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Citation: Farkas, E.; Varga, N.; Veres,
K.; Matus, G.; Sinigla, M.; L˝okös, L.
Distribution Types of Lichens in
Hungary That Indicate Changing
Environmental Conditions. J. Fungi
2022,8, 600. https://doi.org/
10.3390/jof8060600
Academic Editors: Pradeep
K. Divakar and Lourdes Morillas
Received: 26 April 2022
Accepted: 1 June 2022
Published: 3 June 2022
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4.0/).
Fungi
Journal of
Article
Distribution Types of Lichens in Hungary That Indicate
Changing Environmental Conditions
Edit Farkas 1, * , Nóra Varga 1, Katalin Veres 1, Gábor Matus 2, Mónika Sinigla 3and LászlóL˝okös 4
1Institute of Ecology and Botany, Centre for Ecological Research, H-2163 Vácrátót, Hungary;
varga.nora@ecolres.hu (N.V.); veres.katalin@ecolres.hu (K.V.)
2Department of Botany, Faculty of Science and Technology, University of Debrecen,
H-4010 Debrecen, Hungary; matus.gabor@science.unideb.hu
3Bakony Museum of the Hungarian Natural History Museum, H-8420 Zirc, Hungary;
monikasinigla@gmail.com
4Department of Botany, Hungarian Natural History Museum, H-1431 Budapest, Hungary;
lokos.laszlo@nhmus.hu
*Correspondence: farkas.edit@ecolres.hu
Simple Summary:
As the occurrences of lichens are strongly correlated to background environmental
conditions (e.g., air pollution, global warming), the analysis of their distribution has a great value for
bioindication. Distribution data are originating from earlier herbarium collections, recent field and
literature studies. The distribution analyses in lichen species with different ecological requirements
allowed comparisons and showed clear trends. Five distribution types were introduced—presented
by characteristic examples—according to lichen distribution maps prepared in different periods of
time (representing changing environmental conditions): (1) species of decreasing occurrences by time
(acidic pollution sensitive species), (2) species with no or few former records but with increasing
occurrences in recent decades (sub-Mediterranean species), (3) species with increasing and then (from
c. 2000) decreasing occurrences (acidofrequent species), (4) species with widely increasing occurrences
in recent decades (nitrofrequent species), and (5) species with rapidly increasing occurrences (rapidly
spreading species of uncertain reasons). The discussed trends are known for some species at a global
scale or European level, other examples are characteristic for Central Europe or Hungary. By studying
the distribution maps of lichen bioindicators, tendencies of climate change and type of pollution can
be determined and further changes can be predicted.
Abstract:
Distribution data originating from earlier herbarium collections and recent biodiversity
records form the basis of distribution analyses in lichen species with different ecological require-
ments, where the records allowed comparisons or showed clear trends. As the occurrences of lichens
are strongly correlated to background environmental conditions (e.g., air pollution, global warm-
ing), confirmed by Wirth’s ecological indicator values, the analysis of distribution types has a great
value for bioindication and the establishment of current and future climatic and pollution situations.
Five distribution types were introduced—presented by characteristic examples (13)—according to
lichen distribution maps prepared in different periods of time (representing changing environmen-
tal conditions): (1) species of decreasing occurrences by time (e.g., Lobaria pulmonaria,Menegazzia
terebrata, suboceanic, acidic pollution sensitive species), (2) species with no or few former records
but with increasing occurrences in recent decades (e.g., Flavoparmelia soredians,Hyperphyscia adgluti-
nata,Solenopsora candicans, sub-Mediterranean species), (3) species with increasing and then (from c.
2000) decreasing occurrences (e.g., Scoliciosporum chlorococcum,Straminella conizaeoides, acidofrequent
species), (4) species with widely increasing occurrences in recent decades (e.g., Physcia aipolioides,
Piccolia ochrophora,Xanthoria parietina, nitrofrequent species), and (5) species with rapidly increasing
occurrences (e.g., Absconditella lignicola,Coenogonium pineti,Evernia divaricata, rapidly spreading
species). The proposed distribution types of lichen species may be applied to wider regions (the
European or the global level).
J. Fungi 2022,8, 600. https://doi.org/10.3390/jof8060600 https://www.mdpi.com/journal/jof
J. Fungi 2022,8, 600 2 of 17
Keywords:
acidofrequent; air pollution bioindication; biodiversity; climate; environmental changes;
land use; nitrofrequent; rapidly spreading; substrate; time scale
1. Introduction
Lichens are generally known as extremotolerant, cosmopolitan organisms found from
the Equator to the poles and from sea level to the highest montane vegetation zones [
1
,
2
].
Even the simplest lichen consists of two different organisms [
3
]. Others (those of three to
five partner symbioses) live together with additional fungal (parasitic or parasymbiont) or
photosynthetic partners (green algae or cyanobacteria). Recently basidiomycete yeasts [4]
and specific coexisting bacteria (most frequently Alphaproteobacteria) have also been
discovered [5] to form a close connection with lichen association [6].
The indicator nature of lichens was recognized almost two centuries ago in relation
to the environmental effects of Britain’s strong industrialization. It was observed [
7
] that
the cover and the composition of lichen assemblages are related to air quality. Mapping of
lichens revealed that the distribution of lichens according to their ecological requirements
appears as variously polluted zones (central lichen desert, struggle zone, normal/natural
zone) in cities and around factories and industrial plants [
8
–
10
]. The occurrence of par-
ticular species could be related to the concentration of SO
2
, being the most characteristic
pollutant in the 1970s [
11
]. The study of lichens and air pollution soon became one of
the leading directions of lichenological research; later investigations proceeded from the
community and the species levels toward the cell and the molecular levels [
12
–
15
]. With
the decline of SO
2
pollution, it soon became obvious that lichens recolonize former lichen
deserts [16,17],
but the new assemblages consisted of species tolerating changed types of
pollutants (e.g., NH3) [18–22].
Earlier the focus was on the composition and the distribution of species, later changes
were due to time and environmental quality (corresponding to the quantitative and the
qualitative nature of the pollution) [20–25].
Investigations concerning the indicator nature of lichens were extended to the mon-
itoring of land use intensity, and they were studied in a wider context [
26
–
29
], with due
attention also given to the effect of the substrate [
30
–
34
]. For monitoring studies, the
time passed has a key importance [
35
]. Recognizing that global environmental changes
mean an increasing pressure to the living organisms of the earth [
36
–
39
], it was shown
that lichenized associations represent a sensitive model ecosystem in the changing envi-
ronment [
1
,
26
,
28
]. Comparing distribution maps of various species in different periods
of time and geographic scales is a tool that easily presents and illustrates background
environmental changes [
40
–
42
]. The analysis of Hungarian data may result in similar
examples [9,10,26,28].
