The Community of Fine Root Fungi of Silver Fir ( Abies alba Mill.) Saplings

Abstract

This study aimed to assess the biodiversity of fungi colonizing the fine roots (diameter up to 2 mm) of 3-year-old silver fir saplings from areas of Międzylesie Forest District in Poland. It was hypothesized that quantitatively and qualitatively, mycorrhizal fungi would be the dominant fungi in root communities of silver fir. DNA extraction was performed using Plant Genomic DNA purification. The internal transcribed spacer1 (ITS1) rDNA region was amplified using specific primers, and the amplicons were purified and sequenced using sequencing by synthesis (SBS) Illumina technology. The obtained sequences were compared with reference sequences in the UNITE database (https://unite.ut.ee/) using the basic local alignment search tool (BLAST) algorithm to facilitate species identification. A total of 307,511 OTUs was obtained from each sample. There were 246,477 OTUs (80.15%) of fungi known from cultures. The genera Tuber spp. (7.51%) and Acephala spp. (3.23%) accounted for the largest share of the fungal communities on the fine roots of fir trees. Hence our results indicate the dominance of mycorrhizal fungi in these communities and reflect the excellent quality of the saplings that were assessed. Pathogenic fungi constituted a much smaller share of the fungal communities.

Full text

The Community of Fine Root Fungi of Silver Fir (Abies alba Mill.) Saplings
hps://www.seefor.eu SEEFOR 14(2): early view 1
ISSN 1847-6481
eISSN 1849-0891
© 2023 by the Croaan Forest Research Instute. This is an Open Access paper distributed under the terms
of the Creave Commons Aribuon License (hp://creavecommons.org/licenses/by/4.0).
Marlena Baranowska1*, Władysław Barzdajn1, Robert Korzeniewicz1, Wojciech Kowalkowski1,
Adrian Łukowski1, Jolanta Behnke-Borowczyk2, Mirzeta Memišević Hodžić3
(1) Poznań University of Life Sciences, Faculty of Forestry and Wood Technology, Department
of Silviculture, Wojska Polskiego 71A, PL-60-625 Poznań, Poland; (2) Poznań University of Life
Sciences, Faculty of Forestry and Wood Technology, Department of Entomology and Forest
Pathology, Wojska Polskiego 71C, PL-60-625 Poznań, Poland; (3) University of Sarajevo, Faculty
of Forestry, Department of Culvaon of Forests and Urban Greens, Zagrebačka 20, BA-71000
Sarajevo, Bosnia and Herzegovina
* Correspondence: e-mail: marlenab@up.poznan.pl
This study aimed to assess the biodiversity of fungi colonizing the ne roots (diameter up to 2 mm) of 3-year-old silver
r saplings from areas of Międzylesie Forest District in Poland. It was hypothesized that quantavely and qualitavely,
mycorrhizal fungi would be the dominant fungi in root communies of silver r. DNA extracon was performed using
Plant Genomic DNA puricaon. The internal transcribed spacer1 (ITS1) rDNA region was amplied using specic primers,
and the amplicons were puried and sequenced using sequencing by synthesis (SBS) Illumina technology. The obtained
sequences were compared with reference sequences in the UNITE database (hps://unite.ut.ee/) using the basic local
alignment search tool (BLAST) algorithm to facilitate species idencaon. A total of 307,511 OTUs was obtained from each
sample. There were 246,477 OTUs (80.15%) of fungi known from cultures. The genera Tuber spp. (7.51%) and Acephala
spp. (3.23%) accounted for the largest share of the fungal communies on the ne roots of r trees. Hence our results
indicate the dominance of mycorrhizal fungi in these communies and reect the excellent quality of the saplings that were
assessed. Pathogenic fungi constuted a much smaller share of the fungal communies.
Keywords: Lower Silesia; Międzylesie Forest District; nursery; restoraon; Sudeten Mountains
ABSTRACT
INTRODUCTION
The silver r tree (Abies alba Mill.) used to be one of the
most important forest-forming species in the mountainous
and upland regions of forest stands in Poland, in which r is
the dominant species, occupying 2% of the forest area and
having a 2.7% share in the volume in the forests managed
by the State Forests (Bis and Dobrowolska 2012). However,
since 1998, the share of r trees in the forests of the Sudeten
Mountains in Poland has fallen below 0.4%, and the species
had already lost the possibility of connued existence and
self-regeneraon in the compeve environment of the
forests of the region. For this reason, the State Forests
(Poland) undertook a program to restore r trees to forests
of the Sudeten Mountains (Barzdajn 2000, 2012, Barzdajn
and Kowalkowski 2012), but with me it was noced that
fungi-related issues should not be overlooked. The studies
of fungal communies inhabing r roots have focused
mainly on mycorrhizae (within and outside their natural
occurrence) (Kowalski 1982, Comandini et al. 2001, Laganà
et al. 2002, Rudawska et al. 2016) and pathogenic fungi
that infect their root systems (Puddu et al. 2003, Oliva and
Colinas 2007, Chomicz-Zegar et al. 2016). However, no
research has been undertaken in the mountainous regions
of Poland. In addion to pathogenic and mycorrhizal fungi,
these communies are known to include antagonists
of other microorganisms and neutral organisms. Root-
associated fungal communies are essenal components
in ecosystem processes, impacng plant growth and vigour
by inuencing the quality, direcon, and ow of nutrients
and water between plants and fungi (Unuk et al. 2019).
