The family Hymenochaetaceae (Agaricomycetes, Basidiomycota) in the islands of the Aegean Archipelago (Greece)

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1 This article was downloaded by: [Athens Agricultural University ] On: 19 June 2013, At: 03:28 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: Registered office: Mortimer House, Mortimer Street, London W1T 3JH, UK Plant Biosystems - An International Journal Dealing with all Aspects of Plant Biology: Official Journal of the Societa Botanica Italiana Publication details, including instructions for authors and subscription information: The family Hymenochaetaceae (Agaricomycetes, Basidiomycota) in the islands of the Aegean Archipelago (Greece) E. Polemis a, D. M. Dimou a b & G. I. Zervakis a a Laboratory of General and Agricultural Microbiology, Agricultural University of Athens, Greece b Korytsas 10, 15343, Agia Paraskevi, Greece Accepted author version posted online: 26 Feb 2013.Published online: 11 Mar To cite this article: E. Polemis, D. M. Dimou & G. I. Zervakis (2013): The family Hymenochaetaceae (Agaricomycetes, Basidiomycota) in the islands of the Aegean Archipelago (Greece), Plant Biosystems - An International Journal Dealing with all Aspects of Plant Biology: Official Journal of the Societa Botanica Italiana, DOI: / To link to this article: PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material.

2 Plant Biosystems, The family Hymenochaetaceae (Agaricomycetes, Basidiomycota) in the islands of the Aegean Archipelago (Greece) E. POLEMIS 1, D. M. DIMOU 1,2, & G. I. ZERVAKIS 1 1 Laboratory of General and Agricultural Microbiology, Agricultural University of Athens, Greece and 2 Korytsas 10, Agia Paraskevi, Greece Abstract In the frame of studying the diversity of wood-rotting macrofungi in Aegean islands, a notable Mediterranean biodiversity hotspot, the following hymenochaetoid species are reported for the first time in Greece: Fomitiporia erecta, Fomitiporia rosmarini, Hymenochaete fuliginosa, Inonotus cuticularis and Phylloporia ribis. The records of F. erecta and F. rosmarini are the first ever on Quercus coccifera in a worldwide context. Furthermore, they extend the known distribution of both species to the east Mediterranean region as well. The detected occurrence of P. ribis constitutes the first report of this genus for Greece; it was found on living trees and shrubs of Crataegus monogyna and Rhamnus alaternus in Andros and on Spartium junceum in Naxos. I. cuticularis was recorded on Acer sempervirens living tree trunk in Andros and on Quercus ithaburensis subsp. macrolepis in Agios Efstratios, while H. fuliginosa was detected in Ikaria on Quercus ilex. Detailed descriptions are provided for all five taxa, and the findings are critically discussed in conjunction with the relevant taxonomical literature. Furthermore, 10 additional hymenochaetoid taxa are reported together with information on their hosts and distribution in the Aegean islands. Keywords: Fungal diversity, Mediterranean vegetation, polypores, species distribution, wood-decay fungi The Mediterranean Basin is among the world s biodiversity hotspots (Myers et al. 2000), including several types of habitats that are highly diverse but endangered due to the extended anthropogenic pressure. In particular, the plant diversity in the Aegean Archipelago is one of the richest in the Mediterranean area with a high rate of species endemism that is well documented and thoroughly explored (Tzanoudakis & Panitsa 1995; Strid 1996). Each Aegean island is distinct in many aspects due to its size, landscape, altitude, geology, period of geographic isolation, proximity to the adjacent mainland and local environmental conditions. The vegetation often consists of relict Quercus forests and stands of Quercus pubescens, Quercus ithaburensis subsp. macrolepis and Quercus ilex, of Pinus brutia forests, of Platanus orientalis, Alnus glutinosa, Salix alba and Fraxinus ornus communities by wet ravines and of Tamarix spp. stands by the littoral zone. However, sclerophyllous maquis (e.g. Quercus coccifera, Acer