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1. Introduction
Corticioid fungi are homobasidiomycete fungi that form resupi-
nate, eused, crust-like basidiomata, mostly on woody substrates.
Hymenophore congurations vary widely (smooth, odontoid, rad-
uloid, hydnoid, merulioid, and poroid) within genera and even
within several species. The corticioid fungi belong to Agaricomy-
cotina in Basidiomycota, and they have long been grouped in a
single family, Corticiaceae (sensu lato). Recent advances in se-
quencing technology, however, have permitted the gathering of
DNA sequence data for many basidiomycetes, including corticioid
fungi, and molecular phylogenetic analyses have revealed that cor-
ticioid fungi are polyphyletic (K.-H. Larsson, E. Larsson, & Kõljalg,
2004; Binder et al., 2005). Hibbett et al. (2007) proposed a compre-
hensive phylogenetic classication of Basidiomycota in which
Agaricomycotina contains 3 classes and 21 orders. In this classica-
tion, corticioid fungi were categorized into 13 orders: Agaricales,
Atheliales, Boletales, Cantharellales, Corticiales, Dacrymycetales,
Gloeophyllales, Gomphales, Hymenochaetales, Polyporales, Russu-
lales, Thelephorales, and Trechisporales. Subsequently, on the basis
of a six-locus nuclear dataset, Binder, Larsson, Matheny, and Hib-
bett (2010) introduced two new orders; Amylocorticiales—sister to
Agaricales—and Jaapiales, which contains only the genus Jaapia
Bres., comprising two resupinate species. Furthermore, phyloge-
netic analyses indicate that several corticoid genera are polyphylet-
ic: Aleurodiscus Rabenh. ex J. Schröt. (Wu, Hibbett, & Binder,
2001), Gloeocystidiellum Donk (E. Larsson & K.-H. Larsson, 2003),
Hyphoderma Wallr. (K.-H. Larsson, 2007), Hyphodontia J. Erikss.
(Binder et al., 2005; K.-H. Larsson et al., 2006), Leucogyrophana
Pouzar (Jarosch & Besl, 2001), Phanerochaete P. Karst. (De Koker,
Nakasone, Haarhof, Burdsall, & Janse, 2003; Wu, Nilsson, Chen,
Yu, & Hallenberg, 2010), Phlebia Fr. (Kuuskeri, Mäkelä, Isotalo,
Oksane, & Lundell, 2015) and Sistotrema Fr. (Moncalvo et al.,
2006). On the basis of these molecular phylogenetic analyses, each
of these genera has been subdivided into several genera, some of
which are new.
Most corticioid fungi live on woody substrates such as fallen
trunks, branches, and twigs; they decompose cellulose, hemicellu-
loses, and lignin. Ectomycorrhizal associations have been demon-
strated in several corticioid genera, such as Amphinema P. Karst.,
Piloderma Jülich, Sistotrema, and Tomentella Pers. ex Pat. (Dahl-
berg, Jonsson, & Nylund, 1997; Erland & Taylor, 1999; Kõljalg,
Tammi, Timonen, Agerer, & Sen, 2002; Nilsson, K.-H. Larsson, E.
Larsson, & Kõljalg, 2006). Thus, in forest ecosystems, they play an
important role not only in nutrient recycling but also in tree
growth. Some corticioid species are economically important. Sever-
al species—particularly Phanerochaete chrysosporium Burds.—are
being used in basic and applied researches on lignin degradation,
e.g., in pulping, bleaching, and converting lignin to useful products
(Kirk & Farrell, 1987). Corticioid fungi also include several plant
pathogens, such as Athelia rolfsii (Curzi) Tu & Kimbrough (Tu &
Kimbrough, 1978) and Chondrostereum purpureum (Pers.) Pouzar
(Pouzar, 1959), the causal agents of southern Sclerotium rot and
silver-leaf disease, respectively. Some species occur on bedlogs
used for cultivation of shiitake mushroom, Lentinula edodes
Taxonomy of corticioid fungi in Japan: Present status and future prospects
Nitaro Maekawa*
Fungus/Mushroom Resource and Research Center, Faculty of Agriculture, Tottori University, 4-101 Koyama, Tottori 680-855, Japan
Review
ABSTRACT
Corticioid fungi form effused basidiomata with various hymenophore configurations, mostly on woody substrates, and they are present-
ly classified into 15 orders in Agaricomycotina in the phylum Basidiomycota. In this review, the corticioid fungi of Japan are treated on
the basis of the current classification system, and all currently known species are listed and classified by order. Japanese corticioid fungi
number 442 species in 160 genera belonging to 14 orders. Analysis of the distribution of Japanese corticioid species reveals that 375
species (85% of the total) have wide distributions, and that the Japanese corticioid fungal flora is composed of species with diverse pat-
terns of distribution. In the subtropical regions, 146 corticioid species belonging to 77 genera are listed, including 100 species from the
Nansei Islands and 76 from the Ogasawara Islands. Although these two island groups are located at similar latitudes, they share only 30
corticioid species—21% of the total reported from the subtropical regions. Along with this summary of the current status, this study
presents future directions for the taxonomic research of the Japanese corticioid fungi.
Keywords: Agaricomycetes, Basidiomycota
Article history: Received 9 April 2020, Revised 4 October 2021, Accepted 5 October 2021, Available online 26 October 2021.
* Corresponding author.
E-mail address: kin-maek@tottori-u.ac.jp
This is an open-access paper distributed under the terms of the Creative Commons Attribution-NonCommercial-NoDerivative 4.0 international license
(CC BY-NC-ND 4.0: https://creativecommons.org/licenses/by-nc-nd/4.0/).
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(Berk.) Pegler, reducing fruiting body yield (Maekawa & Arita,
1984).
In this review, my primary purpose is to evaluate Japanese corti-
cioid fungi on the basis of the current classication system. Second,
I summarize the bioclimatic features and geographical distribution
patterns of the species making up the Japanese corticioid fungal
ora. Third, I discuss the current status of taxonomic research on
Japanese corticioid fungi, including reclassications based on the
phylogenetic analysis of combined morphological and molecular
data. Finally, I discuss future directions for taxonomic studies of
Japanese corticioid fungi.
2. History of taxonomic research on Japanese corti-
cioid fungi by using morphological features
Hennings (1901) wrote the rst report on Japanese corticioid
fungi. Subsequently, many species were reported by Yasuda (1909,
1912, 1913, 1914a, 1914b, 1918a, 1918b, 1918c, 1919a, 1919b, 1920a,
1920b, 1921a, 1921b, 1923a, 1923b), Lloyd (1920, 1921, 1922), Ito
(1929a, 1929b), and Ito and Imai (1940). The comprehensive publi-
cation “Mycological Flora of Japan Vol. II (4)” by Ito (1955) re-
viewed the species of corticioid fungi that had been recorded in
Japan up to that time. Subsequently, Japanese mycologists have
classied many corticioid species on the basis of morphological
features (Aoshima & Hayashi, 1964; Hayashi & Aoshima, 1964,
1966; Aoshima & Furukawa, 1966a, 1966b; Furukawa & Aoshima,
1966; Hayashi, 1969; Kobayashi, 1971; Furukawa, 1974; Hayashi,
1974), placing them in genera such as Corticium Pers., Odontia
Pers., and Peniophora Cooke on the basis of the Friesian system
(Fries, 1874). Parmasto (1968) introduced a new system for classi-
cation of corticioid fungi, and Japanese species were reclassied
accordingly (Maekawa, 1993b, 1994). New and previously unre-
ported Japanese species continued to be described (Maekawa, Ari-
ta, & Hayashi, 1982; Maekawa & Arita, 1984; Maekawa, 1987, 1990,
1991a, 1991b, 1992, 1993a, 1997a, 1997b, 1998, 1999, 2000a, 2000b,
2003; Núñez & Ryvarden, 1997; Nakasone & Gilbertson, 1998;
Maekawa & Hasebe, 2002; Maekawa & Nordén, 2002; Suhara,
Maekawa, Kubayashi, Sakai, & Kondo, 2002; Maekawa, Suhara,
Kinjo, & Kondo, 2003; Hjortstam & Ryvarden, 2004; Maekawa, Su-
hara, Kinjo, Kondo, & Hoshi, 2005; Wu, Wang, & Yu, 2010; Suhara,
Maekawa, Ushijima, Kinjo, & Hoshi, 2010; Suhara, Maekawa, &
Ushijima, 2011; Chen, Wu, & Chen, 2017, 2018; Ushijima & Maeka-
wa, 2018; Ushijima, Sotome, & Maekawa, 2019; Yagame & Maeka-
wa, 2019; Maekawa et al., 2020; Yurchenko, Wu, & Maekawa, 2020).
The currently known species of Japanese corticioid fungi are listed
in Table 1; there are 442 species in 160 genera belonging to 14 or-
ders. Among the previously reported corticioid fungi, several new
taxa were subsequently placed in synonymy with other taxa
(Maekawa, 1993b), and these taxa were excluded from Table 1.