Hence, the aim of the current study was to assess the
diversity of fungi occurring in the rhizosphere of the root
systems of small cungs and r saplings. We tested the
hypothesis that mycorrhizal fungi should dominate these
communies
ORIGINAL SCIENTIFIC PAPER DOI: hps://doi.org/10.15177/seefor.23-13
Citaon: Baranowska M, Barzdajn W,
Korzeniewicz R, Kowalkowski W, Łukowski
A, Behnke-Borowczyk J, Memišević Hodžić
M, 2023. The Community of Fine Root
Fungi of Silver Fir (Abies alba Mill.) Saplings.
South-east Eur for 14(2): early view. hps://
doi.org/10.15177/seefor.23-13.
Received: 22 Feb 2022; Revised: 9 Apr
2023, 16 May 2023; Accepted: 22 May
2023; Published online: 2 Sep 2023
The Community of Fine Root Fungi of Silver Fir (Abies alba
Mill.) Saplings

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Baranowska M, Barzdajn W, Korzeniewicz R, Kowalkowski W, Łukowski A, Behnke-Borowczyk J, Memišević Hodžić M
2 SEEFOR 14(2): early view
MATERIALS AND METHODS
Fine roots of 6 saplings of silver r from the forest nursery
of Międzylesie Forest District (16°66'23"E, 50°14'86"N,
south-west part of Poland) were randomly selected for
the study. The samples were collected in June 2017. The
research material comprised ne roots (roots up to three
rows to idenfy all types of mycorrhizae) (McCormack et al.
2015). The ne roots were washed on sieves under running
tap water and dried on sterile paper. Aer drying, the roots
were ground in a mortar frozen to -70
о
C. The DNA extracted
from dried roots was separated under the microscope.
We composited one sample from six trees. Environmental
DNA was extracted with Plant Genomic DNA Puricaon
Kit (Thermo Fisher Scienc). The internal transcribed
spacer1 (ITS1), 5.8S rDNA region was used to idenfy the
fungal species, and the analysis was carried out with primers
ITS1FI2 5′-GAACCWGCGGARGGATCA-3′ (Schmidt et al. 2013)
and 5.8S 5`-CGCTGCGTT CTTCATCG-3` (Vilgalys and Gonzalez
1990). Each amplicaon reacon was carried out in a nal
volume of 25.0 μl containing 2 ml of DNA, 0.2 ml of each
primer, 10.1 ml of deionized water and 12.5 mL of 2X PCR
MIX (A&A Biotechnology, Gdynia, Poland). The amplicaon
reacon was carried out in a thermocycler. This included:
inial denaturaon (94°C, 5 min), 35 cycles of denaturaon
(94°C, 30 s), annealing (56°C, 30 s), elongaon (72°C, 30 s)
and nal elongaon (72°C, 7 min). The product was then
checked on a 1% agarose gel stained with Midori Green
Advance DNA (Genecs, Dueren, Germany). The obtained
product was puried and sequenced using sequencing by
synthesis (SBS) technology from Illumina (Genomed S.A.
Warsaw, Poland). The results were subjected to bioinformac
and stascal analysis according to Behnke-Borowczyk et al.
(2019). The resulng sequences were compared with the
reference sequences deposited in the UNITE community
database (Nilsson et al. 2018, UNITE community 2020) using
the basic local alignment search tool (BLAST) algorithm.
The abundance of fungi was dened as the number of
OTUs in a sample. A total amount of OTUs was obtained
from six samples collected from each of the 3-year-old small
roots of A. alba. The frequency of an individual taxon was
dened as the percentage (%) of OTUs in the total number
of OTUs. Diversity was dened as the number of species in
a sample. The trophic role of the detected fungal species in
the community was determined based on literature data
and listed in Appendix (Table A1).
RESULTS
A total of 307 511 OTUs were obtained. There were
246 477 OTUs (80.15%) of fungi known from cultures, 1 876
OTUs (0.61%) of non-cultured fungi, 47 572 (15.47%) OTUs
of non-fungal organisms (mainly plants, including silver r),
and 1 814 OTUs (0.59%) of organisms with no reference
sequence in UNITE database. The total number of taxa
obtained was 1612.
The community comprised taxa belonging to
Ascomycota (46.27%), Basidiomycota (33.64%), Zygomycota
(2.52%), Rozellomycota (0.65%), and we also obtained some
sequences that are not represented in the UNITE database
(0.59%) (Table A1).
The Thelephoraceae (8.22%), Tuber spp. (7.51%) and
Acephala spp. (3.23%) had the largest share of ne roots of
common r trees, therefore accounng for the largest share
of the fungal community (Table A1).
Figure 1. Percentage share (%) of the most abundant types of fungi in the community (share > 0.5%).
