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Basidiomycete diversity within Calabrian pine (Pinus brutia) ecosystems on the island of Cyprus

July 2021
Mycotaxon 136(2):543

July 2021
136(2):543

Authors:

Cyprus Mycological Association

Extended areas on the island of Cyprus including the vast majority of forested land, are occupied by Pinus brutia, an ecologically important tree forming ectomycorrhizal associations with a wide array of fungi. In this third installment of macromycete inventories from selected ecosystems in Cyprus, the basidiomycete diversity within P. brutia communities is reported following a 12-y-inventory. Two hundred and thirty-one taxa are compiled in an annotated checklist, 95 of which constitute new records for the country. Most notable of these are Clitopilus daamsii, Hygrophorocybe nivea, Lepiota lepida, Leucoagaricus georginae, Rhodocybe matesina, Russula werneri, Tephrocybe striaepilea, Tephroderma fuscopallens, and Tricholoma chrysophyllum, which are rarely reported in published literature. Notes on the altitude, phenology and estimated abundance are provided, accompanied by selected imagery and a review of previously published records.

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Basidiomycete diversity within Calabrian pine (Pinus brutia)

ecosystems on the island of Cyprus

ΜICHAEL LOIZIDES

P.O. BOX 58499, 3734 LIMASSOL, CYPRUS

CORRESPONDENCE: michael.loizides@yahoo.com

ABSTRACT—Extended areas on the island of Cyprus including the vast majority of forested land, are occupied by Pinus brutia, an

ecologically important tree forming ectomycorrhizal associations with a wide array of fungi. In this third installment of macromycete

inventories from selected ecosystems in Cyprus, the basidiomycete diversity within P. brutia communities is reported following a 12-y-

inventory. Two hundred and thirty-one taxa are compiled in an annotated checklist, 95 of which constitute new records for the country.

Most notable of these are Clitopilus daamsii, Hygrophorocybe nivea, Lepiota lepida, Leucoagaricus georginae, Rhodocybe matesina,

Russula werneri, Tephrocybe striaepilea, Tephroderma fuscopallens, and Tricholoma chrysophyllum, which are rarely reported in

published literature. Notes on the altitude, phenology and estimated abundance are provided, accompanied by selected imagery and a

review of previously published records.

KEY WORDS—biogeographical distribution, checklist, East-Mediterranean pine, fungi, inventory, island biogeography, Turkish pine

Introduction

Pinus brutia Ten., the Calabrian or East-Mediterranean pine, along with its sister-species the Aleppo pine (P.

halepensis Mill.), are ecologically important constituents of the Mediterranean flora jointly occupying some 7

million hectares (Quézel 2000). Both species have developed efficient regeneration mechanisms to deal with

wildfires in the region, and grow prolifically on dry and infertile substrates in the thermo-, meso- and supra-

Mediterranean belts (Nahal 1983; Thanos & Doussi 2000; Mauri et al. 2016). On the island of Cyprus,

Calabrian pine occupies some 175,000 hectares, accounting for ~90% of all woodland coverage on the country

(Meikle 1977; Tsintides et al. 2002). Although pure stands sometimes occur, the majority of P. brutia

populations are found in mixed ecosystems, with Kermes oaks [Quercus coccifera subsp. calliprinos (Webb)

Holmboe], strawberry trees (Arbutus andrachne L.), and rockroses (Cistus L. spp.) usually present in the

understorey. Throughout the meso- and supra-Mediterranean belts of the Troodos massif (400–1400 m. asl),

P. brutia forms extensive forests with the narrow-endemic golden oak (Quercus alnifolia Poech), while at

higher elevations it is replaced by P. nigra subsp. pallasiana (Lamb.) Holmboe, although both black- and

Calabrian pines can co-exist within a narrow phytogeographical belt from 1200 to 1400 m. asl.

Pinus brutia forms ectomycorrhizal (EcM) associations with a wide array of basidiomycetes and

ascomycetes. Such multifunctional mutualisms are fundamental for terrestrial ecosystems, enhancing plant

productivity, alleviating abiotic and oxidative stress, and regulating carbon, nitrogen and phosphorus cycling

(Smith & Read 2010; van der Heijden et al. 2015). The logs and branches of P. brutia moreover provide suitable

substrates for a number of cellulolytic decomposers (Bernicchia 2005; Bernicchia & Gorjón 2010; Loizides

2018), while several terrestrial saprotrophs are commonly encountered among the tree litter (Athanasiou &

Theohari 2001; Loizides et al. 2011; Polemis et al. 2012; Dimou et al. 2016). Although the physiology and

biogeography of P. brutia have been extensively studied (Panetsos 1981; Thanos & Marcou 1991; Petrou &

Milios 2012), little is known of the composition, structure and dynamics of its associated fungal diversity.

Several basidiomycetes have been reported from P. brutia ecosystems in genus-specific taxonomic treatments

and popular field guides from Cyprus, including a number of taxa described as new to science, such as

Clavulina iris Loizides et al. (Crous et al. 2019), Hydnellum dianthifolium Loizides et al. (Loizides et al. 2016),

Hygrophorus meridionalis Loizides et al. (Moreau et al. 2018), and Russula hobartiae Loizides & Vidal (Vidal

et al. 2019). However, comprehensive inventories from P. brutia ecosystems are lacking, resulting in a

fragmented understanding of the full range of macromycete diversity present in these important Mediterranean

habitats. The basidiomycete diversity in Calabrian pine ecosystems is extensively documented in this paper,

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Loizides, M.

following a 12-y-inventory carried out in Cyprus between 2007 and 2019. A total of 231 taxa are overall

compiled in an annotated checklist, 95 of which are reported from Cyprus for the first time. Some interesting

finds and previously published records are discussed, and imagery, accompanied by brief taxonomic notes for

critical taxa, are included.

Materials & Methods

Material was collected as part of a general inventory in Cyprus carried out by the author from 2007 to 2019. A large number

of sites across an extended altitudinal gradient were systematically surveyed following a sampling protocol described in

Loizides (2016, 2018), and Loizides et al. (2019a, 2019b). Collections were photographed in situ and details on the

macromorphological aspect, including size, colour, odour and oxidation of the flesh, were observed on freshly collected

specimens. Iron sulfate (FeSO4), Guiac tincture and Phenol were applied to observe chemical reactions on the flesh of

species of the genus Russula Pers. Potassium hydroxide (5% KOH) was applied to observe chemical reactions on the pileus

and flesh of Cortinarius (Pers.) Gray species. Microscopic studies were performed under a LEICA BM E binocular and an

AM T360B trinocular microscopes, at ×40, ×100, ×400 and ×1000 magnifications. Congo Red in 10% ammonia (NH3),

5% potassium hydroxide (KOH), lactophenol cotton blue (LPCB), aqueous Phloxine, Melzer’s solution and tap water were

used accordingly as mounting mediums and diagnostically important microscopic structures were observed. The checklist

is arranged in alphabetical order and includes previously reported taxa, as well as species newly identified in the course of

this survey marked below with an asterisk (*). Doubtful records, or species identified only to genus are excluded (see also

“Discussion”). Wood-inhabiting aphyllophoraceous basidiomycetes and ascomycetes are included in separate dedicated

checklists (Loizides 2018 & in prep.). Unless otherwise noted, specimens were collected, photographed and identified by

the author. Index Fungorum (http://www.indexfungorum.org), Mycobank (http://www.mycobank.org), and recently

published systematic revisions were consulted to provide most recently updated names for the cited taxa. Exsiccatae are

kept in the private collection of the author, with selected collections deposited at ACAM (Agricultural University of

Athens), BCN (Universitat de Barcelona Herbari), CEFE (Centre d’Ecologie Fonctionnelle et Evolutive, Montpellier), K

(Royal Botanic Gardens at Kew), and LIP (Université de Lille).

Species recorded

Agaricus augustus Fr.

Occasionally encountered in black-and Calabrian pine forests, with one collection from a mixed Pinus brutia/Quercus

alnifolia/Arbutus andrachne forest in Amíantos, 21-X-2011, ca 1300 m asl.

Agaricus bisporus (J.E. Lange) Imbach

Common in pastures and along Cupressus sempervirens L. groves, but once collected in a coastal P. brutia forest in

Akrotíri, 11-II-2014, ca 20 m asl.

Agaricus brunneolus (J.E. Lange) Pilát

This bitter-almond scented wood dweller is frequently seen at various elevations, e.g., Saittás ,9-XII-2007 & 6-I-2017,

ca 700 m asl; Platánia, 27-XI-2008, 1-XII-2009 & 23-XII-2018, ca 1200 m asl; Lysós, 18-I-2009, ca 600 m a.s.l; Mávro

Dásos, 5-XII-2012, ca 1200 m asl; Troodítissa, 17-XI-2017, ca 1400 m asl. Also reported by Viney (2005) with its old

name “A. porphyrrhizon P.D. Orton”, from the pine woods of Pentadáktylos, Dec. 2001.

Agaricus impudicus (Rea) Pilát

Probably the most common representative of the genus in Cyprus, with numerous sightings in all kinds of forests,

including Calabrian pine forests in Amíantos, 19-X-2008 & 10-X-2009, ca 1300 m asl; Troodítissa, 13-X-2008, 26-X-

2011, 31-X-2011 & 9-XI-2018, ca 1400 m asl; Plátres, 17-XI-2008, ca 1000 m asl; Saittás, 25-XI-2009 & 10-I-2017,

ca 700 m asl; Kannavioú, 29-XI-2017, ca 800 m asl.

*Agaricus litoralis (Wakef. & A. Pearson) Pilát

Three collections in different seasons, all from the same locality in a P. brutia forest at Agros, 4-IV-2007, 8-IV-2008

& 24-III-2011, ca 1000 m asl. Rare.

Agaricus macrocarpus F.H. Møller

Reported by Loizides et al. (2011) based on a collection from a mixed P. brutia/Q. alnifolia forest in Plátres, 29-XII-

2009, ca 1200 m asl, but several new species have been described in section Arvenses since (Parra Sánchez 2013),

whose morphological identification is far from straight forward. Several species of this group seem to occur in Cyprus,

including some not comfortably fitting available descriptions, that ultimately need to be resolved by DNA sequencing.

A collection from Syrianóchori, Jan. 2001, was also reported as “A. aff. macrocarpus” by Viney (2005).

Agaricus moelleri Wasser

Occasionally seen in mixed P. brutia forests (Loizides et al. 2011): e.g., Amíantos, 19-X-2008, ca 1300 m asl; Saittás,

Basidiomycetes within Pinus brutia ecosystems in Cyprus

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4-XII-2008 & 30-XI-2016, ca 700 m asl; or Platánia, 10-X-2009 & 7-XI-2009, ca 1100 m asl.

*Agaricus pampeanus Speg.

One collection fitting the description of this sparingly reported species, from a P. brutia/Quercus alnifolia forest

clearing at Mésa Potamós, 12-XII-2011, ca 800 m asl. Rare.

Agaricus pseudopratensis (Bohus) Wasser

Common in the lowlands under C. sempervirens and open fields, but rarely also seen in Cistus matorral (Loizides 2016)

and forest clearings: Saittás, 30-XI-2016, ca 650 m asl; Stavrós tis Psókas, 29-XI-2017, ca 1200 m asl.

*Agaricus sylvaticus Schaeff.

Rarely seen in riparian broadleaved forests and once in a Calabrian pine forest in Péra Pedí, 10-I-2017, ca 55 asl.

Agaricus sylvicola (Vittad.) Peck sensu lato

Occasionally encountered in black pine and broadleaved riparian forests (Loizides et al. 2011), with a couple of

collections from a Calabrian pine forest in Trimíklini, 23-XII-2007 & 17-XII-2011, ca 650 m asl.

Agaricus xanthodermus Genev.

The “yellow stainer” is rare in Cyprus (Loizides et al. 2011), with a couple of records from mixed P. brutia forest

clearings: Platánia, 16-XI-2014, ca 1100 m asl; Kannavioú, 29-XII-2016, ca 800 m asl.

Amanita ovoidea (Bull.) Link

Probably the most widespread and ecologically versatile Amanita Pers. species on the island, occurring from near-sea

level to ~1700 m asl in association with Pinus L., Quercus L., and occasionally Eucalyptus L’Hér. First reported on the

island by Nattrass (1937) from Ktíma (Paphos), Dec. 1932; and later also by Viney (2005) from Kaló Chorió, Nov.

1999, Loizides (2011), and Loizides et al. (2011, 2018) from Trimíklini, 13-X-2009, 10-XI-2012 & 14-XI-2012, ca

650 m asl; Kouká, 15-X-2009, ca 500 m asl; Karvounás, 16-X-2011 & 23-X-2011, ca 1200 m asl; Plátres, 19-XI-2011,

ca 1200 m asl; Platánia, 16-XI-2014, ca 600 m asl; and Akrotíri, 24-XI-2014 & 10-XII-2017, ca 20 m asl. Sometimes

collected for the table by the locals, which has led to a number of poisonings in the past likely due to confusion with A.

proxima (Loizides et al. 2018).

Amanita proxima Dumée

Several collections from Trimíklini, 30-X-2008, 10-XI-2012 & 1-XII-2017, ca 650 m asl; Kouka, 15-X-2009, ca 500

m asl; Karvounás, 13-X-2011, 16-X-2011 & 3-XI-2015, ca 1200 m asl; Kélefos, 31-X-2012, ca 500 m asl; and Akrotíri,

24-XI-2014 & 10-XII-2017, ca 20 m asl. Previously reported by Viney (2005) from Haléfka, Nov. 1997, and by

Loizides (2008), and Loizides et al. (2011 & 2018) from collections cited above. This fungus contains allenic

norleucine, a potentially deadly nephrotoxin (De Haro et al. 1998) and can be confused with A. ovoidea, with which it

shares the same habitat and distribution. Most likely the unidentified “white Amanita” species briefly discussed and

sketched by Willimott (1933), responsible for a number of fatalities early last century.

Arrhenia rickenii (Hora) Watling

Frequently seen on coastal and inland dunes (Loizides 2016), but also in mossy P. brutia forests in the thermo-

Mediterranean belt: Mosfilotí, 23-I-2011, ca 250 m asl; Politikó, 9-I-2012, ca 400 m asl.

Arrhenia spathulata (Fr.) Redhead

Sparingly encountered in matorral and mossy forests in the dunal and thermo-Mediterranean belts, such as Trimíklini,

26-XI-2008, 9-XII-2011 & 31-XII-2013, ca 650 m asl.

*Atheniella flavoalba (Fr.) Redhead, Moncalvo, Vilgalys, Desjardin & B.A. Perry

One collection matching the description of this taxon, previously placed in genus Mycena (Pers.) Roussel. Found among

P. brutia litter in Kélefos, 2-I-2009, ca 500 m asl. Distribution unclear.

*Atractosporocybe inornata (Sowerby) P. Alvarado, G. Moreno & Vizzini

Previously placed in the genus Clitocybe (Fr.) Staude, this species was recently transferred to the new genus

Atractosporocybe P. Alvarado et al. (Alvarado et al. 2015). Fairly widespread in upland P. nigra subsp. pallasiana

forests and Cedrus brevifolia A. Henry ex Elwes & A. Henry plantations, but occasionally also encountered in mixed

P. brutia/Q. alnifolia forests in Plátres, 17-XII-2007, ca 900 m asl; Álona, 5-XI-2009, ca 1300 m asl.; Mávro Dásos,

1-XII-2012, ca 1200 m asl; and Platánia, 30-XII-2014, ca 1100 m asl.

*Baeospora myosura (Fr.) Singer

Sparingly seen on the island, this species inhabits fallen cones of conifers, particularly of P. brutia. Collected in

Amíantos ,9-XI-2009, ca 1300 m asl; Kélefos, 29-XI-2009, ca 500 m asl; and Platánia, 21-XII-2011, ca 1100 asl.

*Bovista aestivalis (Bonord.) Demoulin

A handful of collections, one from a P. brutia forest in Moniátis, 29-IX-2011, ca 750 m asl. Uncommon.

Chalciporus amarellus (Quél.) Bataille

Previously reported by Viney (2005) from Haléfka, Jan. 1998, this striking bolete is known only from three additional

collections in Saittás, 17-XI-2007, ca 700 m asl; Platánia, 26-XI-2009, ca 1100 m asl; and Kélefos, 28-XI-2009, ca 450

m asl (Loizides et al. 2019a). Although all collections originate from P. brutia forests, this species is probably a

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Loizides, M.

mycoparasite (Tedersoo et al. 2010; Nuhn et al. 2013).

