This is the author's final version of the contribution published as:
A. Saitta; A. Bernicchia; S.P. Gorjón; E. Altobelli; V.M. Granito; C. Losi; D.
Lunghini; O. Maggi; G. Medardi; F. Padovan; L. Pecoraro; A. Vizzini; A.M.
Persiani. Biodiversity of wood-decay fungi in Italy. PLANT BIOSYSTEMS.
145(4) pp: 958-968.
DOI: 10.1080/11263504.2011.633114
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http://www.tandfonline.com/doi/abs/10.1080/11263504.2011.633114
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Biodiversity of wood-decay fungi in Italy
A. Saitta , A. Bernicchia , S. P. Gorjón , E. Altobelli , V. M. Granito , C. Losi , D.
Lunghini , O. Maggi , G. Medardi , F. Padovan , L. Pecoraro , A. Vizzini & A. M.
Persiani
Abstract
Current knowledge about Italian wood-decay fungi (Basidiomycota and Ascomycota) is surveyed:
1582 taxa belonging to Ascomycota (341) and Basidiomycota (1241) have been reported, including
23 species new to science of Basidiomycota described from Italy within the last five years. Evaluating
diversity of wood-decay fungi can provide a more accurate estimation of species richness for fungi
which are an important functional component of ecosystems. Aphyllophoroid and Ascomycota
species play an important role in habitat conservation and management. Sardinia, Sicily, the Alps and
the Apennines are “hot spots” for wood-decay Basidiomycota in Italy.
Keywords: Ascomycota, Basidiomycota, biodiversity, Italy, wood-decay fungi,
Introduction
Dead wood constitutes a large part of total biomass of native forests, approximately 20–30% in
mature forest ecosystems (Boddy & Watkinson 1995 Boddy, L and Watkinson, S C. 1995. Wood
decomposition, higher fungi, and their role in nutrient redistribution. Can J Bot, 73(Suppl. 1): S1377–
S1383. [CrossRef]), containing large pools of carbon and nutrients (Allen et al. 1997 Allen, R B,
Clinton, P W and Davis, M R. 1997. Cation storage and availability along a Nothofagus forest
development sequence in New Zealand. Can J For Res, 27: 323–330. [CrossRef], [Web of Science
®]). Studies carried out in the northern hemisphere have shown that logs host an extremely diverse
range of fungi (e.g. Andersen & Ryvarden 2001 Andersen, H and Ryvarden, L. 2001. Wood
inhabiting fungi on Populus tremula. Windahlia, 24: 37–48. ; Boddy 2001 Boddy, L. 2001. Fungal
community ecology and wood decomposition processes in angiosperms: From standing tree to
complete decay of coarse woody debris. Ecol Bull, 49: 43–56. ; Edmonds & Lebo 1998 Edmonds, R
L and Lebo, D S. 1998. Diversity, production, and nutrient dynamics of fungal sporocarps on logs in
an old-growth temperate rain forest, Olympic National Park, Washington. Can J For Res, 28: 665–
673. [CrossRef], [Web of Science ®]; Heilmann-Clausen & Christensen 2005 Heilmann-Clausen, J
and Christensen, M. 2005. Wood-inhabiting fungi in Danish beech-forests – conflicting diversity
patterns and their implications in a conservation perspective. Biol Conserv, 122: 633–642.
[CrossRef], [Web of Science ®]). The presence of fungi in dead trees can also directly facilitate the
colonization by species dependent on fungal mycelia or fruiting bodies, such as, for example many
insects and their parasites (Komonen et al. 2000 Komonen, A, Penttila, R, Lindgren, M and Hanski,
I. 2000. Forest fragmentation truncates a food chain based on an old-growth forest bracket fungus.
Oikos, 90: 119–126. [CrossRef], [Web of Science ®]; Komonen 2003 Komonen, A. 2003.
Distribution and abundance of insect fungivores in the fruiting bodies of Fomitopsis pinicola. Ann
Zool Fenn, 40: 495–504. [Web of Science ®]; Jonsell & Nordlander 2004 Jonsell, M and Nordlander,
G. 2004. Host selection patterns in insects breeding in bracket fungi. Ecol Entomol, 29: 697–705.;
Jonsell et al. 2005 Jonsell, M, Schroeder, M and Weslien, J. 2005. Saproxylic beetles in high stumps
of spruce: Fungal flora important for determining the species composition. Scand J For Res, 20: 54–
62.) and other species of fungi (Niemelä et al. 1995 Niemelä, T, Renvall, P and Penttilä, R. 1995.
Interactions of fungi at late stage of wood decomposition. Ann Bot Fen, 32: 141–152.). The structural
diversity of forest ecosystems is clearly an important factor in maintaining forest biodiversity
(Franklin et al. 2002 Franklin, J F, Spies, T A, Van Pelt, R, Carey, A B, Thornburgh, D ARae Berg,
D. 2002. Disturbances and structural development of natural forest ecosystems with silvicultural
implications, using Douglas_fir forests as an example. Forest Ecol Manag, 155: 399–423.;
Lindenmayer et al. 2006 Lindenmayer, D B, Franklin, J F and Fischera, J. 2006. General management
principles and a checklist of strategies to guide forest biodiversity conservation. Biol Conserv, 131:
433–445. [CrossRef], [Web of Science ®]). Wood-decay fungi are defined in this article as fungi
which are found on or inhabiting woody substrata. The majority (∼98%) of described fungal species
are members of the dikarya clade, which includes the two phyla Ascomycota and Basidiomycota
(James et al. 2006 James, T Y, Kauff, F, Schoch, C, Matheny, P B, Hofstetter, VCox, C J. 2006.
Reconstructing the early evolution of Fungi using a six-gene phylogeny. Nature, 443: 818–822.).
These will be analyzed in this article.
The evaluation of woody fungal biodiversity in Italy is still in progress. Ecological and distributive
data on lignicolous macromycetes in Italy are not organized and still need a review analysis
(Venturella et al. 2011 Venturella, G, Altobelli, E, Bernicchia, A, Di Piazza, S, Donnini, DGargano,
M L. 2011. Fungal biodiversity and in situ conservation in Italy. Plant Biosyst, 145: 953–960. ). A
positive step in reaching a more consistent organization of this information was the publication of the
Check-list of Italian Fungi (Basidiomycota) by Onofri et al. (2005 Onofri, S, Bernicchia, A, Filipello
Marchisio, V, Padovan, F, Perini, CRipa, C. 2005. Checklist dei funghi italiani – Checklist of Italian
fungi, Basidiomycetes 1–380. Sassari C. Delfino ed) which includes a high number of wood-decay
fungi, and the publication of other checklists (Bernicchia 2001 Bernicchia, A. 2001. A checklist of
Corticioid, Polyporoid and Clavarioid fungi (Basidiomycetes) from the Emilia-Romagna region,
Italy. Sydowia, 53: 1–33. ; Saitta et al. 2004 Saitta, A, Bernicchia, A and Venturella, G. 2004.
Contributo alla conoscenza dei funghi lignicoli della Sicilia. Inform Bot Ital, 36: 192–202. ; Gorjón
et al. 2006 Gorjón, S P, Bernicchia, A and Gibertoni, B T. 2006. Aphyllophoraceous wood-inhabiting
fungi on Arbutus unedo in Italy. Mycotaxon, 98: 159–162.; Bernicchia et al. 2007b Bernicchia, A,
Savino, E and Gorjón, S P. 2007b. Aphyllophoraceous wood-fungi on Pinus spp. in Italy. Mycotaxon,
101: 5–8.,c,d, 2008a,b), monographs (Polyporaceae s.l. by Bernicchia 2005 Bernicchia, A. 2005.
Polyporaceae s.l. Fungi Europaei , 10. Ed, 808Alassio, Italy: Candusso. ; Corticiaceae s.l. by
Bernicchia & Gorjón 2010 Gorjón, S P and Bernicchia, A. 2010. The genus Dendrothele
(Basidiomycota) in Italy, an update with notes on European species. Nova Hedwigia, 90: 233–250.;
Ascomycota by Medardi 2006 Medardi, G. 2006. Ascomiceti d'Italia. Trento: A.M.B. Centro Studi
Micologici, 454Trento: Associazione Micologica Bresadola. ) and papers (Venturella et al. 2007
Venturella, G, Bernicchia, A and Saitta, A. 2007. Contribution to the knowledge of diversity and
distribution of lignicolous fungi from Sicily (southern Italy). Bocconea, 21: 291–295. ; Saitta &
Venturella 2010 Saitta, A and Venturella, G. 2010. La biodiversità dei funghi lignicoli in Sicilia.