Tuber
Acephala
Gyoeryella
Tylospora
Hymenogaster
Morerella
Amphinema
Hydnotrya
Archaeorhizomyces
Ophiostoma
Cadophora
Athelopsis
Trichosporon
Russula
Lactarius
Rhizoscyphus
Ilyonectria
Inocybe
Penicillium
Oidiodendron
Leucosporidium
0 1 2 3 4 5 6 7 8
Share (%)

The Community of Fine Root Fungi of Silver Fir (Abies alba Mill.) Saplings
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Mycorrhizal fungi dominated the fungal community of
the ne root (57,62%). The most common taxa idened
among mycorrhizal fungi were: Hydnotrya spp., Tuber
spp., Amphinema spp., Hymenogaster spp., Tylospora
spp., endophytes including Acephala spp., Cadophora
spp., pathogenic fungi including Ophiostoma spp. and
saprotrophs Athelopsis lembospora, Archaeorhizomyces
borealis and Morerella spp. (Table 1, Table A1).
DISCUSSION
The results of our analysis support our hypothesis that
mycorrhizal fungi dominate the fungal communies on the
ne root systems of silver r saplings. Taxa, which belong
to the fungi, forming mycorrhizal communies of silver r,
include Tuber spp. and Acephala spp., and these accounted
for the largest share of the fungal communies on the ne
Table 1. Genus of ectomycorrhizal (ECM) fungi, whose frequency in the collecon in ne roots of silver r exceeded 0.03%. and which
denotes the presence of a taxa in roots or soil in previous scienc reports.
Genus Root Soil Reference
Amanita + + Ważny (2014), Rudawska et al. (2016), Unuk et al. (2019), Behnke-Borowczyk et al. (2020)
Amphinema +Ważny (2014), Unuk et al. (2019)
Cenococcum + + Unuk et al. (2019), Ważny (2014), Behnke-Borowczyk et al. (2020)
Cornarius +Ważny (2014), DEEMY (2014-2020), Unuk et al. (2019), Behnke-Borowczyk et al. (2020)
Elaphomyces + + Unuk et al. (2019), Ważny (2014), Behnke-Borowczyk et al. (2020)
Hydnotrya +- Ważny (2014)
Inocybe +Ważny (2014), Behnke-Borowczyk et al. (2020)
Lactarius + + Ważny (2014), Rudawska et al. (2016), Unuk et al. (2019), Behnke-Borowczyk et al. (2020)
Meliniomyces + + Unuk et al. (2019), Behnke-Borowczyk et al. (2020)
Piloderma +Ważny (2014), Rudawska et al. (2016)
Russula + + DEEMY (2014-2020), Ważny (2014), Rudawska et al. (2016), Unuk et al. (2019), Behnke-
Borowczyk et al. (2020)
Tricholoma +DEEMY (2014-2020), Rudawska et al. (2016), Behnke-Borowczyk et al. (2020)
Tuber + + Ważny (2014), Rudawska et al. (2016), Behnke-Borowczyk et al. (2020)
Tylospora + + Ważny (2014), Rudawska et al. (2016), Unuk et al. (2019), Behnke-Borowczyk et al. (2020)
Xerocomellus +Rudawska et al. (2016)

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4 SEEFOR 14(2): early view
root systems of r saplings in our study. These results are
slightly dierent from the studies of soil fungal communies
in nurseries producing silver r seedlings conducted by
Behnke-Borowiczyk et al. (2020), where saprotrophic fungi
dominated. However, Behnke-Borowczyk et al. (2020)
also idened mycorrhizal fungi. We also idened some
saprotrophs which accounted for just a small share of the
fungal community (not exceeding 1.5%) on the roots of
silver r tree saplings in our study.
Smutek et al. (2010) obtained the following mycorrhizal
families and species of fungi on old silver r trees (90-105
years old): Cornarius serpes, Sebacina sp., Amphinema
byssoides, Russula puellaris, Clavulina cristata, Cornarius
sp., Tomentella sublilacina, Russula fellea, Laccaria
amesna and Tylospora asteropcumum. Our results concur,
conrming the occurrence of fungi belonging to all the taxa
recorded by Smutek et al. (2010). Similarly, we have found
the presence of some of the taxa detected by Wojewoda
(2003) in the fungal community on r trees, including
Amanita, Amphinema, Boletus, Cenococcum, Clavulina,
Cornarius, Elaphomyces, Hydnotrya, Hydnum, Inocybe,
Lactarius, Leoomycetes, Piloderma, Pseudotomentella,
Russula, Sebacina, Tomentella, Tuber, and Tylospora.
However, unlike Ważny (2014) and Schirkonyer et al.
(2013), the fungi genera Byssocorcium, Laccaria, Paxillus,
Thelephora, Tomentellopsis and Xerocomus were not
idened in our study. Ważny (2014) found that the
mycorrhizal fungi which dominated on the youngest r trees
examined were: Clavulina cristata (25.2%), Tomentella sp.
(10.5%), Tuber puberulum (8.9%), and Clavulina sp. (5.1%).