sempervirens, Pistacia terebinthus, Pistacia lentiscus, Arbutus unedo and Erica arborea) and phrygana (e.g. Sarcopoterium spinosum, Calycotome vilosa, Genista acanthoclada, Corydothymus capitatus, Satureja thymbra and Cistus spp.) are dominating throughout this region. Vegetation of the area is classified within the order Quercetalia ilicis which is divided into two subzones: Oleo-Ceratonion alliance below the elevation of m and Quercion ilicis alliance above this elevation level. The family Hymenochaetaceae Donk includes wood-decaying fungi of great ecological importance, comprising saprotrophs and parasites. Due to their enzymatic properties, most species are efficient lignin degraders (Pelaéz et al. 1995) and several of them are known to infest broad-leaved and coniferous trees by causing wood heart rot, canker and root diseases. In addition, some other taxa are known to produce basidiomata with medicinal, antioxidant and antitumour properties (Cui et al. 2005; Dai et al. 2010). Members of the family form resupinate, pileate or stipitate basidiomata with distinct types of hymenial Correspondence: G. I. Zervakis, Laboratory of General and Agricultural Microbiology, Agricultural University of Athens, Iera Odos 75, Athens, Greece. Tel: þ Fax: þ zervakis@aua.gr q 2013 Societá Botanica Italiana

3 2 E. Polemis et al. structure (corticioid, hydnoid or poroid); their main common characters are the darkening of tissues with KOH (xanthochroic reaction), the absence of clamps and the presence of a continuous parenthosome in the dolipore apparatus, while setae or setiform hyphae are found in many taxa (Parmasto & Parmasto 1979; Moore 1980; Larsen & Cobb- Poule 1990; Corner 1991; Ryvarden 2005). Molecular data showed that the family Hymenochaetaceae forms a distinct phylogenetic entity, i. e. hymenochaetoid clade (Hibbett & Thorn 2001) and provided valuable evidence for circumscribing it (Larsson et al. 2006). According to Kirk et al. (2008), the family includes 27 genera and 487 species. Noteworthy is that the major genera of Hymenochaetaceae, such as Inonotus P. Karst. and Phellinus Quél., have undergone extended segregation during the last decades under the scope of a more natural classification system. Fiasson and Niemelä (1984) used several chemical, morphological and anatomical characters to divide taxa within the family; subsequent mating compatibility and phylogenetic studies supported those subdivisions (Fischer 1996; Wagner & Fischer 2001; Fischer & Binder 2004; Jeong et al. 2005). In view of such evidence, Fomitiporia was reappraised at the genus level and the new taxonomic scheme is currently largely accepted (Fischer & Binder 2004; Decock et al. 2005, 2007; Fischer et al. 2005). In Greece, knowledge on the diversity of the family Hymenochaetaceae is still poor, and only six genera are known to exist: Coltricia Gray, Hymenochaete Lév., Inonotus s.l. (incl. the genus Inocutis Fiasson & Niemelä), Onnia P. Karst., Phaeolus (Pat.) Pat. and Phellinus s.l. including the segregated genera Fomitiporia Murrill, Fuscoporia Murrill and Porodaedalea Murrill (Zervakis et al. 1998, 2002; Elena et al. 2006). Moreover, very few records of hymenochaetoid fungi were previously reported from the islands of the Aegean Archipelago (Petrak 1943; Klán 1978; Plank 1980). Recent long-term studies focused on the inventory and taxonomy of macrofungi in selected Aegean islands (Polemis & Noordeloos 2007; Noordeloos & Polemis 2008; Polemis et al. 2012a, 2012b). Along this line of research, rare and interesting specimens of hymenochaetoid fungi were collected from the islands of Andros, Naxos, Amorgos, Agios Efstratios, Ikaria and Nisyros. The aims were to conduct a thorough study of their morphological characters, to compare them with pertinent literature descriptions for contributing towards the elucidation of the complex taxonomic relationships within this fungal group and to provide information on their geographic distribution and associated plants in selected Aegean islands. Materials and methods Data presented in this study are based on specimens