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Type studies in Hebeloma. Unravelling a taxonomic-nomenclatural problem: what is Hebeloma leucosarx?

January 2007

January 2007
22(2):133-176

Authors:

Abstract: The type collections of Hebeloma leucosarx, H. velutipes, H. favrei and H. lutense, as well as some additional original and historically-relevant material were studied. The type of H. leucosarx turned out to contain parts belonging to two different taxa; accordingly, it represents a case of admixture. One element is characterized by clavatocylindraceous (rarely subcapitato-cylindraceous) cheilocystidia, the other by clavato- to capitato/swollen-pedicellate cheilocystidia, with pedicel typically sinuous. Sporal traits, however useful, do not seem to play a major role in the separation of the two elements. H. velutipes and H. favrei appear to be morphologically very similar to the first element, while H. lutense, to the second. In conformity with the ICBN, the first element is designated as the lectotype to which the epithet leucosarx must be attached. All the taxa presented are illustrated by micrographs, H. leucosarx and H. lutense are also illustrated by colour photographs. The present study and others in preparation are meant to lay the morphological groundwork for a collaborative research tackling also H. leucosarx and its numerous satellite taxa from a molecular perspective.

Hebeloma leucosarx. Holotype, P.D.Orton 583 (K). On le left " Specimen 1 " (lectotypus), on the right " Specimen 2 " (H. lutense). Bar 2 cm.

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H. leucosarx. Holotype, "Specimen 1". Spores & cheilocystidia. Bar 10 µm (spores); bar 30 µm (cheilocystidia).

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H. leucosarx. Holotype. Marginal pileocystidia, " specimen 1 " (top), " specimen 2 " (bottom). Bar 30 µm.

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H. leucosarx. P. D. Orton 593 (E). Spores & cheilocystidia. Bar 10 µm (spores); bar 30 µm (cheilocystidia).

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+10

H. leucosarx. M. Moser 81/424 (IB). Spores & cheilocystidia. Bar 10 µm (spores); bar 30 µm (cheilocystidia).

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Manoscritto ricevuto il 12.12.2007

TYPE STUDIES IN HEBELOMA.

UNRAVELLING A TAXONOMIC-NOMENCLATURAL TANGLE:

WHAT IS HEBELOMA LEUCOSARX ?

Edmondo Grilli

Via dei Pioppi, 2

I-65026 POPOLI (PE)

E-mail: grillie@alice.it

Grilli E., 2007: Studio dei tipi nel genere Hebeloma. Proposta di soluzione di un garbu-

glio tassonomico. Che cosa è Hebeloma leucosarx?

Key words: /hebelomatoid clade, Hebeloma subsect. Velutipes, H. leucosarx, H. favrei, H.

lutense, H. velutipes, revisionary taxonomy.

Abstract: The type collections of Hebeloma leucosarx, H. velutipes, H. favrei and H.

lutense, as well as some additional original and historically-relevant material were stud-

ied. The type of H. leucosarx turned out to contain parts belonging to two different taxa;

accordingly, it represents a case of admixture. One element is characterized by clavato-

cylindraceous (rarely subcapitato-cylindraceous) cheilocystidia, the other by clavato- to

capitato/swollen-pedicellate cheilocystidia, with pedicel typically sinuous. Sporal traits,

however useful, do not seem to play a major role in the separation of the two elements. H.

velutipes and H. favrei appear to be morphologically very similar to the first element,

while H. lutense, to the second. In conformity with the ICBN, the first element is desig-

nated as the lectotype to which the epithet leucosarx must be attached. All the taxa pre-

sented are illustrated by micrographs, H. leucosarx and H. lutense are also illustrated by

colour photographs.

The present study and others in preparation are meant to lay the morphological ground-

work for a collaborative research tackling also H. leucosarx and its numerous satellite

taxa from a molecular perspective.

Riassunto: Le raccolte tipo di Hebeloma leucosarx, H. velutipes, H. favrei e H. lutense,

così come materiale aggiuntivo costituito da raccolte originali o rilevanti da un punto di

vista storico-tassonomico, sono state studiate. Il typus di H. leucosarx si è rivelato conte-

nere due taxa diversi e quindi rappresenta un caso di commistione (raccolta mista). Uno

dei due elementi è caratterizzato da cheilocistidi clavato-cilindracei (raramente subcapita-

to-cilindracei), l’altro da cheilocistidi clavato- a capitato/rigonfio-pedicellati, con pedicel-

lo tipicamente sinuoso. I caratteri sporali, per quanto utili, sembrano avere soltanto una

funzione di supporto nella separazione dei due elementi. H. velutipes ed H. favrei appaio-

no morfologicamente molto simili al primo elemento, mentre H. lutense, al secondo. In

conformità alle regole dell’ ICBN, il primo elemento è designato come lectotipo al quale

l’epiteto leucosarx deve essere collegato. Tutti i taxa trattati sono illustrati da micrografie,

H. leucosarx ed H. lutense sono rappresentati anche da fotografie a colori.

Micol. e Veget. Medit., 22 (2): 133-176. 2007

134 Grilli E.

Il presente lavoro ed altri in preparazione intendono fornire le necessarie premesse mor-

fologiche per uno studio molecolare in collaborazione riguardante, tra l’altro, H. leucosarx

ed i suoi numerosi taxa satellite.

INTRODUCTION

This contribution, as well as some

companion articles to follow, expands on

the problem touched on in GRILLI (2004)

while dealing with H. bulbiferum Maire,

specifically, the fact that H. crustulini-

forme var. bulbiferum (Maire) J. Favre

1960 (= Hebeloma favrei Romagn. &

Quadr. 1984, ? H. bulbosum Romagn.

1983) shares its diagnostic microanatomi-

cal traits with numerous earlier taxa. Asit-

uation which demands careful considera-

tion for the possibility that they refer to

one and the same taxon, or alternatively

that at least some of them are autonomous

but very close and insufficiently circum-

scribed taxa. In particular, the present

paper gets down to specifics regarding the

circumstance – already hinted at in

VESTERHOLT (2000; 2005) and GRILLI

(2004) – that the type material of H. leu-

cosarx P. D. Orton (ORTON, 1960), a taxon

central to the problem in hand, proved to

be mixed. On revising it some years ago,

it was, in fact, immediately apparent –

from an alpha taxonomist’s point of view

at least – that there were two elements

involved, one with clavato-cylindraceous

cheilocystidia which, as shall be seen

below, has generally been referred to as H.

leucosarx ss Vesterholt and more recently

as H. velutipes Bruchet, the other, with

clavato- to capitato/swollen-pedicellate

cheilocystidia with pedicel typically sinu-

ous, which, conversely, can be referred to

H. lutense Romagn. A circumstance

which, incidentally, provided a reasonable

explanation for the conflicting interpreta-

tions of the taxon offered by specialists in

the genus like BRUCHET (1970),

QUADRACCIA (1993), BOEKHOUT &

KUYPER (1995) on one side, and

VESTERHOLT (1992; 1995; 2000; 2005) on

the other; interpretations that have coex-

isted for some time leading to consider-

able confusion and recently to the explicit

rejection of H. leucosarx (AANEN &

KUYPER, 2004).