While the genus Tuber accounted for a similar share (7.51%)
of the community of fungi in our work, the presence of
the other genera/species did not exceed 1%. Apart from
Geopora sp. and Imleria sp., we idened similar taxa of
fungi to those recorded by Rudawska et al. (2016), who
studied mycorrhizal fungi of common r trees outside their
natural range, but from mature forest stands. To date, 13
species of ectomycorrhizal fungi (ECM) associated with
the genus Abies have been included in DEEMY: Abierhiza
fascicularis, A. tomentosa, Cornarius odorifer, Lactarius
caespitosus, L. deliciosus, L. intermedius, L. salmonicolor,
L. subsericatus, Polyporoletus sublividus, Russula brevipes,
R. ochroleuca, R silvicola, and Tricholoma bufonium, of
which only R. ochroleuca was found in the sampled fungal
community. The root community diers signicantly from
the fungal community associated with r analyzed by
Behnke-Borowczyk et all (2020), who isolated 13 taxa of
ectomycorrhizal fungi (ECM) associated with the genus
Abies.
The presence of Cenococcum geophilum and Thele-
phora stuposa was insignicant in the study, which is in
opposion to the research done by Rudawska et al. (2016).
The list of mycorrhizal species detected in silver r roots
include Amphinema byssoides, Clavulina cristata, Lactarius
auranacus, L. salmonicolor, Piloderma fallax, Tuber
puberulum, T. asterophora, T. stuposa Boletus pruinatus,
Cenococcum geophilum, and Laccaria amethysna, which
have been conrmed in other studies (Eberhardt et al.
2000). In addion, C. geophilum, A. byssoides, T. stuposa,
Amanita, Boletus, Cenococcum, Cornarius, Inocybe,
Laccaria, Lactarius, Russula, Sebacina, Tomentella, and
Tuber also form mycorrhizal compounds with other r
species (Matsuda and Hijii 1999, 2004, Ishida et al. 2007,
Kranabeer et al. 2009).
Acephala applanata, which we detected in ne roots
of A. alba, was previously almost exclusively isolated from
Picea abies (L.) H. Karst (Grünig et al. 2006). While second
species from Acephala genus A. macroscleroorum formed
ectomycorrhizas on Pinus sylvestris (Münzenberger et al.
2009) was detected in a study of the community of fungi on
silver r ne roots. These results are consistent with those
obtained by Behnke-Borowczyk et al. (2020) in soil research
related to nurseries producing silver r seedlings. However,
the share of Acephala spp. in the previously studied soil
was lower than in the roots. The greater share of these
fungi in the root community is not surprising, because both
species inhabit the roots: A. applanata is included in the
DSE (dark septate endopyhtes) (Stroheker et al. 2021), while
A. macroscleroorum is classied as ectomycorrhizal fungi
(Münzenberger et al. al. 2009).
For young silver r trees, similarly to Unuk et all (2019),
we detected endophyc root-associated fungal genera
Oidiodendron, Phialocephala, and Rhizoscyphus. Some
consider fungi of these genera to be mycorrhizal, and
therefore they are treated as fungi in Appendix A. However,
their role in the silver r root community has not yet been
clearly dened.
In fungal community of silver r ne roots idened
cosmopolitan fungi from the genera Trichoderma and
Penicillium as well, which are antagonists of the pathogens
Armillaria and Heterobasidion (Behnke-Borowczyk and
Kwaśna 2010; Granna-Ievina et al. 2013; Baranowska et al.
2023).
Our study also idened pathogenic fungi in the
ne roots of silver r tree saplings, namely Ophiostoma
nigrocarpum, which, together with O. novo-ulmi and
Ophiostoma sp., accounted for 1.11% of the fungal
community. Fungi of the Ophiostoma genus are pathogens
whose vectors are bark beetles on older trees (for example,
Pissodes piceae), which cause white discolouraon of r’s
wood, thus reducing its economic value (Six and Bentz 2003,
Kirisits 2004).
CONCLUSIONS
This study led to recognizing the spectrum of mycorrhizal,
saprotrophic, and pathogenic fungi characterisc for ne
roots of the 3-year-old Abies alba. Relavely low proporon
of pathogens in these fungal communies also supports the
conclusion that they were healthy trees. To fully conrm
the roles and funcons of the idened taxa, further
idencaon of communies should be carried out. Parallel
to the analysis of the communies of fungi inhabing silver r
ne roots, it is necessary to study the content of nutrients and
soil pH and determine their impact on these communies. In
addion, research should be connued at a later stage of tree
development, i.e. in young forest stands, to determine the
formaon of these communies and the spectrum of specic
species of fungi associated with young silver r trees.

The Community of Fine Root Fungi of Silver Fir (Abies alba Mill.) Saplings
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Appendix A
Table A1. Taxa occurring in the communies of fungi in the roots of r trees, whose frequency in the collecon exceeded 0.03%.
The legend of colours in the "Frequency" column:
Dominant taxa are marked in bold. Symbols of trophic
groups are: M - mycorrhizal fungi, S - saprotrophic fungi,
A - antagonisc fungi, P - pathogenic fungi, L - lichens, E -
entomopathogenic fungi, U - unknown. The "Frequency, %"
column contains the collected taxa data in the community,
red represenng the smallest share, and green represenng
the largest share of the taxon in the community (the scale is
aached above).
