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Comptes rendus de l’Acad´emie bulgare des Sciences
Tome 76, No 5, 2023
BIOLOGY
Mycology
MACROFUNGI IN STANDS OF THE ENDEMIC PINE
PINUS PEUCE AS INFERRED FROM MORPHOLOGICAL
AND MOLECULAR DATA
Boris Assyov#, Monica Slavova∗
Received on February 13, 2023
Presented by D. Ivanov, Corresponding Member of BAS, on March 28, 2023
Abstract
Pinus peuce (Balkan, Macedonian or Molika pine) is an endemic tree for
the Balkans and a Tertiary relict, one of the two European five-needled pines of
the subgenus Strobus (white pines). This work summarizes the data on macro-
fungal diversity in natural forests and artificial plantations of Macedonian pine
in Bulgaria, based on three year survey of epigeous and hypogeous fungi in
selected sampling plots, and combining morphology-based and molecularly in-
ferred identification. The study registered 190 species in total. Higher diversity
of macrofungi was recorded on sampling plots in natural stands compared to
artificial plantings. Nine species are reported for the first time for the Bulgarian
mycobiota. The paper provides the first public set of sequences of macrofungi
of P. peuce stands, comprising 62 accessions.
Key words: Balkan Peninsula, ascomycetes, basidiomycetes, biogeogra-
phy, DNA-barcoding, nrITS, macromycetes
Introduction. Pinus peuce Griseb. is а member of Pinus subgenus Strobus
(D. Don) Lemmon, encompassing the so-called white pines, also colloquially
known as five-needled pines [1]. It is a Tertiary relict, endemic for the Balkan
Peninsula, where it has relatively limited range, building stands in some of the
#Corresponding author
This study was supported by the Bulgarian Science Fund (Grant no. KP-06-N36/13/
17.12.2019).
DOI:10.7546/CRABS.2023.05.06
707
highest mountains in Albania, Bosnia and Herzegovina, Bulgaria, Greece, Mon-
tenegro, North Macedonia, and Serbia [1, 2 ]. In Bulgaria its natural range is re-
stricted to the highest mountains, namely Central Stara Planina, Pirin, Rila and
Rodopi, where it builds up monodominant or mixed stands with other conifers [3].
The habitats dominated by this pine are deemed to be ‘endangered’ at national
level in this country [3].
As one of the only two European five-needled pines, next to the Alpine-
Carpathian P. cembra L., P. peuce attracted the attention of several mycologists,
although this interest developed relatively recently and the number of focused
studies on its fungi is still limited. Data on macrofungi of Macedonian pine stands
were published from North Macedonia, where the locus classicus (Pelister Mt.) of
the species is situated [1, 4–6 ]. Further on, some data on larger fungi from stands
with this pine have been published from Bulgaria [7–9] and a list of characteristic
fungi of P. peuce habitats was presented [3]. Large parts of the information in
those publications were gathered during relatively short field trips and part of
them refer to mixed stands with Macedonian pine. Very little is known about
the macromycetes of artificial P. peuce plantations, with merely one study from
Bulgaria and one from Poland [8, 10]. Besides, no focused studies with application
of DNA methods are known so far and a limited number of sequences of fungi of
P. peuce was obtained in the context of other publications [11].
Materials and methods. The present study was held on selected perma-
nent sampling plots of 10 000 m2in stands of P. peuce in Pirin, Rila, Stara Planina
and Vitosha Mts. Coordinates and other data on the sampling plots are presented
in Table 1. In the list below, the numbers after species names refer to those plots.
Sampling was done between June and November during three consecutive years,
starting in 2020 and held in accordance with previously published protocol [12].
Epigeous and hypogeous fungi were sampled by random transects during each
visit, covering the entire plot area; the search for hypogeous fungi was done with
the aid of trained dogs. The material was documented in the field and is preserved
in air-dried state in the Mycological Collection of the Institute of Biodiversity and
Ecosystem Research (SOMF). The identification of the collected macrofungi was
achieved with the use of broad range of contemporary monographs, keys and at-
lases. The names of fungi in the text below follow the Index Fungorum database
(http://www.indexfungorum.org/).