The large amount of data accumulated during the past centuries makes a distribution
analysis at a wide scale in time and in space possible. Altogether 55,000 lichen specimens
are deposited in Hungarian herbaria collected between 1762 and 2022. Although these
collections and additional literature records are not homogeneous in space and in time, the
distribution maps of each species—prepared according to these records—are obviously
applicable for recognizing characteristic distribution types in correlation with tendencies in
the changing environment because of the indicator nature of lichens [7–25].
The main aims of this study are to prepare distribution maps of lichens in Hungary
in various time periods and to introduce distribution types by comparing these maps and
the direction of environmental changes. If possible, indicator groups of species are also
going to be established on the basis of characteristic distribution types. We hypothesize that
environmental changes are reflected in lichen distributions since lichens are regarded as
sensitive environmental bioindicators. Though lichens as a group have a wide distribution,
lichen species are characterized with specific and contrasting environmental requirements.
J. Fungi 2022,8, 600 3 of 17
2. Materials and Methods
Hungary has a varied relative relief ranging from 0 m (at lowland) to c. 1000 m (at
lower montane regions) with geomorphological origin–mostly marine, fluvial, and aeolian
sediment of various ages and in smaller areas volcanic rocks covering the formations of
older geographical periods. The six main geomorphological districts are the Alföld (the
Great Hungarian Plain), the Kisalföld (the Little Hungarian Plain), the Alpokalja (the Foot
of the Alps), the Transdanubian Hills, the Transdanubian Range, and the North Hungarian
Range (as a part of the Northwestern Carpathians) [
43
,
44
]. The natural Pannonian veg-
etation consists of various deciduous forests (supplying a large number of tree species
for epiphytic lichens) on mountainous regions and montane rocky and lowland sandy
grasslands with special microclimatic conditions allowing lichen colonization on various
soil and rock surfaces [45].
The macroclimate is predominantly continental with oceanic and Mediterranean
effects. The annual sunshine duration is between 1900 and 2100 h increasing from northwest
to southeast with an average monthly duration between 50 and 280 h (maximum in June,
minimum in January [
46
]. The difference in monthly mean temperature is more than
1◦C
if the current (1981–2010) 30-year period is compared to 1960–1990 (which is often
considered as a reference period in modelling future climate) [46]. The mean temperature
is currently (1981–2010) 10.35
◦
C. It has increased by 1.6
◦
C (reaching even 2
◦
C in summer)
in this 30-year period. According to predicting models, the mean temperature will continue
to increase both annually and seasonally by 1–2
◦
C for 2021 to 2050 and by 3–4
◦
C for
2071–2100 and from west-northwest to east-southeast. The highest increase 3.5–4.5
◦
C is
expected in summer (even by 6
◦
C in August) [
46
]. The frequency of days with frost will
be significantly lower than in the past. Annual precipitation is now 580 mm (1981–2010),
ranging from 700–800 mm (in mountains and at the southwestern border) to 500 mm in the
Alföld. It is predicted to decrease slightly (by c. 15–20 mm) for 2021–2050, particularly in
July–August and January–February. However, there is a precipitation increase in autumn.
Furthermore, different predictions exist for the period 2071–2100, showing either a further
decrease or an increase in precipitation.
The atmospheric concentration of sulphur dioxide, creating the major part of acidic
deposition, increased until the middle of the 1980s (reaching 25–30
µ
g/m
3
SO
2
) in Hungary,
after that it has been continuously decreasing (reaching 1–3
µ
g/m
3
SO
2
in 2015) [
47
]. The
household emission of nitrogen-containing pollutants—with a significant effect on lichen
colonization in and around settlements—increased considerably (from 1955 t NH
3
in 1990
to 5672 t NH3in 2017) [48].
A detailed study on lichen-forming fungi of Hungary was published by Klára Verseghy
in 1994 [
49
]; however, this identification key contained distribution data of lichens by
geographic regions only, still it was very useful since it collected data scattered in a large
number of literature sources. In 2009, a revised checklist of species [
50
] recorded from
Hungary was compiled from literature and herbarium records (from the late 1800s to
2022). The checklist was continuously updated, and the species number has now reached
926 due to our revision and mainly our own collections from 1979. Most of the species
(49%) are saxicolous on various rocks, 15% are terricolous, 33% are corticolous, and 3%
are lignicolous. These species and their records (c. 55,000) form the basis of our studies.
The number of references checked is c. 180 from 1869 to 2021. New collections were
identified with the aid of various literature sources mainly [
51
,
52
]. The morphology and the
anatomy were studied by using a NIKON Eclipse/NiU (DIC, epifluorescence) compound
microscope, Nikon SMZ18 stereo microscope as well as Olympus SZX9 and Olympus BX50
(DIC) microscopes. Micrographs were prepared by Olympus E450 camera (with Quick
Photo Camera 2.3 software) and Nikon DS-Fi1c and Fi3 camera (with NIS-Elements BR
ML software), with the indicated microscopes. HPTLC analysis was carried out according
to standard methods for analyzing lichen samples [
53
,
54
], where it was necessary. The
nomenclature mainly follows the IndexFungorum [55].
J. Fungi 2022,8, 600 4 of 17
Voucher specimens are deposited in herbaria BP, BTM, DE, EGR, GODO, JPU, SZE,
SAMU, and VBI (abbreviations mainly follow [
56
]). Most of the specimens studied are in
BP (c. 40,000 specimens), and a c. 15,000 specimens are found in other herbaria.
Distribution maps were constructed by a computer program for geographical infor-
mation systems, QGIS 3.18.2 ‘Zürich’, released in 2020, applying an adaptation of the
Central European grid system [
57
,
58
]. The illustrated symbols (dots) represent units of
c.
5 km ×6 km
. In 1989, a European Lichen Mapping project was initiated [
59
–
61
]. Data
before and after 1975 were distinguished, considering changes in conditions of air pollution
in Europe. Therefore, these periods are shown on maps, although the decline of acidic
pollution started 10–15 years later in Hungary (cf., [46]).