Tax on Order Frequency
(%)
Similarity
(%)
Trophic
group Reference
Acephala applanata Grünig & T.N.
Sieber + A. macroscleroorum
Münzenb. & Bubner + Acephala
sp.
Heloales 3.231 97-100 M Münzenberger et al. (2009)
Apiognomonia errabunda
(Roberge ex Desm.) Höhn. Diaporthales 0.063 99 PMańka (2005)
Archaeorhizomyces borealis
Menkis, T.Y. James & Rosling +
Archaeorhizomyces sp.
Incertae sedis 1.356 98-100
Ascomycota 1.821 100
Beauveria pseudobassiana S.A.
Rehner & Humber + Beauveria sp. Hypocreales 0.039 98-100 EÁlvarez-Baz et al. (2015)
Cadophora nlandica (C.J.K. Wang
& H.E. Wilcox) T.C. Harr. & McNew
+ C. orchidicola (Sigler & Currah)
M.J. Day & Currah + Cadophora sp.
Incertae sedis 1.049 97-100 PYak et al. (2019)
Capnodiales 0.060 99
Cenococcum geophilum Fr. +
Cenococcum sp. Mylinidiales 0.355 99-100 M Spatafora et al. (2012)
Chaetothyriales 0.271 100
Chalara hyalocuspica Koukol + Ch.
pseudoanis Koukol + Chalara sp. Heloales 0.482 97-100 S/P Koukol (2011), Coker et al.
(2019)
Cheirosporium triseriale L. Cai &
K.D. Hyde Pleosporales 0.181 99 SAbdel-Aziz (2016)
Cladophialophora minussima
M.L. Davey & Currah + C.
chaetospira (Grove) Crous &
Arzanlou + Cladophialophora sp.
Chaetothyriales 0.211 98-100 P/S Badali et. al. (2008)
Clonostachys rosea (Link)
Schroers, Samuels, Seifert & W.
Gams + Clonostachys sp.
Hypocreales 0.038 98-100 A Cota et al. (2009)
Coccomyces australis P.R. Johnst. Rhysmatales 0.088 99 PJohnson (1986)
Coniochaeta sp. Coniochaetales 0.042 100 P Damm et al. (2010)
Coniochaetaceae Coniochaetales 0.035 100
Dermateaceae Heloales 0.532 99
Didymella dactylidis (Aveskamp,
Gruyter & Verkley) Qian Chen &
L. Cai + D. protuberans (Lév.) Qian
Chen & L. Cai
Pleosporales 0.211 98-100 P/ S Chen et al. (2015)
Dothideomycetes 0.289 100
Elaphomyces muricatus Fr. + E.
granulatus Fr. Euroales 0.176 98-99 M Paz et al. (2017)
Erysiphe euonymicola U. Braun+
E. hypophylla (Nevod.) U. Braun &
Cunningt.
Erysiphales 0.104 99-100 PSepúlveda-Chavera et al.
(2013)

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6 SEEFOR 14(2): early view
Tax on Order Frequency
(%)
Similarity
(%)
Trophic
group Reference
Exophiala sp. Chaetothyriales 0.033 100 S Feng et al. (2013)
Fimetariella sp. Sordariales 0.102 100 U
Fusarium sp. + F. oxysporum
Schltdl. Hypocreales 0.087 99-100 S/P Karim et al. (2016)
Geomyces asperulatus Sigler &
J.W. Carmich. + G. auratus Traaen
+ Geomyces sp.
Heloales 0.387 98-100
Gyoeryella entomobryoides
(Boerema & Arx) Marvanová +
Gyoeryella sp.
incertae sedis 2.285 99-100 PJankowiak et al. (2016)
Halokirschsteiniothelia marima
(Linder) Boonmee & K.D. Hyde 0.046 100 S Wilson (1951)
Halosphaeriaceae Microascales 0.067 99
Heloaceae Heloales 0.685 99
Heloales 1.878 100
Hydnotrya cerebriformis Harkn. +
H. michaelis (E. Fisch.) Trappe + H.
tulasnei (Berk.) Berk. & Broome
Pezizales 1.405 99-100 M Hobbie et al. (2001)
Hymenoscyphus sp. Heloales 0.073 100 S Gumińska and Wojewoda
(1985)
Hypocreales 2.001 100
Ilyonectria morspanacis (A.A.