Chroogomphus mediterraneus (Finschow) Vila, Pérez-De-Greg. & G. Mir

One of the most widespread and consistently occurring fungi in Cyprus, present from the dunal zone to the peaks of

the Troodos massif. Numerous sightings, including Péra Pedí, 21-XI-2007, ca 700 m asl; Pissouri, 25-I-2009, ca 200

m asl; Kélefos, 20-XI-2009, 28-XI-2009, 28-I-2010 & 2-XII-2014, ca 500 m asl; Kannavioú, 15-XII-2009, ca 800 m

asl; Mosfilotí, 23-I-2011, ca 250 m asl; Akámas, 18-XII-2014, ca 150 m asl; Akrotíri, 12-I-2015 & 10-II-2017, ca 20

m asl. Collections reported by Viney (2005) and Loizides et al. (2011) as “C. rutilus (Schaeff.) O.K. Mill.”, all seem to

belong to this widespread species (see Scambler et al. 2018 and “Discussion” for further remarks).

*Clathrus ruber P. Micheli ex Pers.

The “latticed stinkhorn” is a rare species in Cyprus, known only from four collections in Saittás, 13-XII-2007, ca 700

m asl; Plátres, 17-X-2008 & 11-XII-2012, ca 900 m asl; and Kélefos, 1-I-2019, ca 500 m asl, all in mixed habitats.

Clavulina coralloides (L.) J. Schröt. sensu lato

Reported by Viney (2005) as “C. cristata”, from Haléfka, Jan. 1998, and later by Loizides et al. (2011) from collections

in Platánia, 5-XII-2009, ca 1150 m asl, and Loizides (2016) from collections under Cistus, but DNA sequencing

suggests that more than one species might be involved (unpubl. data).

Clavulina iris Loizides, Bellanger & P.-A. Moreau

Numerous collections of this recently described and widespread species, appearing from mid-winter to early spring in

a variety of calcareous habitats in association with Cistus, P. brutia and Quercus, e.g. Péra Pedí, 24-III-2009 & 2-III-

2019, ca 550 m asl; Soúni, 16-III-2012, 25-II-2013, 21-II-2015, 2-III-2015, 4-III-2019 & 6-III-2019, ca 400 m asl;

Anogyra, 17-II-2015, ca 400 m asl; Platánia, 26-XII-2018, ca 1100 m asl; Kélefos, 1-I-2019, ca 500 m asl; and

Trimíklini, 28-II-2019, ca 650 m asl (for sequenced collections and a detailed description see Crous et al. 2019).

*Clitocella fallax (Quél.) Kluting, T.J. Baroni & Bergemann

Two collections from mixed forests in Cedar Valley, 18-XI-2009, ca 1100 m asl; and Kélefos, 20-I-2011, ca 450 m asl,

matching the descriptions of this rarely reported species but featuring rather variable spores, ranging from 5–7 × 3.5–

4.5 μm to 6–8 × 4–5 μm. In the phylogeny provided by Kluting et al. (2014), European collections from Norway and

Spain form a well-supported clade distinct from a North American collection identified as this taxon, indicating the

presence of more than one species under this name. European collections likely correspond to C. fallax ss. str.

Clitocella popinalis (Fr.) Kluting, T.J. Baroni & Bergemann sensu lato

Reported by Viney (2005) as “Rhodocybe popinalis” from pine forests of Haléfka, Jan. 2001, and subsequently

collected by the author in Kakomállis, 29-X-2009, ca 750 m asl; and Palechori, 2-XI-2009, ca 900 m asl. At present,

collections identified as C. popinalis and the closely related C. mundula (Lasch) Kluting et al., seem to nest in multiple

clades in published phylogenies (Kluting et al. 2014; Vizzini et al. 2016 & 2018), therefore deeper phylogenetic

investigations are needed to clarify their precise taxonomic status.

Clitocybe diatreta (Fr.) P. Kumm.

Reported by Viney (2005) from the pine woods of Áyios Amvrósios, Nov. 2000, and by Loizides et al. (2011) from a

P. brutia/Quercus coccifera subsp. calliprinos forest clearing in Kélefos, 24-XII-2009, ca 500 m asl.

*Clitocybe fragrans (With.) P. Kumm

One of several Clitocybe species with an aniseed odour, with a couple of collections from Kélefos, 2-XII-2008 & 2-I-

2009, ca 500 m asl. More frequently seen in high elevation P. nigra subsp. pallasiana forests.

*Clitocybe lituus (Fr.) Métrod

A couple of collections matching the description of this taxon from Álona, 5-XI-2009, ca 1300 m asl; and Saittás, 18-

XII-2009, ca 650 m asl. Uncommon or rare.

*Clitocybe metachroa (Fr.) P. Kumm.

Few collections from mixed P. brutia and Q. alnifolia forests in Mesa Potamos 2-XII-2007, ca 800 m asl; Platania 22-

XII-2007 & 28-XI-2008, ca 1100 m asl; and Mavro Dasos 1-XII-2012, ca 1200 m asl. Probably common, but a

remarkably variable taxon frequently confused with similar-looking species.

Clitocybe obsoleta (Batsch) Quél.

One collection matching the description of this species from a P. brutia forest in Péra Pedí, 10-I-2009, ca 550 m asl.

Also reported by Viney (2005) from collections under Cistus in Karavas, Dec. 2001, but these may belong to C.

cistophila Bon & Contu, instead (Loizides 2016). A number of aniseed-scented Clitocybe species seem to occur in

Cyprus, whose identity is not clear since several early-described taxa have yet to be phylogenetically defined.

Clitocybe odora (Bull.) P. Kumm.

Frequent under both broadleaved and coniferous trees, with several sightings from Calabrian pine ecosystems: Saittás,

9-XII-2007, ca 650 m asl; Péra Pedí, 13-X-2008, ca 650 m asl; Kannavioú, 15-XII-2009, ca 900 m asl; Kelláki, 21-

XII-2009, ca 600 m asl; Kélefos, 8-II-2011, 29-XI-2014, 10-XI-2017 & 1-I-2019, ca 500 m asl.

Clitocybe umbilicata P. Kumm.

Reported by Viney (2005) from Panagra, Dec. 2001.

Basidiomycetes within Pinus brutia ecosystems in Cyprus

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Clitopaxillus alexandri (Gillet) G. Moreno, Vizzini, Consiglio & P. Alvarado

A distinctive species recently transferred to the new genus Clitopaxillus G. Moreno et al. (Alvarado et al. 2018a),

appearing gregariously in both Calabrian and black pine ecosystems sometimes forming fairy rings: e.g., Trimíklini, 1-

XII-2007, 9-XII-2011, ca 650 m asl; Moniátis, 13-XII-2007, 800 m asl; Mésa Potamós, 18-XII-2007, ca 800 m asl;

Kélefos, 28-XI-2009, ca 500 m asl; Platánia, 26-XII-2009, ca 1100 m asl; Péra Pedí, 14-I-2017, ca 550 m asl. Also

reported by Viney (2005) from Áyios Amvrósios, Dec. 1998, and Loizides et al. (2011) from collections cited above.

*Clitopilus daamsii Noordel.

One collection fruiting on a fallen P. brutia log in Amíantos, 6-I-2014, ca 1100 m asl, matching the description of this

rarely reported taxon. This species is morphologically close to C. hobsoni (Berk.) P.D. Orton, from which it differs by

larger spores, reported as (7–)8–11.5(–12.5) × 4.8–6.6(–7) μm in the type collection (Noordeloos 1984), and measuring

(7.5–)8–10.5(–11.5) × (4–)4.5–6(–7) μm in the Cypriot specimens.

Clitopilus geminus (Paulet) Noordel. & Co-David sensu lato

This widely reported taxon was previously placed in Rhodocybe Maire, but transferred to genus Clitopilus (Fr. ex

Rabenh.) P. Kumm., based on phylogenetic inferences (Co-David et al. 2009). Reported in Cyprus by Viney (2005) as

“Rhodocybe gemina”, from pine woods in Panagra, Dec. 2001, and later confirmed by the author from Calabrian pine

woods in Platánia, 20-XI-2008 & 25-XI-2009, ca 1100 m asl, and Akámas, 14-XII-2014, 200 m asl. A couple of variants

seem to occur in Cyprus, whose exact phylogenetic and taxonomic status are not yet clear. Collections from Cistus

matorral are generally smaller, mild scented and feature white mycelial threads, matching the concept of C. geminus

var. subvermicularis (Maire) Noordel. & Co-David, originally described from Mauretania (Maire 1924).

Conocybe albipes Hauskn.

Perhaps better known as “C. apala (Fr.) Arnolds”, with one collection from Pissouri, 25-I-2009, ca 200 m asl, among

manured grass, in a P. brutia forest opening. First reported by Viney (2005) as “Bolbitius tener”, fruiting in manure-

rich grass in Kalni Dec. 1998.

*Conocybe semiglobata Kühner & Watling

One collection from Calabrian pine woods in Péra Pedí, 2-III-2014, ca 550 m asl. Distribution unclear.

*Coprinellus disseminatus (Pers.) J.E. Lange

A handful of records from mixed forests in Plátres, 20-X-2007, ca 900 m asl; Moniátis, 20-XII-2010 & 14-X-2012, ca

800 m asl; and Platánia, 25-XI-2013, ca 1100 m asl. Uncommon.

Coprinellus domesticus (Bolton) Vilgalys, Hopple & Jacq. Johnson

Just a couple of collections, one from a burned P. brutia forest near Koúrdali, 15-III-2017, ca 800 m asl. Previously

reported by Viney (2005) from Karmi, Nov. 1997, and Loizides et al. (2011). Apparently rare.

Coprinopsis cortinata (J.E. Lange) Gminder

Two collections, one among P. brutia/Cistus litter in Kelláki, 15-III-2012, ca 650 m asl. Also seen in Cistus debris

(Loizides 2016). Apparently rare, but can be easily overlooked.

Coprinus comatus (O.F. Müll.) Pers.

Only sporadically seen in Cyprus, mostly in high elevation P. nigra subsp. pallasiana forest clearings and along forest

paths, with a couple of sightings of from a P. brutia, Platanus orientalis L. and Q. infectoria subsp. veneris forest in

Plátres, 20-X-2007 and 17-X-2008, ca 900 m asl. Also reported by Viney (2005), and Loizides et al. (2011).

Coprinus ovatus (Schaeff.) Fr.

Sparingly seen in late winter and spring, with one record from a mixed forest clearing at Platánia, 27-V-2018, ca 1100

m asl. Also reported by Viney (2005) at Mándres, Feb. 2001. Uncommon.

Coprinus spadiceisporus Bogart

Occasionally encountered in and out of forests, with a couple of collections from open P. brutia forests in Pissouri, 30-

I-2011, ca 200 m asl; and Soúni, 26-III-2012, ca 400 m asl. Also reported by Loizides et al. (2011) from thermo-

Mediterranean matorral at Konia, 12-XII-2009.

*Cortinarius anthracinus Fr.

One collection from Mávro Dásos, 5.XII-2012, ca 1100 m asl. Distribution not yet understood.

*Cortinarius decipiens (Pers.) Fr.

One collection from Mávro Dásos, 1-XII-2012, ca 1100 m asl, matching the description of C. atrocoeruleus M.M.

Moser ex M.M. Moser (1967). However, in a revision of sections Hydrocybe and Fraternii, sequenced collections

identified as “C. atrocoeruleus”, including the type collection, nested in the same clade as C. decipiens (Pers.) Fr.

(Suárez-Santiago et al. 2009). Until the situation is further clarified, the latter binomial is here tentatively adopted as a

prioritary synonym of the former.

Crinipellis scabella (Alb. & Schwein.) Murrill

A couple of collections on fallen twigs and herbal residue, one in a mossy P. brutia forest in Kélefos, 31-XII-2008, ca

500 m asl. Also reported by Viney (2005) from Geunyeli, Nov. 1997. Rare.

…

Loizides, M.

Cuphophyllus virgineus (Wulfen) Kovalenko

Reported by Viney (2005) from pine forests in Áyios Amvrósios, without date.

*Cystoderma carcharias (Pers.) Fayod

Frequently encountered in Calabrian and black pine forests with collections from Kannavioú, 15.XII-2009, ca 800 m

asl; Mávro Dásos 1-XII-2012, ca 1100 m asl; Plátres 1-XII-2017, ca 1200 m asl.

*Cystoderma fallax A.H. Sm. & Singer

Numerous sightings in both Calabrian and black pine forests, e.g., Plátres, 23-XII-2009, 26-XII-2009 & 19-I-2015, ca

1400 m asl; Platánia, 7-XII-2009, 9-XII-2012 & 2-I-2015, ca 1100 m asl; Mésa Potamós, 2-XII-2017, ca 750 m asl;

Karvounás, 3-XI-2015 & 14-XI-2018, ca 1300 m asl. Collections from Haléfka (Jan. 1998) reported by Viney (2005)

as “Cystoderma amianthinum”, probably also belong to this widespread species.

Cystodermella cinnabarina (Alb. & Schwein.) Harmaja

A couple of collections from Platánia, 2-XII-2008 & 9-XII-2009, ca 1100 m asl. Also reported by Viney (2005), as

“Cystoderma terreyi”, from Haléfka and Syrianóchori, Dec. 1998. Occassional or uncommon.

*Cystodermella granulosa (Batsch) Harmaja

Occasionally seen Calabrian pine forests, e.g., Moniátis, 13-XII-2007, ca 700 m asl; Péra Pedí, 26-XII-2007, 6-I-2011

& 10-I-2017, ca 550 m asl; Plátres, 1-XII-2017, ca 1200 m asl.

Cystolepiota cystophora (Malençon) Bon

One collection from a mossy Calabrian pine forest at Platánia, 27-XI-2009, ca 1100 m asl. Also reported by Viney

(2005) from Haléfka, Jan. 1999. Distribution not yet understood.

*Cystolepiota seminuda (Lasch) Bon

Just one collection from Platánia, 27-XI-2009, ca 1100 m asl. Apparently rare.

*Entoloma atromadidum A.M. Ainsw. & B. Douglas

Two collections from a mixed P. brutia/Q. alnifolia forest at Kannavioú, 13-XII-2009 & 15-XII-2009, ca 800 m asl,

previously reported as “E. bloxamii (Berk. & Broome) Sacc.” (Loizides 2011). Cypriot collections are more likely to

represent the recently described E. atromadidum (Ainsworth et al. 2018), which produces darker basidiomes with

somewhat smaller spores than the epitype of E. bloxamii as reported in Morgado et al. (2013). Rare.

Entoloma cinereo-opacum (Noordel.) Vila, Català & Noordel.s

Sparingly encountered in mossy forest clearings and matorral, such as Moniátis, 9-XII-2007, ca 800 m asl; and Kélefos,

9-XII-2015, ca 500 m asl. Cypriot collections were previously reported as “E. sericeum var. cinereo-opacum Noordel.”

in Loizides (2016). However, as shown by Vila et al. (2013), collections previously ascribed to this variant are

phylogenetically distinct and have since been elevated to species rank.

*Entoloma fuscohebes Vila, J. Carbó & F. Caball.

One collection matching the description of this seldomly reported species from Kélefos, 23-XII-2011, ca 500 m asl.

Distribition not yet understood.

Entoloma hirtipes (Schumach.) M.M. Moser sensu lato

A couple of collections keying out as this highly variable taxon, likely to represent a species-complex: Lysós, 18-1-

2009, ca 600 m a.s.l; Kélefos, 8-II-2011, ca 500 m asl. Collections from the pine woods of Haléfka were also reported

by Viney (2005) as “Entoloma aff. hirtipes”, without dates.

*Entoloma incanum (Fr.) Hesler

Two collections from Plátres, 16-X-2012, ca 1250 m asl; and Platánia, 19-VI-2018, ca 1100 m asl. Rare.

Entoloma jubatum (Fr.) P. Karst.

Reported by Viney (2005) from pine woods near Halefka, Jan. 1998, but not verified by the author.

*Entoloma mougeotii (Fr.) Hesler

Two collections matching the description of this taxon, one from a mixed P. brutia/P. orientalis/Q. infectoria subsp.

veneris forest in Plátres, 21-XII-2012, ca 900 m asl. Rare.

*Entoloma pleopodium (Bull.) Noordel.

Three collections of this scented species, two of them from mixed Calabrian pine forests in Plátres, 18-IV-2011 & 1-

V-2017, ca 900 m asl. Rare.

*Entoloma rusticoides (Gillet) Noordel.

One collection in Pissouri, 25-I-2009, ca 200 m asl, matching the description of this taxon. Distribution unclear.

Faerberia carbonaria (Alb. & Schwein.) Pouzar

Reported by Viney (2005) near Áyios Ilaríon, Dec. 2000, but not seen during this 12-y inventory.

*Galerina badipes (Pers.) Kühner

One identified collection from Platánia, 30-XII-2011, ca 1100 m asl. Perhaps widespread but overlooked, as large

numbers of similar-looking Galerina Earle species occur in Cyprus late in the season.

