Betula
H7049337 H. Kotiranta 28986 23 August 2007 Russia Sakhalin Conifer Abies
H7044873 H. Kotiranta 21344 16 August 2006 Russia Perm region Conifer Picea
H7029500 D. Hildebrandt K-05-09 & I. Stepanchikova 5 August 2009 Russia Kamtschatca Conifer Larix
H7050168 Y.C. Dai 2062 13 September 1995 China NA Conifer Pinus
H7050209 Z. Lu 37. J.-D. Zhao & X.-Q. Zhang 1979 China Heilongjiang Unknown Unknown
TFU.72720 H. Karlsted 9 May 1972 Sweden NA Deciduous Alnus
G. oregonense H7009574 O. Miettinen 19004.1 21 October 2014 USA Washington Conifer Tsuga
H7050171 T. Ahti 51047 & F.M. Rhoades 29 March 1992 USA Washington Conifer Tsuga
G.tsugae H7050170 J. H. Ginns 8800 2 September 1986 Canada Ontario Conifer Tsuga
H7008236 O. Miettinen 16813 14 April 2022 USA Massachusetts Conifer Tsuga
1050 CORTINA-ESCRIBANO ET AL.: PHYLOGENETICS AND MORPHOLOGY OF LACCATE GANODERMA IN FINLAND
PCR product]. The thermal cycling conditions followed 
the protocol of the manufacturer. The PCR products 
were precipitated with ethanol and 3 M pH 4.6 sodium 
acetate and dried in a laminar flow overnight. DNA 
sequencing was performed with an ABI Prism 3100 
DNA analyzer (Applied Biosystems, Foster City, 
California) at the DNA Sequence Facility of the 
University of Pretoria.
The ITS and rpb2 gene regions for some of the iso-
lates and all the gene regions of strain CBS 176.30 were 
sequenced in the Viikki laboratory at the Natural 
Resources Institute Finland (Helsinki). The protocol 
used was similar to that described above, but Phusion 
Green High-Fidelity DNA Polymerase (Thermo Fisher 
Scientific, Vilnus, Lithuania) was used rather than 
KAPA Taq in the PCR master mix. The PCR cycling 
parameters and PCR cleanup protocol were the same as 
those mentioned above, and the PCR products were 
sequenced at Macrogen (Amsterdam, The Netherlands).
DNA sequence and phylogenetic analyses.—The 
forward and reverse sequences were assembled using 
Geneious 10.2.6 (Biomatters, Auckland, New 
Zealand). The consensus sequences were queried 
against the GenBank nucleotide database (https:// 
www.ncbi.nlm.nih.gov/) using BLASTn search for pre-
liminary identification of the isolates. Consensus 
sequences were deposited in GenBank (TABLE 1). 
Data sets were generated for each of the four loci 
(ITS, tef1, β-tub, and rpb2) separately. The data set 
included sequences from this study together with 
reference sequences from previous studies obtained 
from GenBank (Cao et al. 2012; Loyd, Barnes et al. 
2018; Loyd, Richter et al. 2018; Park, Kwon, Son, 
Yoon, Han, Nam et al. 2012; Park, Kwon, Son, Yoon, 
Han et al. 2012; Wang et al. 2012; Zhou et al. 2015). 
Where available, the data sets included sequences 
from type strains and the closely related species 
G. curtisii (Berk.) Murrill, G. lingzhi, G. oregonense 
Murrill, and G. tsugae Murrill as well as strains from 
different geographic locations to show the heterogene-
ity among Ganoderma species (TABLE 1). Available 
sequences of G. lucidum from Europe, North America, 
and East Asia were included in all data sets. Available 
sequences representing G. tsugae from isolates origi-
nating in Canada, USA, China, and India (only ITS 
region) were also included in all data sets. Where 
available, representative sequences for G. adspersum, 
G. annulare (Fr.) Gilb., G. applanatum, Ganoderma 
australe (Fr.) Pat., G. boninense Pat., G. flexipes Pat., 
G. gibbosum (Blume & T. Nees) Pat., G. mirabile 
(Lloyd) C.J. Humphrey, G. multipileum Ding Hou, 
G. resinaceum, and G. sichuanense J.D. Zhao & X.Q. 
Zhang were included in all data sets.