Hildebr.) A. Cabral & Crous + I.
robusta (A.A. Hildebr.) A. Cabral
& Crous
Hypocreales 0.744 99-100 PFarh et al. (2017)
Infundichalara minuta Koukol +
Infundichalara sp. Heloales 0.081 99-100 SKoukol (2012)
Leoa lubrica (Scop.) Pers. Heloales 0.081 100 S Kuo (2012)
Leoomycetes 1.099 100
Leptosphaeria sp. Pleosporales 0.079 100 P Brachaczek et. al. (2016)
Lopadostoma polynesium (Berk. &
M.A. Curs) Rappaz Xylariales 0.033 99 PMehrabi and Hemma
(2015)
Lophodermium conigenum
(Brunaud) Hilitzer + L. pinastri
(Schrad.) Chevall. + L. sediosum
Minter, Staley & Millar
Rhysmatales 0.386 99-100 PBurdekin and Phillips
(1992)
Maasoglossum sp. Geoglossales 0.182 100 Hustad and Miller (2015)
Mariannaea elegans (Corda)
Samson Hypocreales 0.032 99 U Wang and Zabel (1990)
Meliniomyces bicolor Hambl. &
Sigler + M. variabilis Hambl. &
Sigler + M. vraolstadiae Hambl.
& Sigler
incertae sedis 0.285 99-100 M Marno et al. (2018)
Metapochonia bulbillosa (W. Gams
& Malla) Kepler, S.A. Rehner &
Humber
0.040 100 E Adachi et al. (2015)
Mycosphaerellaceae Capnodiales 0.073 99
Nectriaceae Hypocreales 0.055 100
Neobulgaria pura (Pers.) Petr. +
Neobulgaria sp. Heloales 0.113 99-100 SGumińska and Wojewoda
(1985)
Neonectria sp. Hypocreales 0.035 100 P Kryczyński and Weber
(2010)
Oidiodendron maius G.L. Barron
+ O. rhodogenum Robak +
Oidiodendron sp.
Heloales 0.696 99-100 M/S Rice and Currah (2006a)
Ophiosphaerella sp. Pleosporales 0.063 100 P Dernoeden (2000)
Table A1. (counnue) Taxa occurring in the communies of fungi in the roots of r trees, whose frequency in the collecon exceeded
0.03%. The legend of colours in the "Frequency" column:

The Community of Fine Root Fungi of Silver Fir (Abies alba Mill.) Saplings
hps://www.seefor.eu SEEFOR 14(2): early view 7
Ophiostoma nigrocarpum (R.W.
Davidson) de Hoog + Ophiostoma
novo-ulmi Brasier + Ophiostoma
sp.
Ophiostomatales 1.108 98-100 P/S Marcinkowska (2012)
Penicillium citreonigrum Dierckx
+ P. penicillioides (Delacr.) Vuill.
+ P. subrubescens Houbraken,
Mansouri, Samson & Frisvad +
Penicillium sp.
Euroales 0.727 98-100 A Pi et al. (2000)
Petriella sordida (Zukal) G.L.
Barron & J.C. Gilman Microascales 0.033 99 A Lee and Gloer (1995)
Pezizaceae Pezizales 0.210 100
Phaeosphaeriaceae Pleosporales 0.044 100 S
Phialocephala fornii CJK. Wang
& H.E. Wilcox + Phialocephala sp. Heloales 0.414 99-100 M/U Jumpponen et al. (1998)
Phoma boeremae Gruyter +
Phoma sp. Pleosporales 0.070 98-99 Chen et al. (2015)
Pleosporaceae Pleosporales 0.033 100
Pleosporales 0.119 100
Pleotrichocladium opacum (Corda)
Hern.0Restr., R.F. Castañeda &
Gené
Pleosporales 0.248 99 SHernández-Restrepo et al.
(2017)
Podospora appendiculata (Auersw.
ex Niessl) Niessl + Podospora sp. Sordariales 0.023 99-100 SDoveri (2008)
Proliferodiscus sp. Heloales 0.090 100 Han et al. (2014)
Pseudogymnoascus verrucosus
A.V. Rice & Currah Incertae sedis 0.368 99 SRice and Currah (2006b)
Pyronemataceae Pezizales 1.135 100
Rhizoscyphus sp. Heloales 0.786 100 MHambleton and Sigler
(2005)
Rhodoveronaea varioseptata
Arzanlou, W. Gams & Crous incertae sedis 0.053 99 M Tedersoo et al. (2010)
Rhysmataceae Rhysmatales 0.045 100
Sporormiaceae Pleosporales 0.032 100
Stagonospora pseudovitensis
Quaedvl., Verkley & Crous Pleosporales 0.034 99 PQuaedvlieg et al. (2013)
Strumella sp. Pezizales 0.068 100 P/S Zeur and Kullman (2011)
Sympodiella acicola W.B. Kendr. incertae sedis 0.065 99 SShen et al. (2020)
Tetracladium segerum (Grove)
Ingold + Tetracladium sp. Heloales 0.097 99-100 SAnderson and Marvanová
(2020)
Trichoderma sp. Hypocreales 0.120 100 A Benítez et al. (2004)
Tuber anniae W. Colgan & Trappe
+ T. cistophilum P. Alvarado, G.
Moreno, Manjón, Gelpi & Jaime
Muñoz + Tuber sp.
Pezizales 7.508 99-100 M (2004-2020)
Umbilicaria americana Poelt &
T.H. Nash Umbilicariales 0.032 100 L Poelt and Nash (1993)
Venturia hystrioides (Dugan, R.G.
Roberts & Hanlin) Crous & U.
Braun + Venturia sp.