3. Results
Distribution maps of a wide selection of species from the Hungarian lichen flora [
49
,
50
]
were prepared in recent decades. Former versions of some of these maps have been pub-
lished in scattered publications on various taxa, e.g., [
62
,
63
]. Species with the best-known
distribution records were selected to be presented here, where data from different time
periods allowed reliable comparisons or recent data showed clear trends. The following
distribution types are proposed for species on the basis of an analysis of historical and
recent data:
(a)
species with decreasing occurrences;
(b)
species with no or few former records but with increasing occurrences in recent
decades;
(c)
species with increasing, then decreasing occurrences;
(d)
species with widely increasing occurrences in recent decades;
(e)
species with rapidly increasing occurrences.
In order to evaluate all of the 926 species into these distribution types, a few examples
were analyzed in more detail. Nevertheless, a considerable number (c. 40%) were catego-
rized as “distribution types to be established” due to the lack of sufficient data or having
only uncertain records.
Thirteen examples of the distribution types are presented and illustrated below
(
Figures 1–5
; Figures S1–S8 in Supplementary File) and the possible reasons for these
characteristic distributions are discussed later in Section 4.
3.1. Species of Decreasing Occurrences
3.1.1. Lobaria pulmonaria L.
Lobaria pulmonaria is a very conspicuous, large, foliose species (Figure 1) [
63
]. In the
past, it was widely distributed in Hungary in the mountain ranges (mainly the Bükk and
Mátra Mts) occurring dominantly on bark of Fagus sylvatica, and to a lesser extent on
other broad-leaved tree species (Acer,Betula,Quercus,Ulmus spp.) as well as on mossy
andesitic rock surfaces. In older times, it was considered as a medicinal plant (as drug
Herba pulmonariae arboreae,Lichen pulmonarius, or Muscus pulmonarius), and it was used
against lung diseases, but no information is available regarding how extensively it was
collected in Hungary for this purpose [64].
Altogether, 94 data were recorded from herbaria (BP, EGR, GODO, PECS, SZE, SZO)
and literature up to 1967, but only three recent occurrences were found in the Bükk Mts in
2008 and 2016 [64].
3.1.2. Menegazzia terebrata (Hoffm.) A. Massal.
M. terebrata is a characteristic foliose species having large, round perforations on its
upper surface (Figure S1). Until 1960 only nine old records were known from suboceanic
habitats in the mountain range of Hungary (e.g., Bükk, Mátra, Sopron, and Zemplén Mts),
and it was recently confirmed at the former, single locality in the Zemplén Mts. It grows on
bark (Fagus) and on mossy siliceous rocks in Hungary.
J. Fungi 2022,8, 600 5 of 17
J. Fungi 2022, 8, x FOR PEER REVIEW 4 of 18
time periods allowed reliable comparisons or recent data showed clear trends. The fol-
lowing distribution types are proposed for species on the basis of an analysis of historical
and recent data:
(a) species with decreasing occurrences;
(b) species with no or few former records but with increasing occurrences in recent
decades;
(c) species with increasing, then decreasing occurrences;
(d) species with widely increasing occurrences in recent decades;
(e) species with rapidly increasing occurrences.
In order to evaluate all of the 926 species into these distribution types, a few examples
were analyzed in more detail. Nevertheless, a considerable number (c. 40%) were catego-
rized as “distribution types to be established” due to the lack of sufficient data or having
only uncertain records.
Thirteen examples of the distribution types are presented and illustrated below ( Fig-
ure 1 Figure 2 Figure 3 Figure 4 Figure 5; Figures S1–S8 in Supplementary File) and the
possible reasons for these characteristic distributions are discussed later in chapter 4.
3.1. Species of Decreasing Occurrences
3.1.1. Lobaria pulmonaria L.
Lobaria pulmonaria is a very conspicuous, large, foliose species (Figure 1) [63]. In the
past, it was widely distributed in Hungary in the mountain ranges (mainly the Bükk and
Mátra Mts) occurring dominantly on bark of Fagus sylvatica, and to a lesser extent on other
broad-leaved tree species (Acer, Betula, Quercus, Ulmus spp.) as well as on mossy andesitic
rock surfaces. In older times, it was considered as a medicinal plant (as drug Herba pulmo-
nariae arboreae, Lichen pulmonarius, or Muscus pulmonarius), and it was used against lung
diseases, but no information is available regarding how extensively it was collected in
Hungary for this purpose [64].
Figure 1. Lobaria pulmonaria (a) habit (scale 1 cm); (b) its distribution in Hungary (94 records). Dots
represent c. 5 km × 6 km areas. (Photo: L. Lőkös).
Figure 1.
Lobaria pulmonaria (
a
) habit (scale 1 cm); (
b
) its distribution in Hungary (94 records). Dots
represent c. 5 km ×6 km areas. (Photo: L. L˝okös).
3.2. Species with No or Few Former Records but with Increasing Occurrences in Recent Decades
3.2.1. Flavoparmelia soredians (Nyl.) Hale
F. soredians is originally an Atlanto-Mediterranean foliose species (Figure S2), which
seems to be spreading in Hungary [
65
]. Its 19 Hungarian localities were discovered between
2011 and 2022. Contrary to its earlier known habitat requirements (sub-Mediterranean), it
was collected mostly in anthropogenic, urban habitats. F. soredians is corticolous on different
primarily eutrophicated phorophytes in Hungary, e.g., Acer sp., Ailanthus altissima,Prunus
sp., Quercus cerris,Q. petraea,Robinia pseudoacacia,Tilia sp., and decaying wood.
3.2.2. Hyperphyscia adglutinata (Flörke) H. Mayrhofer et Poelt
H. adglutinata is a small foliose species, with narrow (0.3–0.7(–2) mm wide) lobes and
maculiform or capitate soralia (Figure 2). In addition to its four old occurrences from 1925
to 1960, it has been reported from several localities recently from 2000 to 2022, including
urban habitats (e.g., [
66
–
68
]). It is a corticolous species growing on various tree species,
frequently on eutrophicated bark (Acer campestre,A. pseudoplatanus,A. tataricum,Alnus
glutinosa,Betula pendula,Cornus mas,Corylus avellana,Cotinus coggygria,Fraxinus ornus,
Morus alba,Populus canescens,Prunus cerasus,Pyrus pyraster,Quercus pubescens,Robinia
pseudoacacia, and Sambucus nigra).
3.2.3. Solenopsora candicans (Dicks.) J. Steiner
S. candicans has a pale grey, thickly white-pruinose, placodioid, rosette-like thallus
and dark brown to black apothecia (Figure S3). Two old localities were known from the
Buda Mts (1934) and the Balaton Upland (1947). Between 2007 and 2015, it was found at
several localities in the Balaton Upland and the Buda Mts predominantly on calcareous
rocks and also on geyserite near Tihany [69].