Basidiomycetes within Pinus brutia ecosystems in Cyprus

…

Galerina marginata (Batsch) Kühner

This deadly poisonous fungus inhabits decomposing logs of P. brutia in the thermo- and supra-Mediterranean belts,

but sometimes also found among wooden debris on the ground: Plátres, 24-XI-2008, 6-XI-2011 & 10-XI-2012, ca 800

m asl; Kélefos, 2-XI-2009, ca 500 m asl; Platánia, 10-I-2014 & 2-I-2015, ca 1150 asl; Plátres, 15-XII-2018, 1300 m

asl. Also reported by Loizides et al. (2011), and by Viney (2005) north of Haléfka, Jan. 2002. Occasional.

Galerina vittiformis (Fr.) Singer

Reported by Viney (2005) as “G. vittaeformis”, under pine and Cistus, Dec. 1999, without locality. Yet to be verified.

*Gamundia leucophylla (Gillet) H.E. Bigelow

A couple of collections from P. brutia stands in Kelláki, 25-I-2011, ca 600 m asl; and Politikó, 9-I-2012, ca 400 m asl.

Distribution not yet understood.

Geastrum berkeleyi Massee

Reported by Viney (2005) among pine litter at Sirianochóri, Dec. 2001, and by Loizides (2012) from Prastió, 5-III-

2011, ca 550 m asl; and Péra Pedí, 25-III-2011, ca 530 m asl. Probably uncommon.

Geastrum campestre Morgan

Known from a handful of collections under Cistus (Loizides 2012 & 2016), and two collections under P. brutia from

Platánia, 3-XII-2009 & 16-XI-2014, ca 1100 m asl. Occasional.

Geastrum fimbriatum Fr.

Frequently seen in mixed forests, such as Platánia, 22-XII-2007, 7-XII-2009 & 20-XI-2011, ca 1100 m asl; or Plátres,

18-XII-2010, ca 1250 asl. Also reported by Viney (2005) under pines between Áyios Amvrósios and Sirianochóri,

without date, and by Loizides (2012) from collections cited above. Widespread.

Geastrum elegans Vittad.

Sparingly encountered in Cistus matorral (Loizides 2012 & 2016) and open P. brutia forests, such as Kélefos, 21-I-

2010 & 8-I-2011, ca 500 m asl.

*Geastrum michelianum (Sacc.) W.G. Sm.

Previously reported as “G. triplex” by Viney (2005), from pine woods north of Haléfka, Jan. 1999; and by Loizides

(2012) from P. brutia woods in Trimíklini, 24-III-2009, ca 650 m asl; and Prastió, 25-II-2011, ca 600 m asl. Genetic

studies revealed that “collared” collections previously identified as “G. triplex” constitute a species complex, with

European collections corresponding to G. michelianum instead (Kasuya et al. 2012; Zamora et al. 2014). Uncommon.

*Gymnopus androsaceus (L.) Della Magg. & Trassin.

Sparingly seen in black- and Calabrian pine forests, with a couple collections from Moniátis, 23-XI-2007, in riparian

P. brutia, A. orientalis, P. orientalis and Q. alnifolia forest, ca 900 m asl; and Episkopí, 26-I-2018, in an open P.

brutia/C. sempervirens/J. phoenicaea forest, ca 150 m asl. Might be widespread but overlooked.

Gymnopus brassicolens (Romagn.) Antonín & Noordel.

A number of morphologically diverse collections from various habitats currently key out as this taxon, including

collections from mixed Calabrian pine forests in Saittás, 9-XII-2007 & 6-I-2017, ca 600 m asl; Stavrós tis Psókas, 25-

X-2008, ca 1200 m asl; and Kakomállis, 2-XI-2009, ca 900 m asl. Also reported by Viney (2005), as “Micromphale

brassicolens”, from the pine woods of Haléfka, Nov. 2000. More than one species might be involved, but available

sequences of this widely reported taxon are surprisingly scant in public databases.

Gymnopus dryophilus (Bull.) Murrill

A common and widespread species, with collections e.g., from Trimiklini 9-XII-2007, ca 650 m asl; Saittas 1-X-2009,

30-XII-2010 & 29-IX-2011; Kakomallis 31-X-2009, ca 800 m asl; Pera Pedi 10-I-2017, ca 550 m asl. Reported by

Viney (2005), as “Collybia dryophila”, without date and locality.

*Gymnopus hybridus (Kühner & Romagn.) Antonín & Noordel.

Not often reported in literature but frequently seen in Cyprus, mostly in mixed P. brutia/Q. alnifolia forests in Plátres,

10-XII-2009 & 11-XII-2012, ca 1200 m asl; Cedar Valley, 27-XII-2010 & 22-XI-2011, ca 1200 m asl; or Platánia, 20-

XI-2011 & 16-XI-2014, ca 1100 m asl.

Hebeloma crustuliniforme (Bull.) Quél.

One of several Hebeloma (Fr.) P. Kumm. species occurring on the island, frequently encountered in all kinds of forests,

e.g., Mávro Dásos, 1-XII-2012, ca 1200 m asl. See Beker et al. (2016) for sequenced collections.

Hebeloma cylindrosporum Romagn.

Known from just one collection from Platánia 3-XII-2012, ca 1100 asl (Beker et al. 2016). Rare.

Hebeloma sinapizans (Paulet) Gillet

Perhaps the most widespread Hebeloma species on the island, seen in large numbers in P. nigra subsp. pallasiana

forests, but also in mixed P. brutia/Q. alnifolia forests: e.g., Kyperoúnda, 3-XII-2012, ca 1200 m asl; Platánia, 9-XII-

2012, 1100 m asl; Mávro Dásos, 20-XI-2015, ca 1200 m asl (for further collection details and a full description see

Beker et al. 2016).

…

Loizides, M.

Hebeloma subtortum P. Karst.

Perhaps better known as “H. sordidum Maire”, this ecologically versatile fungus was reported by Beker et al. (2016)

and further collected by the author in mixed P. brutia forests in Platánia, 25-XI-2013, ca 1100 m asl; and Plátres, 3-

XII-2017, ca 1200 m asl. Occasional.

*Hemimycena lactea (Pers.) Singer

Few collections, e.g., Platánia, 25-XI-2009 & 20-XI-2011, ca 1100 m asl; Amíantos, 2-X-2009, ca 1300 m asl. Probably

widespread but overlooked.

*Hemimycena pseudogracilis (Kühner & Maire) Singer

Two collections from Kélefos, 8-I-2011, ca 450 m asl; and Platánia, 25-X-2012, ca 1100 m asl key out as this poorly

known and variously interpreted taxon, although neither the macro- nor micromorphological features are a perfect fit.

Cypriot collections might represent an undescribed Mediterranean lookalike.

Hydnellum dianthifolium Loizides, Arnolds & P.-A. Moreau

A new collection and locality of this very rare species, seen in Kélefos, 5-III-2019, ca 500 m asl, leg. C. Hobart.

Previously known only from three collections in Cyprus and one collection in southern Italy (Puglia). See Loizides et

al. (2016) for a detailed description and previous collections’ details.

Hydnellum ferrugineum (Fr.) P. Karst. sensu lato

Common in P. nigra subsp. pallasiana forests in years with early rainfall, but sometimes also seen in mixed P. brutia/Q.

alnifolia forests, e.g., Karvounás, 13-X-2011, ca 1300 m asl; and once documented in a P. brutia/Q. coccifera subsp.

calliprinos forest in Kélefos, 2-XI-2012, ca 500 m asl (Loizides 2011). Multiple phylogenetic lineages are currently

identified as this taxon in public databases (Loizides et al. 2016; Larsson et al. 2019), which are currently the object of

a dedicated study (Loizides et al. in prep.).

*Hygrocybe calciphila Arnolds

One collection from a mossy P. brutia forest clearing at Kélefos, 23-XII-2012, ca 450 m asl. A rare species, proposed

for conservation in the Global Fungal Red List Initiative.

Hygrocybe conica (Schaeff.) P. Kumm. sensu lato

This widely applied binomial currently lumps multiple phylospecies yet to be taxonomically resolved. Several species

of this complex are apparently present in Cyprus, one of them regularly occurring in black- and Calabrian pine forests

e.g., in Platánia, 2-XII-2008 & 23-XII-2018, ca 1150 m asl; or Madarí, 30-XI-2015, ca 1000 m asl.

*Hygrocybe konradii R. Haller Aar.

One collection matching the concept of this species from a mixed forest in Kannavioú, 13-XII-2009, ca 900 m asl.

*Hygrophorocybe nivea (Velen.) Vizzini & Contu

A couple of collections from P. brutia/Q. alnifolia forests at Platánia, 27-XI-2009 & 16-XI-2012, ca 1100 m asl. The

monotypic genus Hygrophorocybe Vizzini & Contu was proposed to accommodate Clitocybe nivea Velen. (Vizzini

2014), a striking species with ellipsoid to sublacrymoid spores and hygrophoroid basidia. Spores measure 8–10 × 4.5–

5.5 μm in the Cypriot collections. Rare, known only from one locality.

Hygrophorus meridionalis Loizides, P.-A. Moreau, Athanassiou & Athanasiades

Numerous sightings of this common Mediterranean species, described as new to science in 2018. See Moreau et al.

(2018) for a detailed description and extensive collection details.

Infundibulicybe costata (Kühner & Romagn.) Harmaj

A couple of collections from Calabrian pine forests in Kélefos, 31-XII-2008 & 29-XI-2014, ca 500 m asl. Previously

reported by Viney (2005), as “Clitocybe costata”, under pines between Haléfka and Syrianóchori, Dec. 2000; and by

Loizides et al. (2011, also as “C. costata”) from collections cited above. Uncommon.

Infundibulicybe geotropa (Bull.) Harmaja

Occasionally appearing late in the autumn and winter in wet riparian forests and mixed forest clearings, with few

collections from Plátres, e.g., 9-XII-2007 & 21-XI-2012, ca 900 m asl; and Platánia, 20-XI-2007, 3-XII-2009 & 30-

XII-2011, ca 1100 m asl. Infundibulicybe gigas (Harmaja) Harmaja, a taxon described from Finland (as “Clitocybe

gigas”, Harmaja 1978), is reported to have a deeply infundibuliform pileus, denser lamellae, a squat stipe and

±lacrymoid spores, and does not appear to occur in Cyprus. The phylogenetic and taxonomic status of Clitocybe maxima

(P. Gaertn., G. Mey. & Scherb.) P. Kumm., sometimes consedered a synonym of either I. geotropa or I. gibba, is as yet

unclear.

*Infundibulicybe mediterranea Vizzini, Contu & Musumeci

A widespread species commonly seen in high-altitude P. nigra subsp. pallasiana forests, but frequently also in meso-

and supra-Meditarranean P. brutia forests, e.g., Stavrós tis Psókas, 25-X-2008, ca 1200 m asl; Platánia, 30-XII-2011,

ca 1100 m asl; Cedar Valley, 29-XI-2017, ca 1100 m asl; and Plátres, 30-XII-2017, ca 1200 m.

*Infundibulicybe meridionalis (Bon) Pérez-De-Greg.

Frequently encountered in Calabrian pine forests, such as Plátres, 21-XI-2007, ca 1200 m asl; Moniátis, 28-XI-2008 &

Basidiomycetes within Pinus brutia ecosystems in Cyprus

…

26-I-2009, ca 900 m asl; Mávro Dásos, 5-XII-2012, ca 1000 m asl; Péra Pedí, 14-I-2017, ca 600 m asl. Very similar to

I. squamulosa but with considerably larger spores, measuring 7–9 × 4.5–6 μm in the Cypriot collections.

*Infundibulicybe squamulosa (Pers.) Harmaja

A handful of collections from Platánia, 30-XII-2011 & 6-I-2014, ca 1100 m asl; Pýrgos Tyllirías, 10-I-2011, ca 350 m

asl; and Plátres, 19-ΧΙ-2011 & 1-XII-2017, ca 1250 m asl. Distribution not quite clear, as a number of morphologically

very similar species co-occur on the island, difficult to distinguish in the field.

Inocybe flocculosa Sacc. sensu lato

A number of collections from diverse habitats match descriptions of this taxon, currently encompassing multiple

phylospecies: e.g., Péra Pedí, 29-XI-2011, ca 450 m asl; and Plátres, 21-XI-2012, ca 900 m asl. Previously reported by

Viney (2005), Loizides & Kyriakou (2011), Loizides et al. (2011), and Loizides (2016). It is unclear whether a

collection reported by Viney (2005) as “I. flocculosa var. crocifolia” corresponds to the same taxon or not.

*Inocybe fuscidula Velen.

One tentatively identified collection from a mossy bank in a P. brutia forest in Péra Pedí, 18-III-2011, ca 600 m asl.

Distribution unclear.

Inocybe geophylla (Bull.) P. Kumm. sensu lato

This widely applied binomial is polyphyletic and yet to be taxonomically resolved (Ryberg et al. 2008). More than one

species of this complex are present in Cyprus, with collections from, e.g. Calabrian pine forests in Moniátis, 25-XI-

2009, ca 900 m asl; Platánia, 27-XI-2009, ca 1100 m asl; or Kélefos, 2-XI-2009, 29-XI-2009 & 9-XII-2014, ca 450–

500 m asl. A lilac counterpart also occurs on the island, previously reported as “I. geophylla var. lilacina (Peck) Gillet”

in Loizides et al. (2011), e.g., from Amíantos, 10-X-2009, ca 1300 m asl. The latter variant, however, was recently

recombined as a distinct species following sequencing the holotype and appears to be restricted to the American

continent (Matheny & Swenie 2018). Large-scale genetic studies are needed to ultimately clarify this difficult species-

complex in Europe.

*Inocybe griseolilacina J.E. Lange

One collection from a P. brutia/Q. coccifera subsp. calliprinos forest in Kélefos, 19-XI-2009, ca 450 m asl.

Inocybe griseovelata Kühner

Reported by Viney (2005) under pines near Mýrtou, Jan. 2000, but not so far confirmed by the author.

*Inocybe hirtella Bres.

Several collections of this bitter-almond scented species, equally frequent in Calabrian and black pine forests, such as

Plátres, 23-XII-2009, ca 1300 m asl; Soúni, 28-VI-2010, ca 400 m asl; Pýrgos Tyllirías, 10-I-2011, ca 350 m asl; or

Pissouri, 2-I-2012, ca 200 m asl. Widespread.

Inocybe lacera (Fr.) P. Kumm. sensu lato

A couple of collections key out as this variously interpreted taxon, e.g., from Calabrian pine forests at Kélefos, 2-I-

2009, ca 450 m asl. Also reported by Viney (2005) from pine woods west of Haléfka, Feb. 1998. Several forms and

varieties have been described in literature, whose systematic position are not yet clarified.

*Inocybe langei R. Heim

Few late winter or early spring sightings in thermo-Mediterranean Calabrian pine forests, e.g., Prastió, 22-III-2011, ca

600 m asl; and Pissouri, 2-I-2012, ca 200 m asl. Occasional or locally frequent in some years.

*Inocybe muricellata Bres.

One collection found in a mixed P. brutia/Q. coccifera subsp. calliprinos forest in Kélefos, 23-XII-2012, ca 500 m asl.

Likely rare.

*Inocybe obscura (Fr.) Quél. sensu auct.

One record on a mossy bank under P. brutia and P. pinea L. near Yiolou, ca 300 m asl. This collection fits the invalidly

published taxon “I. obscura Gillet” ss. auct., tentatively treated as a synonym of I. cincinnata. Taxonomy within this

group of violet-stiped Inocybes is in need of phylogenetic revision. Distribution unclear.

Inocybe obscurobadia (J. Favre) Grund & D.E. Stuntz

Reported by Viney (2005) from pine woods near Mýrtou, Jan. 2000, but basidiomes depicted in this photograph

resemble rather I. furfurea Kühner, erroneously regarded as a synonym in the past. It is also unclear whether collections

matching the description under Cistus (Loizides 2016), sometimes reported as “I. tenuicystidiata E. Horak & Stangl”

(e.g., Esteve-Raventós et al. 2002), are conspecific or distinct from I. obscurobadia ss. str., as nomenclature within this

cluster has not yet been resolved.

*Inocybe phaeoleuca Kühner

One collection from a mossy P. brutia/P. orientalis forest in Plátres, 21-XI-2012, ca 900 m asl. Distribution unclear.

Inocybe posterula (Britzelm.) Sacc.

Several collections from Calabrian and black pine forests, e.g., Saittás, 4-XII-2008, ca 700 m asl; Kélefos, 27-XII-

2008, ca 450 m asl; and Platánia, 27-XII-2009 & 2-I-2015, ca 1100 m asl, previously reported in Loizides et al. (2011).

…

Loizides, M.

Widespread.