Tropical Ganoderma species were not included in the 
final data sets after running preliminary phylogenetic 
analyses (data not shown), as they are very distinct from 
the European Ganoderma spp. The names of species 
shown in TABLE 1 correspond with those given in the 
GenBank records. Tomophagus colossus (Fr.) Murrill 
and Trametes suaveolens (L.) Fr. were selected as out-
group taxa (Tchotet Tchoumi et al. 2018). Individual 
data sets were compiled and edited using Molecular 
Evolutionary Genetic Analysis (MEGA) 10.0.5 (Kumar 
et al. 2018). The data sets were aligned using the online 
version of MAFFT 7 (Katoh and Standley 2013). The 
alignment strategies consisted of FFT-NS-i for the ITS 
data set and G-INS-i (default parameters) for the other 
data sets.
The phylogenetic analyses were performed using 
maximum likelihood (ML), maximum parsimony 
(MP), and Bayesian inference (BI) methods for each 
data set. ML analysis was conducted with the online 
version of PhyML 3.0 (http://www.atgc-montpellier.fr/ 
phyml/; Guindon et al. 2010), using Smart Model 
Selection (SMS; Lefort et al. 2017) and Akaike informa-
tion criterion (AIC) for substitute model selection. 
Approximate likelihood-ratio test (aLRT; Anisimova 
and Gascuel 2006) was selected for branch support 
estimation.
MP trees were inferred using PAUP 4.0a169 
(Swofford 2003) with default parameters for parsi-
mony analysis. A heuristic search with tree bisection 
reconnection (TBR) branch swapping was used to 
search tree space. The starting trees were obtained by 
random stepwise additions of sequences. Branches 
having a maximum length of zero were collapsed, and 
the gaps were considered as missing data. The max-
imum most parsimonious trees to be retained were set 
to 100.
BI analyses were carried out with MrBayes 3.2.7a 
(Ronquist et al. 2003) using a Markov chain Monte 
Carlo (MCMC) simulation. The GTR+G+I substitution 
model was preselected using the AIC in MrModeltest 
3.0 (Nylander 2004). Four runs of MCMC chains for 
5 million generations were done using a sample fre-
quency of every 100th generation. The first 25% of the 
trees sampled were discarded as burn-in, and the poster-
ior probabilities were calculated for the remaining trees. 
Effective sampling (ESS) and convergence of trees and 
posterior probabilities were assessed in Tracer 1.7.2 
(Rambaut et al. 2018). The phylogenetic trees generated 
using the different methods were visualized in MEGA 
and post-edited using Inkscape 1.1.1 (https://inkscape. 
org/).
MYCOLOGIA 1051
Morphological characterization.—The microscopic 
structures of the selected specimens (TABLE 2) were 
studied with a Leica DMLB 100T light microscope 
(Mannheim, Germany). The spore measurements were 
made using phase-contrast optics at ×1000 magnifica-
tion in oil immersion (Leica  Immersion Oil). Melzer’s 
reagent was used to mount the spores before they were 
measured. The spore measurements included the myx-
osporium, the external layer of the spore wall structure. 
Fifty spores per specimen were measured, and 80% 
confidence was used for reporting the variation limits. 
Variation in spore size is reported as in Miettinen et al. 
(2006): number of spores measured (n), mean length 
(L), mean width (W), average length divided by average 
width (Q), and length and width ratio of individual 
spores (Q’). The whole range variation of L, W, and Q’ 
is reported in parentheses, and the 80% confidence 
range is given without parentheses.
The terminology used to describe the spore wall 
structure, spore shape, and spore ornamentation fol-
lowed Clémençon (2012), Niemela (2016), and Torres- 
Torres and Guzmán-Dávalos (2012), respectively. The 
shape of the dermis and the arrangements of the pilei-
pellis dermal elements were described following the 
terminology of Torres-Torres and Guzmán-Dávalos 
(2012) and Clémençon et al. (2012). The general 
descriptive terms for the macroscopic features followed 
Niemela (1982), and Furtado (1981).