Micologia Italiana, 39: 12–17. ) concerning wood-decay fungi at both regional (Sicily, Sardinia and
Emilia Romagna) and national levels. Besides these works, published data still need to be organized
to better define the geographic pattern of this group of fungi, to demonstrate the ecological diversity
within the group and to provide a more accurate estimation of the richness of these species in Italy.
By using more recent published and unpublished data, the aim of this study is to contribute to the
knowledge of wood-decay fungal biodiversity in Italy. The systematic arrangement in this article
follows Kirk et al. (2008 Kirk, P M, Cannon, P F, Minter, D W and Stalpers, J A. 2008. Dictionary
of the fungi. , 10th ed, 771UK: CABI Europe. ) and is also based on Blackwell et al. (2006 Blackwell,
M, Hibbett, D S, Taylor, J W and Spatafora, J W. 2006. Research coordination networks: A phylogeny
for kingdom Fungi (Deep Hypha). Mycologia, 98: 829–837.), James et al. (2006 James, T Y, Kauff,
F, Schoch, C, Matheny, P B, Hofstetter, VCox, C J. 2006. Reconstructing the early evolution of Fungi
using a six-gene phylogeny. Nature, 443: 818–822.) and Hibbett et al. (2007 Hibbett, D S, Binder,
M, Bischoff, J F, Blackwell, M, Cannon, P FEriksson, O E. 2007. A higher-level phylogenetic
classification of the Fungi. Mycol Res, 111: 509–547.).
Ecology, systematics and
basidiomycetes in Italy
distribution
of
wood-decay
The phylum Basidiomycota includes 16 classes, 52 orders, around 180 families and more than 31,000
species. Three subphyla are accepted, Agaricomycotina, Pucciniomycotina and Ustilaginomycotina
(Kirk et al. 2008 Kirk, P M, Cannon, P F, Minter, D W and Stalpers, J A. 2008. Dictionary of the
fungi. , 10th ed, 771UK: CABI Europe. ). In this study, we focus on two groups of wood-decay
Basidiomycota: aphyllophoroid and non-aphyllophoroid. The aphyllophoroid group includes the
Amylocorticiales K.H. Larss., Manfr. Binder & Hibbett, Atheliales Jülich, Cantharellales Gäum.,
Corticiales K.H. Larss., Gloeophyllales Thorn, Gomphales Jülich, Hymenochaetales Oberw.,
Jaapiales Manfr. Binder, K.H. Larss. & Hibbett, Polyporales Gäum., Thelephorales Corner ex
Oberw., Trechisporales K.H. Larss. and some species belonging to the Agaricales Underw., Boletales
E.-J. Gilbert and Russulales Kreisel ex P.M. Kirk, P.F. Cannon & J.C. David. The non-aphyllophoroid
group includes the Agaricales, Auriculariales J. Schröt., Cantharellales (not aphyllophoroid),
Dacrymycetales Henn., Exobasidiales Henn., Helicobasidiales R. Bauer, Begerow, J.P. Samp., M.
Weiss & Oberw., Platygloeales R.T. Moore, Russulales Kreisel ex P.M. Kirk, P.F. Cannon & J.C.
David, Septobasidiales Couch ex Donk and Tremellales Fr.
The up-to-date total number of wood-decay Basidiomycota in Italy are 1241 taxa, which represent
78% of the fungal taxa listed in this work. Among them, 770 species are aphyllophoroid and 471
species are non-aphyllophoroid. This number is continuously changing and at least 23 taxa new to
science of Basidiomycota have been described during the last five years in Italy.
The study of aphyllophoroid wood-inhabiting fungi enjoyed a golden period in Bresadola's time and
the town of Trento became a datum point for all the contemporary mycologists. Later, this branch of
mycological research was neglected for many decades. Only since the beginning of the 1980s of the
last century this important and numerous group of fungi has been consistently studied again. Some
mycologists, both academics and amateurs, kept alive the tradition begun by the Italian mycologists.
Before 2005, we only have reprints listing observations of species from forests and reserves, although
these cover many Italian regions. The aphyllophoroid fungi are included in a heterogeneous group
composed generally of species ranging from resupinate to effuse-reflexed or pileate. During the last
decades, with the introduction of the use of molecular data in taxonomy, the classification of the
Aphyllophorales has dramatically changed. They form highly polyphyletic groups distributed over
several families and orders among the homobasidiomycetes and most of them form basal clades
among the major groups of Agaricales and Boletales (Binder & Hibbett 2002 Binder, M and Hibbett,
D S. 2002. Higher-level phylogenetic relationships of homobasidiomycetes (mushroom-forming
fungi) inferred from four rDNA regions. Mol Phylogenet Evol, 22: 76–90.; Hibbett et al. 2007
Hibbett, D S, Binder, M, Bischoff, J F, Blackwell, M, Cannon, P FEriksson, O E. 2007. A higherlevel phylogenetic classification of the Fungi. Mycol Res, 111: 509–547.; Larsson 2007 Larsson, K
H. 2007. Re-thinking the classification of corticioid fungi. Mycol Res, 111: 1040–1063.; Binder et al.
2010 Binder, M, Larsson, K H, Matheny, P B and Hibbett, D S. 2010. Amylocorticiales ord. nov. and
Jaapiales ord. nov.: Early diverging clades of Agaricomycetidae were dominated by corticioid forms.
Mycologia, 102: 865–880.).
In this article, we update the Italian data of aphyllophoroid wood-decay fungi to 770 species (online
Appendix A) (456 corticioid, 265 polypores, 45 tomentelloid, 4 cyphelloid). They are distributed in
the following orders: Agaricales (49), Amylocorticiales (8), Atheliales (42), Boletales (12),
Cantharellales (43), Corticiales (15), Gloeophyllales (8), Gomphales (3), Hymenochaetales (137),
Jaapiales (1), Polyporales (290), Russulales (89), Thelephorales (41), and Trechisporales (32).
The aphyllophoroid fungi include organisms with several ecological strategies. Most are saprobic
while some are mycorrhizal (some tomentelloid species, the genera Albatrellus Gray and Boletopsis
Fayod) or parasites. Among the polypores, it is worth highlighting some species because of their
pathogenicity. These include Heterobasidion annosum (Fr.) Bref., H. abietinum Niemelä &
Korhonen, Inonotus hispidus (Bull.) P. Karst., Laetiporus sulphureus (Bull.) Murrill, Phaeolus
schweintzii (Fr.) Pat., Phellinus chrysoloma (Fr.) Donk, P. tuberculosus (Baumg.) Niemelä, and
Porodaedalea pini (Brot.) Murrill. Polypores and corticioid fungi colonize wood of a very wide range
of plants during all decay stages; some are strictly specific and grow only on a single plant family or
more rarely, on a sole species. They have been recorded mainly on the following substrata (online
Appendix B): 267 species on Quercus spp., 238 on Pinus spp., 210 on Abies alba Mill., 186 on Fagus
sylvatica L., 170 on Picea abies (L.) H. Karst., 70 on Castanea sativa Mill., 115 on Juniperus spp.,
67 on Ulmus spp., 64 on Salix spp., 55 on Arbutus unedo L., 35 on Taxus baccata L.
More than 20 species new to science have been described during the last five years; they have been
recorded in just a few Italian regions and several species have been described and successively
recorded exclusively in Italy, such as Aleurodiscus ilexicola Bernicchia & Ryvarden, Antrodia
macrospora Bernicchia & De Domincis, Antrodiella semistipitata Bernicchia & Ryvarden,
Botryobasidium sassofratinoense Bernicchia & G. Langer, Byssomerulius pirottae (Bres.) Hjortstam,
Ceriporia sulphuricolor Bernicchia & Niemelä, Ceriporiopsis guidella Bernicchia & Ryvarden,
Cristinia artheniensis Baici & Hjortstam, Echinodontium ryvardenii Bernicchia & Piga, Fomitopsis
labyrinthica Bernicchia & Ryvarden, Hyphoderma etruriae Bernicchia, Lindtneria brevispora
Bernicchia & Gorjón, L. hydnoidea Bernicchia & Ryvarden, Phanerochaete parvispora Sheng H.