Phlebiella athelioides N. Maek. (1993b) has been treated as an inval-
id name because Phlebiella P. Karst. is not regarded as a validly
published genus (Duhem, 2010), and the species referred to that
genus have been transferred to Xenasmatella Oberw. Morphologi-
cal characters clearly place P. athelioides in Xenasmatella. There-
fore, the following combination is proposed: Xenasmatella atheli-
oidea (N. Maek.) N. Maek., comb. nov., MycoBank no. MB840571
(Basionym: Phlebiella athelioides N. Maek., Reports of the Tottori
Mycological Institute, 31: 124, 1993). In addition, a few species re-
ported as Japanese corticioid fungi—e.g., Gloeopeniophora auranti-
aca (Pers.) J. (Bres.) Höhn. & Litsch. (Hayashi, 1974), Peniophora
farlowii Burt (Hayashi, 1974), Phlebia rufa (Pers.) M.P. Christ.
(Maekawa, 1993b), and Xylodon papillosa (Fr.) Riebesehl, Yurchen-
ko & Langer (as Odontia papillosa (Fr.) P. Karst) (Furukawa,
1974)—were excluded from this table because their voucher speci-
mens were revealed to have been misidentied (Maekawa, 1993b,
2000b). On the other hand, 55 species (asterisks, Table 1) previous-
ly unreported in Japan were added, and their data were shown in
Supplementary Table S1.
3. Mycofloristics of the corticioid fungi in Japan
3.1. Diversity of the Japanese corticioid fungi
The Japanese Archipelago extends over approximately 3,000 km
from northeast to southwest between lat. 24°N and 46°N, and it has
many high mountains >2,000 m in altitude. Its climate encompass-
es subarctic, cool-temperate, warm-temperate, and subtropical
zones. The mycooral diversity of Agaricales in Japan reects this
geographical and climatic variation (Hongo & Yokoyama, 1978).
Bioclimatically, the Japanese corticioid fungi are composed of spe-
cies distributed in (A) subarctic to temperate zones (141 species,
32% of the total), (B) temperate to subtropical zones (252 species,
57% of the total), and (C) the subtropical zone (49 species, 11% of
the total) (see footnote to Table 1). According to global distribution
records, 105 species of Japanese corticioid fungi have been record-
ed on ve continents, and 79, 92, and 99 species have been recorded
on four, three, and two continents, respectively. Thus a total of 375
species—85% of known Japanese corticioid species—are distribut-
ed on multiple continents. The remaining 67 species are reported
only from Eurasia, and of these 22 species (5% of the total) are en-
demic to Japan and 33 species (7% of the total) are distributed only
in Far East Asia. Many species of Japanese corticioid fungi there-
fore have a wide global distribution, and Japanese corticioid fungi
have diverse distribution patterns.
Analysis of the distributions of the Japanese corticioid species
accepted in the present study reveals that they can be divided into
12 groups: (a) cosmopolitan (Africa, Eurasia, North America, Oce-
ania, and South America), (b) Northern Hemisphere (Eurasia and
North America), Africa, and Oceania, (c) Northern Hemisphere,
Africa, and South America, (d) Northern Hemisphere and Africa,
(e) Northern Hemisphere and Oceania, (f) Northern Hemisphere
and South America, (g) Northern Hemisphere, (h) Eurasia, (i) Far
East Asia and North America, (j) Far East Asia, (k) Japan (endem-
ic), and (m) others. The main distribution patterns and their repre-
sentative species are described below (see footnote to Table 1).
(a) Cosmopolitan: 112 species. Most of these species have been
collected from both temperate and (sub)tropical regions. Represen-
tative species include Amphinema byssoides (Pers.) J. Erikss., Chon-
drostereum purpureum (Pers.) Pouzar, Dacryobolus sudans (Alb. &
Schwein.) Fr., Laxitextum bicolor (Pers.) Lentz, Peniophorella pu-
bera (Fr.) P. Karst., Phanerochaete sordida (P. Karst.) J. Erikss. &
Ryvarden, Scytinostroma portentosum (Berk. & M.A. Curtis) Donk,
Subulicystidium longisporum (Pat.) Parmasto, and Trechispora fari-
nacea (Pers.) Liberta. These species have been collected from both
subtropical and temperate regions of Japan.
(b) Northern Hemisphere, Africa, and Oceania: 20 species. All
have been found in both temperate and (sub)tropical regions. Rep-
resentative species include Brevicellicium olivascens (Bres.) K.H.
Larss. & Hjortstam, Cerocorticium molle (Berk. & M.A. Curtis)
Jülich, Coniophora puteana (Schumach.) P. Karst., Dacryobolus
karstenii (Bres) Oberw. ex Parmasto, Erythricium salmonicolor
(Berk. & Broome) Burds., Gloeocystidiellum porosum (Berk. & M.A.
Curtis) Donk, Hyphodontia pallidula (Pers.) J. Erikss., and Trechis-
pora alnicola (Bourdot & Galzin) Liberta.
(c) Northern Hemisphere, Africa, and South America: 35 spe-
cies. These have been collected from temperate and (sub)tropical
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Species
Acanthophysium oakesii (Berk. & M.A. Curtis) Parmasto12, A, g
Aleurocystidiellum subcruentatum (Berk. & M.A. Curtis) P.A. Lemke12, A, g
Aleurocystidiellum tsugae (Yasuda) S.H. He & Y.C. Dai12, A, i, **
Aleurodiscus aurantius (Pers.) J. Schröt.12, B, a
Aleurodiscus disciformis (DC.) Pat.12, B, c
Aleurodiscus grantii Lloyd12, A, i
Aleurodiscus mirabilis (Berk. & M.A. Curtis) Höhn.12, B, a, **
Aleurodiscus parvisporus Núñez & Ryvarden12, A, j
Aleurodiscus subglobosporus Ginns & Bandoni12, A, k
Aleurodiscus subroseus S.H. He & Y.C. Dai12, C, i, *, **
Alutaceodontia alutacea (Fr.) Hjortstam & Ryvarden10, A, m, **
Amphinema arachispora Burds. & Nakasone3, A, g
Amphinema byssoides (Pers.) J. Erikss.3, B, a, O
Amylocorticium canadense (Burt) J. Erikss. & Weresub2, B, g
Amylocorticium cebennense (Bourdot) Pouzar2, B, a, **
Amylocorticium indicum K.S. Thind & S.S. Rattan2, B, i, *, **
Amylocorticium suaveolens Parmasto2, A, g
Amylocorticium subincarnatum (Peck) Pouzar2, B, d
Amylofungus globosporus (N. Maek.) Sheng H. Wu12, A, j
Amylostereum areolatum (Chaillet ex Fr.) Boidin12, A, e
Amylostereum chailletii (Pers.) Boidin12, B, a, *, **
Amylostereum laevigatum (Fr.) Boidin12, A, g
Amylostereum orientale S.H. He & Hai J. Li12, A, j, **
Amyloxenasma allantosporum (Oberw.) Hjortstam & Ryvarden2, A, m
Amyloxenasma grisellum (Bourdot) Hjortstam & Ryvarden2, A, c
Aphanobasidium pseudotsugae (Burt) Boidin & Gilles1, B, d