J. Fungi 2022,8, 600 6 of 17
3.3. Species with Increasing, then Decreasing Occurrences
3.3.1. Scoliciosporum chlorococcum (Stenh.) Vˇezda
S. chlorococcum is a corticolous crustose species with a green granulose thallus and
shiny blackish or reddish-brown apothecia of 0.2–0.3 mm diam. (Figure 3). Only
15 records
are known between 1912 and 1974, while 292 records were made between 1976 until
2004 [
70
], but only 15 records between 2005 and 2016. In Hungary it was collected predom-
inantly from phorophytes of acidic bark (Fagus sylvatica,Juniperus communis,Larix decidua,
Picea abies,Pinus nigra,P. sylvestris, and Quercus cerris) and a great number of other species
(Acer campestre,A. platanoides,A. pseudoplatanus,Aesculus hippocastanum,Alnus glutinosa,
Armeniaca vulgaris,Betula pendula,B. pubescens,Carpinus betulus,Castanea sativa,Cerasus
avium,Cotinus coggygria,Crataegus monogyna,Euonymus verrucosus,Fraxinus excelsior,F.
ornus,Juglans regia,Malus domestica,Morus alba,Persica vulgaris,Populus alba,P. tremula,
Prunus domestica,Pyrus achras,P. communis,Quercus farnetto,Q. petraea,Q. pubescens,Q.
robur,Robinia pseudoacacia,Salix sp., Sorbus sp., S. torminalis and Tilia spp.) as well as from
decaying wood.
J. Fungi 2022, 8, x FOR PEER REVIEW 6 of 18
Figure 2. Hyperphyscia adglutinata (a) habit (scale 0.5 cm); (b) its distribution in Hungary (56 records).
Dots represent c. 5 km × 6 km areas. (Photo: L. Lőkös).
3.2.3. Solenopsora candicans (Dicks.) J. Steiner
S. candicans has a pale grey, thickly white-pruinose, placodioid, rosette-like thallus
and dark brown to black apothecia (Figure S3). Two old localities were known from the
Buda Mts (1934) and the Balaton Upland (1947). Between 2007 and 2015, it was found at
several localities in the Balaton Upland and the Buda Mts predominantly on calcareous
rocks and also on geyserite near Tihany [69].
3.3. Species with Increasing, then Decreasing Occurrences
3.3.1. Scoliciosporum chlorococcum (Stenh.) Vězda
S. chlorococcum is a corticolous crustose species with a green granulose thallus and
shiny blackish or reddish-brown apothecia of 0.2–0.3 mm diam. (Figure 3). Only 15 rec-
ords are known between 1912 and 1974, while 292 records were made between 1976 until
2004 [70], but only 15 records between 2005 and 2016. In Hungary it was collected pre-
dominantly from phorophytes of acidic bark (Fagus sylvatica, Juniperus communis, Larix
decidua, Picea abies, Pinus nigra, P. sylvestris, and Quercus cerris) and a great number of other
species (Acer campestre, A. platanoides, A. pseudoplatanus, Aesculus hippocastanum, Alnus glu-
tinosa, Armeniaca vulgaris, Betula pendula, B. pubescens, Carpinus betulus, Castanea sativa,
Cerasus avium, Cotinus coggygria, Crataegus monogyna, Euonymus verrucosus, Fraxinus excel-
sior, F. ornus, Juglans regia, Malus domestica, Morus alba, Persica vulgaris, Populus alba, P.
Figure 2.
Hyperphyscia adglutinata (
a
) habit (scale 0.5 cm); (
b
) its distribution in Hungary (56 records).
Dots represent c. 5 km ×6 km areas. (Photo: L. L˝okös).
J. Fungi 2022,8, 600 7 of 17
J. Fungi 2022, 8, x FOR PEER REVIEW 7 of 18
tremula, Prunus domestica, Pyrus achras, P. communis, Quercus farnetto, Q. petraea, Q. pu-
bescens, Q. robur, Robinia pseudoacacia, Salix sp., Sorbus sp., S. torminalis and Tilia spp.) as
well as from decaying wood.
Figure 3. Scoliciosporum chlorococcum (a) habit (scale 0.5 mm); (b) its distribution in Hungary (322
records altogether). Dots represent c. 5 km × 6 km areas. (Photo: E. Farkas).
3.3.2. Straminella conizaeoides (Cromb.) S. Y. Kondr., Lőkös et Farkas
S. conizaeoides is a corticolous crustose species with a greyish-green granular often
sorediate thallus containing fumarprotocetraric acid. The apothecia are frequent, discs are
pale green-grey to grey-brown, with a thalline margin (Figure S4). Only 46 records were
known between 1920 and 1961, with 191 records between 1978 until 2004, and only 4 be-
tween 2005 and 2021. In Hungary it has been collected mainly from phorophytes of acidic
bark (Fagus sylvatica, Larix decidua, Pinus nigra, P. sylvestris, and Quercus cerris) and a great
number of other species (Acer campestre, A. platanoides, A. pseudoplatanus, Aesculus hippo-
castanum, Alnus glutinosa, Armeniaca vulgaris, Betula pendula, Carpinus betulus, Castanea sa-
tiva, Cerasus avium, Crataegus monogyna, Euonymus verrucosus, Fraxinus excelsior, F. ornus,
Juglans regia, Malus domestica, Persica vulgaris, Populus sp., Pyrus achras, Quercus petraea,
Robinia pseudoacacia, Sorbus torminalis, Tilia argentea, and T. platyphyllos) as well as from
decaying wood. The species grows frequently together with Hypogymnia physodes and
Scoliciosporum chlorococcum.
3.4. Species with Widely Increasing Occurrences
3.4.1. Physcia aipolioides (Nádv.) Breuss et Türk
P. aipolioides is a nitrofrequent species with a relatively large and thick thallus and
slightly pruinose lobes [71] (Figure S5). Formerly, it was collected from the Sopron Mts
Figure 3.
Scoliciosporum chlorococcum (
a
) habit (scale 0.5 mm); (
b
) its distribution in Hungary
(322 records altogether). Dots represent c. 5 km ×6 km areas. (Photo: E. Farkas).
