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Review
Occurrence of indoor wood decay basidiomycetes
in Europe
Ji
r
ı GABRIEL*, Karel
SVEC
Laboratory of Environmental Microbiology, Institute of Microbiology AS CR, V
ıde
nsk
a 1083, 142 20 Prague 4, Kr
c,
Czech Republic
article info
Article history:
Received 1 March 2017
Received in revised form
9 May 2017
Accepted 9 May 2017
Keywords:
Basidiomycetes
Fungi
Indoor
Serpula lacrymans
abstract
Brown-rot fungi are considered to be the most important wood-inhabiting fungi econom-
ically, as they also deteriorate the wood that has been used in buildings. In the northern
hemisphere, coniferous wood is the main source of interior structural timber. White-rot
fungi, which degrade lignin and preferentially attack hardwood, are less common.
Emphasis is usually placed on Serpula lacrymans or Coniophora puteana, which are the
most common indoor basidiomycetes found in buildings in Europe. In this review, we sum-
marize available data on the occurrence of wood decay fungi in the Czech Republic, Poland,
Germany (both former East and West), Belgium, France, Norway, Denmark, Finland, Latvia,
Estonia, Romania and Albania reported in the past few decades. The total number of occur-
rences was near 20,000; original data were collected between 1946 and 2008. The most
abundant basidiomycetes were S. lacrymans and C. puteana, with the exception of Norway,
where the genus Antrodia was the most frequent.
ª2017 British Mycological Society. Published by Elsevier Ltd. All rights reserved.
1. Introduction
Indoor wood decay fungi cause many problems worldwide;
fungi that invade roofs, walls, ceilings, etc., represent a group
of various basidiomycetes that are in many cases resistant to
currently used fungicides. These fungi attack and damage
wooden houses and other wooden constructions, and the
most well-known of these, Serpula lacrymans (often regarded
as the “cancer of buildings”), is responsible for many millions
of USD of damage each year (Palfreyman, 1995). For example,
the cost of fungal damage in France was estimated to be
approximately V30 million yearly (Maurice et al. 2011), and
in the UK, the cost of repairing fungal damage to timber
used in construction amounted to £3 million per week
(Rayner and Boddy 1988). The dry rot remediation business
in the UK was estimated to be worth an excess of £400 million
(Krzyzanowski et al. 1999).
The decay of wood and wood-based products usually be-
gins when the spores or mycelial fragments adhere to the
wood surface. Wood moisture and temperature are the most
important features in terms of the “building habitat”.
Humphrey and Siggers (1933) previously studied the effect of
temperature on the growth rate of 56 wood-decay fungi and
found that none would grow below 12 C and that most would
not grow above 40 C. According to many authors and guide-
lines for the protection of wood and wood products from
attack by decay fungi, it is important and necessary to keep
wood or wooden constructions at a moisture content below
* Corresponding author.
journal homepage: www.elsevier.com/locate/fbr
fungal biology reviews 31 (2017) 212e217
http://dx.doi.org/10.1016/j.fbr.2017.05.002
1749-4613/ª2017 British Mycological Society. Published by Elsevier Ltd. All rights reserved.
20 % (Carll and Highley 1999). Schmidt (2007) reported mini-
mum and maximum humidity requirements for fungi identi-
fied by means of ITS sequencing (to avoid unreliable data for
incorrectly identified species; Table 1). As stated by Carll and
Highley (1999), the spores of wood decay fungi do not germi-
nate and fungal hyphae do not grow at moisture levels
much below the fibre saturation point, which is at approxi-
mately 30 % moisture. However, only the part of water that
is not bound by dissolved substances (salts, sugars) is avail-
able to fungi; a detailed description of the optimum conditions
for wood decay in terms of water activity (aw), water potential
or relative air humidity (RH) can be found, e.g., in the above-
mentioned review of Schmidt (2007). However, data on fungal
development under fluctuating moisture conditions that are
more common in nature are not yet available. Other factors,
such as wood species, local climate, design details, exposure
conditions (esp. in roofs and trusses, cellars, and door frames)
and coatings might have an indirect effect on wood decay
(e.g., Thybring 2017, Meyer and Brischke 2015). Under ideal
moisture and temperature conditions, fungal growth may
occur even within days.
In theory, wood decay fungi need free water as a diffusion
medium for their extracellular digestive enzymes. In the case
of brown-rot (or dry-rot) fungi, other factors in addition to
enzymatic processes are involved in wood degradation. Low
molecular weight compounds, such as oxalate, veratryl
alcohol, variegatic acid and others (Goodell et al. 1997;
Eastwood et al. 2011; Watkinson and Eastwood 2012),
contribute to lignocellulose decay, as do Fe and most likely
other bivalent ions. These chemicals generate hydroxyl, per-
oxyl and hydroxylperoxyl radicals in Fenton and Fenton-like
systems. Inorganic elements play an important role in the
physiology and control of at least S. lacrymans growth
(Schilling 2010; Watkinson and Eastwood 2012). As has been
demonstrated (Low et al. 2000), S. lacrymans removes calcium,
silicon and iron from sandstone and calcium, sulphur and iron
from traditional plaster. The sequestered elements are located
in its hyphae, particularly in the form of calcium oxalate.