RESULTS
DNA sequences and phylogenetic analyses.—The 
ITS, tef1, β-tub, and rpb2 gene regions were successfully 
amplified for most of the isolates. The amplicon size was 
approximately 650 bp long for the ITS region, 550 bp 
long for tef1, 410 bp long for β-tub, and 575 bp long for 
rpb2 genes. The ITS, tef1, and β-tub gene regions were 
identical for all the Finnish isolates, and nearly identical 
for the rpb2 gene region. ITS sequences had high 
sequence similarity (99–100%) to sequences of 
G. lucidum and G. tsugae on GenBank using BLASTn 
searches. The ITS sequence from the UK isolate (CBS 
170.30) showed high sequence similarity to G. carnosum 
and G. oregonense. All Finnish strains sequenced in this 
study grouped together in the phylogenetic trees (FIGS. 
1–4). The topology was similar for all the phylogenetic 
trees generated by ML, MP, and BI. Therefore, only the 
ML tree for each data set is presented, together with 
aLRT, bootstrap, and posterior probability values.
The ITS sequence data set included 82 taxa with two 
outgroup taxa (TABLE 3). The Finnish and UK 
sequences generated in this study formed a strongly 
supported clade (ML-aLRT and MP bootstrap values 
≥95% and BI-PP ≥0.95) together with 26 isolates of 
G. lucidum from Europe, North America, and East 
Asia, 13 sequences for G. tsugae from East Asia and 
North America, and one sequence of G. oregonense 
(FIG. 1). Within this large clade, sequences of 
G. tsugae originating from India, USA, and Canada 
grouped in a subclade with strong statistical support 
from ML (aLRT = 98%) and BI (PP = 0.95) analyses. 
Two isolates, also labeled as G. tsugae, one from north-
ern Arizona, USA (Loyd, Barnes et al. 2018a), and one 
from Konkuk University, also of USA origin (Park, 
Kwon, Son, Yoon, Han et al. 2012), and three isolates 
from China (Xing et al. 2018) grouped outside of this 
subclade. The phylogeny based on ITS sequence data 
was the only one that separated G. tsugae originating 
from USA from the Chinese G. tsugae. The remainder of 
the subclades in the phylogenetic tree were only sup-
ported in the ML analysis. Closely related species there-
fore could not be separated with confidence based on 
ITS sequence data.
The tef1 sequence data set included 36 taxa and one 
outgroup taxon (TABLE 3). The phylogenetic tree gen-
erated from tef1 sequence data (FIG. 2) grouped the UK 
isolate (CBS 176.30) within a major clade strongly sup-
ported by the three phylogenetic methods (aLRT branch 
support = 100%, MP bootstrap = 100%, and PP = 1) and 
included sequences of G. tsugae, G. lucidum, and 
G. oregonense. Within this clade, the species were sepa-
rated into respective monophyletic clades with strong 
statistical support. Sequences for the Finnish isolates 
grouped with those of G. tsugae within a strongly sup-
ported clade. This clade included G. tsugae isolates from 
Connecticut, northern Arizona, North Carolina, and 
Michigan, USA, and from Jilin, China. Sequences of 
G. lucidum from UK, USA, France, Greece, China, and 
Czech Republic grouped in a strongly supported mono-
phyletic clade. The third clade included G. oregonense 
isolates originating from USA. In contrast to the ITS 
phylogeny, the tef1 phylogeny successfully separated 
closely related species. However, the UK isolate (CBS 
176.30) was not placed in any of these clades and its 
identity remains unclear.
The rpb2 data set consisted of 34 taxa and one out-
group taxon (TABLE 3). Like the tef1 phylogeny, the 
rpb2 analyses placed the Finnish and the UK G. lucidum 
isolates within a larger clade that included sequences 
from G. lucidum, G. tsugae, and G. oregonense (FIG. 3). 
The phylogenetic analyses also separated the sequences 
of G. lucidum in a subclade that was strongly supported 
by the three methods (aLRT branch support = 98%, MP 
bootstrap = 88%, and PP = 0.96). The Finnish isolates 
formed a clade with isolates representing G. tsugae from 
1052 CORTINA-ESCRIBANO ET AL.: PHYLOGENETICS AND MORPHOLOGY OF LACCATE GANODERMA IN FINLAND
Figure 1. Maximum likelihood analyses based on the ITS sequence data. Branch support (aLRT) for ML and bootstrap values for MP 
higher than 70% from 1000 replicates are indicated in the nodes. Values below 70% are indicated with an asterisk. BI posterior 
probabilities higher than 0.95 are indicated by thick branch lines. Isolates sequenced in this study are indicated in bold.