collected through the fieldwork which was mainly performed during the past 15 years in various islands of the Aegean Archipelago. Particular emphasis was placed on the long-term research of macrofungi in the islands of Andros, Naxos and Amorgos (Cyclades complex) and of the north Aegean islet of Agios Efstratios; in addition, some preliminary sampling was performed in the island of Ikaria (east Aegean) and the volcanic island of Nisyros (southeast Aegean, Figure 1). Specimens were collected by E. Polemis (EP) and D.M. Dimou (DD), and they were then deposited in the herbarium of the Laboratory of General and Agricultural Microbiology of the Agricultural University of Athens. Dried samples were examined following the methods described in the relevant literature (Ryvarden & Gilbertson 1993; Léger 1998; Bernicchia 2005; Ryvarden 2005), using light and phase-contrast microscopy. All sections were mounted and observed in 3 5% KOH and selectively in cotton blue and Melzer s reagent. In all cases, a minimum of 30 spores were measured. Line drawings were made after editing and printing of microphotographs. The transliteration of Greek geographical names follows the International Standard ISO 843 (1997). Results In the frame of this study, five species of the family Hymenochaetaceae are recorded for the first time in Greece; in addition, one of them (Phylloporia ribis) constitutes a new genus record. The respective detailed annotated descriptions based on specimens collected from Aegean islands are provided below. Furthermore, the occurrence of other 10 species is reported from the Aegean islands, 6 of them were found for the first time in this particular area. In addition, several specimens were collected from plant hosts with which were never before associated, while the occurrence of certain species is now evidenced in the eastern Mediterranean. Fomitiporia erecta (A. David, Dequatre & Fiasson) Fiasson ; Phellinus erectus A. David, Dequatre & Fiasson Description. Basidiomata annual or perennial, effused to shallowly ungulate, erect to almost clavate (in this case having a pileus of 6 cm in diameter and 3 cm in height), or pileate up to cm, encircling cut plant stems at the ground level in a fanshaped manner; upper surface dark sepia brown to almost black and rimose; sterile margin rounded bright yellow to yellow-brown, becoming darker

4 Hymenochaetoid fungi in Aegean islands 3 Figure 1. Greece and its phytogeographical zones according to the Flora Hellenica project (Strid & Tan 1997). Areas under investigation appear as Kik (Cyclades), NAe (North Aegean) and EAe (East Aegean). Islands that were studied appear highlighted in black. rusty-brown when bruised; pore surface rusty to fulvous, with a greyish to yellowish-brown pruina, darkening when bruised, pores circular, 4 6 mm 21 ; context hard and corky to woody, radially fibrous, and evidently concentrically zonate, yellowishbrown, fulvous; tube layers up to 4 mm thick, stratified, concolorous with pore surface. Hyphal system dimitic, generative hyphae hyaline, thin walled, simple-septate, 2 4 mm in diameter; skeletal hyphae thick-walled, brown, with adventitious septa and rarely branched, mm in diameter (Figure 2). Setal hyphae observed only in the specimen from Amorgos, in trama and subhymenium, 4.5 8( 10) mm broad, brown, thick-walled, subulate or rarely enlarged at apex. Hymenial setae ( 9) mm, present in lower parts of the tubes but rare or absent close to the apices, subulate, slender and often flexuous, or curved, brownish; sterile cystidioid elements rare and inconspicuous, mostly lageniform with short neck. Tramal setae (again observed only in the Amorgos specimen) more elongated, then the hymenial setae reaching 67 mm in length. Basidia 11 15( 18) 5 7( 9) mm, clavate or subclavate, four-spored, simple-septate. Basidiospores (6 ) (5 ) ( 7) mm, broadly ellipsoid, ovoid to subglobose, hyaline, smooth, thickwalled, distinctly dextrinoid, cyanophilous. Figure 2. F. erecta: (a) generative and skeletal tramal hyphae, (b) basidiospores, (c) basidia and (d) hymenial and tramal setae.