In light of some surprising results of

recent molecular research (AANEN &

KUYPER, 2004; BOYLE & al, 2006), the aim

of this study was to present a tentative

type-based morphological unravelling of

some of the many strands that enter into

the taxonomic tangle evoked in the title in

order to (i) provide data on which to base

an informed choice among competing

names, and (ii) present a morphologically

well-grounded starting point for a

DNA-based approach. Hopefully, this

could contribute to the reduction of the

risk of misidentifications, a problem with

which, apparently, public DNA databases

are presently plagued (BRIDGE & al,

2003; VILGALYS, 2003; BRIDGE & al,

2004; but see HOLST-JENSEN & al, 2004;

HAWKSWORTH, 2004;).

Recognition criteria based on mating

experiments (biological species concept)

and molecular phylogeny (phylogenetic

species concept) are powerful tools which

illuminate relationships which would oth-

erwise remain cryptic with traditional

methods (TAYLOR & al, 2000; AGAPOW &

al, 2004) and are having a tremendous

impact on taxonomy and systematics, but

taxonomic knowledge remains a crucial

requirement for both the reliable identifi-

cation of the collections used to generate

sequences for phylogenetic studies

(PEINTNER & al, 2003) and as a practical

means of fungal identification in general

(descriptive taxonomy), especially in view

of the prospective loss of biodiversity on a

global scale and the need to document and

135

understand it (WHEELER, 2004; WHEELER

& al, 2004; KORF, 2005).

The traditional morphology-based

inclusion of the genus Hebeloma

(Agaricales, Agaricomycetidae, Basidio-

mycota) in the family Cortinariaceae R.

Heim ex Pouzar (KÜHNER, 1980; MOSER,

1983; SINGER, 1986; BAS, 1988) has lately

been questioned on the basis of molecular

evidence (MONCALVO & al, 2000;

MONCALVO & al, 2002) and KIRK & al,

(2001) soon incorporated it in their classi-

fication scheme, transferring the genus

into the family Bolbitiaceae Singer.

However, the real family affinities of the

genus are still in a state of flux, because if

the close kinship of Hebeloma with

Alnicola Kühner and Anamika K.A.

Thomas, Peintner, M. M. Moser &

Manim., among the agaricoid genera, and

some species of Hymenogaster Vittad.,

among the sequestrate ones, (AANEN & al,

2000; PEINTNER & al, 2001; THOMAS & al,

2002; FRANCIS & BOUGHER, 2002; YANG &

al, 2005), is widely recognized, PEINTNER

& al (2001) still admit that based on

sequence data “a close relationship”

between Cortinarius and Hebeloma can-

not be excluded, while in a multilocus

phylogenetic overview of the Agaricales

by MATHENY & al (2006) Hebeloma clus-

ters in the family Hymenogastraceae

Vittad., in a subclade with Anamika and

Alnicola - having Galerina,Phaeocollybia

and Psilocybe as a sister group – and at a

significant remove from the

Cortinariaceae and Bolbitiaceae clades.

Since the problem warrants much more

phylogenetic scrutiny, for the time being it

is better to err on the side of caution and

opt for an informal clade-based classifica-

tion.

According to my unpublished type

studies, H. bulbiferum ss Favre, as pre-

sented, discussed and illustrated in GRILLI

(2004), exhibits shared core micro-

anatomical features with, in order of

chronological priority, Hebeloma longi-

caudum (Pers.: Fr.) P. Kumm. 1871 (as

interpreted by FAVRE (1948), BRUCHET

(1970), KEIZER & ARNOLDS (1995) and

probably also LANGE (1938) and KONRAD

& MAUBLANC (1924-1937) - but this is

only an educated guess because the last

two authors do not seem to have left any

voucher material – H. bakeri Earle 1902,

H. exiguifolium Murrill 1917, H. bulbosum

Bres. ined. (in sched.), H. leucosarx P. D.

Orton 1960 p. p., H. leucosarx P. D. Orton

(as interpreted by VESTERHOLT, 1992;

1995; 2000), Hebeloma crustuliniforme

(Bull.) Quél. 1872 (as interpreted by

BRUCHET, 1970), H. velutipes Bruchet

1970, H. crustuliniforme var. minus Cooke

em. Rea (as interpreted by BOHUS, 1972),

H. favrei Romagn. & Quadr. 1984, H.

tenuifolium Romagn. 1985 (= H. angusti-

folium Romagn. 1983, ? = H. angustifoli-

um Britzelm. 1894), Hebeloma stenocystis

Favre ex Quadr. 1988, and, to a much less-

er extent, with H. claviceps (Fr.) P. Kumm.

1871, [as interpreted by Moser (KELLER &

MOSER, 2001)], H. stenocystis Favre 1960

inv., H. incarnatulum A. H. Sm. 1984

(= H. bryogenes Vesterh. 1993), H. calvini

Hesler & A.H. Sm. 1984 (= H. crustulini-

forme f. sphagnophilum Kauffman 1918),

morphologically separable from the

above-mentioned taxa on account of the

narrower cystidia, and with H. crustulini-

forme var. brevipes Murril 1946 and H.

subconcolor Bruchet 1970 which, in turn,

differ at least in field characters.

The above long list might also

include H. subtestaceum (Batsch 1789)

Kuyper 1986 non Murrill 1917, as sug-

gested by VESTERHOLT (2000; 2005), and

H. bulbosum Fayod 1892. However, apart

from the fact that the former is legally

unavailable, the interpretation of both

taxa based solely on the information

elicited from the relative protologues

(and the iconotype for H. subtestaceum)

is destined to remain largely speculative

What is Hebeloma leucosarx ?

136 Grilli E.

and consequently easily challenged by

disagreeing researchers.

Some of the above putative syn-

onymies have already been suggested for

H. leucosarx by VESTERHOLT (1992: H.

longicaudum; 1995 and 2000: H. velutipes,

H. favrei) , and LEGON & HENRICI (2005:

H. velutipes); for H. velutipes by AANEN

& KUYPER (2004: H. longicaudum, H.

bulbiferum ss Favre, H. bulbosum and

H. favrei), while P. Sammler & F. Gröger

(in KREISEL, 1987) and KELLER & MOSER

(2001) cite H. crustuliniforme ss Bruchet

and H. velutipes as possible synonyms of

H. claviceps.

MATERIAL AND METHODS

This contribution is based on the study

of about 40 collections, including the type

material of all the taxa discussed.

The presentations of quantitative data

as well as the methods of observations are

the same as those standardized in GRILLI

(2004; 2005; 2006). In the species under

dicussion, cheilocystidia are of the long,

large-sized type and, in variable propor-

tions, dmorphic, i.e. more or less clavate

and lageniform. Subtle variations in their

general morphology, apparently too slight

and hardly significant for the untrained

eye are, as learnt from experience, diag-

nostic. The following descriptive termi-

nology was used to cover the taxonomi-

cally useful variations observed in the

clavate and in the lageniform morphology

respectively:

clavato-cylindraceous: body of the

cystidium comparatively thick (mean

median breadth ≥ 4.0 µm) with more or

less parallel sides and apex progressively

enlarged into a club; the degree of apical

inflation may be specified by pre-modify-

ing adverbs: i.e. narrowly, broadly; apex at

times also dichotomously forked (epi-

physiform or bifid); measures refer to total

length, head and median body breadth;

capitato-cylindraceous: as above, but

apex abruptly swollen into a more or less

spherical head; measures as above;

clavato-pedicellate: body of the cystid-

ium comparatively slender (mean median

breadth ≤4.0 µm) with sides tapering

towards the base and apex progressively

enlarged into a club. The pedicel may be

more or less straight, or typically sinuous

or even tortuous; measures as above;

swollen-pedicellate: as above but with

progressively and or irregularly swollen

apex; measures as above;

capitato-pedicellate: as above but with

abruptly, regularly swollen apex; measures

as above.

lageno-(sub)cylindraceous: lageniform

(a basal swelling surmounted by a more or

less long neck, which is more or less thin-

ner than half the breadth of the basal

swelling) with more or less cylindraceous

neck and a hardly clavate apex; measures

refer to total length, apex, neck base and

basal swelling breadth;

lageno-(sub)clavate: lageniform with

more or less clavate apex; measures as

above;

lageno-(sub)capitate/swollen: lageni-

form with more or less capitate (abruptly

inflated) or swollen apex; measures as

above.