*Inocybe tarda Kühner

Three collections from mixed woodlands at Kélefos, 21-XI-2011, ca 450 m asl; Ayía Paraskeví, 23-XI-2011, ca 600 m

asl; and Plátres, 21-XI-2012, ca 900 m asl, match the description of this species. It is unclear whether a collection

labelled as “Inocybe aff. nitidiuscula” by Viney (2005) corresponds to the same species, since some authors (e.g.,

Kuyper 1986; Stangl 1999) consider the two taxa as synonymous, and others (e.g., Kühner R. 1955; Eyssatier & Roux

2011) as distinct.

*Inocybe tenebrosa Quél.

Just two collections of this distinctive species, notable for its unusual banana odour, one from a P. brutia forest opening

at Platánia, 27-XI-2009, ca 1100 m asl. Rare.

Inocybe vulpinella Bruyl.

Occasionally seen in thermo-Mediterranean pine forests and matorral, such as Klirou, 5-II-2012, ca 400 m asl.

Previously reported by Loizides (2016) from collections under Cistus.

Inosperma bongardii (Weinm.) Matheny & Esteve-Rav.

Recently transferred from Inocybe (Fr.) Fr. to genus Inosperma (Kühner) Matheny & Esteve-Rav., based on multigene

phylogenies (Matheny et al. 2020). One collection from Archimandrita, 2-VI-2011, ca 450 m asl, leg. T. Alexandridis,

previously reported as “Inocybe bongardii” in Loizides et al. (2011). A collection from pine forests in Haléfka, Feb.

1999, was reported by Viney (2005) as “Inocybe aff. bongardii”. Rare.

*Inosperma cervicolor (Pers.) Matheny & Esteve-Rav.

A number of ecologically diverse collections currently key out as this species, originating from mixed P. brutia/Q.

alnifolia forests in Plátres, 2-I-2010, 1200 m asl; a grassy P. brutia/Ceratonia siliqua forest opening in Pissouri, 30-I-

2011, ca 200 m asl; as well as mixed P. brutia/Q. coccifera subsp. calliprinos forests in Kélefos, 8-II-2011, ca 550 m

asl; and Prastió, 25-II-2011, ca 600 m asl. Distribution not yet understood.

Lacrymaria lacrymabunda (Bull.) Pat.

The “weeping widow” is rarely seen in Cyprus, with one collection originating from a recently burned P. brutia forest

in Pelendri, 11-III-2010, ca 800 m asl. Also reported by Viney (2005) as “Psathyrella lacrymabunda”, among pine

litter in Karmi, Dec. 2000.

Lactarius deliciosus (L.) Gray

One of the most widespread and commonly occurring fungi in Cyprus, frantically collected for the table by the locals

and sold in markets throughout the country. Fruiting prolifically on serpentine and acidic substrates in high-altitude P.

nigra subsp. pallasiana forests, but is also locally frequent in lower altitudes under P. brutia, e.g. in Moniátis ,13-XII-

2007, ca 700 m asl; Millomeri, 13-XI-2008, ca 800 m asl; Mésa Potamós, 24-XII-2008, 17-IX-2009 & 20-XII-2009,

ca 900 m asl; Amíantos, 3-X-2009, ca 1300 m asl; Plátres, 20-XI-2007, 23-XII-2009, 17-XII-2011, 17-XI-2012, 19-I-

2015, 1-XII-2017 & 17-X-2018, ca 1200 m asl; Stavrós tis Psókas, 22-XI-2011, ca 1200 m asl; Mávro Dásos, 1-XII-

2012, 28-XI-2015 & 3-XII-2015, ca 1200 m asl; Platánia, 1-XII-2009, 21-XII-2011 & 26-XII-2018, ca 1100 m asl;

Karvounás, 14-XI-2018, ca 1300 m asl. First reported in Cyprus by Nattrass (1937), and subsequently by Viney (2005),

Loizides (2008), and Loizides et al. (2011).

Lactarius sanguifluus (Paulet) Fr.

Mostly occurring in P. nigra subsp. pallasiana ecosystems in Cyprus, with only rare sightings in P. brutia forests, such

as Platánia, 9-XII-2012, ca 1150 m asl; and Plátres, 17-XII-2012, ca 1200 m asl. Common in some years.

Lactarius semisanguifluus R. Heim & Leclair

More widespread than L. deliciosus, growing prolifically in a variety of calcareous habitats in the thermo-, meso- and

supra-Mediterranean belts, typically (but not exclusively) in the presence of P. brutia: e.g. Kélefos, 23-X-2007, 29-X-

2007, 29-XI-2009, 9-II-2011, 2-XII-2014, 24-XI-2017 & 1-I-2019, ca 450 m asl; Péra Pedí, 20-XI-2007, 13-X-2008,

7-I-2016, 10-I-2017 & 19-I-2019, ca 600 m asl; Kakomállis, 29-X-2009, ca 700 m asl; Pareklisiá, 11-I-2011, ca 150 m

asl; Mosfilotí, 23-I-2011, ca 250 m asl; Mávro Dásos, 1-XII-2012, ca 1100 m asl; Akámas, 14-XII-2014, ca 200 m asl;

Akrotíri, 17-I-2015, ca 20 m asl. Very common.

*Lentinellus cystidiosus R.H. Petersen

Frequently seen on fallen P. brutia twigs and branches, such as Péra Pedí, 24-XII-2008, 7-XI-2009 & 10-I-2017, ca

550 m asl; Akámas, 14-I-2009, ca 200 m asl; Kakomállis, 2-XI-2009, ca 900 m asl.

*Lepiota alba (Bres.) Sacc.

Documented in Kélefos, 10-XI-2017, ca 500 m asl. It is as yet uncertain whether L. erminea (Fr.) P. Kumm., often

segregated on the basis of a raphanoid odour and longer spores (e.g., Huijsman 1943), is a synonym of this taxon.

Collections from high altitude P. nigra subsp. pallasiana forests in Troodos were previously reported under the latter

binomial (Loizides et al. 2011) and appear to be morphologically distinct.

Lepiota brunneoincarnata Chodat & C. Martín

This deadly poisonous species is not occasionally seen in Cyprus, with collections from various habitats (Loizides et

Basidiomycetes within Pinus brutia ecosystems in Cyprus

…

al. 2011) including P. brutia forests in Platánia, 27-XI-2008 & 7-XII-2009, ca 1100 m asl; and Kélefos, 29-X-2012, ca

500 m asl.

Lepiota castanea Quél.

Also a deadly poisonous species, with few records from Calabrian pine woods in Moniátis, 13-XII-2007, ca 700 m asl;

Plátres, 24-XI-2007, ca 900 m asl; Platánia, 20-XI-2008 & 27-XI-2008, ca 1100 m asl; and Mosfilotí, 23-I-2011, ca

250 m asl. Occasional.

Lepiota clypeolaria (Bull.) P. Kumm. sensu lato

Few collections match this frequently reported taxon, which presently occupies more than one clade in phylogenetic

analyses (Caballero et al. 2015; Vizzini et al. 2019). Found in a variety of habitats in Cyprus, mostly mixed Calabrian

pine forests in Platánia, 3-XII-2009 & 16-XI-2014, ca 1100 m asl; and Karvounás, 16-X-2011, ca 1200 m asl. A

collection north of Haléfka, Jan. 2002, was also reported as “Lepiota aff. clypeolaria” by Viney (2005).

Lepiota cristata (Bolton) P. Kumm.

Sparingly appearing in black- and Calabrian pine forests, including a collection from Platánia, 27-XI-2008, ca 1100 m

asl. Quite rare.

Lepiota echinella Quél. & G.E. Bernard

A couple of collections from Calabrian pine woods in Ayía Paraskeví, 11-XI-2009, ca 550 m asl; and Péra Vása, 7-XI-

2012, ca 700 m asl. Also seen once in black pine forests, but generally uncommon or rare.

*Lepiota forquignoni Quél.

Known only from one collection at Mosfilotí, 23-I-2011, ca 250 m asl. Rare.

Lepiota griseovirens Maire

Reported by Loizides et al. (2011) from two collections in different localities at Kélefos, both on 20-XI-2009 in P.

brutia/Platanus orientalis L. forests, ca 450 m asl. Rare.

*Lepiota lepida Guinb. & M. Bodin

Two collections of this rarely reported species from Platánia, 1-XII-2009 & 3-X-2018, in mixed P. brutia forest

opening, ca 1100 m asl. The spores of the Cypriot collections measure 6–7 × 3–4 μm.

*Lepiota pseudohelveola Kühner

A couple of collections match the concept of species from Kélefos, 28-XI-2009, in mixed P. brutia/Q. coccifera subsp.

calliprinos forest, ca 500 m asl; and Péra Pedí, 14-I-2017, in P. brutia forest, ca 600 m asl. More than one species of

this cluster are present in Cyprus, but it is yet unclear whether L. pseudolilacea is a synonym of this taxon.

Lepiota pseudolilacea Huijsman

Seen in a grassy P. brutia forest clearing in Saittás, in 5-X-2008, 7-X-2009, 30-XII-2010, 29-IX-2011 and 16-X-2012.

Lepiota lilacea Bres. also occurs in Cyprus and is very similar, but has smaller spores and is typically found outside of

forests (Loizides et al. 2011).

Lepiota subgracilis Kühner

Previously reported by Loizides et al. (2011) from a couple of collections in mixed P. brutia/Q. coccifera subsp.

calliprinos forests in Kélefos, ca 500 m asl; and Péra Pedí, 28-XI-2009, ca 550 m asl. Lepiota kuehneri Huijsman ex

Hora, L. kuehneriana Locq., Lepiota latispora (Kühner ex Wasser) Bon, and L. wasseri Bon are probably all synonyms

of this taxon (Vellinga 2001). Infrequently seen.

Lepiota subincarnata J.E. Lange

An occasionally encountered woodland dweller, with collections from Kélefos, 27-XI-2007, 29-XI-2009 & 8-I-2011,

in a riparian P. brutia/Platanus orientalis forest, ca 450 m asl; Platánia, 5-XII-2009, in P. brutia forest, ca 1200 m asl;

and Stavrós tis Psókas, 20-XI-2017, 1100 m asl in mixed P. brutia forest. Deadly poisonous.

Lepiota sublaevigata Bon & Boiffard

Occasionally seen in thermo-Mediterranean P. brutia forests and Cistus matorral, e.g., Péra Pedí, 7-XI-2009, ca 550 m

asl; Kélefos, 10-XI-2017, ca 500 m asl.

Lepista nuda (Bull.) Cooke sensu lato

A common and widespread woodland dweller found in various habitats in Cyprus, with numerous collections from P.

brutia forests in Plátres, 21-XI-2007, 1-XII-2009, 24-XI-2012 & 3-XII-2018, ca 1100 m asl; Moniátis, 9-XII-2009, ca

900 m asl; Platánia, 20-XI-2008, 23-XI-2009, 27-XI-2009, 30-XII-2014, 20-XI-2015, 26-XII-2018, ca 1100 m asl;

Stavrós tis Psókas, 18-XI-2009, ca 1200 m asl; Saittás, 6-I-2017, ca 700 m asl. Also reported by Viney (2005) from

pine woods around Kyrenia and Haléfka. See further notes under L. sordida.

*Lepista panaeolus (Fr.) P. Karst.

Just two collections of this rare species in Cyprus, both from grassy P. brutia forest openings in Akámas, 4-I-2011 &

14-ΧΙΙ-2014, ca 200 m asl.

Lepista sordida (Schumach.) Singer sensu lato

Very common in a variety of habitats, especially grassy, low-elevation Calabrian pine wood openings: e.g., Pissouri,

…

Loizides, M.

12-II-2007, 30-I-2011 & 2-I-2012, ca 200 m asl; Saittás, 13-XII-2007, 700 m asl; Mosfilotí, 6-I-2009 & 23-I-2011, ca

250 asl; Péra Vása, 8-II-2015, ca 600 m asl. Multiple lineages identified as L. nuda and L. sordida seem to be be present

in phylogenetic trees, therefore these two widely applied binomials might further split in the future and taxonomic

revisions are to be expected (Alvarado et al. 2015).

*Leucoagaricus georginae (W.G. Sm.) Candusso

One collection from Platánia, 25-X-2012, in a mossy P. brutia/C. sempervirens forest clearing, ca 1100 m asl. Rare.

Leucoagaricus melanotrichus (Malençon & Bertault) Trimbach

One collection from a P. brutia forest in Ayía Paraskeví, 11-XI-2009, ca 600 m asl. Also reported in Loizides (2016)

from a collection under Cistus in a P. brutia/C. brevifolia forest at Tripilos, 18-XI-2009, ca 1100 m a.s.l. Rare.

Leucopaxillus gentianeus (Quél.) Kotl.

Among the commonest fungi in Cyprus, occurring abundantly in a variety of habitats including mixed P. brutia forests

in Platánia 3-XII-2009, 3-XII-2012 & 26-I-2018, ca 110 m asl; Plátres, 23-XII-2009, 7-XII-2012, 11-XII-2012, 17-

XII-2012 & 1-XII-2017, ca 1200 m asl; Stavrós tis Psókas, 18-XI-2009 & 22-XI-2011, ca 1200 m asl; or Mávro Dásos,

5-XII-2012, ca 1000 m asl.

Leucopaxillus malenconii Bon

Two collections fit the description of this Mediterranean species previously reported in Loizides et al. (2011): one from

a riparian P. orientalis, Alnus orientalis and P. brutia forest in Moniátis, 9-XII-2007, ca 850 m asl; and one from a P.

brutia forest in Platánia, 5-XII-2009, ca 1200 m asl. Rare.

Limacella illinita (Fr.) Maire

Few sightings in Plátres 15-XI-2007 & 25-XI-2009, ca 1150 m asl; Trimíklini, 20-XII-2008, ca 650 m asl; Saittás, 10-

I-2017, ca 700 m asl, all in P. brutia forests, with some collections corresponding to “f. ochracea” characterized by

ochraceous tinges on the pileus. Also reported by Viney (2005) from pine woods near Áyios Ilaríon, Dec. 2000.

Limacella subfurnacea Contu sensu auct.

Occasionally seen in and out of forests, with one collection from a P. brutia forest edge in Finikária, 20-I-2011, ca 200

m asl. Previously reported by Viney (2005) from pine woods in Haléfka (Dec. 1998), and Ayía Iríni (Dec. 2000).

Limacella grisea Singer (1989) might be an earlier name for this species.

Lycoperdon atropurpureum Vittad.

One collection in a mossy P. brutia forest in Kélefos, 31-X-2012, ca 500 m asl. Also reported from Áyios Mámas, 10-

III-2012, ca 500 m asl, among Cistus litter in a P. brutia forest clearing (Loizides 2016). Distribution unclear.

Lycoperdon excipuliforme (Scop.) Pers.

One collection from an open P. brutia forest in Pareklisiá, 14-II-2009, ca 200 m asl. Probably rare.

*Lycoperdon lividum Pers.

Occasionally encountered, such as in open P. brutia forests at Kélefos, 10-XI-2017, ca 500 m asl.

Lycoperdon molle Pers.

A couple of collections from P. brutia forests in Saittás, 30-XII-2010 & 28-X-2011, ca 700 m asl. Distribution not quite

understood, since its distinction from L. umbrinum, L. subumbrinum and L. umbrinoides is not straight forward.

Lycoperdon perlatum Pers.

Frequently encountered in black- and Calabrian pine forests, e.g., Saittás, 13-XII-2007, ca 700 m asl; Plátres, 27-X-

2008, ca 1200 m asl; Trimíklini, 9-XII-2008, ca 650 m asl; Kélefos, 29-XI-2014 & 24-XI-2017, ca 500 m asl. Also

reported by Viney (2005), without date, from the pine woods of Haléfka; by Loizides et al. (2011) from collections

cited above; and by Loizides (2016) from Cistus matorral in Kalavasyós.

Lycoperdon pratense Pers.

Previously reported by Viney (2005) as “Vascellum pratense”, this puffball is usually found in fields or woodland

edges. A couple of sightings from P. brutia forest clearings in Prastió, 5-III-2011, ca 550 m asl; and Akámas, 18-XII-

2014, ca 200 m asl.

*Lycoperdon umbrinum Pers.

A couple of collections from a mixed P. brutia/Q. coccifera subsp. calliprinos forest in Kelefos, 2-XI-2012, ca 500 m

asl; and a P. brutia forest in Prastio 6-III-2012, ca 550 m asl. Distribution not yet understood.

Lyophyllum fumosum (Pers.) P.D. Orton

A morphologically variable and ecologically versatile species found in diverse ecosystems in Cyprus, including P.

brutia forests in, e.g., Saittás, 25-XI-2007, 1-XII-2011 & 12-XII-2011, ca 700 m asl; Amíantos, 10-X-2009, ca 1300 m

asl; and Plátres, 20-XII-2009 & 11-XII-2012, ca 1200–1300 m asl. Previously reported by Loizides et al. (2011) from

collections cited above, and Loizides (2016) from Cistus matorral. Lyophyllum cistophilum Vila & Llimona, L.

phaeophyllum Vila & Llimona, and L. subglobisporum Consiglio & Contu, might be synonyms of this taxon.