DNA extraction, amplification and sequencing of the nrITS region were out-
sourced to ALVALAB (Spain) and were performed in accordance with the protocol
communicated by the laboratory and presented in previously published work [13].
In addition, small number of LSU and tef -1αsequences was also produced. The
obtained sequences were subjected individually to BLAST search of the Gen-
Bank database [14] and the resulting alignments were critically examined in order
to achieve or confirm taxonomic assignment. In a few cases when unambiguous
identification could not be reached by BLAST alignments, phylogenetic analysis
708 B. Assyov, M. Slavova
T a b l e 1
Sampling plots in stands of Pinus peuce
No Mountain and
plot name Coordinates Age∗
(yrs) Elev. (m) Tree composition
and stand origin
1Pirin Mts
Vihren
41◦45.7150N
23◦25.0170E140 1969 PP(100%)
Natural stand
2Rila Mts
Malyovitsa
42◦12.5380N
23◦23.4020E100 1760 PP(50%), PS(30%), PA(20%)
Natural stand
3Rila Mts
Treshtenik
42◦04.9300N
23◦37.0820E90 1915 PP(80%), PS(10%), PA(10%)
Natural stand
4Vitosha Mts
Goli Vrah
42◦35.5180N
23◦17.5570E80 1814 PP(90%), PA(10%)
Artificial planting
5Balkan Mts
Petrohan
43◦06.8570N
23◦08.0070E90 1472 PP(70%), PS(20%), PA(10%)
Artificial planting
∗In case of mixed stands the age refers to the age in years of the dominant species, Pinus
peuce. Abbreviations of tree species, used in the last column: “PP” – Pinus peuce, “PS” –
Pinus sylvestris, “PA” – Picea abies.
of suitable datasets (not shown) was performed on the portal Phylogeny.fr [15]. In
the list below asterisked numbers indicate the plots from where the sequenced ma-
terials were collected; these are followed by the accession numbers of the sequences
in GenBank (beginning with the prefix ‘OQ’).
Results.
List of the species. Ascomycetes: Choiromyces meandriformis Vittad. –
2*, OQ398590; Elaphomyces granulatus Fr. – 3; Gyromitra esculenta Fr. – 2;
Helvella corium (O. Weberb.) Massee – 4; H. elastica Bull. – 4; Basidiomycetes:
Agaricus augustus Fr. – 2; A. sylvaticus Schaeff. – 2; A. sylvicola (Vittad.)
L´ev. – 1,2; Aleurodiscus amorphus (Pers.) J. Schrot. – 2; Amanita cf. brun-
neofuliginea – 4; A. gemmata (Fr.) Bertill. – 1,2,3,4; A. muscaria (L.) Pers. –
3,4,5; A. rubescens Pers. – 2,4; A. spissa (Fr.) P. Kumm. – 2,3,4; A. submem-
branacea (Bon) Gr¨oger – 1,2,3,4,5; Amylocystis lapponica (Romell) Singer – 2;
Armillaria ostoyae (Romagn.) Herink – 1,4*, OQ398576; Auriscalpium vulgare
Gray – 2; Baeospora myosura (Fr.) Singer – 1,4; Boletus edulis Bull. – 1,2,3,5; B.
pinophilus Pil´at & Dermek – 2,3,5; Bovista plumbea Pers. – 1,3; Calocera cornea
(Batsch) Fr. – 2,3,4,5; C. viscosa (Pers.) Fr. – 2,3,4,5; Cantharellus cibarius
Fr. – 1*,2, OQ388266; Chalciporus piperatus (Bull.) Bataille – 2; Chlorophyllum
rhacodes (Vittad.) Vellinga – 2; Chroogomphus helveticus (Singer) M. M. Moser –
1*,2,3,4,5, OQ398587; C. mediterraneus (Finschow) Vila et al. – 2*, OQ456634;
Clavulina rugosa (Bull.) J. Schr¨ot. – 4*, OQ398559; Climacocystis borealis (Fr.)
