The gilled fungi and boletes of Iran: diversity, systematics, and nrDNA data

Abstract

A first annotated checklist of agarics and boletes in Iran is presented based on literature and new collections. A total number of 128 new DNA sequences, obtained from the nrDNA ITS region as well as the nrLSU, is provided. Based on vouchered specimens, 19 species are newly recorded from Iran, all provided with nrDNA data from basidiomata. Overall, 585 species (comprising 556 agarics and 29 bolete species) are recorded from Iran, representing 147 genera and 34 families. The order Agaricales encompasses 82% of the species. Twenty-eight species are excluded from the Iranian mycobiota. The Hyrcanian forests harbor 79% of the species, with Mazandaran being the most species-rich province in the country. The three largest genera are Russula, Cortinarius, and Inocybe. We also scrutinized the available DNA sequences of Iranian agarics and boletes deposited in public databases. It is shown that 64 species of agarics and boletes in Iran are from environmental sequences, and nine species have been retrieved as plant endophytes. In total, only 24% of Iranian species are shown to have at least one nrITS sequence in GenBank or UNITE. Our analyses reveal that 42% of records in Iran arise merely from abstracts presented at various conferences, most of which lack sufficient characterization and are suggested to be considered tentatively. General recommendations are given to avoid dissemination of low quality and ambiguous species records. This study fills in some of the gaps in our knowledge of agarics and boletes in Iran and provides a framework for biodiversity and phylogenetic studies.
Key words: Agaricomycetes, checklists, Hyrcanian forests, macrobasidiomycetes, mushrooms, Zagros forests

Full text

Mycologia Iranica 7(1): 1 – 43, 2020 DOI: 10.22043/mi.2021.123456
Submitted 16 April 2020, accepted for publication 21 June 2020
 Corresponding Author E-mail: ghobadnejhad@irost.ir
© 2020, Published by the Iranian Mycological Society
http://mij.areeo.ac.ir
Original Article
The gilled fungi and boletes of Iran: diversity, systematics,
and nrDNA data
M. Ghobad-Nejhad✉
Department of Biotechnology, Iranian Research
Organization for Science and Technology (IROST),
Tehran, Iran
E. Langer
Department of Ecology, University of Kassel, Heinrich-
Plett-Str. 40, D-34132 Kassel, Germany
V. Antonín
Department of Botany, Moravian Museum, Zelný trh 6,
659 37 Brno, Czech Republic
G. Gates
Tasmanian Institute of Agriculture, Private Bag 98,
Hobart, Tasmania 7001, Australia
J. Noroozi
Department of Botany and Biodiversity Research,
University of Vienna, Austria
R. Zare
Department of Botany, Iranian Research Institute of
Plant Protection (IRIPP), Agricultural Research,
Education and Extension Organization (AREEO),
Tehran, Iran
Abstract: A first annotated checklist of agarics and
boletes in Iran is presented based on literature and new
collections. A total number of 128 new DNA sequences,
obtained from the nrDNA ITS region as well as the
nrLSU, is provided. Based on vouchered specimens, 19
species are newly recorded from Iran, all provided with
nrDNA data from basidiomata. Overall, 585 species
(comprising 556 agarics and 29 bolete species) are
recorded from Iran, representing 147 genera and 34
families. The order Agaricales encompasses 82% of the
species. Twenty-eight species are excluded from the
Iranian mycobiota. The Hyrcanian forests harbor 79% of
the species, with Mazandaran being the most species-
rich province in the country. The three largest genera are
Russula, Cortinarius, and Inocybe. We also scrutinized
the available DNA sequences of Iranian agarics and
boletes deposited in public databases. It is shown that 64
species of agarics and boletes in Iran are from
environmental sequences, and nine species have been
retrieved as plant endophytes. In total, only 24% of
Iranian species are shown to have at least one nrITS
sequence in GenBank or UNITE. Our analyses reveal
that 42% of records in Iran arise merely from abstracts
presented at various conferences, most of which lack
sufficient characterization and are suggested to be
considered tentatively. General recommendations are
given to avoid dissemination of low quality and
ambiguous species records. This study fills in some of
the gaps in our knowledge of agarics and boletes in Iran
and provides a framework for biodiversity and
phylogenetic studies.
Keywords: Agaricomycetes, checklists, Hyrcanian
forests, macrobasidiomycetes, mushrooms, Zagros
forests
INTRODUCTION
Gilled fungi and boletes (often referred to as
‘mushrooms’) are among the most prevalent groups of
macrofungi. Traditionally they are the best-studied fungi
in many parts of the world probably on account of their
potential value as a food source but also their
conspicuous and often eye-catching fruitbodies. The
number of described species worldwide now exceeds the
ca. 14,500 species of Chang & Miles (2004) and Kirk et
al. (2008). Gilled fungi, also commonly known as
agarics, share a structural characteristic, i.e., vertical
plates, denoted as lamellae or more colloq-uially as gills
(with varying shapes, sizes, thicknesses, distances from
each other) which serve as spore-producing tissue on the
undersurface of the cap and are a highly efficient way of
increasing the surface area of the fertile part of the
fungus. Boletes have tubes which are, in effect, simply
rolled up gills, with some members of the Boletales
having true gills e.g., the genus Phylloporus. According
to molecular evolutionary studies, the formation of gills
has evolved several times and therefore, is considered to
have been a characteristic worth preserving to assist in
spore dispersal and species survival (Hibbett et al. 1997).

2 Mycologia Iranica - Vol. 7, 2020
Gilled fungi and boletes are very diverse,
morphologically, chemically and ecologically. They
range in size from a few millimeters in diameter (e.g.,
species of Mycena, Marasmius and Coprinellus), to
more than a metre across the cap (Macrocybe titans,
Phlebopus marginatus). Important variable character-
istics include the presence or absence of a stipe, of
veils, or veil remnants. The species exhibit numerous
colors, odors and flavours. Chemically, some gilled
taxa e.g., in the genera Amanita, Galerina, and
Cortinarius contain a number of the most toxic natural
compounds namely amatoxins, phallotoxins and
orellanine; boletes do not have such lethal compounds
although the genus Rubroboletus has a toxin
(bolesatine) that causes severe gastrointestinal
problems. In contrast, various species, e.g. Agaricus
campestris, Amanita caesarea, and Boletus edulis are
considered edible. There are other taxa of the genera
Psilocybe, Copelandia, Gymnopilus, Inocybe,
Panaeolus, Pholiotina and Pluteus, that produce
hallucinogenic substances such as psilocybin and/or
psilocin. Ecologically, as symbiotic partners involved
in plant development (mycorrhizae), and as decomp-
osers, with enzymes breaking down lignin and
cellulose in woody and herbaceous debris, they are
critical for carbon recycling and ecosystem
functioning. Some species e.g. Armillaria luteobubalina
and Moniliophthora perniciosa are pathogenic and/or
parasitic and cause destruction of eucalypt forests and
cacao crops respectively (Shearer & Tippett 1988,
Meinhardt et al. 2008). The substrates also vary, as
they grow on soil, wood, dung and litter. Many agarics
and boletes also provide food and medicines for
humans, and nutrition and habitat for other organisms.
Biogeographically, they are widespread in temperate,
equatorial, arctic and sub-antarctic ecosystems,
possibly wherever a vegetation layer exists.
From a systematic viewpoint, the first compreh-
ensive phylogenetic studies on agarics were published
by Moncalvo et al. (2002) and Matheny et al. (2006).
They were, and still are, the basis for the next studies
of individual clades and groups of agarics worldwide.
Concerning boletoid genera, the study by Wu et al.
(2014) opened a broad field for defining many new
genera. A ten-year phylogenetic overview of the
Basidiomycota and related phyla in relation to DNA-
based phylogeny was published by Zhao et al. (2017).
An extensive study dealing with global patterns of
mushroom evolution was recently published by Varga
et al. (2019). According to the most recent account of
a systematic outline of the Basidiomycota (He et al.
2019), the class Agaricomycetes currently contains 22
orders, of which agarics and boletes are distributed across
the six orders Agaricales, Boletales, Cantharellales,
Hymenochaetales, Polyporales, and Russulales.
Iran, with a wide elevational range, from 26 m b.s.l.
along the southern coastline of the Caspian Sea, to
5,671 m a.s.l. at Damavand Mt. in Central Alborz,
harbors diverse climates and habitats. Vast hot and dry
deserts with a precipitation of less than 25 mm/yr in
central Iran, coincide with narrow sub-tropical humid
regions at the northern parts of the country with
precipitation exceeding 1,800 mm/yr. In Iran three
macrobioclimates have been recognized, namely,
mediterranean, tropical and temperate, largely
correlating with the three biogeographical regions
Irano-Turanian, Saharo-Sindian and Euro-Siberian,
respectively (Djamali et al. 2011). Large parts of the
country are characterized by continentality, with hot
and dry summers, cold and harsh winters, and low
precipitation, while forests are mainly confined to the
northern and western borders (Zohary 1973, Djamali
et al. 2012).
From a biodiversity viewpoint, Iran is at the
crossroad of two global biodiversity hotspots, i.e. the
Irano-Anatolian and the Caucasus (Mittermeier et al.
2005, Manafzadeh et al. 2017, Noroozi et al. 2019),
covering the major mountain ranges of the country.
Diverse topography and climate have resulted in the
development of diverse flora and vegetation types.
More than 8,000 vascular plant species are known
from Iran with ca. 32% endemism (Noroozi et al.
2019). The number of woody plants in Iran is about
956 species (Mozaffarian 2005). Desert and semi-
desert steppes, halophytes, montane grasslands,
shrublands and woodlands, deciduous temperate
forests, wetlands and alpine habitats are among the
major vegetation types of the country (Noroozi 2020).
Hyrcanian forests, on the northern slopes of the
Alborz mountain range, are at the southeastern reaches
of the Caucasus biodiversity hotspot. Despite its small
area (ca. 5% of the total area of Iran), the majority of
the Iranian macromycetes come from the Hyrcanian
region (e.g. Ghobad-Nejhad 2011, Ghobad-Nejhad &
Hallenberg 2012). Broad-leaved dense forests have
developed due to the high precipitation (ca. 700 to
2000 mm/yr) and mild climate of the region. These
forests are known to belong to the northern hemisphere
Pleistocene glacial refugia, and contain several Arcto-
Tertiary elements, such as Celtis australis, Parrotia
persica, Pterocarya fraxinifolia and Zelkova carpinifolia
as well as evergreen plants (Zohary 1973, Frey et al.
1999, Akhani et al. 2010, Gholizadeh et al. 2020). The
Zagros forests in western Iran are comprised of
xerophytic open woodlands of deciduous trees and
shrubs mainly dominated by Quercus species and
constitute less than 40% of the forest cover in Iran
(Sagheb Talebi et al. 2014). Among the few coniferous
forests native to Iran, Juniperus excelsa woodlands are
noteworthy, sparsely covering the subalpine zone of
the Irano-Turanian parts of the Iranian mountains,
mostly distributed at 2000 to 3000 m a.s.l.
Macrofungi in Iran are far less studied compared to
microfungi. Historically, mycology in Iran has been
divided up into five distinct periods, with publications
starting from 1860 (Ershad & Zare 2014). While the
majority of studies are characterized by records on
microascomycetes, the second period in Iranian
mycology (1860–1941) also harbors publications
containing some agarics and boletes. Interestingly, the
oldest publication related to the fungi of Iran (Buhse
1860) contains eight species of agarics and boletes

GHOBAD-NEJHAD ET AL.: The gilled fungi and boletes of Iran 3
from the country, most of them later recorded in other
publications (Table 1). The second oldest literature on
Iranian fungi (Rabenhorst 1871) enumerates two
species of this group. Later, publications by Petrak
(1941, 1940, 1949) alone and jointly with the first
Iranian mycologist E. Esfandiari (Petrak & Esfandiari
1941), appear, recording a few species of agarics and
boletes from Iran (Table 1).
Most of the fungi of Iran recorded until 2008
(excluding lichenized fungi, clinical fungi, and some
other taxa) have been listed in a book by Ershad
(2009). In this book, a list of 3229 fungal species has
been sorted in alphabetical order, but with no attempt
at taxonomic classification. Therefore, to figure out the
number of any fungal groups from this book, a full
systematic categorization of the records is needed.
With regard to the number of macrobasidiomycetes
in Iran, no clear figure is available, but up to now ca.
430 aphyllophoroid species are known from the
country, covering polypores (poroids), corticioids,
hydnoids, clavarioids and heterobasidiomycetes
(Ghobad-Nejhad & Hallenberg 2012, Ghobad-Nejhad
& Langer 2017); of these, 132 species are polypores
(Ghobad-Nejhad & Bernicchia 2019). Statistics on
agarics and boletes recorded from the country are
largely lacking. Therefore, the aim of this study is to
provide a first account on the diversity of gilled fungi
and boletes in Iran, with annotations on their
distribution, classification, and nrDNA.
MATERIALS AND METHODS
Species data
The study group includes species commonly
known as gilled fungi (agarics) and boletes. Basically,
the taxa such as Schizophyllum commune and
Daedalea/Daedaleopsis/Lenzites which have some
sort of 'gills' (but studied among aphyllophoroid
fungi), are not included. Data on agarics and boletes in
this study were obtained from a thorough survey of
scientific journal articles and books published up to the
year 2020 and specimens collected by the first author.
Moreover, sequences of Iranian material in GenBank
and UNITE databases were incorporated after full
identity check (see below). The basidiomata specimens
were collected during 2000–2019 in different
provinces of Iran. Samples were dehydrated using a
mushroom drier or were air-dried. Macro- and micro-
morphological characters of the specimens were
studied under a binocular and a light microscope, using
bright field or phase contrast optics. Microscopy
routines generally followed Largent et al. (1977).
Several identification keys (e.g., Knudsen &
Vesterholt 2008, Justo et al. 2011, Antonín et al. 2013,
Zervakis et al. 2014, Mahdizadeh et al. 2016, Voto et
al. 2019, etc.), and some of the literature in He et al.
(2019) for specific genera were used. A full list of
examined specimens is available in Suppl. Table.
Vouchers were deposited at Iranian Cryptgamic
Herbarium (Index Herbariorum acronym ICH).
Species current names and species authorities mostly
follow MycoBank (www.mycobank.org), and Index
Fungorum (www.indexfungorum.org). Recent or
otherwise relevant taxonomic revisions and phylogenetic
publications were also consulted. Records published in
journals not approved by the Iranian Ministry of
Science, Research and Technology (‘black-listed
journals’) are not included here.
Molecular study
A clean portion of the spore-producing part of the
dried basidiomata was taken for molecular study. For
DNA extraction, PCR and sequencing, we generally
followed the protocol by Bußkamp et al. (2020): from
each basidioma, 1–2 mg of tissue was suspended in
100 μl TE buffer in a 1.5 ml tube. Immediately, for cell
lysis, a microwave (600 W) was used twice for 1 min
each, with a pause of 30 s followed by cooling the
tubes to −20 °C for 20 minutes. The tubes were
centrifuged at 10,000 rpm for 5 minutes. For PCR, 100
times diluted portion of the supernatant was used. The
PCR primers for amplification of ITS1, 5.8S and ITS2
regions were ITS1F/ITS4 or ITS1/ITS4 (White et al.
1990, Gardes & Bruns 1993). For PCR, a 45 μl master
mix (QIAGEN, Hilden, Germany) was used, adding 5
μl of extracted DNA. PCR was performed with initial
denaturation at 94°C for 3 min, followed by 29 cycles
of denaturation at 94°C for 30 s, annealing at 55°C for
45 s, and extension at 72°C for 60 s. Final elongation
was performed at 72°C for 7 min. The D1–D4 domain
of nuclear ribosomal large subunit (LSU; 28S) was
amplified using primers LR0R and LR5 (Vilgalys &
Hester 1990). The PCR thermal cycling for LSU
differed from the ITS only by an increased cycle
extension time (90 s per cycle). PCR products were
visualized on a 1% agarose gel stained with GelRed
fluorescence dye (Biotium, Hayward, CA, USA).
Bands were cut out and cleaned with QIAquick PCR
Purification Kit (QIAGEN, Hilden, Germany).
Purified products were sequenced using Sanger
sequencing method (Sanger et al. 1977). The obtained
DNA sequences were edited using MEGA6 (Tamura
et al. 2013) and DNA Sequence Assembler v4 (Heracle
BioSoft 2013, www.DnaBaser.com), checked with
BLASTn (Altschul et al. 1990) and UNITE (Nilsson et
al. 2019), and submitted to GenBank.
We also searched through GenBank and UNITE
databases extracting DNA sequences of agarics and
boletes obtained from Iranian material, with a special
focus on the ITS sequences. The sequences’ associated
metadata were carefully checked, and BLASTn
searches were performed against sufficiently identified
and reliable sequences in those databases. In UNITE,
the Species Hypothesis (SH) with 0–1.5% threshold
was accepted. Accession numbers with incorrect,
ambiguous, or otherwise unverifiable identity were
excluded. Selected ITS accession numbers were
inserted in Table 1.

4 Mycologia Iranica - Vol. 7, 2020
Table 1. Agarics and boletes in Iran. Species names in bold italics are current names, followed by synonyms in the
corresponding literature. Records new to Iran are marked with an asterisk (*). The black circle (●) means that at least one DNA
sequence exists from Iranian material; for these, representative ITS sequence(s) from GenBank or UNITE is presented. Accession
numbers in bold were obtained in this study. Records with an exclamation mark (!) are recommended to be regarded as tentative
only (see the text). Abbreviations: SH: Species Hypothesis; env.: environmental sequence.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
●Agaricus arvensis Schaeff.
Agaricales,
Agaricaceae
Alborz, East Azerbaijan,
Gilan, Golestan,
Hamadan, Mazandaran,
Tehran, West
Azerbaijan, Zanjan
Saber 1994a, Fadavi et al. 2015,
Mahdizadeh 2016, this study
MT535720
●Agaricus bisporus (J.E. Lange)
Imbach
Agaricales,
Agaricaceae
Ardabil, East
Azerbaijan, Hormozgan,
Khuzestan, Kurdistan,
Lorestan, Markazi,
Tehran, West
Azerbaijan
Saber 1994a, Fadavi et al. 2015,
Mahdizadeh 2016
MK287948
●Agaricus bitorquis (Quél.) Sacc.
Agaricales,
Agaricaceae
Alborz, Chaharmahal
and Bakhtiari, East
Azerbaijan, Fars,
Isfahan, Khuzestan,
Kohgiluyeh and Boyer-
Ahmad, Markazi,
Razavi Khorasan,
Tehran, West
Azerbaijan
Watling & Gregory 1977, Zokaei
2001, Asef 2011b, Mahdizadeh
2016, this study
MT535709
!Agaricus bresadolanus Bohus
Agaricales,
Agaricaceae
Golestan
Mahdizadeh 2016
!Agaricus brunneolus (J.E. Lange)
Pilát
Agaricales,
Agaricaceae
Golestan
Mahdizadeh 2016
Agaricus campestris L.
Agaricales,
Agaricaceae
Ardabil, East
Azerbaijan, Golestan,
Mazandaran
Asef 2014, Mahdizadeh 2016
!Agaricus depauperatus (F.H.
Møller) Pilát
Agaricales,
Agaricaceae
West Azerbaijan
Mahdizadeh 2016
●Agaricus devoniensis P.D. Orton
Agaricales,
Agaricaceae
Hormozgan,
Kermanshah
Mahdizadeh 2016,
Seidmohammadi et al. 2018b
MH173888
!Agaricus gennadii (Chatin &
Boud.) P.D. Orton
Agaricales,
Agaricaceae
West Azerbaijan
Mahdizadeh 2016
●Agaricus iodosmus Heinem.
Agaricales,
Agaricaceae
East Azerbaijan, Tehran
Mahdizadeh et al. 2016, this study
KT824784,
MT535702
●Agaricus iranicus Mahdizadeh,
Safaie, Goltapeh, L.A. Parra &
Callac
Agaricales,
Agaricaceae
Gilan
Mahdizadeh et al. 2017a
KY474556 (also
several sequences
available from
different DNA
regions)
!Agaricus langei (F.H. Møller)
F.H. Møller
Agaricales,
Agaricaceae
Golestan
Saber 1994a, Mahdizadeh 2016
●Agaricus litoralis (Wakef. & A.
Pearson) Pilát
Agaricus biberi Hlaváček
Agaricales,
Agaricaceae
Alborz, East Azerbaijan,
Mazandaran, Semnan
Mahdizadeh 2016, this study
MT535711
●Agaricus moelleri Wasser
Agaricales,
Agaricaceae
Gilan, Golestan,
Mazandaran
Mahdizadeh et al. 2016
KT824787
(several more in
GenBank)
●Agaricus nevoi Wasser
Agaricales,
Agaricaceae
Bushehr, Fars,
Hormozgan,
Kermanshah,
Khuzestan, Kohgiluyeh
and Boyer-Ahmad
Mahdizadeh 2016,
Seidmohammadi et al. 2018b
MH173866
●Agaricus phaeolepidotus (F.H.
Møller) F.H. Møller
Agaricales,
Agaricaceae
Mazandaran
Mahdizadeh et al. 2016
KT824790
! Agaricus pseudolutosus (G.
Moreno, Esteve-Rav., Illana &
Heykoop) G. Moreno, L.A. Parra,
Esteve-Rav. & Heykoop
Agaricales,
Agaricaceae
Gilan
Mahdizadeh 2016
*●Agaricus pseudopratensis
(Bohus) Wasser
Agaricales,
Agaricaceae
Tehran
this study
MT535735,
MT535748
●Agaricus subrufescens Peck
Agaricales,
Agaricaceae
Gilan
Mahdizadeh et al. 2017b
KT983412
●Agaricus xanthodermus Genev.
Agaricales,
Agaricaceae
Gilan, Golestan
Mahdizadeh et al. 2016
KT824789