3.3.2. Straminella conizaeoides (Cromb.) S. Y. Kondr., L˝okös et Farkas
S. conizaeoides is a corticolous crustose species with a greyish-green granular often
sorediate thallus containing fumarprotocetraric acid. The apothecia are frequent, discs
are pale green-grey to grey-brown, with a thalline margin (Figure S4). Only 46 records
were known between 1920 and 1961, with 191 records between 1978 until 2004, and only
4 between 2005 and 2021. In Hungary it has been collected mainly from phorophytes of
acidic bark (Fagus sylvatica,Larix decidua,Pinus nigra,P. sylvestris, and Quercus cerris) and
a great number of other species (Acer campestre,A. platanoides,A. pseudoplatanus,Aesculus
hippocastanum,Alnus glutinosa,Armeniaca vulgaris,Betula pendula,Carpinus betulus,Castanea
sativa,Cerasus avium,Crataegus monogyna,Euonymus verrucosus,Fraxinus excelsior,F. ornus,
Juglans regia,Malus domestica,Persica vulgaris,Populus sp., Pyrus achras,Quercus petraea,
Robinia pseudoacacia,Sorbus torminalis,Tilia argentea, and T. platyphyllos) as well as from
decaying wood. The species grows frequently together with Hypogymnia physodes and
Scoliciosporum chlorococcum.
3.4. Species with Widely Increasing Occurrences
3.4.1. Physcia aipolioides (Nádv.) Breuss et Türk
P. aipolioides is a nitrofrequent species with a relatively large and thick thallus and
slightly pruinose lobes [
71
] (Figure S5). Formerly, it was collected from the Sopron Mts and
in the Balaton region, and recently from several new localities near settlements, including
several localities also in Budapest. It is corticolous on various tree species (Acer platanoides,
A. pseudoplatanus,Aesculus hippocastanum,Fraxinus sp., Gleditsia triacanthos,Juglans regia,
J. Fungi 2022,8, 600 8 of 17
Populus
×
euramericana,P. nigra,Quercus pubescens,Q. robur,Robinia pseudoacacia, and
Ulmus campestre).
J. Fungi 2022, 8, x FOR PEER REVIEW 8 of 18
and in the Balaton region, and recently from several new localities near settlements, in-
cluding several localities also in Budapest. It is corticolous on various tree species (Acer
platanoides, A. pseudoplatanus, Aesculus hippocastanum, Fraxinus sp., Gleditsia triacanthos, Ju-
glans regia, Populus × euramericana, P. nigra, Quercus pubescens, Q. robur, Robinia pseudoaca-
cia, and Ulmus campestre).
3.4.2. Piccolia ochrophora (Nyl.) Hafellner
P. ochrophora is a corticolous crustose lichen species with convex brownish or ochra-
ceous orange apothecia covered by thick orange pruina. It has only been collected from
1987 to 2018 [72]. Being moderately nitrofrequent [52], it grows under ruderal conditions
on eutrophicated bark of Acer campestre, Populus canescens, Populus spp., Robinia pseudoaca-
cia, Salix alba or Sambucus nigra (Figure 4).
Figure 4. Piccolia ochrophora (a) habit (scale 0.5 mm); (b) its distribution in Hungary (16 records).
Dots represent c. 5 km × 6 km areas. (Photo: E. Farkas).
3.4.3. Xanthoria parietina (L.) Th. Fr.
X. parietina is a widespread, nitrofrequent, conspicuous, foliose species with yellow
to orange thallus and large lecanorine apothecia, with an orange disc (Figure S6). It was
also widely distributed throughout Hungary in older times (more than half of the records
are between 1870 and 1975). Later it was also observed in several localities, even in the
former lichen deserts of big cities. It mainly colonizes tree bark, but it is often found on
wood, rocks, and concrete. It grows on a great number of phorophytes (mostly on eu-
trophicated bark), e.g., Acer campestre, A. negundo, A. platanoides, A. pseudoplatanus, Aescu-
lus hippocastanum, Ailanthus altissima, Alnus sp., Amygdalus communis, Armeniaca vulgaris,
Artemisia monogyna, Berberis vulgaris, Betula pendula, Carpinus betulus, Castanea sativa, Celtis
Figure 4.
Piccolia ochrophora (
a
) habit (scale 0.5 mm); (
b
) its distribution in Hungary (16 records). Dots
represent c. 5 km ×6 km areas. (Photo: E. Farkas).
3.4.2. Piccolia ochrophora (Nyl.) Hafellner
P. ochrophora is a corticolous crustose lichen species with convex brownish or ochra-
ceous orange apothecia covered by thick orange pruina. It has only been collected from
1987 to 2018 [
72
]. Being moderately nitrofrequent [
52
], it grows under ruderal conditions on
eutrophicated bark of Acer campestre,Populus canescens,Populus spp., Robinia pseudoacacia,
Salix alba or Sambucus nigra (Figure 4).
3.4.3. Xanthoria parietina (L.) Th. Fr.
X. parietina is a widespread, nitrofrequent, conspicuous, foliose species with yellow to
orange thallus and large lecanorine apothecia, with an orange disc (Figure S6). It was also
widely distributed throughout Hungary in older times (more than half of the records are
between 1870 and 1975). Later it was also observed in several localities, even in the former
lichen deserts of big cities. It mainly colonizes tree bark, but it is often found on wood, rocks,
and concrete. It grows on a great number of phorophytes (mostly on eutrophicated bark),
e.g., Acer campestre,A. negundo,A. platanoides,A. pseudoplatanus,Aesculus hippocastanum,
Ailanthus altissima,Alnus sp., Amygdalus communis,Armeniaca vulgaris,Artemisia monogyna,
Berberis vulgaris,Betula pendula,Carpinus betulus,Castanea sativa,Celtis occidentalis,Cerasus
avium,Cotinus coggygria,Crataegus monogyna,Elaeagnus angustifolia,Euonymus sp., Fagus
sylvatica,Fraxinus excelsior,F. ornus,Fumana procumbens,Gleditsia triacanthos,Juglans nigra,J.
regia,Juniperus communis,Kochia prostrata,Koelreuteria paniculata,Maclura pomifera,Malus
pumila,Morus alba,M. nigra,Picea abies,Pinus nigra,P. sylvestris,Populus alba,P. canadensis,
J. Fungi 2022,8, 600 9 of 17
P. ×euramericana
,P. italica,P. nigra,P. pyramidalis,P. tremula,Prunus armeniaca,P. cerasus,P.
domestica,P. mahaleb,P. spinosa,Pyrus achras,P. pyraster,Quercus cerris,Q. pubescens,Q. robur,
Q. rubra,Ribes grossularia,Robinia pseudoacacia,Salix alba,S. fragilis,Sambucus nigra,Sorbus
torminalis,Syringa vulgaris,Tamarix gallica,Taxodium distichum,Thuja occidentalis,Tilia sp.,
Ulmus campestris, and U. glabra.