It is hardly necessary to specify that

some of the above variations may at times

occur in the same mount, in which case the

dominating pattern is emphasized. A visu-

al representation of the above cystidial ter-

minology is shown in the illustrations

accompanying the microanatomical

descriptions.

As usual, herbaria are cited in con-

formity with Index Herbariorum Part

I: The Herbaria of the World

<http://www.nybg.org/bsci/ih/> (The

New York Botanical Garden). The materi-

al housed in the author’s herbarium is

referred to as HG. Author citation follows

137

the updated electronic version of

Fungal name authors by P. M. Kirk

<http://www.indexfungorum.org/Authors

OfFungalNames.htm> (CABI Bioscience

UK). Index fungorum

<http://www.indexfungorum.org/Names/

NAMES.ASP> (CABI Bioscience UK)

was also accessed.

Standard abbreviations are taken from

STEARN (2004). The descriptive terminolo-

gy, unless otherwise stated, follows mostly

FAN (BAS et al, 1988; NOORDELOOS & al,

2005), LARGENT (1973) and LARGENT & al

(1977). Colour terminology is the one

adopted in the Colour Identification Chart

of the Flora of British Fungi (Edinburgh,

1969). Spore colour and dextrinoidity

degree is coded with reference to

KORNERUP & WANSCHER (1978). The sys-

tematic framework of reference for the

genus Hebeloma is that offered in

VESTERHOLT (2005), higher taxa are named

after HIBBETT & al (2007).

Results, notes and comments

Type studies

HEBELOMA LEUCOSARX P. D. Orton

1960 (Fig.s 1, 7).

Basyonym: Hebeloma leucosarx P. D .

Orton, Trans. Brit. Mycol. Soc. 43(2): 244-

245; 1960

Holotypus: Scotland, Loch Loy,

Morayshire, 25.9.1955 D.P.Orton 583 (K)

Description of type collection (P.D.Orton

583; K) (Fig. 1)

ORTON (1960): “Pileus 22-58 mm, e con-

vexo expansus vulgo late obtuse umbona-

tus, senectute plano orbicularis vel ad

marginem undulato-lobatus, sordide

incarnato-ochraceus vel pallide fulvo-

ochraceus, vulgo ad marginem pallidiore

vel ad discum obscuriore coloratus, jove

pluvio viscidus, jove sicco glaber, ad mar-

ginem primo eleganter albo-tomentosum

glabrescens. Lamellae adnatae plus

minusve emarginatae, pallide argillaceae

vel sordide incarnato-ochraceae (colo-

rem marginis pilei eccedentes) postremo

argillaceo-luteolobrunneae, confertae, L

40-66, l (1)-3-(7), ad aciem juventute

conspicue albo-floccosis jove pluvio plo-

rantes. Stipes 25-90 / 3,5-11 mm (infra 5-

18), aequalis vel clavato-bulbosus, albus

dein praecipue ad basim pallide sordido-

ochraceus, primo totus albo-pruinosus

dein intra albo-sericeo striatus, e farcto

cavus. Caro alba, in cortice stipitis sordi-

de ochraceotincta, intus solum vulnerato

leviter decolorans. Odor fortis, raphanoi-

deus. Sapor mitis dein acris. Sporae limo-

niformes, punctatae, 9-12-(14) / 5,5-6,5

µm. Basidia 4-sporigera 24-30 / 8-9 µm.

Cystidia aciei lamellarum plus minusve

clavata 40-60 / 4-6 µm, ad apicem (5)-7-

13 µm; cellulae similes ad apicem stipitis

a-dsunt.

Gregarius, ad terram humidam in salicetis

betuletisque. Loch Loy, Morayshire, 15

Sept. 1955 (typus in Herb.Kew). A carne

plerumque alba, acie lamellarum plorante,

margine pilei primo albotomentoso et

coloribus insignis”.

Observations on the material: The type

(UK. Loch Loy, Morayshire (Grampian),

Scotland, 25 Sep. 1955, P. D. Orton 583,

K), consists of two, well-preserved speci-

mens, one entire and the other a lengthwise

section. Even though very similar in

stature and colours (pileus dark brick to

purplish date at disc, fulvous in the peridis-

cal area and much lighter, pale buff to

dingy whitish at the margin and stipe con-

colorous with the pileal margin and pale

brownish downwards), their micro-

anatomical features, cystidial morphology

in particular, suggest that they be referred

to different morphological species.

Therefore, as it appears to be a case of

admixture the two specimens are analyzed

separately.

What is Hebeloma leucosarx ?

138 Grilli E.

Fig. 1 - Hebeloma leucosarx. Holotype, P.D.Orton 583 (K). On le left “Specimen 1”

(lectotypus), on the right “Specimen 2” (H. lutense). Bar 2 cm.

Before returning the material to K, the

two specimens were separated and num-

bered: “1” the entire basidiome and “2”

the sectioned one (Fig. 1).

Personal study

Specimen 1 (entire basidiome) (Fig. 2).

Spores (11)-11.4-12.6 x (5.8)-6.3-6.8-(7.2)

µm., Q = (1.69)-1.75-1.91-(2.03), on aver-

age [32/1] mL 11.92 µm, mW 6.47 µm.,

mQ = 1.83; morphology prevalently sub-

fusiform (56.7 %), less frequently amyg-

daliform (36.7 %), occasionally fusiform;

apex rather rounded (obovoid in front

view), not or rarely mucronate; ornamen-

tation mostly subdistinct (but most spores

smooth or hardly ornamented in light

microscopy); myxosporium undilating;

light-coloured, greenish-white [30A2,5];

subdextrinoid [4C6]. Apical spore wall

structure modified by the occurrence of a

reduced pore.

Basidia 25-35 x 8.0-10-(11) µm, clavate,

four-spored.

Subhymenium subcellular; hymenophoral

trama subregular.

Lamellar edge sterile. Cheilocystidia

dimorphic, prevalently lageno-clavate,

(30)-42-66-(78) x (5.0)-6.5-8.0-(9.0) x

(3.0)-4.0-5.5-(6.6) x (5.5)-6.0-9,6-(12)

µm; on average (33/1) mL 54.1 µm, maB

7.1 µm, mmB 4.7 µm, mbB 7.5 µm; less

frequently clavato-cylindraceous (48)-60-

75-(94) x (7.0)-7.8-9.0-(10.5) x (3.6)-4.0-

5.0-(5.8) µm; on average (20/1) mL 69.3

µm, maB 8.3 µm, mmB 4.5 µm; both

types with distinctly congophilous walls

and with conspicuous, hypercongophilous,

plaque-like thickenings especially in the

apical part.

Pleurocystidia none.

Pileal margin with numerous pileocystidia,

139

morphologically similar to hymenial cys-

tidia, (36)-62-97-(102) x (3.7)-4.0-7.0-

(8.0) x (3.6)-4.0-5.0 µm (Fig. 4).