Asterostroma andinum Pat.12, B, a, O
Asterostroma boninense Suhara & N. Maek.12, C, k, O, **
Asterostroma cervicolor (Berk. & M.A. Curtis) Massee12, B, a, O, **
Asterostroma macrosporum N. Maek. & Suhara12, C, k, N, **
Asterostroma muscicola (Berk. & M.A. Curtis) Massee12, B, a, N, O, **
Asterostroma praeacutosporum Boidin, Lanq. & Gilles12, C, m, N, *
Athelia acrospora Jülich3, B, a, **
Athelia binucleospora J. Erikss. & Ryvarden3, A, h
Athelia decipiens (Höhn. & Litsch.) J. Erikss.3, B, b, **
Athelia epiphylla Pers.3, B, a, **
Athelia bulata M.P. Christ.3, A, f
Athelia repetobasidiifera N. Maek.3, A, k
Athelia rolfsii (Curzi) C.C. Tu & Kimbr.3, B, a, N, O
Athelia salicum Pers.3, B, d
Athelia sibirica (Jülich) J. Erikss. & Ryvarden3, A, h
Athelia termitophira N. Maek., Yokoi & Sotome3, A, k, **
Atheliachaete galactites (Bourdot & Galzin) Ţura, Zmitr., Wasser & Spirin11, A, g
Atheliachaete sanguinea (Fr.) Spirin & Zmitr.11, A, f
Atheloderma orientale Parmasto10, A, h
Baltazaria galactina (Fr.) Leal-Dutra, Dentinger & G.W. Gri.12, B, m
Boidinella cystidiolophora (Boidin & Gilles) Nakasone14, B, m
Boidinia borbonica Boidin, Lanq. & Gilles12, C, m, N
Boidinia furfuracea (Bres.) Stalpers & Hjortstam12, A, f
Boreostereum radiatum (Peck) Parmasto8, A, g
Boreostereum vibrans (Berk. & M.A. Curtis) Davydkina & Bondartseva8, B, i
Botryobasidium aureum Parmasto5, B, b
Botryobasidium bondarcevii (Parmasto) G. Langer5, B, c
Botryobasidium candicans J. Erikss.5, B, a, O
Botryobasidium conspersum J. Erikss.5, B, b, N, **
Botryobasidium curtisii Hallenb.5, B, m, O, *, **
Botryobasidium globosisporum (Boidin & Gilles) G. Langer5, C, m, N, *
Botryobasidium isabellinum (Fr.) D.P. Rogers5, B, c
Botryobasidium latisporum (N. Maek.) G. Langer5, A, j, **
Botryobasidium medium J. Erikss.5, B, c
Botryobasidium obtusisporum J. Erikss.5, B, a, **
Botryobasidium piliferum Boidin & Gilles5, B, m, N, **
Botryobasidium pruinatum (Bres.) J. Erikss.5, B, b, N
Botryobasidium sordidulum Boidin & Gilles5, B, m
Botryobasidium subcoronatum (Höhn. & Litsch.) Donk5, B, a, **
Botryobasidium vagum (Berk. & M.A. Curtis) D.P. Rogers5, B, a
Brevicellicium olivascens (Bres.) K.H. Larss. & Hjortstam14, B, a
Bulbillomyces farinosus (Bres.) Jülich11, B, a
Byssocorticium ebulatum Hjortstam & Ryvarden3, A, h, N
Byssocorticium pulchrum (S. Lundell) M.P. Christ.3, A, g
Species
Byssomerulius corium (Pers.) Parmasto11, B, a, N, **
Byssomerulius hirtellus (Burt) Parmasto11, B, a, *, **
Byssomerulius tropicus (Sheng H. Wu) Zmitr.11, C, j, N
Cabalodontia queletii (Bourdot & Galzin) Piątek11, B, b
Candelabrochaete langloisii (Pat.) Boidin11, C, m, O
Candelabrochaete verruculosa Hjortstam11, B, m
Ceraceomyces tessulatus (Cooke) Jülich2, B, b
Ceratobasidium cornigerum (Bourdot) D.P. Rogers5, B, a
Ceratobasidium gramineum (Ikata & T. Matsuura) Oniki, Ogoshi & T. Araki5, A, i
Ceratobasidium setariae (Sawada) Oniki, Ogoshi & T. Araki5, A, j
Cerinomyces albosporus Boidin & Gilles7, B, m, **
Cerinomyces altaicus Parmasto7, A, j, *, **
Cerinomyces canadensis (H.S. Jacks. & G.W. Martin) G.W. Martin7, A, e, **
Cerinomyces pallidus G.W. Martin7, A, m
Cerocorticium molle (Berk. & M.A. Curtis) Jülich11, B, b, N, O, **
Chondrostereum purpureum (Pers.) Pouzar1, B, a, N, **
Conferticium karstenii (Bourdot & Galzin) Hallenb.12, A, e
Conferticium ochraceum (Fr.) Hallenb.12, B, f
Coniophora arida (Fr.) P. Karst.4, B, a, **
Coniophora fusispora (Cooke & Ellis) Cooke4, B, d
Coniophora olivacea (Fr.) P. Karst.4, B, a, **
Coniophora puteana (Schumach.) P. Karst.4, B, b
Coniophorafomes matsuzawae (Lloyd) Rick12, A, k
Corticium boreoroseum Boidin & Lanq.6, A, g
Corticium lombardiae (M.J. Larsen & Gilb.) Boidin & Lanq.6, A, g, **
Corticium meridioroseum Boidin & Lanq.6, A, h
Corticium roseocarneum (Schwein.) Hjortstam6, A, f, **
Cristinia helvetica (Pers.) Parmasto1, B, b
Crustodontia chrysocreas (Berk. & M.A. Curtis) Hjortstam & Ryvarden11, B, a, N, **
Crustomyces subabruptus (Bourdot & Galzin) Jülich1, A, g
Crystallicutis serpens (Tode) El-Gharabawy, Leal-Dutra & G.W. Gri.10, B, a
Cyanobasidium microverrucisporum (N. Maek.) Hjortstam & Ryvarden1, A, m
Cylindrobasidium argenteum (Kobayasi) N. Maek.1, C, k, N, **
Cylindrobasidium laeve (Pers.) Chamuris1, B, a, **
Cytidia salicina (Fr.) Burt6, A, g, **
Dacryobolus gracilis H.S. Yuan11, C, i, *, **
Dacryobolus karstenii (Bres.) Oberw. ex Parmasto11, B, b, N
Dacryobolus sudans (Alb. & Schwein.) Fr.11, B, a, N
Dendrocorticium polygonioides (P. Karst.) M.J. Larsen & Gilb.6, B, d
Dendrodontia bispora (Burds. & Nakasone) Guerrero & C.L.M. Rodrigues6, C, m, N, *
Dendrothele acerina (Pers.) P.A. Lemke1, B, a
Dendrothele arachispora Nakasone & Burds.1, A, m
Dendrothele strumosa (Fr.) P.A. Lemke1, B, m
Dentipellicula leptodon (Mont.) Y.C. Dai & L.W. Zhou12, B, a, *, **
Dentipellicula taiwaniana Sheng H. Wu12, C, j, N, *
Dentipellis dissita (Berk. & Cooke) Maas Geest.12, B, m, *
Dentipellis fragilis (Pers.) Donk12, B, m
Dentocorticium sulphurellum (Peck) M.J. Larsen & Gilb.11, A, i, **
Dichostereum granulosum (Pers.) Boidin & Lanq.12, A, e
Dichostereum kenyense Boidin & Lanq.12, B, m, *, **
Dichostereum rhodosporum (Wakef.) Boidin & Lanq.12, B, m
Duportella miranda Boidin, Lanq. & Gilles12, C, m, N
Echinodontiellum japonicum (Imazeki) S.H. He & Nakasone12, A, j
Echinodontium tsugicola (Henn. & Shirai) Imazeki12, A, k
Ebula corymbata (G. Cunn.) Zmitr. & Spirin11, B, m
Ebula tropica Sheng H. Wu11, C, j, N
Epithele ebulata Boidin, Lanq. & Gilles11, C, m, N, *, **
Epithele nikau G. Cunn.11, B, m, N
Epithele typhae (Pers.) Pat.11, A, f, N
Erythricium salmonicolor (Berk. & Broome) Burds.6, B, b, N
Erythromyces crocicreas (Berk. & Broome) Hjortstam & Ryvarden10, C, m, N, **
Fibricium rude (P. Karst.) Jülich10, B, d
Fibricium subcarneum Y. Hayashi10, A, k
Fibrodontia gossypina Parmasto10, B, c, N, O, **
Fibulomyces mutabilis (Bres.) Jülich3, B, m, **
Galzinia incrustans Parmasto6, B, f
Ginnsia viticola (Schwein.) Sheng H. Wu & Hallenb.10, A, m, **
Gloeocystidiellum convolvens (P. Karst.) Donk12, B, c, **
Gloeocystidiellum formosanum Sheng H. Wu12, B, j
Table 1
Corticioid fungi reported from Japan.
