This article was downloaded by: [University of Perugia]On: 14 January 2014, At: 00:13Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954 Registered office: MortimerHouse, 37-41 Mortimer Street, London W1T 3JH, UK

Plant Biosystems - An International Journal Dealingwith all Aspects of Plant Biology: Official Journal ofthe Societa Botanica ItalianaPublication details, including instructions for authors and subscription information:http://www.tandfonline.com/loi/tplb20

Macrofungi in Mediterranean maquis along seashoreand altitudinal transectsLorenzo Pecoraroa, Paola Angelinib, Andrea Arcangelib, Giancarlo Bistocchib, MariaLetizia Garganoc, Alfonso La Rosac, Dario Lunghinid, Elias Polemise, Andrea Rubinif,Alessandro Saittac, Roberto Venanzonib & Georgios I. Zervakise

a Institute of Agricultural and Environmental Sciences, Estonian University of LifeSciences, Estoniab Department Applied Biology, University of Perugia, Italyc Department of Agricultural and Forest Science, University of Palermo, Italyd Dipartimento di Biologia Ambientale, Università di Roma “La Sapienza”, Italye Laboratory of General and Agricultural Microbiology, Agricultural University of Athens,Greecef CNR - Institute of Plant Genetics, ItalyAccepted author version posted online: 08 Jan 2014.Published online: 08 Jan 2014.

To cite this article: Plant Biosystems - An International Journal Dealing with all Aspects of Plant Biology(2014): Macrofungi in Mediterranean maquis along seashore and altitudinal transects, Plant Biosystems - AnInternational Journal Dealing with all Aspects of Plant Biology: Official Journal of the Societa Botanica Italiana, DOI:10.1080/11263504.2013.877535

To link to this article: http://dx.doi.org/10.1080/11263504.2013.877535

Disclaimer: This is a version of an unedited manuscript that has been accepted for publication. As a serviceto authors and researchers we are providing this version of the accepted manuscript (AM). Copyediting,typesetting, and review of the resulting proof will be undertaken on this manuscript before final publicationof the Version of Record (VoR). During production and pre-press, errors may be discovered which couldaffect the content, and all legal disclaimers that apply to the journal relate to this version also.

PLEASE SCROLL DOWN FOR ARTICLE

Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) containedin the publications on our platform. However, Taylor & Francis, our agents, and our licensors make norepresentations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose ofthe Content. Any opinions and views expressed in this publication are the opinions and views of the authors,and are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be reliedupon and should be independently verified with primary sources of information. Taylor and Francis shallnot be liable for any losses, actions, claims, proceedings, demands, costs, expenses, damages, and otherliabilities whatsoever or howsoever caused arising directly or indirectly in connection with, in relation to orarising out of the use of the Content.

This article may be used for research, teaching, and private study purposes. Any substantial or systematicreproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any

form to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http://www.tandfonline.com/page/terms-and-conditions

Dow

nloa

ded

by [

Uni

vers

ity o

f Pe

rugi

a] a

t 00:

13 1

4 Ja

nuar

y 20

14

Accep

ted M

anus

cript

Cover page Running title: Macrofungi in Mediterranean maquis Corresponding author: Lorenzo Pecoraro PhD, Estonian University of Life Sciences, Institute of Agricultural and Environmental Sciences, Kreutzwaldi 5, 51014 Tartu, Estonia; phone +372 7311873; fax +372 7313982; e-mail lorenzo.pecoraro@gmail.com

Dow

nloa

ded

by [

Uni

vers

ity o

f Pe

rugi

a] a

t 00:

13 1

4 Ja

nuar

y 20

14

Accep

ted M

anus

cript

Macrofungi in Mediterranean maquis along seashore and altitudinal transects Lorenzo Pecoraro1, Paola Angelini2, Andrea Arcangeli2, Giancarlo Bistocchi2, Maria Letizia Gargano3, Alfonso La Rosa3, Dario Lunghini4, Elias Polemis5, Andrea Rubini6, Alessandro Saitta3, Roberto Venanzoni2, Georgios I. Zervakis5

1Estonian University of Life Sciences, Institute of Agricultural and Environmental Sciences, Kreutzwaldi 5, 51014 Tartu, Estonia

2University of Perugia, Department Applied Biology, Borgo XX Giugno, 74 – 06121 Perugia, Italy 3University of Palermo, Department of Agricultural and Forest Science, Viale delle Scienze 11, 90128, Palermo, Italy 4Università di Roma “La Sapienza”, Dipartimento di Biologia Ambientale, P.le Aldo Moro 5, 00185 Roma, Italy 5Agricultural University of Athens, Laboratory of General and Agricultural Microbiology, Iera Odos 75, 11855 Athens, Greece 6CNR - Institute of Plant Genetics, Perugia Division, Via Madonna Alta 130, 06128 Perugia, Italy. Abstract In semiarid Mediterranean environments, fungal activity is fundamental for buffering biotic and abiotic stress to the plant and for sustaining a vegetation cover. Despite the important role that fungi play in habitats stability, mycological data from Mediterranean ecosystems are scarce and fragmentary. We investigated fungal diversity in several areas characterized by Mediterranean maquis, from continental Italy, Sicily and Greece in order to contribute to the analysis of distribution, ecology and diversity of macrofungi in evergreen sclerophyllous shrublands at different elevation and distance from the seashore across the Mediterranean Basin. Several fungal taxa that are remarkable due to their ecology, rarity and limited geographical distribution were recovered. Among them, the ectomycorrhizal species Amanita dunensis and A. valens were recorded for the first time in Italy. The wood-inhabiting macrofungi, Aleurodiscus dextrinoideocerussatus, Peniophorella tsugae, Perenniporia meridionalis, Phanerochaete martelliana, Vararia ochroleuca and six Peniophora species were reported for the first time in Greece. Some species, such as Gloeodontia columbiensis, Ceriporia aurantiocarnescens, Peniophora pithya and P. tamaricicola were collected on new substrates. For the two rare basidiomycetes Amanita eliae and Battarrea phalloides, molecular identification and phylogenetic analysis gave valuable information on their relationship with similar taxa. Keywords: Ectomycorrhizal species, macromycetes, Mediterranean ecosystems, molecular analysis, saprotrophs, wood-inhabiting fungi

Dow

nloa

ded

by [

Uni

vers

ity o

f Pe

rugi

a] a

t 00:

13 1

4 Ja

nuar

y 20

14

Accep

ted M

anus

cript

Introduction

Mediterranean zones, occurring at latitudes between 30° to 40°, are characterized by dry summers, wet winters and a spring growing season and represent a major ecosystem within the semiarid regions (Mazzoleni et al. 1992; Pereira et al. 2002). The Mediterranean Basin, where 13,000 endemic plants grow, is considered a “hot spot” of biodiversity (Brooks et al. 2002). About 42,000 km2 of this area (i.e. 38.3% of the hot spot) are protected (Turrini et al. 2010) since they are characterized by particularly high number of living organisms and harbor many ectomycorrhizal trees species growing together with endomycorrhizal herbaceous plants or shrubs. These ecosystems, however, are still largerly unexplored, especially as far as fungal diversity and ecology are concerned (Bergero et al. 2000). Although fungal activity, among other biological processes within the soil-plant system, is considered essential for sustaining a plant cover in natural habitats (Requena et al. 1996), mycological data from Mediterranean ecosystems are scarce and fragmentary (Zervakis & Venturella 2007; Saitta et al. 2011; Venturella et al. 2011, suppl. ref. online). Some papers have dealt with the composition, succession and seasonal variation in fungal species colonizing leaf litter of Mediterranean evergreen species such as Myrtus communis L. (Pasqualetti et al. 1999a), Pistacia lentiscus L. and Quercus ilex L., showing that is possible to identify groups of microfungi characterizing the different matrices (Pasqualetti et al. 1999b).