GHOBAD-NEJHAD ET AL.: The gilled fungi and boletes of Iran 5
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
!Agrocybe acericola (Peck) Singer
Agaricales,
Strophariaceae
Golestan, West
Azerbaijan
Saber 1993f, Saber &
Mehravaran 2004
●Agrocybe dura (Bolton) Singer
Pholiota vermiflua (Peck) Sacc.
Agaricales,
Strophariaceae
Gilan, Golestan,
Kermanshah,
Qazvin, Tehran,
West Azerbaijan
Saber 1991b, Saber 1993f, Saber
& Mehravaran 2004, Fadavi et
al. 2013, this study
MT535714 (more
in Table 2)
Agrocybe ochracea Nauta
Agaricales,
Strophariaceae
Kermanshah
Fadavi et al. 2013
!Agrocybe paludosa (J.E. Lange)
Kühner & Romagn. ex Bon
Agaricales,
Strophariaceae
Gilan
Saber 1995f
!Agrocybe pediades (Fr.) Fayo
Agrocybe semiorbicularis (Bull.)
Fayod
Agaricales,
Strophariaceae
Tehran
Saber 1993f, Saber & Zangeneh
2002
●Agrocybe praecox (Pers.) Fayod
Agaricus togularis Bull. ex Pers.,
Agrocybe sphaleromorpha (Bull. ex
Pers.) Fayod, Conocybe togularis
(Bull. ex Pers.) Kühner, Pholiotina
togularis (Bull. ex Pers.) Fayod
Note: some synonyms are tentative
(unpublished research)
Agaricales,
Strophariaceae
East Azerbaijan,
Gilan, Golestan,
Mazandaran, West
Azerbaijan
Buhse 1860, Saber 1993f, Saber
& Mehravaran 2004, Asef
2007b, this study
MT535701
!Agrocybe pusiola (Fr.) R. Heim
Agaricales,
Strophariaceae
Tehran
Saber 1993f
!Agrocybe tabacina (DC.) Konrad &
Maubl.
Agaricales,
Strophariaceae
East Azerbaijan
Asef 2007b
!Agrocybe vervacti (Fr.) Singer
Agaricales,
Strophariaceae
Gilan,
Kermanshah
Saber 1993f
!Alnicola escharioides (Fr.) Romagn.
Naucoria escharoides (Fr.) P. Kumm.
Agaricales,
Hymenogastraceae
Gilan
Saber 1995f
!Amanita atkinsoniana Coker
Agaricales,
Amanitaceae
Mazandaran
Saber & Zangeneh 2004
Amanita battarrae (Boud.) Bon
Agaricales,
Amanitaceae
Mazandaran
Bahram et al. 2006b, Asef &
Etemad 2016
Amanita caesarea (Scop.) Pers.
Agaricales,
Amanitaceae
Mazandaran
Bahram et al. 2006a, 2006b,
Asef & Etemad 2016
Amanita ceciliae (Berk. & Broome)
Bas
Agaricales,
Amanitaceae
Mazandaran
Bahram et al. 2006b, Asef &
Etemad 2016
Amanita crocea (Quél.) Singer
Agaricales,
Amanitaceae
Mazandaran
Bahram et al. 2006b, Asef &
Etemad 2016
Amanita eliae Quél.
Agaricales,
Amanitaceae
Mazandaran
Saber 1995a
Amanita excelsa (Fr.) Bertill.
Amanita spissa (Fr.) Bertill.
Agaricales,
Amanitaceae
Golestan,
Mazandaran
Saber 1991a, Saber 1995a, Asef
& Etemad 2016
Amanita gemmata (Fr.) Bertill.
Agaricales,
Amanitaceae
Mazandaran
Saber 1991a, Saber 1995a
●Amanita lividopallescens (Secr. ex
Boud.) Kühner & Romagn. agg.
Notes: regarded here as aggregate.
Agaricales,
Amanitaceae
Ilam, Lorestan
Yousefshahi et al. 2020
(wrongly as A. crocea), this
study
MT535691
●Amanita pantherina (DC.) Krombh.
Agaricales,
Amanitaceae
East Azerbaijan,
Golestan,
‘Hyrcan’,
Mazandaran
Saber 1991a, Saber 1995a, Asef
& Tavanaei 2004, Bahram et al.
2012, Asef & Etemad 2016
FR852274 (env.,
SH1932544.08FU)
Amanita phalloides (Fr.) Link
Agaricales,
Amanitaceae
Gilan,
Mazandaran, West
Azerbaijan
Fallahyan 1973, Saber 1995a,
Asef & Etemad 2016
●Amanita rubescens Pers.
Agaricales,
Amanitaceae
Gilan, ‘Hyrcan’
Bahram et al. 2006b, 2012
FR852273 (env.,
SH2271424.08FU)
Amanita strobiliformis (Paulet ex
Vittad.) Bertill.
Agaricales,
Amanitaceae
Mazandaran
Saber & Zangeneh 2000
Amanita umbrinolutea (Secr. ex
Gillet) Bataille
Agaricales,
Amanitaceae
Lorestan
Saber 1997a, Bahram et al.
2006b, Asef & Etemad 2016
Amanita vaginata (Bull.) Lam. s.l.
Notes: more species may be included
under this name.
Agaricales,
Amanitaceae
East Azerbaijan,
Golestan,
Mazandaran
Saber 1991a and 1995a, Asef &
Tavanaei 2004, Asef & Etemad
2016
Amanita verna (Bull.) Lam.
Agaricales,
Amanitaceae
Gilan, Mazandaran
Saber 1991a, Saber 1995a
Ampulloclitocybe clavipes (Pers.)
Redhead, Lutzoni, Moncalvo &
Vilgalys
Clitocybe clavipes (Pers.) P. Kumm.
Agaricales,
Hygrophoraceae
East Azerbaijan,
Golestan,
Mazandaran
Buhse 1860, Saber 2002a

6 Mycologia Iranica - Vol. 7, 2020
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
Armillaria borealis Marxm. &
Korhonen
Agaricales,
Physalacriaceae
Mazandaran
Asef & Mohammadi Goltapeh
2002, Dalili et al. 2008
Armillaria cepistipes Velen.
Agaricales,
Physalacriaceae
East Azerbaijan
Asef & Mohammadi Goltapeh
2002
●Armillaria gallica Marxm. &
Romagn.
Agaricales,
Physalacriaceae
Alborz, East
Azerbaijan,
Golestan,
Mazandaran
Asef et al. 2003, Dalili et al. 2008,
Keča et al. 2014
KJ200954 (tef1),
KJ200948 (LSU-
IGC)
●Armillaria mellea (Vahl) P. Kumm.
Armillariella mellea (Vahl) P. Karst.,
Clitocybe mellea (Vahl) Ricken
Agaricales,
Physalacriaceae
Alborz, East
Azerbaijan, Gilan,
Golestan, Isfahan,
Khuzestan,
Markazi,
Mazandaran,
Qazvin, Razavi
Khorasan,
Semnan, West
Azerbaijan
Sharif & Ershad 1966, Khabiri
1968, Saber 1973, Soleimani 1974
and 1976, Behboudi & Sabahi
1983, Jafarpour et al. 1983,
Mohammadipour 2000, Razavi et
al. 2000, Razzaz-Hashemi &
Zakeri 2000, Zokaei 2001, Asef &
Mohammadi Goltapeh 2002, Asef
& Tavanaei 2004, Coetzee et al.
2005, Amirahmadi et al 2006,
Asef 2007b, Taheri et al. 2008,
Asef & Etemad 2016
AF163583,
MG835859
!Arrhenia griseopallida (Desm.)
Watling
Omphalina griseopallida (Desm.) Quél.
Agaricales,
Hygrophoraceae
Golestan
Saber 2002a
Asterophora lycoperdoides (Bull.)
Ditmar
Agaricales,
Lyophyllaceae
Mazandaran
Mohammadi Goltapeh 2000a
!Atheniella flavoalba (Fr.) Redhead,
Moncalvo, Vilgalys, Desjardin & B.A.
Perry
Mycena flavoalba (Fr.) Quél.
Agaricales,
Mycenaceae
Gilan
Saber 1995h
!Aureoboletus gentilis (Quél.) Pouzar
Boletales,
Boletaceae
Mazandaran
Bahram & Asef 2008a
!Baeospora myosura (Fr.) Singer
Agaricales,
Marasmiaceae
Mazandaran
Saber 1998
Battarrea stevenii (Libosch.) Fr.
Agaricales,
Agaricaceae
Golestan, Tehran
Saber 1986, Saber 2002b
Bolbitius reticulatus (Pers.) Ricken
Bolbitius aleuriatus (Fr.) Singer
Agaricales,
Bolbitiaceae
Golestan,
Mazandaran
Saber 1993f
!Bolbitius titubans (Bull.) Fr.
Agaricales,
Bolbitiaceae
Golestan
Saber 1993f
!Boletus aestivalis (Paulet) Fr.
Boletales,
Boletaceae
Golestan
Saber 1995d
Boletus edulis Bull.
Boletales,
Boletaceae
East Azerbaijan
Asef 2007b, 2008
!Butyriboletus appendiculatus
(Schaeff.) D. Arora & J. L. Frank,
Boletus appendiculatus Schaeff.
Boletales,
Boletaceae
Tehran
Zangeneh & Maivan 1998
Butyriboletus fechtneri (Velen.) D.
Arora & J.L. Frank
Boletus fechtneri Velen.
Boletales,
Boletaceae
East Azerbaijan
Asef 2007b, 2008
●Butyriboletus pseudoregius (Heinr.
Huber) D. Arora & J.L. Frank
Boletales,
Boletaceae
‘Hyrcan’
Bahram et al. 2012
FR852279 (env.,
SH1933335.08FU)
!Calocybe carnea (Bull.) Donk
Agaricales,
Lyophyllaceae
Tehran
Saber & Zangeneh 2004
!Calocybe chrysenteron (Bull.) Singer
Calocybe cerina (Pers.) Donk
Agaricales,
Lyophyllaceae
Gilan
Saber 2000d
!Calocybe gambosa (Fr.) Donk
Agaricales,
Lyophyllaceae
Tehran
Saber 2000d
!Calocybe ionides (Bull.) Donk
Agaricales,
Lyophyllaceae
Gilan
Saber 2000d
!Calocybe persicolor (Fr.) Singer
Agaricales,
Lyophyllaceae
Mazandaran
Saber 2000d
Cantharellus alborufescens
(Malençon) Papetti & S. Alberti
Cantharellales,
Hydnaceae
Golestan,
Mazandaran
Parad et al. 2018
Cantharellus cibarius Fr.
Cantharellales,
Hydnaceae
Golestan
Ershad 1977, Saber 1987
Cantharellus ferruginascens P.D.
Orton
Cantharellales,
Hydnaceae
Gilan, Golestan,
Mazandaran
Parad et al. 2018
Chalciporus piperatus (Bull.) Bataille
Boletus piperatus Bull.
Boletales,
Boletaceae
Golestan
Saber 1995d

GHOBAD-NEJHAD ET AL.: The gilled fungi and boletes of Iran 7
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
!Chlorophyllum brunneum (Farl. &
Burt) Vellinga
Lepiota brunnea Farl. & Burt
Agaricales,
Agaricaceae
?
Saber 1994a
Chlorophyllum rhacodes (Vittad.)
Vellinga
Macrolepiota rhacodes (Vittad.)
Singer
Agaricales,
Agaricaceae
East Azerbaijan
Saber 1994a, Mohammadi
Goltapeh 2001
Clitocybe angustissima (Lasch) P.
Kumm.
Agaricales,
incertae sedis
Golestan
Saber & Zangeneh 2004
!Clitocybe barbularum (Romagn.)
P.D. Orton
Agaricales,
incertae sedis
Gilan
Saber 2002a
!Clitocybe diatreta (Fr.) P. Kumm.
Agaricales,
incertae sedis
West Azerbaijan
Saber & Mehravaran 2004
!Clitocybe metachroa (Fr.) P. Kumm.
Clitocybe dicolor (Pers.) Murrill
Agaricales,
incertae sedis
Mazandaran
Saber 2002a
!Clitocybe nebularis (Batsch) P.
Kumm.
Agaricales,
incertae sedis
Gilan, East Azerbaijan,
Mazandaran
Saber 2002a, Asef 2007b
!Clitocybe phyllophila (Pers.) P.
Kumm.
Agaricales,
incertae sedis
Mazandaran
Saber 2002a
Clitocybe rufuloalutacea Métrod ex
Bon
Agaricales,
incertae sedis
East Azerbaijan, Razavi
Khorasan
Asef 2007b and 2009b
!Clitocybe vibecina (Fr.) Quél.
Clitocybe langei Singer ex Hora,
Tephrocybe langei (Singer) Raithelh.
Agaricales,
incertae sedis
Gilan
Saber & Zangeneh 2004
!Clitopaxillus alexandri (Gillet) G.
Moreno, Vizzini, Consiglio & P.
Alvarado
Clitocybe alexandri (Gillet) Gillet
Agaricales,
incertae sedis
Gilan
Saber 2002a
Clitopilus prunulus (Scop.) P.
Kumm.
Clitopilus orcellus (Bull. ex Pers.) P.
Kumm.
Agaricales,
Entolomataceae
Gilan, Golestan,
Mazandaran
Saber 1993d and 1997a
Clitopilus scyphoides (Fr.) Singer
Clitopilus cretatus (Berk. & Broome)
Quél., Clitopilus intermedius
Romagn., Clitopilus scyphoides var.
intermedius (Romagn.) Noordel.
Agaricales,
Entolomataceae
Golestan
Saber 1995e and 1997a
!Collybia tuberosa (Bull.) P. Kumm.
Agaricales,
incertae sedis
Gilan
Saber 1998
Conocybe albipes (G.H. Otth)
Hauskn.
Conocybe lactea (J.E. Lange) Métrod
Agaricales,
Bolbitiaceae
Alborz, Golestan,
Kermanshah, Razavi
Khorasan, Tehran
Saber 1993f, Zokaei 2001,
Fadavi et al. 2013
*●Conocybe apala (Fr.) Arnolds
Agaricales,
Bolbitiaceae
Tehran
this study
MT535728
Conocybe dunensis T.J. Wallace
Agaricales,
Bolbitiaceae
Kermanshah
Fadavi et al. 2013
Conocybe juniana (Velen.) Hauskn.
& Svrček
Conocybe magnicapitata P.D. Orton
Agaricales,
Bolbitiaceae
Golestan
Watling & Gregory 1977
Conocybe leucopus Kühner ex
Kühner & Watling
Agaricales,
Bolbitiaceae
Hamadan
Jadidian et al. 2015
!Conocybe macrocephala Kühner &
Watling
Agaricales,
Bolbitiaceae
Mazandaran
Saber 1993f
!Conocybe microspora (Velen.)
Dennis
Conocybe brunneola Kühner ex
Kühner & Watling
Agaricales,
Bolbitiaceae
Mazandaran
Saber 1995f
!Conocybe ochracea Kühner ex
Singer
Agaricales,
Bolbitiaceae
Golestan, Tehran
Saber 1993f
!Conocybe pilosella (Pers.) Kühner
Agaricales,
Bolbitiaceae
Mazandaran
Saber 1995f
!Conocybe rickenii (Jul. Schäff.)
Kühner
Agaricales,
Bolbitiaceae
Gilan, Tehran
Saber 1993f
!Conocybe subovalis Kühner &
Watling
Agaricales,
Bolbitiaceae
Alborz
Saber 1993f
!Conocybe tenera (Schaeff.) Fayod
Agaricales,
Bolbitiaceae
Sistan and Baluchestan,
Tehran
Saber 1993f, Saber &
Esmaeili Taheri 2004

8 Mycologia Iranica - Vol. 7, 2020
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
!Contumyces rosellus (M.M.
Moser) Redhead, Moncalvo,
Vilgalys & Lutzoni
Marasmiellus rosellus (M.M.
Moser) Kuyper & Noordel.,
Omphalina rosella (M.M. Moser)
M.M. Moser ex Redhead,
Ammirati & Norvell
Hymeochaetales,
Rickenellaceae
Mazandaran
Saber 2002a
Coprinellus angulatus (Peck)
Redhead
Coprinus angulatus Peck
Agaricales,
Psathyrellaceae
Fars, Tehran
Mohammadi Goltapeh 2003,
Saber & Esmaeili Taheri 2004
●Coprinellus disseminatus
(Pers.) J.E. Lange
Coprinus disseminatus (Pers.)
Gray
Agaricales,
Psathyrellaceae
Alborz, Isfahan,
Mazandaran, Razavi
Khorasan, Sistan and
Baluchestan, Tehran
Saber 1994b, Mohammadi
Goltapeh 2000b and 2001, Zokaei
2001, Saber & Esmaeili Taheri
2002 and 2004, Asef & Etemad
2016, this study
MK050584
!Coprinellus domesticus (Bolton)
Vilgalys, Hopple & Jacq. Johnson
Coprinus domesticus (Bolton)
Gray
Agaricales,
Psathyrellaceae
?
Saber 1994b
Coprinellus flocculosus (DC.)
Vilgalys, Hopple & Jacq. Johnson
Coprinus flocculosus (DC.) Fr.
Agaricales,
Psathyrellaceae
Tehran
Saber 1994b, Mohammadi
Goltapeh 2000b and 2001
!Coprinellus impatiens (Fr.) J.E.
Lange
Coprinus impatiens (Fr.) Quél.
Agaricales,
Psathyrellaceae
?
Saber 1994b
●Coprinellus micaceus (Bull.)
Vilgalys, Hopple & Jacq. Johnson
Coprinus micaceus (Bull.) Fr.
Agaricales,
Psathyrellaceae
Alborz, Ilam, Isfahan,
Kohgiluyeh and Boyer-
Ahmad, Mazandaran,
Razavi Khorasan,
Tehran, West
Azerbaijan
Petrak 1940, Petrak & Esfandiari
1941, Esfandiari 1946, Khabiri
1968, Soleimani 1974 and 1976,
Zokaei 2001, Saber & Esmaeili
Taheri 2002, Saber & Mehravaran
2004, Asef & Etemad 2016, this
study
MT535694
●Coprinellus radians (Desm.)
Vilgalys, Hopple & Jacq. Johnson
Agaricales,
Psathyrellaceae
Mazandaran
Ziaie Juybari et al. 2019
KU375662 (as
endophyte)
!Coprinellus silvaticus (Peck)
Gminder
Coprinellus tardus (P. Karst.) P.
Karst., Coprinus silvaticus Peck
Agaricales,
Psathyrellaceae
?
Saber 1994b
Coprinellus subimpatiens (M.
Lange & A.H. Sm.) Redhead,
Vilgalys & Moncalvo
Coprinus subimpatiens M. Lange
& A.H. Sm.
Agaricales,
Psathyrellaceae
Alborz
Mohammadi Goltapeh 2003
!Coprinellus truncorum (Scop.)
Redhead, Vilgalys & Moncalvo
Agaricales,
Psathyrellaceae
East Azerbaijan
Asef 2007b
!Coprinellus xanthothrix
(Romagn.) Vilgalys, Hopple &
Jacq. Johnson
Coprinus xanthothrix Romagn.
Agaricales,
Psathyrellaceae
?
Saber 1994b
●Coprinopsis atramentaria
(Bull.) Redhead, Vilgalys &
Moncalvo
Coprinus atramentarius (Bull.)
Fr.
Agaricales,
Psathyrellaceae
Alborz, Gilan, Isfahan,
Kermanshah,
Mazandaran, Razavi
Khorasan, Tehran, West
Azerbaijan
Khabiri 1968, Soleimani 1974 and
1976, Saber 1994b, Zokaei 2001,
Saber & Esmaeili Taheri 2002,
2004, Saber & Mehravaran 2004,
GenBank direct submission, this
study
MH259864
Coprinopsis brunneofibrillosa
(Dennis) Redhead, Vilgalys &
Moncalvo
Coprinus brunneofibrillosus
Dennis
Agaricales,
Psathyrellaceae
Mazandaran, Tehran
Mohammadi Goltapeh 2000b and
2003
●Coprinopsis cinerea (Schaeff.)
Redhead, Vilgalys & Moncalvo
Coprinus cinereus (Schaeff.)
Gray, Coprinus fimetarius sensu
auct. brit.
Agaricales,
Psathyrellaceae
Alborz, Mazandaran,
Tehran
Khabiri 1968, Mohammadi
Goltapeh 2003, GenBank direct
submission
MF161131 (as
endophyte)
Coprinopsis ephemeroides (DC.)
G. Moreno
Coprinus ephemeroides (DC.) Fr.
Agaricales,
Psathyrellaceae
Alborz
Mohammadi Goltapeh 2003
!Coprinopsis friesii (Quél.) P.
Karst.
Coprinus friesii Quél.
Agaricales,
Psathyrellaceae
?
Saber 1994b

GHOBAD-NEJHAD ET AL.: The gilled fungi and boletes of Iran 9
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
!Coprinopsis gonophylla (Quél.)
Redhead, Vilgalys & Moncalvo
Coprinus gonophyllus Quél.
Agaricales,
Psathyrellaceae
?
Saber 1994b
Coprinopsis lagopides (P. Karst.)
Redhead, Vilgalys & Moncalvo
Coprinus lagopides P. Karst.
Agaricales,
Psathyrellaceae
Tehran
Mohammadi Goltapeh 2003
Coprinopsis lagopus (Fr.) Redhead,
Vilgalys & Moncalvo
Coprinus lagopus (Fr.) Fr.
Agaricales,
Psathyrellaceae
Alborz, Mazandaran
Saber 1994b, Mohammadi
Goltapeh 2003
Coprinopsis macrocephala (Berk.)
Redhead, Vilgalys & Moncalvo
Coprinus macrocephalus (Berk.)
Berk.
Agaricales,
Psathyrellaceae
Tehran
Mohammadi Goltapeh 2003
!Coprinopsis marcescibilis
(Britzelm.) Örstadius & E. Larss.
Psathyrella marcescibilis
(Britzelm.) Singer
Agaricales,
Psathyrellaceae
?
Saber 1994b
!Coprinopsis martinii (P.D. Orton)
Redhead, Vilgalys & Moncalvo
Coprinus martinii P.D. Orton
Agaricales,
Psathyrellaceae
Isfahan
Saber & Esmaeili Taheri 2004
Coprinopsis nivea (Pers.) Redhead,
Vilgalys & Moncalvo
Coprinus niveus (Pers.) Fr.
Agaricales,
Psathyrellaceae
Alborz, East Azerbaijan
Saber 1994a, Mohammadi
Goltapeh 2003, Asef 2007b
Coprinopsis patouillardii (Quél.)
Gminder
Coprinus patouillardii Quél.
Agaricales,
Psathyrellaceae
Tehran
Mohammadi Goltapeh 2003
Coprinopsis picacea (Bull.)
Redhead, Vilgalys & Moncalvo
Coprinus picaceus (Bull.) Gray
Agaricales,
Psathyrellaceae
Alborz, East Azerbaijan,
Lorestan, Razavi
Khorasan
Saber 1994b, Zokaei 2001,
Mohammadi Goltapeh 2003,
Saber & Esmaeili Taheri 2004
●Coprinopsis sclerotiger (Watling)
Redhead, Vilgalys & Moncalvo
Agaricales,
Psathyrellaceae
?
GenBank direct submission
MF161132 (as
endophyte)
Coprinopsis scobicola (P.D. Orton)
Redhead, Vilgalys & Moncalvo
Coprinus scobicola P.D. Orton
Agaricales,
Psathyrellaceae
Tehran
Mohammadi Goltapeh 2001
●Coprinopsis urticicola (Berk. &
Broome) Redhead, Vilgalys &
Moncalvo
Agaricales,
Psathyrellaceae
Kermanshah,
Mazandaran
Asef et al. 2015, Gholami et al.
2019
MH300615 (as
endophyte)
Coprinus comatus (O.F. Müll.)
Pers.
Agaricales,
Agaricaceae
Kermanshah,
Mazandaran, Razavi
Khorasan, Tehran
Khabiri 1968, Fallahyan 1973,
Mohammadi Goltapeh 2000b and
2001, Zokaei 2001, Fadavi et al.
2015, Asef & Etemad 2016, this
study
Coprinus sterquilinus (Fr.) Fr.
Agaricales,
Agaricaceae
Kermanshah
Saber & Dadkhahipour 2000,
Fadavi et al. 2015
●Cortinarius bivelus (Fr.) Fr.
Agaricales,
Cortinariaceae
‘Hyrcan’
Bahram et al. 2012
FR852016 (env.,
SH2224383.08FU)
●Cortinarius caesiocortinatus Jul.
Schäff.
Agaricales,
Cortinariaceae
‘Hyrcan’
Bahram et al. 2012
FR852020 (env.,
SH1504045.08FU)
●Cortinarius casimirii (Velen.)
Huijsman
Agaricales,
Cortinariaceae
‘Hyrcan’
Bahram et al. 2012
FR851999 (env.,
SH1917131.08FU)
Cortinarius causticus Fr.
Agaricales,
Cortinariaceae
East Azerbaijan
Asef 2007a, b
●Cortinarius cinnabarinus Fr.
Agaricales,
Cortinariaceae
‘Hyrcan’
Bahram et al. 2012
FR852005 (env.,
SH1710033.08FU)
!Cortinarius cinnamomeus (L.)
Gray
Agaricales,
Cortinariaceae
Gilan
Saber 1994c, Saber 2000c
●Cortinarius cotoneus Fr.
Agaricales,
Cortinariaceae
‘Hyrcan’
Bahram et al. 2012
FR852015 (env.,
SH1855917.08FU)
●Cortinarius decipiens Fr.
Agaricales,
Cortinariaceae
Mazandaran
Saber & Zangeneh 2004, Bahram
et al. 2013
HE687043 (env.,
SH1917130.08FU)
●Cortinarius diasemospermus
Lamoure
Agaricales,
Cortinariaceae
Mazandaran?
Bahram et al. 2013
HE687042 (env.,
SH2224452.08FU
Cortinarius erumpens Rob. Henry
Agaricales,
Cortinariaceae
East Azerbaijan
Asef 2007a and 2007b
●Cortinarius ferrugineovelatus
Kytöv., Liimat. & Niskanen
Agaricales,
Cortinariaceae
‘Hyrcan’
Bahram et al. 2012
FR852002 (env.,
SH2224500.08FU)
Cortinarius fluryi (M.M. Moser)
M.M. Moser
Agaricales,
Cortinariaceae
East Azerbaijan
Asef 2007b and 2009a
*●Cortinarius hildegardiae
Schmidt-Stohn, Brandrud & Dima
Agaricales,
Cortinariaceae
Golestan
this study
MT535704