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Species
Gloeocystidiellum kenyense Hjortstam12, B, m, *, **
Gloeocystidiellum leucoxanthum (Bres.) Boidin12, B, b
Gloeocystidiellum luridum (Bres.) Boidin12, B, d
Gloeocystidiellum moniliforme Sheng H. Wu12, C, m, N
Gloeocystidiellum porosum (Berk. & M.A. Curtis) Donk12, B, a, O, **
Gloeodontia discolor (Berk. & M.A. Curtis) Boidin12, B, m, O, **
Gloeohypochnicium analogum (Bourdot & Galzin) Hjortstam12, B, b
Gloeomyces moniliformis (N. Maek.) Sheng H. Wu12, A, j
Gloeostereum incarnatum S. Ito & S. Imai1, A, i, **
Gloiothele citrina (Pers.) Ginns & G.W. Freeman12, A, g, **
Gloiothele citrinoidea Sheng H. Wu12, C, j, *, **
Gloiothele globosa Sheng H. Wu12, C, m, O, *, **
Gloiothele lactescens (Berk.) Hjortstam12, B, c, **
Grammothele fuligo (Berk. & Broome) Ryvarden11, C, m, N, O
Granulobasidium vellereum (Ellis & Cragin) Jülich1, B, f
Gyrophanopsis polonensis (Bres.) Stalpers & P.K. Buchanan11, B, a, **
Haloaleurodiscus mangrovei N. Maek., Suhara & K. Kinjo12, C, k, N, **
Hastodontia hastata (Litsch.) Hjortstam & Ryvarden10, A, f
Hydnocristella himantia (Schwein.) R.H. Petersen9, A, e, **
Hydnophanerochaete odontoidea (Sheng H. Wu) Sheng H. Wu & C.C. Chen11, C, j
Hydnophlebia chrysorhiza (Torr.) Parmasto11, B, m, N, **
Hyphoderma lapponicum (Litsch.) Ryvarden11, A, f
Hyphoderma litschaueri (Burt) J. Erikss. & Å. Strid11, B, m, N
Hyphoderma medioburiense (Burt) Donk11, B, a
Hyphoderma microcystidium Sheng H. Wu11, C, m, N
Hyphoderma nudicephalum Gilb. & M. Blackw.11, B, a, N, O, **
Hyphoderma obtusumJ. Erikss.11, A, e
Hyphoderma pilosum (Burt) Gilb. & Budington11, A, i
Hyphoderma roseocremeum (Bres.) Donk11, B, c
Hyphoderma setigerum (Fr.) Donk (s. l.)11, B, a, N, O, **
Hyphoderma sibiricum (Parmasto) J. Erikss. & Å. Strid11, B, f
Hyphoderma subsetigerum Sheng H. Wu11, A, j, **
Hyphoderma transiens (Bres.) Parmasto11, B, m, **
Hyphodermella corrugata (Fr.) J. Erikss. & Ryvarden11, B, a
Hyphodontia alienata (S. Lundell) J. Erikss.10, B, a
Hyphodontia alutaria (Burt) J. Erikss.10, B, a
Hyphodontia arguta (Fr.) J. Erikss.10, B, a
Hyphodontia pallidula (Bres.) J. Erikss.10, B, b, O, **
Hyphodontia sphaerospora (N. Maek.) Hjortstam10, A, m
Hyphodontia stipata (Fr.) Gilb.10, A, g
Hyphodontia subspathulata (H. Furuk.) N. Maek.10, A, k
Hyphodontia zhixiangii L.W. Zhou & Gaorov10, A, g, *, **
Hypochnicium eichleri (Bres. ex Sacc. & P. Syd.) J. Erikss. & Ryvarden11, B, d
Hypochnicium globosum Sheng H. Wu11, A, j, **
Hypochnicium longicystidiosum (S.S. Rattan) Hjortstam & Ryvarden11, B, m
Hypochnicium pini Y. Jang & J.J. Kim11, A, i, *, **
Hypochnicium punctulatum (Cooke) J. Erikss.11, B, a, **
Hypochnicium sphaerosporum (Höhn. & Litsch.) J. Erikss.11, B, c
Hypochnicium subrigescens Boidin11, B, m, *
Kavinia vivantii Boidin & Gilles9, C, i, O, *
Kneiella alutacea (Fr.) Jülich & Stalpers10, A. d
Kneiella barba-jovis (Bull.) P. Karst.10, A, a, N
Kneiella byssoidea (H. Furuk.) Hjortstam & Ryvarden10, A, k
Kneiella cineracea (Bourdot & Galzin) Jülich10, A, h
Kneiella microspora (J. Erikss. & Hjortstam) Jülich & Stalpers10, B, c, O
Kneiella serpentiformis (Langer) Riebesehl & Langer10, A, m, **
Kneiella subalutacea (P. Karst.) Bres.10, A, g, **
Kneiella subglobosa (Sheng H. Wu) Hjortstam10, C, m, N
Kurtia argillacea (Bres.) Karasiński6, B, a, N, O
Kurtia macedonica (Litsch.) Karasiński6, A, m
Laurilia sulcata (Burt) Pouzar12, A, g
Lauriliella taxodii (Lentz & H.H. McKay) S.H. He & Nakasone12, A, i
Lawrynomyces capitatus (J. Erikss. & Å. Strid) Karasiński10, A, g
Laxitextum bicolor (Pers.) Lentz12, B, a, N
Leptocorticium cyatheae (S. Ito & S. Imai) Hjortstam & Ryvarden6, C, m, O
Leptocorticium sasae (Boidin, Cand. & Gilles) Nakasone6, A, h, *, **
Leptosporomyces fuscostratus (Burt) Hjortstam3, A, g
Leptosporomyces septentrionalis (J. Erikss.) Krieglst.3, B, c
Leucogyrophana mollusca (Fr.) Pouzar4, B, b, O, **
Leucogyrophana pseudomollusca (Parmasto) Parmasto4, A, g, O, *, **
Licrostroma subgiganteum (Berk.) P.A. Lemke6, B, m
Lindtneria thujatsugina M.J. Larsen1, B, i, O, *
Species
Lindtneria trachyspora (Bourdot & Galzin) Pilát1, B, c, O, *
Litschauerella clematidis (Bourdot & Galzin) J. Erikss. & Ryvarden14, B, d, N
Lloydella okabei S. Ito & S. Imai11, C, k, O
Lopharia ayresii (Berk. ex Cooke) Hjortstam11, C, m, N, O
Lopharia cinerascens (Schwein.) G. Cunn.11, B, a, **
Lyoathelia laxa (Burt) Hjortstam & Ryvarden3, A, i
Lyomyces bisterigmatus (Boidin & Gilles) Hjortstam & Ryvarden10, C, i, *, **
Lyomyces boninensis (S. Ito & S. Imai) Hjortstam & Ryvarden10, C, m, N, O
Lyomyces crustosus (Pers.) P. Karst.10, B, c, N, **
Lyomyces griseliniae (G. Cunn.) Riebesehl & Langer10, B, m, N
Lyomyces mascarensis Riebesehi, Yurchenko & Langer10, C, c, *, **
Lyomyces orientalis Riebesehi, Yurchenko & Langer10, A, i, *, **
Lyomyces pruni (Lasch) Riebesehi & Langer10, B, c
Lyomyces sambuci (Pers.) P. Karst.10, A, c, O
Megalocystidium wakullum (Burds., Nakasone & G.W. Freeman) E. Larss. & K.H.
Larss.12, C, m, N
Melzericium rimosum Bononi & Hjortstam3, A, m
Metulodontia nivea (P. Karst.) Parmasto11, A, a
Mutatoderma mutatum (Peck) C.E. Gómez6, B, c
Mycoacia aurea (Fr.) J. Erikss. & Ryvarden11, B, a, N, O, **
Mycoacia chrysella (Berk. & M.A. Curtis) H. Furuk.11, A, i
Mycoacia fuscoatra (Fr.) Donk11, B, a, **
Mycoacia kurilensis Parmasto11, B, i
Mycoacia nothofagi (G. Cunn.) Ryvarden11, C, m, *
Neoaleurodiscus fujii Sheng H. Wu12, A, k, **
Odonticium abelliradiatum (J. Erikss. & Hjortstam) Zmitr.–, B, g, *, **
Odontoebula orientalis C.C. Chen & Sheng H. Wu11, C, j
Peniophora bicornis Hjortstam & Ryvarden12, C, m, O
Peniophora cinerea (Pers.) Cooke12, B, a, N, **
Peniophora erikssonii Boidin12, A, g
Peniophora incarnata (Pers.) P. Karst.12, B, a
Peniophora lycii (Pers.) Höhn. & Litsch.12, B, a
Peniophora manshurica Parmasto12, A, j, **
Peniophora nuda (Fr.) Bres.12, B, a
Peniophora ovalispora Boidin, Lanq. & Gilles12, C, m, N, **
Peniophora pithya (Pers.) J. Erikss.12, B, c
Peniophora versicolor (Bres.) Sacc. & P. Syd.12, B, m
Peniophora versiformis (Berk. & M.A. Curtis) Bourdot & Galzin12, A, g
Peniophora violaceolivida (Sommerf.) Massee12, A, e
Peniophorella crystallifera Yurchenko, Sheng H. Wu & N. Maek.–, B, j, O, **
Peniophorella echinocystis (J. Erikss. & Å. Strid) K.H. Larss.–, A, f
Peniophorella odontiiformis (Boidin & Berthier) K.H. Larss.–, B, a, N, O, **
Peniophorella pertenuis (P. Karst.) Hallenb. & H. Nilsson–, A, e, **
Peniophorella praetermissa(P. Karst.) K.H. Larss.–, B, a, O, **
Peniophorella pubera(Fr.) P. Karst.–, B, a, N, O, **
Penttilamyces olivascens (Berk. & M.A. Curtis) Zmitr., Kalinovskaya & Myas-
nikov4, B, d
Phaeophlebiopsis himalayensis (Dhingra) Zmitr.11, A, j, *
Phaeophlebiopsis peniophoroides (Gilb. & Adask.) Floudas & Hibbett11, B, i, *, **
Phaeophlebiopsis ravenelii (Cooke) Zmitr.11, B, a, N, O
Phanerochaete australis Jülich11, B, m, N, *, **
Phanerochaete brunnea Sheng H. Wu11, C, j, N