Exogenous or environmental factors, regulating fungal colonization of organic substrates in Mediterranean maquis characterized by low water availability and low soil organic matter content, have been rarely analyzed (Ormeño et al. 2006). With regard to endogenous factors, plant secondary metabolite content has been suggested to be an important limiting parameter of microorganisms activity. Indeed, evergreen sclerophyllous species, dominating shrublands vegetation, produce high amounts of leaf phenolic compounds in response to different stress factors (Gershenzon 1984).

Concerning mycorrhizal symbioses, recent studies confirmed the occurrence of all known mycorrhizal types in Mediterranean biomes (Bedini et al. 2000; Pecoraro et al. 2013). Arbuscular mycorrhizal fungi have been described as essential components of Mediterranean ecosystems, for both revegetation of degraded lands and maintenance of soil structure, by increasing the supply of nutrients for plants, reducing water stress and improving soil aggregation (Caravaca et al. 2005). Diversity and specificity of ectomycorrhizal fungi have been analysed in a few forest ecosystems dominated, for instance, by Quercus ilex (Richard et al. 2005) or Q. suber L. (Aponte et al. 2010). Interesting results have been produced by assessing ectomycorrhizal diversity in typical Mediterranean maquis shrubs like Cistus species which are associated with more than 230 macrofungal taxa, including 35 Cistus-specific epigeous basidiomycetes such as Amanita cistetorum Contu & Pacioni, Hebeloma cistophilum Maire, Inocybe cistobulbipes Esteve-Rav. & Vila, Lactarius cistophilus Bon & Trimbach, Russula cistoadelpha M.M. Moser & Trimbach and Russula monspeliensis Sarnari (Comandini et al. 2006). Moreover, Cistus species are able to form both ectomycorrhizas and arbuscular mycorrhizas, simultaneously or over time (Smith & Read 1997). Mycorrhizal plasticity from one side and relationships with narrow host range fungi from the other side, may help plants to thrive in difficult environmental conditions (Perotto et al. 2013), and may represent an important adaptative trait in many Mediterranean maquis plant species that should receive more attention with regard to ecological effects.

Dow

nloa

ded

by [

Uni

vers

ity o

f Pe

rugi

a] a

t 00:

13 1

4 Ja

nuar

y 20

14

Accep

ted M

anus

cript

The role of soil microorganisms is fundamental in Mediterranean environments, where water is the major limiting factor to plant productivity, and summer drought limits the accumulation of organic matter and humus (Persiani et al. 2008). In these semiarid ecosystems, man-mediated degradative activities and environmental stress, including wildfires which represent the major disturbance affecting both plant and fungal communities (Hernández-Rodríguez et al. 2013), can promote desertification processes (Azcón-Aguilar et al. 2003). Fungal diversity plays an important role in Mediterranean habitats stability by buffering potential biotic and abiotic adverse effects to plants (Smith & Read 1997). Knowledge of the identities and roles of fungi colonizing Mediterranean habitats is of pivotal importance for conservation strategy, protection and management of natural areas.

In this work, we assessed fungal diversity in several areas characterized by Mediterranean maquis from continental Italy, Sicily and Greece. The primary objective of this study was to contribute to the analysis of biodiversity, ecology and distribution of macrofungi in evergreen sclerophyllous shrublands at different altitudes and distances from the seashore across the Mediterranean Basin. Macrofungi associated with evergreen sclerophyllous shrublands from Latium littoral (Italy)

Mycological research aiming at improving knowledge of the fungal biodiversity associated with Mediterranean vegetation were carried out from November 1996 to January 2001 in different ecosystems included in Macchiagrande Natural Reserve (suppl. ref. online). This area is situated in the northern part of Latium littoral (Central Italy), about 30 km far from Rome, has an extension of 280 ha and is constituted by a flatland that lies around the sea level. Macchiagrande Natural Reserve is covered by different Mediterranean vegetation types, from the sand dunes shrubland (Juniperetum macrocarpae-phoeniceae) to the plain deciduous oaks forest (Lathyro-Quercetum cerris) and the Mediterranean forest dominated by Quercus ilex belonging to Asplenio-Quercetum ilicis association. This Natural Reserve characterized by typical Mediterranean climate, with hot summers, a dry period of about four months and a wet period between October and April, represents a rare example of the natural coastal environment survived to anthropogenic disturbance and degradation in the Latium region. It has previously been described as an important center for conservation of rare fungal species (Pecoraro & Lunghini 2012).

During the above mentioned study period, sporomata were collected in seashore transects from the sand dunes to the Mediterranean woodlands. Fungal species were identified based on macro- and microscopic observations on fresh basidiomata. Notes on morphological characters and ecological conditions were recorded in the field, and samples were taken to the laboratory for microscopic examination. The specimens are deposited in the Herbarium D.L. of Rome (ROHB). A total of 381 macrofungal species belonging to 116 genera, included in Basidiomycota and Ascomycota, were identified (suppl. ref. online).

The main object of our study was the Mediterranean maquis, colonizing the inner core of the coastal dune system consisting of typical evergreen Mediterranean bush, where species exhibit a xeromorphic habit (e.g., Quercus ilex, Pistacia lentiscus, Arbutus unedo L., Cistus incanus L., C. salviifolius L., Phillyrea latifolia L., P. angustifolia L., Smilax aspera L., Juniperus phoenicea L., J. oxycedrus L.) due to existing abiotic conditions. In this ecosystem we collected some fungal taxa that are remarkable due to their ecology, rarity and limited geographical distribution.

Dow

nloa

ded

by [

Uni

vers

ity o

f Pe

rugi

a] a

t 00:

13 1

4 Ja

nuar

y 20

14

Accep

ted M

anus

cript

Two rare Amanita species with sabulicolous habit, A. dunensis R. Heim ex Bon & Andary and A. valens (E.-J. Gilbert) Bertault were collected for the first time in Italy. A. dunensis, described from France and reported in Spain (suppl. ref. online), is characterized by a slender aspect, non-bulbous stipe, very well-developed saccate volva, subcellular subhymenial trama, and ellipsoid to elongate-ellipsoid spores. It is the only known Amanita species with both a striate cap margin and amyloid spores (suppl. ref. online). A. dunensis differs morphologically from A. phalloides (Fr.) Link in the more ellipsoid spores and distinctly striate cap margin and from A. phalloides var. larroquei F. Massart & Beauvais (Neville & Poumarat 2004) by a different stipe : pileus ratio, absence of marginal pileus striations, the obvious bulbous stipe, and the pine-wood habitat (Pecoraro & Lunghini 2012).

A. valens is morphologically very similar to A. curtipes E.-J. Gilbert and A. ponderosa Malençon & R. Heim. Castro (1997) included A. curtipes and A. valens in a single species, A. curtipes, and proposed A. ponderosa as a variety of A. curtipes, A. curtipes var. ponderosa. Pinho-Almeida (1994) related phylogenetically A. ponderosa and A. curtipes with an evolutionary origin in A. valens. Neville & Poumarat (2004) proposed a new combination for A. valens as A. ponderosa f. valens. Molecular phylogenetic analysis revealed that A. ponderosa and A. curtipes are clearly distinct species, but it did not clarify the taxonomic status of A. valens (Moreno et al. 2008). A. valens has been reported in France, Spain and Morocco (Neville & Poumarat 2004).