Wu & Losi, Phlebia capitata Bernicchia & Gorjón, Tomentellopsis pulchella K[otilde]ljalg &
Bernicchia and Vararia maremmana Bernicchia (Bernicchia 2005 Bernicchia, A. 2005.
Polyporaceae s.l. Fungi Europaei , 10. Ed, 808Alassio, Italy: Candusso. ; Bernicchia & Gorjón 2010
Gorjón, S P and Bernicchia, A. 2010. The genus Dendrothele (Basidiomycota) in Italy, an update
with notes on European species. Nova Hedwigia, 90: 233–250.). Some species, described for the first
time in Italy, have a distribution limited to certain Mediterranean or central European countries.
Antrodia sandaliae Bernicchia & Ryvarden has been collected only from dead branches of Arbutus
unedo L. in Mediterranean forests of Sardinia and central-west Spain. Antrodiella ichnusana
Bernicchia, Renvall & Arras has been recorded also from Finland and France. Dendrothele
wojewodae Pouzar was recorded in Italy, the Czech Republic and Ukraine (Gorjón & Bernicchia
2010 Gorjón, S P and Bernicchia, A. 2010. The genus Dendrothele (Basidiomycota) in Italy, an
update with notes on European species. Nova Hedwigia, 90: 233–250.). The new species Fibroporia
bohemica Bernicchia, Vampola & Prodi has been recently described morphologically and on a
molecular basis, using specimens previously ascribed to F. radiculosa (Peck.) Gilbn. & Ryvarden
(Bernicchia et al. in press); so far, it is known from Bohemia in the Czech Republic and the Italian
Alpes. Lindtneria panphyliensis Bernicchia & M.J. Larsen has a well-documented distribution in
Czech Republic, France, Italy and UK. Fibroporia citrina (Bernicchia & Ryvarden) Bernicchia &
Ryvarden, Hyphoderma crustulinum (Bres.) Nakasone, Neolentiporus squamosellus (Bernicchia &
Ryvarden) Bernicchia & Ryvarden, Phellinus juniperinus Bernicchia & S. Curreli and P. rosmarini
Bernicchia have also been recorded from France.
Among the group of species sharing a Mediterranean distribution and thus penetrating only into the
warmer European areas the following are present in Italy: Acanthophysellum dextrinoideocerussatum
Manjón, M.N. Blanco & G. Moreno, Botryobasidium asperulum (D.P. Rogers) Boidin,
Byssomerulius hirtellus (Burt) Parmasto, Ceriporia camaresiana (Bourdot & Galzin) Bondartsev &
Singer, C. griseoviolascens M. Pieri & B. Rivoire, Corticium meridioroseum Boidin & Lanq.,
Dendrophora versiformis (Berk. & M.A. Curtis) Chamuris, Duportella malenconii (Boidin & Lanq.)
Hjortstam, Hyphoderma malençonii (Manjón & G. Moreno) Manjón, G. Moreno & Hjortstam,
Hyphodermella rosae (Bres.) Nakasone, Hexagonia nitida Durieu & Mont., Inocutis tamaricis (Pat.)
Fiasson & Niemelä, Lenzites warnieri Mont. & Durieu, Lenzitopsis oxycedri Malençon & Bertault,
Oligoporus inocybe (A. David & Malençon) Ryvarden & Gilb., Peniophora meridionalis Boidin,
Phanerochaete aculeata Hallenb., P. martelliana (Bres.) J. Erikss. & Ryvarden, P. juniperinus, P.
rosmarini, Perenniporia meridionalis Decock & Stalpers, Polyporus meridionalis (A. David) H.
Jahn, Skeletocutis percandida (Malençon & Bertault) Jean Keller, Stereum reflexulum D.A. Reid,
Trametes junipericola Manjón, G. Moreno & Ryvarden and Vuilleminia pseudocystidiata Boidin,
Lanq. & Gilles. A few species show a surprising distribution. Xanthoporus syringae (Parmasto)
Audet, a species typically known from Scandinavia growing on Syringa vulgaris, has been collected
in a mixed Picea and Alnus forest in north Italy. Fibricium gloeocystidiatum Rajchenb. has been
described from the Andean Patagonian forests growing on the endemic conifer Austrocedrus chilensis
(D. Don) Pic. Serm. & Bizzarri and was collected in Sardinia on dead leaves of Osmunda regalis L.
(Arras et al. 2007 Arras, L, Bernicchia, A and Piga, A. 2007. Fibricium gloeocystidiatum
(Polyporales, Basidiomycetes) new to Europe. Mycotaxon, 100: 342–347. ). Dendrothele tetracornis
Boidin & Duhem is present in some Scandinavian countries and in two Mediterranean ones, Italy and
France. Gyrodontium sacchari (Spreng.) Hjortstam, recorded from Sardinia is the only European
collection while its main distributional pattern is Central and South America. Trechispora clancularis
(Park.-Rhodes) K.H. Larss., known in Brasil and Tanzania, is now recorded in a very small number
of European countries and in two Italian regions. Some corticioid and polyporoid species recently
collected (Bernicchia pers. comm.), have such peculiar microscopic characteristics as to be
undoubtedly new species and they will be described in further publications.
Most non-aphyllophoroid Basidiomycota belong to the order Agaricales. More than 13,000 species
representing 413 genera and 33 families have been described (Kirk et al. 2008 Kirk, P M, Cannon, P
F, Minter, D W and Stalpers, J A. 2008. Dictionary of the fungi. , 10th ed, 771UK: CABI Europe. ).
Most lignicolous species of the Agaricales are white rot agents. The production of basidiomata on
living trees or rotting wood is not necessarily indicative of saprotrophic nutrition. Some
ectomycorrhizal Agaricales may also produce basidiomata on rotten wood. Recent phylogenetic
research has shown that Singer's Agaricales sensu stricto (Singer 1986 Singer, R. 1986. The
Agaricales in modern taxonomy. , 4th ed, Koenigstein: Koeltz Scientific Book. ) roughly corresponds
to the euagarics clade (Hibbett et al. 1997 Hibbett, D S, Grimaldi, D and Donoghue, M J. 1997. Fossil
mushrooms from Cretaceous and Miocene ambers and the evolution of homobasidiomycetes. Am J
Bot, 84: 981–991.; Moncalvo et al. 2000 Moncalvo, J M, Lutzoni, F, Rehener, S A, Johnson, J and
Vilgalys, R. 2000. Phylogenetic relationships of agaric fungi based on nuclear large subunit ribosomal
DNA sequences. Syst Biol, 49: 278–305., 2002). However, molecular data have shown that an
overemphasis on spore print color, basidiome developmental pattern and some anatomical and
cytological traits has led to the establishment of many artificial groups, while some cyphelloid,
aphyllophoroid and gasteroid fungi should be included. Hibbett et al. (1997 Hibbett, D S, Grimaldi,
D and Donoghue, M J. 1997. Fossil mushrooms from Cretaceous and Miocene ambers and the
evolution of homobasidiomycetes. Am J Bot, 84: 981–991.) and Hibbett (2004 Hibbett, D S. 2004.
Trends in morphological evolution in homobasidiomycetes. Systematic Biol, 53: 889–903.), using
nuclear and mitochondrial ribosomal DNA sequences of representatives of agarics, aphyllophoroid
and gasteroid fungi (i.e. gilled, non-gilled agarics and puffballs) have shown that basidiome form and
hymenophore type are not reliable phylogenetic markers. Phylogenetic analyses focused on
Agaricales (euagarics clade) (Moncalvo et al. 2000 Moncalvo, J M, Lutzoni, F, Rehener, S A,
Johnson, J and Vilgalys, R. 2000. Phylogenetic relationships of agaric fungi based on nuclear large
subunit ribosomal DNA sequences. Syst Biol, 49: 278–305., 2002) have pointed out that some
traditional taxonomic groups are artificial (for example Cortinariaceae, Tricholomataceae,
Strophariaceae, Hygrophoraceae, Clitocybe, Omphalina and Marasmius), revealing 117
monophyletics clades. Some non-gilled resupinate, cyphelloid, and gasteroid taxa were classified as
Agaricales (e.g. Peintner et al. 2001 Peintner, U, Bougher, N L, Castellano, M, Moncalvo, M, Moser,
MTrappe, J. 2001. Multiple origins of sequestrate fungi related to Cortinarius (Cortinariaceae). Am
J Bot, 88: 2168–2179.; Binder & Bresinsky 2002 Binder, M and Bresinsky, A. 2002. Derivation of a
polymorphic lineage of Gasteromycetes from boletoid ancestors. Mycologia, 94: 85–98.;
Bodensteiner et al. 2004 Bodensteiner, P, Binder, M, Moncalvo, J-M, Agerer, R and Hibbett, D S.