Kotl. & Pouzar – 2*, OQ398588; Clitocybe fragrans (With.) P. Kumm. – 1,2,4;
C. nebularis (Batsch) P. Kumm. – 2,3; C. odora (Bull.) P. Kumm. – 2,4; C.
C. R. Acad. Bulg. Sci., 76, No 5, 2023 709
vibecina (Fr.) Qu´el. – 1,2,3,5; Clitopilus prunulus (Scop.) P. Kumm. – 2; Colly-
bia cirrhata (Schumach.) Quel. – 3,5; Collybiopsis ramealis (Bull.) Millsp. – 2;
Coprinopsis nivea (Pers.) Redhead et al. – 2; Cortinarius acutus (Pers.) Fr. –
1; C. croceus (Schaeff.) Gray – 2; C. jacobi-langei Bidaud – 2*, OQ398581; C.
malicorius Fr. – 2; C. mucosus (Bull.) J. Kickx f. – 2,3; C. sommerfeltii Høil.
– 2*, OQ398585; C. variecolor (Pers.) Fr. – 3; C. varius (Schaeff.) Fr. – 3; C.
venetus (Fr.) Fr. – 2*, OQ398572; Cyathus striatus (Huds.) Willd. – 2; Cys-
toderma amianthinum (Scop.) Fayod – 1,3,4; Cystodermella granulosa (Batsch)
Harmaja – 1; Dacrymyces stillatus Nees. – 2,3,4,5; Deconica coprophila (Bull.) P.
Karst. – 2; D. merdaria (Fr.) Noordel. – 2; Ditiola peziziformis (L´ev.) D.A. Reid
– 2; Entoloma cetratum (Fr.) M.M. Moser – 4; E. hirtipes (Schumach.) M.M.
Moser – 2,3; E. vernum S. Lundell – 1,2,3; Fomitopsis pinicola (Sw.) P. Karst. –
2,3,4,5; Galerina marginata (Batsch) K¨uhner – 2,5; Geastrum fimbriatum Fr. – 2;
Gloeophyllum abietinum (Bull.) P. Karst. – 2; G. sepiarium (Wulfen) P. Karst.
– 1,2,4,5; Gomphidius glutinosus (Schaeff.) Fr. – 2,3; Guepiniopsis cf. estonica
– 2; Gymnopilus penetrans (Fr.) Murrill – 2,5; Gymnopus androsaceus (L.) Della
Magg. & Trassin. – 2,3,5; G. dryophilus (Bull.) Murrill – 1,2,3,4,5; Heteroba-
sidion annosum (Fr.) Bref. – 3; Hydnum repandum L. – 2,3,4*,5, OQ398557; H.
rufescens Pers. – 2*, OQ398575; Hygrocybe conica s. l. – 2*, OQ398606; Hy-
grophoropsis aurantiaca (Wulfen) Maire – 2; Hygrophorus agathosmus (Fr.) Fr.
– 2*, OQ398568; H. cf. persicolor – 2*, OQ398580; H. discoideus (Pers.) Fr.
– 1; Hypholoma capnoides (Fr.) P. Kumm. – 2,3; H. fasciculare (Huds.) P.
Kumm. – 2,5; Imleria badia (Fr.) Vizzini – 1,2*,3,5, OQ398556; Infundibuli-
cybe gibba (Pers.) Harmaja – 1*,2,3,4, OQ398592; Inocybe asterospora Qu´el. –
1; I. geophilla (Pers.) P. Kumm. – 2*,3,4,5, OQ398594; I. lilacina (Peck) Kauff-
man – 2*,3, OQ398578; Inosperma maculatum (Boud.) Matheny & Esteve-Rav.
– 1; Ischnoderma benzoinum (Wahlenb.) P. Karst. – 1; Laccaria amethystina
Cooke – 2,5; L. bicolor (Maire) P.D. Orton – 1,2*, OQ398565, OQ398583; L.
laccata (Scop.) Cooke – 3; L. ’macrocystidia’ – 2*, OQ398566; Lactarius auran-
tiacus (Pers.) Gray – 1*,2*,3*,4, OQ398573, OQ398574, OQ398586; L. borzianus
(Cavara) Verbeken & Nuytinck – 1,2*, OQ398554; L. deliciosus (L.) Gray – 2,3,4;
L. deterrimus Gr¨oger – 2,4*, OQ398570; L. picinus Fr. – 2,3,4*, OQ398555; L.
piperatus (L.) Pers. – 2; L. rufus (Scop.) Fr. – 1,3*,5, OQ398561; L. scrobiculatus
(Scop.) Fr. – 2; L. zonarioides K¨uhner & Romagn. – 1*,4, OQ398569; Laeti-
cutis cristata (Schaeff.) Audet – 2; Lepiota clypeolaria (Bull.) P. Kumm. – 2;
Lepista nuda (Bull.) Cooke – 1,2,5; Leucocortinarius bulbiger (Alb. & Schwein.)