10 Mycologia Iranica - Vol. 7, 2020
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
●Cortinarius hinnuleus Fr.
Agaricales,
Cortinariaceae
‘Hyrcan’
Bahram et al. 2012
FR851998 (env.,
SH1917246.08FU)
●Cortinarius infractus (Pers.) Fr.
Agaricales,
Cortinariaceae
‘Hyrcan’
Bahram et al. 2012
FR852022 (env.,
SH1855756.08FU)
●Cortinarius olivaceofuscus
Kühner
Agaricales,
Cortinariaceae
‘Hyrcan’
Bahram et al. 2012
FR851993 (env.,
SH1855819.08FU)
Cortinarius paracephalixus
Bohus
Agaricales,
Cortinariaceae
East Azerbaijan
Asef 2007b and 2009a
●Cortinarius parvannulatus
Kühner
Agaricales,
Cortinariaceae
Mazandaran?
Bahram et al. 2013
HE687041 (env.,
SH1709965.08FU
*●Cortinarius persoonianus
Bidaud
Agaricales,
Cortinariaceae
Golestan
this study
MT535741
Cortinarius pluviorum Jul.
Schäff. ex M.M. Moser
Agaricales,
Cortinariaceae
East Azerbaijan
Asef 2007a and 2007b
●Cortinarius uraceonemoralis
Niskanen, Liimat., Dima, Kytöv.,
Bojantchev & H. Lindstr.
Agaricales,
Cortinariaceae
‘Hyrcan’
Bahram et al. 2012
FR852021 (env.,
SH1917211.08FU)
!Cortinarius valgus Fr.
Agaricales,
Cortinariaceae
East Azerbaijan
Asef 2007b
●Cortinarius vernus H. Lindstr.
& Melot
Agaricales,
Cortinariaceae
Mazandaran?
Bahram et al. 2013
HE687034 (env.,
SH1917268.08FU)
Cortinarius vespertinus (Fr.) Fr.
Agaricales,
Cortinariaceae
East Azerbaijan
Asef 2007b and 2009a
Cortinarius vibratilis (Fr.) Fr.
Agaricales,
Cortinariaceae
Mazandaran
Bahram & Asef 2008a, Asef &
Etemad 2016
Cortinarius violaceus (L.) Gray
Agaricales,
Cortinariaceae
Mazandaran
Khabiri 1968, Saber 1994c
Craterellus cinereus (Pers.: Fr.)
Maire
Cantharellus cinereus Pers.
Cantharellales,
Hydnaceae
Gilan, Mazandaran
Saber 1989b
Craterellus cornucopioides (L.)
Pers.
Pleurotus cornucopioides (L.)
Gillet
Cantharellales,
Hydnaceae
Gilan, Mazandaran
Khabiri 1968, Saber 1987
Craterellus tubaeformis (Fr.)
Quél.
Cantharellales,
Hydnaceae
'forests of northern Iran'
Fallahyan 1973
Crepidotus applanatus (Pers.) P.
Kumm.
Agaricales,
Inocybaceae
Gilan, Golestan,
Mazandaran
Saber 1990, Asef & Etemad
2016
Crepidotus caspari Velen.
Crepidotus amygdalosporus
Kühner, Crepidotus inhonestus P.
Karst. [sensu Orton, Pegler &
Young, and Nordstein (non s. P.
Karst.)], Crepidotus lundellii Pilát
Agaricales,
Inocybaceae
Gilan, Golestan,
Mazandaran
Saber 1990, Saber 1997a, Saber
& Zangeneh 2002, Saber &
Esmaeili Taheri 2004, Asef &
Etemad 2016
Crepidotus cesatii (Rabenh.)
Sacc.
Agaricales,
Inocybaceae
East Azerbaijan,
Mazandaran
Watling & Sweeney 1974, Asef
2007b
Crepidotus crocophyllus (Berk.)
Sacc.
Agaricales,
Inocybaceae
Golestan, Mazandaran
Saber 1997a, Asef & Etemad
2016
Crepidotus mollis (Schaeff.)
Staude
Agaricales,
Inocybaceae
Gilan, Golestan,
Mazandaran
Saber 1990
*●Crepidotus subverrucisporus
Pilát
Agaricales,
Inocybaceae
East Azerbaijan
this study
MT535745
!Crinipellis scabella (Alb. &
Schwein.) Murrill
Crinipellis stipitaria (Fr.) Pat.
Agaricales,
Marasmiaceae
Golestan
Saber 1995g
*●Cuphophyllus virgineus
(Wulfen) Kovalenko Hygrocybe
virginea (Wulfen) P.D. Orton &
Watling
Agaricales,
Hygrophoraceae
Mazandaran
this study
MT535688
Cyanoboletus pulverulentus
(Opat.) Gelardi, Vizzini &
Simonini
Boletus pulverulentus Opat.
Boletales,
Boletaceae
East Azerbaijan, Gilan
Saber 1995d, Asef 2007b and
2008
●Cyclocybe cylindracea (DC.)
Vizzini & Angelini
Agrocybe cylindracea (DC.)
Maire, Pholiota aegerita (V.
Brig.) Quél.
Agaricales,
Tubariaceae
Gilan, Golestan,
Mazandaran, Razavi
Khorasan
Khabiri 1968, Soleimani 1976,
Saber 1993f, Zokaei 2001, Asef
& Etemad 2016, GenBank direct
submission
MT573399

GHOBAD-NEJHAD ET AL.: The gilled fungi and boletes of Iran 11
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
Cystoderma aureum (Matt.)
Kühner & Romagn.
Phaeolepiota aurea (Matt.) Maire
ex Konrad & Maubl.
Agaricales, incertae
sedis
Golestan,
Mazandaran
Saber 1994a, Asef & Etemad
2016
●Deconica crobula (Fr.) Romagn.
Psilocybe crobula (Fr.) Singer
Agaricales,
Strophariaceae
East Azerbaijan,
Razavi Khorasan
Saber 1991a, Zokaei 2001, this
study
MT535747
!Delicatula integrella (Pers.) Fayod
Agaricales, incertae
sedis
Mazandaran
Saber 1995h
!Dermoloma cuneifolium (Fr.)
Singer ex Bon
Dermoloma atrocinereum (Pers.)
Herink, Tricholoma atrocinereum
(Pers.) Quél.
Agaricales, incertae
sedis
Golestan
Saber 2000d
Echinoderma asperum (Pers.) Bon
Amanita aspera (Pers.) Pers.,
Lepiota aspera (Pers.) Quél.,
Lepiota acutesquamosa (Weinm.)
P. Kumm.
Agaricales,
Agaricaceae
Golestan,
Mazandaran
Saber 1994a, Saber & Zangeneh
2002, Bahram et al. 2006b, Asef
& Etemad 2016
!Entoloma clypeatum (L.) P.
Kumm.
Agaricales,
Entolomataceae
Mazandaran
Saber 1993d
Entoloma griseoluridum (Kühner)
M.M. Moser
Agaricales,
Entolomataceae
East Azerbaijan
Asef 2007b, Asef & Muradov
2014
Entoloma griseorubellum (Lasch)
Kalamees & Urbonas
Agaricales,
Entolomataceae
Kermanshah
Fadavi et al. 2013
!Entoloma hirtipes (Schumach.)
M.M. Moser
Agaricales,
Entolomataceae
Gilan
Saber 2000c
Entoloma incanum (Fr.) Hesler
Agaricales,
Entolomataceae
Golestan, Razavi
Khorasan
Zokaei 2001, Saber & Zangeneh
2002
!Entoloma majaloides P.D. Orton
Agaricales,
Entolomataceae
Golestan
Saber & Zangeneh 2002
!Entoloma mammosum (L.) Hesler
Agaricales,
Entolomataceae
Golestan
Saber 1993d
Entoloma niphoides Noordel.
Agaricales,
Entolomataceae
East Azerbaijan
Asef & Muradov 2014
!Entoloma rhodopolium (Fr.) P.
Kumm.
Agaricales,
Entolomataceae
Gilan, Mazandaran
Saber 1993d
!Entoloma sericellum (Fr.) P.
Kumm.
Agaricales,
Entolomataceae
Gilan
Saber 1993d
Entoloma sinuatum (Bull. ex
Pers.) P. Kumm.
Agaricales,
Entolomataceae
Golestan,
Mazandaran
Saber 1995f, Asef & Etemad
2016
●Entoloma subcollariatum
(Kühner) Bon
Agaricales,
Entolomataceae
?
GenBank direct submission
MH453494
!Entoloma vernum S. Lundell
Agaricales,
Entolomataceae
Ardabil
Saber & Zangeneh 2002
!Flammula alnicola (Fr.) P.
Kumm.
Pholiota alnicola (Fr.) Singer
Agaricales,
Hymenogastraceae
?
Saber 1991b
!Flammulaster erinaceellus (Peck)
Watling
Agaricales,
Inocybaceae
Mazandaran
Saber 1993a
Flammulaster ferrugineus (Maire)
Watling
Agaricales,
Inocybaceae
Golestan
Watling & Gregory 1977
!Flammulaster gracilis (Quél.)
Watling
Agaricales,
Inocybaceae
Gilan, Mazandaran
Saber 1993a
!Flammulaster granulosus (J.E.
Lange) Watling
Agaricales,
Inocybaceae
Tehran
Saber 1995f
●Flammulina velutipes (Curtis)
Singer
Collybia velutipes (Curtis) P.
Kumm., Collybia velutipes var.
lactea (Quél.) Rea, Flammulina
velutipes var. lactea (Quél.) Bas
Agaricales,
Physalacriaceae
Alborz, East
Azerbaijan, Gilan,
Isfahan,
Mazandaran, Razavi
Khorasan, Tehran
Soleimani 1974 and 1976, Saber
1998, Zokaei 2001, Asef 2007b,
Asef & Etemad 2016, this study
MT535715
!Galerina hypnorum (Schrank)
Kühner
Agaricales,
Hymenogastraceae
?
Saber 1994c
Galerina marginata (Batsch)
Kühner
Galerina unicolor (Vahl) Singer
Agaricales,
Hymenogastraceae
Golestan,
Mazandaran
Watling & Sweeney 1974, Saber
1994c
!Galerina mniophila (Lasch)
Kühner
Agaricales,
Hymenogastraceae
?
Saber 1994c

12 Mycologia Iranica - Vol. 7, 2020
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
!Galerina pumila (Pers.) M. Lange
Galerina mycenopsis (Fr.) Kühner
Agaricales,
Hymenogastraceae
?
Saber 1994c
!Galerina sphagnorum (Pers.)
Kühner
Agaricales,
Hymenogastraceae
?
Saber 1994c
!Gymnopilus penetrans (Fr.)
Murrill
Agaricales,
Strophariaceae
?
Saber 1994c
●Gymnopilus spectabilis (Weinm.)
A.H. Sm.
Gymnopilus junonius (Fr.) P.D.
Orton
Agaricales,
Strophariaceae
Golestan
Saber 1994c, Vahidi et al.
2006, this study
MT535703
!Gymnopus androsaceus (L.) J.L.
Mata & R.H. Petersen
Marasmius androsaceus (L.) Fr.
Agaricales,
Omphalotaceae
Gilan, Golestan,
Mazandaran
Saber 1995g
*●Gymnopus aquosus (Bull.)
Antonín & Noordel.
Agaricales,
Omphalotaceae
Mazandaran
this study
MT535700,
MT535699
!Gymnopus brassicolens
(Romagn.) Antonín & Noordel.
Micromphale brassicolens
(Romagn.) P.D. Orton
Agaricales,
Omphalotaceae
Gilan
Saber 1995g
Gymnopus dryophilus (Bull.)
Murrill
Collybia dryophila (Bull.) P.
Kumm.
Agaricales,
Omphalotaceae
Gilan, Golestan,
Mazandaran
Buhse 1860, Saber 1998
!Gymnopus erythropus (Pers.)
Antonín, Halling & Noordel.
Collybia acervata sensu auct.,
Collybia erythropus (Pers.) P.
Kumm.
Agaricales,
Omphalotaceae
East Azerbaijan, Gilan,
Mazandaran, Tehran
Saber 1998, Saber &
Zangeneh 2004, Asef
2007b
!Gymnopus foetidus (Sowerby)
J.L. Mata & R.H. Petersen
Micromphale foetidum (Sowerby)
Singer
Agaricales,
Omphalotaceae
East Azerbaijan, Gilan,
Mazandaran
Saber 1995g, Asef 2007b
Gymnopus fusipes (Bull.) Gray
Collybia fusipes (Bull.) Quél.
Agaricales,
Omphalotaceae
Alborz, Gilan
Saber 1998, Soleimani
1974 and 1976
*●Gymnopus hybridus (Kühner &
Romagn.) Antonín & Noordel.
Agaricales,
Omphalotaceae
Ilam
this study
MT535705
!Gymnopus inodorus (Pat.)
Antonín & Noordel.
Micromphale inodorum (Pat.)
Svrček
Agaricales,
Omphalotaceae
Gilan, Golestan,
Mazandaran
Saber 1995g
!Gymnopus terginus (Fr.) Antonín
& Noordel.
Marasmius terginus (Fr.) Fr.
Agaricales,
Omphalotaceae
Gilan, Mazandaran
Saber 1995g
*●Gyroporus castaneus (Bull.)
Quél.
Boletales,
Gyroporaceae
Golestan
this study
MT535725
!Hebeloma birrus (Fr.) Gillet
Agaricales,
Hymenogastraceae
?
Saber 1994c
!Hebeloma crustuliniforme (Bull.)
Quél.
Agaricales,
Hymenogastraceae
?
Saber 1994c
!Hebeloma hiemale Bres.
Hebeloma oculatum Bruchet
Agaricales,
Hymenogastraceae
East Azerbaijan
Asef 2007b
●Hebeloma incarnatulum A.H. Sm.
Agaricales,
Hymenogastraceae
‘Hyrcan’
Bahram et al. 2012
FR852303 (env.,
SH2290416.08FU)
!Hebeloma mesophaeum (Pers.)
Quél.
Agaricales,
Hymenogastraceae
?
Saber 1994c
●Hebeloma sinapizans (Paulet)
Gillet
Agaricales,
Hymenogastraceae
?
Saber 1994c
KX266254
(original
submission named
wrongly)
!Hemileccinum impolitum (Fr.)
Šutara
Boletus impolitus Fr., Xerocomus
impolitus (Fr.) Quél.
Boletales,
Boletaceae
Kohgiluyeh and Boyer-
Ahmad
Saber 1995d
!Hemimycena cucullata (Pers.)
Singer
Agaricales, incertae
sedis
Mazandaran
Saber 1995h
!Hodophilus hymenocephalus
(A.H. Sm. & Hesler) Birkebak &
Adamčík
Camarophyllopsis hymenocephala
(A.H. Sm. & Hesler) Arnolds
Agaricales,
Clavariaceae
East Azerbaijan
Asef 2007b

GHOBAD-NEJHAD ET AL.: The gilled fungi and boletes of Iran 13
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
Hohenbuehelia atrocoerulea (Fr.)
Singer
Agaricales, Pleurotaceae
Gilan
Saber 1990
Hohenbuehelia auriscalpium
(Maire) Singer
Agaricales, Pleurotaceae
Golestan, Mazandaran
Shahtahmasebi et al.
2018
Hohenbuehelia petaloides (Bull.)
Schulzer
Hohenbuehelia geogenia (DC. ex
Fr.) Singer
Agaricales, Pleurotaceae
Gilan, Mazandaran
Saber 1990
!Homophron spadiceum (P.
Kumm.) Örstadius & E. Larss.
Psathyrella sarcocephala (Fr.)
Singer
Agaricales,
Psathyrellaceae
?
Saber 1994b
!Hydropus marginellus (Pers.)
Singer
Agaricales,
Marasmiaceae
Mazandaran
Saber 2002a
!Hygrocybe acutoconica (Clem.)
Singer
Hygrocybe persistens (Britzelm.)
Singer
Agaricales,
Hygrophoraceae
Mazandaran
Saber 1995f
!Hygrocybe chlorophana (Fr.)
Wünsche
Agaricales,
Hygrophoraceae
Mazandaran
Saber 1993d
Hygrophoropsis aurantiaca
(Wulfen) Maire
Cantharellus aurantiacus (Wulfen)
Fr.
Boletales,
Hygrophoropsidaceae
Gilan, Mazandaran
Khabiri 1968, Saber
1993d
Hygrophorus eburneus (Bull.) Fr.
Agaricales,
Hygrophoraceae
Gilan
Saber 1995f
!Hygrophorus mesotephrus Berk. &
Broome
Agaricales,
Hygrophoraceae
East Azerbaijan
Asef 2007b
!Hygrophorus persoonii Arnolds
Hygrophorus dichrous Kühner &
Romagnesi (invalid name)
Agaricales,
Hygrophoraceae
Gilan, Mazandaran
Saber 2000c
●Hymenopellis radicata (Relhan)
R.H. Petersen
Oudemansiella radicata (Relhan)
Singer, Xerula radicata (Relhan)
Dörfelt
Agaricales,
Physalacriaceae
East Azerbaijan, Gilan,
Golestan, Mazandaran
Saber 1998, Asef &
Etemad 2016, this study
MT535683
!Hypholoma capnoides (Fr.) P.
Kumm.
Agaricales,
Strophariaceae
Mazandaran
Saber & Zangeneh 2004
●Hypholoma fasciculare (Huds.) P.
Kumm.
Hypholoma elaeodes (Fr.) Gillet
Agaricales,
Strophariaceae
East Azerbaijan, Gilan,
Golestan, Mazandaran,
Razavi Khorasan
Buhse 1860, Watling &
Gregory 1977, Zokaei
2001, Saber & Zangeneh
2002, Asef & Tavanaei
2004, Asef 2007b, Asef
& Etemad 2016, Bari et
al. 2019, Ziaie Juybari et
al. 2019, this study
MT535706,
KX169250 (as
endophyte)
Hypholoma lateritium (Schaeff.) P.
Kumm.
Hypholoma sublateritium (Fr.) Quél.
Agaricales,
Strophariaceae
Mazandaran
Asef & Etemad 2016
!Hypholoma radicosum J.E. Lange
Hypholoma epixanthum (Fr.) Quél.
Agaricales,
Strophariaceae
East Azerbaijan, Gilan
Saber 1993a, Asef 2007b
!Hypholoma subericaeum (Fr.)
Kühner
Agaricales,
Strophariaceae
Gilan
Saber 1993a
Hypsizygus ulmarius (Bull.)
Redhead
Agaricales,
Lyophyllaceae
Golestan
Saber 1995e, Saber 1997a
●Imleria badia (Fr.) Vizzini
Boletus badius (Fr.) Fr.
Boletales, Boletaceae
‘Hyrcan’
Bahram et al. 2012
FR852283 (env.,
SH2143392.08FU)
Infundibulicybe geotropa (Bull. ex
DC.) Harmaja
Clitocybe geotropa (Bull. ex DC.)
Quél.
Agaricales, incertae
sedis
East Azerbaijan
Asef & Tavanaei 2004,
Asef 2007b and 2009b
Infundibulicybe gibba (Pers.)
Harmaja
Agaricales, incertae
sedis
Mazandaran
Asef & Etemad 2016
Infundibulicybe trulliformis (Fr.)
Gminder
Clitocybe trulliformis (Fr.) P. Karst.
Agaricales, incertae
sedis
East Azerbaijan
Asef 2009b
●Inocybe amethystina Kuyper
Agaricales, Inocybaceae
Mazandaran?
Bahram et al. 2013
HE687066 (env.,
SH1975517.08FU)
●Inocybe asterospora Quél.
Agaricales, Inocybaceae
Gilan, ‘Hyrcan’,
Mazandaran
Saber 1994c and 1999,
Bahram et al. 2012
FR852223 (env.,
SH2563972.08FU)