MYCOLOGIA 1053
China and USA, but it had statistical support only from 
the aLRT analysis (89%). Similar to the tef1 phylogeny, 
the sequence generated in this study for the G. lucidum 
UK isolate did not group within the G. lucidum clade.
The β-tub data set included 24 taxa and one outgroup 
taxon (TABLE 3). The β-tub phylogeny formed 
a strongly supported major clade (aLRT branch sup-
port = 100%, MP bootstrap = 97%, and PP = 1) includ-
ing isolates of G. lucidum from Korea, Thailand, China, 
and Bangladesh, a G. curtisii isolate from USA, and 
a G. tsugae isolate also from USA (FIG. 4). Isolate CBS 
176.30 from the UK grouped within a well-supported 
major clade that included sequences from strains of 
G. tsugae and a G. lucidum isolate from Canada (aLRT 
branch support = 100%, MP bootstrap = 95%, and PP = 
0.97). Within this major clade, the Finnish isolates 
formed a strongly supported subclade (aLRT = 100%, 
MP bootstrap = 96%, and PP = 1) with isolates repre-
senting G. tsugae from Canada and China. Only one β- 
tub sequence from European G. lucidum material was 
available in GenBank (strain M9720 from France), and 
it did not group within the East Asian G. lucidum clade 
nor the G. tsugae clade.
Morphological characterization.—The Finnish iso-
lates were identical based on DNA sequence compari-
sons, and the basidiocarps collected displayed only 
expected phenotypic variation (i.e., color range and 
shape). The basidiocarps of the Finnish specimens are 
clearly stipitate, large, and flabelliform. The context of 
the Finnish specimens presents creamy white color on 
the stipe and subdermal part, and light brown to beige 
in the layer next to the concolourous tubes. The speci-
men H6135370 (southern Finland) is macroscopically 
representative of G. lucidum as understood in Finland 
(FIG. 6); however, it is also a good example of the 
problem associated with morphological variation in 
Ganoderma spp. Specimen H6135370 has consistently 
more narrow spores than is usual for Finnish 
G. lucidum, with average width of only 6.0 microns, 
and average Q-value of 1.7. This is likely due to the 
specimen being in the beginning of prime sporulation 
phase at the time of the collection. The other end of the 
spectrum is Karsten’s oldest collection, H6049741, with 
spore size with an average of 10.3 × 6.9 Microns µm, 
Q 1.5 (TABLE 4). The width of the spore in H6049741 
is, however, still within the range of typical Finnish 
laccate Ganoderma (6–7 microns).
The Finnish isolates grouped together with 
G. tsugae in the phylogenetic trees, but comparisons 
of spore and basidiocarp morphology showed that 
they are slightly distinct from each other. G. tsugae 
does not differ much in spore dimensions from the 
Finnish specimens (TABLE 4; FIG. 5) but often 
appears to have slightly longer echinulate. Moreover, 
G. tsugae may have on average slightly longer hyme-
nodermis terminal elements. This is, however, highly 
dependent on basidiocarp stage. Macroscopically, 
G. tsugae has often a deeper color and shinier appear-
ance than the Finnish herbarium specimens and all 
the specimens collected in this study. G. carnosum 
fruiting bodies have a darker color and wider spores 
(>7 microns width) compared with laccate 
Ganoderma originating in Finland.
DISCUSSION
The G. lucidum complex is a taxonomically complicated 
group that invokes considerable debate regarding spe-
cies boundaries. This arises from a lack of type material, 
an absence of cultures and, thus, reliable sequence data, 
as well as the fact that the complex includes several 
species with overlapping morphological characteristics. 
In this study, we focused on the phylogenetic analysis 
and morphological descriptions of north European bor-
eal-temperate material resembling G. lucidum. The stu-
died gene regions were nearly identical for all the 
Finnish isolates and grouped together in the phyloge-
netic trees. Surprisingly, the Finnish material, widely 
assumed to correspond with the European G. lucidum, 
grouped with G. tsugae in the phylogenetic analyses. 