5 4 E. Polemis et al. Specimens examined. Amorgos, on dead stems of Q. coccifera L. cut on the ground level, 3 December 2005, EP.05-M186. Naxos, on dead stems of Q. coccifera cut on the ground level, 29 November 2005, EP.05-N356. Fomitiporia rosmarini (Bernicchia) Ghob.-Nejh. & Y. C. Dai ; Phellinus rosmarini Bernicchia Description. Basidiomata annual, pileate, emerging as tuberculate outgrowths, ungulate or cushionshaped, up to 5 cm long and 2.5 cm thick, upper surface velutinous, yellowish towards the margin, dark brown to almost black and rimose close to the substrate; pore surface yellowish-brown, pores circular, 5 7 per mm; context hard and woody, yellowish-brown; tube layers up to 4 mm thick, stratified, concolorous with pore surface. Hyphal system dimitic, generative hyphae hyaline, thin-walled, simple-septate, 2 3 mm in diameter; skeletal hyphae thick-walled, brown, without septa and rarely branched, 2 6 mm in diameter (Figure 3). Hymenial setae mm, rare but found in all specimens, subulate, brownish. Cystidioles rarely conspicuous, ventricose, fusiform or with a filiform projection up to 20 mm long. Basidia mm, broadly clavate, four-spored, simpleseptate. Basidiospores (4.5 )5 6.5( 7) mm, globose to subglobose, hyaline, smooth, thick-walled, distinctly dextrinoid, cyanophilous. Specimens examined. Naxos, on the base of a dead Q. coccifera stump, 29 November 2005, EP.05-N205. Amorgos, on dead wood of Pistacia lentiscus L., 03 December 2005, EP.05-M186. Hymenochaete fuliginosa (Pers.) Lév (Hymenochaete rubiginosa subsp. subfuliginosa (Bourdot & Galzin) Bourdot & Galzin, and H. subfuliginosa Bourdot & Galzin) Description. Basidiomata resupinate, closely adnate, up to 0.5 mm thick, effused on substrate for more than 15 cm in length, fawn to dark purplish brown with a greyish hue; hymenophore smooth to distinctly tuberculate, irregularly cracked; margin thin, clearly delimited, concolorous with hymenium or with a darker zone; tomentum absent, cortex indistinct; context absent and the basidioma consists of several setal layers with overlapping rows of setae. Hyphal system monomitic, setal hyphae absent; generative hyphae 2 4 mm in diameter, yellowish to brownish, thin- to thick-walled (Figure 4). Setae abundant (45 )60 100( 110) (5 )6 10( 11) mm, subulate, with acute apex, straight, not incrusted. Hyphidia present, hyaline or yellowish, without incrustation; basidia clavate, ( 5) mm; spores cylindrical, slightly curved, (4.5 )5 6.5( 7) 2 3mm. Specimen examined. Ikaria, on stems and trunks of dead Quercus ilex L., 10 February 2011, EP.11- K102. Inonotus cuticularis (Bull.) P. Karst Description. Basidiomata annual, but remaining attached to the substrate for one or more years, imbricate, in large clusters; pilei dimidiate to applanate of up to 10 cm in diameter and up to 2 cm thick; upper surface velutinous to hirsute, Figure 3. F. rosmarini: (a) generative and skeletal tramal hyphae, (b) basidiospores, (c) basidia and (d) hymenial setae. Figure 4. H. fuliginosa: (a) hymenial setae, (b) basidiospores, (c) basidia and (d) hyphidia.

6 azonate, becoming glabrous, reddish-brown to yellow-brown towards the margin; pore surface greyish-brown, pores angular, 4 5 per mm; context relatively soft when fresh, then firm and corky, azonate, yellowish-brown. Hyphal system monomitic, contextual hyphae of two types: thin-walled pale yellowish, and thickwalled brown; subhymenial hyphae thin- to thickwalled, hyaline to yellowish-brown, all types of hyphae simple-septate (Figure 5A). Setal hyphae from pileal surface mm, firm to thick-walled, branched in various patterns, monopodial to dichotomous, branches often curved or hooked, with pointed apices; hyaline thin-walled hyphae, with clavate terminal elements of up to 14 mm broad also present on pileal surface. Tramal setae abundant, mm, subulate, thickwalled (Figure 5B). Hymenial setae mm, abundant, subulate, conical, ventricose, with straight or curved apices, thick-walled to almost solid, brownish. Basidia mm, short clavate, four-spored, simple-septate. Basidiospores (6 )6.5 7( 8) (4 ) mm, broadly ellipsoid to ovoid, yellowish-brown, smooth, thick-walled, negative in Melzer s reagent. Figure 5a. I. cuticularis: Setal hyphae and thin-walled clavate elements from the pileal surface. Hymenochaetoid fungi in Aegean islands 5 Figure 5b. I. cuticularis: (a) basidiospores, (b) thin- and thickwalled contextual hyphae, (c) tramal setae, (d) hymenial setae and (e) cross-section of the hymenium with basidia cystidioles and hymenial setae. Specimens examined. Agios Efstratios, on the trunk of living Q. ithaburensis Decne. subsp. macrolepis (Kotschy) Hedge & Yalt. tree, 20 August 1999, DD Andros, on the trunk of living A. sempervirens L. tree, 01 November 2002, EP.02-A507. Nisyros, on the trunk of living Q. ithaburensis subsp. macrolepis tree, 24 December 2010, EP.10-S011. P. ribis (Schumach.) Ryvarden ; Phellinus ribis (Schumach.) Quél. Description. Basidiomata perennial, imbricate, pilei dimidiate to semicircular, encircling stem and branches of shrubs near ground surface, up to 15 cm in diameter and up to 8 cm thick at base, gradually thinning towards the almost acute margin; upper surface velutinous to tomentose, concentrically sulcate, brown at centre, velutinous or smooth, yellowish-brown at margin; pore surface yellowishbrown, dark brown when bruised, tube layers up to 2 cm thick, pores circular, 5 7 per mm; context firm and corky, yellowish-brown. Hyphal system monomitic, contextual hyphae thick-walled, brown, up to 6 mm wide; subhymenial hyphae thin to thick-walled, hyaline, 2 4 mm wide, simple-septate. Basidia mm, clavate, four-spored, simple-septate. Basidiospores mm, broadly ellipsoid, yellowish-brown in KOH, smooth, slightly thick-walled, negative in

7 6 E. Polemis et al. Melzer s reagent. Cystidia, setae and other sterile elements lacking. Specimens examined. Andros, on the base of the stem of living Crataegus monogyna Jacq. shrub, 08 October 2002, EP.02-A416; on the base of the stem of dead unidentified shrub, 01 December 2002, EP.02-A635; on the base of the stem of living Rhamnus alaternus L. shrub, 24 April 2011, EP.11- A1045. Naxos, on the base of the stem of living Spartium junceum L. shrub, 13 December 2004, EP.04-N224. Apart from the preceding thoroughly described species which were not previously reported in Greece, the following taxa belonging to the family Hymenochaetaceae were also recorded in various Aegean islands during the last 15 years. In many cases, the substrate/host or substrate association constitutes a new record for Greece and this is indicated with a cross ( ). Fomitiporia mediterranea M. Fisch Specimen examined. Andros, on Citrus sp., 12 April 2005, DD Remarks. This specimen was originally examined by D.M. Dimou, and its identity was subsequently confirmed through the use of molecular methods (Elena et al. 2006). Fomitiporia punctata (P. Karst.) Murrill (Phellinus punctatus (Fr.) Pilát) Specimens examined. Naxos, on Q. coccifera, 15 January 1999, EP.99-N084. Andros, on Olea europaea L., 19 April 2001, EP.01-A210, on Laurus nobilis L. ( ), 29 August 2012, EP.12-A1121. Amorgos, on Calycotome villosa (Poir.) Link ( ), 20 December 2004, EP.04-M048. Nisyros, on Q. coccifera, 26 December 2010, EP.10-S058. Fomitiporia robusta (P. Karst.) Fiasson & Niemelä ; Phellinus robustus (P. Karst.) Bourdot & Galzin Specimens examined. Naxos, on Q. coccifera, 15 December 2004, EP.04-N261. Fuscoporia torulosa (Pers.) T. Wagner & M. Fisch ; Phellinus torulosus (Pers.) Bourdot & Galzin Specimens examined. Naxos, on Quercus pubescens Willd. ( ), 13 January 1999, EP.99-N057; on Anthyllis hermaniae L. ( ), 28 January 2002, EP.02- N165; on Ceratonia siliqua L., 11 December 2004, EP.04-N191; on Q. coccifera, 29 November 2005, EP.05-N357; on A. glutinosa L., 30 November 2005, EP.05-N398. Andros, on a standing Prunus sp. tree, 20 April 2001, EP.01-A216; on Q. coccifera ( ) stump, 7 October 2002, EP.02-A397; on C. monogyna ( ) stump, 29 October 2002, EP.02- A485; on A. glutinosa ( ), 15 October 2004, EP-04- A668. Amorgos, on C. siliqua, 8 January 