Phanerochaete burtii (Romell ex Burt) Parmasto11, B, m
Phanerochaete carnosa (Burt) Parmasto11, B, m
Phanerochaete eburnea Sheng H. Wu11, A, m, N, **
Phanerochaete laevis (Fr.) J. Erikss. & Ryvarden11, B, a
Phanerochaete leptoderma Sheng H. Wu11, A, j, N, **
Phanerochaete salmoneolutea Burds. & Gilb.11, B, i, *, **
Phanerochaete sordida (P. Karst.) J. Erikss. & Ryvarden11, B, a, N, O, **
Phanerochaete stereoides Sheng H. Wu11, C, j, N
Phanerochaete subceracea (Burt) Burds.11, B, m
Phanerochaete subglobosa Sheng H. Wu11, B, j
Phanerochaete velutina (DC.) P. Karst.11, B, c, **
Phanerodontia magnoliae (Berk. & M.A. Curtis) Hjortstam & Ryvarden11, B, f, O, **
Phlebia acanthocystis Gilb. & Nakasone11, B, m, N, O, **
Phlebia acerina Peck11, B, a, N, **
Phlebia brevispora Nakasone11, A, i, O, **
Phlebia caspica Hallenb.11, A, h, *
Phlebia coccineofulva Schwein.11, B, c
Phlebia lilascens (Bourdot) J. Erikss. & Hjortstam11, B, a
Phlebia livida (Pers.) Bres.11, B, a, N, O, **
Phlebia ludoviciana (Burt) Nakasone & Burds.11, B, f
― 4 ―
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Species
Phlebia radiata Fr.11, B, a, **
Phlebia subserialis (Bourdot & Galzin) Donk11, B, a, N
Phlebia tremellosa (Schrad.) Nakasone & Burds.11, B, a, **
Phlebia tuberculata (Hallenb. & E. Larss.) Ghob.-Nejh.11, B, e
Phlebiopsis crassa (Lév.) Floudas & Hibbett11, B, a, N, **
Phlebiopsis avidoalba (Cooke) Hjortstam11, B, f, N
Phlebiopsis gigantea (Fr.) Jülich11, B, a
Phlebiopsis roumeguerei (Bres.) Jülich & Stalpers11, B, d
Phlyctibasidium polyporoideum (Berk. & M.A. Curtis) Jülich–, B, a
Piloderma byssinum (P. Karst.) Jülich3, A, b
Piloderma lanatum (Jülich) J. Erikss. & Hjortstam3, A, g
Porostereum spadiceum (Pers.) Hjortstam & Ryvarden11, B, a, O, **
Pseudolagarobasidium calcareum (Cooke & Massee) Sheng H. Wu11, B, m, N, **
Pseudomerulius aureus (Fr.) Jülich4, B, e, **
Punctularia atropurpurascens (Berk. & Broome) Petch6, B, c, *, **
Punctularia strigosozonata (Schwein.) P.H.B. Talbot6, B, a, **
Radulodon casearius (Morgan) Ryvarden11, A, g
Radulodon copelandii (Pat.) N. Maek.11, A, e, **
Radulomyces conuens (Fr.) M.P. Christ.1, B, a, N, **
Radulomyces molaris (Chaillet ex Fr.) M.P. Christ.1, A, g
Ramaricium albo-ochraceum (Bres.) Jülich9, B, c, O, *
Repetobasidium intermedium Oberw.5, A, j
Repetobasidium miricum J. Erikss.5, B, c, O, *
Repetobasidium vile (Bourdot & Galzin) J. Erikss.5, A, g
Resinicium bicolor (Alb. & Schwein.) Parmasto–, B, m, N, O
Resinicium luteosulphureum (Rick) Baltazar & Rajchenb.–, B, m, N, O, **
Resinicium mutabile Nakasone–, B, m, *
Resinicium saccharicola (Burt) K.K. Nakasone–, C, c, N, *, **
Rhizochaet e lamentosa (Berk. & M.A. Curtis) Gresl., Nakasone & Rajchenb.11, B, m, N, **
Rhizochaete radicata (Henn.) Gresl., Nakasone & Rajchenb.11, B, a, **
Rhizochaete sulphurina (P. Karst.) K.H. Larss.11, A, e
Rhizoctonia anceps (Bres., Syd. & P. Syd.) Oberw., R. Bauer, Garnica & R.
Kirschner5, A, g
Rhizoctonia fusispora (J. Schröt.) Oberw., R. Bauer, Garnica & R. Kirschner5, A, e, *, **
Rhizoctonia ochracea (Massee) Oberw., R. Bauer, Garnica & R. Kirschner5, B, m
Rhizoctonia solani J.G. Kühn5, B, a
Rhizoctonia sterigmatica (Bourdot) Oberw., R. Bauer, Garnica & R. Kirschner5, B, m, N
Sceptrulum inatum (Burt) K.H. Larss.12, A, i, *
Scopuloides rimosa (Cooke) Jülich11, B, c, N, O, **
Scotoderma viride (Sacc.) Jülich12, A, m
Scytinostroma africanogalactinum Boidin, Lanq. & Gilles12, C, m, O, *
Scytinostroma duriusculum (Berk. & Broome) Donk12, B, a, N, O, **
Scytinostroma lusitanicum (Trotter) P.M. Kirk12, B, c
Scytinostroma odoratum (Fr.) Donk12, B, a, O, **
Scytinostroma portentosum (Berk. & M.A. Curtis) Donk12, B, a, N, O
Scytinostromella nannfeldtii (J. Erikss.) G.W. Freeman & R.H. Petersen12, A, g, O
Serpula himantioides (Fr.) P. Karst.4, B, a, **
Serpula lacrymans (Wulfen) J. Schröt.4, B, d, **
Serpula pulverulenta (Sowerby) Bondartsev4, A, g
Sertulicium granuliferum(Hallenb.) Spirin & Volobuev.14, B, d, O, *, **
Sertulicium niveocremeum (Höhn. & Litsch.) Spirin & K.H. Larss.14, B, a, O
Sistotrema athelioides Hallenb.5, A, f, **
Sistotrema biggsiae Hallenb.5, B, d
Sistotrema brinkmannii (Bres.) J. Erikss.5, B, a, **
Sistotrema conuens Pers.5, B, d
Sistotrema coronilla (Höhn.) Donk ex D.P. Rogers5, B, d, O
Sistotrema diademiferum (Bourdot & Galzin) Donk5, B, c, O
Sistotrema microsporum N. Maek.5, A, j
Sistotrema muscicola (Pers.) S. Lundell5, B, b
Sistotrema oblongisporum M.P. Christ. & Hauerslev5, B, c, **
Sistotrema porulosum Hallenb.5, A, f
Sistotrema subtrigonospermum D.P. Rogers5, B, a
Sistotremella perpusilla Hjortstam14, B, m
Skvortzovia furfuracea (Bres.) G. Gruhn & Hallenberg–, B, d
Skvortzovia furfurella (Bres.) Bononi & Hjortstam–, A, m, N
Skvortzovia pinicola (J. Erikss.) G. Gruhn & Hallenb.–, A, m, O, *, **
Sphaerobasidium minutum (J. Erikss.) Oberw. ex Jülich14, A, g, **
Steccherinum ciliolatum (Berk. & M.A. Curtis) Gilb. & Budington11, B, f
Steccherinum mbriatum (Pers.) J. Erikss.11, B, a, N, **
Steccherinum helvolum (Zipp. ex Lév.) S. Ito11, A, j
Steccherinum laeticolor (Berk. & M.A. Curtis) Banker11, B, c
Steccherinum ochraceum (Pers. ex J.F. Gmel.) Gray11, B, a, N, **
Species
Steccherinum rawakense (Pers.) Banker11, B, a
Stereofomes palmicola S. Ito & S. Imai12, C, k, O
Stereum boninense Yasuda12, C, k, O
Stereum gausapatum (Fr.) Fr.12, B, c
Stereum hirsutum (Willd.) Pers.12, B, c, N, **
Stereum kurilense Yasuda12, A, k
Stereum ochraceoavum (Schwein.) Sacc.12, A, g
Stereum ostrea (Blume & T. Nees) Fr.12, B, m, N, **
Stereum pendulum R. Sasaki12, A, k
Stereum rimosum Berk.12, B, m
Stereum rugosum Pers.12, B, a, **
Stereum sanguinolentum (Alb. & Schwein.) Fr.12, B, a, N, **
Stereum spectabile Klotzsch12, B, m, N, O, **
Stereum vellereum Berk.12, B, a
Subulicium lautum (H.S. Jacks.) Hjortstam & Ryvarden10, A, e
Subulicystidium brachysporum (P.H.B. Talbot & V.C. Green) Jülich14, B, m, N, O
Subulicystidium longisporum (Pat.) Parmasto14, B, a, N, O
Subulicystidium meridense Oberw.14, B, m, N, O
Subulicystidium nikau (G. Cunn.) Jülich14, B, m, N
Subulicystidium perlongisporum Boidin & Gilles14, B, d
Terana coerulea (Lam.) Kuntze11, B, a
Thanatephorus repetosporus (G. Langer & Ryvarden) P. Roberts5, C, m, O, *
Thelephora ellisii (Sacc.) Zmitr., Shchepin, Volobuev & Myasnikov13, A, a, **
Theleporus calcicolor (Sacc. & P. Syd.) Ryvarden11, B, m, N, O, **
Theleporus membranaceus Y.C. Dai & L.W. Zhou11, C, j, N, *, **
Tomentella crinalis (Fr.) M.J. Larsen13, A, e, **
Tomentella ferruginea (Pers.) Pat.13, A, e
Trechispora cohaerens (Schwein.) Jülich & Stalpers14, B, f, N, O, **
Trechispora dimitica Hallenb.14, B, m, O
Trechispora farinacea (Pers.) Liberta14, B, a, N, O, **
Trechispora microspora (P. Karst.) Liberta14, B, a, **
Trechispora mollusca (Pers.) Liberta14, B, a
Trechispora nivea (Pers.) K.H. Larss.14, B, a, N, O, **
Trechispora praefocata (Bourdot & Galzin) Liberta14, A, m, N, O
Trechispora regularis (Murrill) Liberta14, B, a