However, the morphology of the Finnish material was 
slightly different from that of G. tsugae from North 
America and northeast Asia.
There are two nomenclatural outcomes for the 
Finnish laccate Ganoderma species. If the laccate 
Ganoderma material from Finland is indeed conspecific 
with North American G. tsugae, G. valesiacum should be 
considered the valid name, with G. tsugae as a later 
synonym, as was previously suggested (Adaskaveg and 
Gilbertson 1986; Stalpers 1978). This nomenclatural 
proposal requires further study of authentic 
G. valesiacum specimens. Alternatively, the Finnish lac-
cate specimen represents G. lucidum sensu stricto. 
Given the lack of type material from the original type 
locality (UK), an epitype should be selected for this 
species. The basionym of G. lucidum is a sanctioned 
name by Fries (1821), P. lucidus. Thus, morphological 
examination of material noted by Fries and obtaining 
DNA sequence data for the material could aid in pro-
viding an epitype of G. lucidum. If the P. lucidus mate-
rial collected by Fries exists, sequence data from that 
specimen would resolve the relationship with the 
Finnish laccate Ganoderma species.
1054 CORTINA-ESCRIBANO ET AL.: PHYLOGENETICS AND MORPHOLOGY OF LACCATE GANODERMA IN FINLAND
Ganoderma tsugae was first described by Murrill 
(1902) as a species occurring predominantly on Tsuga 
canadensis (Pinaceae). This tree species is confined to 
North America; hence, it has been assumed that 
G. tsugae is native to North America (Loyd, Barnes 
et al. 2018a). Ganoderma tsugae has been predominantly 
reported on conifers such as Abies spp. and Larix spp., 
and it is assumed being strictly associated with these 
Figure 2. Maximum likelihood phylogenetic tree derived from the tef1 sequence data including the isolates used for DNA sequencing 
in this study (in bold). Branch support (aLRT) and bootstrap values ≥70% from 1000 replicates are indicated in the nodes for ML and 
MP, respectively. Values below 70% are indicated with an asterisk. BI posterior probabilities ≥0.95 are shown in thick lines.
MYCOLOGIA 1055
hosts (Loyd, Barnes et al. 2018a; Xing et al. 2018). 
Adaskaveg and Gilbertson (1988) reported G. tsugae 
occurring also on Betula spp. in North America. In his 
description of G. tsugae, Murrill (1902) noted that the 
specimens found by Karsten (1881) growing on Picea 
excelsa could have been G. tsugae rather than 
“Ganoderma pseudoboletus.” However, Karsten (1881) 
reported G. lucidum (“G. pseudoboletus”) occurring on 
both conifer and deciduous trees. Our results support 
the observation of Karsten (1881), in which he reported 
a single species (G. lucidum in Fungi Fenniae Exsiccati) 
growing on both hardwoods and conifers in Finland. As 
G. lucidum commonly grows on conifer substrates in 
Finland, G. carnosum, a European laccate species 
growing preferably in conifer hosts, has been actively 
looked for (Niemela and Kotiranta 1986). However, to 
date, no basidiocarps having spore sizes of G. carnosum 
have been recorded from Finland.
The basidiocarp morphology of the Finnish speci-
mens considered in this study was slightly different 
from that of G. tsugae. Basidiocarps of the G. tsugae 
specimens examined often presented a deeper color and 
shinier appearance than the Finnish G. lucidum mate-
rial. Overholts (1953) noted similar morphological dif-
ferences when comparing specimens thought to 
represent P. tsugae and P. lucidus in North America 
and concluded that they are indeed separate species. 
The spore characteristics showed that the North 
Figure 3. Maximum likelihood phylogenetic tree constructed from the rpb2 data set. Isolates sequenced in this study are in bold. 
Branch support (aLRT) and bootstrap values higher than 70% are indicated in the nodes (ML/MP, respectively). Values below 70% are 
indicated with an asterisk. BI posterior probabilities higher than 0.95 are indicated by thick lines in the branches.