2. Detection and identification of decay fungi
Fruiting bodies are normally preferentially used for field iden-
tification (e.g., Abrego and Salcedo 2015, Nicolotti et al. 2010).
Some species rarely fructify in buildings but form mycelial
strands (cords). Frankl (2014) found vital mycelia or active
fruiting bodies only in only 7 % of his observations (but the re-
mains were found in 95 %). Many studies address the diagnos-
tics of wood decaying fungi based on their macromorphology
and micromorphology; i.e., the typical shape and colour of
fruiting bodies or spores, cell wall thickness of hyphae, type
of branching, presence of dolipore septa, clamp connections
or aggregates on the surface or inside the cells, etc. Typical
visible properties (brown or white discolouration, eventually
cracking into roughly cubical pieces) of degraded wood are
also very important. Following the crucial work of Falck
(1912), newer diagnostic keys including drawings or colour
photographs have been published, e.g., by Doma
nski (1972),
Stalpers (1978), Hanlin (1998), Schmidt (2006), Huckfeldt and
Schmidt (2006), Buchalo et al. (2009) and Stancheva et al. (2009).
Precise molecular methods were not available for the iden-
tification of indoor wood decay fungi until the 1980s. These
methods include species-specific priming PCR (SSPP), rDNA
ITS region sequence analysis, restriction fragment length
polymorphism analysis (RFLP), random amplified polymor-
phic DNA analysis (RAPD), amplified fragment length poly-
morphism analysis (AFLP) and sequence-specific
oligonucleotide probe analysis (SSO). For more information,
see the work of Maurice et al. (2011) or the recent paper by
Raja et al. (2017) and references cited herein. Methods based
on DNA analyses can provide efficient, sensitive and rapid
diagnostic tools for the detection and identification of wood
decay fungi without requiring a prior fungal isolation step
(Glaeser and Lindner 2011, Gonthier et al. 2015). In the case
of wood decay basidiomycetes, methods based on ribosomal
DNA (ITS 1 or 2 rDNA region) sequencing have been estab-
lished as routine techniques for the identification of fungi to
the species level, esp. for those that are hardly or not at all
distinguishable by species, such as Antrodia,Coniophora and
Leucogyrophana (Schmidt 2006, Jarosch and Besl, 2001; Binder
and Hibbett, 2006, Coetzee et al., 2003; Schmidt et al., 2012).
In the last ten years, sequencing technologies have changed
dramatically, offering multiple options in throughput, accu-
racy and cost for answering different biological questions.
Some other alternative techniques are based on the pro-
duction of typical volatile organic compounds (VOC) by fungi
(e.g., Schmidt and Kallow, 2005). In addition to 1-octen-3-ol
(Ewen et al. 2004), which causes the typical smell of mush-
rooms, several other compounds typical of fungi have been
described (e.g., Anton et al. 2016, Konuma et al. 2015, Korpia
Table 1 eHumidity requirements (wood moister content; %) of selected fungi with respect to the colonization and decay of
wood (after Schmidt 2007).
Species Minimum for
colonization
Minimum
for decay
Optimum
for decay
Maximum
for decay
Serpula lacrymans 21 26 45e140 240
Leucogyrophana pinastri 30 37 44e151 184
Coniophora puteana 18 22 36e210 262
Antrodia vaillantii 22 29 52e150 209
Donkioporia expansa 21 26 34e126 256
Gloeophyllum abietinum 20 22 40e208 256
Gloeophyllum sepiarium 28 30 46e207 225
Gloeophyllum trabeum 25 31 46e179 191
Occurrence of indoor wood decay 213
et al. 1998). In a study focused on wood decay fungi, Konuma
et al. (2015) reported 110 organic compounds, some of them
produced only when fungi were cultivated on wood. Based
on this, the use of dogs trained to sniff out S. lacrymans in
buildings should also be noted (Watkinson and Eastwood,
2012), mostly as a curiosity.
3. Abundance of wood decay basidiomycetes
in Europe
The abundance of indoor wood decay basidiomycetes re-
ported in Europe in the past few years is summarized in
Table 2.Schmidt (2007) also listed species found in Danish
buildings (after Bech-Andersen 1995) but without percentage
incidence; the list includes the white-rot basidiomycetes
Coprinus domesticus, Fomes fomentarius, Hyphoderma puberum,
Hypholoma fasciculare, Perenniporia medulla-panis, Phellinus
nigrolimitatus, Phlebiopsis gigantea, Physisporinus vitreus and Sis-
toterma brinkmannii; the only brown-rot fungi mentioned are
Dacromyces tortus, Daedalea quercina, Laetiporus sulphureus and
Oligoporus caesius.Alfredsen et al. (2005) also give a detailed
survey of fungi found in publications from several Nordic
countries.