2004. Phylogenetic relationships of cyphelloid homobasidiomycetes. Mol Phylogenet Evol, 33: 501–
515.; Larsson et al. 2004 Larsson, K H, Larsson, E and K[otilde]ljalg, U. 2004. High phylogenetic
diversity among corticioid basidiomycetes. Mycol Res, 108: 983–1002.; Binder et al. 2005 Binder,
M, Hibbett, D S, Larsson, K H, Larsson, E, Langer, E and Langer, G. 2005. The phylogenetic
distribution of resupinate forms across the major clades of mushroom-forming fungi
(Homobasidiomycetes). Syst Biodivers, 3: 113–157.; Matheny & Bougher 2006a Matheny, P B and
Bougher, N L. 2006a. The new genus Auritella from Africa and Australia (Inocybaceae, Agaricales):
Molecular systematics, taxonomy and historical biogeography. Mycol Prog, 5: 2–17.; Larsson 2007
Larsson, K H. 2007. Re-thinking the classification of corticioid fungi. Mycol Res, 111: 1040–1063.).
The most recent phylogenetic analysis of Agaricales was conducted by Matheny et al. (2006b
Matheny, P B, Curtis, J C, Hofstter, V, Aime, M C, Moncalvo, J MGe, Z W. 2006b. Major clades of
Agaricales: A multi-locus phylogenetic overview. Mycologia, 98: 982–995.) using a multigenic
approach on 146 genera and 238 species. Bayesian analysis recognized six monophyletic groups
which were named as Agaricoid, Tricholomatoid, Marasmioid, Pluteoid, Hygrophoroid and
Plicaturopsidoid clades. According to Hibbett et al. (2007 Hibbett, D S, Binder, M, Bischoff, J F,
Blackwell, M, Cannon, P FEriksson, O E. 2007. A higher-level phylogenetic classification of the
Fungi. Mycol Res, 111: 509–547.) and to Matheny et al. (2006b Matheny, P B, Curtis, J C, Hofstter,
V, Aime, M C, Moncalvo, J MGe, Z W. 2006b. Major clades of Agaricales: A multi-locus
phylogenetic overview. Mycologia, 98: 982–995.), the order Agaricales belongs to the subclass
Agaricomycetidae, along with the Atheliales and Boletales.
The total number of wood-decay non-aphyllophoroid taxa in Italy is 471 (online Appendix C),
belonging to the orders Agaricales (351 taxa), Auriculariales (36), Dacrymycetales (14), Tremellales
(14), Cantharellales (11), Polyporales (9), Russulales (8), Boletales (8), Gloeophyllales (4),
Gomphales (4), Exobasidiales (3), Platigloeales (6), Septobasidiales (2) and Geastrales (1). Genera
with higher numbers of taxa are Mycena (Pers.) Roussel (56), Pluteus Fr. (34), Pholiota (Fr.) P.
Kumm. (24), Psathyrella (Fr.) Quél. (17), Crepidotus (Fr.) Staude (16), Marasmiellus Murrill (11),
Exidia Fr. (10), Gymnopilus P. Karst. (10), Tremella Pers. (10), and Dacrymyces Nees (9).
Three species new to the science have been described during the last few years, Crinipellis
pedemontana A. Vizzini, Antonín & Noordel. (Vizzini et al. 2007 Vizzini, A, Antonin, V and
Noordeloos, M E. 2007. Crinipellis pedemontana sp. nov. (Agaricomycetes), a new basidiomycete
from Italy. Mycologia, 99: 786–791.), Gymnopilus maritimus Contu, Guzm.-Dáv., A. Ortega &
Vizzini (Guzmán-Dávalos et al. 2009 Guzmán-Dávalos, L, Contu, M, Ortega, A, Vizzini, A and
Herrera, M. 2009. A new species of Gymnopilus (Basidiomycota, Agaricales) from coastal sand dunes
of northern Sardinia. Mycol Prog, 8: 195–205.) and Gymnopus inexpectatus Consiglio, Vizzini,
Antonín & Contu (Vizzini et al. 2008 Vizzini, A, Consiglio, G, Antonin, V and Contu, M. 2008. A
new species within the Gymnopus dryophilus complex (Agaricomycetes, Basidiomycota) from Italy.
Mycotaxon, 105: 43–52.). Several species show a restricted national and European distribution. These
include Baeospora myriadophylla (Peck) Singer, Buchwaldoboletus lignicola (Kallenb.) Pilát,
Chaetocalathus craterellus (Durieu & Lév.) Singer, Cheimonophyllum candidissimum (Berk. &
Curt.) Singer, Chromosera cyanophylla (Fr.) Redhead, C. subbulbipes Murrill, Chrysomphalina
chrysophylla (Fr.) Clémençon, C. strombodes (Berk. & Mont.) Clémençon, Cleistocybe pleurotoides
(J. Favre) Vizzini (Vizzini 2009 Vizzini, A. 2009. Due nuove combinazioni nel genere Cleistocybe
(Basidiomycota, Agaricomycetes), con validazione di Hygrophorus pleurotoides. Micologia e
Vegetazione Mediterr, 24: 95–98. ), Clitocybe truncicola (Peck) Sacc., Clitopilus hobsonii (Berk. &
Broome) P.D. Orton, C. rhodophyllus (Bres.) Singer, Ammirati & Norvell, Coprinopsis strossmayeri
Schulzer, Crepidotus cinnabarinus Peck, C. roseoornatus Pöder & E. Ferrari, C. pedemontana, C.
sardoa Candusso, Cystoderma subvinaceum A.H. Sm., Flammulina fennae Bas, G. maritimus, G.
purpureosquamulosus Høil. (Guzmán-Dávalos et al. 2008 Guzmán-Dávalos, L, Contu, M, Ortega, A,
Vizzini, A, Herrera, M, Ovrebo, CRodríguez, A. 2008. New morphological and molecular data on
Gymnopilus purpureosquamulosus and its phylogenetic relationships among similar species.
Sydowia, 60: 41–56.), Haasiella venustissima (Fr.) Kotl. & Pouzar, Heliocybe sulcata (Berk.)
Redhead & Ginns, Hemistropharia albocrenulata (Peck) Jacobsson & E. Larss., Hydropus
atramentosus (Kalchbr.) Kotl. & Pouzar, H. trichoderma (Joss.) Singer, Marasmiellus virgatocutis
Robich, Esteve-Rav. & G. Moreno, Mycetinis epidryas (Kühner) Antonín & Noordel.,
Phaeomarasmius rimulincola (Rabenh.) P.D. Orton, Pholiota henningsii (Bres.) P.D. Orton, P. jahnii
Tjall.-Beuk. & Bas, P. subochracea (A.H. Sm.) A.H. Sm. & Hesler, P. tubercolosa (Schaeff.:Fr.) P.
Kumm., Pleurocybella porrigens (Pers.) Singer, Pluteus aurantiorugosus (Trog) Sacc., Rhodotus
palmatus (Bull.:Fr.) Maire, Simocybe maritima (Bon) Bon, S. rubi (Berk.) Singer, Sparassis brevipes
Krombh., Trichocybe puberula (Kuyper) Vizzini (Vizzini et al. 2010), and Tricholomopsis flammula
Métrod.
Analysis of available data on distribution of wood-decay basidiomycetes in Italy suggests that some
regions are very well explored while others such as the Valle d'Aosta, Umbria, Marche and Molise
are poorly investigated and further studies and inventorial field work are needed. It is not possible to
establish a conclusion about the distribution of the biodiversity due to the fragmentary survey.
Candidate species included in the draft European red list
Taking into account their distribution and ecological value, a significant number of these species, can
be considered for inclusion in the draft European red list.