2005, EP.05-M124 and on O., 2 December 2005, EP.05- M170. Ikaria, on Q. coccifera, 3 December 2010 EP.10-K020. Nisyros, on Q. coccifera, 26 December 2010, EP.10-S036. Hymenochaete rubiginosa (Dicks.) Lév Specimen examined. Ikaria, on Q. ilex, 10 February 2011, EP.11-K106. Inocutis tamaricis (Pat.) Fiasson & Niemelä ; Inonotus tamaricis (Pat.) Maire Specimens examined. Naxos, on living tree of Tamarix sp., 27 January 2002, EP.02-N148. Amorgos, on living tree of Tamarix sp., 6 January 2005, EP.05-M113. Agios Efstratios, on living tree of Tamarix sp., 28 February 2010, DD Andros, on living tree of Tamarix sp., 19 November 2012, EP.12-A1143. Phellinus igniarius (L.) Quél Specimen examined. Amorgos, on Prunus dulcis (Mill.) D.A.Webb ( ), 19 December 2004, EP.04-M041. Phellinus laevigatus (Fr.) Bourdot & Galzin Specimen examined. Andros, on F. ornus L. ( ) living tree, 30 October 2002, EP.02-A490. Phellinus rimosus (Berk.) Pilát Specimens examined. Andros, on P. dulcis standing tree, 13 November 2002, EP.02-A521. Amorgos, on Q. coccifera, 18 December 2004, EP.04-M023; on Q. coccifera, 9 January 2005, EP.05-M142. Naxos, on P. lentiscus ( ), 1 December 2005, EP.05-N402. Porodaedalea pini (Brot.) Murrill ; Phellinus pini (Brot.) Bondartsev & Singer Specimen examined. Andros, on living Pinus halepensis Miller, 24 September 1998, EP.98-A180. Discussion F. erecta is a relatively rare Mediterranean species known to date from Portugal, Spain, France, Italy and the former Yugoslavia (Ryvarden & Gilbertson 1994; Telleria et al. 1997; Bernicchia 2005; Saitta et al. 2011). It was previously recorded on Q. ilex, P. dulcis, as well as on various species of the genera

8 Arbutus, Rosmarinus, Ulex and Viburnum (Larsen & Cobb-Poulle 1990; Ryvarden & Gilbertson 1994; Bernicchia 2005). Our findings represent the only known records from Q. coccifera. F. erecta belongs to the F. robusta complex, differing from all other pileate species of the complex by the presence of hymenial setae. Phellinus juniperinus Bernicchia & Curreli is a closely related taxon known only from Sardinia and a few localities in Mediterranean France; it is characterized by the constant presence of setal hyphae and by its host preference, i.e. it grows exclusively on Juniperus spp. (Bernicchia 2005). In contrast, tramal setae and setal hyphae in F. erecta are reported to be either completely absent (David et al. 1982; Ryvarden & Gilbertson 1994; Bernicchia 2005) or present albeit inconsistent (Melo 1988; Pieri & Rivoire 2000). Since our Amorgos F. erecta specimens possessed both setal hyphae and tramal setae, it seems that the presence of such morphological characters is not of diagnostic value in discriminating between the two species. Furthermore, the spores of F. erecta are referred to as subglobose or globose (David et al. 1982; Ryvarden & Gilbertson 1994; Bernicchia 2005), whereas in our specimens several ovoid to broadly ellipsoid spores were also observed. In addition, basidia were longer and typically clavate or subclavate as opposed to the subglobose or globose basidia reported in the previous pertinent studies. F. rosmarini is another rare Mediterranean species known to date from Portugal, France, Italy and the former Yugoslavia (Ryvarden & Gilbertson 1994, Saitta et al. 2011); however, recently it was also reported from Iran on Quercus sp. (Ghobad-Nejhad & Dai 2007). It belongs to the F. punctata complex, differing from F. punctata by the presence of hymenial setae. Three other closely related species, F. pseudopunctata, F. erecta and Ph. juniperinus, possess larger spores and broader setae (Bernicchia 2005). F. rosmarini was previously recorded on various Mediterranean shrubs and sclerophyllous trees including P. lentiscus (Larsen & Cobb-Poule 1990; Ryvarden & Gilbertson 1994; Bernicchia 2005), but our finding from Naxos constitutes the first ever record on Q. coccifera. H. fuliginosa is a widely distributed species in temperate and Mediterranean Europe, in north America and Cuba and in New Zealand (Léger 1998; Bernicchia & Gorjón 2010). In