Pileipellis not studied. (Regrettably, at the

time of the revision caulocystidia and

pileipellis structure were not routinely ana-

lyzed in case of scanty material).

Specimen 2 (section) (Fig. 3).

Spores (9.6)-10.4-11.4-(12.2) x 5.6-6.4-

(6.6) µm, Q = (1.62)-1.71-1.96-(2.0); on

average (69/1) mL 11.06 µm, mW 6.07

µm, mQ 1.82; morphology prevalently

subfusiform (56.3 %), less frequently

amygdaliform (37.1 %), occasionally

fusiform; apex more or less acute, not

mucronate; ornamentation mostly distinct;

myxosporium undilating; light- to medi-

um-coloured, greenish-white to yellowish-

greyish [29A2; 30A/B3; 1A/B3]; subdex-

trinoid [4B/C4]. Apical spore wall modi-

fied by the occurrence of a reduced pore.

Basidia 26-38 x 7.0-9.0 µm, clavate, four-

spored.

Subhymenium subcellular; hymenophoral

trama subregular.

Lamellar edge sterile. Cheilocystidia

dimorphic, prevalently (65.7 %) clavato-

to swollen-pedicellate (28)-40-66-(76) x

(5.0)-6.0-10.5-(15) x (2.0)-2.8-4.0-(5.0)

µm, on average (71/1) mL 55.8 µm, maB

7.9 µm, mmB 3.3 µm, less frequently

(34.3 %) lageno-clavate (30)-40-51-(68) x

(4.5)-6.0-7.5-(9.5) x (2.0)-2.5-3.8-(4.5) x

(4.5)-5.0-6.3-(7.5) µm, on average (37/1)

mL 46.2 µm, maB 6.9 µm, mmB 3.1 µm,

mbB 5.7 µm, pedicel or lower portion

often typically sinuous; no plaque-like

thickenings observed, apex often hyper-

congophilous.

Pleurocystidia none.

Pileal margin with numerous pileocystidia,

morphologically similar to hymenial cys-

tidia, generally with less inflated apex,

(33)-38-89-(98) x (4.6)-5.5-8 x 2-3.5 µm

and 38-46 x 4.8-7.8 x 2.7-4.0 x 4.5-5.5 µm

(Fig. 4).

Pileipellis not studied.

Additional collections (Fig.s 5, 7).

Given the situation above, it was

judged advisable to revise four additional

collections, now stored at E, which have

the status of “original material”. However,

two of them, namely PD Orton 1967 and

2611, were excluded from the present

study because showing a doubtful

microanatomical picture suggesting H.

pusillum J. E. Lange and H. helodes J.

Favre respectively.

One more collection of H. leucosarx

from the UK (M. Moser 81/424 [IB]) was

considered worth including in this study

because its identification was confirmed

by PD Orton (authentic), and the relative

photograph was published in MOSER &

PEINTNER (1985-2006).

1 - UK. Loch Loy, Morayshire

(Grampian), Scotland, Sept. 25, 1955, P.

D. Orton 593 (E00076310; E) (Fig. 5).

Observations on the material: the collec-

tion, badly fragmented, is in rather poor

conditions; also, most spores are rather

anomalous in appearance (smooth, larger

and sometimes morphologically deviating)

with very few spores exhibiting a normal-

ly developed myxosporium. The sporal

data, as usual, refers to the minority of

spores judged “normal”.

Spores (9.6)-10.4-12.2-(13) x (5.6)-6.0-

7.0-(7.2) µm, Q = (1.60)-1.71-1.87-(1.96);

on average (71/1) mL 11.51 µm, mW 6.42

µm, mQ 1.79; morphology subfusiform

(53.5 %) or amygdaliform 46.5 %); not

mucronate; ornamentation subdistinct;

myxosporium undilating; medium-

coloured, pale yellowish-greyish [1A/B2];

subdextrinoid [4-5B3]. Reduced pore

hardly visible.

Basidia 25-32 x 9.0-10 µm, clavate,

four-spored.

What is Hebeloma leucosarx ?

140 Grilli E.

Fig. 2 - H. leucosarx. Holotype, “Specimen 1”. Spores & cheilocystidia. Bar 10 µm

(spores); bar 30 µm (cheilocystidia).

141

Fig. 3 - H. leucosarx. Holotype, “Specimen 2”. Spores & cheilocystidia. Bar 10 µm

(spores); bar 30 µm (cheilocystidia).

What is Hebeloma leucosarx ?

142

Fig. 4 - H. leucosarx. Holotype. Marginal pileocystidia, “specimen 1” (top), “specimen 2”

(bottom). Bar 30 µm.

Grilli E.

143

What is Hebeloma leucosarx ?

Lamellar edge sterile. Cheilocystidia

dimorphic, lageno-clavate, rarely lageno-

capitate (36)-43-62-(77) x (6.0)-6.5-10-

(11.5) x (4.0)-4.4-5.0-(6.0) x (5.5)-6.0-9.0-

(11) µm, on average (20/1) mL 51.6 µm,

maB 8.3 µm, mmB 4.7 µm, mbB 7.1 µm;

less frequently clavato-cylindraceous,

occasionally with epiphysiform (bifid)

apex, (46)-57-79-(105) x (7.0)-8.0-8.5-

(10) x (3.5)- 4.4-5.5-(6.5) µm; on average

(15/1) mL 67.2 µm, maB 8.2 µm, mbB

4.8 µm; both types with occasional hyper-

congophilous, plaque-like thickenings.

Pleurocystidia none.

Pileipellis: suprapellis a 120-150 µm

thick, loosely tangled ixotomentocutis,

hyphae (1.6)-2.0-3.5 µm wide, in the

lower layer becoming slightly broader,

embedded in a complete gel matrix; termi-

nal hyphal cells rather long, 50-102 µm,

with apex 2.0-4.0-(5.5) µm wide, hardly

differentiated, rarely dermacystidioid; pig-

ment mostly intraparietal and, in the thin-

ner hyphae, intracellular (pale yellowish),

occasionally also epiparietal. Subpellis

somewhat chromatically distinct [c 3/4B4]

from the pale pileitrama [1A3], pseudo-

parenchymatic in structure.

2 - UK. Bookham Common, Surrey, Oct.

15,1955, P. D. Orton 596 (E00076300; E)

(Fig. 6).

Observations on the material: the collec-

tion includes c. 5 basidiomata, entire or in

section, all in fairly good conditions, 4 of

which were studied. All the specimens

show a rather bicolorous pileus, with dis-

cal zone more strongly coloured: dark red-

dish brown to reddish brown, than the

marginal one that is dingy ochraceous to

dingy whitish.

Spores (9.8)-10.4-12-(13.6) x (5.5)-6.0-

6.4-(7.0) µm, Q = (1.55)-1.72-1.96-(2.14);

on average (133/4) mL 11.25 µm, mLr

10.93-11.53 µm, mW 6.13 µm, mWr

6.01-6.21 µm, mQ 1.83, mQr 175-1,90;

morphology subfusiform (55%) to amyg-

daliform (35.9%), less frequently fusiform

(10%), occasionally also broadly amygdal-

iform; not mucronate; ornamentation sub-

distinct to (distinct); myxosporium undi-

lating to rugulose, occasionally shallowly

vesiculate; light-coloured, pale greenish-

grey [28A/B2] to greenish-white

[29A/B2]; subdextrinoid [4C5; 4B/C5].

Reduced pore hardly visible.

Basidia 26-34-(37) x 7.0-8.5 µm, more or

less clavate, four-spored.