Of the wood-decay Basidiomycota (online Appendix D) 194 species are among the 1644 candidates
for the European red list (http://www.wsl.ch/eccf/). Among them, the following species, because of
their environmental importance and rarity in Italy, should be included in the future European red list:
Aleurodiscus ilexicola, Amphinema diadema K.H. Larss. & Hjortstam, Amyloathelia amylacea
(Bourdot & Galzin) Hjortstam & Ryvarden, Antrodia albobrunnea (Romell) Ryvarden, A. alpina
(Litsch.) Gilb. & Ryvarden, A. pulvinascens (Pilát) Niemelä, A. sandaliae, Antrodiella ichnusana,
Atheloderma mirabile Parmasto, B. myriadophylla, Botryobasidium robustius Pouzar & Hol.-Jech.,
Ceriporia alachuana (Murrill) Hallenb., C. alba M. Pieri & B. Rivoire, C. sulphuricolor,
Ceriporiopsis guidella, C. cyanophylla, Clitocybe subbulbipes, C. truncicola, Coprinus strossmayeri,
C. cinnabarinus, C. roseo-ornatus, Crinipellis sardoa, Crustoderma longicystidiatum (Litsch.)
Nakasone, Cystoderma subvinaceum, Dendrothele amygdalispora Hjortstam, D. dryina (Pers.) P.A.
Lemke, Dentipellis fragilis (Pers.) Donk, E. ryvardenii, Fomitopsis labyrinthica, H. venustissima
(Fr.) Kotl. & Pouzar, Hjortstamia crassa (Lév.) Boidin & Gilles, Hyphoderma etruriae, Hypochnella
violacea Auersw. ex J. Schröt., Inonotus andersonii (Ellis & Everh.) Cerný, Laurilia sulcata (Burt)
Pouzar, Lenzitopsis oxycedri, Lindtneria flava Parmasto, Odonticium helgae Hjortstam & Ryvarden,
Oligoporus simanii (Pilát) Bernicchia, Peniophora tamaricicola Boidin & Malençon, Peniophorella
guttulifera (P. Karst.) K.H. Larss., Phellinus rimosus (Berk.) Pilát, Ramaricium alboochraceum
(Bres.) Jülich, Scotomyces subviolaceus (Peck) Jülich, Steccherinum robustius (J. Erikss. & S.
Lundell), Trametes junipericola, Trechispora kavinioides de Vries, Trichaptum laricinum (P. Karst.)
Ryvarden, Vuilleminia megalospora Bres., and V. pseudocystidiata, Lanq. & Gilles. In addition, there
are some very rare and interesting species absent from the preliminary list, including Chaetoderma
luna (Romell ex D.P. Rogers & H.S. Jacks.) Parmasto, C. candidissimum, Dendrothele nivosa (Berk.
& M.A. Curtis ex Höhn. & Litsch.) P.A. Lemke, Dendrothele wojewodae, F. fennae, Lindtneria
panphyliensis, M. virgatocutis, P. porrigens and Vararia maremmana.
Focal species for habitat conservation
Given the importance of fungi in ecosystem processes, several studies have suggested use of fungi as
focal species, or indicators of ecological continuity in forest ecosystems worldwide (e.g. SverdrupThygeson & Lindenmayer 2003 Sverdrup-Thygeson, A and Lindenmayer, D B. 2003. Ecological
continuity and assumed indicator fungi in boreal forest: The importance of the landscape matrix.
Forest Ecol Manag, 174: 353–363.; Blasi et al. 2010 Blasi, C, Marchetti, M, Chiavetta, U, Aleffi, M,
Audisio, PAzzella, M M. 2010. Multi-taxon and forest structure sampling for identification of
indicators and monitoring of old growth forest. Plant Biosyst, 144: 160–170.).
A group of species growing on specific endangered substrata are the most useful way to drive the
conservation of forest ecosystems. In Italy, many interesting and rare fungal species are linked to a
specific substratum and can be used, as usually happens for mammals or birds, as “umbrella species”,
which provide for conservation of the entire ecosystem.
A significant number of Basidiomycota were found in plant communities included in the priority
Habitats of Annex I of the Directive 92/43/EEC and its interpretation manual for Italy (Blasi et al.
2007 Blasi, C, Boitani, L, La Posta, S, Manes, F and Marchetti, M. 2007. “Biodiversity in Italy”. In
Contribution to the national biodiversity strategy, 460Roma: Palombi & Partner S.r.L. ) and most of
these belong to the aphyllophoroid group. The most significant habitats for their conservation are
“Endemic forests with Juniperus spp. (*9560)” “Quercus suber forests (9330)”, “Quercus ilex and
Quercus rotundifolia forests (9340)”, “Alpine Larix decidua and/or Pinus cembra forests (9420)” and
“Apennine beech forests with Abies alba and beech forests with Abies nebrodensis (*9220)”.
The cases of juniper and cork oak forests are highlighted as follow. Old Sardinian juniper (Juniperus
phoenicea L. and J. oxycedrus L.) forests very often are selective substrata for very rare species such
as E. ryvardenii, Hyphoderma etruriae, Lenzitopsis oxycedri, Neolentiporus squamosellus, Piloporia
sajanensis (Parmasto) Niemelä and Trametes junipericola. Most species associated with ancient
juniper forests occur in the Urzulei-Piraonni-Orgosolo-Lanaittu area, where some outstanding
examples have been recorded. Juniper trees are an irreplaceable substratum, and for this reason
survival of many wood-inhabiting Italian aphyllophoraceous species has become uncertain. Many
polypores restricted to old specimens of Juniperus L. follow the host genus wherever it occurs, with
a scattered distributional pattern. They may therefore be considered threatened species, and old
juniper trees should be protected (Bernicchia & Ryvarden 1988 Bernicchia, A and Ryvarden, L. 1988.
Aleurodiscus ilexicola Bernicchia & Ryvarden sp. nov. from Sardinia. Mycol Helv, 3: 83–88. ).
Mediterranean oak forests and shrublands are fragile ecosystems and some aphyllophoroid species
connected with these habitat need to be protected. A species growing specifically or preferentially on
Quercus suber L., Odonticium flavicans, is a good indicator of cork oak forests which should be
conserved.
Hot spots for wood-decay Basidiomycota in Italy
Some areas such as the Alps and the Apennines, present a notable diversity of polypores and
corticioids. Several aphyllophoroid fungi have been recorded exclusively from the Alps. These
include Antrodia alpina, Antrodiella semistipitata, A. serpula (P. Karst.) Spirin & Niemelä,
Asterodon ferruginosus Pat., Atheloderma mirabile, Athelopsis lembospora (Bourdot) Oberw.,
Ceratobasidium anceps (Bres. & Syd.) H.S. Jacks., Dendrothele amygdalispora and Xanthoporus
syringae. Some areas of the Alps show a high fungal diversity (Bernicchia 2005 Bernicchia, A. 2005.
Polyporaceae s.l. Fungi Europaei , 10. Ed, 808Alassio, Italy: Candusso. ; Bernicchia & Gorjón 2010
Gorjón, S P and Bernicchia, A. 2010. The genus Dendrothele (Basidiomycota) in Italy, an update
with notes on European species. Nova Hedwigia, 90: 233–250.), for example “Val Casies”, “Val
d'Ultimo” and “Val di Genova” in the Trentino Alto-Adige Region. From the last of these, recently,
Antrodiella semistipitata has been described as a species new to the science (Bernicchia et al. 2007a
Bernicchia, A, Ryvarden, L and Gibertoni, B T. 2007a. Antrodiella semistipitata (Basidiomycetes,
Polyporales) a new species from Italy. Mycotaxon, 99: 231–237.). Some species have been
exclusively recorded from the Apennines. The “Sasso Fratino” Reserve, for example, is a very
interesting area for aphyllophoroid fungi, with two endemic polypores, Ceriporiopsis guidella
(Bernicchia & Ryvarden 2003 Bernicchia, A and Ryvarden, L. 2003. A new white-rot polypore from
Italy. Mycotaxon, 88: 219–224.), Fomitopsis labyrinthica, one corticioid, Botryobasidium
sassofratinoense Bernicchia & G. Langer (Bernicchia et al. 2010 Bernicchia, A, Langer, G and
Gorjón, S P. 2010. Botryobasidium sassofratinoense sp. nov. (Chantarellales, Basidiomycota) from
Italy. Mycotaxon, 111: 403–409.), and several rare species such as Ceriporia herinkii Vampola,
Crustomyces expallens (Bres.) Hjortstam, C. subabruptus (Bourdot & Galzin) Jülich, Dentipellis
fragilis, Hyphodontia latitans (Bourdot & Galzin) Ginns & M.N.L. Lefebvre, Parvobasidium
cretatum (Bourdot & Galzin) Jülich, Podofomes trogii (Fr.) Pouzar, Steccherinum robustius, and
Xylodon nudisetus (Warcup & P.H.B. Talbot) Hjortstam & Ryvarden.