Trechispora stevensonii (Berk. & Broome) K.H. Larss.14, B, a, N, **
Trechispora subsphaerospora (Litsch.) Liberta14, A, f
Trechispora tenuicula (Litsch.) K.H. Larss.14, A, h, *
Tubulicium curvisporum Ushijima & N. Maek.14, A, k, **
Tubulicium raphidisporum (Boidin & Gilles) Oberw., Kisim.-Hor. & L.D. Gó-
mez14, C, m, N, O
Tubulicium vermiferum (Bourdot) Oberw. ex Jülich14, A, f
Tubulicrinis angustus (D.P. Rogers & Weresub) Donk10, A, g, *
Tubulicrinis borealis J. Erikss.10, A, f
Tubulicrinis calothrix (Pat.) Donk10, B, a, **
Tubulicrinis glebulosus (Fr.) Donk10, B, a, **
Tubulicrinis hamatus (H.S. Jacks.) Donk10, A, m
Tubulicrinis hirtellus (Bourdot & Galzin) J. Erikss.10, A, g, *, **
Tubulicrinis inornatus (H.S. Jacks. & D.P. Rogers) Donk10, A, f
Tubulicrinis medius (Bourdot & Galzin) Oberw.10, A, g, *, **
Tubulicrinis orientalis Parmasto10, A, h
Tubulicrinis subulatus (Bourdot & Galzin) Donk10, B, e, **
Tubulicrinis yunnanensis C.L. Zhao10, B, j, *, **
Tylospora asterophora (Bonord.) Donk3, A, g
Tylospora brillosa (Burt) Donk3, A, g, O
Vararia gittonii Boidin & Lanq.12, C, m, N
Vararia investiens (Schwein.) P. Karst.12, B, m
Vararia ochroleuca (Bourdot & Galzin) Donk12, A, m, **
Vararia phyllophila (Massee) D.P. Rogers & H.S. Jacks.12, B, m, O
Veluticeps abietina (Pers.) Hjortstam & Tellería8, A, g
Veluticeps berkeleyana Cooke8, B, g
Vuilleminia comedens (Nees) Maire6, B, b **
Vuilleminia cystidiata Parmasto6, B, m, * **
Waitea circinata Warcup & P.H.B. Talbot6, A, m
Xenasma pulverulentum (H.S. Jacks.) Donk11, B, a
Xenasma rimicola (P. Karst.) Donk11, B, a
Xenasma tulasnelloideum (Höhn. & Litsch.) Donk11, A, a, N, **
Xenasmatella alnicola (Bourdot & Galzin) K.H. Larss. & Ryvarden14, B, b, O, **
Xenasmatella athelioidea (N. Maek.) N. Maek.11, A, k, O, **
Xenasmatella christiansenii (Parmasto) Stalpers11, A, f, **
Xenasmatella brillosa (Hallenb.) K.H. Larss. & Hjortstam11, A, g
Xenasmatella vaga (Fr.) Stalpers11, B, a, **
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regions of Africa, Eurasia, and North and South America, exclud-
ing Oceania. Representative species include Amyloxenasma grisel-
lum (Bourdot) Hjortstam & Ryvarden, Fibrodontia gossipina Par-
masto, Gloeocystidiellum convolvens (P. Karst.) Donk, and
Kneiella microspora (J. Erikss. & Hjortstam) Jülich & Stalpers.
(d) Northern Hemisphere and Africa: 21 species. Most of them
are known from both temperate regions in the Northern Hemi-
sphere and tropical regions of Africa. Litschauerella clematidis
(Bourdot & Galzin) J. Erikss. & Ryvarden (Maekawa, 1993b) has
been collected from subtropical regions of Japan (the Okinawa Is-
lands in the Nansei Islands). Representative species include Amylo-
corticium subincarnatum (Peck) Pouzar, Aphanobasidium pseudot-
sugae (Burt) Boidin & Gilles, Gloeocystidiellum luridum (Bres.)
Boidin, and Sistotrema oblongisporum M.P. Christ. & Hauerslev.
(e) Northern Hemisphere and Oceania: 13 species. These have
been collected primarily in the temperate regions of Eurasia, North
America, and Oceania and are not found in Africa or South Amer-
ica. In Japan, these species are known only from the temperate re-
gions so far. Representative species include Amylostereum areola-
tum (Fr.) Hjortstam & Ryvarden, Byssomerulius albostramineus
(Torrend) Hjortstam, Hydnocristella himantia (Schwein.) R.H. Pe-
tersen, Pseudomerulius aureus (Fr.) Jülich, Radulodon copelandii
(Pat.) N. Maek., Tomentella ferruginea (Pers.) Pat., and Tubulicrinis
subulatus (Bourdot & Galzin) Donk.
(f) Northern Hemisphere and South America: 25 species. These
are known mainly from temperate regions. In Japan, Phanerodon-
tia magnoliae (Berk. & M.A. Curtis) Hjortstam & Ryvarden, Phlebi-
opsis avidoalba (Cooke) Hjortstam, and Trechispora cohaerens
(Schwein.) Jülich have also been collected in subtropical regions,
whereas the other species are known only from temperate regions
(Maekawa, 1993b, 2010).
(g) Northern Hemisphere: 35 species. Most have been found in
subarctic to temperate zones. In Japan, all of these species have
been collected only from the temperate regions, except Tylospora
brillosa (Burt) Donk, which has been collected not only from tem-
perate regions but also from the subtropical Ogasawara Islands
(Maekawa, 2010). Representative species include Aleurocystidiel-
lum subcruentatum (Berk. & M.A. Curtis) P.A. Lemke, Amylocorti-
cium canadense (Burt) J. Erikss & Weresub, Athelia salicum Pers.,
Byssocorticium pulchrum (S. Lundell) M.P. Christ., Cytidia salicina
(Fr.) Burt., Gloiothele citrina (Pers.) Ginns & G.W. Freeman, Kne-
iella subalutacea (P. Karst.) Bres., Peniophora erikssonii Boidin,
and Repetobasidium vile (Bourdot & Galzin) J. Erikss.
(h) Eurasia: 7 species. These species are known from with distri-
butions in the subarctic to temperate zones of Asia and Europe, but
not from other continents. In Japan, all of these species have been
collected only from the temperate regions (Honshu and Shikoku
islands), except for Byssocorticium ebulatum Hjortstam & Ryvar-
den, which has been collected not only from the temperate regions
but also from subtropical lowland of Yakushima Island (Maekawa,
2010). Representative species include Athelia sibilica (Jülich) J.
Erikss. & Ryvarden, Atheloderma orientale Parmasto, Byssocortici-
um ebulatum Hjortstam & Ryvarden, Kneiella cineracea (Bour-
dot & Galzin) Jülich, Phlebia caspica Hallenb., and Tubulicrinis
orientalis Parmasto.
(i) Far East Asia and North America: 24 species. Most species in
this group are distributed mainly in the boreal to temperate zones
of Far East Asia and North America, but Kavinia vivantii Boidin &
Gilles is known from Guadeloupe in the West Indies (Boidin &
Gilles, 2000) and from the subtropical Ogasawara Islands (Maeka-
wa, 2010). Representative species include Aleurodiscus grantii
Lloyd, Gloeostereum incarnatum S. Ito & S. Imai, Hyphoderma pilo-
sum (Burt) Bilb. & Budington, and Lyoathelia laxa (Burt) Hjorts-
tam & Ryvarden.
(j) Far East Asia: 33 species. Of these species, Byssomerulius
tropicus (Sheng H. Wu) Zmitr., Dentipellicula taiwaniana Sheng H.
Wu, Ebula tropica Sheng H. Wu, Phanerochaete stereoides Sheng
H. Wu, and Theleporus membranaceus Y.C. Dai & L.W. Zhou are
known only from subtropical regions. In Japan, these four species
are distributed in the Nansei Islands.
(k) Japan (endemic): 22 corticioid species are considered poten-
tially endemic to Japan (Fig. 1). Cyanobasidium microverrucispo-
rum (N. Maek.) Hjortstam & Ryvarden, Leptocorticium cyatheae (S.
Ito & S. Imai) Hjortstam & Ryvarden, Lyomyces boninensis (S. Ito &
S. Imai) Hjortstam & Ryvarden, and Xylodon pelliculae (H. Furuk.)
Riebesehl, Yurchenko & Langer were also recognized as endemic
to Japan, but they were subsequently found in Colombia (Hjorts-
tam & Ryvarden, 2005), Reunion Island (Boidin & Gilles, 1998),
Vanuatu (Maekawa, 2002), and China (Dai, 2011), respectively.
(m) Others: 95 corticioid species possess distribution patterns
that are dierent from patterns (a) to (k), above.