Collections previously reported by Viney (2005) as “Lyophyllum decastes (Fr.) Singer”, probably also correspond to

this widespread species.

Basidiomycetes within Pinus brutia ecosystems in Cyprus

…

*Lyophyllum loricatum (Fr.) Kühner

Just three collections all late in the season, two of them from mixed P. brutia/P. orientalis forests at Platánia, 25-XI-

2013 & 26-XII-2018, ca 1100 m asl. Apparently rare.

*Macrocystidia cucumis (Pers.) Joss.

The “cucumber cap” is sparingly seen in black- and Calabrian pine forests: Platánia, 9-XII-2009, ca 1100 m asl; Ayía

Paraskeví, 18-II-2011, ca 550 m asl; Akámas, 15-I-2016, ca 200 m asl; Plátres, 1-XII-2017, ca 1200 m asl.

Macrolepiota excoriata (Schaeff.) Wasser

Frequently encountered in thermo-Mediterranean Calabrian pine wood edges and grassy wood openings, e.g., in

Kélefos, 3-XII-2008, ca 500 m asl; Akámas, 14-I-2009, 200 m asl; Kalavasos, 14-II-2011, ca 180 m asl; Péra Pedí, 10-

I-2017, ca 600 m asl (Loizides et al. 2011).

*Macrolepiota mastoidea (Fr.) Singer

So far only seen in Calabrian pine forests of the Akámas Peninsulla where it appears in large numbers locally, e.g., 28-

XI-2009, ca 200 m asl. Rare or altogether absent elsewhere on the island.

*Macrolepiota procera f. permixta (Barla) Vizzini & Contu

Just one collection from a P. brutia forest opening in Karvounás, 20-XI-2015, ca 1200 m asl. This form, characterised

by the vinaceous-red bruising of basidiomes, was previously regarded as a distinct species (Pacioni 1979; Breitenbach

& Kränzlin 1991; Vellinga EC. 2001). However, as shown by Vizzini et al. (2011), this character has no phylogenetic

basis and collections identified as this species (including the epitype collection), nest in the M. procera (Scop.) Singer

clade. Evidently very rare in Cyprus, not a species to go unnoticed.

*Macrolepiota subsquarrosa (Locq.) Bon

Several collections match the description of this species, e.g., from P. brutia forests in Plátres, 20-XI-2007 & 27-X-

2012, ca 1000–1300 m asl; P. brutia/Q. coccifera subsp. calliprinos forests in Kélefos, 20-XI-2009 & 21-I-2015, ca

500 m asl; and P. brutia forests at Kykkos, 29-XI-2017, ca 1200 m asl. Perhaps widespread, but the phylogenetic

identity of this taxon has not yet been fully clarified and is often difficult to distinguish from the similar M. excoriata

and M. mastoidea.

Mallocybe dulcamara (Pers.) Vizzini, Maggiora, Tolaini & Ercole. sensu auct.

A variously interpreted spring taxon whose taxonomic identity is yet to be fully resolved (Matheny et al. 2020).

Reported by Viney (2005) from Karmi as “Inocybe dulcamara”, May 2000. Collections from a mixed P. brutia/A.

andrachne/Q. coccifera subsp. calliprinos forest in Prastió, 22-III-2011, ca 550 m asl, match the description of this

taxon, but the current combination might be invalid as per Art. 41(1) of ICN (Turland et al. 2018).

Marasmius wynneae Berk. & Broome

Very common and widespread, found in various habitats and elevations including P. brutia forests in, e.g., Pissouri,

15-I-2008 & 2-I-2012, ca 200 m asl; or Pareklisiá, 14-II-2009 & 25-I-2011, ca 200 m asl. Also reported by Viney

(2005) from Haléfka, Dec. 1998.

Melanoleuca exscissa (Fr.) Singer

Often seen in grassy areas at lower elevations, with at least one collection in a P. brutia wood opening in Péra Pedí, 1-

III-2013, ca 550 asl. Previously reported by Viney (2005) from Panagra, Dec. 2004. Possibly widespread, but the

morphological identification of Melanoleuca Pat. species is far too complicated to draw reliable conclusions.

*Melanoleuca iris Kühner

One collection from a grassy P. brutia forest in Pareklisiá, 18-II-2009, ca 200 m asl. Distribution unclear.

Mycena amicta (Fr.) Quél.

Commonly seen among pine litter, or on logs, pine cones and wooden debris, e.g., in Calabrian pine woods in Saittás,

29-X-2008, ca 700 m asl; Akámas, 4-I-2011, ca 200 m asl; or Moniátis, 12-XII-2011, ca 700 m asl. Also reported by

Viney (2005) from Kyrenia, Jan. 2000.

Mycena arcangeliana Bres.

Reported by Viney as “M. oortiana” (2005) and occasionally seen on fallen logs and tree stumps, e.g., in a mixed

Cedrus brevifolia/P. brutia/Q. alnifolia forest in Cedar Valley, 27-XII-2010, ca 1100 m asl; or in mixed P. brutia/Q.

alnifolia/A. andrachne forests in Platánia, 9-XII-2012, ca 1100 m asl.

*Mycena capillaripes Peck

Many sightings from mixed Calabrian pine forests, e.g., in Saittás, 24-XII-2010, ca 650 m asl; Akámas, 4-I-2011, ca

200 m asl; and Amíantos, 19-XI-2013, ca 1200 m asl. Common and widespread.

*Mycena clavicularis (Fr.) Gillet

Appearing in vast numbers in wet P. brutia forests during the autumn and winter months, such as: Moniátis, 22-XII-

2010, ca 900 m asl; Kannavioú, 27-XII-2010, ca 900 m asl; Kelláki, 25-I-2011, ca 600 m asl; or Plátres, 11-XII-2012,

26-XI-2014 & 1-XII-2017, ca 1200 m asl. Very common and widespread.

…

Loizides, M.

*Mycena epipterygia (Scop.) Gray sensu lato

Several varieties of this fungus have been described in literature, whose phylogenetic and taxonomic status remain

unclear. Few collections corresponding to some of these variants originate from P. brutia forests in Cyprus, such as

Mésa Potamós, 20-XII-2007, 7-I-2009 & 17-XII-2009, ca 700 m asl; Platánia, 7-XII-2009, 9-XII-2009 & 30-XII-2011,

ca 1100 m asl; and Plátres, 6-I-2017, ca 1200 m asl.

*Mycena flavescens Velen.

One collection in a mixed P. brutia/Q. alnifolia/P. orientalis forest in Platánia, 30-XII-2011, ca 1100m asl. Distribution

unclear.

Mycena galopus (Pers.) P. Kumm.

Reported by Viney (2005), without dates or localities, and only sparingly seen by the author: Trimíklini, 30-XII-2007,

in P. brutia forest, ca 650 m asl; Kélefos, 21-XI-2011, in a riparian P. brutia/P. orientalis/A. orientalis forest, ca 450

m asl. Distribution unclear, as several forms and varieties are described in literature.

*Mycena haematopus (Pers.) P. Kumm.

Rare, known only from two collections, one in a riparian P. brutia/P. orientalis/A. orientalis forest at Kélefos, 21-XI-

2011, ca 450 m asl.

*Mycena mirata (Peck) Sacc.

One collection keying out as this species, albeit with larger spores than those mentioned in literature, measuring 11–15

× 5.5–7.5 μm in the Cypriot specimens. Found in a mixed P. brutia/Q. alnifolia/A. andrachne forest in Platánia, 21-

XII-2011, ca 1100 m asl. Distribution unknown.

*Mycena pseudopicta (J.E. Lange) Kühner

One collection from Kélefos, 2-I-2009, in a mossy P. brutia forest, ca 550 m asl. Apparently rare.

Mycena pura (Pers.) P. Kumm. sensu lato

Numerous collections belonging to this species-cluster, shown by Harder et al. (2010, 2013) to encompass multiple

phylospecies yet to be taxonomically resolved. More than one species of this complex are present in Cyprus, with

numerous collections from Calabrian pine forests in Kélefos, 2.I-2009, 20-XI-2009, 25-XI-2009 & 28-XI-2009, ca 500

m asl; Platánia, 3-XII-2009, ca 1100 m asl; Karvounás, 21-X-2011, 1200 m asl; Kouka, 29-XI-2011, ca 600 m asl;

Madarí, 3-XII-2012, ca 1300 m asl; Mésa Potamós, 10-I-2017, ca 800 m asl; Kannavioú, 29-XI-2017, ca 900 m asl.

Also reported by Viney (2005) in Larnaka tis Lapithou, Dec. 1998.

*Mycena renati Quél.

This striking species was only seen once on a P. brutia log at Stavrós tis Psókas, 18-XI-2009, ca 1200 m asl. Rare.

*Mycena rosea Gramberg

Occasionally seen in a variety of habiatas: Plátres, 27-X-2008, in P. brutia forest, ca 1200 m asl; Péra Pedí, 28-X-2009,

in P. brutia/ Q. coccifera subsp. calliprinos forest, ca 550 m asl; Platánia, 21-XII-2011, in P. brutia/Q. alnifolia forest,

ca 1100 m asl. Uncommon or rare.

Mycena sanguinolenta (Alb. & Schwein.) P. Kumm.

Few sightings, one among P. brutia/Crataegus azarolus/Cistus litter in Saittás, 24-XII-2010, ca 650 m asl. Previously

reported by Viney (2005) from Haléfka, Dec. 2001. Probably widespread but often unnoticed due to its tiny size.

Mycena seynii Quél.

One of the commonnest basidiomycetes in Cyprus, fruiting throughout the island where fallen P. brutia cones are

present. Hundreds of sightings, from Trimíklini, 29-XI-2007, 17-XII-2011 & 31-XII-2013, ca 650 m asl; to Saittás, 30-

XI-2007, 24-XII-2011 & 30-XII-2013, ca 700 m asl; Kélefos, 2-I-2009, 2-XI-2009, 25-XI-2009, 8-I-2011 & 9-XII-

2015, ca 550 m asl; Stavrós tis Psókas, 18-XI-2009, ca 1200 m asl; Moniátis, 20-XII-2010, ca 800 m asl; Pissouri, 2-I-

2011, ca 200 m asl; Kelláki, 25-XI-2011, ca 600 m asl; Kannavioú, 28-XII-2016, ca 900 m asl; or Plátres, 1-XII-2017,

ca 1200 m asl.

*Mycena cf. stipata Maas Geest. & Schwöbel

A couple of collections keying out as this species, both found on carbonized P. brutia logs and twigs in Asgata, 24-

XII-2011 & 26-XII-2011, ca 200 m asl. Cypriot collections feature partially encrusted and considerably larger cystidia

than those reported in literature, measuring 70–115 × 13–16 μm, while the spores measure (7–)8–11 × 4.5–5.5 μm.

Either a highly variable taxon, or a Mediterranean counterpart to be clarified by future DNA sequencing.

*Panaeolus acuminatus (P. Kumm.) Quél.

Sparingly encountered in and out of forests, with one collection originating from a grassy P. brutia forest clearing in

Platánia, 21-XII-2011, ca 1100 m asl. Occasional.

*Phaeoclavulina myceliosa (Peck) Franchi & M. Marchetti

Few collections key out as this species, from a mixed riparian forest in Plátres, 23-IX-2008, 5-X-2009 & 10-XI-2012,

ca 900 m asl; as well as calcareous Calabrian pine forests in Kélefos, 23-XII-2012, ca 500 m asl; and Péra Vása, 29-

XII-2012, ca 700 m asl. Probably widespread, though presently unclear whether European collections identified as this

Basidiomycetes within Pinus brutia ecosystems in Cyprus

…

taxon are conspecific to the North American ones.

Phallus impudicus L.

The notorious “stinkhorn” is only sparingly encountered in Cyprus with some sightings originating from Calabrian pine

forests in Saittás, 20-XII-2009 & 12-ΧΙΙ-2011, ca 700 m asl; and Péra Vása, 9-XII-2012, ca 700 m asl.

Pholiota gallica Holec & M. Kolařík

This species was previously known as “P. lubrica var. obscura Bon & Chevassut”, “P. highlandensis var.

citrinosquamulosa Maire ex Bidaud & Borgarino”, and occasionally misidentified as “Pholiota brunnescens A.H. Sm.

& Hesler”, before it was recombined under its current name (Holec & Kolařík 2014). In Cyprus, it has already been

reported in Cistus mattoral in Asgata (Loizides 2016), but was also collected in P. brutia forests in Akámas, 23-II-

2016, ca 200 m asl.

Pholiota mixta (Fr.) Kuyper & Tjall.-Beuk.

Reported by Viney (2005) on buried conifer wood, beside the Buffavento dirt road, Jan. 2001. Not seen by the author.

Pholiotina filaris (Fr.) Singer

Reported from Kaló Chorió by Viney (2005), but this may correspond to the recently described P. mediterranea Siquier

& Salom (Siquier & Salom 2017).

*Pisolithus tinctorius (Pers.) Coker & Couch

A couple of sightings by a dirt road in Calabrian pine forests at Kélefos, 19-X-2008 & 4-XII-2016, ca 450–500 m asl;

a grassy P. brutia forest clearing in Kato Plátres, 21-X-2008, ca 800 m asl; and a P. brutia/Q. alnifolia forest in Spília,

3-XII-2015, ca 1100 m asl. Less common than P. arhizus (Scop.) Rauschert, the latter typically found outside of forests

and lacking a well-defined stalk (Loizides et al. 2011). A couple of the images identified as “P. arrhizus” in Viney

(2005) also seem to depict P. tinctorius. The two taxa were widely regarded as synonymous in the past, but in recent

years have shown to be genetically distinct (e.g., Phosri et al. 2012).

Psathyrella spadiceogrisea (Schaeff.) Maire

Reported by Viney (2005) east of Haléfka, Jan. 2001, in mixed tree litter.

*Psathyrella tephrophylla (Romagn.) Bon

A couple of collections match the description of this species, one of them from a grassy P. brutia forest opening in

Kelláki, 25-I-2011, ca 600 m asl. Distribution unclear.

Psilocybe crobula (Fr.) Singer

Rarely encountered in Cyprus, with just a handful of collections from P. brutia forests in Péra Pedí, 7-XI-2009, ca 550

m asl; Madarí, 28-XI-2015, ca 1300 m asl; and Kykkos, 29-XI-2017, ca 1200 m asl. Also reported in Áyios Ilaríon,

Dec. 2000, on fallen pine bark (Viney 2005).

*Ramaria apiculata (Fr.) Donk

A couple of collections from mixed P. brutia/Q. alnifolia/Cistus forests in Ayía Paraskeví, 14-XI-2009, ca 600 m asl;

and Amíantos, 23-X-2011, ca 1200 m asl. Distribution not yet understood.

*Ramaria rubella (Schaeff.) R.H. Petersen

One collection on a decorticated P. brutia log at Kélefos, 23-XII-2012, ca 500 m asl. A somewhat faded collection

from Ayía Paraskeví, 28-III-2009, previously reported as “Ramaria bataillei (Maire) Corner” in Loizides et al. (2011),

may also represent this taxon. Rare.

Ramaria sanguinea (Pers.) Quél.

One collection of strongly rufescent basidiomes matching the description of this species, found in a mixed P. brutia/Q.

alnifolia forest at Stavrós tis Psókas, 25-X-2008, ca 1150 m asl (Loizides et al. 2011). Distribution unclear.

*Ramariopsis subtilis (Pers.) R.H. Petersen

Two collections from Kannavioú matching the descriptions of this taxon on 13-XII-2009 & 15-XII-2009, found in

mixed Calabrian pine forests, 900 m asl. Rare.

Rhizocybe vermicularis (Fr.) Vizzini, P. Alvarado, G. Moreno & Consiglio

Possibly the most widespread basidiomycete appearing in spring, fruiting in large numbers in Calabrian pine forests at,

e.g. Trimíklini, 3-III-2008, 4-III-2009, 24-II-2010, 1-III-2010, 21-III-2011, 1-III-2013 & 28-II-2019, ca 600 m asl; Péra

Pedí, 15-III-2008, 4-III-2009, 15-III-2009, 19-III-2009, 22-III-2009, 12-III-2015 & 2-III-2019, ca 550 m asl; Ayía

Paraskeví, 3-IV-2011, 3-III-2013, ca 550 m asl; Pareklisiá, 8-III-2012 & 25-II-2015, ca 200 m asl; Soúni, 13-III-2012,

5-III-2013, 7-III-2015 & 10-III-2018, ca 400 m asl; Prastió, 25-II-2-15, ca 600 m asl; Kélefos, 5-III-2014 & 5-III-2019,

ca 500 m asl; Episkopí, 28-II-2015, ca 150 m asl; Agrós, 30-III-2015 & 16-III-2019, ca 1000 m asl. Also reported by

Viney (2005) as “Clitocybe vermicularis”, above Haléfka, Mar. 1997.