Another sabulicolous basidiomycete observed was Agaricus devoniensis P.D. Orton, which normally grows on bare or lightly vegetated areas of coastal sand dunes, close to the sea (Cappelli 1984). The white or whitish-straw-yellow pileus, always covered with grains of sand, as well as the almost roundish spores 5 - 6 (7) × 4 – 5 (5,5) μm, distinguish A. devoniensis from the similar A. littoralis (Wakef. & A. Pearson) Pilát, also occurring on sand dunes; the latter is characterized by an ochraceous brown pileus and more elongated spores, typically 7 × 4.5 µm (Cappelli 1984). A. devoniensis occurs in several Italian regions and in many European countries (Onofri et al. 2005). This is the first record of A. devoniensis in Latium.

The hypogeous or semi-epigeous, gasteroid basidiomycete Melanogaster variegatus (Vittad.) Tul. & C. Tul. was accidentally found in sandy soil as it was dug out by animals. All species belonging to the genus Melanogaster Corda show the distinctive characters of blackish gleba and dark brown, smooth, truncate spores (Montecchi & Sarasini 2000). M. variegatus is characterized by irregular fruitbodies, ellipsoid, reniform, tuberiform, elastic in consistency, with ochraceous yellow, opaque and lightly tomentose surface, blackening when manipulated. It is well differentiated due to the presence of yellow pigments in peridium hyphae (absence of any reddish pigments), of yellowish (rarely withish) tramal plates and broadly ellipsoid, ovoid, almost rounded (at the base) spores. M. variegatus prefers xerophilous Mediterranean environments like bushy sand dunes, colonized by Juniperus and Cistus species (Montecchi & Sarasini 2000). This species has been reported in some regions of continental Italy and Sicily (Onofri et al. 2005; Venturella et al. 2006), but has not previously been reported in Latium. Rare and noteworthy macrofungi of Agaricales (Agaricomycetes, Basidiomycota) in the Umbria region (Italy)

The European Council for the Conservation of Fungi in 1991 and the Congress of European Mycologists at Kew (UK) in 1992 underlined the necessity to preserve, in

Dow

nloa

ded

by [

Uni

vers

ity o

f Pe

rugi

a] a

t 00:

13 1

4 Ja

nuar

y 20

14

Accep

ted M

anus

cript

each country, the diversity of mycobiota, with special emphasis on rare, threatened and biogeographically important taxa (Varese et al. 2011; Zotti et al. 2013).

The present study reports new chorological data of two rare basidiomycetes in the Umbria region: Battarrea phalloides (Dicks.) Pers., and Amanita eliae Quél. (Agaricales). The above-mentioned macromycetes were registered during mycological field trips in different floristic areas of Umbria region (central Italy): Collestrada forest (PG) and Polvese Island (Lake Trasimeno, PG).

The forest of Collestrada (PG) covers an area of approximately 136 ha (altitudinal range 250-306 m asl) which is characterized by exceptional species richness of plants, animals and macrofungi, and it was declared a Site of Community Importance (SCI) (IT5210077) in 2008. It is mainly characterized by mesophilic continental and submediterranean woodlands (Carpinus betulus L., Quercus cerris L., Quercus frainetto Ten., Quercus petraea (Matt.) Liebl.) and to a lesser extent by Mediterranean maquis belonging to the Rusco aculeati-Quercetum ilicis association. Studies on Collestrada forest macrofungi were conducted from 2011 to today with the sample plot method, mainly throughout the most widespread local plant communities (Angelini et al. 2012a). Until now, the forest mycobiota list includes about 320 fungal species (unpublished data).

The Polvese Island situated in the south eastern part of Lake Trasimeno (PG), has an extension of 69.6 ha (altitudinal range 258-309 m asl). It is the largest of Trasimeno Lake’s three islands and due to its interesting biodiversity, it was included in the SCI of Lake Trasimeno (IT5210018) (Angelini et al., in press). The vegetation is very differentiated and includes: a belt of marsh vegetation distributed on the lake shores, the Mediterranean therophytic grasslands, the olive groves and Mediterranean maquis represented by S. Lorenzo holm oak forest belonging Rusco aculeati-Quercetum ilicis, the same association found in Collestrada forest.

The two rare basidiomycetes (B. phalloides and A. eliae) were identified by morphological and molecular methods (Angelini et al. 2012b).

The rare and endangered specimens are supplied with description of macro- and micromorphological characteristics in order to allow further revision, as well as appropriate field notes.

B. phalloides and A. eliae are two rare species, in Umbria but also in Italy, and require management or conservation of their habitats to secure their long-term survival. In a few countries these species are on the red list data book and/or on the list of fungi to be protected by law. Battarrea phalloides, Syn. Meth. Fung. (Göttingen) 1: xiv, 129 (1801)

Description. Basidioma at first rounded, with exoperidium white, gelatinous, splitting horizontally into two parts leaving on the ground a gelatinous volva-like and on the apex of the stipe, a pileus-like, 20-40 mm wide, rusty brown, convex and fibrous. Endoperidium flattened-hemispherical at the top of the stipe, concave in the portion below, white, and caduceus (Figure 1a on line). Stipe 15-25 × 1.5-3 cm, pale brown to brown, woody, surface fibrous-scaly, hollow; volva-like ovate, formed of two white membranes with an internal gelatinous olive-yellow layer; subterranean, then superficial (Figure 1b on line).

Gleba brown-ferruginous, free after laceration of the concave part of the endoperidium, soon reduced to a dusty pile (Figure 1c on line); with two types of threads: pseudocapillitium with hyphae up to 5 µm diam, mostly thin walled, smooth, septate, sparsely branched, hyaline to pale yellow, with clamps (Figure 1d on line);

Dow

nloa

ded

by [

Uni

vers

ity o

f Pe

rugi

a] a

t 00:

13 1

4 Ja

nuar

y 20

14

Accep

ted M

anus

cript

elaters 3.5-7 µm diam, 32-70 µm long, cylindrical, tapered, with spiral thickenings, pale yellow, aseptate, not branched (Figure 1e on line). Spores: globose to subglobose, 5.5-6.5 µm, yellowish to yellow-brown; densely verruculose (Figure 1f on line).

Material examined. Italy, Perugia: Polvese Island, Lake Trasimeno (43.116723°N, 12.145988°E, 256 m asl), 01 June 2012, on soil associated with Cupressus sempervirens L. Representative voucher specimen were deposited at the herbarium of Sect. ‘Plant Biology and Geobotany’ of University of Perugia (Italy).

Note. Year of the first record in Polvese Island: 1998. Molecular identification. The internal transcribed spacer (ITS) region of the

ribosomal DNA was amplified by PCR and sequenced. BlastN search of the GenBank database with the 713 bp ITS sequence (GB accession no. KF780873) identified several similar sequences ascribed to the species B. phalloides and B. stevenii.