1056 CORTINA-ESCRIBANO ET AL.: PHYLOGENETICS AND MORPHOLOGY OF LACCATE GANODERMA IN FINLAND
American G. tsugae spores have similar dimensions to 
those of the Finnish specimens in this study, but with 
coarser ornamentation. Ganoderma oregonense, 
a species closely related to G. tsugae, has generally larger 
spores (Adaskaveg and Gilbertson 1988; Atkinson 1908; 
Murrill 1908) compared with the Finnish material 
examined in this study. However, the spore dimensions 
may differ at different sporulation phases of the same 
specimen. Moreover, the natural variation within the 
populations and the influence of the environmental 
conditions could also affect morphological characteris-
tics of the basidiocarps (Adaskaveg and Gilbertson 
1986). Such variation is well known, and Nuss (1982), 
for example, reported differences in size and appearance 
of early and later spores from several Ganoderma spe-
cies, including G. lucidum.
Previous phylogenetic studies have separated the 
North American G. tsugae from European G. lucidum 
(Loyd, Barnes et al. 2018a; Sun et al. 2022; Zhou et al. 
2015). The ITS data generated in the present study did 
not delimit strains of European G. lucidum, East Asian 
G. lucidum, North American G. tsugae, and Chinese 
G. tsugae in distinct clades. The tef1, rpb2, and β-tub 
phylogenies separated the European G. lucidum and the 
Finnish strains in different clades, the latter grouping 
together with G. tsugae. This is consistent with a recent 
taxonomic revision of Ganodermataceae by Sun et al. 
(2022) in which it was shown that sequence data for the 
Figure 4. Maximum likelihood phylogenetic tree generated from the β-tub data set. Isolates used for DNA sequencing in this study are 
in bold. ML branch support (aLRT) followed by MP bootstrap values above 70% are indicated in the nodes. Values below 70% are 
indicated with an asterisk. BI posterior probabilities above 0.95 are indicated in the branches (bold lines).
MYCOLOGIA 1057
ITS region alone do not differentiate Ganoderma species 
sufficiently. Previous studies have demonstrated that for 
species in the G. lucidum complex, tef1 and rpb2 loci are 
more informative than the ITS region (Loyd, Barnes 
et al. 2018a; Loyd, Richter et al. 2018b; Sun et al. 2022).
There are some constraints when constructing phy-
logenetic trees from tef1, rpb2, and β-tub gene regions 
for taxonomic purposes. Most important is the lack of 
sequence data from most or all loci for some crucial 
species belonging to the G. lucidum complex (i.e., 
G. carnosum, G. mongolicum Pilát, and G. valesiacum). 
This results in phylogenies based on only the ITS or one 
additional locus. Rokas et al. (2003) showed that a single 
locus or low number of loci does not provide support in 
resolving phylogenetic questions due to incongruent 
tree topologies that can be generated from different 
DNA regions. Thus, we suggest employing the genealo-
gical concordance phylogenetic species recognition 
Figure 5. Spore morphology variation of specimens representing Ganoderma lucidum (1) H7704873, (2) H6049741, (3) H7029500, (4) 
H7044337, (5) H7050168, (6) H7050209, (7) H6135371/MUS11, (8) TFU.186580, and (9) kalske2017; Ganoderma tsugae (10) H7050170; 
Ganoderma oregonense (11) H7050171; and Ganoderma carnosum (12) H7055041.
Table 3. Statistics resulting from the phylogenetic analyses.
Data set No. of taxa No. of bp
Maximum parsimony Maximum likelihood
PCC PIC No. of trees Tree length CI RI RC HI Substitution model NST
ITS 84 616 423 137 100 383 0.68 0.898 0.61 0.321 HKY85+G+I 4
tef1 37 583 414 121 100 338 0.68 0.853 0.58 0.32 GTR+G 4
rpb2 35 590 428 122 100 315 0.65 0.832 0.54 0.352 GTR+I 1
β-tub 25 413 396 59 4 188 0.73 0.822 0.6 0.271 GTR+G 4
Note. PIC = number of parsimony informative characters; CI = consistency index; RI = retention index; RC = rescaled consistency index; HI = homoplasy index; 
NST = number of substitution rate categories.
