Biogeography of polypores in Malesia, Southeast Asia

Abstract

Biogeography of polypores distributed in the Malesian region is reviewed. Some of the species reported from this region are pantropical or paleotropical and widely distributed along the equator. Several species are restricted to lowland areas of Malesia and adjacent regions, and can be classified as Asian or Asian-Oceanian tropical species. Some species have only been recorded from lowland areas of Malesia at present, but detailed distributions are unclear for many of them. Some lowland Malesian species are also distributed in temperate areas of East Asia; among them, few species are circumglobal in the Northern Hemisphere, whereas many others are tropical species showing continuous distributions up to the warm temperate areas of East Asia. Species recorded from highland areas of Malesia include tropical species common to lowland Malesia, temperate species common to temperate Asia, and endemic species in highland areas of Malesia and adjacent regions. Regarding the factors that delimitate distribution of the Malesian polypores, I discuss polypore host preference, response to physical environmental factors, and dispersal ability. Systematics and ecological characteristics are briefly discussed for selected Malesian polypores: Corneroporus subcitrinus, Paratrichaptum accuratum, Postia stellifera and Roseofavolus eos.

Full text

Review
Biogeography of polypores in Malesia, Southeast Asia
Tsutomu Hattori
Forestry and Forest Products Research Institute, Tsukuba, Ibaraki 305-8687, Japan
Corresponding author:
T. Hattori
Tel: +81 29 829 8247
Fax: +81 29 873 1543
E-mail: hattori@affrc.go.jp
Text: 28 pages; 2 tables; 2 figures:
1

Abstract
Biogeography of polypores distributed in the Malesian region is reviewed. Some of the species
reported from this region are pantropical or paleotropical and widely distributed along the
equator. Several species are restricted to lowland areas of Malesia and adjacent regions, and
can be classified as Asian or Asian-Oceanian tropical species. Some species have only been
recorded from lowland areas of Malesia at present, but detailed distributions are unclear for
many of them. Some lowland Malesian species are also distributed in temperate areas of East
Asia; among them, few species are circumglobal in the Northern Hemisphere, whereas many
others are tropical species showing continuous distributions up to the warm temperate areas of
East Asia. Species recorded from highland areas of Malesia include tropical species common
to lowland Malesia, temperate species common to temperate Asia, and endemic species in
highland areas of Malesia and adjacent regions. Regarding the factors that delimitate
distribution of the Malesian polypores, I discuss polypore host preference, response to
physical environmental factors, and dispersal ability. Systematics and ecological
characteristics are briefly discussed for selected Malesian polypores:
Corneroporus subcitrinus, Paratrichaptum accuratum, Postia stellifera and Roseofavolus
eos.
Keywords:
Biodiversity, Host specificity, Polyporales, Tropics, Wood-decaying fungi
2

1. Introduction
“The Malesian region” refers to a biogeographic region that includes the Malay Peninsula,
Java, Sumatra, Borneo, the Philippines, and their surrounding islands, and that is categorized
under the Indo-Malesian Subkingdom in the Paleotropical Kingdom (Takhtajan 1961; Fig. 1).
Tree species richness in this region is extremely high. The numbers of the tree species recorded
in a 52-ha plot in Lambir Hills National Park (Northern Borneo) and in a 50-ha plot in Pasoh
Forest Reserve (the Malay Peninsula), both of which are primary lowland rainforest, are 1,175
and 818, respectively, whereas only 1,166 species are collectively known from all temperate
forests in the Northern Hemisphere (Wright 2002; Davis et al. 2003). Many trees are endemic
to this region, and areas including the Malay Peninsula, Northern Borneo, and the Philippines
are considered biodiversity hotspots in need of biodiversity conservation (Myers 1988).
Polypores are a group of fungi that includes important wood decomposers in forest
ecosystems. Their species richness and composition are often influenced by tree species
composition in forests, because many show host specificity or preference for certain trees.
High species diversity of forest trees in this region may result in high polypore species
richness.
Yamashita et al. (2015) surveyed a 2-ha plot survey in a lowland primary area of Pasoh
Forest Reserve and estimated 188 species of polypores, which is much higher than in
temperate areas. It is expected that the polypore species throughout the Malesian region should
be much richer, because different species should be distributed in other forest types including
hill forests, mountain and subalpine forests, young secondary forests, plantations, swamp
forests, mangrove forests, and geographically isolated primary lowland forests.
Ryvarden (1991) reviewed polypore biogeography, mainly at the genus level. He
mentioned that only the following five genera are endemic in tropical Asia, although some of
them are now known to accommodate non-Asian species: Elmerina Bres., Flabellophora G.
Cunn., Hymenogramme Mont. & Berk., Paratrichaptum Corner, and Sparsitubus Xu & Zhao.
This number of genera is substantially less than 22, which is the number of pantropic polypore
genera (Ryvarden 1991).
In contrast, polypore mycobiota in tropical Asia, including the Malesian region, is peculiar
3

compared with other regions at the species level. E. J. H. Corner intensively collected in the
Malesian and adjacent Papuan regions, and described nearly 280 new polypore species from
these regions (Corner 1983, 1984, 1987, 1989a, b, 1991, 1992), although several are now
considered synonyms of previously described or dubious species (Hattori 2000, 2001a, b,
2002, 2003a, b, 2005a; Dai and Li 2012; Hattori and Sotome 2013). Many of the species newly
described by Corner are still unknown outside of these two regions. Hattori and Lee (2003)
also concluded that many of the polypores recorded in Pasoh Forest Reserve are likely
endemic to tropical Asia.
In addition, recent phylogenetic studies have revealed that several pantropic and
paleotropic “species” actually represent species complexes that contain cryptic species
restricted to tropical Asia. This indicates that more species are endemic in tropical Asia,
including the Malesian and adjacent regions, than was once considered.
Although the polypore mycobiota in these regions is species-rich and peculiar, no
discussion has been made regarding the factors that influence distribution of these polypores.
As mentioned above, tree flora can be a primary factor that delimitates distribution of
polypores with host specificity. Climate, including temperature and precipitation, is another
factor that affects wood-decaying fungus distribution (Heilmann-Clausen and Boddy 2008).
Changes in mycobiota along latitudinal and altitudinal gradients may be partially influenced by
the response of each species to temperature. Dispersal ability may also influence the
distribution of each species.
In this article, I review important mycologists and literatures that described polypores
from the Malesian region. Then, I discuss polypore distribution patterns in this region, and
biological characteristics that might affect their distribution. Finally, I review some peculiar
species of polypores reported from this region.
2. Important taxonomical studies on Malesian polypores
Polypores in general produce less watery and highly persistent basidiomata, and are more
easily dried for keeping as specimens than agarics and many other macrofungi. This made it
possible for mycologists to describe considerable new polypore species during the 19th and the
4

early 20th centuries based on specimens collected during field expeditions to the Malesian and
adjacent Papuan regions (Table 1).
Among these regions, the largest number of new species was described from the
Philippines during this time, where significant contributions were made by W. A. Murrill and
C. G. Lloyd. Murrill (1907, 1908a, b) described a total of 58 polypore species from the
Philippines based on collections made by his coworkers including E. D. Merrill, R. S.
Williams, and A. D. E. Elmer (Ryvarden 1985). Later, Lloyd described 41 new species based
on specimens sent from his coworkers including E. D. Merrill (Ryvarden 1989, 1990, 1992).
Graff (1916) summarized newly described fungi from the Philippines in the 19th and early
20th centuries. However, despite intensive reports during this time, only limited studies have
been conducted on Philippine polypores after the late 20th century.
Java Island, Indonesia is another area from which many polypores were described. Most
species from “Indonesia excluding Borneo, Celebes, and New Guinea”, shown on Table 1,
were actually described from Java, except for some limited species that were mainly from
Sumatra. F. W. Junghuhn stayed for an extended amount of time in Java, and described 30 new
polypore species from Java (Ryvarden 1981a). J-H. Léveillé is another mycologist who
described various polypores from Java and Sumatra; type specimens of species he described
were mainly collected by H. Zollinger, C. E. O. Kuntze, and Junghuhn (Ryvarden 1981b).
More recently, Corner (1984, 1987, 1989a, 1989b, 1991) reported nearly 20 species from
mountain areas in Sumatra; subsequently, Suhirman and Núñez (1998b) made more
comprehensive collections from western Sumatra.
Only limited studies have been conducted on Celebes and its surrounding islands that
belong to the Papuan region, which neighbors the Malesian region to the east. P. C. Hennings
reported 13 species, including three new species from Celebes (Hennings 1900; Ryvarden
2012). Núñez et al. (2002) discussed polypore distribution patterns and included species
records from the Lesser Sunda Islands and Celebes. Suhirman and Núñez (1995, 1998a)
further summarized the literature and herbarium specimens of Indonesian polypores.
M. C. Cooke was the first mycologist who intensively reported on polypores from the
Malay Peninsula, mainly from Perak. He described nine new species based on the collection of
G. King (Ryvarden 1988a). C. G. Lloyd described several polypores from Singapore and other
5