14 Mycologia Iranica - Vol. 7, 2020
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
●Inocybe castaneicolor A. La
Rosa, Bizio, Saitta & Tedersoo
Agaricales,
Inocybaceae
‘Hyrcan’,
Mazandaran?
Bahram et al. 2012, 2013
FR852226 (env.,
SH2183488.08FU)
, HE687070 (env.,
SH2183488.08FU
●Inocybe cincinnata (Fr.) Quél.
Agaricales,
Inocybaceae
‘Hyrcan’
Bahram et al. 2012
FR852255 (env.,
SH2338401.08FU)
!Inocybe corydalina Quél.
Agaricales,
Inocybaceae
Isfahan
Saber & Esmaeili Taheri 2002
●Inocybe decemgibbosa (Kühner)
Vauras
Agaricales,
Inocybaceae
Mazandaran?
Bahram et al. 2013
HE687073 (env.,
SH2187666.08FU)
Inocybe dulcamara (Pers.) P.
Kumm.
Agaricales,
Inocybaceae
Gilan, Mazandaran
Saber 1994c and 1999
Inocybe erubescens A. Blytt
Inocybe patouillardii Bres.
Agaricales,
Inocybaceae
Golestan
Saber 1999
●Inocybe flocculosa Sacc.
Agaricales,
Inocybaceae
Mazandaran?
Bahram et al. 2013
HE687062 (env.,
SH1952027.08FU)
●Inocybe geophylla (Sowerby) P.
Kumm.
Agaricales,
Inocybaceae
Mazandaran
Saber 1994c and 1999, Bahram
et al. 2013
HE687075 (env.,
SH2187809.08FU)
●Inocybe godeyi Gillet
Agaricales,
Inocybaceae
Mazandaran?
Bahram et al. 2013
HE687071 (env.,
SH2183506.08FU)
●Inocybe hirtella Bres.
Agaricales,
Inocybaceae
‘Hyrcan’, Isfahan
Saber & Esmaeili Taheri 2002,
Bahram et al. 2012
FR852249 (env.,
SH2081453.08FU)
●Inocybe huijsmanii Kuyper
Agaricales,
Inocybaceae
‘Hyrcan’
Bahram et al. 2012
FR852248 (env.,
SH1928024.08FU)
●Inocybe langei R. Heim
Agaricales,
Inocybaceae
Mazandaran?
Bahram et al. 2013
HE687072 (env.,
SH1932718.08FU)
!Inocybe leptocystis G.F. Atk.
Agaricales,
Inocybaceae
East Azerbaijan
Asef 2007b
Inocybe lilacina (Peck) Kauffman
Inocybe geophylla var. lilacina
(Peck) Gillet
Agaricales,
Inocybaceae
Gilan, Golestan,
Mazandaran
Saber 1994c and 1999
●Inocybe mixtilis (Britzelm.) Sacc.
Agaricales,
Inocybaceae
Mazandaran?
Bahram et al. 2013
HE687061 (env.,
SH2125306.08FU)
●Inocybe mystica Stangl &
Glowinski
Agaricales,
Inocybaceae
‘Hyrcan’
Bahram et al. 2012
FR852253 (env.,
SH2258584.08FU)
Inocybe napipes J.E. Lange
Agaricales,
Inocybaceae
Mazandaran
Saber 1999
Inocybe paludinella (Peck) Sacc.
Agaricales,
Inocybaceae
Gilan
Saber 1999
Inocybe praetervisa Quél.
Agaricales,
Inocybaceae
Golestan
Saber 1994c and 1999
●Inocybe pusio P. Karst.
Agaricales,
Inocybaceae
‘Hyrcan’
Bahram et al. 2012
FR852266 (env.,
SH1891582.08FU)
Inocybe subnudipes Kühner
Agaricales,
Inocybaceae
Kermanshah
Fadavi et al. 2015
Inocybe tabacina Furrer-Ziogas
Agaricales,
Inocybaceae
Mazandaran
Saber 1994c and 1999
Inocybe terrifera Kühner
Agaricales,
Inocybaceae
Kermanshah
Fadavi et al. 2015
Inosperma adaequatum (Britzelm.)
Matheny & Esteve-Rav.
Inocybe adaequata (Britzelm.)
Sacc.
Agaricales,
Inocybaceae
Mazandaran
Saber 1994c and 1999
Inosperma bongardii (Weinm.)
Matheny & Esteve-Rav.
Inocybe bongardii (Weinm.) Quél.
Agaricales,
Inocybaceae
Isfahan, Mazandaran
Saber 1999, Saber & Esmaeili
Taheri 2002
Inosperma cookei (Bres.) Matheny
& Esteve-Rav.
Inocybe cookei Bres.
Agaricales,
Inocybaceae
Gilan, Mazandaran
Saber 1994c and 1999
Inosperma maculatum (Boud.)
Matheny & Esteve-Rav.
Inocybe maculata Boud.
Agaricales,
Inocybaceae
Gilan, Mazandaran
Saber 1994c and 1999
●Laccaria amethystina Cooke
Agaricales,
Hydnangiaceae
Gilan, ‘Hyrcan’
Saber 1998, Bahram et al. 2012
FR852306 (env.,
SH2252734.08FU)
!Laccaria bicolor (Maire) P.D.
Orton
Agaricales,
Hydnangiaceae
Gilan, Golestan,
Mazandaran
Saber 1998

GHOBAD-NEJHAD ET AL.: The gilled fungi and boletes of Iran 15
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
Laccaria laccata (Scop.) Cooke
Agaricales,
Hydnangiaceae
East Azerbaijan,
Mazandaran
Asef 2007b, Asef & Etemad 2016
!Laccaria tortilis (Bolton) Cooke
Agaricales,
Hydnangiaceae
Mazandaran
Saber 1998
Lacrymaria lacrymabunda (Bull.)
Pat.
Drosophila velutina (Pers.) Kühner
& Romagn., Psathyrella velutina
(Pers.) Singer
Agaricales,
Psathyrellaceae
Mazandaran
Soleimani 1974, Saber 1994b
●Lactarius acris (Bolton) Gray
Russulales,
Russulaceae
‘Hyrcan’
Bahram et al. 2012
FR852031 (env.
SH2040289.08FU)
●Lactarius circellatus Fr.
Russulales,
Russulaceae
‘Hyrcan’
Bahram et al. 2012
FR852038 (env.,
SH2160217.08FU)
Lactarius deliciosus (L.) Gray
Russulales,
Russulaceae
Gilan
Saber 1989a
●Lactarius fulvissimus Romagn.
Russulales,
Russulaceae
‘Hyrcan’
Bahram et al. 2012
FR852027 (env.,
SH2160802.08FU)
●Lactarius rubrocinctus Fr.
Russulales,
Russulaceae
Mazandaran
Bahram et al. 2012, 2013
UDB005361 (env.,
SH2160166.08FU)
Lactarius scrobiculatus (Scop.) Fr.
Russulales,
Russulaceae
Gilan, Golestan
Saber 1989a
Lactarius serifluus (DC.) Fr.
Russulales,
Russulaceae
East Azerbaijan
Asef 2007b and 2011a
●Lactarius subdulcis (Pers.) Gray
Russulales,
Russulaceae
Gilan,
Mazandaran
Saber 1989a, Bahram et al. 2012,
2013
KX395722,
UDB005356
●Lactarius tabidus Fr.
Lactarius theiogalus (Bull.) Gray
Russulales,
Russulaceae
Golestan,
Mazandaran
Saber 1989a, Bahram et al. 2012,
2013
UDB005472
●Lactarius zonarius (Bull.) Fr.
Russulales,
Russulaceae
‘Hyrcan’
Bahram et al. 2012
FR852035 (env.,
SH1878819.08FU)
●Lactifluus glaucescens (Crossl.)
Verbeken
Lactarius glaucescens Crossl.
Russulales,
Russulaceae
‘Hyrcan’,
Mazandaran
Saber 1997c, Bahram et al. 2012,
this study
MT535681,
FR852034 (env.,
SH1939607.08FU)
Lactifluus piperatus (L.) Roussel
Lactarius piperatus (L.) Pers.
Russulales,
Russulaceae
Gilan, Golestan,
Isfahan?
Khabiri 1968, Saber 1989a
●Lactifluus vellereus (Fr.) Kuntze
Lactarius vellereus (Fr.) Fr.
Russulales,
Russulaceae
Gilan, ‘Hyrcan’,
Mazandaran
Saber 1989a, Bahram et al. 2012
FR852039 (env.,
SH1873315.08FU)
Lactifluus volemus (Fr.) Kuntze
Lactarius volemus (Fr.) Fr.
Russulales,
Russulaceae
East Azerbaijan,
Golestan
Saber 1989a, Asef 2007b and
2011a
●Leccinellum pseudoscabrum
(Kallenb.) Mikšík
Leccinum griseum (Quél.) Singer,
Leccinum pseudoscabrum
(Kallenb.) Šutara
Boletales, Boletaceae
East Azerbaijan,
Mazandaran
Saber 1995d, Asef 2008, Bahram
et al. 2013
HE687078 (env.,
SH1726911.08FU)
!Leccinum scabrum (Bull.) Gray
Boletales, Boletaceae
Golestan
Saber 1995d
!Leccinum versipelle (Fr. & Hök)
Snell
Leccinum testaceoscabrum Secr. ex
Singer
Boletales, Boletaceae
Gilan
Saber 1995d
Lentinellus cochleatus (Pers.) P.
Karst.
Russulales,
Auriscalpiaceae
Gilan
Saber 1990
Lentinellus ursinus (Fr.) Kühner
Russulales,
Auriscalpiaceae
Golestan
Saber 1997a
Lentinellus vulpinus (Sowerby)
Kühner & Maire
Russulales,
Auriscalpiaceae
Golestan
Saber 1990
Lentinus cyathiformis (Schaeff.)
Bres.
Neolentinus schaefferi (Weinm.)
Redhead & Ginns
Polyporales,
Polyporaceae
Gilan
Saber 1990
Lentinus lepideus (Fr.) Fr.
Neolentinus suffrutescens (Brot.)
T.W. May & A.E. Wood
Polyporales,
Polyporaceae
Alborz,
Mazandaran
Soleimani 1974 and 1976
Lentinus sajor-caju (Fr.) Fr.
Pleurotus sajor-caju (Fr.) Singer
Polyporales,
Polyporaceae
Golestan
Bari et al. 2019
Lentinus strigosus Fr.
Panus rudis Fr.
Polyporales,
Polyporaceae
Gilan, Golestan,
Mazandaran
Soleimani 1974 and 1976; Saber
1990 and 1997a
Lepiota anthomyces (Berk. &
Broome) Sacc.
Agaricales,
Agaricaceae
Khuzestan
Mohammadi Goltapeh 2002

16 Mycologia Iranica - Vol. 7, 2020
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
●Lentinus tigrinus (Bull.) Fr.
Panus tigrinus (Bull.) Singer
Polyporales,
Polyporaceae
Alborz, East
Azerbaijan,
Gilan,
Kermanshah,
Kohgiluyeh and
Boyer-Ahmad,
Lorestan,
Mazandaran,
Tehran
Petrak 1940, Watling & Sweeney
1974, Soleimani 1974 and 1976,
Saber 1990, Saber & Esmaeili
Taheri 2004, Grand et al. 2011,
Fadavi et al. 2015, Ghobad-
Nejhad et al. 2017 (as endophyte),
this study
MT535686,
MG208016 (as
endophyte)
Lepiota brunneoincarnata Chodat
& C. Martin
Agaricales,
Agaricaceae
Gilan
Asef 2015
Lepiota castanea Quél.
Agaricales,
Agaricaceae
East Azerbaijan,
Mazandaran
Saber 1994a, Asef & Tavanaei
2004, Asef 2007b, Asef & Etemad
2016
Lepiota cristata (Bolton) P. Kumm.
Agaricales,
Agaricaceae
Alborz, East
Azerbaijan,
Gilan, Isfahan,
Razavi Khorasan
Saber 1995f, Zokaei 2001,
Mohammadi Goltapeh 2002,
Saber & Esmaeili Taheri 2002,
Asef 2007b
Lepiota echinella Quél. & G.E.
Bernard
Agaricales,
Agaricaceae
Golestan
Asef 2015
!Lepiota felina (Pers.) P. Karst.
Agaricales,
Agaricaceae
?
Saber 1994a
Lepiota helveola Bres.
Agaricales,
Agaricaceae
West Azerbaijan
Fallahyan 1973
Lepiota leprica (Berk. & Broome)
Sacc.
Agaricales, Agaricaceae
Mazandaran
Mohammadi Goltapeh
2001
!Lepiota lilacea Bres.
Agaricales, Agaricaceae
Isfahan
Saber 1994a, Saber &
Esmaeili Taheri 2002
Lepiota metulispora (Berk. &
Broome) Sacc.
Agaricales, Agaricaceae
Khuzestan
Mohammadi Goltapeh
2002
!Lepiota micropholis (Berk. &
Broome) Sacc.
Agaricales, Agaricaceae
?
Saber 1994a
!Lepiota subalba Kühner ex P.D.
Orton
Agaricales, Agaricaceae
?
Saber 1994a
Lepiota subincarnata J.E. Lange
Agaricales, Agaricaceae
Mazandaran
Asef 2015
!Lepista irina (Fr.) H.E. Bigelow
Agaricales, incertae sedis
Golestan
Saber 2002a
!Lepista nuda (Bull.) Cooke
Agaricales, incertae sedis
East Azerbaijan,
Gilan
Saber 2002a
●Lepista saeva (Fr.) P.D. Orton
Lepista personata (Fr.) Cooke
Agaricales, incertae sedis
Mazandaran, Razavi
Khorasan, Zanjan
Zokaei 2001, Saber 2002a,
MK785234
!Leratiomyces squamosus (Pers.)
Bridge & Spooner
Stropharia squamosa (Pers.) Quél.
Agaricales,
Strophariaceae
Lorestan
Saber & Zangeneh 2004
●Leucoagaricus americanus
(Peck) Vellinga
Lepiota americana (Peck) Sacc,
Leucoagaricus bresadolae
(Schulzer) Bon & Boiffard
Agaricales, Agaricaceae
Mazandaran, West
Azerbaijan
Saber 1994a, Saber &
Mehravaran 2004,
GenBank
MT573394
!Leucoagaricus badhamii (Berk. &
Broome) Singer
Leucocoprinus badhamii (Berk. &
Broome) Locq.
Agaricales, Agaricaceae
Mazandaran
Saber 2000c
Leucoagaricus carneifolius
(Gillet) Wasser
Agaricales, Agaricaceae
Kermanshah
Fadavi et al. 2015
Leucoagaricus holospilotus (Berk.
& Broome) Bon
Leucocoprinus holospilotus (Berk.
& Broome) D.A. Reid, Lepiota
holospilota (Berk. & Broome)
Sacc.
Agaricales, Agaricaceae
Mazandaran
Mohammadi Goltapeh
2001
●Leucoagaricus leucothites
(Vittad.) Wasser
Lepiota naucina (Fr.) P. Kumm.,
Leucoagaricus naucinus (Fr.)
Singer
Agaricales, Agaricaceae
Isfahan, Mazandaran,
Tehran
Saber 1994a, Saber &
Esmaeili Taheri 2002, Asef
& Etemad 2016, this study
MT535696
!Leucoagaricus nympharum
(Kalchbr.) Bon
Macrolepiota puellaris (Fr.) M.M.
Moser
Agaricales, Agaricaceae
Gilan
Saber 1995f

GHOBAD-NEJHAD ET AL.: The gilled fungi and boletes of Iran 17
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
Leucoagaricus roseoalbus (Henn.)
Heinem. Lepiota roseoalba Henn.
Agaricales, Agaricaceae
Tehran
Mohammadi Goltapeh
2002
Leucoagaricus serenus (Fr.) Bon
& Boiffard
Lepiota serena (Fr.) Quél.
Agaricales, Agaricaceae
Khuzestan
Mohammadi Goltapeh
2002
!Leucocoprinus birnbaumii
(Corda) Singer
Agaricales, Agaricaceae
?
Saber 1994a
!Leucocoprinus brebissonii
(Godey) Locq.
Agaricales, Agaricaceae
?
Saber 1994a
!Leucocoprinus cepistipes
(Sowerby) Pat.
Lepiota cepistipes (Sowerby) P.
Kumm.
Agaricales, Agaricaceae
?
Saber 1994a
!Leucocoprinus magnusianus
(Henn.) Singer
Agaricales, Agaricaceae
Mazandaran
Saber 2000c
!Leucocybe candicans (Pers.)
Vizzini, P. Alvarado, G. Moreno &
Consiglio
Clitocybe candicans (Pers.) P.
Kumm.
Agaricales, incertae sedis
Mazandaran
Saber 2002a
Leucocybe houghtonii (W.
Phillips) Halama & Pencakowski
Clitocybe houghtonii (W. Phillips)
Dennis
Agaricales, incertae sedis
East Azerbaijan
Asef 2009b
!Leucopaxillus compactus (P.
Karst.) Neuhoff
Agaricales,
Tricholomataceae
Mazandaran
Saber 2002a
!Leucopaxillus giganteus
(Sowerby) Singer
Agaricales,
Tricholomataceae
East Azerbaijan
Saber 2002a
!Leucopaxillus pinicola J. Favre
Agaricales,
Tricholomataceae
Mazandaran
Saber 2002a
!Lyophyllum atratum (Fr.) Singer
Tephrocybe atrata (Fr.) Donk
Agaricales,
Lyophyllaceae
Mazandaran
Saber 2002a
!Lyophyllum baeospermum
Romagn.
Tephrocybe baeosperma (Romagn.)
M.M. Moser
Agaricales,
Lyophyllaceae
Mazandaran
Saber 2002b
●Macrocybe gigantea (Massee)
Pegler & Lodge
Tricholoma giganteum Massee
Agaricales,
Biannulariaceae
Ilam
Yousefshahi et al. 2020
MG867660
!Macrolepiota excoriata (Schaeff.)
Wasser
Agaricales, Agaricaceae
?
Saber 1994a
!Macrolepiota mastoidea (Fr.)
Singer
Macrolepiota konradii (Huijsman
ex P.D. Orton) M.M. Moser,
Macrolepiota prominens (Sacc.)
M.M. Moser
Agaricales, Agaricaceae
East Azerbaijan
Saber 1994a, Saber &
Zangeneh 2002, Asef
2007b
!Macrolepiota permixta (Barla)
Pacioni
Agaricales, Agaricaceae
Mazandaran
Saber & Zangeneh 2002
Macrolepiota procera (Scop.)
Singer
Agaricales, Agaricaceae
East Azerbaijan,
Mazandaran, Razavi
Khorasan
Saber 1994a, Mohammadi
Goltapeh 2001, Asef &
Etemad 2016
Mallocybe terrigena (Fr.) Matheny,
Vizzini & Esteve-Rav.
Inocybe terrigena (Fr.) Kuyper
Agaricales, Inocybaceae
Hamadan
Jadidian et al. 2015
●Marasmiellus candidus (Fr.)
Singer
Agaricales,
Omphalotaceae
Gilan, Golestan,
Mazandaran
Saber 1995g, Asef &
Etemad 2016
MT573397
!Marasmiellus confluens (Pers.)
J.S. Oliveira
Collybia confluens (Pers.) P.
Kumm., Gymnopus confluens
(Pers.) Antonín, Halling & Noordel.
Agaricales,
Omphalotaceae
Gilan
Saber 1998
!Marasmiellus peronatus (Bolton)
J.S. Oliveira
Collybia peronata (Bolton) P.
Kumm., Gymnopus peronatus
(Bolton) Gray
Agaricales,
Omphalotaceae
Gilan, Mazandaran
Saber 1998
!Marasmiellus ramealis (Bull.)
Singer
Marasmius ramealis (Bull.) Fr.
Agaricales,
Omphalotaceae
Gilan, Golestan,
Mazandaran
Saber 1995g

18 Mycologia Iranica - Vol. 7, 2020
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
!Marasmius atrorubens (Berk.)
Mont.
Agaricales,
Marasmiaceae
Gilan, Mazandaran,
Tehran
Mohammadi Goltapeh 2004
!Marasmius corrugatiformis
Singer
Agaricales,
Marasmiaceae
Gilan, Mazandaran,
Tehran
Mohammadi Goltapeh 2004
!Marasmius epiphyllus (Pers.) Fr.
Agaricales,
Marasmiaceae
Gilan
Saber 1995g
!Marasmius favoloides Henn.
Agaricales,
Marasmiaceae
Gilan, Mazandaran,
Tehran
Mohammadi Goltapeh 2004
!Marasmius ferrugineus Berk. &
M.A. Curtis
Agaricales,
Marasmiaceae
Gilan, Mazandaran,
Tehran
Mohammadi Goltapeh 2004
!Marasmius haematocephalus
(Mont.) Fr.
Agaricales,
Marasmiaceae
Gilan, Mazandaran,
Tehran
Mohammadi Goltapeh 2004
!Marasmius oreades (Bolton) Fr.
Agaricales,
Marasmiaceae
Gilan, West
Azerbaijan
Saber 1995g, Saber &
Mehravaran 2004
Marasmius rotula (Scop.) Fr.
Agaricales,
Marasmiaceae
Gilan, Golestan,
Mazandaran
Watling & Sweeney 1974,
Soleimani 1974 and 1976, Saber
1995g
!Marasmius rubroflavus (Theiss.)
Singer
Agaricales,
Marasmiaceae
Gilan, Mazandaran,
Tehran
Mohammadi Goltapeh 2004
Marasmius wynneae Berk. &
Broome
Agaricales,
Marasmiaceae
Mazandaran
Saber 1995g, Ershad 1995
●Megacollybia platyphylla (Pers.)
Kotl. & Pouzar
Oudemansiella platyphylla (Pers.)
M.M. Moser
Agaricales,
Marasmiaceae
Gilan, Golestan,
Mazandaran
Saber 2000d, Saber & Zangeneh
2002, Asef & Etemad 2016, this
study
MT535698
!Melanoleuca cognata (Fr.)
Konrad & Maubl.
Agaricales, incertae
sedis
Lorestan
Saber 2000d
*●Melanoleuca exscissa (Fr.)
Singer
Agaricales, incertae
sedis
East Azerbaijan
this study
MT535742
!Melanoleuca graminicola
(Velen.) Kühner & Maire
Agaricales, incertae
sedis
Tehran
Saber 2000d
!Melanoleuca grammopodia
(Bull.) Fayod
Agaricales, incertae
sedis
Tehran
Saber 2000d
!Melanoleuca strictipes (P. Karst.)
Jul. Schäff.
Agaricales, incertae
sedis
Tehran
Saber 2000d
!Melanoleuca subpulverulenta
(Pers.) Métrod
Agaricales, incertae
sedis
Golestan, Tehran
Saber 2000d
Montagnea arenaria (DC.) Zeller
Montagnea arenaria var.
macrospora D.A. Reid & Eicker,
Montagnites arenarius (DC.)
Morse
Agaricales,
Agaricaceae
Fars, Kerman, Qom,
Razavi Khorasan,
Semnan, Sistan and
Baluchestan
Petrak 1940 and 1949, Watling
& Gregory 1977, Esfandiari
1951, Saber 1986 and 2000b and
2002b, Saber & Esmaeili Taheri
2004, this study
Montagnea haussknechtii Rabenh.
Agaricales,
Agaricaceae
Gilan
Rabenhorst 1871
Mycena acicula (Schaeff.) P.
Kumm.
Agaricales,
Mycenaceae
East Azerbaijan
Asef & Muradov 2014
Mycena clavicularis (Fr.) Gillet
Agaricales,
Mycenaceae
East Azerbaijan
Asef & Muradov 2014
Mycena crocata (Schrad.) P.
Kumm.
Agaricales,
Mycenaceae
Gilan, Golestan,
Mazandaran
Saber 1995h, Asef & Etemad
2016, this study
!Mycena filopes (Bull.) P. Kumm.
Mycena amygdalina (Pers.) Singer
(s. Singer)
Agaricales,
Mycenaceae
Gilan, Golestan
Saber 1995h
!Mycena galericulata (Scop.) Gray
Agaricales,
Mycenaceae
Gilan, Golestan,
Mazandaran
Saber 1995h
Mycena galopus (Pers.) P. Kumm.
Agaricales,
Mycenaceae
Razavi Khorasan
Zokaei 2001
Mycena haematopus (Pers.) P.
Kumm.
Agaricales,
Mycenaceae
Gilan, Golestan,
Mazandaran
Saber 1995h, this study
!Mycena inclinata (Fr.) Quél.
Agaricales,
Mycenaceae
Gilan, Golestan,
Mazandaran
Saber 1995h
!Mycena metata (Fr.) P. Kumm.
Agaricales,
Mycenaceae
Golestan
Saber 1995h
!Mycena pearsoniana Dennis ex
Singer
Agaricales,
Mycenaceae
Gilan
Saber 1995h
!Mycena pelianthina (Fr.) Quél.
Agaricales,
Mycenaceae
Gilan
Saber 1995h
!Mycena polygramma (Bull.) Gray
Agaricales,
Mycenaceae
Gilan
Saber 1995h