Venturiales 0.146 99-100 PGonzález-Domínguez et
al. (2017)
Venturiaceae Venturiales 0.076 100
Wilcoxina sp. Pezizales 0.049 99 M DEEMY (2004-2020)
Xenopolyscytalum pinea Crous Heloales 0.368 100 P Koukol (2019)
Frequency of Ascomycota isolates 46.27
Agaricales 2.261 100
Table A1. (counnue) Taxa occurring in the communies of fungi in the roots of r trees, whose frequency in the collecon exceeded
0.03%. The legend of colours in the "Frequency" column:
Tax on Order Frequency
(%)
Similarity
(%)
Trophic
group Reference

hps://www.seefor.eu
Baranowska M, Barzdajn W, Korzeniewicz R, Kowalkowski W, Łukowski A, Behnke-Borowczyk J, Memišević Hodžić M
8 SEEFOR 14(2): early view
Table A1. (counnue) Taxa occurring in the communies of fungi in the roots of r trees, whose frequency in the collecon exceeded
0.03%. The legend of colours in the "Frequency" column:
Tax on Order Frequency
(%)
Similarity
(%)
Trophic
group Reference
Agaricomycetes 0.064 100
Amanita rubescens Pers. +
Amanita sp. Agaricales 0.159 99-100 M DEEMY (2004-2020)
Amphinema byssoides (Pers.) J.
Erikss. + Amphinema sp. Atheliales 1.448 99-100 M DEEMY (2004-2020)
Amphistereum leveilleanum (Berk.
& M.A. Curs) Spirin & Malysheva Auriculariales 0.131 100 U
Atheliaceae Atheliales 0.078 100
Atheliales 0.181 100
Athelopsis lembospora (Bourdot)
Oberw. Amylocorciales 1.030 99 U
Auriculariales 0.444 99
Basidiodendron caesiocinereum
(Höhn. & Litsch.) Luck-Allen Auriculariales 0.097 100 S Koranta and Saarenoksa
(2005)
Basidiomycota 0.565 100
Cantharellales 0.096 100
Ceratobasidiaceae Cantharellales 0.351 100
Clavariaceae Agaricales 0.155 99
Clavulina coralloides (L.) J. Schröt. Cantharellales 0.245 99 SWojewoda (2003)
Clavulinopsis sp. Agaricales 0.077 100 S Kautmanová et al. (2012)
Cornarius croceus (Schae.) Gray
+ C. semisanguineus (Fr.) Gillet Agaricales 0.032 99-100 M DEEMY (2004-2020)
Cryptococcus sp. Tremellales 0.056 100 P/S Springer et al. (2017)
Curvibasidium cygneicollum J.P.
Samp. incertae sedis 0.064 100 P/ S Kaitera et al. (2019)
Deconica phyllogena (Sacc.)
Noordel. Agaricales 0.061 99 SNoordeloos (2011)
Galerina nana (Petri) Kühner +
Galerina sp. Agaricales 0.349 99-100 SGulden et al. (2005)
Hydnaceae Cantharellales 0.308 100
Hydnodontaceae Trechisporales 0.054 100
Hygrophoraceae Agaricales 2.149 100
Hymenochaetales 0.085 100
Hymenogaster boozeri Zeller &
C.W. Dodge + H. huthii Selow,
Bratek & Hensel + H. olivaceus
Viad.
Agaricales 1.625 99-100 M Selow et al. (2011)
Inocybe rufoalba Sacc. + Inocybe
sp. Agaricales 0.733 99-100 M DEEMY (2004-2020)
Itersonilia perplexans Derx + I.
pannonica (Niwata, Tornai-Leh., T.
Deák & Nakase) Xin Zhan Liu, F.Y.
Bai, M. Groenew. & Boekhout
Cystolobasidiales 0.063 99-100 PMcGovern et al. (2006)
Kockovaella sacchari M. Takash.
& Nakase Tremellales 0.032 99 U
Lactarius sp. + L. necator (Bull.)
Pers. + L. tabidus Fr. Russulales 0.797 98-100 M DEEMY (2004-2020)
Leucosporidium drummii Yurkov,
A.M. Schäfer & Begerow +
Leucosporidium sp.
Leucosporidiales 0.508 99-100 U Sampaio et al. (2003)
Malassezia restricta E. Guého, J.
Guillot & Midgley + Malassezia sp. Malasseziales 0.068 99 PSaunders et al. (2012)
Microbotryomycetes 0.050 100
Mycena sanguinolenta (Alb. &
Schwein.) P. Kumm. + Mycena sp. Agaricales 0.244 99-100 SPerry (2002)

The Community of Fine Root Fungi of Silver Fir (Abies alba Mill.) Saplings
hps://www.seefor.eu SEEFOR 14(2): early view 9
Phellinus castanopsidis BK. Cui,
Y.C. Dai & Decock Hymenochaetales 0.022 100 P Cui and Decock (2012)
Piloderma sp. Atheliales 0.216 100 M DEEMY (2004-2020)
Piskurozyma sp. Filobasidiales 0.016 100 U
Pterula sp. Agaricales 0.117 100 S Leal-Dutra et al. (2020)
Ramariopsis sp. Agaricales 0.021 99 SMatouš et al. (2017)
Rhodotorula sp. Sporidiobolales 0.034 100 AAkhtyamova and Saarova
(2013)
Rigidoporus sanguinolentus (Alb.