areas of the Malay Peninsula based on specimens sent from various coworkers (Ryvarden
1989, 1990, 1992). An intensive study in Borneo (mainly Sarawak) was started by Cesati
(1879), who described 20 new species from this area (Saccardo 1888). Corner (1983, 1984,
1987, 1989a, b, 1991, 1992) reported a large number of polypores from West Malaysia,
Singapore and Sarawak. Yamashita et al. (2009) provided a list of polypores collected in the
Lambir Hills National Park, Sarawak followed by Lee et al. (2012), who made a
comprehensive list of fungi reported from Malaysia with important references.
3. Malesian polypore distribution patterns
3.1. Pantropic and paleotropic species
Hattori and Lee (2003) listed 33 pantropic and 24 paleotropic polypore species collected in
Pasoh Forest Reserve. Some of these species, however, contain cryptic species after
morphological and phylogenetic investigations, and tropical Asian populations represent
distinct species from those in the other regions. Polyporus tenuiculus (P. Beauv.) Fr. was once
considered a pantropic species, but it actually represents a species complex, and tropical Asian
populations include Favolus spatulatus (Jungh.) Lév. and F. roseus Lloyd, whereas a tropical
American population represents F. brasiliensis (Fr.) Fr. (Sotome et al. 2013). Fomitopsis feei
(Fr.) Kreisel was considered a species distributed in both tropical America and Asia, but the
tropical Asian population was shown to represent a distinct species,
F. rubida (Berk.) A. Roy & A.B. De (Han and Cui 2015).
Among species listed as pantropic or paleotropic species by Hattori and Lee (2003),
Ganoderma australe (Fr.) Pat., Perenniporia tephropora (Mont.) Ryvarden and Tinctoporellus
epimiltinus (Berk. & Broome) Ryvarden have also been shown to consist of species complexes
or have morphologically similar species (Moncalvo and Buchanan 2008; Yuan and Wan 2012;
Zhao and Cui 2013). Further molecular studies using tropical Asian specimens may reveal
additional cryptic Asian species among those that were regarded as pantropic or paleotropic
species.
3.2. Species restricted to lowland areas of Malesia and adjacent regions
6

The Malesian region neighbors the Indo-Chinese region to the north/northwest and the Papuan
region to the west (Takhtajan 1961). The Indo-Chinese region includes areas such as southern
continental China, Thailand, Hainan, Taiwan, and Sakishima, and numerous polypores have
been newly described from these areas based on morphological and phylogenetic
investigations (e.g., Sotome et al. 2009; Tokuda et al. 2009; Hattori et al. 2014; Zhao et al.
2014; Zhou 2014; Bian and Dai 2015; Han and Cui 2015). Many of these species may also be
distributed in the Malesian region, because species reported from Sakishima and other
subtropical areas in Japan by Aoshima and Ogimi (1974) are also known from the Malesian
region and/or other tropical regions in Asia; this is also true for most species known from
tropical areas of China, excluding recently described species without detailed information
about their distributions (Dai 2012). Datroniella subtropica B.K. Cui, Hai J. Li & Y.C. Dai,
described from subtropical areas of China by Li et al. (2014), is actually a common species in
lowland Malesia based on my personal observations, and was recorded from Singapore as
Trametes scutellata (Schwein.) G.H. Cunn. by Corner (1989b). Intensive collections in the
Malesian and Papuan regions with molecular evaluations will reveal if those species described
from the Indo-Chinese region are also distributed in the Malesian region.
However, only a few species described from lowland areas of the Malesian region by
Corner have been recorded from Hainan, Taiwan, and Sakishima, despite intensive collections
in these areas by Asian mycologists. Hyphodontia ovispora (Corner) T. Hatt. and
Ceriporiopsis aurantitingens (Corner) T. Hatt. are the limited examples that are also found in
Sakishima and/or other subtropical area of Japan (Hattori 2002, 2003a). Some more, but still
limited species, are reported from southern/central Thailand, situated in the south of the Indo-
Chinese region: Coriolopsis lacunosa (Corner) T. Hatt., Neofomitella fumosipora (Corner)
Y.C. Dai, Hai J. Li & Vlasák, Perenniporia corticola (Corner) Decock, P. marmorata (Corner)
T. Hatt., and Trametes pallidilusor Corner [= T. daedaleoides Corner non (Berk.) Berk.]
(Choeyklin et al. 2011). A northward distribution is likely restricted for many species described
from lowland areas of the Malesian region.
Although most parts of the lowland forests in both the Malesian and Papuan regions are
classified as tropical rain forest, the fauna in the Malay Peninsula, Borneo, Java, Sumatra, and
their surrounding islands is distinct from that in New Guinea, as delimited by Wallace’s Line.
7

Núñez et al. (2002), however, concluded that polypore mycobiota did not significantly differ
on either side of Wallace’s line after examining specimens collected from Bali, Lombok, and
Sumbawa, which are members of the Lesser Sunda Islands that are distributed across the
Wallace’s line. In fact, most of the common polypores distributed in the Malesian region are
also recorded from New Guinea (Quanten 1997), and many of those reported from the
Malesian region but unreported from the Papuan regions, are only known from singleton or
limited specimens, which indicates that they are rare and their distributions are poorly known.
However, it should be noted that some species recorded from New Guinea are previously
or widely known from New Zealand and/or Northeast Australia, but unknown from the
Malesian region: Australoporus tasmanicus (Berk.) P.K. Buchanan & Ryvarden,
Ceriporia otakou (G. Cunn.) P.K. Buchanan & Ryvaren, Fomes hemitephrus (Berk.) Cooke,
Heterobasidion araucariae P.K. Buchanan, Phellinus nothofagi (G. Cunn.) Ryvarden,
Piptoporus portentosus (Berk.) G. Cunn., and Skeletocutis novaezealandiae (G. Cunn.) P.K.
Buchanan & Ryvarden (Buchanan 1988; Quanten 1997). On the contrary,
Daedalea aurora (Ces.) Aoshima and Perenniporia corticola (Corner) Decock, which are
specific to Dipterocarpaceae trees, are common in Malesia (Hattori et al. 2012), but at present
are unknown from the Papuan region, where Dipterocarpaceae trees are sparse.
As mentioned above, several polypore species are shared between the lowland areas of the
Malesian and Papuan regions; some are restricted to these areas and considered tropical
Southeast Asian species. The following are examples of species restricted to the lowland areas
of these two regions: Antrodiella aurantiaca (Corner) T. Hatt., A. flava (Corner) T. Hatt.,
A. depauperata (Corner) T. Hatt., Elmerina cladophora (Berk.) Bres., Fomitopsis
pseudopetchii (Lloyd) Ryvarden, Nigrofomes nigrivineus (Corner) T. Hatt., Perenniporia
permacilenta (Corner) T. Hatt., Rigidoporus cystidioides (Lloyd) Ryvarden, and Tyromyces
corticola Corner (Corner 1987, 1989a, b; Hattori 2000, 2001a, 2002, 2003a, 2005a; Hattori and
Sotome 2013; Sotome et al. 2014).
Some species are also known from the Indian region (Malabar coast and southern India,
Deccan, Ganges Plain flanks of the Himalayas, Ceylon) in South Asia, and/or from the North
and East Australian region (northern forests, Queensland forests, south-eastern forests,
Tasmania), the Polynesian region (e. g., Mariana, Palau, and Caroline Islands) and the Fijian
8