1058 CORTINA-ESCRIBANO ET AL.: PHYLOGENETICS AND MORPHOLOGY OF LACCATE GANODERMA IN FINLAND
approach to further study the relationships between the 
Finnish G. lucidum and related species.
The concept of G. lucidum sensu stricto is ambigu-
ous, leading to a difficult delimitation between closely 
related species. Host-tree specificity, morphological 
characteristics, interfertility between isolates, and phy-
logenetic and phylogenomic analyses are some aspects 
that need to be considered for species recognition. The 
Figure 6. Basidiocarp morphology of specimen H6135370 collected from Betula sp. stump in the region of Helsinki.
Table 4. Basidiospore characteristics of Ganoderma specimens.
Species Specimen Country n Microns (µm) L variation Microns (µm) L avg. Microns (µm) W variation Microns (µm) W avg. Q’ variation Q avg. Host genus
G. carnosum H7050121 Romania 37 (9.7)10.4–11.7(12.3) 11.1 (6.6)7–7.8(8) 7.4 (1.3)1.4–1.6 1.5 Quercus
H7055041 Slovakia 42 (8.9)10.6–12.6(13.2) 11.6 (6.5)7.1–7.9(8.3) 7.5 (1.3)1.4–1.7(1.8) 1.5 Abies
G. lucidum H6049741 Finland 50 (8.8)9.6–10.9(11.3) 10.3 (5.9)6.5–7.3(7.5) 6.9 (1.4)1.42–1.6(1.7) 1.5 Quercus
TFU.186924 Finland 50 (10)10.1–11.8(12.7) 11 (5.8)6.4–7.5(9) 7 (1.4)1.44–1.7(1.9) 1.6 Picea
H6135371 Finland 50 (9.4)9.8–11.2(11.5) 10.6 (6.3)6.5–7.2(7.6) 6.9 (1.4)1.5–1.6(1.7) 1.5 Picea
TFU.186580 Finland 50 (10)10.4–11.8(12.4) 11.2 (6.3)6.6–7.5(7.8) 7.1 (1.4)1.5–1.6(1.8) 1.6 Picea
TFU.110754 Finland 50 (8.8)10–11.7(12.8) 10.9 (5.8)6.4–7.2(7.5) 6.8 (1.4)1.5–1.7(1.8) 1.6 Picea
TFU.205608 Finland 50 (8.4)9.2–10.5(10.8) 9.8 (5.6)6.2–7.1(7.6) 6.7 (1.3)1.4–1.6(1.7) 1.5 Prunus
H6135369 Finland 50 (8.7)9.3–11.6(12.9) 10.6 (6.1)6.4–7.2(7.5) 6.8 (1.2)1.4–1.7(1.9) 1.6 Larix
H6135370 Finland 30 (8.8)9.0–11.0(11) 10.0 (5.3)5.8–6.1(6.3) 6.0 (1.4)1.5–1.8(1.9) 1.7 Betula
TFU.34356 Estonia 50 (9.6)10.1–11.6(12.4) 10.9 (6)6.5–7.3(8) 7.0 (1.3)1.5–1.7(1.8) 1.6 Picea
TFU.34357 Latvia 30 (8.9)9.3–10.9(11) 10.1 (5.9)6.3–7.2(7.4) 6.8 (1.3)1.4–1.6 1.5 Betula
H7049337 Russia 40 (8.7)9.3–11(11.4) 10.1 (5.4)5.6–6.9(7.2) 6.2 (1.4)1.5–1.7(1.9) 1.6 Abies
H7044873 Russia 50 (8.6)9.1–11.1(12.3) 10.0 (5.5)6–7.3(8.2) 6.6 (1.3)1.4–1.6(1.8) 1.5 Picea
H7029500 Russia 50 (8.9)9.5–11.3(12) 10.4 (6)6.2–7.1(7.6) 6.7 (1.4)1.45–1.7 1.6 Larix
H7050168 China 50 (8.7)9.2–10.5(10.8) 9.8 (5.7)6–6.7(6.9) 6.4 (1.4)1.5–1.7(1.9) 1.6 Pinus
H7050209 China 50 (8.6)9.9–11.5(12.3) 10.7 (5.3)5.9–7(7.2) 6.5 (1.4)1.6–1.8(2) 1.7 Unknown
TFU.72720 Sweden 50 (9.1)9.9–11.2(13.2) 10.5 (6.1)6.6–7.6(7.7) 7 (1.3)1.4–1.6(1.7) 1.5 Alnus
G. oregonense H7009574 USA 34 (11)12–13(14) 12.4 (7)7.3–8(8.4) 7.8 (1.4)1.5–1. 8(1.8) 1.6 Tsuga
H7050171 USA 50 (10.4)11.1–13.1(13.6) 12.3 (6.6)6.7–7.9(8.3) 7.3 (1.4)1.6–1.8(1.9) 1.7 Tsuga
G. tsugae H7050170 Canada 50 (9.3)9.9–11.1(11.7) 10.5 (6.4)6.5–7.2(7.4) 6.9 (1.4)1.44–1.6(1.7) 1.5 Tsuga
H7008236 USA 34 (9.5)10–12(12) 10.9 (6)6.5–7.4(7.5) 7 (1.3)1.4–1.7(1.8) 1.6 Tsuga