*Rhizopogon graveolens f. pomaceus Vaček

A sequenced collection from a P. brutia forest in Kélefos, 28-I-2010, ca 500 m asl, nested with the isotype of this

poorly known taxon (det. & pers. comm. M. Martín), which according to Martín & García (2009) probably corresponds

to R. rubescens (Tul. & C. Tul.) Tul. & C. Tul. ss. str. Distribution unclear.

…

Loizides, M.

*Rhizopogon occidentalis Zeller & C.W. Dodge

Few collections matching the description of this taxon, characterized by prominent rhizomorphs engulfing the entire

exoperidium. Seen in both black- and Calabrian pine forests at Kélefos, 6-II-2011 & 12-II-2011, ca 450m asl; and Péra

Pedí, 10-XI-2012, ca 550 m asl.

Rhizopogon vulgaris (Vittad.) M. Lange sensu auct.

An ambiguous and variously interpreted taxon, sometimes regarded as a synonym of R. roseolus (Corda) Th. Fr.

Collections identified as the latter in Cyprus, however, appear to be morphologically distinct and restricted to the higher

elevations of Troodos, found in strict association with P. nigra subsp. pallasiana. There are numerous tentatively

identified collections of R. vulgaris sensu auct. from P. brutia forests in Trimíklini, 17-II-2010, ca 600 m asl; Péra

Pedí, 1-III-2014, ca 550 m asl; Akrotíri, 24-XI-2014, ca 20 m asl; or Soúni, 9-III-2014, ca 400 m asl. Further reported

by Viney (2005) in Kharcha, Nov. 1997, and by Loizides (2016) from Cistus matorral. A collection under Cistus in

Kormakitis reported by Nattrass (1937) as “R. luteolus Fr.”, also likely belongs to this species. A difficult complex in

need to be resolved by dedicated phylogenetic investigations.

*Rhodocollybia butyracea (Bull.) Lennox

Common and widespread in Cyprus, frequently seen in black- and Calabrian pine forests. Numerous sightings from,

e.g., Kélefos, 27-XI-2007, 29-XI-2009 & 10-XI-2017, ca 450–500 m asl; Platánia, 7-XI-2008, 20-XI-2008, 3-XII-

2009, 5-XII-2011 & 5-I-2012, ca 1100 m asl; or Mésa Potamós, 10-I-2017, ca 800 m asl.

*Rhodocybe matesina Picillo & Vizzini

One collection of this recently described species (Crous et al. 2017), until now known only from Italy: Platánia, 26-

XII-2018, ca 1100 m asl, among P. brutia and C. sempervirens litter. The spores of the Cypriot collection measure 5–

7 × 4–5 μm. Rare.

*Rimbachia neckerae (Fr.) Redhead

This minute cyphelloid basidiomycete was documented once growing among mosses, in a P. brutia forest at Kélefos,

22-XII-2012, ca 500 m asl. Distribution unclear, but probably overlooked.

Rugosomyces onychinus (Fr.) Raithelh.

Occasionally seen in black and Calabrian pine ecosystems, with collections from Moniátis, 13-XII-2007, ca 800 m asl;

Platánia, 23-XI-2009, ca 1100 m asl; Plátres, 26-XII-2009, ca 1300 m asl; and Péra Pedí, 10-I-2017, ca 550 m asl.

Russula acrifolia Romagn.

Fruiting prolifically in P. nigra subsp. pallasiana forests, but occasionally also seen in mixed P. brutia forests, such as

Amíantos, 15-X-2008, ca 1300 m asl; Madarí, 20-XI-2015, ca 1200 m asl; Troodítissa, 11-VI-2018, ca 1300 m asl.

Very common.

*Russula atropurpurea (Krombh.) Britzelm.

Occasionally seen, with a couple of collections from P. brutia/Q. alnifolia forests in Plátres, 17-XI-2011, ca 1200 m

asl; and Platánia, 2-XII-2012, ca 1100 m asl.

Russula aurea Pers.

Widespread in P. nigra subsp. pallasiana forests in years with early rainfall (Loizides et al. 2011), but occasionally

also seen in mixed P. brutia/Q. alnifolia forests in, e.g., Platánia, 20-XI-2009 & 25-VI-2018, ca 1150 m asl.

*Russula badia Quél.

One collection in a pure P. brutia forest in Saittás, 21-XI-2012, ca 700 m asl. Distribution not yet understood.

Russula chloroides (Krombh.) Bres.

Very common, appearing prolifically in mixed black- and Calabrian pine forests, typically at altitudes above 1000 m.

Numerous sightings, e.g., from Amíantos, 15-X-2008, ca 1300 m asl; Troodítissa, 8-IX-2009, 25-X-2011, 31-X-2011,

16-XI-2014 & 14-VI-2018, ca 1300 m asl; Platánia, 25-VI-2018, ca 1150 m asl; Madarí, 20-XI-2015, ca 1350 m asl.

Russula delica Fr.

One of the most common and ecologically versatile fungi on the island, fruiting in vast numbers from ~200 m asl to

the peaks of Troodos. Equally abundant on both calcareous and serpentine soil, with hundreds of sightings from mixed

or pure Calabrian pine forests e.g. in Plátres, 21-XI-2007, 15-X-2008, 19-XI-2011, 1-XII-2017 & 16-X-2018 ca 1100–

1300 m asl; Péra Pedí, 21-XI-2007, ca 600 m asl; Saittás, 25-XI-2007, 29-IX-2011, 16-X-2011 & 20-X-2018 ca 700 m

asl; Cedar Valley, 18-XI-2009, 29-XI-2017 & 6-XII-2019, ca 1100 m asl; Mésa Potamós, 9-XII-2007, 6-I-2017, 2-XII-

2017 & 1-XII-2019, ca 700–850 m asl; Amíantos, 19-X-2008, 10-X-2009, 16-X-2011 & 21-X-2011, ca 1200–1300 m

asl; Troodítissa, 12-VIII-2010 & 16-VI-2014, ca 1300 m asl; Karvounás, 13-X-2011 & 13-XI-2018, ca 1250 m asl;

Platánia, 23-X-2011 & 19-VI-2018, ca 1100–1200 m asl; Stavrós tis Psókas, 22-XI-2011, ca 1200 m asl; Germasógeia,

9-XI-2012, ca 200 m asl; Mávro Dásos, 1-XII-2012 & 3-XII-2012, ca 1000 m asl. Also reported in Syrianóchori, Nov.

2000 (Vibey 2005).

Russula foetens Pers.

Occasional in mixed forests, with a couple of collections from a P. brutia/Q. coccifera subsp. calliprinos forest in

Trimíklini, 25-XI-2012, ca 650 m asl; and P. brutia/Q. alnifolia forest in Platánia, 25-VI-2018, ca 1150 m asl.

Basidiomycetes within Pinus brutia ecosystems in Cyprus

…

Russula hobartiae Loizides & J.M. Vidal

This sequestrate endemic species was recently described from the peaks of Troodos, where it occurs in almost exclusive

association with P. nigra subsp. pallasiana (Vidal et al. 2019). Seen only once under P. brutia in Platánia, 16-VI-2014,

ca 1100 m asl, leg. M. Tordelli & C. Hobart.

Russula sanguinea Fr.

Sparingly encountered, seen in Calabrian pine forests at Péra Pedí, 8-XII-2007 & 10-I-2017, ca 550 m asl; Mávro

Dásos, 5-XII-2012, ca 1000 m asl; Plátres, 17-X-2018, ca 1250 m asl; and Péra Vása, 7-XI-2012, ca 600 m asl.

Previously reported by Viney (2005) from Syrianóchori, Dec. 1998, by Loizides et al. (2011) from collections cited

above, and by Loizides (2016) from Cistus matorral.

Russula torulosa Bres. sensu lato

Several collections match various interpretations of this taxon, most likely encompassing more than one phylospecies.

The phylogenetic and taxonomic status of some satellite taxa such as R. fuscorubra (Bres.) J. Blum, or R. fuscorubroides

Bon, also remains unclear, therefore this complex needs to be resolved by molecular phylogenetics and the appointment

of neo/epitypes. Few collections from Calabrian pine forests in Mandriá, 25-XI-2007, ca 700 m asl; Péra Pedí 8-XII-

2007 & 10-I-2017, ca 550 m asl; Saittás, 15-XI-2007 & 30-XII-2010, ca 700 m asl; Platánia, 10-X-2009 & 25-XI-2013,

ca 1100 m asl. Also reported by Viney (2005), without localities given.

*Russula werneri Maire

Few collections from P. brutia forests match the concept of this rare taxon described from Morocco (Maire & Werner

1938): Saittás, 17-XI-2007 & 5-X-2008, ca 700 m asl; Trimíklini, 10-XI-2012 & 21-XI-2012, ca 650 m asl.

Sarcodon leucopus (Pers.) Maas Geest. & Nannf.

Frequently encountered in Cyprus but very rare elsewhere, and considered extinct in large parts of western and central

Europe. Mostly appearing in high-altitude P. nigra subsp. pallasiana forests (Loizides et al. 2011) in years with early

rainfall, but seen only once under P. brutia in the Platánia picnic area, 19-VI-2018, ca 1100 m asl.

Scleroderma polyrhizum (J.F. Gmel.) Pers.

Occasionally encountered in P. brutia forest clearings and road edges, such as Plátres, 2-X-2007, ca 1150 m asl;

Moniátis, 13-XII-2007, ca 750 m asl; Trypilos, 8-XI-2009, ca 1250 m asl (Loizides et al. 2011).

Scleroderma verrucosum (Bull.) Pers.

An uncommon species first reported by Viney (2005) in Karmi, Nov. 1997. Mostly seen in mixed forests and cultivated

land, with collections from a riparian P. brutia/P. orientalis forest in Moniátis, 29-X-2008, ca 900 m asl; a P. brutia

forest edge in Agrós, 12-III-2012, ca 1000 m asl; a P. brutia/Q. coccifera subsp. calliprinos forest in Kélefos, 23-XII-

2012, ca 500 m asl; and a grassy P. brutia forest opening also in Kélefos, 29-XII-2014, ca 450 m asl.

*Singerocybe phaeophthalma (Pers.) Harmaja

Previously placed in Clitocybe, this species has been transferred to back to genus Singerocybe Harmaja based on

phylogenetic data (Qin et al. 2014; Alvarado et al. 2015, 2018a, 2018b). Just a couple of records in Cyprus, one from

a P. brutia forest clearing at Platánia, 5-XII-2011, ca 1100 m asl. Uncommon or rare.

Sparassis crispa (Wulfen) Fr.

The “cauliflower fungus” is rare and highly localised in Cyprus (Loizides et al. 2011), largely restricted to the western

provinces of the island, particularly the Paphos forest: Cedar Valley, 18-XI-2009, ca 1100 m asl; Stavrós tis Psókas,

22.XI.2011, ca 1250 m asl; Kannavioú, 28-XII-2016, ca 800 m asl.

Stropharia aeruginosa (Curtis) Quél.

Rarely encountered, with one of the collections coming from a mixed Calabrian pine forest in Moniátis, 9-ΧII-2009,

ca 800 m asl. Also reported by Viney (2005) in Lapithos, Dec. 2000.

Suillus bellinii (Inzenga) Kuntze

One of the most prolifically occurring fungi on the island, fruiting in huge numbers in black- and Calabrian pine forests

throughout the autumn and winter months. Hundreds of sightings, e.g. in Saittás, 15-XI-2007, 30-XII-2010, 29-IX-

2011 & 6-I-2017, ca 700 m asl; Moniátis, 13-XII-2007, 25-XI-2009 & 6-I-2017, ca 700 m asl; Mésa Potamós, 23-XII-

2007, 12-XII-2011 & 10-I-2017, ca 800 m asl; Kélefos, 25-XI-2009, 6-XII-2013, 21-I-2015, 9-XII-2015, 10-I-2017,

14-I-2017 & 3-I-2019, ca 450–500 m asl; Platánia, 10-X-2009, 3-XII-2009, 20-XII-2014, 20-XI-2015 & 16-XI-2018,

ca 1100 m asl; Plátres, 26-XII-2009, 2-I-2010, 8-XII-2015, 1-XII-2017, 26-X-2018 & 3-XII-2018, ca 1200 m asl;

Pýrgos Tyllirías, 10-I-2011, ca 200 m asl; Amíantos, 1300 m asl; 21-X-2011; Akámas, 18-XII-2014, ca 200 m asl;

Karvounás, 13-XI-2018, ca 1300 m asl. Also reported by Viney (2005) in Áyios Amvrósios, Nov. 1999 & Jan. 2000,

and Loizides et al. (2011) from collections cited above. A collection reported by Nattrass (1937) as “Boletus boudieri

Quél.”, from Kórnos, 1931, probably also corresponds to this species.

Suillus collinitus (Fr.) Kuntze

Extremely common throughout the island, found in association with various pine species from the dunal zone, to the

peaks of Troodos. Dozens of sightings in Saittás, 17-XI-2007, 18-XII-2009, 28-X-2011, 18-XI-2016 & 3-XII-2018, ca

700 m asl; Kélefos, 11-XI-2007, 20-X-2009, 25-XI-2009, 2-XI-2012, 31-XII-2012, 2-XII-2014, 21-I-2015 & 1-I-2019,

…

Loizides, M.

ca 450–550 m asl; Plátres, 21-XI-2011, ca 1100–1200 m asl; Platánia, 10-X-2009, 16-XI-2014, 2-I-2015 & 16-XI-

2018, ca 1100 m asl; Aphrodite’s Rock, 2-II-2009, ca 100 m asl; Péra Pedí, 4-X-2009, ca 550 m asl; Amíantos, 10-X-

2009, ca 1300 m asl; Pýrgos Tyllirías, 10-I-2011, ca 200 m asl; Trimíklini, 10-XI-2012, ca 650 m asl; Soúni, 9-III-

2014, ca 400 m asl; Akrotíri, 24-XI-2014, ca 20 m asl; Akámas, 14-XII-2104 & 19-XII-2104, ca 200 m asl.

Tapinella panuoides (Fr.) E.-J. Gilbert

Reported by Loizides et al. (2011) as “Paxillus ionipus Quél.”. Occasionally seen on logs of P. nigra subsp. pallasiana

and P. brutia, with collections from Péra Pedí, 12-III-2008, ca 550 m asl; and Archimandrita, 16-II-2013, ca 450 m asl.

*Tephrocybe striipilea (Fr.) Donk

Two collections matching descriptions of this taxon in Cedar Valley, 18-XI-2009, ca 1000 m asl, in P. brutia and C.

brevifolia forest; and Platánia, 21-XII-2011, ca 1100 m asl, in P. brutia and C. brevifolia forest clearing. Rare.

*Tephroderma fuscopallens Musumeci & Contu

The monotypic genus Tephroderma Contu & Musumeci, was in recently proposed to accommodate T. fuscopallens, a

species originally collected from North France (Musumeci & Contu 2014) and subsequently documented in Turkey

(Sesli & Topçu Sesli 2016). One collection found among lichens in a mixed P. brutia/C. sempervirens/Cistus forest in

Episkopí, 26-I-2018, ca 150 m asl. Distribution unclear.

Tricholoma batschii Gulden ex Mort. Chr. & Noordel.

Previously reported by its synonym “T. fracticum (Britzelm.) Kreisel” in Viney (2005), and Loizides et al. (2011), this

widespread species appears in large numbers in Calabrian pine forests, e.g. Mandriá, 25-XI-2007, ca 700 m asl; Péra

Pedí, 31-XII-2007 & 10-I-2017, ca 550 m asl; Lysós, 20-I-2008, ca 600 m asl; Plátres, 22-XII-2009, 26-XII-2009, 2-I-

2010, 8-XII-2015, 1-XII-2017 & 3-XII-2018, ca 1200 m asl; Platánia, 3-XII-2012, ca 1100 m asl; Akámas, 14-XII-

2012, ca 200 m asl; Kélefos, 2-XII-2014, ca 450 m asl. See “Discussion” for further notes.

Tricholoma caligatum (Viv.) Ricken

A typically Mediterranean species, considered a delicacy and widely collected for pickling by the locals (Loizides 2008,

2011; Loizides et al. 2011). Found in various habitats across the Troodos mountain range, including mixed Calabrian

pine forests in Saittás, 20-XII-2007, ca 700 m asl; Plátres, 22-XII-2009, 26-XII-2009, 2-I-2010, 8-XII-2015, 1-XII-

2017 & 3-XII-2018, ca 1200 m asl; Platánia, 1-XII-2009, 3-XII-2009, 9-XII-2009, 30-XII-2011 & 16-XI-2018, ca 1100

m asl; Mésa Potamós, 20-XII-2009, 2-XII-2017 & 20-XII-2018, ca 800–900 m asl; Mávro Dásos, 1-XII-2012, ca 1000

m asl; Moniátis, 20-XII-2012, ca 800 m asl; Karvounás, 14-XI-2018, ca 1300 m asl. Common.