region (Santa Cruz, Banks, New Hebrides, Fiji, Tonga, Samoa) in Oceania. These species can
be classified as tropical Asian or tropical Asian and Oceanian species.
3.3. Lowland Malesia endemic species
At present, several species described by Corner from lowland areas of the Malesian region are
reported only from this region (Corner 1983, 1984, 1987, 1989a, b, 1991, 1992; Hattori 2000,
2001a, b, 2002, 2003a, b, 2005a; Dai and Li 2012; Hattori and Sotome 2013). As mentioned
above, however, exact distribution patterns are still unknown for many of them, because only
limited specimens are available for each. The following species are possible examples of
endemic species in lowland Malesia because of the multiple known localities and/or frequent
collections that are limited to this region: Antrodiella diffluens (Corner) T. Hatt., Coltricia
salpincta Corner, Corneroporus subcitrinus (Corner) T. Hatt., Coriolopsis suberosifusca
(Corner) T. Hatt., Daedalea aurora, D. fulvirubida (Corner) T. Hatt., D. lusor (Corner) T.
Hatt., Datronia perstrata (Corner) T. Hatt., Fomitiporella caryophylii (Racib.) T. Wagner &
M. Fisch., Inonotus duostratosus (Lloyd) P.K. Buchanan & Ryvarden, Microporellus
grandiporus Corner, P. decurrata Corner, Roseofavolus eos (Corner) T. Hatt., Skeletocutis
basifusca (Corner) T. Hatt., Trichaptum suberosum Corner, and Tyromyces armeniacus
(Corner) T. Hatt. (Ryvarden and Johansen 1980; Corner 1987, 1989a, b, 1991; Hattori 2001b,
2002, 2003b; Dai and Li 2012; Lee et al. 2012; Hattori and Sotome 2013).
3.4. Species common to lowland Malesia and temperate East Asia
Some polypores have been reported both from lowland Malesia and cool temperate East Asia:
Bjerkandera adusta (Willd.) P. Karst., Coltricia cinnamomea (Jacq.) Murrill, Gloeoporus
dichrous (Fr.) Bres., Polyporus arcularius (Batsch) Fr., Skeletocutis nivea (Jungh.) Jean
Keller, Trametes hirsuta (Wulfen) Lloyd, and Trichaptum biforme (Fr.) Ryvarden (Hattori and
Lee 2003). These species show circumglobal distribution in the Northern Hemisphere, and
their distribution centers likely exist in temperate areas.
More species are shared by warm temperate East Asia and lowland Malesia, and most of
the shared species show continuous distribution from tropical to warm temperate areas of East
Asia. Many of them are more frequently and widely recorded in tropical than temperate areas,
9

and it is thought that their distribution centers exist in tropical areas. These widespread species
are considered as pantropic, paleotropic, or Asian tropical species and include species such as:
Coriolopsis glabrorigens (Lloyd) Núñez & Ryvarden, C. strumosa (Fr.) Ryvarden,
Hymenochaete poroides T. Wagner & M. Fisch. [= Cyclomyces tabacinus (Mont.) Pat.],
Lenzites elegans (Spreng.) Pat., L. vespaceus (Pers.) Pat., Neofomitella rhodophaea (Lév.)
Y.C. Dai, Hai J. Li & Vlasák, Nigroporus vinosus (Berk.) Murrill, Perenniporia ochroleuca
(Berk.) Ryvarden, Phellinus lamaensis (Murrill) Pat., Rigidoporus lineatus (Pers.) Ryvarden,
Trametes lactinea (Berk.) Sacc., Trichaptum byssogenum (Jungh.) Ryvarden, and T. durum
(Jungh.) Corner (Aoshima and Ogimi 1974; Corner 1987, 1989a, b, 1991; Núñez and
Ryvarden 2001; Hattori 2005b; Dai 2012).
Microporus affinis (Blume & T. Nees) Kuntze is one of the most frequent species in both
warm temperate to subtropical Castanopsis forests of Japan and lowland rainforests of Malesia
(Hattori 2005b; Fukasawa et al. 2012; Hattori et al. 2012). This species is usually found on
evergreen species of Fagaceae in Japan (Imazeki and Hongo 1989) and is also frequently on
these trees in lowland Malaysia (Hattori et al. 2012). Distribution centers for species found on
evergreen Fagaceae trees may exist in warm temperate to subtropical areas where
Castanopsis Spach, Quercus L. subgenus Cyclobalanopsis (Oersted) Schneider, and
Lithocarpus Blume species dominate the natural forests.
Many of the species found in both temperate and tropical areas, however, produce simple
and/or resupinate basidiomata with limited distinctive morphological characters.
Fomitiporia torreyae Y.C. Dai is a temperate Asian species known from China and Japan, and
has a morphologically similar but phylogenetically distinct population in Malaysia (Ota et al.
2014). Tinctoporellus epimiltinus (Berk. & Broome) Ryvarden was considered a widely
distributed species in tropical to temperate areas in Asia, but at least some of the temperate
population represents a distinct species (Yuan and Wan 2012).
Other “species” with a disjunct distribution in tropical and cool temperate areas may
actually include cryptic species with different distribution patterns.
Aporpium hexagonoides A. David & Jaq. was once considered a species with a disjunct
distribution in tropical and cool temperate Asia (Núñez 1998), but the temperate population
was shown to be a distinct species, A. strigosum Sotome & T. Hatt., after molecular and
10

morphological investigations (Sotome et al. 2014). Reevaluations based on morphological and
phylogenetic characteristics are desirable for polypores that are widespread from tropical to
temperate areas to ascertain exact distributions of these fungi.
3.5. Highland Malesian species
Fungal species associated with temperate trees can be expected from highland areas above
1200 m of the Malesian region regardless of latitude (Hattori et al. 2012). Consequently,
mycobiota in these areas are expected to be distinct from those of lowland areas in the same
region. Corner (1983, 1984, 1987, 1989a, b, 1991, 1992) made intensive collections in
highland Malesia including Mt. Kinabalu (Borneo), Fraser’s Hill (the Malay Peninsula),
Cameron Highland (the Malay Peninsula), and Brastagi (Sumatra), and recorded 96 polypore
species from these areas. Of these species, 49 are common to lowland areas of this region,
whereas 18 are common to temperate Asia and 29 are currently known to be endemic to
highland areas of the Malesian and the Papuan regions.
Several species show disjunct distributions in highland Malesia and cool temperate areas
of East Asia. Antrodiella aurantilaeta (Corner) T. Hatt. and A. brunneimontana (Corner) T.
Hatt. (= A. ussurii Y.C. Dai) are known from singleton specimens in the type locality (Mt.
Kinabalu), but both are frequently collected in cool temperate to boreal areas in Japan and
Northeast China (Hattori and Ryvarden 1993; Dai and Niemelä 1997; Hattori 2002).
Tyromyces incarnatus Imazeki, which is widely distributed in temperate areas in Japan, was
also recorded from Mt. Kinabalu as T. roseipileus Corner based on a specimen collected in Mt.
Kinabalu (Hattori 2003a). Bondarcevomyces taxi (Bondartsev) Parmasto was reported from
Mt. Kinabalu under the name of Hapalopilus olivascens Corner, although Hattori (2002)
overlooked this synonymy. This species is also known from boreal to temperate areas of
Eurasia, but its occurrence is very rare (Bondartsev 1953). Paratrichaptum accuratum
Corner is another species recorded both from highland Malesia and Japan, but has a wider
distribution in the latter; it was first described from a highland forest in Sumatra, and later
collected from boreal to subtropical areas in Japan [Hokkaido Pref., Yubari, Numaosawa, 21
Sep 1990, leg. T. Hattori (TFM: F-15818); Kagoshima Pref., Amami Is., Kamiya, 28 Oct
1989, leg. T. Hattori (TFM: F-15268); Nara Pref., Mt. Wasamata, 30 Aug 1994, leg. N. Iwata
11

(TFM: F-17203); Yamagata Pref., Asahi Kosen, 28 Oct 1992, leg. T. Hattori (TFM: F-16692)].
Cerrena zonata (Berk.) H.S. Yuan [= Antrodiella zonata (Berk.) Ryvarden] was considered
a species with a disjunct distribution in temperate areas of East Asia to India, Australia, and
New Zealand (Núñez and Ryvarden 2001). This species, however, was also reported from a
mountain area of Mt. Kinabalu under the name of Heteroporus kinabaluensis Corner (Corner
1987) and by my personal observation at Cameron Highland, Malaysia [23 Nov 2005, alt.
1600–1900 m (TFM: F-23806)], which indicates that this may be widely distributed in
highland areas of the Malesian region. Accordingly, C. zonata was revealed to be a species
distributed in temperate and low latitude highland areas along the West Pacific Rim. The above
species are currently unknown from tropical lowland areas, and considered to be adapted to
lower temperature.
Several species have only been reported from highland areas in the Malesian and Papuan
regions. Exact distribution patterns are unknown for many of them because only limited
specimens are available, but some species including Meripilus applanatus Corner,
Buglossoporus malesianus Corner [= Piptoporus malesianus (Corner) T. Hatt.], Tyromyces
dolosus Corner, and T. friabilis Corner were recorded from multiple highland areas but are
unknown from lowland areas within these regions; they are more likely endemic species in
highland areas in tropical Southeast Asia.
However, some possible “endemic” species in highland Malesia and New Guinea have
morphologically similar counterparts in temperate Asia with a similar morphology (Table 2).
Buglossoporus malesianus recorded from highland areas of West Malaysia (Cameron
Highland) and Borneo (Mt. Kinabalu) is very similar to B. quercinus (Schrad.) Kotl. & Pouzar,
but has larger pores and may lack rufescence (Hattori 2000). Meripilus applanatus (Corner) T.
Hatt. is recorded from the Cameron Highlands, Mt. Kinabalu, and Sumatra (Brastagi, alt. 1700
m) in addition to the type locality in the Solomon Islands (alt. 600–700 m). This species is
characterized by smaller pores and more globose and shorter basidiospores than
M. giganteus (Pers.) P. Karst. (Hattori 2000). Detailed comparative studies including
morphological and phylogenetic investigations are needed to confirm species delineations.
About half of the species recorded from highland Malesia are also seen in lowland areas.
Among these fungi, several species, such as the following, are not seen, even in subtropical
12