Note. Spores measured (n), mean length (L), mean width (W), average length divided by average width (Q), and length and width ratio of individual spores (Q’). 
The whole range variation of L, W, and Q’ is reported in parentheses, and the 80% confidence range is given without parentheses.
MYCOLOGIA 1059
present phylogenetic and morphological analyses could 
not confirm the identity of the laccate Ganoderma spe-
cies of Finland, other than the fact that it is a single 
taxon that resides in the G. lucidum complex. Regarding 
the nomenclature related to G. lucidum sensu stricto, 
the most convenient approach would be to select an 
epitype following the recommendations of the 
International Code of Nomenclature for algae, fungi, 
and plants. We thus recommend that interfertility tests 
be performed between G. tsugae originating from North 
America and China, the selected G. lucidum sensu 
stricto epitype, and G. lucidum originating from 
Finland, Siberia, and the UK. Morphological and multi-
locus phylogenetic studies of less scrutinized 
Ganoderma species (i.e., G. carnosum, G. mongolicum 
Pilát, and G. valesiacum) as well as those originating 
from undersampled boreal regions, such as Siberia, are 
also needed to resolve the identity of the laccate 
Ganoderma species occurring in Finland.
ACKNOWLEDGMENTS
We thank the University of Turku Herbarium (Turku, 
Finland) and the Finnish Museum of National History 
Botanical Herbarium for allowing us to examine their fungal 
collections. We are also grateful to Prof. Tuan Duong and 
Dr. Suvi Sutela for their help with our PCRs and to the citizens 
and field workers who collected the specimens used in this 
study, and to Luke’s and FABI’s laboratories and laboratory 
staff.
DISCLOSURE STATEMENT
No potential conflict of interest was reported by the author(s).
FUNDING
This work was supported by Niemi Foundation under grant 
20180020 and by the Wihuri Foundation under grant 
200042/3b. This work was also supported by Luke  Leads 
under MushValue project 41007-00098000 and InnoFungi 
project 41007-00158400; by Business Finland Research to 
Business funding under Natural Antivirals project 42431/ 
31/2020; by Academy of Finland under Antivirals from 
Forest Biomasses: Structure, Function, and Applicability 
project 342250; by the Regional Council of North Karelia 
EAFRD program under MushroomHarvester project 
EURA2014/8860/09020101/2019/PKARJALA; and by the 
Regional Council of Lapland EAFRD program under 
FeedFUNK project EURA2014/10739/09020101/2020/LL. 
The strains used in this work were collected in MushValue 
project; Sievi project 68873 funded by EU Rural 
Development Programme for Mainland Finland 2007– 
2013; and LUMO-INKA project 2845/31/2015 funded by 
Tekes.
ORCID
Marta Cortina-Escribano http://orcid.org/0000-0003- 
1862-4391
Martin Petrus Albertus Coetzee http://orcid.org/0000- 
0001-7848-4111
Henri Vanhanen http://orcid.org/0000-0002-5165-0236
Riikka Linnakoski http://orcid.org/0000-0002-3294-8088
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