Molecular analyses of the ITS region previously performed on samples collected from different localities failed to resolve B. phalloides and B. stevenii into two separate species wich have been regarded as conspecific (Martín & Joannesson 2000). The ITS analysis however have shown a high level of genetic variability within this species with the identification of four clusters/clades (Martín et al. 2013). Phylogenetic analysis (Figure 2 on line) performed by using sequences retrieved from GenBank indicates that the Battarrea sample collected in Umbria belong to the clade A described by Martín et al. (2013).

Geography. Global distribution: Africa, Asia, Australia, Europe and North and South America (suppl. ref. online). Distribution in Europe: The fungus is rather widespread from the Mediterranean to the nemoral zone. It is mainly restricted to the lowlands. Distribution in Italy: Calabria, Emilia Romagna, Latium, Lombardy, Sardinia, Sicily, Tuscany and Trentino Alto Adige (Onofri et al. 2005, suppl. ref. online). This is the first report for this species from Umbria region.

Ecology. On dry, sandy localities, sunny edges, and in clearings of different types of decidous, mixed, and coniferous woodlands. It often colonises anthropogenic biotopes as hedges, roadsides, gardens, parks, and cemeteries. Fruiting from spring to autumn.

Status of conservation. The species is red listed in Armenia, Austria, Czech Republic, France, Germany, Hungary, FYR Macedonia, Poland, Serbia, Spain, and United Kingdom. Amanita eliae, Mém. Soc. Emul. Montbéliard, Sér. 2(5): 230 (1872)

Description. Pileus 70-100 mm wide, convex to plano-convex, pale brown tints in young specimens, becoming flesh-pink to brownish pink at least in part in apparently older basidiomata (dried in situ), otherwise color unchanging when bruised; context fleshy, up to 3 mm thick at center, unchanging, thinning toward margin, white; margin short and indistinctly striate; universal veil as numerous white patches scattered over almost entire surface, soon becoming fixed to drying pileipellis (Figure 3a online). Lamellae free in older basidiomata, however with lines on stipe apex, subcrowded, white in fresh basidiomata, becoming ochraceous to dark ochraceous in exsiccata; lamellulae truncate to subtruncate, of diverse lengths, plentiful (Figure 3b online). Stipe 10-12 × 0.5-1 cm, cylindric above bulb, white, with small squamules turning brownish with age, with more than half of its length buried in substrate; bulb abruptly or gradually turbinate, with short pseudorhiza; context fleshy; membranous, white and fugacious annulus; universal veil often as collar above bulb, sometimes

Dow

nloa

ded

by [

Uni

vers

ity o

f Pe

rugi

a] a

t 00:

13 1

4 Ja

nuar

y 20

14

Accep

ted M

anus

cript

becoming broken as a collar ring of verrucose irregular scales or warts where stipe and bulb come in contact (Figure 3b, c online). Odor not distinct.

Basidiospores 9.8-14 × 5.5-8 µm, inamyloid, hyaline, colorless, ellipsoid to elongate, occasionally cylindrical, smooth, thin-walled, usually at least somewhat adaxially flattened; apiculus broad, rounded, sublateral; contents including at least one or two guttules (Figure 3d). Basidia, clavate, with 4 and sometimes 2 sterigmata; size can be very variable depending on the degree of maturation; clamp connections not observed (Figure 3e). Cystidia absents. Lamellar trama bilateral, with narrow or inflated elements gradually diverging from central stratum. Universal veil on pileus: in contact with suprapellis; filamentous hyphae 3.5-10 µm wide, frequently branching, plentiful, with frequently incrusted walls; plentiful subisodiametric to ovoid and (dominantly) isodiametric elements 31-80 × 14-35 µm, hyaline, colorless, with a septum at the base, peduncolate (Figure 3f). On bulb: similar to elements of universal veil on pileus surface, with more interwoven filamentous hyphae 5-12 µm wide, often with thickened walls, frequently with incrustation.

Material examined. Italy, Perugia: Collestrada forest (43.07893889 °N, 12.46621944° °E, 251 m asl), 16 May 2013 (date of the first collection), on soil associated with Q. frainetto Ten. Representative voucher specimens were deposited at the herbarium of Sect. ‘Plant Biology and Geobotany’ of University of Perugia (Italy).

Molecular identification. Molecular identification was performed by sequencing the ITS rDNA region. A 230 bp sequence containing the ITS1 ribosomal spacer was obtained (GB accession no. KF780872) and used to search for similar sequences in the GenBank database by using BlastN. The species assignment was confirmed since the best match was found with sequence JF907763 identified as A. eliae (Osmundson et al. 2013). It is worth of noting, however, that a single partial ITS sequence relative to this species is present in the database sharing only 94% identity with the sequence obtained from our sample. Preliminary phylogenetic analysis (Figure 4 on line), using sequences representative of the subgenus Amanita according to Justo et al. (2010), suggests that A. eliae is related to species in the Amanitopsis section. It will be intersting to analyze more samples from different provenances to better evaluate phylogenetic relationship with other Amanita species and the extent of intraspecific variability in A. eliae.

Ecology. Associated with plant species in clayey-siliceous, usually hardwood habitats: oaks (Quercus spp.), beech (Fagus sylvatica L.), chestnut (Castanea sativa Mill.), hornbeam (Carpinus betulus), heart (Corylus avellana L.) sometimes mixed with Scots pine (Pinus sylvestris L.), appearing as early as May, but mainly in June and July, sometimes until October, in hills (300-400 m asl), or plains, usually 1-2 or 3 basidiomata.

Geography. Global distribution: southern Europe, Mediterranean Africa, and Mexico (suppl. ref. online). Distribution in Europe: France, Germany, Greece, Italy, Netherlands, Spain, Switzerland (suppl. ref. online). Distribution in Italy: Calabria, Campania, Emilia Romagna, Latium, Liguria, Lombardy, Piedmont, Apulia, Sicily, Tuscany, Veneto (Onofri et al. 2005, suppl. ref. online). This is the first report for this species from Umbria region.

Status of conservation. The species is red listed in Switzerland. Target species in arid and semi-arid environments of Sicily

Dow

nloa

ded

by [

Uni

vers

ity o

f Pe

rugi

a] a

t 00:

13 1

4 Ja

nuar

y 20

14

Accep

ted M

anus

cript

Sicily is an area of high phytogeographic and mycological interest in the Mediterranean Region. The most represented type of vegetation is the Mediterranean maquis which occupies many coastal areas and inland areas. Usually such vegetation is attributed to the Mediterranean arid and the Temperate Mediterranean belts. As regards fungi in particular, this environment has been partially studied and data are available through studies dealing with biodiversity of epigeous and hypogeous macromycetes (Venturella et al. 2006; Saitta et al. 2011).

In order to increase the knowledge on the ecology and distribution of macrofungi growing in the Mediterranean maquis the distributional and ecological data of some rare or uncommon species along an altitudinal transect from the sea level up to 550 m a.s.l. are here reported.

In particular four different areas located at different altitudinal levels were investigated:

a) “Capo Rama Natural Reserve” (province of Trapani, N.E. Sicily, 30 m a.s.l.);

b) “Birribaida wood” (province of Trapani, N.E. Sicily 90 m a.s.l.);

c) “Monte Catalfano Natural Reserve” (province of Palermo, N. Sicily, 300-400 m a.s.l.);

d) ”Monte Petroso wood” (province of Palermo, N. Sicily, 450-550 m a.s.l.)