areas of Asia, which indicates that they are species restricted to equatorial regions:
Abundisporus roseoalbus (Jungh.) Ryvarden, Flavodon cervinogilvus (Jungh.) Corner,
Microporellus fuliginosus Corner, M. inusitatus (Lloyd) Corner, and Trichaptum sprucei
(Berk.) Rajchenb. & Bianchin (Corner 1987, 1989a). In highland Malesia, such tropical
species show sympatric distributions with temperate species that are completely unknown from
tropical lowland areas, which indicates that temperature and other climatic conditions are not
the only factors that determine polypore distribution.
4. Ecological and biological characteristics related to polypore distribution
4.1. Host preference
It is widely believed that host and substrata distribution primarily controls fungal distribution
(Bisby 1943). As in other temperate to boreal areas, polypore distribution in temperate Asia is
related to that of their host trees, because many polypores in this area have host specificity or
preference (Hattori 2005b; Yamashita et al. 2010). Alhough host specificity of polypores and
other wood-inhabiting aphyllophoraceous fungi was widely considered to be low in tropical
areas, some polypores have specificity or preference for certain tree families or genera in
Malaysia. Among the 45 frequent species in Pasoh Forest Reserve, Daedalea aurora, D.
dochmia (Berk.) T. Hatt., Erythromyces crocicreas (Berk. & Broome) Hjortstam & Ryvarden,
Perenniporia corticola, and Phellinus fastuosus (Lév.) S. Ahmad were always associated with
Dipterocarpaceae trees (Hattori et al. 2012). Some additional specialists of this tree family
may be expected on less frequent species in this area. Dipterocarpaceae specialist presence
may produce distinct polypore mycobiota in the lowland Malesian region compared with those
in the Papuan region and highland forests in the Malesian region where Dipterocarpaceae trees
are sparse or absent.
Fagaceae is the most important hardwood tree family that influences polypore distribution
in temperate East Asia. Several polypores have preference for Fagus, Quercus and
Castanopsis spp., and distributions of these tree genera influence the polypore mycobiota in
temperate Japan (Hattori 2005b; Yamashita et al. 2010). In highland areas of the Malesian
region, Castanopsis, Lithocarpus, and Quercus are frequent tree genera. As already
mentioned, Microporus affinis show some preference for Fagaceae trees, even in lowland
13

Malesia. Host trees of Hymenochaete cyclolamellata T. Wagner & M. Fisch. [= Cyclomyces
fuscus Kuntze ex Fr.] and Fomitiporia robusta (P. Karst.) Fiasson & Niemelä recorded from
highland Malesia by Corner (1991) are unknown but are possibly associated with Fagaceae
trees because they are mostly found on Castanopsis and other Fagaceae trees in East Asia.
Many polypores have high specificity to coniferous trees in temperate to boreal areas
(Ryvarden and Gilbertson 1993, 1994). Presence of abundant specialists on pine trees makes
the polypore mycobiota in pine forests distinct from those of other forest types (Hattori
2005b). Pinus L. (Pinaceae), Agathis Salisb. (Araucariaceae), Podocarpus Persoon
(Podocarpaceae), and Nageia Gaertn. (Podocarpaceae) are important conifer tree genera
distributed in the Malesian region, although their distributions are mostly restricted to highland
areas. Pinus merkusii Jungh. & de Vriese is naturally distributed in highland areas of Sumatra,
but no Pinus species are distributed in other areas of Malaysia and Indonesia (Ashton 2014),
other than several Pinus spp. that are artificially planted in highland forests.
Pinus spp. including P. merkusii and P. kesiya Royle ex Gordon are widely distributed in the
Indo-Chinese region, and several species with high preference for
Pinus spp. were collected in Thailand (Hjortstam and Ryvarden 1982). Lack of
Pinus spp. in most parts of the Malesian region may be a factor that characterizes polypore
mycobiota in this region.
Agathis is a characteristic tree genus that is widespread in highland forests of the Malesian
and the Papuan regions. This genus is species-rich in northern Borneo, but they as well as
Araucaria spp. can form distinct stands east of Wallace's Line (Ashton 2014). Corner listed
only a limited number of polypore species on Agathis, but he described Inonotus agathidis
Corner based on a collection made on stumps of Agathis in a highland forest of Mt. Kinabalu
(Corner 1991). He also reported Heterobasidion annosum (Fr.) Bref. on trunks of
Agathis and Podocarpus (Corner 1989a), although it is possible that this was a
misidentification of H. araucariae P.K. Buchanan, which are known to occur on Araucariaceae
trees and Pinus spp. (Buchanan 1988).
Strong host preference was also reported for some of the aphyllophoraceous fungi on
mangrove trees in tropical areas (Gilbert and Sousa 2002; Gilbert et al. 2008); therefore, tree
species composition in mangrove forests may influence polypore mycobiota in forests. Three
14

species of Fulvifomes Murrill newly described from Thailand are only known to occur on
Xylocarpus granatum K.D. Koenig., which indicates that they also have strong preference for
this tree species (Hattori et al. 2014). No information is available for the host range of
polypores on mangrove in the Malesian region yet, but similar phenomena can be expected for
this region.
In addition, limited polypores may have strong preference for less frequent trees in the
Malesian region (Hattori et al. 2012). Distributions of some polypores are limited as a result
of their host specificity in this region, and tree flora is considered a biotic environmental
variable that defines polypore mycobiota.
4.2. Response to physical environmental factors
As described above, polypore mycobiota changes along latitudinal and altitudinal gradients.
Host tree distribution is at least partly responsible for this change regarding polypores with
strong host preferences. Many of the wood-inhabiting polypores, however, do not show
distinct host specificity in lowland Malesia. Hattori et al. (2012) showed that most of the
frequent wood-inhabiting polypores were recorded on multiple tree families except for the
Dipterocarpaceae tree specialists in Pasoh Forest Reserve. Among these species, 17 were
recorded on at least five different tree families, and most can be regarded as generalists
(Hattori et al. 2012). Regarding latitudinal distribution, all of the 17 species are also known
from subtropical to tropical areas of the Indo-Chinese region, but the following 10 species are
unknown or sparsely recorded in temperate areas of East Asia, and likely restricted to tropical
and subtropical areas: Antrodiella liebmanii (Fr.) Ryvarden, Coriolopsis albobadia (Lloyd) T.
Hatt. & Sotome [= C. retropicta (Lloyd) Teng], Earliella scabrosa (Pers.) Gilb. & Ryvarden,
Ganoderma mastoporum (Lév.) Pat., Grammothele lineata Berk. & M.A. Curtis, Hyphodontia
ovispora (Corner) T. Hatt., Rigidoporus hypobrunneus (Pers.) Corner, R. microporus (Sw.)
Overeem, Stecchericium seriatum (Lloyd) Maas Geest., and Trichaptum durum (Jungh.)
Corner (Aoshima and Ogimi 1974; Núñez and Ryvarden 2001; Dai 2012). Distribution of
generalists restricted to tropical and subtropical areas is possibly delimitated by temperature
and other climatic conditions instead of host distribution.
Mswaka and Magan (1999) examined the impact of temperature and osmotic potential on
15

growth of nine Trametes spp. and five other wood-inhabiting fungi, and concluded that species
distributed in hot and dry areas have tolerance against a wide temperature range, and some
show very high thermotolerance. Alternatively, Humphrey and Siggers (1933) studied the
impact of temperature on several wood-inhabiting fungi, and classified them into low-
temperature, intermediate, and high-temperature groups. Several species included in the high-
temperature group, growth best above 32 ˚C, are actually distributed in temperate to boreal
areas, which indicates that optimal growth temperature is not always the factor that
differentiates temperate and tropical species. For some tree pathogenic fungi, winter survival
linked to low temperatures is a limiting factor for their distribution (Desprez-Loustau et al.
2007b). Minimum temperature for survival can be another factor that delimitates distribution
of polypores and other fungi mainly distributed in tropical and/or low-latitude regions.
Some polypores are physiologically adapted to hot and dry environments (Mswaka and
Magan 1999). In the Malesian region, Earliella scabrosa (Pers.) Gilb. & Ryvarden,
Pycnoporus sanguineus (L.) Murrill and Trametes lactinea (Berk.) Pat. are more commonly
and frequently seen in open areas than in primary forests (Corner 1989b), and may have a
preference for dry environments. Regarding higher plants in this region, the flora in Java is
distinct from those in other parts of the Sunda Shelf region, such as Borneo, Sumatra, and the
Malay Peninsula, but more similar to west Wallacea, including Sulawesi and the Lesser Sunda
Islands, because of the dry monsoon climate in Java (Raes and van Welzen 2009; Welzen and
Raes 2011). Therefore, it is possible that amount and pattern of precipitation can influence
polypore distribution in the Malesian region as in higher plants.
4.3. Dispersal ability
Distribution patterns of some higher plants are explained by vicariance due to continental
drift, glacier distributions, and other geographical histories (Pielow 1992). Some macrofungi
show Gondwanan and Grayan distributions, although they are not always directly related to
those of the geographical histories (Petersen and Hughes 2007). Ancient long-distance
dispersal is another possible factor that explains their wide distributions. Moncalvo and
Buchanan (2008) examined isolates of the Ganoderma applanatum-australe complex
worldwide, and suggested that this complex diversified much later than the break-up of
16