older literature reports, H. fuliginosa was considered to grow only on conifers, and it was hence distinguished from H. subfuliginosa, colonizing broad-leaved trees only, and especially Quercus spp. (Jahn 1971). However, these two taxa are now considered as conspecific (Hansen & Knudsen 1997; Léger 1998). Léger (1998) states that the taxon known under the name Hymenochaetoid fungi in Aegean islands 7 H. subfuliginosa is merely an eco-climatic form of H. fuliginosa, which is characterized by thicker basidiomata and the occasional presence of indistinct cortex; furthermore, the color of hymenium and the size of spores are not considered as significant diagnostic characters. This interpretation fits well to the features of our specimens collected in a montane thermo-mediterranean Q. ilex forest. I. cuticularis is a widespread species found throughout Europe, and it is also reported to occur in Asia, North America and North Africa (Ryvarden & Gilbertson 1993). It is considered as infrequent in Italy (Bernicchia 2005), while it is known by a few collections from Turkey as well (Solak et al. 2007). Similarly, it is rather rare in Greece since it has not been found to date in the mainland, still being known by our three collections from the Aegean islands of Agios Efstratios, Andros and Nisyros. Although it was reported to colonize dead wood of various Quercus and Acer spp., it is the first time that this species was recorded on living Q. ithaburensis subsp. macrolepis and A. sempervirens. I. cuticularis is an easily recognized species due to its unique branched setal hyphae on the pileal surface, and it is the only European species possessing such elements. The macroscopical and microscopical features of our specimens are in accordance with the pertinent literature. However, the thin-walled elements from pileus surface as well as the tramal setae were not included in previous descriptions (Ryvarden & Gilbertson 1993; Bernicchia 2005; Ghobad-Nejhad & Kotiranta 2008). P. ribis is the only known species of the genus that is widespread in the whole Northern Hemisphere (Wagner & Ryvarden 2002). Although it is a rather common species in Mediterranean countries as well (Doğan et al. 2005; Denchev & Assyov 2010), this is the first record of the occurrence of the respective genus in Greece. It grows mainly on Ribes spp. (north Europe), but it has also been recorded on a large number of living shrubs belonging to various genera of angiosperms including Crataegus and Spartium. Our record of P. ribis on Rhamnus is the first one on this plant. The annual basidiomata arising always from the ground level by encircling stems of shrubs, the monomitic system, the small ellipsoid, yellowishbrown basidiospores and the absence of setae are the main diagnostic features of P. ribis. Basidiomata in one of our collections possessed a pileus diameter of 15 cm, thickness at base 8 cm and tube layers up to 2 cm thick. These dimensions are considerably larger, if compared to those provided for this species by Wagner and Ryvarden (2002), i.e. pileus diameter of 10 cm, thickness at base 4 cm and tube layers up to 0.5 cm thick.

9 8 E. Polemis et al. Among the other 10 species of Hymenochaetaceae previously recorded in Greece, the occurrence of 4 was reported in the past from Aegean islands: F. punctata in Rodos (Plank 1980), F. torulosa in Crete on C. siliqua, O. europaea and P. dulcis (Plank 1980), P. rimosus in Chios on P. lentiscus var. chia Duhamel (Sarejannis & Dimitriadis 1951), in Hydra on P. vera L. (Sarejannis et al. 1954) and in Rodos on P. terebinthus L. (Kotlaba & Pouzar 1978), and I. tamaricis on Tamarix spp. in several islands (Klán 1978; Plank 1980; Kotlaba & Klan 1994). In conclusion, new information is provided on the occurrence of wood-rotting macrofungi of the family Hymenochaetaceae in the islands of the Aegean Archipelago. This particular area, which is very significant as a biodiversity hotspot, was neglected in the past with respect to its mycobiota. 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