3.2. Corticioid fungi from subtropical regions of Japan
The Nansei and Ogasawara Islands are located in the subtropi-
cal zone of Japan. The Ogasawara Islands (also known as the Bonin
Islands), located about 1,000 km south of Tokyo (lat. 26°N–27°N,
long. 142°E), are oceanic islands with a characteristic forest ecosys-
tem that has many endemic species of trees and small woody
plants. The Nansei Islands are 1,600 km west of the Ogasawara Is-
lands (lat. 24°N–27°N, long. 124°E–127°E), are at almost the same
latitude as the former. In contrast to the Ogasawara Islands, the
Nansei Islands are continental islands and have forest vegetation
types, including mangrove forests, similar to those of Taiwan and
Species
Xylobolus frustulatus (Pers.) P. Karst.12, B, a, N, **
Xylobolus hiugensis (Imazeki) Imazeki & Hongo12, A, k
Xylobolus illudens (Berk.) Boidin12, B, a
Xylobolus princeps (Jungh.) Boidin12, B, m, N, **
Xylobolus subpileatus (Berk. & M.A. Curtis) Boidin12, B, a
Xylodon asper (Fr.) Hjortstam & Ryvarden10, B, b, **
Xylodon brevisetus (P. Karst.) Hjortstam & Ryvarden10, B, a
Xylodon aviporus (Berk. & M.A. Curtis ex Cooke) Riebesehl & Langer10, B, a, N, **
Xylodon nespori (Bres.) Hjortstam & Ryvarden10, B, a, N, **
Xylodon niemelaei (Sheng H. Wu) Hjortstam & Ryvarden10, C, m, O, **
Xylodon nudisetus (Warcup & P.H.B. Talbot) Hjortstam & Ryvarden10, A, m, O
Xylodon ovisporus (Corner) Riebesehl & Langer10, B, m, N, **
Xylodon pelliculae (H. Furuk.) Riebesehl, Yurchenko & Langer10, A, j
Xylodon quercinus (Pers.) Gray10, A, g
Xylodon radula (Fr.) Ţura, Zmitr., Wasser & Spirin10, A, m, N
Xylodon raduloides Riebesehl & Langer10, B, a, N
Xylodon reticulatus (C.C. Chen & Sheng H. Wu) C.C. Chen & Sheng H. Wu10, A, j
Xylodon spathulatus (Schrad.) Kuntze10, B, c, **
Xylodon subaviporus C.C. Chen & Sheng H. Wu10, C, i, *, **
* First report from Japan. ** Species identied on the basis of morphological and ITS
sequence data
Superscript numbers indicate the orders to which the species belong: 1, Agaricales;
2, Amylocorticiales; 3, Atheliales; 4, Boletales; 5, Cantharellales; 6, Corticiales; 7, Da-
crymycetales; 8, Gloeophyllales; 9, Gomphales; 10, Hymenochaetales; 11, Polyporales;
12, Russulales; 13, Thelephorales; 14, Trechisporales; and –, Unknown.
Superscript uppercase letters indicate the climatic zones or islands in which species
are distributed: A, Subarctic to temperate zones; B, Subarctic to subtropical zones; C,
Subtropical zone; N, Nansei Islands; and O, Ogasawara Islands.
Superscript lowercase letters indicate the regions where species are distributed: a,
Cosmopolitan; b, Northern Hemisphere, Africa, and Oceania; c, Northern Hemi-
sphere, Africa, and South America; d, Northern Hemisphere and Africa; e, Northern
Hemisphere and Oceania; f, Northern Hemisphere and South America; g, Northern
Hemisphere; h, Eurasia; i, Far East Asia and North America; j, Far East Asia; k, Ja-
pan (endemic); and m, Other.
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subtropical areas of the mainland China.
Taxonomic studies on corticioid fungi distributed in both archi-
pelagos have been carried out by several Japanese mycologists (Ito
& Imai, 1940; S. Ito, 1955; Maekawa, 1993b, 1994, 1997a, 1997b,
1998, 1999, 2000a, 2000b, 2010; Maekawa & Hasebe, 2002; Maeka-
wa et al., 2003, 2005; Sato, Uzuhashi, Hosoya, & Hosaka, 2010; Su-
Fig. 1 – Basidiomata of corticioid species endemic to Japan. A: Asterostroma boninense (TUMH 63841, holotype). B: Asterostroma macrospo-
rum (TUMH 60992, holotype). C: Athelia repetobasidiifera (TUMH 63503, holotype). D: Athelia termitophila (TUMH 40433, holotype). E:
Cylindrobasidium argenteum (TUMH 62012). F: Haloaleurodiscus mangrovei (TUMH 63842, holotype). G: Xenasmatella athelioidea (TUMH
63844, holotype). H: Tubulicium curvisporum (TUMH 63048, holotype). All specimens are deposited in TUMH, Tottori University. Bars: A, B,
E, H 10 mm; C, D, G 1 mm; F 10 cm.
AB
C
D
E
F
GH
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hara et al., 2010, 2011; Hosaka, 2018). These reports indicate that
146 corticioid species belonging to 77 genera are recorded in the
subtropical regions of Japan; 100 species in 58 genera are recorded
in the Nansei Islands, and 76 species in 50 genera are recorded in
the Ogasawara Islands (see footnote to Table 1). Although the Nan-
sei and Ogasawara Islands are located at similar latitudes, only 30
species (21% of the total corticioid species reported from the sub-
tropical regions) are found on both island groups. The dierences
in the corticioid fungi found on the Nansei and Ogasawara Islands
may be due to the following factors: (1) since the Nansei Islands are
continental and the Ogasawara Islands are oceanic, they oer dif-
ferent types of vegetation as substrates; (2) the Nansei and Ogas-
awara Islands are separated by 1,600 km, east to west; (3) there are
mangrove and Fagaceae trees on the Nansei Islands, whereas these
trees are absent and also limited distribution of coniferous trees on
the Ogasawara Islands.
3.3. Corticioid fungi in mangrove forests
Mangrove forests are distributed in coastal and riverine intertid-
al zones of the subtropics and tropics in many parts of the world.
According to the checklist for mangrove xylophilous basidiomy-
cetes compiled by Baltazar, Trierveiler-Pereira, and Louguer-
cio-Leite (2009), a total of 23 corticioid species are known from
mangroves around the world. In Japan, mangrove trees thrive
mainly in intertidal zones of the mouths of rivers in the southwest-
ern subtropical region. Before 2000, however, no corticioid species
were known from the Japanese mangrove forests, although 98
species of corticioid fungi had been collected from the subtropical
inland forests (Maekawa, 1993b, 1994, 1997a, 1997b, 1998, 2000a).
Since the beginning of this century, several surveys of corticioid
fungi have been conducted in the mangrove forests of the islands of
Okinawa and Iriomote, in the Ryukyu Islands (part of the Nansei
Islands), and the following 19 corticioid species, including one new
genus and two new species, have been reported (Maekawa et al.,
2003, 2005; Suhara et al., 2010): Asterostroma macrosporum N.
Maek. (Fig. 1B), A. muscicola (Berk. & M.A. Curtis) Massee, Bys-
somerulius tropicus (Sheng H. Wu) Zmitr. (as Phanerochaete tropica
(Sheng H. Wu) Hjortstam), Cerocorticium molle, Fibrodontia gos-
sypina Parmasto, Gloeocystidiellum moniliforme Sheng H. Wu,
Haloaleurodiscus mangrovei N. Maek., Suhara & K. Kinjo (Fig. 1F),
Hyphoderma nudicephalum Gilb. & M. Blackw., Lopharia ayresii
(Berk. ex Cooke) Hjortstam. (as Hyphoderma ayresii (Berk. ex
Cooke) Boidin & Gilles), Lyomyces crustosus (Pers.) P. Karst., Mega-
locystidium wakullum (Burds., Nakasone & G.W. Freeman) E.
Larss. & K.H. Larss. (as Gloeocystidiellum wakullum Burds.), Penio-
phorella odontiiformis (Boidin & Bethier) K.H. Larss. (as Hyphoder-
ma rude (Bres.) Hjortstam & Ryvarden), Phanerochaete sordida,
Phlebia acanthocystis Gilb. & Nakasone, Radulomyces conuens
(Fr.) M.P. Christ., Subulicystidium longisporum, Trechispora farina-
cea, T. nivea (Pers.) K.H. Larss., and Tubulicium raphidisporum
(Boidin & Gilles) Kisim.-Hor. & L.D. Gómez. Of these, Haloaleuro-
discus N. Maek., Suhara & Kinjo was described as a new genus; H.
mangrovei and A. macrosporum were each described as new spe-
cies; and A. muscicola, B. tropicus, C. molle, G. moniliforme, L. ayre-
sii, M. wakullum, and P. acanthocystis were reported from Japan for
the rst time (Maekawa et al., 2003, 2005; Suhara et al., 2010). None
of these 19 species has been reported from mangroves outside Ja-
pan.
The genus Haloaleurodiscus is morphologically characterized as
having resupinate basidioma, nodose-septate hyphae, sulphoalde-
hyde-positive gloeocystidia, dendrohyphidia, and amyloid basidio-
spores with minute warts. These morphological features are similar
to those of Aleurodiscus s. l., but ribosomal DNA sequence data in-
dicate that H. mangrovei is not closely related to Aleurodiscus spp.