*Tricholoma chrysophyllum A. Riva, C.E. Hermos. & Jul. Sánchez

A handful of sightings matching the profile of this rarely reported Mediterranean taxon (Riva 1998), including a

collection from a mixed P. brutia/Q. alnifolia/A. andrachne forest in Platánia, 1-XII-2009, ca 1000 m asl; and

collections from P. brutia forests at Péra Pedí, 10-I-2017, ca 550 m asl. Less common than its lookalike T. equestre

(L.) P. Kumm., which is mostly seen in P. nigra subsp. pallasiana forests in Cyprus (Loizides et al. 2011).

Tricholoma squarrulosum Bres.

Previously regarded a variety of T. atrosquamosum Sacc. (and reported as such in Loizides 2011), this has now been

shown to be a phylogenetically distinct species (Heilmann-Clausen et al. 2017). Common in mixed P. brutia/Q.

alnifolia forests, e.g., in Platánia, 1-XII-2009, 3-XII-2009, 9-XII-2009 & 30-XII-2014, ca 1100 m asl; Plátres, 23-XII-

2009, 26-XII-2009, 17-XII-2012 & 12-XII-2015; or Mávro Dásos, 1-XII-2012, ca 1000 m asl.

Tricholoma sulphureum (Bull.) P. Kumm. sensu lato

Collections identified as this taxon formed multiple clades in the phylogenies of Heilmann-Clausen et al. (2017).

Probably more than one species of this cluster occur in Cyprus, including a collection from a mixed P. brutia/Q.

alnifolia/A. andrachne forest at Mésa Potamós, 10-I-2017, ca 800 m asl. A collection from a Q. alnifolia/P. brutia/A.

orientalis riparian forest reported under this name in Loizides (2011), appears to be morphologically distinct, as is an

odorless collection from a P. brutia forest in Kélefos, 20-XI-2019, ca 450 m asl. Widespread DNA sequencing and

appointment of neo/epitypes for T. sulphureum ss. str. are necessary before any new taxa can be proposed.

Tricholoma terreum (Schaeff.) P. Kumm.

Common in Cyprus, fruiting abundantly from the thermo-Mediterranean belt (Viney 2005; Loizides 2016) to the

subalpine peaks of Troodos (Loizides et al. 2011). Seen in Calabrian pine forests e.g. in Mandriá, 21-XI-2017, ca 700

m asl; Saittás, 29-XI-2007, 30-XII-2013 & 6-I-2017, ca 700 m asl; Péra Pedí, 31-XII-2007, 6-I-2011 & 10-I-2017, ca

550 m asl; Platánia, 22-XII-2007, 1-XII-2009, 3-XII-2009, 9-XII-2009 & 15-XII-2018, ca 1100 m asl; Plátres, 23-XII-

2009, ca 1200 m asl; Stavrós tis Psókas, 22-XI-2011, ca 1200 m asl; Mávro Dásos, 5-XII-2012, ca 1000 m asl; Panayiá,

23-I-2013, ca 1000 m asl; Kannavioú, 29-XII-2016, ca 900 m asl; Moniátis, 10-I-2017, ca 800 m asl.

*Tricholomopsis rutilans (Schaeff.) Singer

Known from a single basidiome found on a P. brutia stump at Platánia, 9-XII-2012, ca 1150 m asl. Rare.

Xerocomellus redeuilhii A.F.S. Taylor, U. Eberh., Simonini, Gelardi & Vizzini

Occasional, with several collections from lowland P. brutia forests. See Loizides et al. (2019) for collection details.

Basidiomycetes within Pinus brutia ecosystems in Cyprus

…

Xerocomellus chrysenteron (Bull.) Šutara

Just a handful of collections from P. nigra subsp. pallasiana forests, and a single collection from a mixed P. brutia/C.

brevifolia/Abies sp. forest at Cedar Valley, 4-IV-2016, ca 1100 m asl (Loizides et al. 2019). Rare in Cyprus.

Xerocomellus cisalpinus (Simonini, H. Ladurner & Peintner) Klofac

A widespread species in Europe but rare in Cyprus, known from a single collection in a mixed P. brutia, C. brevifolia

and Abies sp. stand at Cedar Valley, 15-VII-2012, ca 1000 m asl (Loizides et al. 2019).

Xeromphalina fellea Maire & Malençon sensu lato

Reported by Viney (2005) from Haléfka, Jan. 1999, but several Xeromphalina Kühner & Maire species are present on

the island, whose exact taxonomic position is not entirely clear and needs to be resolved by DNA sequencing.

Discussion

Clearly, the basidiomycete diversity present in P. brutia ecosystems is substantial, with 95 of the 231 taxa

included in the present checklist reported from Cyprus for the first time. Considering that an additional 22

aphyllophoraceous basidiomycetes were associated with P. brutia in a previous checklist (Loizides 2018), but

also that ascomycetes and several dozens of unidentified collections have been excluded, it becomes apparent

that the macromycete diversity in Calabrian pine ecosystems amounts to at least several hundreds of species.

Interestingly, only few taxa seem to exhibit a transcontinental distribution and have been molecularly verified

in more than one continent. These, include mostly opportunistic saprotrophs and wood decomposers such as

Agaricus bisporus, Coprinus comatus, Cystodermella cinnabarina, C. granulosa, Galerina marginata, Phallus

impudicus, or Rhodocollybia butyracea. Lepista nuda and L. sordida have also been reported from several

countries and continents, but appear to be polyphyletic and occupy multiple clades in phylogenetic trees

(Alvarado et al. 2015). Among ectomycorrhizal taxa, Tricholoma batschii has been molecularly confirmed in

both ends of the Atlantic and is widespread in Cyprus, but a number of lookalikes also occur in mixed pine/oak

ecosystems, whose exact taxonomic position is not yet entirely clear. Brown-capped Tricholoma species have

not been adequately sampled in Mediterranean ecosystems and the phylogenetic status of taxa like T.

cedretorum Bon ex A. Riva, T. ezcarayense C.E. Hermos. & Jul. Sánchez, T. striatum (Schaeff.) Quél., and T.

tridentinum Singer, is yet to be fully clarified (Heilmann-Clausen et al. 2017). The widespread distribution and

ecological plasticity of several Hebeloma species present in Cyprus is also notable, particularly H. sinapizans

and H. crustuliniforme, which display a pan-European distribution and are associated with dozens of tree-hosts,

as is H. subtorum whose distribution extends to North Africa (Beker et al. 2016). Few taxa of Inocybe are

reported from different continents (e.g., I. flocculosa, I. geophylla), but these probably involve multiple

biological species (Ryberg et al. 2008; Matheny & Sweney 2018). Similarly, extra-European reports of some

Russula Pers. taxa originally described from Europe (e.g., R. acrifolia, R. atropurpurea, R. foetens, R.

sanguinea), are in need of molecular verification.

Other than the few widely distributed species, the majority of taxa documented exhibit continental

endemism and some are even regional endemics narrowly restricted to the Mediterranean basin. At least one

species, the sequestrate Russula hobartiae, appears to be narrowly endemic in Cyprus and to the Troodos

massif, where it exclusively occurs at elevations above 1000 m (Vidal et al. 2019). The recently described

Clavulina iris and Hygrophorus meridionalis are apparently common and widespread, but again are confined

to southern latitudes and are both strongly associated with Mediterranean flora (Moreau et al. 2018; Crous et

al. 2019). Other typically Mediterranean species documented are Amanita gioiosa, A. ovoidea, A. proxima,

Leucopaxillus malenconii, Russula werneri, Suillus bellinii, Tricholoma caligatum and T. chrysophyllum, in

addition to the recently described Rhodocybe matesina and Tephroderma fuscopallens, whose

phylogeographical distribution is here extended to their second and third countries, respectively. Few other

taxa, such as Clitopilus daamsii, Hygrocybe calciphila, Hygrophorocybe nivea, Lepiota lepida, Leucoagaricus

georginae and Tephrocybe striaepilea are rare, and seldom appear in published literature.

A number of previous reports are considered doubtful or spurious and have been excluded. Chroogomphus

rutilus (Schaeff.) O.K. Mill., previously reported by both Viney (2005) and Loizides et al. (2011), does not

appear to occur on the island and most sequenced collections from Cyprus have turned out to belong to C.

mediterraneus (Scambler et al. 2018). A second species of Chroogomphus (Singer) O.K. Mill., C. subfulmineus

Niskanen et al., is indeed present οn the island, but appears to be confined to the higher eleveations of Troodos

…

Loizides, M.

and is so far known only from P. nigra subsp. pallasiana ecosystems. A collection previously reported as

“Ramaria bataillei” in Loizides et al. (2011), is now thought to represent a faded collection of R. rubella and

is also excluded, as is Entoloma bloxamii (Loizides 2011), whose collections from Kannavioú likely represent

the recently described E. atromadidum. A collection reported by Viney (2005) as “Suillus mediterraneensis

(Jacquet. & J. Blum) Redeuilh”, from Syrianóchori, Jan. 2001, is probably misidentified, since the image

provided appears to depict a typical S. bellini. Although the presence of S. mediterraneensis on the island has

been molecularly verified (unpubl. data), this species is rare and so far confined within coastal P. halepensis

plantations suggesting an anthropogenic introduction. Suillus granulatus (L.) Roussel, on the other hand,

reported by Viney (2005) from the same locality (Syrianóchori), does not occur on the island and the image

provided for this collection again appears to depict the widespread S. bellini. A collection from Áyios

Amvrósios, Dec. 2000, reported by Viney (2005) as “Hygrophorus latitabundus Britzelm.”, depicts a pale, dry

and probably unrelated fungus, while collections identified as “Rugosomyces chrysenteron (Bull.) Bon” are

more likely to represent its Mediterranean lookalike R. fallax Bon. Other doubtful records include Clitocybe

ditopa (Fr.) Gillet, Cystoderma amianthinum (Scop.) Fayod, Lyophyllum decastes, Macrolepiota konradii

(Huijsman ex P.D. Orton) M.M. Moser, Ripartites tricholoma (Alb. & Schwein.) P. Karst., Sericeomyces

serenus (Fr.) Heinem., and Tricholoma portentosum (Fr.) Quél., whose published images by Viney (2005) do

not resemble these species, nor have they been subsequently verified on the island during this 12-y-survey.

Although by no means exhaustive, the present inventory provides the first comprehensive account of

basidiomycete diversity in P. brutia ecosystems and a basis on which future phylotaxonomic investigations

and conservation policies can build. Since the majority of the taxa reported here are yet to be genetically tested

and several species-clusters remain unresolved, it is likely that the taxonomic status of some collections will

change in the future. Generic and infrageneric revisions are thus to be expected, particularly within critical

genera such as Clitocybe s.l., Lepista s.l., or Lyophyllum s.l., which in their current circumscription are not

monophyletic (Alvarado et al. 2015, 2018a & 2018b; Bellanger et al. 2015; Bellanger 2016). Clarification of

old binomials through DNA sequencing of type material or appointment of epitypes, is also necessary for

clarifying critical taxa currently encompassing multiple phylospecies. These include species-clusters currently

lumped under Amanita mairei, Clavulina coralloides, C. rugosa, Clitopilus geminus, Entoloma hirtipes,

Hydnellum ferrugineum, Hygrocybe conica and Tricholoma sulphureum, to name but a few (Olariaga et al.

2009; Lodge et al. 2015; Heilmann-Clausen et al. 2017; Loizides et al. 2018). More importantly, documenting

and understanding fungal diversity in neglected but ecologically important Mediterranean ecosystems has in

recent years become an urgent priority (Richard et al. 2011; Büntgen et al. 2015; Loizides et al. 2019a). Given

the accelerated climate warming and aridification trends in the eastern Mediterranean region (Giorgi and

Lionello 2008; Hoerling et al. 2012; Lelieveld et al. 2012; Zittis et al. 2015, 2018 & 2019; Lionello & Scarascia

2018), an enhanced understanding of the full spectrum of fungal diversity and plant-fungi interactions in

Mediterranean ecosystems will greatly improve our ability to predict shifts in biodiversity, identify urgent

conservation priorities and draft effective conservation policies. Towards this end, the documentation of fungal

diversity in other priority habitats in Cyprus, such as Pinus nigra subsp. pallasiana, Quercus alnifolia, and

Cedrus brevifolia ecosystems, as well as coastal dunes and halophytic wetlands, will follow in future dedicated

checklists.

Acknowledgments

I am grateful to Elias Polemis, Juan-Carlos Zamora and Carmel Sammut for critically reviewing and improving the

manuscript prior to submission. Thanks to María P. Martín for the sequencing and identification of Rhizopogon graveolens f.

pomaceus; to Pablo Alvarado (ALVALAB) for sequencing collections cited in this work; to Pierre-Arthur Moreau and Jean-

Michel Bellenager for taxonomic and phylogenetic advice; and to Caroline Hobart for her help and friendly support. Thanks

also to Theologis Alexandrides and Christos Attashis for providing collections and information.

Basidiomycetes within Pinus brutia ecosystems in Cyprus

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Loizides, M.

Basidiomycetes within Pinus brutia ecosystems in Cyprus

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Loizides, M.

Basidiomycetes within Pinus brutia ecosystems in Cyprus

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Loizides, M.

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Jan 2024
STUD MYCOL

The phylogenetic position of several clitocyboid/pleurotoid/tricholomatoid genera previously considered incertae sedis is here resolved using an updated 6-gene dataset of Agaricales including newly sequenced lineages and more complete data from those already analyzed before. Results allowed to infer new phylogenetic relationships, and propose taxonomic novelties to accommodate them, including up to ten new families and a new suborder. Giacomia (for which a new species from China is here described) forms a monophyletic clade with Melanoleuca (Melanoleucaceae) nested inside suborder Pluteineae, together with the families Pluteaceae, Amanitaceae (including Leucocortinarius), Limnoperdaceae and Volvariellaceae. The recently described family Asproinocybaceae is shown to be a later synonym of Lyophyllaceae (which includes also Omphaliaster and Trichocybe) within suborder Tricholomatineae. The families Biannulariaceae, Callistosporiaceae, Clitocybaceae, Fayodiaceae, Macrocystidiaceae (which includes Pseudoclitopilus), Entolomataceae, Pseudoclitocybaceae (which includes Aspropaxillus), Omphalinaceae (Infundibulicybe and Omphalina) and the new families Paralepistaceae and Pseudoomphalinaceae belong also to Tricholomatineae. The delimitation of the suborder Pleurotineae (= Schizophyllineae) is discussed and revised, accepting five distinct families within it, viz. Pleurotaceae, Cyphellopsidaceae, Fistulinaceae, Resupinataceae and Schizophyllaceae. The recently proposed suborder Phyllotopsidineae (= Sarcomyxineae) is found to encompass the families Aphroditeolaceae, Pterulaceae, Phyllotopsidaceae, Radulomycetaceae, Sarcomyxaceae (which includes Tectella), and Stephanosporaceae, all of them unrelated to Pleurotaceae (suborder Pleurotineae) or Typhulaceae (suborder Typhulineae). The new family Xeromphalinaceae, encompassing the genera Xeromphalina and Heimiomyces, is proposed within Marasmiineae. The suborder Hygrophorineae is here reorganized into the families Hygrophoraceae, Cantharellulaceae, Cuphophyllaceae, Hygrocybaceae and Lichenomphaliaceae, to homogenize the taxonomic rank of the main clades inside all suborders of Agaricales. Finally, the genus Hygrophorocybe is shown to represent a distinct clade inside Cuphophyllaceae, and the new combination H. carolinensis is proposed.

First record of rare boreal-alpine waxcap Hygrocybe calciphila (Hygrophoraceae, Agaricales) in Southeastern Europe

Article

Nov 2021

Hygrocybe calciphila, an uncommon species of the genus was found for the first time in Southeastern Europe in forests of Pinus heldreichii in Bulgaria. Description and illustrations of the finding are presented. The European distribution of the species is summarized and briefly discussed, with emphasis on the scarce previous findings in Southern Europe.