A list of 40 wood decay species found in the roofs or cellars
of damaged buildings in the Czech Republic was published a
few years ago (Vampola 2008). The author did not provide
the percent incidence, but the most abundant species were
Antrodia serialis, Coniophora puteana, Coniophora marmorata,
Donkioporia expansa, Fomitopsis rosea, Gloeophyllum trabeum, P.
gigantea and S. lacrymans. The author also demonstrated the
occurrence of some rare species (Amyloporia xantha, Asteros-
troma ochroleucum, Hypochnicium bombycinum, O. caesius and
Tubulicrinis thermometrus, etc.). Unlike other fungi, A. ochroleu-
cum belongs to the rare species found in buildings; Vampola
(2008) reported only two occurrences in historical buildings
in the Czech Republic. He also stated that Coniophora confluens
is a good indicator of elevated moisture content in buildings,
and roof leaks should primarily be evaluated in this case.
The author also speculated about the origin of F. rosea in
damaged buildings. According to him, wood timbers previ-
ously infected by this fungus were used for building houses,
and F. rosea is able to survive in this substrate for many
(tens or even hundreds) years, but this speculation is of course
questionable. Nevertheless, Vampola (2008) reported the for-
mation of fruiting bodies of F. rosea on a timber removed
from the roof after being deposited in the fcourtyard of a his-
torical building over 2e3 weeks; the timber was estimated to
be ca. 200 y old. The same species was also reported in historic
monuments in Romania (Bucs
¸a and Bucs
¸a 2009b).
The most abundant basidiomycete across Europe is S.
lacrymans, as is clearly shown in Table 2, followed by C.
puteana. The literature about both fungi and their properties
or requirements is so exhaustive that it is neither necessary
nor possible to list them here. Antrodia damage seems to be
more common in Norway than in other countries. One expla-
nation for why Antrodia is most common in Norway might be
climate; other explanations might be building traditions and
the spore rain inoculum potential. Alfredsen et al. (2005)
stated that Antrodia sp. demands much more moisture than
S. lacrymans and slightly more than C. puteana since its opti-
mum lies between 35 % and 55 % wood moisture content. Ac-
cording to Bech-Andersen (1996),A. sinuosa was found mostly
in roof constructions, such as unventilated attics under the
roofing felt, often in competition with Gloeophyllum.The
occurrence of multiple fungi at one site is likely common;
Schultze-Dewitz (1985) reported that S. lacrymans was often
found together with C. puteana or Poria placenta. Recently,
Pottier et al. (2014) studied fungal contamination in homes
located in Low Normandy in France and reported that S. lacry-
mans was sometimes detected with the co-occurrence of
other basidiomycete wood decay species like such as D.
expansa;Maurice et al. (2011) repeatedly detected S. lacrymans,
D. expansa and C. puteana, and in another sample from north-
western France, the occurrence of white-rot species Trametes
versicolor and Hyphodontia sp. In a supplementary table, the
authors listed 76 isolates of S. lacrymans from Norway,
Table 2 eSpecies abundance (%) of indoor basidiomycetes reported in Europe.
Species Poland
a
BRD
b
DDR
c
DDR
c
Belgium
d
Norway
e
Latvia
f
Denmark
e
Finland
e
Estonia
g
Romania
h
Serpula lacrymans 54 27 29 22 60 16 47 20 45 79 32
Coniophora puteana 22 17 18 17 10 16 6 34 12 7 60
Antrodia vaillantii 210ee 118*13*0,1*12*e21
Antrodia sinuosa 92ee 2ee e e e e
Gloeophyllum sp. 2 6 1 e1335 2e11
Donkioporia expansa e10 ee 10 10 ee ee 21
Poria placenta ee613 eeee eee
Total number of occurrences 3050 748 1005 498 749 3434 338 8293 1237 633 n/a
Total number of species 29 31 11 11 26 35 60 n/a n/a n/a 75
Data since 1950 2000 1966 1980 1985 2001 1996 1946 1978 2002 1979
until 1960 2006 1980 1984 1991 2003 2007 1983 1988 2008 2009
Identification method** m n/a n/a n/a n/a n/a m n/a n/a m m
References:
a
Wa _
zny and Czajnik (1963);
b
Schmidt (2007);
c
Schultze-Dewitz (1985);
d
Guillitte (1992);
e
Alfredsen et al. (2005);
f
Irbe and Andersone
(2008);
g
Pilt et al. (2009);
h
Bucs
¸a and Bucs
¸a (2009a).