J. Fungi 2022, 8, x FOR PEER REVIEW 10 of 18
Figure 5. Coenogonium pineti (a) habit (scale 0.5 mm); (b) its distribution in Hungary (101 records,
1954–2022). Dots represent c. 5 km × 6 km areas. (Photo: E. Farkas).
3.5.3. Evernia divaricata (L.) Ach.
E. divaricata is a fruticose species with greyish-green to yellowish-green flattened
lobes, and short, thorn-like, divaricate side branches (Figure S8). Four records were
known between 1866 and 1932, and 31 records between 2000 and 2022, sometimes spread-
ing into unusual habitats. In natural habitats its thalli are pendent, beard-like, however
Hungarian specimens are mostly, compact, shrub-like. It is mostly corticolous growing on
various tree species (Acer platanoides, Ailanthus altissima, Betula pendula, Crataegus monog-
yna, Picea abies, Populus alba, P. nigra, Prunus spinosa, Quercus cerris, Q. petraea, Sorbus do-
mestica), but lignicolous specimens are also known.
4. Discussion
In comparing the various distribution maps based on historical and recent records
by studying (1) the direction of changes, (2) the possible environmental changes, and (3)
the ecological indicator values by Wirth [75], the following explanation for the various
groups can be suggested (Figure 6).
Figure 5.
Coenogonium pineti (
a
) habit (scale 0.5 mm); (
b
) its distribution in Hungary (101 records,
1954–2022). Dots represent c. 5 km ×6 km areas. (Photo: E. Farkas).
3.5. Species with Rapidly Increasing Occurrences
3.5.1. Absconditella lignicola Vˇezda et Pišút
A. lignicola is a crustose, primarily lignicolous species (Figure S7). Its thin thallus
is vivid or dark greenish-brownish, or inconspicuous, usually growing together with a
gelatinous algal biocrust. The apothecia are tiny (0.1–0.3 mm), pale waxy cream or ivory
whitish, scattered, sessile, and usually concave, urn-like [73].
Hungarian specimens were collected from decaying wood of Picea abies,Pinus nigra
and P. sylvestris in both montane and lowland habitats, mostly in lowland pine plantations,
but also in seminatural forests, accompanied by Micarea denigrata,Placynthiella icmalea, and
Trapeliopsis flexuosa.
3.5.2. Coenogonium pineti (Ach.) Lücking et Lumbsch
C. pineti is a corticolous crustose species with tiny whitish, pale orange apothecia of
0.2–0.4 mm diameter [
74
] (Figure 5). Prior to 1975, only two old records were known (1954
Buzsák and 1974 Mátra Mts, Ágasvár); since then, a further 99 records have been added.
J. Fungi 2022,8, 600 10 of 17
Thalli of C. pineti most frequently colonize the base of Alnus glutinosa and Sorbus torminalis
tree trunks, but it is also found on Acer campestre,A. platanoides,Carpinus betulus,Cornus
mas,Fagus sylvatica,Fraxinus excelsior,Larix decidua,Picea abies,Pinus nigra,P. sylvestris,
Pseudotsuga sp., Pyrus pyraster,Quercus cerris,Q. petraea,Q. pubescens, and Salix sp.; they
even occur on Robinia pseudoacacia, where the trunk can be covered up to 2 m.
3.5.3. Evernia divaricata (L.) Ach.
E. divaricata is a fruticose species with greyish-green to yellowish-green flattened
lobes, and short, thorn-like, divaricate side branches (Figure S8). Four records were known
between 1866 and 1932, and 31 records between 2000 and 2022, sometimes spreading into
unusual habitats. In natural habitats its thalli are pendent, beard-like, however Hungarian
specimens are mostly, compact, shrub-like. It is mostly corticolous growing on various
tree species (Acer platanoides,Ailanthus altissima,Betula pendula,Crataegus monogyna,Picea
abies,Populus alba,P. nigra,Prunus spinosa,Quercus cerris,Q. petraea,Sorbus domestica), but
lignicolous specimens are also known.
4. Discussion
In comparing the various distribution maps based on historical and recent records by
studying (1) the direction of changes, (2) the possible environmental changes, and (3) the
ecological indicator values by Wirth [
75
], the following explanation for the various groups
can be suggested (Figure 6).
J. Fungi 2022, 8, x FOR PEER REVIEW 11 of 18
Figure 6. The five changing distribution types of lichen species in time (years) according to possible
explanations represented by the thickness of the red line.
4.1. Species of Suboceanic Origin
Species of decreasing occurrences (chapter 3.1) belonging to this group (e.g., Lobaria
pulmonaria, Menegazzia terebrata–Figures 1 and S1) were considerably or moderately fre-
quent before the period with strong acidic air pollution (caused by SO2 pollution from
traditional heating in households, from industry and from coal heated power stations).
Their natural distribution is also characteristic of more humid climatic conditions than
those currently occurring in Hungary. They are more frequent at higher elevation and
among suboceanic conditions.
Though the decreasing SO2 pollution level (under 30 µg/m3 SO2 according to [76])
could result in the re-colonization of Lobaria pulmonaria–as recorded in 2009 [63], the drier
and warmer climate is not supporting the return of this species in larger abundance in the
country. Because of the general decline of its populations worldwide, various aspects of
its dispersal (cf. also land use history) and population dynamics were studied [77–84]. It
corresponds with ecological indicator values by Wirth [75], especially those of acidity (4–
5) and humidity (7). Its decline from its former habitats was most probably also due to the
decrease in natural forest ecosystems [48], which resulted in changing climatic, especially
microclimatic conditions (e.g., air humidity) important for this species. According to Rose
[85], Lobaria amplissima, L. pulmonaria, and L. scrobiculata are regarded as indicators of eco-
logical continuity. Lobaria pulmonaria was found to be a key species in primeval forest
preservation as an “old forest indicator” in the Eastern Carpathians [86].
Several other oceanic/suboceanic and alpine species show a declining tendency in
Hungary probably due to changes to a drier and a warmer climate; these are: e.g., Bacidia
Figure 6.
The five changing distribution types of lichen species in time (years) according to possible
explanations represented by the thickness of the red line.