GHOBAD-NEJHAD ET AL.: The gilled fungi and boletes of Iran 19
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
!Mycena pura (Pers.) P. Kumm.
Agaricales,
Mycenaceae
Gilan
Saber 1995h
!Mycena rapiolens J. Favre
Agaricales,
Mycenaceae
Mazandaran
Saber 1995h
!Mycena sanguinolenta (Alb. &
Schwein.) P. Kumm.
Agaricales,
Mycenaceae
Mazandaran
Saber 1995h
*●Mycena xantholeuca Kühner
Agaricales,
Mycenaceae
Ilam
this study
MT535719,
MT535733
Mycenastrum corium (Guers.)
Desv.
Agaricales,
Agaricaceae
Mazandaran,
Tehran
Saber 1993b and 1995b
!Mycenella salicina (Velen.)
Singer
Agaricales, incertae
sedis
Mazandaran
Saber 1995h
Mycetinis alliaceus (Jacq.) Earle
Marasmius alliaceus (Jacq.) Fr.
Agaricales,
Marasmiaceae
Gilan, Golestan,
Mazandaran
Watling & Sweeney 1974, Saber
1995g
!Mycetinis scorodonius (Fr.) A.W.
Wilson & Desjardin
Marasmius scorodonius (Fr.) Fr.
Agaricales,
Marasmiaceae
Gilan, Mazandaran
Saber 1995g
!Myxomphalia maura (Fr.) Hora
Agaricales, incertae
sedis
Tehran
Saber 2002a
Neoboletus luridiformis (Rostk.)
Gelardi
Boletus erythropus Pers.,
Neoboletus erythropus (Pers.) C.
Hahn
Boletales, Boletaceae
East Azerbaijan
Asef 2007b and 2008
Neofavolus suavissimus (Fr.) J.S.
Seelan, Justo & Hibbett
Lentinus suavissimus Fr.
Polyporales,
Polyporaceae
Gilan, Sistan and
Baluchestan
Saber 1990
Neolentinus adhaerens (Alb. &
Schwein.) Redhead & Ginns
Lentinus adhaerens (Alb. &
Schwein.) Fr., Lentinus
pulverulentus (Scop.) Fr.
Polyporales,
Polyporaceae
Gilan, Mazandaran
Watling & Sweeney 1974, Saber
1990
!Omphaliaster asterosporus (J.E.
Lange) Lamoure
Hygroaster asterosporus (J.E.
Lange) Singer
Agaricales, incertae
sedis
Golestan
Saber 1993d
!Omphalina mutila (Fr.) P.D.
Orton
Clitocybe josserandii (Singer)
Singer
Agaricales, incertae
sedis
Razavi Khorasan
Saber & Esmaeli Taheri 2004
!Omphalina pyxidata (Bull.) Quél.
Agaricales, incertae
sedis
Gilan
Saber 2002a
Omphalotus olearius (DC.) Singer
Agaricales,
Omphalotaceae
Gilan, Golestan,
Mazandaran
Saber 1990, Bari et al. 2019
Ossicaulis lignatilis (Pers.)
Redhead & Ginns
Clitocybe lignatilis (Pers.) P. Karst.
(as ‘Clitocybe pignatilis’)
Agaricales,
Lyophyllaceae
Mazandaran
Saber 1993c, Saber 1997a
*●Ossicaulis salomii Siquier &
Bellanger
Agaricales,
Lyophyllaceae
Tehran
this study
MT535738
Panaeolus acuminatus (P. Kumm.)
Quél.
Agaricales, incertae
sedis
Mazandaran
Asef & Etemad 2016
Panaeolus campanulatus (L.)
Quél.
Agaricales, incertae
sedis
Gilan
Buhse 1860, Khabiri 1968
!Panaeolus fimicola (Fr.) Quél.
Agaricales, incertae
sedis
East Azerbaijan
Asef 2007b
●Panaeolus olivaceus F.H. Møller
Notes: identity doubtful, with
caution.
Agaricales, incertae
sedis
Kermanshah
Seidmohammadi et al. 2019
MH593015
Panaeolus papilionaceus (Bull.)
Quél.
Panaeolus sphinctrinus (Fr.) Quél.
Agaricales, incertae
sedis
East Azerbaijan,
Mazandaran
Watling & Sweeney 1974, Saber
1995f
!Panaeolus plantaginiformis
(Lebedeva) E.F. Malysheva
Galeropsis plantaginiformis
(Lebedeva) Singer
Agaricales, incertae
sedis
Golestan
Saber 1993f
Panaeolus rickenii Hora
Agaricales, incertae
sedis
Razavi Khorasan
Zokaei 2001

20 Mycologia Iranica - Vol. 7, 2020
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
Panaeolus semiovatus (Sowerby)
S. Lundell & Nannf., Panaeolus
phalaenarum (Fr.) Quél.
Agaricales, incertae
sedis
Mazandaran
Watling & Sweeney 1974
!Panaeolus speciosus P.D. Orton
Panaeolus subfirmus P. Karst.
Agaricales, incertae
sedis
?
Saber 1994b
Panaeolus teutonicus Bride &
Métrod
Agaricales, incertae
sedis
Mazandaran
Watling & Gregory 1977
Panellus stipticus (Bull.) P. Karst.
Panus stipticus (Bull.) Fr.
Polyporales,
Panaceae
Gilan, Golestan,
Mazandaran
Rabenhorst 1871, Soleimani
1974 and 1976, Saber 1990,
Saber 1994b, Asef & Etemad
2016
Panus conchatus (Bull.) Fr.
Polyporales,
Panaceae
Gilan, Mazandaran
Soleimani 1974 and 1976
!Paragymnopus perforans
(Hoffm.) J.S. Oliveira
Micromphale perforans (Hoffm.)
Gray
Agaricales,
Marasmiaceae
Gilan, Golestan,
Mazandaran
Saber 1995g
Paralepista flaccida (Sowerby)
Vizzini
Lepista flaccida (Sowerby) Pat.,
Clitocybe flaccida (Sowerby) P.
Kumm., Clitocybe
infundibuliformis Quél.
Agaricales, incertae
sedis
East Azerbaijan,
Mazandaran
Saber 2002a, Asef & Tavanaei
2004, Asef 2007b and 2009b
!Parasola auricoma (Pat.)
Redhead, Vilgalys & Hopple
Coprinus auricomus Pat.
Agaricales,
Psathyrellaceae
?
Saber 1994b
!Parasola hemerobia (Fr.)
Redhead, Vilgalys & Hopple
Coprinus hemerobius Fr.
Agaricales,
Psathyrellaceae
?
Saber 1994b
!Parasola leiocephala (P.D. Orton)
Redhead, Vilgalys & Hopple
Coprinus leiocephalus P.D. Orton
Agaricales,
Psathyrellaceae
East Azerbaijan
Asef 2007b
!Parasola miser (P. Karst.)
Redhead, Vilgalys & Hopple
Coprinus miser P. Karst.
Agaricales,
Psathyrellaceae
Isfahan
Saber 1994b, Saber & Esmaeili
Taheri 2002
Parasola plicatilis (Curtis)
Redhead, Vilgalys & Hopple
Coprinus plicatilis (Curtis) Fr.,
Coprinus plicatilis var.
microsporus Kühner & Joss.
Agaricales,
Psathyrellaceae
Mazandaran, Razavi
Khorasan, Tehran
Saber 1994b, Mohammadi
Goltapeh 2000b, Zokaei 2001
!Paraxerula caussei (Maire)
Petersen
Oudemansiella renati Clémençon,
Xerula caussei Maire
Agaricales,
Physalacriaceae
East Azerbaijan
Asef 2007b
●Paxillus involutus (Batsch) Fr.
Boletales,
Paxillaceae
East Azerbaijan,
Gilan, Golestan,
Mazandaran
Saber 1993d, Asef 2007b,
Bahram et al. 2013
HE687082 (env.,
SH2308624.08FU)
!Phaeomarasmius erinaceus (Fr.)
Scherff. ex Romagn.
Agaricales,
Inocybaceae
?
Saber 1991b
!Phaeonematoloma myosotis (Fr.)
Bon
Pholiota myosotis (Fr.) Singer
Agaricales,
Strophariaceae
?
Saber 1991b
!Phellorinia herculeana (Pers.)
Kreisel
Agaricales,
Agaricaceae
Golestan
Saber 1993b
Phloeomana speirea (Fr.) Redhead
Mycena speirea (Fr.) Gillet
Agaricales,
Mycenaceae
Gilan, Golestan,
Mazandaran
Saber 1995h, Asef & Etemad
2016
●Pholiota adiposa (Batsch) P.
Kumm.
Pholiota aurivella (Batsch) P.
Kumm.
Agaricales,
Strophariaceae
Alborz, East
Azerbaijan,
Golestan, Isfahan,
Mazandaran, Razavi
Khorasan, Tehran,
West Azerbaijan
Soleimani 1974 and 1976,
Zokaei 2001, Saber & Esmaeili
Taheri 2002, Saber &
Mehravaran 2004, Asef &
Etemad 2016, this study
MT535689
!Pholiota astragalina (Fr.) Singer
Agaricales,
Strophariaceae
?
Saber 1991b
Pholiota gummosa (Lasch) Singer
Pholiota ochrochlora (Fr.) P.D.
Orton, Stropharia punctulata Sacc.
Agaricales,
Strophariaceae
Alborz, Gilan
Soleimani 1974 and 1976, Saber
1991b, Saber & Zangeneh 2002
Pholiota highlandensis (Peck)
A.H. Sm. & Hesler
Pholiota carbonaria (Fr.) Singer
Agaricales,
Strophariaceae
Kermanshah
Saber 1991b, Fadavi et al. 2013
*●Pholiota jahnii Tjall.-Beuk. &
Bas
Agaricales,
Strophariaceae
East Azerbaijan
this study
MT535737

GHOBAD-NEJHAD ET AL.: The gilled fungi and boletes of Iran 21
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
Pholiota populnea (Pers.) Kuyper
& Tjall.-Beuk.
Pholiota destruens (Brond.) Gillet
Agaricales,
Strophariaceae
Alborz, Gilan,
Tehran
Khabiri 1968, Soleimani 1974
and 1976
!Pholiota scamba (Fr.) M.M.
Moser
Agaricales,
Strophariaceae
?
Saber 1991b
!Pholiota spumosa (Fr.) Singer
Agaricales,
Strophariaceae
?
Saber 1991b
Pholiota squarrosa (Oeder) P.
Kumm.
Agaricales,
Strophariaceae
Golestan,
Mazandaran
Saber 1991b, Bari et al. 2019
Pholiota squarrosoides (Peck)
Sacc.
Agaricales,
Strophariaceae
Mazandaran
Saber 1991b, this study
!Pholiotina aporos (Kits van Wav.)
Clémençon
Conocybe aporos Kits van Wav.
Agaricales,
Bolbitiaceae
?
Saber 1993f
!Pholiotina arrhenii (Fr.) Singer
Agaricales,
Bolbitiaceae
Mazandaran
Saber 1993f
!Pholiotina striipes (Cooke) M.M.
Moser
Conocybe striaepes (Cooke) S.
Lundell
Agaricales,
Bolbitiaceae
Gilan
Saber & Zangeneh 2002
!Pholiotina vexans (P.D. Orton)
Bon
Conocybe blattaria (Fr.) Kühner
Agaricales,
Bolbitiaceae
Gilan
Saber 2000c
!Phyllotopsis nidulans (Pers.)
Singer
Agaricales,
Phyllotopsidaceae
Mazandaran
Saber 1993c
Pleurotellus chioneus (Pers.)
Kühner
Agaricales,
Inocybaceae
Gilan, Golestan
Saber 1990
Pleurotus calyptratus (Lindblad ex
Fr.) Sacc.
Agaricales,
Pleurotaceae
Mazandaran
Saber & Abaei 2001
●Pleurotus cornucopiae (Paulet)
Rolland
Agaricales,
Pleurotaceae
Golestan,
Mazandaran,
Tehran, West
Azerbaijan
Soleimani 1976, Saber 1990,
Saber & Mehravaran 2004, this
study
MT535734
Pleurotus djamor (Rumph. ex Fr.)
Boedijn
Pleurotus placentodes (Berk.) Sacc.
Agaricales,
Pleurotaceae
Gilan
Saber 1990
Pleurotus dryinus (Pers.) P.
Kumm.
Agaricales,
Pleurotaceae
Mazandaran
Saber 1997a
Pleurotus elongatipes Peck
Agaricales,
Pleurotaceae
Mazandaran
Saber 1997a
●Pleurotus eryngii (DC.) Quél.
Pleurotus fuscus Battarra ex Bres.
Agaricales,
Pleurotaceae
Alborz, Ardabil,
Chaharmahal and
Bakhtiari, East
Azerbaijan, Fars,
Isfahan, Kohgiluyeh
and Boyer-Ahmad,
Kurdistan, Lorestan,
South Khorasan,
Tehran, West
Azerbaijan
Petrak 1940, Esfandiari 1948,
Heim 1960, Saber 1990, Saber
1997a, Saber & Esmaeili Taheri
2002 and 2004, Saber &
Mehravaran 2004, Ravash et al.
2010, Zervakis et al. 2014, this
study
MT535679,
HM998837
(numerous
sequences in
GenBank)
Pleurotus fossulatus (Cooke) Sacc.
Agaricales,
Pleurotaceae
Chaharmahal and
Bakhtiari, Fars,
Kohgiluyeh and
Boyer-Ahmad,
Kurdistan
Saber 1997a, Ravash et al. 2010
●Pleurotus nebrodensis (Inzenga)
Quél.
Pleurotus eryngii var. nebrodensis
(Inzenga) Sacc.
Agaricales,
Pleurotaceae
Chaharmahal and
Bakhtiari,
Kohgiluyeh and
Boyer-Ahmad,
Kurdistan
Saber 1997a, Ravash et al. 2010,
Zervakis et al. 2014
HM998835
●Pleurotus ostreatus (Jacq.) P.
Kumm.
Agaricus salignus Pers., Agaricus
salignus var. ochraceus Pers.
Agaricales,
Pleurotaceae
Alborz, East
Azerbaijan, Gilan,
Golestan, Hamadan,
Kermanshah,
Mazandaran, North
Khorasan, Razavi
Khorasan, Tehran,
West Azerbaijan
Buhse 1860, Khabiri 1968,
Soleimani 1974 and 1976, Saber
1990, Zokaei 2001, Saber &
Esmaeili Taheri 2004, Saber &
Mehravaran 2004, Asef 2007b,
Asef & Etemad 2016, this study
MT535690
Pleurotus pulmonarius (Fr.) Quél.
Agaricales,
Pleurotaceae
Gilan, Golestan,
Kermanshah,
Mazandaran
Saber 1990, Bari et al. 2019

22 Mycologia Iranica - Vol. 7, 2020
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
!Pluteus aurantiorugosus (Trog)
Sacc.
Agaricales,
Pluteaceae
Gilan, Mazandaran
Saber 1993e
●Pluteus cervinus (Schaeff.) P.
Kumm.
Pluteus atricapillus (Batsch) Fayod
Agaricales,
Pluteaceae
East Azerbaijan,
Golestan,
Mazandaran
Saber 1991a, Saber & Esmaeili
Taheri 2004, Asef 2007b, Asef
& Etemad 2016, Bari et al. 2019,
this study
MT535687
Pluteus chrysophaeus (Schaeff.)
Quél.
Pluteus luteovirens Rea
Agaricales,
Pluteaceae
Golestan,
Mazandaran
Saber 1991a
●Pluteus cinereofuscus J.E. Lange
Agaricales,
Pluteaceae
Kermanshah
Seidmohammadi et al. 2018a
MH595963
!Pluteus depauperatus Romagn.
Agaricales,
Pluteaceae
Gilan, Golestan,
Mazandaran
Saber 1993e
Pluteus exiguus (Pat.) Sacc.
Agaricales,
Pluteaceae
Mazandaran
Watling & Sweeney 1974
Pluteus leoninus (Schaeff.) P.
Kumm.
Agaricales,
Pluteaceae
Mazandaran
Saber 1991a, Asef & Etemad
2016
●Pluteus nanus (Pers.) P. Kumm.
Agaricales,
Pluteaceae
Kermanshah,
Mazandaran, West
Azerbaijan
Saber 1991a, Saber &
Mehravaran 2004,
Seidmohammadi et al. 2018a
MH595974
!Pluteus pellitus (Pers.) P. Kumm.
Agaricales,
Pluteaceae
?
Saber 1991a
!Pluteus petasatus (Fr.) Gillet
Pluteus patricius (Schulzer) Boud.
Agaricales,
Pluteaceae
?
Saber 1991a
!Pluteus punctipes P.D. Orton
Agaricales,
Pluteaceae
Golestan
Saber 1993e
Pluteus romellii (Britzelm.) Sacc.
Agaricales,
Pluteaceae
Golestan, Isfahan
Saber 1991a, Saber & Esmaeili
Taheri 2002
Pluteus salicinus (Pers.) P. Kumm.
Agaricales,
Pluteaceae
Golestan,
Mazandaran
Saber 1995f
Pluteus semibulbosus (Lasch)
Gillet
Agaricales,
Pluteaceae
Mazandaran
Saber 1991a, this study
Pluteus thomsonii (Berk. &
Broome) Dennis
Agaricales,
Pluteaceae
Golestan,
Mazandaran
Saber 2000c
!Pluteus umbrosus (Pers.) P.
Kumm.
Agaricales,
Pluteaceae
?
Saber 1991a
●Pogonoloma macrocephalum
(Schulz.) Sánchez-García
Tricholoma macrocephalum
Schulz.
Agaricales, incertae
sedis
Golestan,
Kermanshah
Watling & Gregory 1977,
GenBank
MH595847
*●Psathyrella bivelata Contu
Agaricales,
Psathyrellaceae
Tehran
this study
MT535693
●Psathyrella candolleana (Fr.)
Maire
Agaricales,
Psathyrellaceae
Fars, Mazandaran,
Razavi Khorasan,
Sistan and
Baluchestan,
Tehran, West
Azerbaijan
Saber 1994b, Saber & Esmaeili
Taheri 2004, Saber &
Mehravaran 2004, Asef &
Etemad 2016, Teimoori-
Boghsani et al. 2020 (as
endophyte), this study
MT535718,
MK367774 (as
endophyte)
Psathyrella clivensis (Berk. &
Broome) P.D. Orton
Agaricales,
Psathyrellaceae
Hamadan
Jadidian et al. 2015
*●Psathyrella fatua (Fr.) Konrad
& Maubl.
Agaricales,
Psathyrellaceae
Kohgiluyeh and
Boyer-Ahmad
this study
MT535695
*●Psathyrella hellebosensis D.
Deschuyteneer & A. Melzer
Agaricales,
Psathyrellaceae
Semnan
this study
MT535716
!Psathyrella laevissima (Romagn.)
Singer
Agaricales,
Psathyrellaceae
?
Saber 1994b
!Psathyrella microrhiza (Lasch)
Konrad & Maubl.
Agaricales,
Psathyrellaceae
?
Saber 1994b
Psathyrella multipedata (Peck)
A.H. Sm.
Agaricales,
Psathyrellaceae
Mazandaran
Saber 1994b, Asef & Etemad
2016
!Psathyrella obtusata (Fr.) A.H.
Sm.
Agaricales,
Psathyrellaceae
Golestan,
Mazandaran, Razavi
Khorasan, South
Khorasan
Saber 1994b, Saber & Emaeili
Taheri 2004
!Psathyrella pennata (Fr.) A.
Pearson & Dennis
Agaricales,
Psathyrellaceae
?
Saber 1994b
!Psathyrella piluliformis (Bull.)
P.D. Orton
Psathyrella hydrophila (Bull.)
Maire
Agaricales,
Psathyrellaceae
?
Saber 1994b

GHOBAD-NEJHAD ET AL.: The gilled fungi and boletes of Iran 23
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
!Psathyrella prona (Fr.) Gillet
Agaricales,
Psathyrellaceae
?
Saber 1994b
!Psathyrella pseudogracilis
(Romagn.) M.M. Moser
Agaricales,
Psathyrellaceae
?
Saber 1994b
!Psathyrella spadiceogrisea
(Schaeff.) Maire Psathyrella
groegeri G. Hirsch, Psathyrella
vernalis (J.E. Lange) M.M. Moser
Agaricales,
Psathyrellaceae
Mazandaran, West
Azerbaijan
Saber 1994b, Saber 1995f, Saber
& Mehravaran 2004
!Psathyrella squamosa (P. Karst.)
A.H. Sm.
Agaricales,
Psathyrellaceae
?
Saber 1994b
Psathyrella tephrophylla
(Romagn.) M.M. Moser
Agaricales,
Psathyrellaceae
Hamadan
Jadidian et al. 2015
●Pseudoclitocybe cyathiformis
(Bull.) Singer
Clitocybe cyathiformis (Bull.) P.
Kumm.
Agaricales, incertae
sedis
Gilan, Golestan,
Ilam, Mazandaran
Buhse 1860, Saber 2002a, this
study
MT535721
Pseudosperma perlatum (Cooke)
Matheny & Esteve-Rav.
Inocybe perlata (Cooke) Sacc.
Agaricales,
Inocybaceae
East Azerbaijan,
Golestan
Saber 1994c, Saber 1999, Asef
2007b
●Pseudosperma rimosum (Bull.)
Matheny & Esteve-Rav.
Inocybe fastigiata (Schaeff.) Quél.,
Inocybe rimosa (Bull.) P. Kumm.
Agaricales,
Inocybaceae
East Azerbaijan,
‘Hyrcan’, Ilam,
Mazandaran
Saber 1994c, Asef 2007b, Saber
1999, Asef & Tavanaei 2004,
Bahram et al. 2012, Yousefshahi
et al. 2020
MF278770,
FR852236 (env.,
SH1951094.08FU)
Psilocybe atrobrunnea (Lasch)
Gillet
Agaricales,
Hymenogastraceae
Kermanshah
Fadavi et al. 2013
!Psilocybe coprophila (Bull.) P.
Kumm.
Agaricales,
Hymenogastraceae
?
Saber 1991b
!Psilocybe cyanescens Wakef.
Agaricales,
Hymenogastraceae
Tehran
Saber 1995f
!Psilocybe serbica M.M. Moser &
E. Horak
Agaricales,
Hymenogastraceae
Tehran
Saber 1995f
Resupinatus applicatus (Batsch)
Gray
Agaricales,
Pleurotaceae
Mazandaran
Saber 1990
Rhodocollybia maculata (Alb. &
Schwein.) Singer
Collybia maculata (Alb. &
Schwein.) P. Kumm.
Agaricales,
Omphalotaceae
Gilan
Khabiri 1968
!Rhodocollybia prolixa (Fr.)
Antonín & Noordel.
Collybia distorta (Fr.) Quél.,
Rhodocollybia prolixa var. distorta
(Fr.) Antonín, Halling & Noordel.
Agaricales,
Omphalotaceae
Mazandaran
Saber 1998
Rhodotus palmatus (Bull.) Maire
Agaricales,
Physalacriaceae
Golestan
Saber 1990
!Rubroboletus satanas (Lenz)
Kuan Zhao & Zhu L. Yang
Boletus satanas Lenz
Boletales, Boletaceae
Golestan,
Mazandaran
Saber 1995d
!Russula acetolens Rauschert
Russula lutea (Huds.) Gray
Russulales,
Russulaceae
Mazandaran
Saber 2000a
!Russula alutacea (Pers.) Fr.
Russulales,
Russulaceae
Golestan
Saber & Ershad 2004
!Russula anthracina Romagn.
Russulales,
Russulaceae
Mazandaran
Bahram & Asef 2008b
●Russula atropurpurea Peck
Russulales,
Russulaceae
Gilan, ‘Hyrcan’,
Mazandaran
Saber & Ershad 2004, Bahram et
al. 2012
FR852111 (env.,
SH1667944.08FU)
!Russula atrorubens Quél.
Russulales,
Russulaceae
Gilan
Saber & Ershad 2004
!Russula brunneoviolacea
Crawshay
Russulales,
Russulaceae
Mazandaran
Saber & Ershad 2004
!Russula carminipes J. Blum
Russulales,
Russulaceae
Golestan
Saber & Ershad 2004
!Russula claroflava Grove
Russulales,
Russulaceae
Gilan
Saber & Ershad 2004
!Russula cyanoxantha (Schaeff.)
Fr.
Russula cyanoxantha f. peltereaui
Singer
Russulales,
Russulaceae
East Azerbaijan,
Golestan
Saber & Ershad 2004, Asef
2007b and Asef 2011a
●Russula delica Fr.
Russulales,
Russulaceae
East Azerbaijan,
‘Hyrcan’
Asef 2007b, Asef 2011a,
Bahram et al. 2012
FR852108 (env.,
SH1865348.08FU)