Gondwana from Laurasia and the final separation of the Gondwanan landmasses from each
other. They concluded that the wide distribution of this complex was due to long-distance
dispersal. Similar results were also shown for the Armillaria mellea complex, which is a
widely distributed group that causes wood decay (Coetzee et al. 2011).
Hibbett (2001) discussed the biogeography of Lentinula Earle, which is a wood-inhabiting
agaric genus, based on molecular clocks. He concluded that most of the dispersal of this genus
can be explained by short dispersal via island hopping and one long-distance dispersal event
instead of vicariance due to the continental drift. It is likely that the wide distributions of many
wood-inhabiting fungi are caused by dispersal rather than migration through land drift.
Alternatively, some polypores are distributed within limited areas, and several species are
restricted to lowland Malesia but unreported even from neighboring regions. Spore dispersal
abilities were examined for selected species in northern Europe, and that of a rare species,
Fomitopsis rosea (Alb. & Schwein.) P. Karst., is much shorter than those of common and
widespread species (Stenlid and Gustafsson 2001). No information is available regarding spore
dispersal ability of Malesian polypores with limited distributions yet, but lack of long-
dispersal ability could also be one factor that explains their distributions.
Human-mediated invasion is another possible factor that modifies fungal distributions.
Several plant and animal pathogenic fungi have been introduced to non-native areas, resulting
in epidemics caused by exotic pathogens (Desprez-Loustau et al. 2007a). Linzer et al. (2008)
showed that Heterobasidion annosum, a pathogenic polypore to trees, underwent human-
mediated dispersal from America to Italy quite recently, which was determined using a
phylogeographic approach. Some saprophytic wood-inhabiting basidiomycetes are also
considered invasive species. Vizzini et al. (2009) revealed that Favolaschia calocera R. Heim
was introduced from Madagascar or Asia to New Zealand and Italy.
At present, there is no clear evidence for any introduced polypores in the Malesian region,
but the species highly associated with exotic trees are potentially also exotic.
Pinus spp. are widely planted in highland areas of Malesia where these trees are not naturally
distributed, and some introduced species are expected to exist on them, although little
information is available for those on Pinus in Malesia.
Regarding plantations, exotic tree crops widely planted in the Malesian region include
17

Hevea brasiliensis Müll. Arg., Acacia mangium Willd., and Elaeis guineensis Jacq. Most
polypores seen in Hevea and Acacia plantations, however, also occur in secondary natural
forests in Malesia (Yamashita et al. 2008; Hattori et al. 2012), and species specific to these
trees are unknown. Elaeis guineensis is a monocotyledon, and polypores on this plant are
distinct from those on most hardwood trees. However, species on Elaeis are also seen on other
domestic palms, bamboos, or other monocotyledons based on my personal observation, and no
specialist on Elaeis is known among Malesian polypores, at present.
5. Characteristic genera/species of polypores in Malesia
Among polypores described from the Malesian region, some species are currently considered
to belong to peculiar monotypic genera that are only known from Asia. Some other species are
now accommodated in widely distributed genera, but are morphologically very distinct or
genetically isolated from other species in the genera. Here, I review some examples of such
species to highlight peculiar polypore mycobiota in this region.
Corneroporus subcitrinus (Corner) T. Hatt. Mycoscience 42: 426 (2001).
Fig. 2A.
Corner (1989a) described this fungus as a member of Boletopsis Fayod because of the
terrestrial and stipitate basidiocarps, and ornamented basidiospores. Hattori (2001b), however,
transferred it to a monotypic genus Corneroporus T. Hatt., because it has echinulate and
hyaline basidiospores, lacks dark coloration when touched, and lacks hyphal swelling in the
context, whereas other Boletopsis spp. have irregularly angular and colored basidiospores,
nigrescent basidiocarps, and swollen context hyphae.
Phylogenetic position of this fungus is still unknown, but most of its morphological
characters are shared with members of Bankeraceae Donk except for the poroid hymenophore
(Hattori 2001b). Another possible relative is Diacanthodes Singer with similar stipitate
basidiocarps, poroid hymenophore and ornamented basidiospores, although
D. novoguineensis (Henn.) O. Fidalgo, the type species, has a dimitic hyphal system with
generative and skeletal hyphae, and basidiospores with a dextrinoid reaction.
This fungus is known from lowland areas of the Malay Peninsula and Borneo (Corner
1989a; Hattori 2001b; Yamashita et al. 2009). It is rare, and possibly associated with old
18

growth forests in the Malesian region. For a detailed description of this species, see Hattori
(2001b).
Paratrichaptum accuratum Corner, Beih. Nova Hedwig. 86: 137 (1987).
Fig. 2B.
Corner described this species based on a single specimen collected at a highland forest in
Sumatra. Later, it was also collected from boreal to subtropical forests in Japan. This fungus is
the type species of a monotypic genus, Paratrichaptum Corner, described by Corner (1987).
This is a peculiar genus with large and irregular pores, dark-colored context, a monomitic
hyphal system, and light brown basidiospores. It is a brown rot fungus based on my personal
observation of Japanese specimens.
Its phylogenetic position is yet to be examined, but most of the above characteristics are
shared with several species that belong to Boletales. Other poroid and wood-inhabiting species
of this order, however, have sappy and mostly brittle context, in contrast to that
P. accuratum, which has a fibrous-corky context. Gloeophyllales also include brown rot fungi
with dark corky context and irregular hymenophore, but no species of this order produce
distinctly colored basidiospores. In addition, they usually have a dimitic hyphal system with
scattered to abundant skeletal hyphae, whereas the present fungus has a purely monomitic
hyphal system both in context and hymenophore. For a detained description of this species, see
Corner (1987).
Postia stellifera T. Hatt. & Sotome, Mycotaxon 114: 154 (2010).
Fig. 2C.
Hattori et al. (2010) described this species based on two specimens collected in lowland forests
of the Malay Peninsula. The macromorphology of this fungus is similar to that of species
belonging to Albatrellus Gray, but a molecular phylogenetic study revealed that it is related to
some Postia spp. that cause a brown rot. This is a very peculiar species within
Postia Fr., and characterized by the presence of a distinct stipe, terrestrial habitat, and
distinctly verrucose chlamydospores in the context.
This fungus was placed in Postia based on nuclear ribosomal RNA large subunit gene
sequences, but “Postia s.l. clade” detected by Hattori et al. (2010) only had weak support, and
it was nested in this clade with a long branch. Comprehensive phylogenetic studies on
19

Postia and its related genera might reveal that this represents a member of a distinct genus
related with Postia.
The type locality is a secondary forest near the road, and the second collection was made at
Taman Negara, a well-preserved forest. Therefore, it is likely a rare species, but not a species
restricted to old-growth areas.
Roseofavolus eos (Corner) T. Hatt., Mycoscience 44: 457 (2003).
Fig. 2D.
Corner (1989a) described this as a member of Grifola Gray because of the stipitate and
lignicolous basidiocarps and a monomitic hyphal system, although G. frondosa (Fr.) Gray, the
type species of this genus, actually has a dimitic hyphal system. Hattori (2003b) transferred it
to a monotypic genus Roseofavolus T. Hatt. because it has basidiocarps with a single pileus, a
monomitic hyphal system, and cylindrical and thin-walled hymenial cystidia, which are
unknown in Grifola and other polypore genera.
The phylogenetic position of this species is still yet to be studied. Its macromorphology is
similar to some species of Polyporus P. Micheli and Favolus Fr., but its monomitic hyphal
system and thin-walled cystidia are unknown from these genera. Fleshy context, somewhat
gelatinous and pinkish hymenophore, monomitic hyphal system with indistinctly thick-walled
generative hyphae in context, and presence of cylindrical cystidia are similar to
Gloeostereum Imai (Cyphellaceae; Petersen and Parmasto 1993), although hymenophore in the
latter is smooth. Phylogenetic studies are desirable to make its taxonomic position clear.
This species is always associated with well-decomposed wood and likely a wood-decaying
fungus at the later stage, but for the time being it was never isolated from context tissue or
basidiospores. For a detailed description of this, see Hattori (2003b).
6. Conclusion
The Malesian region is rich in polypore species with various distribution patterns. Many of
these species are considered tropical species whose distribution centers exist in tropical areas,
whereas some of the species seen in highland areas are widely recorded from temperate areas,
and considered temperate species. Many polypore species are only known from the Malesian
or adjacent regions at present, and several of them are rare and have only been recorded from
20