Target species 1. Disciseda bovista (Klotzsch) P. Henn. Recently collected in the “Capo Rama Natural Reserve”, an area characterized by a

natural population of Chamaerops humilis L. and many endemic plants of the “Sicilian floristic domain” (suppl. ref. online). D. bovista (Figure 5 online) grows on sandy substrates mainly covered by Ephedra fragilis Desf. and Thapsia garganica L. The entire vegetation corresponds to the association Chamaeropo-Quercetum calliprini Brullo et Marcenò. This association is distributed along the coastal belt, in particularly arid environments. D. bovista is a rare species distributed in America and Europe (suppl. ref. online), and it was recently included in several Red-list of rare and threatened macromycetes in Europe (Austria, Denmark, Italy, Poland, Netherland, Sweden, Switzerland). The genus Disciseda Czern. includes semi-hypogeous gasteroid species belonging to the family Agaricaceae Cheval. Until today the distribution in Italy was limited to three regions, i.e. Piedmont, Sardinia and Sicily (suppl. ref. online). The collection from Sicily is a new record for this region and widens the distribution of this taxon in Italy. Besides the type of habitat confirms that D. bovista is strictly linked to xeric environments.

2. The infrequent Colus hirudinosus Cavalier & Séchier (Figure 6 online) was collected in the “Capo Rama Natural Reserve”, the “Birribaida wood”, and the “Monte Catalfano Natural Reserve”. All collected samples were found growing in areas characterized by a mixed vegetation of Pistacia lentiscus and C. humilis. The “Birribaida wood” is a very interesting forest remnant, including a strip of “forest-maquis” dominated by Olea europea L. var. sylvetris Brot., Ceratonia siliqua L. and Pistacia lentiscus. It is really striking, that the Emperor Frederick II used this area as a hunting ground mainly due to its particular orography, the richness of the vegetation, and abundance of animals. Nowadays, the original and impenetrable forest is almost totally destroyed mainly because of human activities. The area of Monte Catalfano

Dow

nloa

ded

by [

Uni

vers

ity o

f Pe

rugi

a] a

t 00:

13 1

4 Ja

nuar

y 20

14

Accep

ted M

anus

cript

Natural Reserve is characterized by the presence of a small group of plants of Quercus auzandrii Gren. & Godr. (=Quercus soluntina Tin. ex Lojac.), a species which was described by Lojacono-Pojero as a hybrid generated by the natural crossing of Q. coccifera L. and Q. ilex. The surrounding vegetation is mainly composed of Quercus ilex, Pistacia lentiscus, P. terebinthus L., Phillyrea latifolia and Chamaerops humilis.

C. hirudinosus is distributed in Asia, Africa, Europe and America. In Italy its occurrence is restricted in the South (Onofri et al. 2005).

3. The infrequent Lyophyllum littoralis (Ballero & Contu) Contu (Figure 7 online) was collected in the “Birribaida wood”. The area of collection is characterized by P. lentiscus and Rhamnus alaternus L. This taxon was previously described as Calocybe littoralis Ballero & Contu and then transferred to the genus Lyophyllum. The distribution of L. littoralis is limited to Sardinia and Sicily (suppl. ref. online).

4. The very rare wood-inhabiting fungi, Ceriporia aurantiocarnescens (Henn.) M. Pieri & B. Rivoire and Gloeodontia columbiensis Burt ex Burds. & Lombard, have been collected in the “Monte Petroso wood”. In this area the vegetation is distributed in discontinuous belts and is mainly characterized by a mixed oak wood with a prevalence of Quercus ilex.

The corticioid genus Gloedontia includes seven taxa, and only G. columbiensis and G. subasperispora (Litsch.) E. Larsson & K.H. Larsson are present in Europe. The distribution of G. columbiensis is limited to USA, Canada, Switzerland, Iran, Turkey, the Canary Islands and Italy (suppl. ref. online). G. columbiensis is very rare in Europe and our record from Sicily represents the second from Italy. Previously it was collected in Sardinia on Erica arborea L. (Bernicchia & Gorjón 2010). The hymenophore is odontioid with aculei up to 1 mm long, monomitic hyphal system, gloeocystidia, amyloid spores, thick-walled and encrusted cystidia which characterize this interesting species. The Sicilian specimens were collected on fallen branches of Quercus ilex.

Ceriporia aurantiocarnescens (Henn.) M. Pieri & B. Rivoire is a rare polyporoid species previously collected in Italy on wood of Abies alba L. (Bernicchia et al. 2007). The European distribution is limited to Estonia and France (suppl. ref. online). C. aurantiocarnescens was collected on a small trunk of Q. ilex. The presence of C. aurantiocarnescens in Sicily confirms the adaptation abilities of this species to arid and semiarid Mediterranean climatic conditions. Wood-inhabiting macrofungi associated with maquis vegetation in two Cyclades islands, Aegean Sea, Greece Naxos is the biggest of the Cyclades islands situated in the central Aegean Sea with a surface area of 442 km2; it is characterized by a rich bas-relief with three high mountains reaching altitudes of ca. 1,000 m asl. Amorgos lies southeast of Naxos and has a surface area of 121 km2. It is a coline and rocky island and the highest peak has an altitude of 823 m. The climate of both islands is characterized as intense hot-summer Mediterranean with a long dry period of almost seven months. The annual precipitation is 300–400 mm and the wet period is between November and March.

Naxos and Amorgos possess a high plant species diversity and several endemics. Local maquis vegetation is mainly characterized by low shrubs and phrygana communities, while Juniperus macrocarpa Sibth. & Sm., J. phoenicea, Quercus coccifera, Pistacia lentiscus and Ceratonia siliqua L. are prominent in the littoral zone and in low elevations.

Dow

nloa

ded

by [

Uni

vers

ity o

f Pe

rugi

a] a

t 00:

13 1

4 Ja

nuar

y 20

14

Accep

ted M

anus

cript

Long-term field work in selected Aegean islands contributed at inventorying specific elements of their fungal diversity (Polemis et al. 2012a, b, 2013). Especially as regards wood-inhabiting macrofungi, 92 corticioid and poroid basidiomata were collected from dead woody parts of shrubs and phrygana in altitudinal transects from sea level to 500 m in Naxos and Amorgos. These collections were identified to 32 species and 20 genera, and they were associated with a total of 28 plant species.