Haloaleurodiscus mangrovei diers from Aleurodiscus primarily by
the association with a white pocket-rot. Maekawa et al. (2005) ex-
amined the phylogenetic relationships of aquatic homobasidiomy-
cetous fungi, namely Calathella mangrovei E.B.G. Jones & Agerer
(from mangrove), Halocyphina villosa Kohlm. & E. Kohlm. (from
mangrove), Nia vibrissa R.T. Moore & Meyers (marine), Physalac-
ria maipoensis Inderb. & Desjardin, and Limnoperdon incarnatum
G.A. Escobar (fresh water). All the examined species were found to
be phylogenetically nested in the euagarics clade, whereas H. man-
grovei was placed at the root of the Peniophorales clade. On the ba-
sis of this result, Maekawa et al. (2005) indicated that H. mangrovei
is phylogenetically distinct from previously reported aquatic ho-
mobasidiomycetous species, supporting the view that it represents
a unique evolutionary transition from terrestrial to marine envi-
ronments in Homobasidiomycetes. This species prefers dead
branches at the lower portions of living Sonneratia alba Sm. as its
substrate, although the collection sites is dominated by Bruguiera
gymhorhiza (L.) Lam. and Rhizophora mucronata Lam. Basidioma-
ta of this species usually occur on dead branches near the high tide
mark, but they sometimes form below the high tide mark. There-
fore, the tide certainly covers the basidiomata at least once a day for
about 1 week during the higher high tides. This habitat resembles
that of the marine basidiomycetous fungi. Mycelia of H. mangrovei
can grow on malt-extract agar (MEA) containing higher concentra-
tions (100‰, w/v) of sea salts than are found (32‰–35‰, w/v) in
seawater, although growth is slower than on freshwater agar media
(MEA). These features lead us to conclude that H. mangrovei is salt
tolerant, as is Physalacria maipoensis collected from mangrove (In-
derbitzin & Desjardin, 1999). They also suggest that basidiomata
can be produced on substrates that are periodically covered by the
tide during development (Maekawa et al., 2005). On the other
hand, all other corticioid species collected in mangrove forests pro-
duce their basidiomata 1.5 m or more above the ground on dead
branches of living mangroves, never forming basidiomata as low as
H. mangrovei. In addition, these species are not able to grow on
MEA containing salt concentrations similar to seawater (32‰–
35‰, w/v). Dierences in the position of basidioma formation be-
tween corticioid species of mangrove trees may be due to dierenc-
es in their tolerance to seawater (unpublished data).
4. Reevaluation of Japanese corticioid fungi on the
basis of combined morphological and molecular
data
Molecular phylogenetic analyses have been performed for sever-
al corticioid taxa, and these studies included some Japanese speci-
mens or isolates, or both. Nilsson, Hallenberg, Nordén, Maekawa,
and Wu (2003) performed molecular analysis, morphological ob-
servation, and crossing tests of Hyphoderma setigerum collected
from the Northern Hemisphere, and they revealed that H. seti-
gerum sensu lato contains nine preliminary taxa. Hyphoderma nu-
dicephalum Gilb. & M. Blackw., H. subsetigerum Sheng H. Wu and
an undescribed species were also detected, but H. setigerum sensu
stricto was not recognized among the Japanese specimens.
Peniophorella praetermissa (P. Karst.) K.H. Larss. has long been
regarded as a morphologically variable species complex. It is dis-
tributed widely in Japan and shows variability in cystidium mor-
phology, basidiospore shape and size, and productivity of stepha-
nocysts. Hallenberg et al. (2007) recognized three major clades,
further categorized into well-supported subclades on the basis of
phylogenetic analysis of internal transcribed spacer (ITS) data from
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worldwide sampling. Japanese specimens identied as P. praeter-
missa sensu lato on the basis of morphological features were placed
in ve (sub)clades; subclade 1A, subclade 1B (= P. praetermissa
sensu stricto), Clade 2 [= P. pertenuis (P. Karst.) Hallenb. & H. Nils-
son], subclade 3A [= P. odontiiformis (Boidin & Berthier) K.-H.
Larss.], and a fth, unnamed clade. Recently, the specimens be-
longing to the subclade 1A were reported as a new species, P. cryst-
allifera Yurchenko, Sheng H. Wu & N. Maek. (Yurchenko et al.,
2020).
Tabata and Abe (1997) were the rst to report Amylostereum
laevigatum (Fr.) Boidin from Japan, associated with the Japanese
horntail, Urocerus japonicus (L. f.) D. Don, a species of sawy. Al-
though Tabata, Harrington, Chen, and Abe (2000) used molecular
and morphological analyses to show Japanese A. laevigatum is dis-
tinct from French A. laevigatum, the scientic name remains in use
in Japan (Tabata & Abe, 1999; Tabata, Harrington, Chen, & Abe,
2000). Recently, however, by using morphological and molecular
data (ITS nrDNA and rpb2 markers), Nogal-Prata et al. (2017)
demonstrated that some specimens of A. laevigatum from Japan
belong to A. orientale S.H. He & Hai J. Li.
These studies suggest that some corticioid species may consist
of several cryptic species. Molecular phylogenetic analyses have
also revealed cryptic species in many corticioid genera such as
Gloeocystidiellum, Hyphoderma, Hyphodontia (sensu lato), and
Hypochnicium J. Erikss. (sensu lato). In a study of G. porosum and
its cryptic species, although we could not identify morphological
dierences between teleomorphs, the two taxa diered in the pro-
duction of anamorphs in culture (Maekawa, 2018). Thus, mycelial
characteristics in culture may be useful as taxonomic criteria for
distinguishing cryptic species.
5. Future prospects
Of the 442 corticioid fungi reported from Japan, sequence data
are available for almost half (218 species), and 168 species have
been identied on the basis of both morphological characteristics
of basidiomata and ITS sequence data (Supplementary Table S2).
However, for 50 species, the sequence collected for this study did
not match those previously deposited in GenBank (unpublished
data). Such discrepancies indicate that species identication by
using morphological characteristics of the specimens alone can be
unreliable. Therefore, the taxonomic positions of these specimens
should be reevaluated. As mentioned in the previous section, cryp-
tic species have also been found among some corticioid species
from Japan. More accurate identication of these species will re-
quire a combination of morphological characteristics and DNA se-
quence data. In addition, mycelial characteristics in culture may be
useful as taxonomic criteria for distinguishing cryptic species.
The remaining 224 species have been identied on the basis of
morphological characteristics alone and lack sequence data. In ad-
dition, some species have been identied from an insucient num-
ber of specimens or by using only a few morphological traits. For
example, taxa with the Japanese names “Akakouyakutake” and
“Kawatake” were rst reported from Japan as Aleurodiscus amor-
phus (Pers.) J. Schröt. by Yasuda (1914a) and Peniophora quercina
(Pers.) Cooke by Hennings (1903), respectively. The scientic
names have been commonly used in Japan for more than 80 years.
However, all of the Japanese specimens labeled “Aleurodiscus
amorphus” and “Peniophora quercina” deposited in several herbar-
ia were identied as A. grantii Lloyd and P. manshurica Parmasto,
respectively (Maekawa, 1994). In addition, A. grantii and P. mans-
hurica have been collected throughout Japan, whereas I have never
encountered A. amorphus or P. quercina. As mentioned in the pre-
vious section, H. setigerum (sensu stricto) was not recognized
among the specimens of H. setigerum (sensu lato) collected in Ja-
pan (Nilsson et al., 2003). Subsequently, many specimens of H. seti-
gerum (sensu lato) have been collected from various localities in
Japan, but no H. setigerum sensu stricto has been found. These re-
sults indicate that A. amorphus, H. setigerum sensu stricto, and P.
quercina, reported as Japanese corticioid species, may not be dis-
tributed in Japan. To verify the identities of those Japanese species
that have been identied by morphological characteristics alone,
detailed morphological observations and phylogenetic analysis are
needed. In addition, the characteristics of cultured mycelia, such as
production of anamorphs and cystidia, may be useful as taxonomic
criteria, depending on the species.
To date, numerous corticioid specimens have been collected
from all over Japan. However, many specimens have yet to be iden-
tied. In particular, more than half of the specimens collected in
subtropical regions, such as the Nansei and Ogasawara Islands, re-
main unidentied. Many of these are likely to be new taxa not list-
ed in Table 1. These specimens warrant detailed taxonomic studies
in the future.
Disclosure
The author declares no conicts of interest. All the experiments
undertaken in this study comply with the current laws of the coun-
try where they were performed.
Acknowledgements
The present article is based on the memorial talk for the recep-
tion of MSJ award. I sincerely thank the late Dr. Naohide Hiratsuka
and the late Dr. Yasuyuki Hiratsuka, of the Tottori Mycological In-
stitute, Japan, for their support of my research on the corticioid
fungi. I also thank Dr. B. Nordén Dr. N. Hallenberg and Dr. R. H.
Nilsson, University of Gothenburg, Göteborg, Sweden, for their
contribution to our cooperative studies and support. I am grateful
to the late Dr. J. Boidin (Lyon, France), Dr. J. Ginns (Centre for
Land and Biological Resources Research, Canada), Dr. K.-H. Lars-
son and Dr. E. Larsson (University of Gothenburg), Dr. K. Naka-
sone (United States Department of Agriculture, USA) and the late
Dr. E. Parmasto (Institute of Zoology and Botany, Estonia) and Dr.
S.-H. Wu (National Museum of Natural Science, Taiwan), for their
valuable comments on the Japanese corticioid specimens and their
loan of precious specimens. Thanks are also due to the many pro-
fessional and amateur researchers who provided their many corti-
cioid specimens. Finally, I would like to thank all my colleagues
and students at the Fungus/Mushroom Resource and Research
Center, Tottori University, for valuable discussions on the taxono-
my of corticioid fungi.
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