4.1. Species of Suboceanic Origin
Species of decreasing occurrences (Section 3.1) belonging to this group (e.g., Lobaria
pulmonaria,Menegazzia terebrata—Figure 1and Figure S1) were considerably or moderately
frequent before the period with strong acidic air pollution (caused by SO
2
pollution from
J. Fungi 2022,8, 600 11 of 17
traditional heating in households, from industry and from coal heated power stations).
Their natural distribution is also characteristic of more humid climatic conditions than
those currently occurring in Hungary. They are more frequent at higher elevation and
among suboceanic conditions.
Though the decreasing SO
2
pollution level (under 30
µ
g/m
3
SO
2
according to [
76
])
could result in the re-colonization of Lobaria pulmonaria–as recorded in 2009 [
63
], the drier
and warmer climate is not supporting the return of this species in larger abundance in the
country. Because of the general decline of its populations worldwide, various aspects of
its dispersal (cf. also land use history) and population dynamics were studied [
77
–
84
]. It
corresponds with ecological indicator values by Wirth [
75
], especially those of acidity (4–5)
and humidity (7). Its decline from its former habitats was most probably also due to the
decrease in natural forest ecosystems [
48
], which resulted in changing climatic, especially
microclimatic conditions (e.g., air humidity) important for this species. According to
Rose [
85
], Lobaria amplissima,L. pulmonaria, and L. scrobiculata are regarded as indicators of
ecological continuity. Lobaria pulmonaria was found to be a key species in primeval forest
preservation as an “old forest indicator” in the Eastern Carpathians [86].
Several other oceanic/suboceanic and alpine species show a declining tendency in
Hungary probably due to changes to a drier and a warmer climate; these are: e.g., Bacidia
rosella,Cetrelia spp., Cladonia portentosa,Fuscopannaria leucophaea,Heterodermia speciosa,
Leptogium cyanescens,Normandina pulchella,Parmeliella triptophylla,Parmotrema crinitum,P.
perlatum,Peltigera collina,Pertusaria hemisphaerica, and Umbilicaria cylindrica.
Some of the species related to this type have a slightly increasing distribution,
e.g., Ochrolechia arborea [
87
], Peltigera leucophlebia [
62
], and the recently found Xanthoparmelia
mougeotii [88].
4.2. Species of Sub-Mediterranean Requirements
Most of these lichens, with no or few former records but with increasing occurrences
in recent decades (Section 3.2), were not collected in the 20th century and several species
only appeared in Hungary during the past 10–20 years (e.g., Flavoparmelia soredians—
Figure S2 [65]; Hyperphyscia adglutinata—Figure 2;Solenopsora candicans—Figure S3 [69]).
Since these lichens are still rarely collected in Hungary, it is difficult to explain their
presence. Some hypotheses, such as that described by Wirth in 1997 [
40
] may provide some
explanation since some lichens may expand their distribution. The background to these
changes may be a joint effect of an as yet unknown cause, such as a minor environmental
change to the microhabitat of the species and changed characters (morphological, phys-
iological, and/or chemical) due to a possible change in the expression of genes [
89
–
92
].
Flavoparmelia soredians was also collected in urbanized places, outside of its preferred sub-
oceanic habitats. Wirth [
40
] also suggested the decrease in the SO
2
level and the change
of climate to a milder one in Germany as a reason. Seaward and Coppins [
93
] mention
hypertrophication as the reason for its spread in the British Isles. Anthropogenic effects are
also confirmed by Nygaard and Tønsberg [94] for its immigrant nature in Norway.
Solenopsora candicans was overlooked in Hungary but collected recently at several new
localities [
69
]. The recent spread of Hyperphyscia adglutinata is even more obvious (Figure 2);
it is regarded as a sub-Mediterranean element, growing on nutrient rich trees frequently in
well illuminated conditions [95].
All these species are of sub-Mediterranean origin, but due to climatic warming they
have found more advantageous conditions and therefore are spreading in Hungary. Further
species with a similar distribution type in Hungary are Leptogium ferax [
96
], Parmelia
submontana Hale [
97
], and Xanthoparmelia verrucigera. Among the ecological indicator
values by Wirth [
75
], temperature (value 9) mostly justifies the change in distribution for
these species.
J. Fungi 2022,8, 600 12 of 17
4.3. Acidofrequent Species
Scoliciosporum chlorococcum,Straminella conizaeoides,Hypogymnia physodes, and Lepraria
incana are more difficult to find recently than a few decades ago. These are species with
increasing, then decreasing occurrences (Section 3.3.); especially, Scoliciosporum chlorococcum
(Figure 3) and Straminella conizaeoides (Figure S4) with hardly any herbarium records since
the beginning of the 20th century in Hungary, while they were reported in lists indicating
acidic air pollution [
11
] and later also discovered in Budapest, and these were among its
most frequent species [
10
,
98
]. Since acidic air pollution has decreased considerably [
46
],
these acidofrequent species, tolerating acidic air pollution, have been disappearing since
2000. Both species have ecological indicator values of 2–3 by Wirth [75], which are charac-
teristic of an acid environment.
4.4. Nitrofrequent Species
While acidic air pollution is decreasing, nitrogen-containing pollutants (NO
x
and
NH
3
) are increasing, especially in and near settlements [
46
,
47
,
93
]. These conditions have
affected the distribution of the so-called nitrophilous species (e.g., Phaeophyscia orbicularis,
Xanthoria parietina–Figure S6), however, we prefer a more neutral term: nitrofrequent. Their
ecological indicator value of 8 by Wirth [
74
] in this respect is high. These species were
already more or less frequent earlier, almost irrelevant of the level of acidic pollution, but
when it started to decrease and nitrogen-containing pollutants increased (cf. Section 3.4), the
nitrofrequent species spread and their distribution increased significantly. Physcia aipolioides
(Figure S5) is an interesting example; in Slovakia, Czechia, and Austria, it demonstrates
a similar continuous distribution type with a few outliers in fairly remote localities. An
increase in the number of localities in Czechia and Slovakia, together with the toxitolerance
of the lichen, led in the 1980s and the 1990s to the assumption that the species spread
invasively [
71
]. The lichen also occurs in Hungary, and the records from Bulgaria and
Montenegro indicate that its distribution area is probably even much larger. The lichen has
the potential to spread further.
The species increasing under nitrogen-rich conditions in Hungary are Amandinea
punctata,Candelariella reflexa,Catillaria nigroclavata,Lecania cyrtella,L. naegelii,Lepraria elobata,
L. lobificans,Phaeophyscia orbicularis,Physcia adscendens,P. aipolioides (Figure S5), Piccolia
ochrophora (Figure 4), and Xanthoria parietina (Figure S6).