Lamellar edge sterile. Cheilocystidia

dimorphic, prevalently (60.7 %) clavato-

to inflato-pedicellate, (28)-37-58-(76) x

(5.0)-6.2-10-(14) x (2.0)-2.3-4.0-(4.5) µm,

on average (99/4) mL 44.8 µm, nLr 42.1-

47.6 µm; maB 7.7 µm, maBr 7.5-8.1 µm;

mmB 3.0 µm, mmBr 2.8-3.2 µm, pedicel

typically sinuous; less frequently lageno-

clavate (39.3 %), (25)-34-54-(74) x (4.5)-

6.0-8.5-(10) x (1.5)-2.4-4.0-(4.4) x (4.0)-

4.8-6.0-(7.4) µm, on average (64/4) mL

42.5 µm, mLr 38.5-46.9 µm; maB 7.2 µm,

maBr 6.6-8.0 µm; mmB 2.9 µm,

mmBr 2.7-3.2 µm; mbB 5.3 µm, mbBr

5.1-5.6 µm.

Pleurocystidia none.

Pileipellis: suprapellis a 90-120 µm deep

loosely interwoven ixotomentocutis,

hyphae (1,6). 2.4-4.0-(5.6) µm wide,

embedded in a complete gel matrix; termi-

nal hyphal cells 34-94 x 3.0-5.0-(6.0) µm,

cylindrical to subclavate, occasionally

dermatocystidioid; pigment mostly intra-

cellular, epiparietal in the lower layers;

subpellis more or less chromatically dis-

tinct [2A/B3; 3B5] in comparison to the

pileitrama [29A2], not distinctly pseudo-

parenchymatic in structure.

3 - UK. Hothfield Common, Kent, Sept.

22, 1981, M. Moser 81/424; IB (Fig. 7).

Observations on the material: The collec-

tion is mixed; it includes 5 basidiomata or

parts of basidiomata (see MOSER &

PEINTNER,1985-2006: III Hebeloma 14

144

Fig. 5 - H. leucosarx. P. D. Orton 593 (E). Spores & cheilocystidia. Bar 10 µm (spores);

bar 30 µm (cheilocystidia).

Grilli E.

145

Fig. 6 - H. leucosarx. P. D. Orton 596 (E). Spores & cheilocystidia. Bar 10 µm (spores);

bar 30 µm (cheilocystidia).

What is Hebeloma leucosarx ?

Grilli E.

146

bottom). Three of them, the small ones in

the photograph, belong in the H.

hiemale/album/fragilipes complex and are

excluded, the two bigger specimens are

described below.

Spores (9.6)-10.4-11.2-(12.4) x (5.8)-6.2-

6.8-(7.2) µm, Q (1.48)-1.60-1.78-(1.90);

on average (65/2) mL 10.85 µm, mLr

10.80-10.91 µm, mW 6.42 µm, mWr

6.38-6.47 µm; mQ 1.69, mQr 1.68-1.69;

morphology mostly (81.3 %) amygdali-

form, rarely (12.3 %) broadly amygdali-

form, occasionally subfusiform; generally

not mucronate; ornamentation subdistinct

to distinct; myxosporium undilating; light-

coloured, pale greenish-white [27-28A2];

subdextrinoid [4B3.5-4.5; 4B/C4-5].

Reduced pore hardly visible.

Lamellar edge sterile. Cheilocystidia

dimorphic, mostly lageno-clavate to

lageno-capitulate (75.7%), (38)-46-62-

(70) x (6.0)-7.5-9.5-(10.3) x (3.5)-4.0-5.5-

(7.0) x (5.8)-6.0-8.0-(10) µm, on average

(50/2) mL 53.7 µm, mLr 52.7-54.6 µm,

maB 8.2 µm, maBr 8.21-8.24 µm, mmB

4.7 µm, mmBr 4.6-4-8 µm, mbB 7.4 µm,

mbBr 7.2-7.6; less frequently (24.3%)

clavate-cylindraceous to subcapitulato-

cylindraceous, (52)-62-67-(78) x (6.5)-

9.0-9.5-(12.5) x (3.5)-4.5-5.0-(7.5) µm, on

average (16/2) mL 64.4 µm, mLr 62.7-

66.1 µm, maB 9.1 µm, maBr 8.6-9.6 µm,

mmB 4.9 µm, mmBr 4.2-5.6 µm; both

types with frequent hypercongophilous

plaque-like thickenings.

Pleurocystidia none.

Caulocystidia and pileipellis not studied.

Comments

The above descriptions and relative

micrographs indicate that two of the addi-

tional collections (i.e. Orton 593 and

Moser 81/424) are in agreement with the

microanatomical data obtained from

“specimen 1”, while the third (i.e. Orton

596), with the data from “specimen 2”.

Therefore, they provide corroborating evi-

dence for the assumption that H. leucosarx

is based on a mixed collection and, at the

same time, present additional data on the

morphological circumscriptions of the two

putative taxa involved.

The micro-anatomical features shown

by specimen 1 (entire basidiome) of the

type of H. leucosarx Orton, plus the addi-

tional collections, correspond closely to

those characterizing H. leucosarx ss.

Vesterholt - lately referred to also as H.

velutipes Bruchet (VESTERHOLT, 2004;

2005) - as observed in material kindly sent

to me by the Danish mycologist some

years ago (JV91-509, Oct. 6, 1991,

Marbaek Plantage, Denmark) and as pre-

sented and illustrated in VESTERHOLT

(1992; 1995; 2000; 2005). As shall be seen

below, apart from H. bulbiferum ss Favre

non Maire (GRILLI, 2004) also the types of

H. velutipes Bruchet and H. favrei

Romagn. & Quadr. exhibit comparable

microscopic traits.

On the other hand, the microanatomi-

cal traits of specimen 2 and of Orton 586

conform to those observable in H. leu-

cosarx ss. Bruchet, in the type of H.

lutense Romagn. (see below) and, on the

basis of literature data (regrettably, despite

repeated requests it has not been possible

to study any voucher material), presum-

ably in H. leucosarx ss. Dutch authors, a

taxon first identified as H. lutense (AANEN

& KUYPER, 1999; AANEN & al, 2000b) and

more recently rejected because variously

interpreted, while H. lutense is considered

a synonym of H. helodes (AANEN &

KUYPER, 2004).

Based on the considerations above and

the macro- and micromorphological cir-

cumscription delineated below, it is pro-

posed that the first element (“specimen 1”)

of Orton’s type be referred to Hebeloma

leucosarx P. D. Orton. The second element

(“specimen 2“), by contrast, shows a

microanatomical characterization which

147

What is Hebeloma leucosarx ?

Fig. 7 - H. leucosarx. M. Moser 81/424 (IB). Spores & cheilocystidia. Bar 10 µm

(spores); bar 30 µm (cheilocystidia).

rhizally associated with a wide variety of

mycotrophic hosts.

The description of the microanatomi-

cal details below is based, unless other-

wise stated, on the type and

original/authentic material analyzed

above, field characters and ecological

preferences are integrated with personal

data.

HEBELOMA LEUCOSARX P. D. Orton

(Fig.s 8, 12).

Basyonym: Hebeloma leucosarx P.D.

Orton, Trans. Brit. Mycol. Soc. 43(2):

244-245; 1960

= Hebelomatis leucosarx (P. D. Orton)

Locquin, Fl. Mycol. 3, Cortinariales-A:

146;

1977 (? 1979)

= H. bulbosum Bres. in sched. (inedit.)