& Schwein.) Donk Polyporales 0.353 100 S Wojewoda (2003)
Russula puellaris Fr. + R. fragilis
Fr. + R. nigricans Fr. + R. veternosa
Fr. + R. badia Quél. + R. ionochlora
Romagn. + Russula sp.
Russulales 0.859 99-100 M DEEMY (2004-2020)
Sebacinaceae Sebacinales 0.174 100
Sebacinales 0.447 100
Solicoccozyma terrea (Di Menna)
Yurkov + Solicoccozyma sp. Filobasidiales 0.224 99-100 A Yurkov et al. (2019)
Sporidiobolales 0.049 100
Stereum sanguinolentum (Alb. &
Schwein.) Fr. Russulales 0.046 100 S Łakomy and Kwaśna (2008)
Thelephoraceae Thelephorales 8.220 100 Richard et al. (2011)
Trechispora sp. Trechisporales 0.069 99 S/M Gumińska (1985), Vanegas-
León et al. (2019)
Trechisporales 0.084 100
Tremellales 0.044 100
Tremellomycetes 0.073 100
Tricholoma fulvum (DC.) Bigeard
& H. Guill. + Tricholoma sp. Agaricales 0.064 99-100 M DEEMY 2004-(2020)
Trichosporon sp. Tremellales 0.949 100 A Bosqueiro et al. (2020)
Tylospora asterophora (Bonord.)
Donk + Tylospora sp. Atheliales 1.689 98-99 M DEEMY (2004-2020)
Vishniacozyma victoriae (M.J.
Montes, Belloch, Galiana, M.D.
García, C. Andrés, S. Ferrer, Torr.
-Rodr. & J. Guinea) Xin Zhan Liu,
F.Y. Bai, M. Groenew. & Boekhout
+ Vishniacozyma sp.
Tremellales 0.067 99-100 A Gramisci et al. (2018)
Xerocomellus pruinatus (Fr. & Hök)
Šutara + X. chrysenteron (Bull.)
Šutara n
Boletales 0.032 99-100 M Šutara (2008)
Frequency of Basidiomycota
isolates 33.64
Absidia sp. Mucorales 0.034 100 S Alastruey-Izquierdo et al.
(2010)
Basidiobolus ranarum Eidam Entomophthorales 0.064 99 SYang (1962)
Endogone sp. Endogonales 0.072 100 S/M Warcup (1990)
Jimgerdemannia lacua (Berk. &
Broome) Trappe, Desirò, M.E. Sm.,
Bonito & Bidartondo
Endogonales 0.030 99-100 S/M Desirò et al. (2018)
Morerella horcola Linnem. +
M. angusta (Linnem.) Linnem. +
Morerella sp.
1.484 99-100 S/A Granna-Ievina et al.
(2014)
Morerellaceae 0.130 100 S
Morerellales Morerellales 0.079 100 S
Umbelopsis dimorpha Mahoney &
W. Gams + U. ramanniana (Möller)
W. Gams + Umbelopsis sp.
Umbelopsidales 0.241 99-100 S/A
Crowther et al. (2012),
Granna-Ievina et al.
(2014)
Table A1. (counnue) Taxa occurring in the communies of fungi in the roots of r trees, whose frequency in the collecon exceeded
0.03%. The legend of colours in the "Frequency" column:
Tax on Order Frequency
(%)
Similarity
(%)
Trophic
group Reference

hps://www.seefor.eu
Baranowska M, Barzdajn W, Korzeniewicz R, Kowalkowski W, Łukowski A, Behnke-Borowczyk J, Memišević Hodžić M
10 SEEFOR 14(2): early view
Table A1. (counnue) Taxa occurring in the communies of fungi in the roots of r trees, whose frequency in the collecon exceeded
0.03%. The legend of colours in the "Frequency" column:
Tax on Order Frequency
(%)
Similarity
(%)
Trophic
group Reference
Frequency of zygomycota isolates 2.52
Frequency of other kingdoms 15.47
Not represented in UNITE
database 0.59
Frequency of uncultured fungi 0.61
Author Contribuons
WB, WK, JBB, MB conceived and designed the research, MB
carried out the eld measurements, JB performed laboratory
analysis, JB and WB processed the data and performed the
stascal analysis, WK, WB secured the research funding, AŁ, RK,
JBB supervised the research and helped to dra the manuscript,
MB wrote the manuscript. The main part of these results was
presented at the internaonal IUFRO Conference - Abies&Pinus
2022, “Fir and pine management in a changing environment: Risks
and opportunies” held 19-22 September 2022 in Sarajevo, Bosnia
and Herzegovina.
Funding
This study was nanced by the State Forests Naonal Forest
Holding, General Directorate of the State Forests in Warsaw
(Poland), as Program to Restore Resources in the Sudetes, part IV,
(Project number OR.271.3.12.2017).
Conicts of Interest
The authors declare no conict of interest.
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