24 Mycologia Iranica - Vol. 7, 2020
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
Russula emetica (Schaeff.) Pers.
Russulales,
Russulaceae
East Azerbaijan,
Golestan, Mazandaran
Saber & Ershad 2004, Asef
2007b and 2011a
*●Russula emeticolor J. Schaeffer
Russulales,
Russulaceae
Golestan, Mazandaran
this study
MT535680,
MT535682
!Russula farinipes Romell
Russulales,
Russulaceae
Mazandaran
Saber & Ershad 2004
●Russula foetens Pers.
Russulales,
Russulaceae
Mazandaran
Saber 2000a, Bahram et al.
2013
HE687097 (env.,
SH1569737.08FU)
!Russula graveolens Romell
Russulales,
Russulaceae
Golestan
Saber & Ershad 2004
!Russula grisea Fr.
Russulales,
Russulaceae
Mazandaran
Saber & Ershad 2004
●Russula heterophylla (Fr.) Fr.
Russulales,
Russulaceae
Golestan, ‘Hyrcan’
Saber & Ershad 2004,
Bahram et al. 2012
FR852114 (env.,
SH1956732.08FU)
Russula integra (L.) Fr.
Russula polychroma Singer ex
Hora
Russulales,
Russulaceae
Golestan, Mazandaran
Watling & Sweeney 1974,
Saber & Ershad 2004
!Russula ionochlora Romagn.
Russulales,
Russulaceae
Gilan
Saber & Ershad 2004
Russula lilacea Quél.
Russulales,
Russulaceae
East Azerbaijan
Asef 2007b and 2011a
●Russula luteotacta Rea
Russulales,
Russulaceae
Mazandaran?
Bahram et al. 2013
HE687092 (env.,
SH1667972.08FU)
!Russula nigricans Fr.
Russulales,
Russulaceae
Mazandaran
Saber 2000a
!Russula ochroleuca Pers.
Russulales,
Russulaceae
Gilan, Golestan
Saber 2000a
!Russula ochroleucoides Kauffman
Russulales,
Russulaceae
Gilan
Saber & Ershad 2004
●Russula olivacea Pers.
Russulales,
Russulaceae
‘Hyrcan’, Mazandaran
Saber & Ershad 2004,
Bahram et al. 2012
FR852106 (env.,
SH1560122.08FU)
!Russula pectinata Fr.
Russulales,
Russulaceae
Mazandaran
Saber & Ershad 2004
!Russula pectinatoides Peck
Russulales,
Russulaceae
Gilan, Golestan
Saber & Ershad 2004
●Russula persicina Krombh.
Russulales,
Russulaceae
Mazandaran?
Bahram et al. 2013
HE687094 (env.,
SH1509032.08FU)
!Russula perlactea Murrill
Russulales,
Russulaceae
Gilan
Saber & Ershad 2004
!Russula puellaris Fr.
Russulales,
Russulaceae
Mazandaran
Saber & Ershad 2004
!Russula queletii Fr.
Russulales,
Russulaceae
Mazandaran
Saber & Ershad 2004
!Russula risigallina (Batsch) Sacc.
Russula chamaeleontina (Lasch)
Fr.
Russulales,
Russulaceae
Golestan
Saber & Ershad 2004
!Russula romellii Maire
Russulales,
Russulaceae
Golestan
Saber & Ershad 2004
●Russula rosea Pers.
Russulales,
Russulaceae
‘Hyrcan’
Bahram et al. 2012
FR852107 (env.,
SH1712188.08FU)
!Russula silvestris (Singer)
Reumaux
Russula emetica var. silvestris
Singer
Russulales,
Russulaceae
Mazandaran
Saber & Ershad 2004
●Russula solaris Ferd. & Winge
Russulales,
Russulaceae
‘Hyrcan’
Bahram et al. 2012
FR852110 (env.,
SH2136150.08FU)
Russula sororia (Fr.) Romell
Russulales,
Russulaceae
East Azerbaijan
Asef 2007b and 2011a
!Russula torulosa Bres.
Russulales,
Russulaceae
Mazandaran
Bahram & Asef 2008b
●Russula versicolor Jul. Schäff.
Russulales,
Russulaceae
‘Hyrcan’
Bahram et al. 2012
FR852102 (env.,
SH2136145.08FU)
●Russula veternosa Fr.
Russulales,
Russulaceae
‘Hyrcan’
Bahram et al. 2012
FR852104 (env.,
SH1905766.08FU)
●Russula vinosopurpurea Jul.
Schäff.
Russulales,
Russulaceae
‘Hyrcan’
Bahram et al. 2012
FR852115 (env.,
SH1702036.08FU)
!Russula virescens (Schaeff.) Fr.
Russula aeruginosa (Pers.) Pers.
Russulales,
Russulaceae
Gilan, Mazandaran
Saber 2000a, Bahram &
Asef 2008b
!Russula xerampelina (Schaeff.)
Fr.
Russulales,
Russulaceae
Golestan
Saber & Ershad 2004

GHOBAD-NEJHAD ET AL.: The gilled fungi and boletes of Iran 25
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
Saproamanita codinae (Maire)
Redhead, Vizzini, Drehmel &
Contu
Amanita codinae (Maire) Bertault
Agaricales,
Amanitaceae
West Azerbaijan
Saber & Mehravaran 2001
Sarcomyxa serotina (Pers.) V.
Papp
Panellus serotinus (Pers.) Kühner
Agaricales,
Sarcomyxaceae
Mazandaran
Saber 1990, Saber 1994b,
Saber and Zangeneh 2004
*●Simocybe centunculus (Fr.: Fr.)
P. Karst.
Agaricales,
Inocybaceae
East Azerbaijan
this study
MT535746
!Sphagnurus paluster (Peck)
Redhead & V. Hofst.
Collybia thelephora (Cooke &
Massee) Sacc., Tephrocybe
palustris (Peck) Donk
Agaricales,
Lyophyllaceae
Mazandaran
Saber & Zangeneh 2004
Strobilomyces strobilaceus (Scop.)
Berk.
Boletales, Boletaceae
Gilan
Asef 2013
!Strobilurus esculentus (Wulfen)
Singer
Agaricales,
Physalacriaceae
Mazandaran
Saber 1998
!Strobilurus tenacellus (Pers.)
Singer
Agaricales,
Physalacriaceae
Gilan, Mazandaran
Saber 1998
!Stropharia aeruginosa (Curtis)
Quél.
Agaricales,
Strophariaceae
East Azerbaijan
Saber 1991b, Asef &
Tavanaei 2004, Asef 2007b
Stropharia coronilla (Bull.) Quél.
Psilocybe coronilla (Bull.) Noordel.
Agaricales,
Hymenogastraceae
Kermanshah
Fadavi et al. 2013
!Stropharia melanosperma (Bull.)
Quél.
Agaricales,
Strophariaceae
West Azerbaijan
Saber & Mehravaran 2004
●Suillellus comptus (Simonini)
Vizzini, Simonini & Gelardi
Boletus comptus Simonini
Boletales, Boletaceae
Ilam, Kermanshah
Seidmohammadi et al. 2017,
Yousefshahi et al. 2020
MF278769 (ident.
97%)
*●Suillellus luridus (Schaeff.)
Murrill
Boletales, Boletaceae
Lorestan
this study
MT535740
!Suillus bovinus (L.) Roussel
Boletales, Suillaceae
Golestan
Saber 1995d
●Suillus collinitus (Fr.) Kuntze
Boletales, Suillaceae
East Azerbaijan, Gilan,
Mazandaran, Tehran,
West Azerbaijan
Saber 1995d, Saber &
Zangeneh 2002, Asef 2007b
and 2008, Bahram et al.
2013
HE687129 (env.,
SH1935285.08FU)
●Suillus granulatus (L.) Roussel
Boletales, Suillaceae
East Azerbaijan,
Mazandaran?,Tehran
Saber 1995d, Asef 2007b
and 2008, Bahram et al.
2013
HE687128 (env.,
SH2260335.08FU)
●Tricholoma acerbum (Bull.)
Quél.
Agaricales,
Tricholomataceae
Mazandaran
GenBank Direct Submission
MH628231
!Tricholoma argyraceum (Bull.)
Gillet
Agaricales,
Tricholomataceae
Tehran
Zangeneh & Maivan 1998
!Tricholoma caligatum (Viv.)
Ricken
Agaricales,
Tricholomataceae
Gilan
Saber 2000d
!Tricholoma equestre (L.) P.
Kumm.
Tricholoma flavovirens (Pers.) S.
Lundell
Agaricales,
Tricholomataceae
East Azerbaijan
Asef 2007b
Tricholoma fulvum (Fr.) Bigeard
& H. Guill.
Agaricales,
Tricholomataceae
Razavi Khorasan
Zokaei 2001
●Tricholoma lascivum (Fr.) Gillet
Agaricales,
Tricholomataceae
Gilan, Golestan,
‘Hyrcan’
Saber 2000d, Bahram et al.
2012
FR852322 (env.,
SH1840391.08FU)
!Tricholoma orirubens Quél.
Tricholoma guttatum (Schaeff.) P.
Kumm.
Agaricales,
Tricholomataceae
East Azerbaijan
Asef 2007b
!Tricholoma psammopus
(Kalchbr.) Quél.
Agaricales,
Tricholomataceae
Mazandaran
Saber 2000d
!Tricholoma robustum (Alb. &
Schwein.) Ricken
Agaricales,
Tricholomataceae
Tehran
Saber 2000d
●Tricholoma scalpturatum (Fr.)
Quél.
Agaricales,
Tricholomataceae
Hamadan,
Mazandaran?
Bahram et al. 2013, Jadidian
et al. 2015
HE687177 (env.,
SH2569733.08FU)
●Tricholoma cingulatum (Almfelt
ex Fr.) Jacobasch
Agaricales,
Tricholomataceae
Mazandaran
GenBank Direct Submission
MH620781
●Tricholoma sulphureum (Bull.)
P. Kumm.
Agaricales,
Tricholomataceae
Golestan, ‘Hyrcan’,
Razavi Khorasan
Zokaei 2001, Bahram et al.
2012
FR852325 (env.,
SH1891870.08FU)

26 Mycologia Iranica - Vol. 7, 2020
Table 1. Continued.
Species
Order, family
Distribution
References
ITS
GenBank/UNITE
accession no.
●Tricholoma terreum (Schaeff.) P.
Kumm. Tricholoma gausapatum
(Fr.) Quél.
Tricholoma myomyces (Pers.) J.E.
Lange
Agaricales,
Tricholomataceae
East Azerbaijan,
Mazandaran?, Tehran
Zangeneh & Maivan
1998, Saber 2000d, Asef
2007b, Bahram et al.
2013
HE687176 (env.,
SH2569728.08FU)
!Tricholoma ustale (Fr.) P. Kumm.
Agaricales,
Tricholomataceae
Golestan,
Mazandaran
Saber 2000d
!Tricholoma ustaloides Romagn.
Agaricales,
Tricholomataceae
Mazandaran
Saber 2000d
!Tricholoma vaccinum (Schaeff.)
P. Kumm.
Agaricales,
Tricholomataceae
Mazandaran
Saber 2000d
!Tricholomopsis formosa (Murrill)
Singer
Agaricales, incertae sedis
Gilan
Saber 2000d
!Tubaria confragosa (Fr.) Harmaja
Agaricales, Tubariaceae
East Azerbaijan
Asef 2007b
!Tubaria conspersa (Pers.) Fayod
Agaricales, Tubariaceae
Mazandaran
Saber & Zangeneh 2002
Tubaria furfuracea (Pers.) Gillet
!Tubaria hiemalis Romagn. ex Bon,
Tubaria pellucida (Bull. & Vent.)
Sacc.
Agaricales, Tubariaceae
Gilan, Golestan,
Tehran, West
Azerbaijan
Saber 1990, Saber &
Mehravaran 2004
!Tubaria pallidospora J.E. Lange
Agaricales, Tubariaceae
Tehran
Saber & Zangeneh 2002
!Tylopilus felleus (Bull.) P. Karst.
Boletales, Boletaceae
Mazandaran
Saber 1995d
Volvariella iranica (Fallahyan)
Szczepka
Volvaria iranica Fallahyan
Agaricales, Pluteaceae
Mazandaran
Fallahyan 1973
!Volvariella bombycina (Schaeff.)
Singer
Agaricales, Pluteaceae
Golestan
Saber 1993f
Volvariella hypopithys (Fr.)
Shaffer
Volvariella plumulosa (Lasch)
Singer
Agaricales, Pluteaceae
Mazandaran, Razavi
Khorasan
Saber 2000c, Zokaei
2001
!Volvariella murinella (Quél.)
M.M. Moser ex Dennis, P.D. Orton
& Hora
Agaricales, Pluteaceae
Golestan
Saber 2000c
!Volvariella pusilla (Pers.) Singer
Agaricales, Pluteaceae
?
Saber 1991a
Volvariella volvacea (Bull.) Singer
Volvaria volvacea (Bull.) P.
Kumm.
Agaricales, Pluteaceae
Kermanshah, West
Azerbaijan
Fallahyan 1973, Fadavi et
al. 2015
Volvopluteus gloiocephalus (DC.)
Vizzini, Contu & Justo
Volvariella gloiocephala (DC.)
Boekhout & Enderle, Volvariella
speciosa (Fr.) Singer,
Agaricales, Pluteaceae
East Azerbaijan,
Kermanshah
Saber 1991a, Asef 2007b,
Fadavi et al. 2015
!Xerocomellus chrysenteron
(Bull.) Šutara
Boletus chrysenteron Bull.
Boletales, Boletaceae
Golestan
Saber 1995d
!Xerocomellus pruinatus (Fr. &
Hök) Šutara
Boletus pruinatus Fr. & Hök,
Xerocomus pruinatus (Fr. & Hök)
Quél.
Boletales, Boletaceae
Mazandaran
Saber 1995d
!Xerocomus moravicus (Vacek)
Herink
Boletus leonis D.A. Reid
Boletales, Boletaceae
Mazandaran
Saber 1995d
●Xerocomus subtomentosus (L.)
Quél.
Xerocomus lanatus (Rostk.) Singer
Boletales, Boletaceae
East Azerbaijan,
Golestan, ‘Hyrcan’
Saber 1995d, Asef 2007b
and 2008, Bahram et al.
2012
FR852284 (env.,
SH1873179.08FU)
●Xerula pudens (Pers.) Singer
Oudemansiella badia M.M. Moser
Agaricales,
Physalacriaceae
Golestan
Saber & Zangeneh 2002,
this study
MT535743
Zhuliangomyces ochraceoluteus
(P.D. Orton) Redhead
Limacella ochraceolutea P.D.
Orton
Agaricales, Amanitaceae
East Azerbaijan
Asef & Muradov 2014

GHOBAD-NEJHAD ET AL.: The gilled fungi and boletes of Iran 27
Table 2. Excluded and doubtful taxa. Names in informal references are marked with an exclamation mark (!).
Name in reference
Reference
Notes
!Agaricus erubescens Mont. (Clitocybe erubescens Sacc.)
Saber 2002a
Illegitime name and also quite unclear taxon
(see Index Fungorum).
!Clitocybe dealbata (Sowerby) P. Kumm.
Saber 2002a
Doubtful name.
!Clitocybe ericetorum Quél.
Saber 2002a, Asef 2007b
Doubtful name. No description provided by
citing references for clarification.
!Clitocybula lacerata (Scop.) Singer ex Métrod
Saber 1998
Quite unlikely in Iran, associated with old-
growth coniferous forests.
!Collybia ocellata (Fr.) P. Kumm.
Asef 2007b
A nomen dubium (Antonín & Noordeloos
2010).
Collybia xylophila (Weinm.) Sacc.
Petrak & Esfandiari 1941,
Esfandiari 1946
A nomen dubium (Antonín & Noordeloos
2010).
Cortinarius arvinaceus Fr.
Asef 2007a and 2007b
Likely a nomen dubium (Kibby et al. 2009).
!Cortinarius mucifluus Fr.
Asef 2007b
Unlikely to occur in Iran. Associated with
Pinus and Picea, in dry Pinus forests and also
in bogs.
Cortinarius pseudonapus (Rob. Henry ex M.M. Moser)
M.M. Moser
Asef 2007b and 2009a
Unlikey in Iran, associated with Picea forests.
Cortinarius sublubricus (Jul. Schäff. ex M.M. Moser)
M.M. Moser
Asef 2007b and 2009a
Poorly known, likely a nomen dubium.
Cortinarius subvalidus Rob. Henry (C. saginus (Fr.) Fr.)
Asef 2007b and 2009a
A typical species of natural spruce forests,
unlikely in Iran.
!Cortinarius traganus (Fr.) Fr.
Saber 1995f
A typical species of natural spruce and pine
forests, unlikely in Iran.
!Entoloma abortivum (Berk. & M.A. Curtis) Donk
(Clitopilus abortivus (Berk. & M.A. Curtis) Sacc.)
Saber 1993d
North American species. Very unlikely to
occur in Iran.
!Galera conferta (Bolton) P. Kumm.
Saber 1994c
Poorly known taxon without a modern
interpretation in Europe.
!Hebeloma versipelle (Fr.) Gillet
Saber 1994c
A nomen dubium.
!Hygrocybe obrussea (Fr.) Wünsche
Saber 1993d
Doubtful name.
!Hygrocybe parvula (Peck) Murrill
Asef 2007b
A North American taxon, unlikely to occur in
Iran.
Lactarius insulsus
Khabiri 1968
Authority not mentioned. Affinity ambiguous.
!Mycena alcalina (Fr.) P. Kumm.
Saber 1995h
Doubtful name, more species now.
!Mycena sepia J.E. Lange
Saber 1995h
Identity of this specis is unclear; it belongs to
the M. filopes/M. metata complex (Aronsen &
Læssøe 2016).
!Oudemansiella ephippium (Fr.) M.M. Moser
Saber 1998
Unclear and doubtful name.
!Pleurotus romellianus Pilát
Saber 2000c
Unclear species. Probably not a member of
Pleurotus.
Psalliota pratensis Schaeff.
Esfandiari 1946
Unclear name, not Hygrocybe (Cuphophyllus)
pratensis as mentioned in MycoBank.
!Psathyrella torpens (Fr.) Konrad & Maubl.
Asef 2007b
Doubtful name. Comparable with P.
panaeoloides R. Maire according to Svrček
(1961).
Rhizomorpha subcorticalis P. Micheli ex Pers.
Buhse 1860
Only rhizomorphs described in old literature
(mostly in 19th cent.). It may represent diverse
fungi (e.g. Armillarias).
Russula sardonia Fr.
Khabiri 1968
Uncertain identification. Growing under Pinus
especially in acid forests.
Stropharia magnivelaris Peck (Leratiomyces
magnivelaris (Peck) Bridge & Spooner)
Fadavi et al. 2013
Unlikely to occur in Iran. Might be
Leratiomyces percevalii (Berk. & Broome)
Bridge & Spooner instead.
!Xerula furfuracea (Peck) Redhead, Ginns & Shoemaker
(Oudemansiella furfuracea (Peck) Zhu L. Yang, G.M.
Muell., G. Kost & Rexer)
Saber 1998
North American species. Very unlikely to
occur in Iran.
RESULTS
Species data
The annotated list of species of agarics and boletes
currently known from Iran is presented in Table 1.
Nineteen species are newly recorded from the country,
based on vouchered specimens all provided with
nrDNA data from basidiomata (Table 3). In situ
basidiomata photographs of selected species are shown
in Fig. 1. Overall, 585 species of agarics and boletes
are recorded from Iran, comprising 556 agarics and 29
bolete species (Fig. 2a).
From a taxonomical viewpoint, the Iranian species
belong to 147 genera and 34 families, distributed in the
six Agaricomycetes orders Agaricales, Russulales,
Boletales, Polyporales, Cantharellales, and
Hymenochaetales (Fig. 2b). The largest order is
Agaricales which encompasses 82% of the species
(Fig. 2b). The three largest families are Agaricaceae
(59 species), Russulaceae (57 species), and
Psathyrellaceae (51 species) (Fig. 3). The family
Polyporaceae is represented with seven agaric species
in Iran, in the genera Lentinus, Neofavolus and
Neolentinus (Fig. 3).