= H. velutipes Bruchet, Bull. Soc. linn.

Lyon (suppl.) 39: 127; 1970

= H. bulbosum Romagn. Sydowia 36: 263;

1983 non Fayod 1892

= H. favrei Romagn. & Quadr., Doc.

Mycol. 14 (56): 31; 1984

Misapplications: H. crustuliniforme var.

bulbiferum (Maire) J. Favre Ergbn. wiss.

Unters. schweiz. NatParks, NF 6: 488;

1960 (invalid); H. crustuliniforme ss

Bruchet, Bull. mens. Soc. linn. Lyon

(suppl.) 6: 127; 1970; H. crustulinifortme

var. minus ss Bohus, Ann. Hist.-nat. Mus.

Nat. Hung. 64: 73; 1972

Lectotypus: hic designatus, Scotland,

Loch Loy, Morayshire, 25.9.1955 P.D.

Orton 583 (1, entire basidiome) (K)

Pileus (20)-30-60-(70) mm in diameter,

smooth, viscid in wet weather, colours

rather variable from whitish or ivory, pos-

sibly slightly darker at disc: pale cream,

cream, yellowish cream to pale ochra-

ceous, and then more or less unicolorous,

to various shades of buff: pale clay pink,

vinaceous buff to clay buff, to cinnamonon

148 Grilli E.

makes it indistinguishable from H. lutense

and shall be assigned to Romagnesi’s

taxon.

Lectotypification of Hebeloma leucosarx

Cases of admixtures are governed by

Art. 9.12 and 9.9 of the ICBN (MCNEAL &

al, 2006) which rule: “When a type speci-

men (herbarium sheet or equivalent prepa-

ration) contains parts belonging to more

than one taxon, the name must remain

attached to that part which corresponds

most nearly with the original description

or diagnosis” and “[…] when the material

designated as type is found to belong to

more than one taxon, a lectotype […] as a

substitute for it may be designated”.

On the basis of the protologue, it is

undoubtedly difficult to make the right

decision about which part of the type

should be designated as the lectotype of H.

leucosarx. Either solution appears to some

extent arbitrary. At any rate, the fact that

two of the three line drawings accompany-

ing the protologue (ORTON , 1960: 391,

figs. 47-49) exhibit a bulbous stipe base

and the absence, in the protologue, of any

hint at the typical sinuous morphology of

the cheilocystidia incline me to choose

“specimen 1” as the lectotype. In future

molecular research involving the type of

H. leucosarx, one should be aware of the

situation of the type and generate

sequences from both elements.

This fungal organism, by far the com-

monest species in the section Velutipes

Vesterh, and one of the most frequent in

the whole genus as well, is normally in the

medium size range, with pileus ranging

considerably in colours and stature also

variable from squat to fairly slender, with

stipe thickly pruinose-velutinous and more

or less distinctly bulbous, at times even

markedly so. As as rule, odour is more or

less distinctly raphanoid, while taste is

usually bitterish. The species is mycor-

149

or sienna, in which cases assuming a dis-

tinctly bicolorous appearance.

Lamellae generally emarginate, crowded,

narrow to fairly broad, (2)-3-6-(7) mm,

exuding watery to opalescent droplets in

wet weather, otherwise hardly beaded to

dry, pale beige to brown, edge distinctly

whitish (under a magnifying glass).

Stipe (20)-30-80-(100) x 5-9-(12) mm,

more or less equal, with base generally

swollen into a more or less distinct bulb

(up to 18-20 mm), thickly pruinose-veluti-

nous, whitish, slightly discolouring

brownish towards base; without visible

cortinate velar remnants. Stuffed to hol-

low, and then often with a sharp pointed

shred of pith hanging down from the apex.

Context whitish, more or less concolorous

with the outer surface in stipe base. Odour

more or less raphanoid, at least on cutting;

taste raphanoid-bitterish.

Spores in the medium-size range:

(140/4/3) (9.6)-10.2-12.2-(13.6) x (5.6)-

6.3-6.8-(7.2) µm, Q (1.48)-1.64-1.93-

(2.04); mL 11.29 µm, mLr 10.80-11.92

µm; mW 6.43 µm, mWr 6.38-6.47 µm;

mQ 1.75, mQr 1.68-1.83. Morphology

predominantly amygdaliform (60 %), less

frequently subfusiform (32 %), occasion-

ally broadly amygdaliform or, very rarely,

fusiform; obovoid in face view (tendency

to low/medium mean Q values with fairly

rounded apex); ornamentation of normally

developed spores mostly subdistinct to

distinct; myxosporium never truly dilat-

ing, generally indistinct, at most rugulose,

only occasionally vesiculate; light- to

medium-coloured; dextrinoidity always

more or less distinct; cyanophily negative

to weak (acyanophilous or hypocyano-

philous).

Apical modification of spore teguments

generally present as a light spot, especial-

ly visible in Melzer’s reagent (Cle-

mençon’s reduced pore).

Cheilocystidia dimorphic: (a) lageno-

clavate or less frequently lageno-capitate:

(36)-44-63-(78) x (5.5)-6.5-9.0-(11.5) x

(3.0)-4.2-5.0-(7.0) x (5.5)-6 .0-8.5-(12)

µm, on average (103/4/3) mLr 51.6-54.6

µm, maBr 7.1-8.3 µm, mmBr 4.6-4.8 µm,

mbBr 7.1-7.6 µm, and (b) clavato-cylin-

draceous to capitato-cylindraceous: (46)-

57-78-(105) x (6.5)-8.0-9.5-(12.5) x (3.5)-

4.0-5.5-(7.5) µm, on average (51/4/3) mLr

62.7-69.3 µm, mBr 8.2-9.1 µm, mbBr

4.5-4.9 µm. Both types typically with

average median breadth ≥ 4 µm, apex at

times also epiphysiform (bifid), especially

in type (b) (in this case the often signifi-

cantly larger apex measures were not

included in the size range). The percentage

of the morphological types is variable, but

generally type (a) is more frequent.

The typical congophilous plaque-like

thickenings (which in some cases appear

more like incrustations) that often modify

the external outline of the cystidium are

often present in either type. Generally they

are located at the apex, but occasionally

they occur also elsewhere. In cystidia with

epiphysiform apex, most of the times, the

thickening is characteristically located in

the saddle-like depression of the epiphysis.

These typical thickenings seem to be age-

dependent. In sections mounted in Congo

red, they are generally numerous and eye-

catching and it is almost impossible to

miss them, but occasionally, especially in

not fully mature basidiomes, it takes some

experience to spot them as they are rare,

small and hardly congophilous.

Caulocystidia similar to cheilocystidia but

normally much longer, e.g. 70-148-(178)

µm (EG-030920.2), (40)-54-145-(194) µm

(Consiglio 96021), occasionally with

sparse plaque-like thickenings (Fig. 12).

Pileocystidia along pileus margin similar

to cheilocystidia, mostly clavate.

Pileipellis: suprapellis a loosely tangled

ixotomentocutis, 120-150 µm thick,

hyphae (1.6)-2.0-3.5 µm wide, in the lower

layer becoming slightly broader, embed-

ded in a complete gel matrix; terminal

hyphal cells rather long, 50-102 µm, with

What is Hebeloma leucosarx ?

150 Grilli E.

apex 2.0-4.0-(5.5) µm wide, hardly differ-

entiated, rarely dermacystidioid; pigment

mostly intraparietal and, in the thinner

hyphae, intracellular (pale yellowish), epi-

parietal pigment rather scarce. Subpellis

somewhat chromatically distinct [c 3/4B4]

from the pale pileitrama [1A3], pseudo-

parenchymatic in structure.