singleton or limited specimens. Phylogenetic studies have revealed that some of the widely
distributed “species” actually include cryptic species restricted to Malesia and adjacent
regions, which indicates that more species are expected to be endemic to tropical Asia than
currently recognized. Molecular phylogenetic studies using specimens collected in and around
this region, morphological investigations on these specimens, and detailed local inventories
within these regions will reveal more accurate distributions for these species. Only limited
information is available at present regarding the factors that delimitate distribution of endemic
polypores to this and adjacent regions, but some polypore species have specificity or
preferences for certain tree families/genera that are frequent in this region, and distribution of
their host trees likely delimitates their distributions. However, for most species without host
preferences, the causes of their restricted distributions are yet to be revealed. Understanding
the factors that delimitate distribution may also contribute to conservation of the rare and
threatened species distributed in and around this region.
Disclosure
The authors declare no conflicts of interest. All the experiments undertaken in this study
comply with the current laws of Japan and other countries where they were performed.
Acknowledgments
I greatly appreciate Prof. Leif Ryvarden (University of Oslo) for his kind guidance for studying
tropical polypores, Prof. Roy Watling (Royal Botanic Garden Edinburgh) and other staff of
RBGE for their kind hospitality during my stay to let me examine polypore specimens
collected by E. J. H. Corner. I also thank Lee Su See (Forest Research Institute Malaysia), E.
B. Gareth Jones (King Saud University), Yuko Ota (Nihon University), Kozue Sotome (Tottori
University), Satoshi Yamashita (Tokushima University), Jariya Sakayaroj (BIOTEC), Rattaket
Choeyklin (Burapha University), Baharuddin Salleh (University of Sains Malaysia),
Vikineswary Sabaratnam (University of Malaya), Thi Bee Kin (Forest Research Institute
Malaysia) and all of my co-workers who work with tropical Asian polypores. Curators of the
21

following herbaria kindly sent polypore specimens on loan that contributed to my
understanding of tropical Asian polypores: BPI, FH, K, NY, O, PC, RO. This research was
partly supported by a Grant-in-Aid for Scientific Research, JSPS Japan (No. 21405023).
22

REFERENCES
Aoshima K, Ogimi C, 1974. Distribution of the Aphyllophorales in the Ryukyu Islands (in
Japanese). Memoirs of the National Science Museum 7: 85–88.
Ashton P, 2014. On the forests of tropical Asia. Kew Publishing, Richmond.
Bian LS, Dai YC, 2015. Coltriciella globosa and C. pseudodependens spp. nov.
(Hymenochaetales) from southern China based on morphological and molecular
characters. Mycoscience 56: 190–197; http://dx.doi.org/10.1016/j.myc.2014.06.001.
Bisby GR, 1943. Geographical distribution of fungi. The Botanical Review 9: 466–482.
Bondartsev AS, 1953. The Polyporaceae of the European USSR and Caucasia
(translated from Russian, 1971). Israel Program for Scientific Translations, Jerusalem.
Buchanan P, 1988. A new species of Heterobasidion (Polyporaceae) from Australasia.
Mycotaxon 32: 325–337.
Cesati V, 1879. Mycetum in itinere Borneensi lectorum a cl. Od. Beccari.
Atti della Reale Accademia della Science Fisiche e Matematiche di Napoli
8: 1–28 (not seen).
Choeyklin R, Hattori T, Jones EBG, 2011. A checklist of aphyllophoraceous fungi in Thailand:
part 1. New records. Mycosphere 2: 161–177.
Coetzee MPA, Bloomer P, Wingfield MJ, Wingfield BD, 2011. Paleogene radiation of a plant
pathogenic mushroom. PLoS ONE 6: e28545;
http://dx.doi.org/10.1371/journal.pone.00228545.
Corner EJH, 1983. Ad Polyporaceas I. Amauroderma and Ganoderma. Beihefte Nova
Hedwigia 75: 1–182.
Corner EJH, 1984. Ad Polyporaceas II & III. Beihefte Nova Hedwigia 78: 1–222.
Corner EJH, 1987. Ad Polyporaceas IV. The genera Daedalea, Flabellophora, Flavodon,
Gloeophyllum, Heteroporus, Irpex, Lenzites, Microporellus, Nigrofomes, Nigroporus,
Oxyporus, Paratrichaptum, Rigidoporus, Scenidium, Trichaptum, Vanderbylia, and
Steccherinum. Beihefte Nova Hedwigia 86: 1–265.
Corner EJH, 1989a. Ad Polyporaceas V. The genera Albatrellus, Boletopsis, Coriolopsis
(dimitic), Cristelloporia, Diacanthodes, Elmerina, Fomitopsis (dimitic), Gloeoporus,
Grifola, Hapalopilus, Heterobasidion, Hydnopolyporus, Ischnoderma, Loweporus,
Parmastomyces, Perenniporia, Pyrofomes, Stecchericium, Trechispora, Truncospora and
Tyromyces. Beihefte Nova Hedwigia 96: 1–218.
Corner EJH, 1989b. Ad Polyporaceas VI. The genus Trametes. Beihefte Nova Hedwigia
23

97: 1–197.
Corner EJH, 1991. Ad Polyporaceas VII. The xanthochroic polypores.
Beihefte Nova Hedwigia 101: 1–175.
Corner EJH, 1992. Additional resupinate non-xanthochroic polypores from Brazil and
Malesia. Nova Hedwigia 55: 119–152.
Dai YC, 2012. Polypore diversity in China with an annotated checklist of Chinese polypores.
Mycoscience 53: 49–80; http://dx.doi.org/10.1007/s10267-011-0134-3.
Dai YC, Li HJ, 2012. Type studies on Coltricia and Coltriciella described by E. J. H. Corner
from Southeast Asia. Mycoscience 53: 337–346; http://dx.doi.org/10.1007/S10267-011-
0174-8.
Dai YC, Niemelä T, 1997. Changbai wood-rotting fungi 6. Study on Antrodiella, two new
species and notes on some other species. Mycotaxon 64: 67–81.
Davies SJ, Noor NSM, LaFrankie JV, Ashton PS, 2003. The trees of Pasoh forest: stand
structure and floristic composition of the 50-ha forest research plot. In:
Okuda T, Manokaran N, Matsumoto Y, Niiyama K, Thomas SC (eds), Pasoh ecology of a
lowland rain forest in Southeast Asia. Springer, Tokyo, pp 35–50.
Desprez-Loustau M-L, Robin C, Buée M, Courtecuisse R, Garbaye J, Suffert F, Sache I, Rizzo
DM, 2007a. The fungal dimension of biological invasions. TRENDS in Ecology and
Evolution 22: 473–480; http://dx.doi.org/10.1016/j.tree.2007.04.005.
Desprez-Loustau M-L, Robin C, Reynaud G, Déqué M, Badeau V, Piou D, Husson C, Marçais
B, 2007b. Simulating the effects of a climate-change scenario on the geographical range
and activity of forest-pathogenic fungi. Canadian Journal of Plant Pathology
29: 101–120; http://dx.doi.org/10.1080/07060660709507447.
Fukasawa Y, Osono T, Takeda H, 2012. Fungal decomposition of woody debris of
Castanopsis sieboldii in a subtropical old-growth forest. Ecological Research 27: 211–
218; http://dx.doi.10.1007/s11284-011-0890-8.
Gilbert GS, Gorospe J, Ryvarden L, 2008. Host and habitat preference of polypore fungi in
Micronesian tropical flooded forests. Mycological Research 112: 674–680;
http://dx.doi.org/10.1016/j.mycres.2007.11.009.
Gilbert GS, Sousa WP, 2002. Host specialization among wood-decay polypore fungi in a
Caribbean mangrove forest. Biotropica 34: 396–404; http://dx.doi.org/10.1111/j.1744-
7429.2002.tb00553.x.
Graff, PW, 1916. Bibliography and new species of Philippine fungi.
24