In the following list, species of macrofungi are presented together with their substrates and are divided in three altitude zones of occurrence, i.e. the littoral (0–50 m), the lowland (51–250 m) and the hillside (251–500 m). The abbreviations “A” and “N” refer to Amorgos and Naxos respectively while the number following each of them indicates the collection site as depicted in the pertinent maps (Figure 8a, b online). Species that are recorded for the first time in Greece are indicated by an asterisk (*), while some first-ever records of substrates in a world-wide context are also noted. *Aleurodiscus dextrinoideocerussatus Manjón, M.N. Blanco & G. Moreno: 0–50 m on Anthyllis hermaniae L. (N5), J. phoenicea (N9), Calycotome villosa L. (A9); 51–250 m on Salvia fruticosa Mill. and P. lentiscus (A12); 251–500 m on C. villosa (N2). Rare species, not previously reported from southeast Europe. Byssomerulius corium (Pers.) Parmasto: 51–250 m on C. siliqua (A8), P. lentiscus (N1). Ceriporia purpurea (Fr.) Donk: 51–250 m on C. siliqua (A8). Fibrodontia gossypina Parmasto: 0–50 m on J. macrocarpa (N9), J. phoenicea (A11); 51–250 m on J. phoenicea and P. lentiscus (A12), S. fruticosa (A3). Gloeophyllum abietinum (Bull.) P. Karst.: 0–50 m and 51–250 m on J. phoenicea (N9 and A12 respectively). Hyphodontia juniperi (Bourdot & Galzin) J. Eriksson & Hjortstam: 0–50 m on Coridothymus capitatus L. (N9, A9); 51–250 m on Q. coccifera (A2), S. fruticosa (A3); 251–500 on C. spinosa (A10), Origanum sp. and S. fruticosa, (A4), Satureja thymbra L. (A8). Hyphodontia quercina (Pers.) J. Erikss.: 51–250 m on Olea europaea L. (A3). Hyphodontia sambuci (Pers.) J. Erikss.: 0–50 m on Centaurea spinosa L. and Juniperus sp. (N5), C. capitatus (A9); 51–250 m on O. europaea (N6), Ballota acetabulosa (A8), Phlomis fruticosa L. (A12), S. fruticosa (A4); 251–500 on C. spinosa and Opuntia ficus-indica (L.) Mill. (A10), Ficus carica (A5), S. fruticosa and S. thymbra (A8). *Peniophora boidinii Reid.: 51–250 m on Cistus sp. (N6). Very rare in Europe not previously recorded from eastern Mediterranean. *Peniophora cinerea (Pers.) Cooke: 51–250 m on C. siliqua (A3). *Peniophora junipericola J. Erikss.: 0–50 m on J. phoenicea (N8). Peniophora lycii (Pers.) Höhn. & Litsch.: 0–50 m on Tamarix sp. (N3, N5); 51–250 m on A. hermaniae and P. lentiscus (N1), C. siliqua and Cistus sp. (A8) and Euphorbia dendroides L. (A3); 251–500 m on C. villosa (N2), Q. coccifera (A1, A5), Spartium junceum L. (A4). *Peniophora piceae (Pers.) J. Erikss.: 0–50 m on J. phoenicea (N9). First-ever record on this substrate.

Dow

nloa

ded

by [

Uni

vers

ity o

f Pe

rugi

a] a

t 00:

13 1

4 Ja

nuar

y 20

14

Accep

ted M

anus

cript

*Peniophora pithya (Pers.) J. Erikss.: 51–250 m on P. lentiscus (N1). First-ever record on this substrate. Peniophora quercina (Pers.) Cooke: 51–250 m and 251–500 m on Q. coccifera (A6). *Peniophora tamaricicola Boidin & Malenç.: 0–50 m on Tamarix sp. (N5); 51–250 m on P. lentiscus (N1). First-ever report from eastern Mediterranean and first record on P. lentiscus. Peniophorella praetermissa (P. Karst.) K.H. Larss.: 51–250 m on Q. coccifera (A4), C. siliqua (N6, A8, A12); 251–500 m on S. fruticosa (A4). *Peniophorella tsugae (Burt) K.H. Larss.: 0–50 m on J. phoenicea (N8). First-ever record on this substrate. *Perenniporia meridionalis Decock & Stalpers: 0–50 m on J. phoenicea (N8). Perenniporia ochroleuca (Berk.) Ryvarden: 251–500 m on Myrtus communis (N2). *Phanerochaete martelliana (Bres.) J. Erikss. & Ryvarden: in 51–250 m on S. junceum (A1). First-ever record on this substrate. Phanerochaete sordida (P. Karst.) J. Erikss. & Ryvarden: 0–50 m on Tamarix sp. (N5). Phlebia rufa (Pers.) M.P. Christ.: 251–500 m on E. dendroides (A4). Polyporus meridionalis (A. David) Jahn: 0–50 m on Erica manipuliflora Salisb. (N9), Q. coccifera (N8); 51–250 m on P. fruticosa (A12). Postia subcaesia (A. David) Jülich: 51–250 m on P. lentiscus (N1), C. siliqua (A8). Radulomyces confluens (Fr.) M.P. Christ.: 251–500 m on Lavandula stoechas L. (N4), S. spinosum (A1). Schizopora paradoxa (Schrad.) Donk: in 51–250 m on O. europaea (A3). Steccherinum fimbriatum (Pers.) J. Erikss.: 51–250 m on Euphorbia sp. (N7), O. europaea (A8); 251–500 on C. villosa (N2). Steccherinum ochraceum (Pers.) Gray: 51–250 m on Cistus sp. (A8), Genista acanthoclada D.C. and Q. coccifera (A6); 251–500 on C. villosa (N2), Q. coccifera (A1). Stereum hirsutum (Willd.) Gray: 251–500 on Q. coccifera (N2). *Vararia ochroleuca (Bourdot & Galzin) Donk: 0–50 m on G. acanthoclada (A9). First-ever record on this substrate. Vuilleminia megalospora Bres.: 51–250 m and 251–500 m on Q. coccifera (A1). Xylodon pruni (Lasch) Hjortstam & Ryvarden: 251–500 m on C. villosa (N2).

Among the fungi listed above, the genera Peniophora Cooke and Hyphodontia J. Erikss. are represented with the highest number of species (eight and three respectively), whereas the most common taxa detected were A. dextrinoideocerussatus, H. juniperi, H. sambuci and P. lycii. These dominant decomposers of woody low shrubs and phrygana show various adaptations in the xerothermic conditions prevalent in Cyclades through the colonization of plant twigs and small stems (rarely exceeding 5 cm), which are particularly exposed to solar radiation and rapid drying.

Growing on restricted substrate surface area, e.g. small diameter wood-piece, is a common feature of drought resistant corticioid (and some resupinate polyporoid) species with effused basidiomata (Junninen et al. 2006). The presence of a dense layer of branched hyphidia (acanthohyphidia, dendrohyphidia or dichohyphidia) covering

Dow

nloa

ded

by [

Uni

vers

ity o

f Pe

rugi

a] a

t 00:

13 1

4 Ja

nuar

y 20

14

Accep

ted M

anus

cript

the hymenial surface is another relevant character observed in A. dextrinoideocerussatus, P. lycii, V. ochroleuca and V. megalospora. This type of hymenial surface was also described as “catahymenium” (Lemke 1964). Basidiomata with a catahymenium are adapted to resist dry spells and to rapidly resume sporulation after adequate rehydration (Larsson & Larsson 2003).

Another adaptation to drought by many corticioid species is the accumulation of polysaccharides and proteins resulting in the formation of a ceraceous, resinous or gelatinous matrix, which protects basidia from desiccation. Genera with gelatinized hyphae such as Phlebia Fr. and Radulomyces M.P. Christ. can dry out and rehydrate very quickly. Characteristic examples of such adaptations could be observed in P. marteliana, a species with a limited distribution in dry Mediterranean or steppe-like habitats (Eriksson et al. 1978; Bernicchia & Gorjón 2010), and in V. megalospora (common in southern Europe; Bernicchia & Gorjón 2010). The latter is adapted to grow initially under the bark by producing very thin, ceraceous or gelatinous basidiomata with numerous dendrohyphidia, very large basidia and spores.