Singer – 1,2; Leucogyrophana mollusca (Fr.) Pouzar – 5; Leucopaxillus gentia-
neus (Qu´el.) Kotl. – 2; Lycoperdon excipuliforme (Scop.) Pers. – 1,2,3,5; L.
molle Pers. – 2,4,5; L. perlatum Pers. – 2,3,4,5; L. pyriforme Schaeff. – 2,5;
Macrolepiota fuliginosa (Barla) Bon – 1,2; Mallocybe terrigena (Fr.) Matheny et
al. – 1; Melanoleuca cf. subpulverulenta – 5*, OQ398593; Melanoleuca melaleuca
(Pers.) Murrill – 4; Mycena aurantiomarginata (Fr.) Qu´el. – 2; M. capillar-
710 B. Assyov, M. Slavova
ipes Peck – 1,2,3; M. crocata (Schrad.) P. Kumm. – 2; M. epipterygia (Scop.)
Gray – 1,2,3,5; M. galopus (Pers.) P. Kumm. – 4; M. pura s. l. – 2,3,4,5;
M. rosea (Schumach.) Gramberg – 5; M. rosella (Fr.) P. Kumm. – 1,2,3,4,5;
Mycetinis alliaceus (Jacq.) A.W. Wilson & Desjardin – 2; Myxomphalia maura
(Fr.) Hora – 2,4; Neoboletus erythropus (Pers.) C. Hahn – 2,3; Onnia tomentosa
(Fr.) P. Karst. – 1; Panaeolus semiovatus (Sowerby) S. Lundell & Nannf. – 2;
Paragymnopus perforans (Hoffm.) J.S. Oliveira – 1,2,3,4,5; Paralepista flaccida
(Sowerby) Vizzini – 2,5; Paxillus involutus (Batsch) Fr. – 2,4; Phaeoclavulina abi-
etina (Pers.) Giachini – 2; Phaeocollybia festiva (Fr.) R. Heim – 2,4; Phlegmacium
balteatum (Fr.) A. Blytt – 3*, OQ398558; Pholiota flammans (Fr.) P. Kumm.
– 4; P. pinicola Jacobsson – 4; P. spumosa (Fr.) Singer – 2*, OQ398597; Poly-
porus melanopus (Pers.) Fr. – 2; Porphyrellus porphyrosporus (Fr. & H¨ok) E.-J.
Gilbert – 1*,2,4, OQ398562; Protostropharia semiglobata (Batsch) Redhead et al.
– 2; Pseudohydnum gelatinosum (Scop.) P. Karst. – 1,2,3; Ramaria aurea (Scha-
eff.) Qu´el. – 1,3; Rhizopogon salebrosus s. l. – 1*, 5*, OQ398602, OQ398603,
OQ398604, OQ398605; Rhodocollybia butyracea (Bull.) Lennox – 2,3,4,5; Rhodo-
phana nitellina (Fr.) Larouz et al. – 2; Rickenella fibula (Bull.) Raithelh. –
2; Russula adusta (Pers.) Fr. – 3,5; Russula amara Kuиera – 1,2*, OQ398584;
R. anthracina Romagn. – 1*, OQ398551; R. chloroides (Krombh.) Bres. – 1*,
OQ398563; R. delica s. lat. – 2,5; R. foetens (Pers.) Fr. – 3,5; R. integra (L.)