Mycologia 6: 253–288.
Han ML, Cui BK, 2015. Morphological characters and molecular data reveal a new species of
Fomitopsis (Polyporales) from southern China. Mycoscience 56: 168–176;
http://dx.doi.org/10.1016.j.myc.2014.05.004.
Hattori T, 2000. Type studies of the polypores described by E.J.H. Corner from Asia and the
West Pacific I. Species described in Polyporus, Buglossoporus, Meripilus, Daedalea, and
Flabellophora. Mycoscience 41: 339–349; http://dx.doi.org/10.1007/BF02463947.
Hattori T, 2001a. Type studies of the polypores described by E.J.H. Corner from Asia and
West Pacific Areas II. Species described in Gloeophyllum, Heteroporus,
Microporellus, Oxyporus, Paratrichaptum, and Rigidoporus. Mycoscience 42: 19–28;
http://dx.doi.org/10.1007/BF02463971.
Hattori T, 2001b. Type studies of the polypores described by E.J.H. Corner from Asia and
West Pacific Areas III. Species described in Trichaptum, Albatrellus, Boletopsis,
Diacanthodes, Elmerina, Fomitopsis and Gloeoporus. Mycoscience 42: 423–431;
http://dx.doi.org/10.1007/BF02464338.
Hattori T, 2002. Type studies of the polypores described by E.J.H. Corner from Asia and West
Pacific Areas IV. Species described in Tyromyces (1). Mycoscience 43: 307–315;
http://dx.doi.org/10.1007/s102670200045.
Hattori T, 2003a. Type studies of the polypores described by E.J.H. Corner from Asia and
West Pacific Areas V. Species described in Tyromyces (2). Mycoscience 44: 265–276;
http://dx.doi.org/10.1007/s10267-003-0114-3.
Hattori T, 2003b. Type studies of the polypores described by E.J.H. Corner from Asia and
West Pacific Areas VI. Species described in Tyromyces (3), Cristelloporia, Grifola,
Hapalopilus, Heterobasidion, Ischnoderma, Loweporus, and Stecchericium. Mycoscience
44: 453–463; http://dx.doi.org/10.1007/s10267-003-0139-7.
Hattori T, 2005a. Type studies of the polypores described by E.J.H. Corner from Asia and
West Pacific Areas VII. Species described in Trametes (1). Mycoscience 46: 303–312;
http://dx.doi.org/10.1007/s10267-005-0250-z.
Hattori T, 2005b. Diversity of wood-inhabiting polypores in temperate forests with different
vegetation types in Japan. Fungal Diversity 18: 73–88.
Hattori T, Lee SS, 2003. Community structure of wood-decaying Basidiomycetes in Pasoh. In:
Okuda T, Manokaran N, Matsumoto Y, Niiyama K, Thomas SC (eds) Pasoh: ecology of a
lowland rain forest in southern Asia. Springer, Tokyo, pp 163–170; http://dx.doi.org/
10.1007/978-4-431-67008-7_12.
Hattori T, Ryvarden L, 1993. Antrodiella aurantilaeta (Polyporaceae, Basidiomycotina), a new
25

polypore to Japan. Transaction of the Mycological Society of Japan 34: 363–368.
Hattori T, Sakayaroj J, Jones EBG, Suetrong S, Preedanon S, Klaysuban A, 2014. Three
species of Fulvifomes (Basidiomycota, Hymenochaetales) associated with rots on mangrove
tree Xylocarpus granatum in Thailand. Mycoscience 55: 344–354;
http://dx.doi.org/10.1016.j.myc.01.001.
Hattori T, Sotome K, 2013. Type studies of the polypores described by E.J.H. Corner from
Asia and West Pacific Areas VIII. Species described in Trametes (2). Mycoscience 54: 297–
308; http://dx.doi.org/10.1016/j.myc.2012.10.008.
Hattori T, Sotome K, Ota Y, Thi BK, Lee SS, Salleh B, 2010. Postia stellifera sp. nov., a
stipitate and terrestrial polypore from Malaysia. Mycotaxon 114: 151–161;
http://dx.doi.org/10.5248/114.151.
Hattori T, Yamashita S, Lee SS, 2012. Diversity and conservation of wood-inhabiting
polypores and other aphyllophoraceous fungi in Malaysia. Biodiversity and Conservation
21: 2375–2396; http://dx.doi.org/10.1007/210531-012-0238-x.
Heilmann-Clausen J, Boddy L, 2008. Distribution patterns of wood-decay Basidiomycetes at
the landscape to global scale. In: Boddy L, Frankland JC, van West P (eds),
Ecology of saprotrophic Basidiomycetes. Academic Press, London, pp 263–275;
http://dx.doi.org/10.1016/S0275-0287(08)80016-1.
Hennings P, 1900. Fungi. Monsunia 1: 1–38.
Hibbett DS, 2001. Shiitake mushrooms and molecular clocks: historical biogeography of
Lentinula. Journal of Biogeography 28: 231–241; http://dx.doi.org/10.1046/j.1365-
2699.2001.00528.x.
Hjortstam K, Ryvarden L, 1982. Aphyllophorales from Northern Thailand.
Nordic Journal of Botany 2: 273–281.
Humphrey CJ, Siggers PV, 1933. Temperature relations of wood-destroying fungi.
Journal of Agricultural Research 47: 997–1008.
Imazeki R, Hongo T, 1989. Colored illustrations of mushrooms of Japan vol. II
(in Japanese). Hoikusha, Osaka.
Lee SS, Alias SA, Jones EBG, Zainuddin N, Chang HT, 2012. Checklist of fungi of Malaysia.
Forest Research Institute Malaysia Research Pamphlet 132: 1–556.
26

Li HJ, Cui BK, Dai YC, 2014. Taxonomy and multi-gene phylogeny of
Datronia (Polyporales, Basidiomycota). Persoonia 32: 170–182;
http://dx.doi.org/10.3767/003158514X681828.
Linzer RE, Otrosina WJ, Gonthier P, Bruhn J, Laflamme G, Bussieres G, Garbelotto M, 2008.
Influences on the phylogeography of the fungal pathogen Heterobasidion annosum,
including evidence of interspecific horizontal genetic transfer and of human-mediated,
long-range dispersal. Molecular Phylogenetics and Evolution 46: 844–862;
http://dx.doi.org/10.1016/j.ympev.2007.12.010.
Moncalvo J-M, Buchanan PK, 2008. Molecular evidence for long distance dispersal across the
Southern Hemisphere in the Ganoderma applanatum-australe species complex
(Basidiomycota). Mycological Research 112: 425–436; http://dx.doi.org/
10.1016/j.mycres.2007.12.001.
Mswaka AY, Magan N, 1999. Temperature and water potential relations of tropical
Trametes and other wood-decay fungi from the indigenous forests of Zimbabwe.
Mycological Research 103: 1309–1317.
Murrill WA, 1907. Some Philippine Polyporaceae. Bulletin of the Torrey Botanical Club
34: 465–481.
Murrill WA, 1908a. A collection of Philippine polypores. Leaflets of Philippine Botany
1: 262–271.
Murrill WA, 1908b. Additional Philippine Fungi. Bulletin of the Torrey Botanical Club
35: 391–416.
Myers N, 1988. Threatened biotas: “hot spots” in tropical forests.
The Environmentalists 8: 187–208.
Núñez M, 1998. The genus Elmerina (Heterobasidiomycetes) in Japan. Folia Cryptogamica
Estonica 33: 99–101.
Núñez M, Ryvarden L, 2001. East Asian polypores vol 2. Synopsis Fungorum 14: 170–522.
Núñez M, Suhirman, Stokland JN, 2002. Patterns of polypore distribution in the Lesser Sunda
Islands, Indonesia. Is Wallace’s Line significant? In: Watling R, Frankland JC, Ainsworth
AM, Isaac S, Robinson CH (eds), Tropical mycology, vol. 1. macromycetes.
27

CAB International, Wallingford, pp 73–86;
http://dx.doi.org/10.1079/9780851995427.0073.
Ota Y, Hattori T, Nakamura H, Terashima Y, Lee SS, Miyuki Y, Sotome K, 2014. Taxonomy
and phylogenetic position of Fomitiporia torreyae, a causal agent of trunk rot on Sanbu-
sugi, a cultivar of Japanese cedar (Cryptomeria japonica) in Japan. Mycologia 106: 66–76;
http://dx.doi.org/10.3852/13-045.
Petersen RH, Hughes KW, 2007. Some agaric distribution patterns involving Pacific
landmasses and Pacific Rim. Mycoscinece 48: 1–44; http://dx.doi.org/10.1007/s10267-006-
0333-5.
Petersen RH, Parmasto E, 1993. A redescription of Gloeostereum incarnatum. Mycological
Research 97: 1213–1216; http://dx.doi.org/10.1016/S0953-7562(09)81287-2.
Pielow EC, 1992. Biogeography. Krieger Publishing Company, Malabar.
Quanten E, 1997. The polypores (Polyporaceae s.l.) of Papua New Guinea.
Opera Botanica Belgica 11: 1–352.
Raes N, van Welzen PC, 2009. The demarcation and internal division of Flora Malesiana:
1857–present. Blumea 54: 6–8; http://dx.doi.org/10.3767/000651909X475888.
Ryvarden L, 1973. Type studies in the Polyporaceae 1. Species described by C. H. Persoon.
Persoonia 7: 305–312.
Ryvarden L, 1975. Type studies in the Polyporaceae 7. Species described by J. M. Berkeley
from 1836 to 1843. Kew Bulletin 31: 81–103.
Ryvarden L, 1976. Type studies in the Polyporaceae 4. Species described by J. F. Klotzsch.
Memoirs of the New York Botanical Garden 28: 109–207.
Ryvarden L, 1977a. Type studies in the Polyporaceae 9. Species described by E. M.
Wakefield. Mycotaxon 5: 331–336.
Ryvarden L, 1977b. Type studies in the Polyporaceaea 10. Species described by J. M.
Berkeley, either alone or with other authors, from 1844 to 1855.
Norwegian Journal of Botany 24: 213–230.
Ryvarden L, 1981a. Type studies in the Polyporaceae 12. Species described by F. W.
Junghuhn. Persoonia 11: 369–372.
Ryvarden L, 1981b. Type studies in the Polyporaceae 13. Species described by J. H. Léveillé.
Mycotaxon 13: 175–186.
Ryvarden L, 1982. Type studies in the Polyporaceae 11. Species described by J. F. C.
Montagne, either alone or with other authors. Nordic Journal of Botany 2: 75–84.
Ryvarden L, 1983. Type studies in the Polyporaceae 14. Species described by N. Patouillard,
28