Many species of the genus Aleurodiscus Rabenh. ex J. Schröt. inhabiting dry branches, possess large sized spores. This feature ensures an ample supply of nutrients for the production of hyphae that can penetrate quickly beneath the wood cortex, where they are protected from drought (Núñez & Ryvarden 1997). Members of the subgenus Peniophora (incl. all Peniophora findings noted here) show a pronounced xerophilic adaptation allowing growth on hanging and/or rapidly drying dead branches (Eriksson et al. 1978). Some resupinate poroid species produce chlamydospores (e.g. Schizopora paradoxa) while other corticioids with filmy basidiomata are capable to resume growth very quickly when moist conditions appear (Rayner & Boddy 1988). Furthermore, the existence of metuloids, and binding and skeletal hyphae with thickened walls confer toughness to basidiomata and permit the colonization of less decayed pieces of wood.

Conclusions This work represents a contribution to the analysis of distribution, ecology and diversity of macrofungi in evergreen sclerophyllous shrublands at different altitudes and distances from the seashore across the Mediterranean Basin. Fungal diversity was investigated with different methodologies in several areas characterized by Mediterranean maquis, from three Italian Regions and two Greek islands. Valuable information about ectomycorrhizal species, saprothrophs and wood-inhabiting fungi that are remarkable due to their ecology, rarity and limited geographical distribution were reported. The finding of several fungal species growing on new substrates and recorded for the first time in new Countries, like Italy and Greece, expands  the known ecology and geographic distribution  in Europe of  the examined  taxa and constitutes, on the other hand, the evidence of the limited knowledge about the biology of these fungi. There is still more to be learnt about fungal diversity in Mediterranean environments and its impact on the stability of these delicate ecosystems. Improved understanding of the the identity and ecology of mycetes associated with Mediterranean maquis may represent a crucial aspect for the conservation of natural habitats that are considered hot spots of biodiversity. Acknowledgements

Dow

nloa

ded

by [

Uni

vers

ity o

f Pe

rugi

a] a

t 00:

13 1

4 Ja

nuar

y 20

14

Accep

ted M

anus

cript

The authors thank the Estonian Science Foundation and the European Social Fund (Mobilitas Postdoctoral Research Grants MJD135) and the Greek General Secretariat of Research and Technology project THALIS (“Metagenomics of ligninolytic microorganisms – Bioconversion of plant by-products into high-added value products”, MIS 377062). for financial support. References Angelini P, Bistocchi G, Arcangeli A, Venanzoni R. 2012a. Preliminary check-list of

the macromycetes from Collestrada forest ecosystems in Perugia (Italy). Mycotaxon 120: 505-506.

Angelini P, Rubini A, Gigante D, Reale L, Pagiotti R, Venanzoni R. 2012b. The endophytic fungal communities associated with the leaves and roots of the common reed (Phragmites australis) in Lake Trasimeno (Perugia, Italy) in declining and healthy stands. Fungal Ecol, 5(6): 683-693.

Angelini P, De Angelis MC, Guerzoni RP, Gigante D, Rubini A, Properzi P, Venanzoni R. Wood identification of pile dwellings from the Bronze Age San Savino site (Lake Trasimeno, central Italy). Plant Biosyst (in press), DOI:10.1080/11263504.2013.814604.

Aponte C, García LV, Marañón T, Gardes M. 2010. Indirect host effect on ectomycorrhizal fungi: Leaf fall and litter quality explain changes in fungal communities on the roots of co-occurring Mediterranean oaks. Soil Biology & Biochemestry 42: 788-796.

Azcón-Aguilar C, Palenzuela J, Roldán A, Bautista S, Vallejo R, Barea JM. 2003. Analysis of the mycorrhizal potential in the rhizosphere of representative plant species from desertification-threatened Mediterranean shrublands. Applied Soil Ecology 22: 29-37.

Bedini S, Maremmani A, Giovannetti M. 2000. Paris-type mycorrhizas in Smilax aspera L. growing in a Mediterranean scherophyllous wood. Mycorrhiza 10: 9-13.

Bergero R, Perotto S, Girlanda M, Vidano G, Luppi AM. 2000. Ericoid mycorrhizal fungi are common root associates of a Mediterranean ectomycorrhizal plant (Quercus ilex). Molecular Ecology 9: 1639-1649.

Bernicchia A, Gorjón SP. 2010. Corticiaceae s.l. Fungi Europaei 12. Alassio (Italy): Candusso. p. 1008.

Bernicchia A, Savino E, Gorjón SP. 2007. Aphyllophoraceous wood-inhabiting fungi on Abies alba in Italy. Mycotaxon 100: 185–188.

Brooks TM, Mittermeier RA, Mitiermeie CG, da Fonseca GAB, Rylands AB, Konstant WR, Flick P, Pilgrim J, Oldfield S, Magin G, Hilton-Taylor C. 2002. Habitat loss and extinction in the hotspots of biodiversity. Conservation Biol 16: 909-923.

Cappelli A. 1984. Agaricus L.: Fr. Ss. Karsten (Psalliota Fr.). Fungi Europaei. Saronno: Libreria editrice Biella Giovanna. p 1-558.

Caravaca F, Alguacil MM, Barea JM, Roldán A. 2005. Survival of inocula and native AM fungi species associated with shrubs in a degraded Mediterranean ecosystem. Soil Biology & Biochemistry 37: 227-233.

Castro ML. 1997. Analyse critique des taxons appartenant au groupe Amanita curtipes Gilbert (Agaricales, Basidiomycotina). Distribution dans la Peninsule Iberique. Doc Mycol 27: 43-51.

Comandini O, Contu M, Rinaldi AC. 2006. An overview of Cistus ectomycorrhizal fungi. Mycorrhiza 16: 381-395.

Dow

nloa

ded

by [

Uni

vers

ity o

f Pe

rugi

a] a

t 00:

13 1

4 Ja

nuar

y 20

14

Accep

ted M

anus

cript

Eriksson J, Hjorstam K, Ryvarden L. 1978. The Corticiaceae of North Europe Volume 5: Mycoaciella–Phanerochaete, Oslo. 887–1047 pp.

Gershenzon J. 1984. Changes in the level of plant secondary metabolites production under water and nutrient stress. In: Loewus FA, Timmermenn BN, Steelink C, editors. Phytochemical Adaptation to Stress, Recent Advances in Phytochemistry. New York: Plenum Press. p 273-320.

Hernández-Rodríguez M, Oria-de-Rueda JA, Martín-Pinto P. 2013. Post-fire fungal succession in a Mediterranean ecosystem dominated by Cistus ladanifer L. Forest Ecology and Management 289: 48-57.

Junninen K., M. Similä M, Kouki J, Kotiranta H. 2006. Assemblages of wood-inhabiting fungi along the gradients of succession and naturalness in boreal pine–dominated forests in Fennoscandia. Ecography 29: 75–83.

Justo A, Morgenstern I, Hallen-Adams HE, Hibbett DS. 2010. Convergent evolution of sequestrate forms in Amanita under Mediterranean climate conditions. Mycologia 102: 675-688.

Larsson E, Larsson K–H. 2003. Phylogenetic relationships of russuloid basidiomycetes with emphasis on aphyllophoralean taxa. Mycologia 95: 1037–1065.

Lemke PA. 1964. The genus Aleurodiscus (sensu stricto) in North America. Can J Bot 42: 213–282.

Martín MP, Johannesson H. 2000. Battarrea phalloides and B. stevenii, insight into a long-standing taxonomic puzzle. Mycotaxon 74: 67-75.

Martín MP, Rusevska K, Duenas M, Karadelev M. 2013. Battarrea phalloides in Macedonia: Genetic variability, distribution and ecology. Acta Mycologica 48: 113-122.