Fr. – 1*,2,3,4,5, OQ398553; R. mustelina Fr. – 1,2*,4*,5, OQ398589, OQ398564;
R. ochroleuca Fr. – 5*, OQ398567; R. puellaris s. l. – 2*, OQ398601; R. rosea
Pers. – 2; R. roseipes Bres. – 1*,4, OQ398579; R. torulosa Bres. – 2; R. turci
Bres. – 1*, OQ398591; R. xerampelina (Schaeff.) Fr. – 2,3; Sarcodon imbricatus
(L.) P. Karst. – 2; Skeletocutis amorpha (Fr.) Kotl. & Pouzar – 2,4; Sparassis
crispa (Wulfen) Fr. – 2*, OQ398560; Stereum hirsutum (Willd.) Gray – 1,2,3,4;
Strobilurus sp. – 1, non-plot: Rila Mts, Panichishte area*, OQ398598, OQ402730;
S. stephanocystis (Hora) Singer – 1,2,3,4*, OQ398550; S. tenacellus (Pers.) Singer
– 2,3; Stropharia aeruginosa (Curtis) Qu´el. – 1,2,3,4; Suillus americanus (Peck)
Snell – 1*,3,4, OQ398582, OQ402729, OQ411036; S. bovinus (L.) Roussel – 5; S.
granulatus (L.) Roussel – 3,5; S. luteus (L.) Roussel – 3,5; S. variegatus (Sw.)
Richon & Roze – 3,5; Tapinella atrotomentosa (Batsch) ˇ
Sutara – 2; T. panuoides
(Fr.) E.-J. Gilbert – 2; Thaxterogaster turmalis (Fr.) Niskanen & Liimat. – 3*,
OQ398577; Thelephora caryophyllea (Schaeff.) Pers. – 3,5; T. palmata (Scop.)
Fr. – 2; T. terrestris Ehrh. – 5; Tremella mesenterica Retz. – 2; Tricholoma
equestre (L.) P. Kumm. – 1; T. imbricatum (Fr.) P. Kumm. – 2; T. orirubens
Qu´el. – 4; T. portentosum (Fr.) Qu´el. – 1,2,3; T. saponaceum (Fr.) P. Kumm. –
2*,3, OQ398571; T. terreum (Schaeff.) P. Kumm. – 2*,3, OQ398552; T. vaccinum
(Schaeff.) P. Kumm. – 1*,2,3, OQ398595; T. virgatum (Fr.) P. Kumm. – 2; Tri-
cholomopsis rutilans s. l. – 2,4*, OQ398596; Xerocomellus chrysenteron (Bull.)
ˇ
Sutara – 1*,2,3, OQ398600; Xerocomus ferrugineus (Schaeff.) Alessio – 3,5*,
OQ398599; Xeromphalina campanella (Batsch) K¨uhner & Maire – 1,2.
C. R. Acad. Bulg. Sci., 76, No 5, 2023 711
Discussion. A total of 190 species were registered during the study. Natural
stands appeared to be richer, compared to artificial plantations, with 59 (plot 1),
69 (plot 3) and 135 species (plot 2) in the former, against 49 (plot 4) and 54
(plot 5) in the latter. Among the plots in natural forests, the one in the area of
Malyovitsa (plot 2) is richest, with twice or more species. This plot has different
stand composition and is the only one where trees other than P. peuce have 50%
participation, while in the remaining sampling stands this figure does not exceed
30% (Table 1). We hypothesize that the observed higher diversity of macrofungi
is at least partly driven by the diversity and abundance of different host trees.
When pooled data from natural and artificial plots are compared, the former host
170 species versus twice less in the latter (85 species). Shared between the two
types of plots are 67 species, of which almost half – 33 species are saprotrophs with
different substrate specialisation, and the rest are ectomycorrhizal fungi. Should
plot 2 is excluded and the pooled data for the remaining natural (plots 1 & 3)
and artificial stands (plots 4 & 5) are examined, the plots in natural stands still
harbour higher diversity (100 species) than those in artificial ones (85 species).
Of the recorded species, 103 (54.2%) were only found in plots in natural forests,
while merely 18 (9.5%) were recorded only on plots in artificial plantings.
A handful of species were found on all sampling plots and some of them
were even common (Fig. 1a-c). Among these is Chroogomphus helveticus (Fig.
1b), species already outlined as a characterizing taxon for stands of P. peuce
[4, 16]. Other such common species appeared to be the saprotrophs Gymnopus
dryophilus, Mycena rosella, Paragymnopus perforans, and the mycorrhizal Russula
integra, which are, however, ubiquitous in coniferous forests in all studied areas.