28 Mycologia Iranica - Vol. 7, 2020
Table 3. DNA sequences from Iranian agarics and boletes newly generated in this study.
Species
Voucher
Locality
GenBank
ITS
LSU
Agaricus arvensis
Ghobad-Nejhad 4295
IRAN: Tehran
MT535720
-
Agaricus bitorquis
Ghobad-Nejhad 4284
IRAN: Tehran
MT535709
MT554302
Agaricus iodosmus
Ghobad-Nejhad 4277
IRAN: Tehran
MT535702
MT554295
Agaricus litoralis
Ghobad-Nejhad 4283
IRAN: Semnan
MT535711
MT554304
Agaricus pseudopratensis
Ghobad-Nejhad 4278
IRAN: Tehran
MT535735
MT554325
Agaricus pseudopratensis
Ghobad-Nejhad 3071
IRAN: Tehran
MT535748
-
Agrocybe dura
Ghobad-Nejhad 4286
IRAN: Tehran
MT535714
MT554306
Agrocybe dura
Ghobad-Nejhad 4296
IRAN: Tehran
MT535722
MT554312
Agrocybe dura
Ghobad-Nejhad 4299
IRAN: Tehran
MT535724
MT554314
Agrocybe dura
Ghobad-Nejhad 4325
IRAN: Tehran
MT535732
MT554322
Agrocybe praecox
Ghobad-Nejhad 4261
IRAN: Mazandaran
MT535701
MT554294
Amanita lividopallescens
Ghobad-Nejhad 4019
IRAN: Lorestan
MT535691
-
Conocybe apala
Ghobad-Nejhad 4302
IRAN: Tehran
MT535728
MT554318
Coprinellus micaceus
Ghobad-Nejhad 3186
IRAN: Ilam
MT535694
MT554289
Coprinellus micaceus
Ghobad-Nejhad 3278
IRAN: Kohgiluyeh and Boyer-Ahmad
MT535744
MT554334
Coprinopsis sp.
Ghobad-Nejhad 4282
IRAN: Tehran
MT535708
MT554301
Cortinarius hildegardiae
Ghobad-Nejhad 4202
IRAN: Golestan
MT535704
MT554297
Cortinarius persoonianus
Ghobad-Nejhad 4206
IRAN: Golestan
MT535741
MT554330
Crepidotus subverrucisporus
Ghobad-Nejhad 387
IRAN: East Azerbaijan
MT535745
-
Cuphophyllus virgineus
Ghobad-Nejhad 4191
IRAN: Mazandaran
MT535688
MT554284
Deconica crobula
Ghobad-Nejhad 117
IRAN: East Azerbaijan
MT535747
-
Flammulina velutipes
Ghobad-Nejhad 4305
IRAN: Tehran
MT535715
MT554307
Gymnopilus spectabilis
Ghobad-Nejhad 4207
IRAN: Golestan
MT535703
MT554296
Gymnopus aquosus
Ghobad-Nejhad 4272
IRAN: Mazandaran
MT535700
MT554293
Gymnopus aquosus
Ghobad-Nejhad 4265
IRAN: Mazandaran
MT535699
MT554292
Gymnopus hybridus
Ghobad-Nejhad 3075
IRAN: Ilam
MT535705
MT554299
Gyroporus castaneus
Ghobad-Nejhad 4220
IRAN: Golestan
MT535725
MT554315
Hymenopellis radicata
Ghobad-Nejhad 4204
IRAN: Golestan
MT535683, MT535684
MT554280
Hypholoma fasciculare
Ghobad-Nejhad 4201a
IRAN: Mazandaran
MT535706, MT535707
MT554300
Lactifluus glaucescens
Ghobad-Nejhad 4205
IRAN: Mazandaran
MT535681
MT554278
Lentinus tigrinus
Ghobad-Nejhad 768
IRAN: East Azerbaijan
MT535686
MT554282
Lentinus tigrinus
Ghobad-Nejhad 4326
IRAN: Tehran
MT535692
MT554287
Leucoagaricus leucothites
Ghobad-Nejhad 4276
IRAN: Tehran
MT535696 MT535697
-
Leucoagaricus leucothites
Ghobad-Nejhad 4292
IRAN: Tehran
MT535726
MT554316
Megacollybia platyphylla
Ghobad-Nejhad 4148
IRAN: Mazandaran
MT535698
MT554291
Melanoleuca exscissa
Ghobad-Nejhad 4375
IRAN: East Azerbaijan
MT535742
MT554331
Mycena sp.
Ghobad-Nejhad 134
IRAN: East Azerbaijan
MT535730
MT554320
Mycena sp.
Ghobad-Nejhad 4122
IRAN: Mazandaran
-
MT554298
Mycena xantholeuca
Ghobad-Nejhad 3074
IRAN: Ilam
MT535719
MT554310
Mycena xantholeuca
Ghobad-Nejhad 3077
IRAN: Ilam
MT535733
MT554323
Ossicaulis salomii
Ghobad-Nejhad 4324
IRAN: Tehran
MT535738
MT554327
Pholiota adiposa
Ghobad-Nejhad 600
IRAN: East Azerbaijan
MT535689
MT554285
Pholiota jahnii
Ghobad-Nejhad 757B
IRAN: East Azerbaijan
MT535737
MT554326
Pleurotus cornucopiae
Ghobad-Nejhad 4308
IRAN: Tehran
MT535734
MT554324
Pleurotus eryngii
Ghobad-Nejhad 137
IRAN: East Azerbaijan
MT535679
MT554276
Pleurotus eryngii
Ghobad-Nejhad 1068
IRAN: Ardabil
MT535685
MT554281
Pleurotus ostreatus
Ghobad-Nejhad 759
IRAN: East Azerbaijan
MT535690
MT554286
Pluteus cervinus
Ghobad-Nejhad 4271
IRAN: Mazandaran
MT535687
MT554283
Psathyrella bivelata
Ghobad-Nejhad 4310
IRAN: Tehran
MT535693
MT554288
Psathyrella bivelata
Ghobad-Nejhad 4293
IRAN: Tehran
MT535710
MT554303
Psathyrella bivelata
Ghobad-Nejhad 4303
IRAN: Tehran
MT535727
MT554317
Psathyrella bivelata
Ghobad-Nejhad 71
IRAN: Tehran
MT535729
MT554319
Psathyrella candolleana
Ghobad-Nejhad 4301
IRAN: Tehran
MT535718
MT554309
Psathyrella cf. candolleana
Ghobad-Nejhad 141
IRAN: Tehran
MT535731
MT554321
Psathyrella fatua
Ghobad-Nejhad 3253
IRAN: Kohgiluyeh and Boyer-Ahmad
MT535695
MT554290
Psathyrella hellebosensis
Ghobad-Nejhad 27
IRAN: Semnan
MT535716 MT535717
MT554308
Psathyrella aff. latispora
Ghobad-Nejhad 3322
IRAN: Kohgiluyeh and Boyer-Ahmad
MT535723
MT554313
Pseudoclitocybe cyathiformis
Ghobad-Nejhad 3166
IRAN: Ilam
MT535721
MT554311
Psilocybe sp.
Ghobad-Nejhad 4306
IRAN: Tehran
MT535712, MT535713
MT554305
Rhizomarasmius sp.
Ghobad-Nejhad 4151
IRAN: Mazandaran
MT535736
-
Russula emeticolor
Ghobad-Nejhad 4213
IRAN: Golestan
MT535680
MT554277
Russula emeticolor
Ghobad-Nejhad 4149
IRAN: Mazandaran
MT535682
MT554279
Simocybe centunculus
Ghobad-Nejhad 125
IRAN: East Azerbaijan
MT535746
-
Suillellus luridus
Ghobad-Nejhad 4016
IRAN: Lorestan
MT535740
MT554329
Volvopluteus cf. earlei
Ghobad-Nejhad 4321
IRAN: Tehran
MT535739
MT554328
Xerula pudens
Sohrabi 30619
IRAN: Golestan
MT535743
MT554333

GHOBAD-NEJHAD ET AL.: The gilled fungi and boletes of Iran 29
Fig. 1. In situ photographs from basidiomata of selected species newly recorded in this study. a. Amanita lividopallescens,
Ghobad-Nejhad 4018. b. Conocybe apala, Ghobad-Nejhad 4384. c. Cortinarius hildegardiae, Ghobad-Nejhad 4202. d. Cortinarius
persoonianus, Ghobad-Nejhad 4206. e. Cuphophyllus virgineus, Ghobad-Nejhad 4191. f. Gymnopus aquosus, Ghobad-Nejhad
4272. g. Gymnopus hybridus, Ghobad-Nejhad 3075. h. Gyroporus castaneus, Ghobad-Nejhad 4220. i. Mycena xantholeuca,
Ghobad-Nejhad 3074. j. Russula emeticolor, Ghobad-Nejhad 4213. k. Suillellus luridus, Ghobad-Nejhad 4016. l, m. Psathyrella
bivelata, Ghobad-Nejhad s.n. Photos: Ghobad-Nejhad.

30 Mycologia Iranica - Vol. 7, 2020
Fig. 2. Figures and facts on agarics and boletes in Iran. a. Total number of agaric species vs. bolete species. b. Number of species
in each order. c. Number of species in Hyrcan vs. Zagros regions; ‘other’ means not found in Hyrcan nor in Zagros, but mainly
found in central Iran. d. Number of species with and without nrITS sequence. e. The source of recorded species being from
basidiomata, from environmental samples, or retrieved as plant endophytes. f. The number of species recorded solely in congress
abstracts. g. The Iranian genera with more than 10 species.
Out of a total of 147 genera in Iran, 20 genera have
≥ ten species (Fig. 2g). The three most species-rich
genera in Iran are Russula (43 species), Cortinarius
(26 species), and Inocybe (25 species) (Fig. 2g); these
genera are ectomycorrhizal. Excluded and doubtful
taxa are presented in Table 2. Based on this, 28 species
are excluded from the Iranian mycobiota (Table 2).
Considering the geographic distribution of the
species, we summarized data on species occurring in
the Hyrcan (northern Iran), Zagros (western Iran), and
other regions in Iran (Fig. 2c). Based on our analyses,
the majority (79%) of agaric and bolete species
recorded from Iran reside in the narrow, small area of
Hyrcanian forests (Fig. 2c). Here, Mazandaran
province is at the top, being the most species-rich
province in the country (244 species), followed by
Gilan (143 species) and Golestan provinces (134
species) (Fig. 4). The Zagros region harbors only 13%
of the species (Fig. 2c). In total, 8% of the species have
not been recorded in Hyrcan/Zagros forests, but were
found mainly in central Iran. Tehran province is shown
to have 65 species, while no species is yet known from
Yazd province. For 89 species, distribution data is

GHOBAD-NEJHAD ET AL.: The gilled fungi and boletes of Iran 31
lacking (Fig. 4). Lists of species for each province of
the country are provided in Table 4.
Regarding the situation with the literature records,
our analyses demonstrate that as much as 42% of
records in Iran arise merely from single abstracts
presented in conferences (Fig. 2f, species marked with
! in the Table 1). This means that such species have
appeared in congress abstract only, with no subsequent
evidence to support them; most of these records lack
sufficient characterization, species name authority, or
vouchers. For many of these, modern interpretation of
the names was difficult, if not impossible. We were not
able to trace the vouchers to re-examine them.
Therefore, it is recommended that these records be
considered tentatively for the time being, until more
data become available about their identity.
Fig. 3. Families of agarics and boletes in Iran and the number of species per family. Family color fonts correspond to the color
of the order names in the lower side of the chart.
Fig. 4. Number of agaric and bolete species in each province of Iran. 89 species lack information on province distribution. (Full
list of species for each province is available in Table 4).

32 Mycologia Iranica - Vol. 7, 2020
Table 4. Lists of agarics and boletes for each province of Iran. For more details see Table 1 and the text.
Province
List of species
Alborz
Agaricus arvensis, Agaricus bitorquis, Agaricus litoralis, Armillaria lutea, Armillaria mellea,
Conocybe albipes, !Conocybe subovalis, Coprinellus disseminatus, Coprinellus micaceus,
Coprinellus subimpatiens, Coprinopsis atramentaria, Coprinopsis cinerea, Coprinopsis
ephemeroides, Coprinopsis lagopus, Coprinopsis nivea, Coprinopsis picacea, Flammulina velutipes,
Gymnopus fusipes, Lentinus lepideus, Lentinus tigrinus, Lepiota cristata, Pholiota adiposa, Pholiota
gummosa, Pholiota populnea, Pleurotus eryngii, Pleurotus ostreatus.
Ardabil
Agaricus bisporus, Agaricus campestris, !Entoloma vernum, Pleurotus eryngii.
Bushehr
Agaricus nevoi.
Chaharmahal and Bakhtiari
Agaricus bitorquis, Pleurotus eryngii, Pleurotus fossulatus, Pleurotus nebrodensis.
East Azerbaijan
Agaricus arvensis, Agaricus bisporus, Agaricus bitorquis, Agaricus campestris, Agaricus iodosmus,
Agaricus litoralis, Agrocybe praecox, !Agrocybe tabacina, Amanita pantherina, Amanita vaginata,
Ampulloclitocybe clavipes, Armillaria cepistipes, Armillaria lutea, Armillaria mellea, Boletus edulis,
Butyriboletus fechtneri, Chlorophyllum rhacodes, !Clitocybe nebularis, Clitocybe rufuloalutacea,
!Coprinellus truncorum, Coprinopsis nivea, Coprinopsis picacea, Cortinarius causticus, Cortinarius
erumpens, Cortinarius fluryi, Cortinarius paracephalixus, Cortinarius pluviorum, !Cortinarius
valgus, Cortinarius vespertinus, Crepidotus cesatii, Crepidotus subverrucisporus, Cyanoboletus
pulverulentus, Deconica crobula, Entoloma griseoluridum, Entoloma niphoides, Flammulina
velutipes, !Gymnopus erythropus, !Gymnopus foetidus, !Hebeloma hiemale, !Hodophilus
hymenocephalus, !Hygrophorus mesotephrus, Hymenopellis radicata, Hypholoma fasciculare,
!Hypholoma radicosum, Infundibulicybe geotropa, Infundibulicybe trulliformis, !Inocybe leptocystis,
Laccaria laccata, Lactarius serifluus, Lactifluus volemus, Leccinellum pseudoscabrum, Lentinus
tigrinus, Lepiota castanea, Lepiota cristata, !Lepista nuda, Leucocybe houghtonii, !Leucopaxillus
giganteus, !Macrolepiota mastoidea, Macrolepiota procera, Melanoleuca exscissa, Mycena acicula,
Mycena clavicularis, Neoboletus luridiformis, !Panaeolus fimicola, Panaeolus papilionaceus,
Paralepista flaccida, !Parasola leiocephala, !Paraxerula caussei, Paxillus involutus, Pholiota
adiposa, Pholiota jahnii, Pleurotus eryngii, Pleurotus ostreatus, Pluteus cervinus, Pseudosperma
perlatum, Pseudosperma rimosum, !Russula cyanoxantha, Russula delica, Russula emetica, Russula
lilacea, Russula sororia, Simocybe centunculus, !Stropharia aeruginosa, Suillus collinitus, Suillus
granulatus, !Tricholoma equestre, !Tricholoma orirubens, Tricholoma terreum, !Tubaria
confragosa, Volvopluteus gloiocephalus, Xerocomus subtomentosus, Zhuliangomyces
ochraceoluteus.
Fars
Agaricus bitorquis, Agaricus nevoi, Coprinellus angulatus, Montagnea arenaria, Pleurotus eryngii,
Pleurotus fossulatus, Psathyrella candolleana.
Gilan
Agaricus arvensis, Agaricus iranicus, Agaricus moelleri, !Agaricus pseudolutosus, Agaricus
subrufescens, Agaricus xanthodermus, Agrocybe dura, !Agrocybe paludosa, Agrocybe praecox,
!Agrocybe vervacti, !Alnicola escharioides, Amanita phalloides, Amanita rubescens, Amanita verna,
Armillaria mellea, !Atheniella flavoalba, !Calocybe chrysenteron, !Calocybe ionides, Cantharellus
ferruginascens, !Clitocybe barbularum, !Clitocybe nebularis, !Clitocybe vibecina, !Clitopaxillus
alexandri, Clitopilus prunulus, !Collybia tuberosa, !Conocybe rickenii, Coprinopsis atramentaria,
!Cortinarius cinnamomeus, Craterellus cinereus, Craterellus cornucopioides, Crepidotus
applanatus, Crepidotus caspari, Crepidotus mollis, Cyanoboletus pulverulentus, Cyclocybe
cylindracea, !Entoloma hirtipes, !Entoloma rhodopolium, !Entoloma sericellum, !Flammulaster
gracilis, Flammulina velutipes, !Gymnopus androsaceus, !Gymnopus brassicolens, Gymnopus
dryophilus, !Gymnopus erythropus, !Gymnopus foetidus, Gymnopus fusipes, !Gymnopus inodorus,
!Gymnopus terginus, Hohenbuehelia atrocoerulea, Hohenbuehelia petaloides, Hygrophoropsis
aurantiaca, ! Hygrophorus eburneus, Hygrophorus persoonii, Hymenopellis radicata, Hypholoma
fasciculare, !Hypholoma radicosum, !Hypholoma subericaeum, Inocybe asterospora, Inocybe
dulcamara, Inocybe lilacina, Inocybe paludinella, Inosperma cookei, Inosperma maculatum,
Laccaria amethystina, !Laccaria bicolor, Lactarius deliciosus, Lactarius scrobiculatus, Lactarius
subdulcis, Lactifluus piperatus, Lactifluus vellereus, !Leccinum versipelle, Lentinellus cochleatus,
Lentinus cyathiformis, Lentinus strigosus, Lentinus tigrinus, Lepiota brunneoincarnata, Lepiota
cristata, !Lepista nuda, !Leucoagaricus nympharum, Marasmiellus candidus, !Marasmiellus
confluens, !Marasmiellus peronatus, !Marasmiellus ramealis, !Marasmius atrorubens, !Marasmius
corrugatiformis, !Marasmius epiphyllus, !Marasmius favoloides, !Marasmius ferrugineus,
!Marasmius haematocephalus, !Marasmius oreades, Marasmius rotula, !Marasmius rubroflavus,
Megacollybia platyphylla, Montagnea haussknechtii, Mycena crocata, !Mycena filopes, !Mycena
galericulata, Mycena haematopus, !Mycena inclinata, !Mycena pearsoniana, !Mycena pelianthina,
!Mycena polygramma, !Mycena pura, Mycetinis alliaceus, !Mycetinis scorodonius, Neofavolus
suavissimus, Neolentinus adhaerens, !Omphalina pyxidate, Omphalotus olearius, Panaeolus
campanulatus, Panellus stipticus, Panus conchatus, !Paragymnopus perforans, Paxillus involutus,
Phloeomana speirea, Pholiota gummosa, Pholiota populnea, !Pholiotina striipes, !Pholiotina vexans,
Pleurotellus chioneus, Pleurotus djamor, Pleurotus ostreatus, Pleurotus pulmonarius, !Pluteus
aurantiorugosus, !Pluteus depauperatus, Pseudoclitocybe cyathiformis, Rhodocollybia maculata,
Russula atropurpurea, !Russula atrorubens, !Russula claroflava, !Russula ionochlora, !Russula
ochroleuca, !Russula ochroleucoides, !Russula pectinatoides, !Russula perlactea, !Russula virescens,
Strobilomyces strobilaceus, !Strobilurus tenacellus, Suillus collinitus, !Tricholoma caligatum,
Tricholoma lascivum, !Tricholomopsis formosa, Tubaria furfuracea.
Hamadan
Agaricus arvensis, Conocybe leucopus, Mallocybe terrigena, Pleurotus ostreatus, Psathyrella
clivensis, Psathyrella tephrophylla, Tricholoma scalpturatum.

GHOBAD-NEJHAD ET AL.: The gilled fungi and boletes of Iran 33
Table 4. Continued.
Province
List of species
Golestan
Agaricus arvensis, !Agaricus bresadolanus, !Agaricus brunneolus, Agaricus campestris, !Agaricus
langei, Agaricus moelleri, Agaricus xanthodermus, !Agrocybe acericola, Agrocybe dura, Agrocybe
praecox, Amanita excelsa, Amanita pantherina, Amanita vaginata, Ampulloclitocybe clavipes,
Armillaria lutea, Armillaria mellea, !Arrhenia griseopallida, Battarrea stevenii, Bolbitius reticulatus,
!Bolbitius titubans, !Boletus aestivalis, Cantharellus alborufescens, Cantharellus cibarius,
Cantharellus ferruginascens, Chalciporus piperatus, Clitocybe angustissima, Clitopilus prunulus,
Clitopilus scyphoides, Conocybe albipes, Conocybe juniana, !Conocybe ochracea, Cortinarius
hildegardiae, Cortinarius persoonianus, Crepidotus applanatus, Crepidotus caspari, Crepidotus
crocophyllus, Crepidotus mollis, !Crinipellis scabella, Cyclocybe cylindracea, Cystoderma aureum,
!Dermoloma cuneifolium, Echinoderma asperum, Entoloma incanum, !Entoloma majaloides,
!Entoloma mammosum, Entoloma sinuatum, Flammulaster ferrugineus, Galerina marginata,
Gymnopilus spectabilis, !Gymnopus androsaceus, Gymnopus dryophilus, !Gymnopus inodorus,
Gyroporus castaneus, Hohenbuehelia auriscalpium, Hymenopellis radicata, Hypholoma fasciculare,
Hypsizygus ulmarius, Inocybe erubescens, Inocybe lilacina, Inocybe praetervisa, !Laccaria bicolor,
Lactarius scrobiculatus, Lactifluus piperatus, Lactifluus volemus, !Leccinum scabrum, Lentinellus
ursinus, Lentinellus vulpinus, Lentinus sajor-caju, Lentinus strigosus, Lepiota echinella, !Lepista
irina, Marasmiellus candidus, !Marasmiellus ramealis, Marasmius rotula, Megacollybia platyphylla,
!Melanoleuca subpulverulenta, Mycena crocata, !Mycena filopes, !Mycena galericulata, Mycena
haematopus, !Mycena inclinata, !Mycena metata, Mycetinis alliaceus, !Omphaliaster asterosporus,
Omphalotus olearius, !Panaeolus plantaginiformis, Panellus stipticus, !Paragymnopus perforans,
Paxillus involutus, !Phellorinia herculeana, Phloeomana speirea, Pholiota adiposa, Pholiota
squarrosa, Pleurotellus chioneus, Pleurotus cornucopiae, Pleurotus ostreatus, Pleurotus
pulmonarius, Pluteus cervinus, Pluteus chrysophaeus, !Pluteus depauperatus, !Pluteus punctipes,
Pluteus romellii, Pluteus salicinus, Pluteus thomsonii, Pogonoloma macrocephalum, !Psathyrella
obtusata, Pseudoclitocybe cyathiformis, Pseudosperma perlatum, Rhodotus palmatus, !Rubroboletus
satanas, !Russula alutacea, !Russula carminipes,!Russula cyanoxantha, Russula emetica, Russula
emeticolor, !Russula graveolens, Russula heterophylla, Russula integra, !Russula ochroleuca,
!Russula pectinatoides, !Russula risigallina, !Russula romellii, !Russula xerampelina, !Suillus
bovinus, Tricholoma lascivum, Tricholoma sulphureum, !Tricholoma ustale, Tubaria furfuracea,
!Volvariella bombycina, !Volvariella murinella, !Xerocomellus chrysenteron, Xerocomus
subtomentosus, Xerula pudens.
Hormozgan
Agaricus bisporus, Agaricus devoniensis, Agaricus nevoi.
Ilam
Amanita lividopallescens, Coprinellus micaceus, Gymnopus hybridus, Macrocybe gigantea, Mycena
xantholeuca, Pseudoclitocybe cyathiformis, Pseudosperma rimosum, Suillellus comptus.
Isfahan
Agaricus bitorquis, Armillaria mellea, Coprinellus disseminatus, Coprinellus micaceus, Coprinopsis
atramentaria, !Coprinopsis martini, Flammulina velutipes, !Inocybe corydalina, Inocybe hirtella,
Inosperma bongardii, Lactifluus piperatus (?), Lepiota cristata, !Lepiota lilacea, Leucoagaricus
leucothites, !Parasola miser, Pholiota adiposa, Pleurotus eryngii, Pluteus romellii.
Kerman
Montagnea arenaria.
Kermanshah
Agaricus devoniensis, Agaricus nevoi, Agrocybe dura, Agrocybe ochracea, !Agrocybe vervacti,
Conocybe albipes, Conocybe dunensis, Coprinopsis atramentaria, Coprinopsis urticicola, Coprinus
comatus, Coprinus sterquilinus, Entoloma griseorubellum, Inocybe subnudipes, Inocybe terrifera,
Lentinus tigrinus, Leucoagaricus carneifolius, !Panaeolus olivaceus, !Pholiota highlandensis,
Pleurotus ostreatus, Pleurotus pulmonarius, Pluteus cinereofuscus, Pluteus nanus, Pogonoloma
macrocephalum, Psilocybe atrobrunnea, Stropharia coronilla, Suillellus comptus, Volvariella
volvacea, Volvopluteus gloiocephalus.
Khuzestan
Agaricus bisporus, Agaricus bitorquis, Agaricus nevoi, Armillaria mellea, Lepiota anthomyces,
Lepiota metulispora, Leucoagaricus serenus.
Kohgiluyeh and Boyer-Ahmad
Agaricus bitorquis, Agaricus nevoi, Coprinellus micaceus, !Hemileccinum impolitum, Lentinus
tigrinus, Pleurotus eryngii, Pleurotus fossulatus, Pleurotus nebrodensis, Psathyrella fatua.
Kurdistan
Agaricus bisporus, Pleurotus eryngii, Pleurotus fossulatus, Pleurotus nebrodensis.
Lorestan
Agaricus bisporus, Amanita umbrinolutea, Amanita lividopallescens, Coprinopsis picacea, Lentinus
tigrinus, ! Leratiomyces squamosus, !Melanoleuca cognata, Pleurotus eryngii, Suillellus luridus.
Markazi
Agaricus bisporus, Agaricus bitorquis, Armillaria mellea.
North Khorasan
Pleurotus ostreatus.
Qazvin
Agrocybe dura, Armillaria mellea.
Qom
Montagnea arenaria.
Razavi Khorasan
Agaricus bitorquis, Armillaria mellea, Clitocybe rufuloalutacea, Conocybe albipes, Coprinellus
disseminatus, Coprinellus micaceus, Coprinopsis atramentaria, Coprinopsis picacea, Coprinus
comatus, Cyclocybe cylindracea, Deconica crobula, Entoloma incanum, Flammulina velutipes,
Hypholoma fasciculare, Lepiota cristata, Lepista saeva, Macrolepiota procera, Montagnea arenaria,
Mycena galopus, !Omphalina mutila, Panaeolus rickenii, Parasola plicatilis, Pholiota adiposa,
Pleurotus ostreatus, Psathyrella candolleana, !Psathyrella obtusata, Tricholoma fulvum, Tricholoma
sulphureum, Volvariella hypopithys.
Semnan
Agaricus litoralis, Armillaria mellea, Montagnea arenaria, Psathyrella hellebosensis.
Sistan and Baluchestan
! Conocybe tenera, Coprinellus disseminatus, Montagnea arenaria, Neofavolus suavissimus,
Psathyrella candolleana.
South Khorasan
Pleurotus eryngii, Psathyrella obtusata.