Sardinia and Sicily, the two major Italian islands, provide a refuge for many interesting species.
Mediterranean forests and shrublands, typically consisting of densely or sparsely growing evergreen
shrubs and trees, constitute a complex ecological system inhabited by some recently recognized
species, many of them rare or very rare lignicolous fungi. In Sardinia, ancient juniper forests are a
most remarkable ecosystem because of their many rare aphyllophoroid fungi already discussed. Some
areas are notable for their fungal diversity. These include “Badde Salighes” (Bernicchia & Ryvarden
1988 Bernicchia, A and Ryvarden, L. 1988. Aleurodiscus ilexicola Bernicchia & Ryvarden sp. nov.
from Sardinia. Mycol Helv, 3: 83–88. ), “Valle di Lanaittu”, “Supramonte di Orgosolo” and “Urzulei”
in Nuoro province. In Ogliastra province, there is the “Montarbu forest”, the only known location of
Antrodia sandaliae, and “Baccu Gerduri forest” where Antrodiella ichnusana can be found. Several
species have been recorded on Juniperus spp.: E. ryvardenii, collected in Sardinia on Juniperus
phoenicea and J. oxycedrus subsp. macrocarpa, Sibth. & Sm., is very interesting because it is the
first and so far the only known collection area of Echinodontium Ellis & Everh. in Europe: all
previously known species are North American or Asiatic (Bernicchia & Piga 1998 Bernicchia, A and
Piga, A. 1998. A new species of Echinodontium from Italy. Mycotaxon, 68: 483–491.). Piloporia
sajanensis, recorded on J. oxycedrus subsp. oxycedrus L. is a boreal species, found in Scandinavia
and central to eastern Russia. Its presence in Sardinia can be regarded as a relic from the last glacial
period when ice reached most of central Europe and coniferous forest covered most of the higher
areas in the Mediterranean region.
In Sicily, assessment of wood-decay fungi is still in progress and new distributional and ecological
information is being acquired with the frequent discovery of previously unrecorded taxa. A huge
number of species have been found in Quercus ilex L. and Q. suber woods. These include Ceriporia
griseoviolascens, Gloeodontia columbiensis Burt ex Burds. & Lombard, Lindtneria chordulata (D.P.
Rogers) Hjortstam, Odonticium flavicans, Phellinus erectus A. David, Dequatre & Fiasson, Phlebia
nothofagi, Sarcodontia crocea (Schwein.) Kotl., Trechispora stevensonii (Berk. & Broome) K.H.
Larss., Vuilleminia coryli Boidin, Lanq. & Gilles, and V. pseudocystidiata. The presence in Sicily of
Oligoporus mappa (Overh. & J. Lowe) Gilb. & Ryvarden on Pinus halepensis Miller reforestation is
noteworthy. Areas with high fungal diversity in Sicily are the “Madonie Park”, “Ficuzza Wood-Rocca
Busambra Reserve”, “Monte Petroso” forest and “Nebrodi Park”.
Ecology, systematics
ascomycetes
and
distribution
of
wood-decay
Ascomycota is the largest phylum of kingdom Fungi and includes approximately 64,000 described
species in 327 families of 68 orders (Kirk et al. 2008 Kirk, P M, Cannon, P F, Minter, D W and
Stalpers, J A. 2008. Dictionary of the fungi. , 10th ed, 771UK: CABI Europe. ). It is characterized by
production of meiospores in specialized sac-shaped meiosporangia (asci). The current classification
of Ascomycota is based on a series of major phylogenetic studies carried out through the “Deep
Hypha” and assembling the fungal tree of life (AFTOL) projects (Blackwell et al. 2006 Blackwell,
M, Hibbett, D S, Taylor, J W and Spatafora, J W. 2006. Research coordination networks: A phylogeny
for kingdom Fungi (Deep Hypha). Mycologia, 98: 829–837.; James et al. 2006 James, T Y, Kauff, F,
Schoch, C, Matheny, P B, Hofstetter, VCox, C J. 2006. Reconstructing the early evolution of Fungi
using a six-gene phylogeny. Nature, 443: 818–822.; Hibbett et al. 2007 Hibbett, D S, Binder, M,
Bischoff, J F, Blackwell, M, Cannon, P FEriksson, O E. 2007. A higher-level phylogenetic
classification of the Fungi. Mycol Res, 111: 509–547.), as well as other resources including Myconet
(e.g. Eriksson 2006a Eriksson, O E. 2006a. Outline of Ascomycota – 2006. Myconet, 12: 1–82. ,b;
http://fieldmuseum.org/myconet/printed.asp). In many instances, molecular and morphological data
are congruent, but integration of these data has proved to be unfeasible in some cases, moreover a
significant number of orders and many families have yet to have any members in them sequenced.
Currently, Ascomycota is divided into three monophyletic subphyla: Pezizomycotina,
Saccharomycotina, and Taphrinomycotina. Pezizomycotina is the largest subphylum and includes the
vast majority of filamentous, fruit-body-producing species (James et al. 2006 James, T Y, Kauff, F,
Schoch, C, Matheny, P B, Hofstetter, VCox, C J. 2006. Reconstructing the early evolution of Fungi
using a six-gene phylogeny. Nature, 443: 818–822.). Within this subphylum, in the Sordariomycetes
clade there are fungi significant in a range of major ecologies including wood-decay (e.g. Xylariales)
(Spatafora et al. 2006 Spatafora, J W, Sung, G H, Johnson, D, Hesse, C, O'Rourke, BSerdani, M.
2006. A five-gene phylogeny of Pezizomycotina. Mycologia, 98: 1018–1028.).
The decay of living trees, snags, fallen branches and logs is critical for the recycling of nutrients, and
is effected primarily by fungi especially basidiomycetes and to a lesser extent ascomycetes and
bacteria. Decayed wood within these structures houses a large portion of forest biodiversity (Berg et
al. 1994 Berg, Å, Ehnström, B, Gustasson, L, Hallingbäck, T, Jonsell, M and Weslien, J. 1994.
Threatened plant, animal, and fungus species in Swedish forests: Distribution and habitat
associations. Conserv Biol, 8: 718–731.; Samuelsson et al. 1994 Samuelsson, J, Gustafsson, L and
Ingelog, T. 1994. Dying and dead trees: A review of their importance for biodiversity, Uppsala,
Sweden: Swedish Threatened Species Unit, Swedish National Environmental Protection Agency
Report 4306. ; Franklin et al. 1997 Franklin, J F, Berg, D R, Thornburgh, D A and Tappeiner, J C.
1997. “Alternative silvicultural approaches to timber harvesting: Variable retention harvest systems”.
In Creating forestry for the 21st century: The science of ecosystem management, Edited by: Kohm,
K A and Franklin, J F. 111–139. Washington, DC: Island Press. ; Butler et al. 2002 Butler, J,
Alexander, K NA and Green, T. 2002. Decaying wood: An overview of its status and ecology in the
United Kingdom and Continental Europe USDA Forest Service General Technical Report PSWGTR-181; Grove 2002 Grove, S J. 2002. Tree basal area and dead wood as surrogate indicators of
saproxylic insect faunal integrity: A case study from the Australian lowland tropics. Ecol Indic, 1:
171–188.).
Many species of wood-decay fungi have preference for trees in a particular age class. In addition, the
fungal communities in decomposing wood differ according to the particular decay classes (e.g.