* Data given for Antrodia sp.; BRD ¼Bundesrepublik Deutschland (former “West Germany”), DDR ¼Deutsche Demokratische Republik (former
“East Germany”).
** m emorphology.
214 J. Gabriel, K.
Svec
Finland, the UK, Germany, Belgium, and France isolated be-
tween 1939 (Berlin, Germany) and 2011 (Brest, France). In
this study, ITS or beta-tubuline analyses were used in some
cases. Frankl (2014) described fungi from sixteen smaller
churches in different places in the Czech Republic
(1999e2012) and reported the combined occurrences of the
genera Coniophora with Gloeophyllum (3), Serpula (8), Ster-
eum (6)andTrametes (19); Gloeophyllum with Serpula (1)
or Trametes (10) and Trametes with Stereum (2)orAntrodia
(1). In some cases, he found a combination of three different
genera (e.g., Coniophora,Gloeophyllum and Trametes e5). The
repeated detection of various fungi in a single place could
make the interpretation of some tables ambiguous. In an
exhaustive work by Bucs
¸a and Bucs
¸a (2009a), the authors re-
ported the results of an investigation of more than 400 histor-
ic monuments (castles, palaces, citadels, churches of all
faiths, etc.) in Romania. The authors presented a list of 74
fungal species found in more than 1200 buildings. With the
exception of Dacrymyces stillatus, which is found mostly on
spruce shingle roofings (in 270 buildings), the most frequent
infections were caused by Hyphodontia breviseta (80), Gl. abie-
tinum (78)andsepiarium (67), T. versicolor (54), C. puteana
(45), S. hirsutum (42) and D. expansa (40). The data for
Romania given in Table 2 were taken from the summary re-
ported for wood, masonry and plastered wood buildings
together.
Wood decay basidiomycetes from unusual locations have
also been sometimes reported. For example, Lentinus suffrutes-
cens was found in mine timbering (N
emec 1941), as well as Pos-
tia stiptica and Postia caesia (Ryp
a
cek 1957). We found S.
lacrymans forming unusual white branching fruiting bodies
on timbering in a uranium mine in Doln
ıRo
z
ınka in the Czech
Republic (unpublished). Wa_
zny and Czajnik (1963) reported
the occurrence of S. lacrymans, Poria vaillantii, Corticium laeve,
Corticium byssinum and Peniophora setigera on wooden and con-
crete structures in the Warsaw subway. In another case study,
Kazartsev et al. (2014) found S.lacrymans and A. xantha, both
stated as common indoor wood decay fungi in St. Petersburg,
in wooden structures of the hotel “Mikhaylovskaya” and in a
wooden pavilion of the Narcological dispensary situated in
the historical city centre of St. Petersburg. Shumka et al.
(2010) investigated 5 post-byzantine churches in the Prespa
area (Albania), where fungal attack was caused only by C.
puteana. In another study, Kozir
og et al. (2014) studied wooden
barracks as well as wooden elements of brick buildings (doors,
floors, bunks, door and window frames, and structural walls
and beams) in the former Auschwitz II eBirkenau camp.
The authors found S. lacrymans,Corticium leave and Poria
vaporaria on bunks, beams and floors.
Although S. lacrymans is found in buildings in temperate re-
gions in Eurasia, North and South America, and Oceania
(Australia/New Zealand), in contrast to the frequent indoor
occurrence of S. lacrymans, its absence in nature has remained
an enigma for many years. In their review on S. lacrymans
(2012), Kauserud et al. reported that some of the first reliable
reports are from wooden sailing vessels in the 17th century,
where it presumably caused severe damage (Ramsbottom,
1937). Up to now, the fungus has been reported in India,
Pakistan, China, the USA, Russia (Kauserud et al. 2012) and
the Czech Republic (Kotlaba 1992, 2012).
Some works describing fungi found in the wood of urban
trees have also been published (e.g., Schmidt et al. 2012,
Guglielmo et al. 2007 and literature cited herein), but from
the viewpoint of possible building damage, they do not seem
to be very important (perhaps with the exception of Gloeophyl-
lum sp.).
4. Summary
S. lacrymans and C. puteana are the most frequently found
fungi reported in damaged buildings in Europe. This is likely
a result of the common knowledge of these fungi, which
form typical fruiting bodies. The occurrence of other wood
decay fungi may not necessarily be reported to specialized
laboratories/facilities and could not be fully included in the
available statistics. Some other questions might arise with
the development of new molecular methods for the identifica-
tion of fungi that were previously difficult to distinguish.
Nevertheless, the damage caused by S. lacrymans and C.
puteana is so well documented that there is no doubt about
their “leading role” across the Europe.
Acknowledgements
This work was supported by the Institute of Microbiology CAS
(RVO61388971) and by the Czech Science Foundation (GA
CR
17-05497S).
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