4.5. Rapidly Spreading Species of Uncertain Reason
The distribution tendencies of certain species, namely the rapid and sudden increase of
occurrences (cf. Section 3.5), makes them very similar to invasive vascular plant species [
99
].
The earliest Hungarian specimens of Coenogonium pineti were collected in Buzsák (Bels˝o-
Somogy) in 1954, almost 20 years later in valley “Csörg˝o-patak völgye,” Mátra Mts [
100
],
then in Barcsi ˝
Osborókás TvT and in Zselic in 1987 [
101
]. From 1991, several new localities
were recognized practically from all regions of the country. These collection data suggest
that changes of environmental conditions in recent decades support the distribution of this
species throughout Hungary, but no closer explanation was found for its spread.
Evernia divaricata also had scattered records in the past, but in recent decades it
is collected more and more frequently in Hungary. Another example is Absconditella
lignicola [
73
]. It was found in 2009 in the ˝
Orség (W Hungary), then it was collected from
34 localities in the Börzsöny, Buda, and Mátra Mts, the Danube–Tisza Interfluve, the
Transdanubian Hills, the Nyírség, and the ˝
Orség.
It is difficult to explain the status of the above species, and most probably a combina-
tion of several reasons (e.g., pollution, preference of climate or substrate) are responsible
for their appearance and spread. The various ecological indicator values (by Wirth [
75
])
themselves do not explain the distributions since the different values characterizing the
species fall in a wide range or they show even opposite trends.
J. Fungi 2022,8, 600 13 of 17
5. Conclusions
The study of a large amount of herbarium material and literature records made it
possible to establish different distribution types of lichens. The analyses—also based on
literature sources of background data—including Wirth’s ecological indicator values [
75
],
have justified the possible reasons for the appearance of these distribution types.
The discussed trends are known for some species at a global scale or European level,
other examples are characteristic for Central Europe or Hungary. Since the distribution
is strongly correlated to background environmental conditions (e.g., air pollution, land
use intensity, global warming, substrate type), the analyses of distribution maps have
a great value in biomonitoring. By studying the distribution maps of lichen bioindica-
tors, tendencies of climate change and type of pollution can be determined and further
changes can be predicted. However, further extended collections with precise collection
design in space and time are necessary to provide current distributional data suitable for a
statistical analysis.
Supplementary Materials:
The following supporting information can be downloaded at: https:
//www.mdpi.com/article/10.3390/jof8060600/s1, Figure S1: Menegazzia terebrata (a) habit (scale
1 cm); (b) its distribution in Hungary (10 records). Dots represent c. 5
×
6 km areas. (Photo:
©
E.
Timdal); Figure S2: Flavoparmelia soredians (a) habit (scale 0.5 cm); (b) its distribution in Hungary
(
19 records
). Dots represent c. 5 km
×
6 km areas. (Photo: E. Farkas); Figure S3: Solenopsora candicans
(a) habit (scale 0.5 cm); (b) its distribution in Hungary (13 records). Dots represent c. 5 km
×
6 km
areas. (Photo: E. Farkas); Figure S4: Straminella conizaeoides (a) habit (scale 0.1 cm); (b) its distribution
in Hungary (241 records altogether). Dots represent c. 5 km
×
6 km areas. (Photo: E. Farkas);
Figure S5: Physcia aipolioides (a) habit (scale 0.5 cm); (b) its distribution in Hungary (31 records). Dots
represent c. 5 km
×
6 km areas. (Photo: L. L˝okös); Figure S6: Xanthoria parietina (a) habit (scale 1 cm);
(b) its distribution in Hungary (1023 records). Dots represent c. 5 km
×
6 km areas. (Photo: E. Farkas);
Figure S7: Absconditella lignicola (a) habit (scale 0.5 cm); (b) its distribution in Hungary (37 records,
2009–2022). Dots represent c. 5 km
×
6 km areas. (Photo: E. Farkas); Figure S8: Evernia divaricata
(a) habit (scale 1 cm); (b) its distribution in Hungary (35 records). Dots represent c. 5 km
×
6 km
areas. (Photo: G. Matus); Table S1: Distribution records of illustrated species with main literature
sources (containing published distribution records) and additional herbarium records (herbarium
code, locality code [57,58], year).
Author Contributions:
Conceptualization, E.F. and L.L.; methodology, E.F. and L.L.; formal analysis,
E.F. and L.L.; investigation, G.M., M.S., K.V. and N.V.; resources, G.M., M.S., K.V. and N.V.; data
curation, G.M., M.S., K.V. and N.V.; writing—original draft preparation, E.F.; writing—review and
editing, L.L.; visualization, E.F. and L.L.; supervision, E.F. and L.L.; project administration, E.F.;
funding acquisition, E.F. All authors have read and agreed to the published version of the manuscript.
Funding:
Our work is supported by the National Research Development and Innovation Fund, grant
number NKFI K 124341 as well as Human Resources Development Operational Programme/Hungarian
Ministry of Human Resources grant number EFOP 3.4.3-16-2016-00021.
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Data Availability Statement:
The following public databases were used: IndexFungorum (http://
www.indexfungorum.org/, accessed on 27 September 2021); Index Herbariorum (http://sweetgum.
nybg.org/science/ih/, accessed on 27 September 2021); MycoBank (https://www.mycobank.org/,
accessed on 27 September 2021); Recent literature on lichens (http://nhm2.uio.no/botanisk/lav/
RLL/RLL.HTM, accessed on 27 September 2021). The data presented in this study are available in
the referred publications [
49
,
63
,
65
,
69
–
72
], if otherwise not, then these are available on request from
the corresponding author [E.F.]. Further records (herbarium and literature) are summarized in Table
S1 [link for supplement of current paper is to be added here]. Data of herbarium records are available
upon request from the curators of the given herbaria [see Index Herbarium at above link].
J. Fungi 2022,8, 600 14 of 17
Acknowledgments:
We express our special thanks to M.R.D. Seaward (Bradford University, UK) for
his advice and revision of the English text. We are indebted to the curators of herbaria (especially
EGR, GODO, PECS, SZE, and SZO) for specimen information. We are grateful for the appreciating
words and the useful comments and questions of our three anonymous reviewers.
Conflicts of Interest:
The authors declare no conflict of interest. The funders had no role in the design
of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript, or
in the decision to publish the results.
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