34 Mycologia Iranica - Vol. 7, 2020
Table 4. Continued.
Province
List of species
Mazandaran
Cortinarius diasemospermus (?), Cortinarius parvannulatus (?), Cortinarius vernus (?), Inocybe
amethystine (?), Inocybe castaneicolor (?), Inocybe decemgibbosa (?), Inocybe flocculosa (?), Inocybe
godeyi (?), Inocybe langei (?), Inocybe mixtilis (?), Russula luteotacta (?), Russula persicina (?),
Suillus granulatus (?), Tricholoma scalpturatum (?), Tricholoma terreum (?), Agaricus arvensis,
Agaricus campestris, Agaricus litoralis, Agaricus moelleri, Agaricus phaeolepidotus, Agrocybe
praecox, !Amanita atkinsoniana, Amanita battarrae, Amanita caesarea, Amanita ceciliae, Amanita
crocea, Amanita eliae, Amanita excelsa, Amanita gemmata, Amanita pantherina, Amanita phalloides,
Amanita strobiliformis, Amanita vaginata, Amanita verna, Ampulloclitocybe clavipes, Armillaria
borealis, Armillaria lutea, Armillaria mellea, Asterophora lycoperdoides, !Aureoboletus gentilis,
!Baeospora myosura, Bolbitius reticulatus, !Calocybe persicolor, Cantharellus alborufescens,
Cantharellus ferruginascens, !Clitocybe metachroa, !Clitocybe nebularis, !Clitocybe phyllophila,
Clitopilus prunulus, !Conocybe macrocephala, !Conocybe microspora, !Conocybe pilosella,
!Contumyces rosellus, Coprinellus disseminatus, Coprinellus micaceus, Coprinellus radians,
Coprinopsis atramentaria, Coprinopsis brunneofibrillosa, Coprinopsis cinerea, Coprinopsis
lagopus, Coprinopsis urticicola, Coprinus comatus, Cortinarius decipiens, Cortinarius vibratilis,
Cortinarius violaceus, Craterellus cinereus, Craterellus cornucopioides, Crepidotus applanatus,
Crepidotus caspari, Crepidotus cesatii, Crepidotus crocophyllus, Crepidotus mollis, Cuphophyllus
virgineus, Cyclocybe cylindracea, Cystoderma aureum, !Delicatula integrella, Echinoderma
asperum, !Entoloma clypeatum, !Entoloma rhodopolium, Entoloma sinuatum, !Flammulaster
erinaceellus, !Flammulaster gracilis, Flammulina velutipes, Galerina marginata, !Gymnopus
androsaceus, Gymnopus aquosus, Gymnopus dryophilus, !Gymnopus erythropus, !Gymnopus
foetidus, !Gymnopus inodorus, !Gymnopus terginus, !Hemimycena cucullata, Hohenbuehelia
auriscalpium, Hohenbuehelia petaloides, !Hydropus marginellus, !Hygrocybe acutoconica,
!Hygrocybe chlorophana, Hygrophoropsis aurantiaca, !Hygrophorus persoonii, Hymenopellis
radicata, !Hypholoma capnoides, Hypholoma fasciculare, Hypholoma lateritium, Infundibulicybe
gibba, Inocybe asterospora, Inocybe dulcamara, Inocybe geophylla, Inocybe lilacina, Inocybe
napipes, Inocybe tabacina, Inosperma adaequatum, Inosperma bongardii, Inosperma cookei,
Inosperma maculatum, !Laccaria bicolor, Laccaria laccata, !Laccaria tortilis, Lacrymaria
lacrymabunda, Lactarius rubrocinctus, Lactarius subdulcis, Lactarius tabidus, Lactifluus
glaucescens, Lactifluus vellereus, Leccinellum pseudoscabrum, Lentinus lepideus, Lentinus strigosus,
Lentinus tigrinus, Lepiota castanea, Lepiota leprica, Lepiota subincarnata, Lepista saeva,
Leucoagaricus americanus, !Leucoagaricus badhamii, Leucoagaricus holospilotus, Leucoagaricus
leucothites, !Leucocoprinus magnusianus, !Leucocybe candicans, !Leucopaxillus compactus,
!Leucopaxillus pinicola, !Lyophyllum atratum, !Lyophyllum baeospermum, !Macrolepiota permixta,
Macrolepiota procera, Marasmiellus candidus, !Marasmiellus peronatus, !Marasmiellus ramealis,
!Marasmius atrorubens, !Marasmius corrugatiformis, !Marasmius favoloides, !Marasmius
ferrugineus, !Marasmius haematocephalus, Marasmius rotula, !Marasmius rubroflavus, Marasmius
wynneae, Megacollybia platyphylla, Mycena crocata, !Mycena galericulata, Mycena haematopus,
!Mycena inclinata, !Mycena rapiolens, !Mycena sanguinolenta, Mycenastrum corium, !Mycenella
salicina, Mycetinis alliaceus, !Mycetinis scorodonius, Neolentinus adhaerens, Omphalotus olearius,
Ossicaulis lignatilis, Panaeolus acuminatus, Panaeolus papilionaceus, Panaeolus teutonicus,
Panellus stipticus, Panaeolus semiovatus, Panus conchatus, !Paragymnopus perforans, Paralepista
flaccida, Parasola plicatilis, Paxillus involutus, Phloeomana speirea, Pholiota adiposa, Pholiota
squarrosa, Pholiota squarrosoides, !Pholiotina arrhenii, !Phyllotopsis nidulans, Pleurotus
calyptratus, Pleurotus cornucopiae, Pleurotus dryinus, Pleurotus elongatipes, Pleurotus ostreatus,
Pleurotus pulmonarius, !Pluteus aurantiorugosus, Pluteus cervinus, Pluteus chrysophaeus, !Pluteus
depauperatus, Pluteus exiguus, Pluteus leoninus, Pluteus nanus, Pluteus salicinus, Pluteus
semibulbosus, Pluteus thomsonii, Psathyrella candolleana, Psathyrella multipedata, !Psathyrella
obtusata, !Psathyrella spadiceogrisea, Pseudoclitocybe cyathiformis, Pseudosperma rimosum,
Resupinatus applicatus, !Rhodocollybia prolixa, !Rubroboletus satanas, !Russula acetolens, !Russula
anthracina, Russula atropurpurea, !Russula brunneoviolacea, Russula emetica, Russula emeticolor,
!Russula farinipes, Russula foetens, !Russula grisea, Russula integra, Russula olivacea, !Russula
pectinata, !Russula puellaris, !Russula queletii, !Russula silvestris, !Russula torulosa, !Russula
virescens, Sarcomyxa serotina, !Sphagnurus paluster, !Strobilurus esculentus, !Strobilurus
tenacellus, Suillus collinitus, Tricholoma acerbum, !Tricholoma psammopus, Tricholoma
cingulatum, !Tricholoma ustale, !Tricholoma ustaloides, !Tricholoma vaccinum, !Tubaria conspersa,
!Tylopilus felleus, Volvariella iranica, Volvariella hypopithys, !Xerocomellus pruinatus, !Xerocomus
moravicus.
Tehran
Agaricus arvensis, Agaricus bisporus, Agaricus bitorquis, Agaricus iodosmus, Agaricus pseudopratensis,
Agrocybe dura, !Agrocybe pediades, !Agrocybe pusiola, Battarrea stevenii, !Butyriboletus appendiculatus,
!Calocybe carnea, !Calocybe gambosa, Conocybe albipes, Conocybe apala, !Conocybe ochracea,
!Conocybe rickenii, !Conocybe tenera, Coprinellus angulatus, Coprinellus disseminatus, Coprinellus
flocculosus, Coprinellus micaceus, Coprinopsis atramentaria, Coprinopsis brunneofibrillosa, Coprinopsis
cinerea, Coprinopsis lagopides, Coprinopsis macrocephala, Coprinopsis patouillardii, Coprinopsis
scobicola, Coprinus comatus, !Flammulaster granulosa, Flammulina velutipes, !Gymnopus erythropus,
Lentinus tigrinus, Leucoagaricus leucothites, Leucoagaricus roseoalbus, !Marasmius atrorubens,
!Marasmius corrugatiformis, !Marasmius favoloides, !Marasmius ferrugineus, !Marasmius
haematocephalus, !Marasmius rubroflavus, !Melanoleuca graminicola, !Melanoleuca grammopodia,
!Melanoleuca strictipes, !Melanoleuca subpulverulenta, Mycenastrum corium, !Myxomphalia maura,
Ossicaulis salomii, Parasola plicatilis, Pholiota adiposa, Pholiota populnea, Pleurotus cornucopiae,
Pleurotus eryngii, Pleurotus ostreatus, Psathyrella bivelata, Psathyrella candolleana, !Psilocybe
cyanescens, !Psilocybe serbica, Suillus collinitus, Suillus granulatus, !Tricholoma argyraceum,
!Tricholoma robustum, Tricholoma terreum, Tubaria furfuracea, !Tubaria pallidospora.

GHOBAD-NEJHAD ET AL.: The gilled fungi and boletes of Iran 35
Table 4. Continued.
Province
List of species
West Azerbaijan
Agaricus arvensis, Agaricus bisporus, Agaricus bitorquis, !Agaricus depauperatus, !Agaricus
gennadii, !Agrocybe acericola, Agrocybe dura, Agrocybe praecox, Amanita phalloides, Armillaria
mellea, !Clitocybe diatreta, Coprinellus micaceus, Coprinopsis atramentaria, Lepiota helveola,
Leucoagaricus americanus, !Marasmius oreades, Pholiota adiposa, Pleurotus cornucopiae,
Pleurotus eryngii, Pleurotus ostreatus, Pluteus nanus, Psathyrella candolleana, !Psathyrella
spadiceogrisea, Saproamanita codinae, !Stropharia melanosperma, Suillus collinitus, !Tubaria
furfuracea, Volvariella volvacea.
Yazd
_
Zanjan
Agaricus arvensis, Lepista saeva.
province not specified
Butyriboletus pseudoregius, !Chlorophyllum brunneum, !Coprinellus domesticus, !Coprinellus
impatiens, !Coprinellus silvaticus, !Coprinellus xanthothrix, !Coprinopsis friesii, !Coprinopsis
gonophylla, !Coprinopsis marcescibilis, Coprinopsis sclerotiger, Cortinarius bivelus, Cortinarius
caesiocortinatus, Cortinarius casimirii, Cortinarius cinnabarinus, Cortinarius cotoneus, Cortinarius
diasemospermus, Cortinarius ferrugineovelatus, Cortinarius hinnuleus, Cortinarius infractus,
Cortinarius olivaceofuscus, Cortinarius parvannulatus, Cortinarius uraceonemoralis, Cortinarius
vernus, Craterellus tubaeformis, Entoloma subcollariatum, !Flammula alnicola, !Galerina
hypnorum, !Galerina mniophila, !Galerina pumila, !Galerina sphagnorum, !Gymnopilus penetrans,
!Hebeloma birrus, !Hebeloma crustuliniforme, Hebeloma incarnatulum, !Hebeloma mesophaeum,
Hebeloma sinapizans, !Homophron spadiceum, Imleria badia, Inocybe amethystina, Inocybe
castaneicolor, Inocybe cincinnata, Inocybe decemgibbosa, Inocybe flocculosa, Inocybe godeyi,
Inocybe hirtella, Inocybe huijsmanii, Inocybe langei, Inocybe mixtilis, Inocybe mystica, Inocybe
pusio, Lactarius acris, Lactarius circellatus, Lactarius fulvissimus, Lactarius zonarius, !Lepiota
felina, !Lepiota micropholis, !Lepiota subalba, !Leucocoprinus birnbaumii, !Leucocoprinus
brebissonii, !Leucocoprinus cepistipes, !Macrolepiota excoriata, !Panaeolus speciosus, !Parasola
auricoma, !Parasola hemerobia, !Phaeomarasmius erinaceus, !Phaeonematoloma myosotis,
!Pholiota astragalina, !Pholiota scamba, !Pholiota spumosa, !Pholiotina aporos, !Pluteus pellitus,
!Pluteus petasatus, !Pluteus umbrosus, !Psathyrella laevissima, !Psathyrella microrhiza,
!Psathyrella pennata, !Psathyrella piluliformis, !Psathyrella prona, !Psathyrella pseudogracilis,
!Psathyrella squamosa, !Psilocybe coprophila, Russula luteotacta, Russula persicina, Russula rosea,
Russula solaris, Russula versicolor, Russula veternosa, Russula vinosopurpurea, !Volvariella pusilla.
Molecular study
In this study, a total number of 128 new nrDNA
sequences was obtained, comprising 70 sequences
from the ITS barcode region, and 58 sequences from
the nrLSU (Table 3). The scrutiny through public
databases GenBank and UNITE was also performed to
extract the available DNA sequences of Iranian agarics
and boletes and selected ITS accession numbers were
inserted in Table 1 after careful checking. Our analyses
show that in total, only 24% of Iranian agarics and
bolete species have at least one publically available
nrITS sequence, while 76% still lack any DNA
sequences (Fig. 2d). We also show that 64 species of
agarics and boletes in Iran are known from
environmental sequences of ectomycorrhizal root tips
(Fig. 2e). Moreover, nine species have been recovered
as plant endophyte (Fig. 2e).
DISCUSSION
In this study, we presented a first annotated
checklist of 585 species of agarics and boletes in Iran.
To our current knowledge, perhaps except for Agaricus
iranicus which is currently known only from Iran,
there is no agaric nor bolete species endemic to the
country. Also no lichenized agaric is yet known in Iran.
The ITS region has been a favored and practically
suitable DNA barcode marker for identification of
mushrooms (Dentinger et al. 2011). Here, we obtained
70 new ITS sequences, and have also made a full
scrutiny of the sequences of Iranian agarics and boletes
available in GenBank and UNITE. We tried to contrast
environmental sequences against well-identified fungi
or the Species Hypothesis. Selected verified ITS
sequences were reported in Table 1. Currently, only
24% of the Iranian agarics and boletes have at least one
reliable ITS sequence in the public databases.
Application of sequence-based records in
checklists and taxonomy studies is not straightforward.
The majority of the extracted GenBank/UNITE
sequences in our study are from environmental
samples (Bahram et al. 2012 and 2013). Many include
insufficiently identified sequences or lack vouchered
basidiomata. Considering fruitbody-based sequences
in GenBank, we also noticed that a perceptible number
of them suffer from a wrong interpretation of Basic
Local Alignment Search Tool (BLAST), with medium
to large ignorance of standards of molecular
identification. In most cases, we carefully did a re-
BLAST, to correct for the identifications.
While sequences from environmental samples are
highly important in ecological studies, using them in
regional species checklists and taxonomy studies is
still not routine. Interestingly, environmental samples
(from ectomycorrhizal root tips) added 64 more
species to our checklist.
Noteworthily, 42% of agaric and bolete species in
Iran (248 out of 585 species) arise merely from
congress abstracts, with no other supporting data in
other sources. Usually, abstracts presented in
congresses are intended as informal announcements of
scientific research and are not regarded citable by most
peer journals. The same also applies to university
theses, dissertations, and research reports. Data
including biodiversity records in such informal
publications are more reliable when they are

36 Mycologia Iranica - Vol. 7, 2020
subsequently supported by other types of evidence and
published in peer reviewed journals or other formal
publications. We noticed that many of these records
lack sufficient characterization, species authority, or
vouchers; so, they are considered here as tentative
records. We preferred to keep these records in our
checklist, to encourage more research on them, but
recommend them to be used with caution. In the
present work, we were able to confirm the presence of
some of the previously recorded species by new
specimens or by DNA sequences.
Recent studies highlight that climate change will be
a serious threat for biodiversity in Iran due to severe
habitat fragmentation and destruction, and land-use
alteration, specifically in the North West, West
(Zagros), and central parts of Iran (Yousefi et al. 2019).
Although no studies have been done to test the impact
of climate change on fungi in Iran, the range reductions
predicted for some plant species (reviewed by Yousefi
et al. 2019) might also affect their associated fungi,
more notably ectomycorrhizae and wood-inhabiting
macromycetes. It is highly probable that many species
will go extinct before they are recorded. In Iran, agaric
specialists are rare, while forest decline is intensive,
which makes it even more imperative to record the
agarics and boletes of the country. Undoubtedly,
standard DNA barcodes would promote identification
and verification of agarics and boletes in Iran; this
would close the gap in the empirical knowledge of
these fungi.
The above discussed problems of unverifiable
literature records in congress abstracts, and the need to
increase expertise in fungal molecular identification,
are not limited to agarics and boletes but are also
affecting other fungal groups in Iran. Tackling these
problems would ultimately promote mycology and
fungal conservation in Iran. Therefore, general
recommendations are given to the mycological society
in Iran, to prevent dissemination of low quality and
ambiguous species records.
General recommendations:
• Biodiversity records in congress abstracts to be
taken as provisional only, and not treated
unequivocally. These records need to be backed
up with proper citability and sufficient
characterization.
• The specimens studied in dissertations, articles,
etc. to be deposited in at least one of the Iranian
herbaria registered in Index Herbariorum.
Samples with unknown or inaccessible deposition
should be ignored by the scientific society.
• For DNA studies, at least two primers should be
used for amplification and sequencing of any
given DNA region and the quality of obtained
sequences’ electropherograms carefully checked.
Sequences resulting from a single primer, or with
low quality trace files may cause serious
misinterpretations in BLAST searches.
• In molecular identification, it is necessary to
carefully follow standard routines. Exemplar
procedures have been described by e.g. Nilsson et
al. (2012), Kõljalg et al. (2013), and Ryberg et al.
(2008).
• Besides BLASTn, to make use of the UNITE
platform to search for the best matching well-
identified fungal taxon (Species Hypothesis).
Conclusions
This study provides a preliminary account on 585
species of agarics and boletes in Iran, with about half
of them being regarded as tentative records. It fills in
some of the gaps in our knowledge of these important
groups of macrofungi in Iran and provides a
framework for biodiversity and phylogenetic studies.
Up to now, no taxonomic monograph/revision studies
have been done for Iranian agarics and boletes, and
among large species-rich genera in Iran (Fig. 2g), only
the genus Agaricus has been taxonomically and
phylogenetically studied (Mahdizadeh et al. 2017a,
2017b, 2016). Therefore, for the time being it is
difficult to figure out the actual number of species of
agarics and boletes in Iran.
The Hyrcanian region comprises only ca. 5% of the
total area of Iran, yet a large proportion of the Iranian
agarics and boletes (79%) come from this region. This
pattern is similar for other macrobasidiomycetes of
Iran such as aphyllophoroid fungi (Ghobad-Nejhad &
Hallenberg 2012). Macrofungi are vital members of
forest ecosystems warranting their persistence and
conservation. This accentuates the importance of
taking immediate habitat protection strategies in the
narrow, fragile Hyrcanian forests of Iran.
ACKNOWLEDGEMENTS
The authors thank Else C. Vellinga (Berkeley,
USA) for providing comments and input in the use of
names of taxa. Jacob Heilmann-Clausen (Copenhagen,
Denmark) is thanked for some taxonomic notes and
suggestions. Mohammad Sohrabi (Tehran, Iran) is
gratefully acknowledged for field surveys accompany.
This work has been supported by the Center for
International Scientific Studies & Collaboration
(CISSC), Ministry of Science, Research and
Technology.
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RussulaCortinariusInocybe
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
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 :
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Email: ghobadnejhad@irost.ir