Habitat: a generalist ECM species, associ-

ated with many ecologically important

families of ectomycorrhizal plants, among

which especially Betulaceae (Betula,

Carpinus, Corylus), Fagaceae (Castanea,

Fagus, Quercus), Malvaceae (Tilia), and

Pinaceae (Abies, Pinus), less frequently

also Salicaceae (Populus, Salix).

VESTERHOLT (2005) also reports associa-

tions with Salix herbacea and Dryas

octopetala (Rosaceae) in alpine areas.

Also recorded among Sphagnum under

Betula (LLISTOSELLA & al, 1999 as H.

helodes; LANZONI, 1988, as H. longi-

caudum).

Comments

The main diagnostic microanatomical

feature is the rather bulky morphology of

cheilocystidia, with main body generally

straight, apex mostly clavate, but including

also the possibility for it to be obscurely to

distinctly capitulate, and, in particular, the

breadth of the median part of the main

body on average ≥ 4 µm - a trait whose

diagnostic nature was first emphasized by

VESTERHOLT (1995). Ancillary characters

for cheilocystidia are the not at all rare

presence of a bifid apex and that of congo-

phylous plaque-like thichenigs (“rare

unique character states” according to

AANEN & KUYPER (2004) definition). This

trait is coupled with the relatively broad,

mostly non-mucronate spores exhibiting a

more or less distinct reaction in Melzer’s

and a non-dilating myxosporium. In most

collections the average quotient is in the

“amygdaliform” range (1.60-1.79), only in

a few cases exceeding the lower limit of

Fig. 8 - Hebeloma leucosarx P. D. Orton (EG-030920.2, Val Fondillo, Opi, AQ).

151

What is Hebeloma leucosarx ?

Fig. 9 - Hebeloma leucosarx P. D. Orton (EG-981013.2, Goriano Sicoli, AQ).

Fig. 10 - Hebeloma leucosarx P. D. Orton (G. Consiglio 98064, Civago, RE).

152 Grilli E.

the “subfusiform” range (1.80-1.99). Even

though not routinely recorded, the remark-

able length of caulocystidia may also turn

out to be diagnostically useful.

Being a generalist ECM species, the

information load carried by habitat data is

rather limited, even though the association

with Pinaceae, Fagaceae and Betulaceae

seems to be statistically more frequent

than with Salicaceae.

The morphological circumscription

above is roughly the same as that estab-

lished by AANEN & KUYPER (2004) and

VESTERHOLT (2005) for H. velutipes

Bruchet, minus, for the former authors, the

inclusion in the synonymy of H. bulbifer-

um Maire (= H. colossus Huijsman), a

taxon which is clearly set apart from H.

velutipes by macro- and microscopic fea-

tures alike, as advocated in GRILLI (2004).

H. leucosarx, as here conceived and

lectotypified, is illustrated by photographs

of some recent collections, chosen in order

to provide some evidence of the broad

range of possible pileus colours. (Fig.s 8,

11). For brevity’s sake, their descriptions

are not included, but the average measures

of the core microanatomical features

(spores and cystidia) of two of the four col-

lections illustrated were included in

SUPPLEMENTARY TABLE 1.

Selected illustrations: VESTERHOLT

(1992, 2000, 2004, 2005), (as H. leucosarx

or H. velutipes); GRILLI (1994), (as H.

favrei); LLISTOSELLA & al (1999), (as H.

helodes); GERHARDT & al, (2000), (as H.

helodes); ENDERLE (2004); MOSER &

PEINTNER (1985-2006), (see above). The

colours of the specimens illustrated in the

photographs range from pale brownish to

bright reddish-ochre. Regarding the photo-

graph published in LLISTOSELLA & al

(1999), my conclusions are based on the

study of the relative voucher material: J.

Llistosella MA 1460, MA 1461, MA 1462

(BBC); the same photograph was also pub-

lished in GERHARDT & al, (2000). Other

Fig. 11 - Hebeloma leucosarx P. D. Orton (G. Consiglio 96021, Val Saviore, BS).

153

What is Hebeloma leucosarx ?

Fig. 12 - H. leucosarx. Caulocystidia, EG-030920.2 (left),G. Consiglio 96021 (right). Bar

30 µm.

154 Grilli E.

photographs have been published in BON

(1993) (as H. velutipes), BREITENBACH &

KRÄNZLIN (2000), BASTARDO DEL VAL & al

(2001), (as H. favrei), and ALBERS &

GRAUWINKEl (2005), but the descriptions

are not exhaustive and I have not studied

the voucher material. The photograph pre-

sented by Albers & Grauwinkel most like-

ly represents H. crustuliniforme (stipe

pruinose-squamulose, not bulbous, spores

hardly dextrinoid; regrettably, cheilocys-

tidia morphology is not described).

Comparison with the types of H. velu-

tipes Bruchet, H. favrei Romagn. &

Quadr. and with the voucher collections

of H. bulbosum Bres. (inedit.) and of

some putative misapplications

HEBELOMA BULBOSUM Bres. nov. sp.

(inedit., in sched.)

Italy.Trento, Piné, Sept.. 10, 1926 G.

Bresadola B/266 (TR); Trento,Villa

Margone, Oct. 13, 1926, G. Bresadola

B/247 (TR).

Habitat: “in pinetis” and “in silvis mixtis”.

Observations on the material: The collec-

tions include two specimens each, all in

good conditions. Inside the first sheet there

is a herbarium slip by L. Quadraccia, who

revised the material on July 20, 1989,

which reads: “H. crustuliniforme (Bull.)

Quél (ut “H. bulbosum Bres. n. sp.” inedi-

tum)”

The collections are in perfect morpho-

logical agreement with H. leucosarx as

delimited in the present study. To avoid

burdening unnecessarily the paper the rel-

ative data is not shown; however, the aver-

age measures of the core features are

included for comparison in SUPPLEMEN-

TARY TABLE I.

Comments

The mycological literature of the past

is replete of homonyms, but much more

rarely have they turned out to be also

synonyms, as in the case of H. bulbosum

Bres. (ined.) and H. bulbosum Romagn;

but there is yet another H. bulbosum, the

one published by Fayod in 1892 (Annali R.

Accad. d’Agric. 35 (6): 95), which could

well refer to the same organism: “Fungus

albus vel lutescens, pileo hemisphaerico,

laevi, dein pulvinato, c. 4-5 cm lato, umbo-

ne obtuso, jove udo subviscoso; carne alba

(centro circiter, 1 cm. crassus), stipite

medullato farcto (6-7 cm longo, 0,5 cm

lato), albo apice farinoso, basi in bulbillo

subsphaerico inflato; lamellis liberis, sub-

confertis, acie crenulatis et albo punctatis,

cinnamomeis. Dispositio lamellarum et

schemate18281 expressa est. Hab.

Catervatim in graminosis, betuletis et

alnetis, in alpibus Pedemontium (Jaco-

punt, Bricherasio), septembri mense. Obs.

micr. Trama sat regularis; subhymenium

pseudoparenchymaticum monostratum.

Paraphysa clavatae. Basidia emergentia

(27 x 9 µ) tetrasterigmatica. Sporae (9 x 6

µ) ovoideo-conicae, leaves vel subtiliter

punctatae, sub micr. ochraceae. – Cuticula

pilei monostrata, glutinosa.”