either alone or with other mycologists. Occasional Papers of the Farlow Herbarium of
Cryptogamic Botany 18: 1–39.
Ryvarden L, 1984. Type studies in the Polyporaceae 16. Species described by J. M. Berkeley,
either alone or with other mycologists from 1856 to 1886. Mycotaxon 20: 329–363.
Ryvarden L, 1985. Type studies in the Polyporaceae 17. Species described by W. A. Murrill.
Mycotaxon 23: 169–198.
Ryvarden L, 1988a. Type studies in the Polyporaceae 19. Species described by M. C. Cooke.
Mycotaxon 31: 45–58.
Ryvarden L, 1988b. Type studies in the Polyporaceae 20. Species described by G. Bresadola.
Mycotaxon 33: 303–327.
Ryvarden L, 1989. Type studies in the Polyporaceae 21. Species described by C. G. Lloyd in
Cyclomyces, Daedalea, Favolus, Fomes and Hexagonia. Mycotaxon 35: 229–236.
Ryvarden L, 1990. Type studies in the Polyporaceae 22. Species described by C. G. Lloyd in
Polyporus. Mycotaxon 38: 83–102.
Ryvarden L, 1991. Genera of polypores nomenclature and taxonomy. Synopsis Fungorum
5: 1–363.
Ryvarden L, 1992. Type studies in the Polyporaceae 23. Species described by C. G. Lloyd in
Lenzites, Polystictus, Poria and Trametes. Mycotaxon 44: 127–136.
Ryvarden L, 2012. Type studies in Polyporaceae 27. Species described by Ch. Hennings.
Czech Mycology 64: 13–21.
Ryvarden L, Gilbertson RL, 1993. European polypores part 1. Synopsis Fungorum 6: 1–387.
Ryvarden L, Gilbertson RL, 1994. European polypores part 2. Synopsis Fungorum 7: 388–
743.
Ryvarden L, Johansen I, 1980. A preliminary polypore flora of East Asia. Fungiflora, Oslo.
Saccardo PA, 1888. Sylloge fungorum omnium hucusque cognitorum vol. 6. Sumptibus
auctoris, Typis seminarii, Patavii.
Sotome K, Akagi Y, Lee SS, Ishikawa NK, Hattori T, 2013. Taxonomic study of
Favolus and Neofavolus gen. nov. segregated from Polyporus (Basidiomycota,
Polyporales). Fungal Diversity 58: 245–266; http://dx.doi.org/10.1007/s13225-012-0213-6.
Sotome K, Hattori T, Ota Y, Lee SS, Vikineswary S, Abdullar N, Kakishima M, 2009.
Taxonomic study of Echinochaete (Polyporaceae, Basidiomycota) and description of
E. maximipora sp. nov. Mycological Progress 8: 123–132;
http://dx.doi.org/10.1007/s11557-009-0584-6.
Sotome K, Maekawa N, Nakagiri A, Lee SS, Hattori T, 2014. Taxonomic study of Asian
species of poroid Auriculariales. Mycological Progress 13: 987–997;
29

http://dx.doi.org/10.1007/s11557-014-0984-0.
Stenlid J, Gustafsson M, 2001. Are rare wood decay fungi threatened by inability to spread?
Ecological Bulletins 49: 85–91.
Suhirman, Núñez M, 1995. Indonesian Aphyllophorales I. A literature survey.
Bulletin Kebun Raya Indonesia 8: 69–78.
Suhirman, Núñez M, 1998a. Indonesian Aphyllophorales II. Species of Polyporaceae in the
Herbarium Bogoriensis. Bulletin Kebun Raya Indonesia 9: 107–109.
Suhirman, Núñez M, 1998b. Indonesian Aphyllophorales 3. Poroid and stereoid species from
Kerinci-Seblat National Park, Western Sumatra. Mycotaxon 68: 273–292.
Takhtajan A, 1961. Flowering plants. Origin and distribution (translated from Russian, 1969).
Oliver & Boyd, Edinburgh.
Tokuda S, Hattori T, Dai YC, Ota Y, Buchanan PK, 2009. Three species of
Heterobasidion (Basidiomycota, Hericiales), H. parviporum, H. orientale sp. nov. and
H. ecrustosum sp. nov. from East Asia. Mycoscience 50: 190–202;
http://dx.doi.org/10.1007/s10267-008-0476-7.
Vizzini A, Zotti M, Mello A, 2009. Alien fungal species distribution: the study case of
Favolaschia calocera. Biological Invasions 11: 417–429; http://dx.doi.org/10.1007/s10530-
008-9259-5.
Welzen PC, Raes N, 2011. The floristic position of Java. Gardens' Bulletin Singapore
63: 329–339.
Wright SJ, 2002. Plant diversity in tropical forests: a review of mechanisms of species
coexistence. Oecologia 130: 1–14; http://dx.doi.org/10.1007/s004420100809.
Yamashita S, Hang SM, Hattori T, 2009. List of polypores and other aphyllophoraceous fungi
collected in the Lambir Hills National Park, Sarawak, Malaysia.
Contributions from the Biological Laboratory, Kyoto University 29: 1–24.
Yamashita S, Hattori T, Abe H, 2010. Host preference and species richness of wood-inhabiting
aphyllophoraceous fungi in a cool temperate area of Japan. Mycologia 102: 11–19;
http://dx.doi.org/10.3852/09-008.
Yamashita S, Hattori T, Lee SS, Okabe K, 2015. Estimating the diversity of wood-decaying
polypores in tropical lowland rain forests in Malaysia: the effect of sampling strategy.
Biodiversity and Conservation 24: 393–406; http://dx.doi.org/10.1007/s10531-014-0815-2.
Yamashita S, Hattori T, Momose K, Nakagawa M, Aiba M, Nakashizuka T, 2008. Effects of
forest use on aphyllophoraceous fungal community structure in Sarawak, Malaysia.
30

Biotropica 40: 354–362; http://dx.doi.org/10.1111/j.1744-7429.2007.00366.x.
Yuan HS, Wan XZ, 2012. Morphological and ITS rDNA-based phylogenetic identification of
two new species in Tinctoporellus. Mycological Progress 11: 947–952;
http://dx.doi.org/10.1007/s11557-012-0810-5.
Zhao CL, Cui BK, 2013. Morphological and molecular identification of four new resupinate
species of Perenniporia (Polyporales) from southern China. Mycologia 105: 945–958;
http://dx.doi.org/10.3852/12-201.
Zhao CL, Shen LL, Cui BK, 2014. Perenniporia cinereofusca sp. nov. (Polyporales,
Basidiomycota) evidenced by morphological characters and phylogenetic analysis.
Mycoscience 55: 417–422; http://dx.doi.org/10.1016/j.myc.2013.11.016.
Zhou LW, 2014. Fulvifomes hainanensis sp. nov. and F. indicus comb. nov.
(Hymenochaetales, Basidiomycota) evidenced by a combination of morphology and
phylogeny. Mycoscience 55: 70–77; http://dx.doi.org/10.1016/j.myc.2013.05.006.
Figure legends
Fig. 1 – Map of the floristic regions of Southeast Asia modified from Takhtajan (1961). A:
The Eastern Asian region. B: The Indo-Chinese region. C: The Malesian region. D: The
Papuan region. E: The North and East Australian region.
Fig. 2 – Basidiocarps of some peculiar polypores distributed in the Malesian region. A:
Corneroporus subcitrinus (Holotype, E00159738). B: Paratrichaptum accuratum (TFM: F-
15818). C: Postia stellifera (Holotype, TFM: F-20668). D: Pore surface of Roseofavolus eos
(TFM: F-17426). Bars: 2 cm.
31