Fungal Planet description sheets: 951–1041

Article

Full-text available

Dec 2019

Novel species of fungi described in this study include those from various countries as follows: Antarctica , Apenidiella antarctica from permafrost, Cladosporium fildesense from an unidentified marine sponge. Argentina , Geastrum wrightii on humus in mixed forest. Australia , Golovinomyces glandulariae on Glandularia aristigera , Neoanungitea eucalyptorum on leaves of Eucalyptus grandis , Teratosphaeria corymbiicola on leaves of Corymbia ficifolia , Xylaria eucalypti on leaves of Eucalyptus radiata . Brazil , Bovista psammophila on soil, Fusarium awaxy on rotten stalks of Zea mays , Geastrum lanuginosum on leaf litter covered soil, Hermetothecium mikaniae-micranthae (incl. Hermetothecium gen. nov.) on Mikania micrantha , Penicillium reconvexovelosoi in soil, Stagonosporopsis vannaccii from pod of Glycine max . British Virgin Isles , Lactifluus guanensis on soil. Canada , Sorocybe oblongispora on resin of Picea rubens . Chile , Colletotrichum roseum on leaves of Lapageria rosea . China , Setophoma caverna from carbonatite in Karst cave. Colombia , Lareunionomyces eucalypticola on leaves of Eucalyptus grandis . Costa Rica , Psathyrella pivae on wood. Cyprus , Clavulina iris on calcareous substrate. France , Chromosera ambigua and Clavulina iris var. occidentalis on soil. French West Indies , Helminthosphaeria hispidissima on dead wood. Guatemala , Talaromyces guatemalensis in soil. Malaysia , Neotracylla pini (incl. Tracyllales ord. nov. and Neotracylla gen. nov.) and Vermiculariopsiella pini on needles of Pinus tecunumanii . New Zealand , Neoconiothyrium viticola on stems of Vitis vinifera , Parafenestella pittospori on Pittosporum tenuifolium , Pilidium novae-zelandiae on Phoenix sp. Pakistan , Russula quercus-floribundae on forest floor. Portugal , Trichoderma aestuarinum from saline water. Russia , Pluteus liliputianus on fallen branch of deciduous tree, Pluteus spurius on decaying deciduous wood or soil. South Africa , Alloconiothyrium encephalarti , Phyllosticta encephalarticola and Neothyrostroma encephalarti (incl. Neothyrostroma gen. nov.) on leaves of Encephalartos sp., Chalara eucalypticola on leaf spots of Eucalyptus grandis × urophylla , Clypeosphaeria oleae on leaves of Olea capensis , Cylindrocladiella postalofficium on leaf litter of Sideroxylon inerme , Cylindromonium eugeniicola (incl. Cylindromonium gen. nov.) on leaf litter of Eugenia capensis , Cyphellophora goniomatis on leaves of Gonioma kamassi , Nothodactylaria nephrolepidis (incl. Nothodactylaria gen. nov. and Nothodactylariaceae fam. nov.) on leaves of Nephrolepis exaltata , Falcocladium eucalypti and Gyrothrix eucalypti on leaves of Eucalyptus sp., Gyrothrix oleae on leaves of Olea capensis subsp. macrocarpa , Harzia metrosideri on leaf litter of Metrosideros sp., Hippopotamyces phragmitis (incl. Hippopotamyces gen. nov.) on leaves of Phragmites australis , Lectera philenopterae on Philenoptera violacea , Leptosillia mayteni on leaves of Maytenus heterophylla , Lithohypha aloicola and Neoplatysporoides aloes on leaves of Aloe sp., Millesimomyces rhoicissi (incl. Millesimomyces gen. nov.) on leaves of Rhoicissus digitata , Neodevriesia strelitziicola on leaf litter of Strelitzia nicolai , Neokirramyces syzygii (incl. Neokirramyces gen. nov.) on leaf spots of Syzygium sp., Nothoramichloridium perseae (incl. Nothoramichloridium gen. nov. and Anungitiomycetaceae fam. nov.) on leaves of Persea americana , Paramycosphaerella watsoniae on leaf spots of Watsonia sp., Penicillium cuddlyae from dog food, Podocarpomyces knysnanus (incl. Podocarpomyces gen. nov.) on leaves of Podocarpus falcatus , Pseudocercospora heteropyxidicola on leaf spots of Heteropyxis natalensis , Pseudopenidiella podocarpi , Scolecobasidium podocarpi and Ceramothyrium podocarpicola on leaves of Podocarpus latifolius , Scolecobasidium blechni on leaves of Blechnum capense , Stomiopeltis syzygii on leaves of Syzygium chordatum , Strelitziomyces knysnanus (incl. Strelitziomyces gen. nov.) on leaves of Strelitzia alba , Talaromyces clemensii from rotting wood in goldmine, Verrucocladosporium visseri on Carpobrotus edulis . Spain , Boletopsis mediterraneensis on soil, Calycina cortegadensisi on a living twig of Castanea sativa , Emmonsiellopsis tuberculata in fluvial sediments, Mollisia cortegadensis on dead attached twig of Quercus robur , Psathyrella ovispora on soil, Pseudobeltrania lauri on leaf litter of Laurus azorica , Terfezia dunensis in soil, Tuber lucentum in soil, Venturia submersa on submerged plant debris. Thailand , Cordyceps jakajanicola on cicada nymph, Cordyceps kuiburiensis on spider, Distoseptispora caricis on leaves of Carex sp., Ophiocordyceps khonkaenensis on cicada nymph. USA , Cytosporella juncicola and Davidiellomyces juncicola on culms of Juncus effusus , Monochaetia massachusettsianum from air sample, Neohelicomyces melaleucae and Periconia neobrittanica on leaves of Melaleuca styphelioides × lanceolata , Pseudocamarosporium eucalypti on leaves of Eucalyptus sp., Pseudogymnoascus lindneri from sediment in a mine, Pseudogymnoascus turneri from sediment in a railroad tunnel, Pulchroboletus sclerotiorum on soil, Zygosporium pseudomasonii on leaf of Serenoa repens . Vietnam , Boletus candidissimus and Veloporphyrellus vulpinus on soil. Morphological and culture characteristics are supported by DNA barcodes.

A multi-model, multi-scenario, and multi-domain analysis of regional climate projections for the Mediterranean

Article

Full-text available

Dec 2019
REG ENVIRON CHANGE

Observation and model-based studies have identified the Mediterranean region as one of the most prominent climate change “hot-spots.” Parts of this distinctive region are included in several Coordinated Regional Downscaling Experiment (CORDEX) domains such as those for Europe, Africa, the Mediterranean, and the Middle East/North Africa. In this study, we compile and analyze monthly temperature and precipitation fields derived from regional climate model simulations performed over different CORDEX domains at a spatial resolution of 50 km. This unique multi-model, multi-scenario, and multi-domain “super-ensemble” is used to update projected changes for the Mediterranean region. The statistical robustness and significance of the climate change signal is assessed. By considering information from more than one CORDEX domains, our analysis addresses an additional type of uncertainty that is often neglected and is related to the positioning of the regional climate model domain. CORDEX simulations suggest a general warming by the end of the century (between 1 and 5 °C with respect to the 1986–2005 reference period), which is expected to be strongest during summer (up to 7 °C). A general drying (between 10 and 40%) is also inferred for the Mediterranean. However, the projected precipitation change signal is less significant and less robust. The CORDEX ensemble corroborates the fact that the Mediterranean is already entering the 1.5 °C climate warming era. It is expected to reach 2 °C warming well within two decades, unless strong greenhouse gas concentration reductions are implemented. The southern part of the Mediterranean is expected to be impacted most strongly since the CORDEX ensemble suggests substantial combined warming and drying, particularly for pathways RCP4.5 and RCP8.5.

Reassessment of the generic limits for Hydnellum and Sarcodon (Thelephorales, Basidiomycota)

Article

Full-text available

Jun 2019

DNA sequences from the nuclear LSU and ITS regions were used for phylogenetic analyses of Thelephorales with a focus on the stipitate hydnoid genera Hydnellum and Sarcodon. Analyses showed that Hydnellum and Sarcodon are distinct genera but that the current division, based on basidioma texture, makes Sarcodon paraphyletic with respect to Hydnellum. In order to make genera monophyletic several species are moved from Sarcodon to Hydnellum and the following new combinations are made: Hydnellumamygdaliolens, H.fennicum, H.fuligineoviolaceum, H.fuscoindicum, H.glaucopus, H.joeides, H.lepidum, H.lundellii, H.martioflavum, H.scabrosum, H.underwoodii, and H.versipelle. Basidiospore size seems to separate the genera in most cases. Hydnellum species have basidiospore lengths in the range 4.45-6.95 µm while the corresponding range for Sarcodon is 7.4-9 µm. S.quercinofibulatus deviates from this pattern with an average spore length around 6 µm. Neotropical Sarcodon species represent a separate evolutionary lineage.

Phylogenetic and distributional data on boletoid fungi (Boletaceae) in Cyprus and description of a new sampling methodology

Article

Full-text available

Jun 2019

The data presented here was obtained during a decade-long macromycete inventory on the island of Cyprus and is supplementary to the research article “Present status and future of boletoid fungi (Boletaceae) on the island of Cyprus: cryptic and threatened diversity unravelled by ten-year study”. A new, rainfall-based sampling protocol for documenting fungal diversity in Mediterranean ecosystems, is described in detail. Keywords: Boletales, Boletus, Fungi, Island biogeography, Inventory, Mediterranean

Present status and future of boletoid fungi (Boletaceae) on the island of Cyprus: Cryptic and threatened diversity unravelled by ten-year study

Article

Full-text available

May 2019
FUNGAL ECOL

Fungi historically placed in the iconic genera of Boletus, Leccinum and Xerocomus have been the subject of major taxonomic revisions in recent years. Yet, despite all advancements in systematics, boletoid fungi in insular ecosystems remain little explored and our knowledge of their diversity, distribution and abundance in Mediterranean ecoregions is far from complete. To shed light on this blind spot, the findings of a ten-year study from the island of Cyprus were analysed, integrating phylogenetic, ecological, morphological , phenological and chorological data. An unexpected diversity of Boletaceae fungi is unveiled, with twenty-five species phylogenetically confirmed to be present on the island, thirteen of them previously unreported. Sequencing of the ITS rDNA region, reveals crypticism within the Butyriboletus fechtneri, Caloboletus radicans, Rubroboletus lupinus and Rheubarbariboletus armeniacus species-complexes and infrageneric relationships are discussed. A strong link between boletoid fungi and Mediterranean oaks of the ilicoid group (Quercus alnifolia, Quercus coccifera subsp. calliprinos) is illustrated, with nineteen species (76%) overall found to be strictly or broadly associated with evergreen oaks. In stark contrast, the semi-deciduous Quercus infectoria subsp. veneris appears to be an unfavorable host for boletoid fungi, with just a single associated species so far. Phenological and chorological records indicate that most species on the island are rare, highly localized and fruit during very brief spells several years apart, mainly in response to increased annual, late summer or early autumn precipitation. The conservation status of these high-profile fungi is hence discussed, particularly in view of alarming climate changes, forecasted to have a dramatic impact on Mediterranean ecosystems in the years to come.

Looking for Lepiota psalion Huijser & Vellinga (Agaricales, Agaricaceae)

Article

Full-text available

May 2019

Lepiota psalion is fully described based on a recent collection from Sardinia (Italy) and the holotype. NrITS-and nrLSU-based phylogeny demonstrates that sequences deposited in GenBank as "L. psalion" and generated from two Dutch and one Chinese collections are not conspecific with the holotype and represent two distinct, undescribed species. These species are here proposed as Lepiota recondita sp. nov. and Lepiota sinorecondita ad int.

A phylogenetic and taxonomic revision of sequestrate Russulaceae in Mediterranean and temperate Europe

Article

Full-text available

Apr 2019
Persoonia

A comprehensive morphological and genetic study of type material and new collections of sequestrate Russulales species formerly belonging to the genera Arcangeliella, Elasmomyces, Gymnomyces, Hydnangium, Hymenogaster, Macowanites, Martellia, Secotium and Zelleromyces is here undertaken, for the purpose of providing a complete taxonomical revision of sequestrate Russulaceae species in the Mediterranean and temperate regions of Europe. As a result, seven distinct taxa in the genus Lactarius and 18 in the genus Russula are identified. Six of them are new species: L. populicola, L. subgiennensis, R. bavarica, R. candidissima, R. hobartiae and R. mediterraneensis, and seven represent new combinations: L. josserandii (≡ Zelleromyces josserandii), L. soehneri (≡ Hydnangium soehneri), R. candida (≡ Hydnangium candidum), R. cerea (≡ Hydnangium cereum), R. messapica var. messapicoides (≡ Macowanites messapicoides), R. meridionalis (≡ Zelleromyces meridionalis) and R. neuhoffii (≡ Hydnangium neuhoffii). Twenty-two of the 25 taxa are illustrated, while descriptions, microscopy images, as well as extensive information on the ecology, chorology and phylogeny for all taxa are provided. A key is further included to facilitate their identification.

Preliminary phylogenetic investigations into the genus Amanita (Agaricales) in Cyprus, with a review of previous records and poisoning incidents

Article

Full-text available

Oct 2018

Early reports on the occurrance of Amanita species in Cyprus, particularly A. muscaria and A. phalloides, remained highly controversial for nearly a century. Published literature dating as far back as the 1930s, implicating these two species in a series of fatal poisonings among the local population, based their assumptions on unconfirmed sightings and post-mortem findings, but provided no detailed descriptions or exsiccatae. Subsequent citations of these early reports by later authors, as well as more recently published doubious records, have added to the confusion. To clarify this longstanding controvercy, a preliminary phylogenetic investigation into the genus Amanita in Cyprus has been carried out, based on ITS rDNA sequencing of representative collections gathered over the past decade. At least thirteen Amanita species are phylogenetically confirmed to be present on the island, including three species in sect. Vaginatae tentatively interpreted as new to science. Another four species, namely A. citrina, A. gioiosa, A. lividopallescens and A. vittadinii, are reported from Cyprus for the first time. Although the presence of A. phalloides is confirmed following a single collection from a Corylus avellana plantation, our data suggests this is an extremely rare and probably introduced species in Cyprus, unlikely to have caused the 1930s fatalities. Formerly published records and poisoning episodes are reviewed and critically discussed, including reports of A. muscaria and A. virosa, which are shown to be spurious.

Diversity of Chroogomphus (Gomphidiaceae, Boletales) in Europe, and typification of C. rutilus

Article

Full-text available

Sep 2018

In this study, eight species of Chroogomphus are recognized from Europe: C. britannicus, C. aff. liformis 1, C. fulmineus, C. cf. helveticus, C. mediterraneus, C. cf. purpurascens, C. rutilus, and C. subfulmineus. Different candidates for the application of the name C. rutilus are evaluated and the best fit to the description is selected; lecto- and epitypes are chosen to fix the name. Chroogomphus fulmineus and C. mediterraneus are also epitypified and a new species, C. subfulmineus, is described. The infrageneric classification is revised and a new subgenus Siccigomphus and three new sections, Confusi, Filiformes, and Fulminei are introduced. The former sections Chroogomphus and Floccigomphus are elevated to subgeneric level. Comparison of the ITS regions (nuc rDNA ITS1-5.8S-ITS2) of all species studied shows that there is a minimum interspecific difference of 1.5 %, with the exception of the two species belonging to sect. Fulminei which differ by a minimum of 0.9 %. Ecological specimen data indicate that species of Chroogomphus form basidiomes under members of Pinaceae, with a general preference for species of Pinus. Five European species have been recorded under Picea, while Abies and Larix have also been recorded as tree associates, although the detailed nutritional relationships of the genus, involving other suilloid fungi in particular, have yet to be fully clarified.

Genera of Inocybaceae: New skin for the old ceremony

Article

Dec 2019

A six-gene phylogeny of the Inocybaceae is presented to address classification of major clades within the family. Seven genera are recognized that establish a global overview of phylogenetic relationships in the Inocybaceae. Two genera—Nothocybe and Pseudosperma—are described as new. Two subgenera of Inocybe—subg. Inosperma and subg. Mallocybe—are elevated to generic rank. These four new genera, together with the previously described Auritella, Tubariomyces, and now Inocybe sensu stricto, constitute the Inocybaceae, an ectomycorrhizal lineage of Agaricales that associates with at least 23 plant families worldwide. Pseudosperma, Nothocybe, and Inocybe are recovered as a strongly supported inclusive clade within the family. The genus Nothocybe, represented by a single species from tropical India, is strongly supported as the sister lineage to Inocybe, a hyperdiverse genus containing hundreds of species and global distribution. Two additional inclusive clades, including Inosperma, Tubariomyces, Auritella, and Mallocybe, and a nested grouping of Auritella, Mallocybe, and Tubariomyces, are recovered but with marginal statistical support. Overall, the six-gene data set provides a more robust phylogenetic estimate of relationships within the family than do single-gene and single-gene-region estimates. In addition, Africa, India, and Australia are characterized by the most genera in the family, with South America containing the fewest number of genera. A total of 180 names are recombined or proposed as new in Inosperma, Mallocybe, and Pseudosperma. A key to genera of Inocybaceae is provided.

Basidiomycete diversity within Calabrian pine (Pinus brutia) ecosystems on the island of Cyprus

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