Heilmann-Clausen & Boddy 2008; Fukasawa et al. 2011 Fukasawa, Y. 2011. Wood decomposing
abilities of diverse lignicolous fungi on nondecayed beech wood. Mycologia, 103: 474–482[Web of
Science ®]). The occurrence of different fungal communities with different abilities to cause decay
and with different ecological specialization defines the trophic fungal succession as a sequential
colonization of the substratum. Fungal succession on wood is indirectly driven by wood properties
(resource quality) and environmental factors (microclimate), and directly driven by fungal ecological
strategies (pioneer, combative) (Boddy 1992 Boddy, L. 1992. “Development and function of fungal
communities in decomposing wood”. In The fungal community: Its organisation and role in the
ecosystem. , 2nd ed, Edited by: Carroll, G C and Wicklow, D T. 749–782. New York, NY: MarcelDekker. , 2001). Many studies have reported that the Basidiomycota and Xylariaceae account for
most lignocellulose decomposition in wood debris. Many authors (e.g. Worrall et al. 1997 Worrall, J
J, Anagnost, S E and Zabel, R A. 1997. Comparison of wood decay among diverse lignicolous fungi.
Mycologia, 89: 199–219.; Fukasawa et al. 2011 Fukasawa, Y. 2011. Wood decomposing abilities of
diverse lignicolous fungi on nondecayed beech wood. Mycologia, 103: 474–482) have investigated
wood decaying abilities of fungi using pure culture tests under laboratory conditions with the aim of
understanding the role of fungi in wood decomposition in nature as well as the biological applications
of these abilities. Weight losses caused by members of Ascomycota were generally moderate to low,
but in the Xylariales and Sordariales approached 40%. All ascomycetes that caused weight loss
exceeding 2% caused decay features characteristic of soft rot (Worrall et al. 1997 Worrall, J J,
Anagnost, S E and Zabel, R A. 1997. Comparison of wood decay among diverse lignicolous fungi.
Mycologia, 89: 199–219.). For several Ascomycota, such as some Xylariales, these authors reported
the same complement of tested phenoloxidase activities (gallic and tannic acid, laccase, and
peroxidase) as did typical white-rot fungi. White-rot fungi can progressively utilize all major cell wall
components, including both the carbohydrates and the lignin.
Members of the Xylariaceae and Chaetomiaceae G. Winter and a few additional species showed other
advanced decay features, including relatively high weight loss and lignin degrading ability. Fukasawa
et al. (2011 Fukasawa, Y. 2011. Wood decomposing abilities of diverse lignicolous fungi on
nondecayed beech wood. Mycologia, 103: 474–482) reported among the Xylariaceae [Hypoxylon
fragiforme (Pers.) J. Kickx f. and Kretzschmaria deusta (Hoffm.) P.M.D. Martin], a greater weight
loss for non-decayed beech wood than for intermediately and well-decayed wood. The weight losses
were higher in intermediately or well-decayed wood in some anamorphic Ascomycota.
This first survey of wood-decay Ascomycota (excluding anamorphs) from Italy is based on species
reported in the available literature which was largely contributed by Medardi (2006 Medardi, G. 2006.
Ascomiceti d'Italia. Trento: A.M.B. Centro Studi Micologici, 454Trento: Associazione Micologica
Bresadola. ), Saitta et al. (2004 Saitta, A, Bernicchia, A and Venturella, G. 2004. Contributo alla
conoscenza dei funghi lignicoli della Sicilia. Inform Bot Ital, 36: 192–202. ) for Sicily, and
unpublished data resulting from surveys carried out on coarse woody debris in the Cilento and Vallo
di Diano National Park, (Campania) southern Italy (A. M. Persiani, personal communication).
Collectively, these constitute a total of 341 taxa within the Pezizomycotina, with information about
their distribution and substratum associations. It is far from exhaustive, but represents a good start for
assessing the status of knowledge about Italian wood-decay Ascomycota diversity, for determining
threats to that diversity, and promoting the most appropriate actions to be taken for its conservation
within with forest ecosystems. The nomenclatural sources adopted were the Dictionary of the Fungi
(Kirk et al. 2008 Kirk, P M, Cannon, P F, Minter, D W and Stalpers, J A. 2008. Dictionary of the
fungi. , 10th ed, 771UK: CABI Europe. ) and IndexFungorum, the de facto global nomenclator for
fungal names, http://www.indexfungorum.org/names/Names.asp.
The 341 species of wood-decay Ascomycota (Pezizomycotina) in Italy (i.e. only those species for
which a woody substratum has been reported, belong in 24 orders, 57 families and 138 genera. The
species are listed in the online Appendix E, in which are also reported their references published
and/or unpublished in the case of research in progress. In respect of the biogeographic regions of
Europe, and in the context of Alpine/Continental regions assigned to Italy, Lombardy is the most
widely studied with 287 taxa, followed by Trentino and Veneto, with 85 and 71 taxa, respectively. In
the Mediterranean region, Tuscany with 38 taxa, and Campania and Sicily, both with 36 taxa are the
regions most studied. However, this dataset may reflect the distribution of researchers and of
investigations carried out as well.
The best represented orders, as a percentage of the 341 species are: Helotiales Nannf. 30.5%,
Xylariales Nannf. 18.8%, Pezizales J. Schröt. 12.3%, Hypocreales Lindau 6.7%, Diaporthales Nannf.
6.7%, Pleosporales Luttr. ex M.E. Barr 5.0% and Orbiliales Baral, O.E. Erikss., G. Marson & E.
Weber 3.5%. The best represented families are: Xylariaceae 11.4%, Helotiaceae Rehm 10.9%,
Hyaloscyphaceae Nannf. 7.9%, Diatrypaceae Nitschke 6.7%, Pezizaceae Dumort. 5.3% and
Dermateaceae Fr. 5.3%. The best represented genera are: Peziza Fr. (14 taxa), Xylaria Hill ex
Schrank (14), Orbilia Fr. (11), Scutellinia (Cooke) Lambotte (10), Hymenoscyphus Gray (9),
Lachnellula P. Karst. (9), Hypocrea Fr. (8), Mollisia (Fr.) P. Karst. (8), Nectria (Fr.) Fr. (7), Lachnum
Retz. (6), Hypoxylon Bull. (6), and Phaeohelotium Kanouse (6).
The high richness in species of Mollisia agrees with reports for boreal forests. In fact Mollisia includes
species growing on plant debris as well as on decaying wood, and some are also obviously biotrophs.
Kauserud et al. (2005 Kauserud, H, Lie, M, Stensrud, O and Ohlson, M. 2005. Molecular
characterization of airborne fungal spores in boreal forests of contrasting human disturbance.
Mycologia, 97: 1215–1224.) sampled fungal spores from air in three boreal forest sites and
phylogenetic analyses were used to obtain a molecular characterization. Mollisia was the most
abundant genus within Helotiales and was especially frequent in old-growth forest sites and lowest
in the most disturbed clear-cut forests.
The results for the Pezizomycotina can also be correlated with various plant taxa (more than 50),
including conifers, and broad-leaved, deciduous and evergreen shrubs, and trees. In most cases, these
represent the most relevant vegetational aspects relative to the “forests, wood and scrubs” macro
category of the Directive's habitats (92/43/EEC) and its interpretation manual for Italy (Blasi et al.
2007 Blasi, C, Boitani, L, La Posta, S, Manes, F and Marchetti, M. 2007. “Biodiversity in Italy”. In
Contribution to the national biodiversity strategy, 460Roma: Palombi & Partner S.r.L. ).
The highest number of Pezizomycotina taxa was associated with Fagus sylvatica wood substrata (73),
and in decreasing order Quercus spp. (70), Alnus spp. (42) and Populus spp. (36). Records from
woody substrata of F. sylvatica and Quercus spp. will now be discussed.
In Europe, the southern limit of F. sylvatica is found in Italian beech forest stands and for this reason,
these stands are of particular interest, particularly since, in Mediterranean Europe, forest exploitation
has been going on for thousands of years, and during the 20th century a lot of deforestation took place
(Gilg 2004 Gilg, O. 2004. Foréts a caractère naturel: Caractéristiques, conservation et suivi, 96Aten,
Montpellier: Cahiers Techniques de l'ATEN, 74. ).
Of the 73 species reported as occurring on dead wood of F. sylvatica, around 70% are attributable to
investigations in beech forests of protected areas in Italy (online Appendix F). With respect to some
of these areas, two priority habitats indicated by the Habitat Directive (European Commission DG
Environment, Nature and Biodiversity, 2003 – Interpretation Manual of European Union
Habitats – EUR25) are relevant, both phytogeographically and ecologically. They are: “Apennine
beech forests with Taxus and Ilex” (*9210) for the PNCVD and “Apennine beech forests with Abies
alba and beech forests with Abies nebrodensis” (*9220) for Sicily (Madonie Park).