Mazzoleni S, Loporto A, Blasi C. 1992. Multivariate analysis of climatic patterns of the Mediterranean basin. Vegetatio 98: 1-12.

Montecchi A, Sarasini M. 2000. Funghi ipogei d’Europa. Vicenza: Associazione Micologica Bresadola, Fondazione Centro Studi Micologici. p. 714.

Moreno G, Platas G, Peláez F, Bernedo M, Vargas A, Daza A, et al. 2008. Molecular phylogenetic analysis shows that Amanita ponderosa and A. curtipes are distinct species. Mycol Progress 7: 41-47.

Neville P, Poumarat S. 2004. Amaniteae. Amanita, Limacella & Torrendia. Fungi Europaei 9. Candusso. p 1-1119.

Núñez M, Ryvarden L. 1997. The genus Aleurodiscus (Basidiomycotina). Sats & Trykk, Oslo. 164 p.

Onofri S, Bernicchia A, Filipello Marchisio V, Padovan F, Perini C, Ripa C, et al. 2005. Checklist dei funghi italiani - Checklist of Italian fungi, Basidiomycetes. Sassari: C. Delfino ed. p 1-380.

Ormeño E, Baldy V, Ballini C, Larchevêque M, Périssol C, Fernandez C. 2006. Effects of environmental factors and leaf chemistry on a leaf litter colonization by fungi in a Mediterranean shrubland. Pedobiologia 50: 1-10.

Osmundson TW, Robert VA, Schoch CL, Baker LJ, Smith A, Robich G, Mizzan L, Garbelotto MM. 2013. Filling gaps in biodiversity knowledge for macrofungi: contributions and assessment of an herbarium collection DNA barcode sequencing project. PLoS ONE 8(4): E62419.

Pasqualetti M, Mulas B, Zucconi L, Rambelli A. 1999a. Succession of mirofungal communities on Myrtus communis leaf litter in a Sardinian Mediterranean maquis ecosystem. Mycol. Res. 103(6): 724-728.

Dow

nloa

ded

by [

Uni

vers

ity o

f Pe

rugi

a] a

t 00:

13 1

4 Ja

nuar

y 20

14

Accep

ted M

anus

cript

Pasqualetti M, Fonck M, Rambelli A, Mulas B. 1999b. Analysis of populations of Beltrania rhombica, a mitosporic fungus, in ecosystems with Mediterranean maquis. Plant Biosyst 133: 265-276.

Pecoraro L, Lunghini D. 2012. First record of A. dunensis in Italy. Mycotaxon 122: 261-263.

Pecoraro L, Girlanda M, Kull T, Perini C, Perotto S. 2013. Fungi from the roots of the terrestrial photosynthetic orchid Himantoglossum adriaticum. Plant Ecology and Evolution 146 (2): 145–152

Pereira PT, de Carvalho MM, Gírio FM, Roseiro JC, Amaral-Collaço MT. 2002. Diversity of microfungi in the phylloplane of plants growing in a Mediterranean ecosystem. J Basic Microbiol 42(6): 396-407.

Perotto S, Angelini P, Bianciotto V, Bonfante P, Girlanda M, Kull T, et al. 2013. Interaction of fungi with other organisms. Plant Biosyst 147: 208-218.

Persiani AM, Maggi O, Montalvo J, Casado MA, Pineda FD. 2008. Mediterranean grassland soil fungi: Patterns of biodiversity, functional redundancy and soil carbon storage. Plant Biosyst 142: 111-119.

Pinho-Almeida F. 1994. Estudos taxonómicos no género Amanita. Secçao Amidella- Complexo Lepiotoides: A. curtipes Gilbert, A. lepiotoides Barla e A. ponderosa Malençon & Heim. Rev Biol (Lisboa) 15: 131-151.

Polemis E, Dimou DM, Tzanoudakis D, Zervakis GI. 2012a. Annotated Checklist of Basidiomycota (Subclass Agaricomycetidae) from the islands of Naxos and Amorgos (Cyclades, Greece). Ann Bot Fenn 49(3): 145–161.

Polemis E, Dimou, DM, Tzanoudakis D, Zervakis GI. 2012b. Diversity of Basidiomycota (subclass Agaricomycetidae) in the island of Andros (Cyclades, Greece). Nova Hedwigia 95: 25–58.

Polemis E, Dimou DM, Zervakis GI. 2013. The family Hymenochaetaceae (Agaricomycetes, Basidiomycota) in the islands of the Aegean Archipelago (Greece). Plant Biosyst 147: 306–314.

Rayner ADM, Boddy L. 1988. Fungal decomposition of wood: its biology and ecology. John Wiley & Sons Ltd., Chichester. 587 p.

Requena N, Jeffries P, Barea JM. 1996. Assessment of natural mycorrhizal potential in a desertified semiarid ecosystem. Appl Environ Microbiol 62: 842-847.

Richard F, Millot S, Gardes M, Selosse M-A. 2005. Diversity and specificity of ectomycorrhizal fungi retrieved from an old-growth Mediterranean forest dominated by Quercus ilex. New Phytol 166: 1011-1023.

Saitta A, Bernicchia A, Gorjón SP, Altobelli E, Granito VM, Losi C, et al. 2011. Biodiversity of wood-decay fungi in Italy. Plant Biosyst 145(4): 958-968.

Smith SE, Read DJ. 1997. Mycorrhizal Symbiosis, 2nd ed. San Diego, California: Academic Press.

Turrini A, Sbrana C, Strani P, Pezzarossa B, Risaliti R, Giovannetti M. 2010. Arbuscular mycorrhizal fungi of a Mediterranean island (Pianosa), within a UNESCO Biosphere Reserve. Biol Fertil Soils 46: 511-520.

Varese GC, Angelini P, Bencivenga M, Buzzini P, Donnini D, Gargano M.L, Maggi O, Pecoraro L, Persiani AM, Savino E, Bigini V, Turchetti B, Vannacci G, Venturella G, Zambonelli A. 2011. Ex-situ conservation and exploitation of fungi in Italy. Plant Biosyst 145: 997-1005.

Venturella G., Pecorella E., Saitta A., Zambonelli A., Morara M. 2006. Ecology and distribution of hypogeous fungi from Sicily (southern Italy). Cryptogamie Mycologie 27(3): 201-217.

Dow

nloa

ded

by [

Uni

vers

ity o

f Pe

rugi

a] a

t 00:

13 1

4 Ja

nuar

y 20

14

Accep

ted M

anus

cript

Venturella G, Altobelli E, Bernicchia A, Di Piazza S, Donnini D, Gargano ML, et al. 2011. Fungal biodiversity and in situ conservation in Italy. Plant Biosystems 145(4): 950-957.

Zervakis GI, Venturella G. 2007. Adverse effects of human activities on the diversity of macrofungi in forest ecosystems. Bocconea 21: 77-84.

Zotti M, Persiani AM, Ambrosio E, Vizzini A, Venturella G, Donnini D, Angelini P, Di Piazza S, Pavarino M, Lunghini D, Venanzoni R, Polemis E, Granito VM, Maggi O, Gargano ML, Zervakis GI. 2013. Macrofungi as ecosystem resources: conservation versus exploitation. Plant Biosyst 147: 219-225.

Dow

nloa

ded

by [

Uni

vers

ity o

f Pe

rugi

a] a

t 00:

13 1

4 Ja

nuar

y 20

14