One species that drew attention during our study was Amanita submembranacea
(Fig. 1a). It was detected on all plots and during all visits, even in unfavourable
seasons, while in rainy periods it created characteristic aspect, dominating all
other mycorrhizal fungi by the number of fruitbodies. This outlines it as one
of the characteristic species of the stands of P. peuce, suggesting also a high
degree of ecological plasticity and possible pioneer behaviour. Suillus americanus
(Fig. 1c), known to bind mycorrhizally only to five-needled pines in Europe,
where it was previously known as S. sibiricus (Singer) Singer [4, 9], appeared in
a few but not all sampling plots. It was, however, found in nearly all studied
areas outside of the plots boundaries, often fruiting under seedlings of P. peuce,
suggesting that tree age may be important for its mycorrhizae. In fact, suilloid
fungi are known to be important for the establishment of pines in native and non-
native ranges [17]. Tricholoma terreum, generally uncommon in the studied plots
where mostly mature trees were present, exhibited somewhat similar behaviour,
occurring prolifically in areas with seedlings of Macedonian pine present. It may
be examined in future whether it might play somewhat similar role as suilloid
fungi.
712 B. Assyov, M. Slavova
Fig. 1. Basidiomata of selected characteristic species of macrofungi for Pinus peuce stands: a –
Amanita submembranacea; b – Chroogomphus helveticus; c – Suillus americanus; d – Rhizopogon
salebrosus s. l.; e – Strobilurus sp.
Several incompletely resolved taxa were recovered in this study, which are
worthy of further observations. Rhizopogon salebrosus (Fig. 1d) occurred in two
of the studied sites, undoubtedly forming mycorrhizae with P. peuce as the only
tree in the surveyed areas. Recently, a new species – R. confusus Koukol &
Valda, was proposed to accommodate European collections of this group [18]. In
C. R. Acad. Bulg. Sci., 76, No 5, 2023 713
general, our ITS sequences seem compatible to and cluster together with those
of R. confusus. However, as the taxonomy of this group relies on the use of
anonymous loci to improve the resolution of the phylogenetic inferences, we prefer
to treat those collections as R. salebrosus s. l. for the time being. A specimen of
the genus Laccaria Berk. & Broome, which could not be plausibly matched to any
taxon in the existing European keys, yielded a sequence, identical to sequences,
dubbed as L. ‘macrocystidia’ (nom. nud.). Apparently further work is necessary
as to show whether some of the existing binomials may be attached to it or it will
represent an undescribed entity. Further on, a morphologically distinct, autumnal
Strobilurus was detected (Fig. 1e), which sequence matches an unnamed species of
the S. wyomingensis complex, uncovered by a recent study [11]. Those sequences
also come from collections on cones of P. peuce, from North Macedonian sites,
and this taxon is pending further judgement. Lastly, a sequenced specimen of the
Tricholomopsis rutilans complex turns out to represent one of the known, but not
yet formally described cryptic species in this group [19].
Nine new species for the Bulgarian mycota [20] were revealed by the present
study, namely Cortinarius jacobi-langei, C. malicorius, C. sommerfeltii, Mallocybe
terrigena, Pholiota pinicola, Rhizopogon salebrosus s. l., Phlegmacium balteatum,
Russula anthracina, and Thaxterogaster turmalis, the majority of them supported
by nrITS sequences. Over half of these are cortinarioid fungi (Cortinarius, Phleg-
macium, Thaxterogaster ), which fact is not surprising, as this group of larger fungi
is among the most diverse in Europe and is apparently understudied in Bulgaria.
Acknowledgements. Dr. Pablo Alvarado (ALVALAB, Spain) is thanked
for obtaining the sequences released in this work. Thanks are due to the staff of
the Pirin National Park Directorate for the provided assistance.
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Institute of Biodiversity
and Ecosystem Research
Bulgarian Academy of Sciences
2 Yurii Gagarin St
1113 Sofia, Bulgaria
e-mail:contact@boletales.com
∗Faculty of Biology
Plovdiv University “Paisii Hilendarski”
24 Tsar Assen St
4000 Plovdiv, Bulgaria
e-mail:helvellasolitaria@gmail.com
C. R. Acad. Bulg. Sci., 76, No 5, 2023 715