Of the Pezizomycotina recorded on F. sylvatica dead wood, four genera are the most represented,
each with four species: Hymenoscyphus calyculus (Sowerby) W. Phillips, H. imberbis (Bull.) Dennis,
H. serotinus (Pers.) W. Phillips, H. sublateritius (Berk. & Broome) Dennis, Hypocrea aureoviridis
Plowr. & Cooke, H. citrina (Pers.) Fr., H. gelatinosa (Tode) Fr., H. rufa (Pers.) Fr., Hypoxylon
fragiforme, H. fuscum (Pers.) Fr., H. rubiginosum (Pers.) Fr., H. rutilum Tul. & C. Tul., Xylaria
hypoxylon (L.) Grev., X. juruensis Henn., X. longipes Nitschke, and X. polymorpha (Pers.) Grev.
Diatrype Fr. is represented by three species: D. bullata (Hoffm.) Fr., D. disciformis (Hoffm.) Fr., and
D. stigma (Hoffm.) Fr.
This survey is the first Italian contribution about species of Pezizomycotina associated with beech
wood to highlight the key importance of dead wood for biodiversity in these forests and its availability
to possible specific species' demand.
The occurrence of wood-inhabiting fungi can be considered as an indicator of the natural value of
European beech forests, and some authors (e.g. Christensen et al. 2004 Christensen, M, HeilmannClausen, J, Walleyn, R and Adamcik, S. Wood-inhabiting fungi as indicators of nature value in
European beech forests. EFI Proceedings No. 51. Edited by: Marchetti, M. pp.526Finland Joensuu:
European Forest Institute. Monitoring and indicators of forest biodiversity in Europe e from ideas to
operationality) have proposed a set of fungal indicator species, predominantly Basidiomycota, to be
considered in setting conservation priorities in European beech forests.
In respect of the Pezizomycotina recorded on Quercus spp. dead wood (online Appendix G) four
genera are strongly represented. Hypoxylon has five species: Hypoxylon fragiforme, H. fuscum, H.
howeanum Peck, H. rubiginosum and H. rutilum. Mollisia has four species: M. cinerea (Batsch) P.
Karst., M. discolor var. longispora Le Gal, M. ligni (Desm.) P. Karst., M. ramealis P. Karst. The
genera Daldinia Ces. & De Not. and Lachnum are both represented by three species: D. concentrica
(Bolton) Ces. & De Not., D. martinii M. Stadler, Venturella & Wollw., D. raimundi M. Stadler,
Venturella & Wollw., L. bicolor (Bull.) P. Karst., L. brevipilosum Baral and L. virgineum (Batsch) P.
Karst. Among Quercus species, Q. ilex is notable as representative of the Mediterranean
sclerophyllous forests so widely characteristic of Italy. Prolonged exploitation of forests in Italy
favors this species which is more adapted to xeric environments than deciduous Quercus species like
Q. pubescens Willd. and Q. cerris L. The highest number of taxa were recorded from Q. ilex (26)
(online Appendix H), compared to 70 species for all species of Quercus combined.
Of the 341 species reported in this study, 20 (online Appendix I) appear in red lists of other European
countries (http://www.wsl.ch/eccf/). Among those 20 species we wish to highlight Biscogniauxia
mediterranea (De Not.) Kuntze, Hymenoscyphus monticola (Berk.) Baral and Chlorociboria
aeruginosa (Oeder) Seaver ex C.S. Ramamurthi, Korf & L.R. Batra, all on woody substrata of F.
sylvatica and Quercus spp.
Focal species for habitat conservation
In Italy, an integrated research project has been carried out in the Cilento and Vallo di Diano National
Park on broad-leaved forests with varying management histories (Burrascano et al. 2009 Burrascano,
S, Rosati, L and Blasi, C. 2009. Plant species diversity in Mediterranean old-growth forests: A case
study from central Italy. Plant Biosyst, 143: 190–200.). As part of this, the diversity of woodinhabiting fungi and beetles was also surveyed (Persiani et al. 2010 Persiani, A M, Audisio, P,
Lunghini, D, Maggi, O, Granito, V MBiscaccianti, A B. 2010. Linking taxonomical and functional
biodiversityof saproxylic fungi and beetles in broad-leaved forests in Southern Italy with varying
management histories. Plant Biosyst, 144: 250–261.).
This project carried out multi-taxon and forest structure sampling to identify indicators and monitor
(Blasi et al. 2010 Blasi, C, Marchetti, M, Chiavetta, U, Aleffi, M, Audisio, PAzzella, M M. 2010.
Multi-taxon and forest structure sampling for identification of indicators and monitoring of old
growth forest. Plant Biosyst, 144: 160–170.). Among other taxa, three Pezizomycotina, Ascocoryne
cylichnium (Tul.) Korf, Xylaria polymorpha and Hymenoscyphus serotinus, were shown to be oldgrowth forest indicators, albeit with an only marginally significant indicator value, using the indicator
species analysis (ISA) and the software PC-ORD 5 (McCune & Mefford 2006 McCune, B and
Mefford, M J. 2006. “PC-ORD. Multivariate analysis of ecological data”. In Version 5.10, Gleneden
Beach, OR, USA: MjM Software. ). Xylaria polymorpha was also considered to be an old-growth
indicator species by Schmid and Helfer (1999 Schmid, H and Helfer, W. 1999. Die Bedeutung der
Naturwaldreservate für den Pilzartenschutz. Seminarbericht der Natur-und Umweltschutzakademie
des Landes Nordrhein-Westfalen (NUA), 4: 140–146. ) and Parmasto (2001 Parmasto, E. 2001.
“Fungi as indicators of primeval and old-growth forests deserving protection”. In Fungal
conservation, issues and solutions, Edited by: Moore, D, Nauta, M N, Evans, S E and Rotheroe, M.
81–88. Cambridge, UK: Cambridge University Press.).
Referring to our dataset, all three are among the species associated with beech wood; their occurrence
thus plays a vital role to indicate the natural value of Italian/European beech forests.
Documenting and conserving Italian fungal biodiversity
Mycology has a long tradition in Europe and hence knowledge of species, their distribution, ecology
and status in Europe is the most extensive in the world. On this basis it would be feasible to analyze
the status of European fungal diversity to establish conservation priorities for fungi as part of national
and the European conservation priorities also applying IUCN red-listing priorities (Dahlberg &
Mueller 2011 Dahlberg, A and Mueller, G M. 2011. Applying IUCN red-listing criteria for assessing
and reporting on the conservation status of fungal species. Fungal Ecol, 4: 147–162.). On the other
hand, the large number of species, and in some cases, a lack of synergy between professional and
amateur mycologists, has meant that, to date, this knowledge (particularly within anamorphic and
teleomorphic ascomycetes) is low, compared with more well-known groups of species (Senn-Irlet et
al. 2007 Senn-Irlet, B, Heilmann-Clausen, J, Genney, D R and Dahlberg, A. 2007. Guidance for
conservation of macrofungi in Europe, 34Strasbourg: ECCF. ).
In Italy there are still limits to the knowledge of wood-decay Pezizomycotina impeding compilation
of the regional and national level check-lists which would allow characterization of declining, rare
and threatened populations. The set of data presented in this survey represents the first major
contribution to the knowledge of the distribution of these species and their relationship with their
substrata in Italy. We believe the present contribution will be useful at a European scale too.
According
to
the
Ascomycete
Conservation
Specialist
Group
(http://www.cybertruffle.org.uk/ascos/index.htm) ascomycetes have been seriously overlooked in
conservation while, at the same time, it is becoming clear that this largest single group of fungi may
be adversely affected by human activities. We also hope for an increase in Italian research to
determine the threats Pezizomycotina are facing and the development of long-term programs to assess
the status of individual species. This will contribute to the conservation of fungi in Europe and will
achieve the aims reported within the Convention on the conservation of European wildlife and natural
habitats (Senn-Irlet et al. 2007 Senn-Irlet, B, Heilmann-Clausen, J, Genney, D R and Dahlberg, A.
2007. Guidance for conservation of macrofungi in Europe, 34Strasbourg: ECCF. ).
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