Regrettably, there is no material refer-

able to this taxon in Fayod’s herbarium at

G (Ph. Clerc pers. comm.); consequently,

in a genus in which microanatomy plays a

major, often decisive role, the issue of its

possible synonymy, and thus of its being

an earlier competing epithet of H. leu-

cosarx, is destined to remain pure specula-

tion, which can only be source of endless

and inconclusive dispute.

HEBELOMA VELUTIPES Bruchet

(Fig. 13).

Basyonym: H. velutipes Bruchet, Bull.

mens.Soc.linn.Lyon Supp.6: 127; 1970.

= Hebelomatis velutipes (Bruchet)

Locquin, Fl. Mycol., 3, Cortinariales-A:

146; 1977 (?1979).

Holotypus: France. Les Echets (Ain), Sept.

24, 1963, BR 63-20 (LY).

Description of type collection (G. Bruchet

63-20; LY)

BRUCHET, 1970: “Cortina nulla; pileo

majore, usque ad 50 mm. lato, convexo vel

e plano convexo, haud mammoso, concol-

ore, jove sicco luteolo, ex ocraceo luteo vel

e gilvo luteo, parum carnoso; stipite 50-70

x 6-8 mm., longiore, haud gracili, subbul-

boso, albido, sub lamellis velutino ac

inferius passim fioccoso; lamellis guttulis

lacteis in acie madidis; odore debiliter

raphanoideo. Sporis 10-12 x 5,5-6 µm,

amygdaliformibus, brevioribus, parum

ventricosis. s. m. brunneis, verrucosis,

pariete firma; pilis marginum 50-65 µm

longis, summis 7,5-10 µm capitatis. Sub

Betulis Salicibus mixtis. Typus in Her. n°

BR 63-20.”

Observations on the material: Holotype

probably lost. An isotypus (00875), consist-

ing of a small pileus section, housed in

Quadraccia’s herbarium at RO (duplum of

G. Bruchet 63.20, France, Les Echets

(Ain), Sept. 24 1963) was studied. Three

topotypic paratypes were also studied

(France, Les Echets (Ain), Sept. 24, 1963,

G.Bruchet 63.21 (LY); ditto, Oct. 9, 1964,

G. Bruchet 64.31 (LY); ditto, Oct. 2, 1965,

G. Bruchet 64.15 (LY) which, as usual with

Bruchet’s material, include few, small frag-

ments. The scarcity of the material allowed

only for the study of spores and cystidia.

Given the quantitative homogeneity of the

microanatomical traits observed in the

study of the three paratypes, the relative

data is presented jointly.

Personal study

Isotype

Spores (9.6)-10.4-12-(12.8) x (5.6)-6.2-

155

7.0-(7.2) µm, Q = (1.55)-1.70-1.87-(1.93);

on average [43/1] mL 11.20 µm, mW 6.38

µm, mQ 1.75; morphology mostly amyg-

daliform (79 %), more rarely subfusiform

(18.6 %), occasionally broadly amygdali-

form; mostly non-mucronate; ornamenta-

tion subdistinct to distinct; myxosporium

undilating, light- to medium-coloured,

greenish-grey to yellowish-greyish

[30A/B2; 1A/B2]; subdextrinoid [4A/B4;

5B3.5]. Apical spore wall structure modi-

fied by the occurrence of a reduced pore.

Basidia 34-40 x 8.0-10 µm, four-spored,

clavate to slightly constricted.

Lamellar edge sterile. Cheilocystidia

dimorphic, lageno-clavate to lageno-capit-

ulate, (48)-53-78-(86) x (7.0)-7.8-10.5-

(11) x 3.5-5.0-(6.0) x 6.0-8.0-(10) µm, on

average [23/1] mL 66.4 µm, maB 8.8 µm,

mmB 4.3 µm, mbB 7.3 µm; and clavato-

cylindraceous or, less frequently, capitula-

to-cylindraceous (a few with bifid apex),

(53)-57-84-(88) x 7.0-10-(12) x (3.8)4.0-

5.5-(5.8) µm, on average [21/1] mL 73.2

µm, maB 8.8 µm, mmB 4.8 µm; both

types with more or less frequent hypercon-

gophilous plaque-like thickenings.

Pleurocystidia none.

Caulocystidia not studied, but BRUCHET

(1970) reports: “(40)-60-85-(100) µm”,

and describes them as “bien plus longs que

les poils marginaux”.

Pileipellis not studied.

Paratypes

Spores (9.2)- 9.6-11.4-(12) x (5.4)-5.6-6.4-

(6.8) µm., Q = (1.45)-1.52-1.83-(1.90),on

average [125/3/3] mL 10.3 µm, mLr 10.2-

10.5 µm, mW 6.19 µm, mWr 6.1-6.3 µm;

mQ 1.66 µm, mQr 163-1.72;1.66 µm,

mQr 163-1.72; morphology prevalently

amygdaliform (68 %), less frequently

broadly amygdaliform (19.2 %) or sub-

fusiform (12.8 %); apex rounded (obovoid

in front view), not or occasionally slightly

mucronate; ornamentation subdistinct to

distinct; myxosporium undilating; light- to

medium-coloured, greenish-grey, pale yel-

What is Hebeloma leucosarx ?

156 Grilli E.

lowish-grey or yellowish-grey [30A/B2-3;

1A/B2; 1B3.5; 2B3]; subdextrinoid

[4A/B2; 4C4; 5B3.5]. Apical spore wall

structure modified by the occurrence of a

reduced pore.

Basidia 26-31 x 8.0-11 µm, four-spored,

clavate to slightly constricted..

Lamellar edge sterile. Cheilocystidia

dimorphic, prevalently lageno-clavate

(28)-35-56-(69) x (4.6)-5.8-8.0-(9.0) x

(3.7)-4.0-5.0-(6.2) x (5.0)-5.8-8.0-(9.0)

µm, on average [62/3] mL 41.8 µm, mLr

38.1-45.8 µm, maB 6.6 µm, maBr 6.1-7.0

µm, mmB 4.5 µm, mmBr 4.4-4.7 µm,

mbB 6.7 µm, mbBr 6.5-7.0 µm; less fre-

quently clavato-cylindraceous to, occa-

sionally, capitulato-cylindraceous (no cys-

tidia with bifid apex), (38)-47-58-(64) x

(5.2)-6.0-9.0-(10) x (3.0)-4.0-5.5-(6.0)

µm, on average (50/3) mL 51.5 µm, mLr

48.4-53.1 µm, maB 7.4 µm, maBr 6.7-8.0

µm, mmB 4.6 µm, mmBr 4.4-4.8 µm;

both types with distinctly congophilous

apical portion and showing more or less

frequent hypercongophilous, plaque-like

thickenings.

Pleurocystidia none.

Caulocystidia and pileipellis not studied.

Comments

The quantitative differences between

the isotype and the three paratypes was con-

sidered worth reporting in full, to give an

idea of the variability pattern. Having stud-

ied the isotype only recently, based on the

paratype data I toyed for some time with the

idea that H. velutipes might be character-

ized by comparatively small spores coupled

with relatively short cystidia. The addition-

al data from the isotype make such a possi-

bility absolutely unrealistic.

Material referring to Hebeloma crus-

tuliniforme ss Bruchet was also studied to

substantiate the hypothetical synonymy

assumed on the basis of the description in

BRUCHET (1970: 73-77).

Hebeloma crustuliniforme ss. Bruchet

1970

France, Forêt de l’Isertan, Pralognan,

Savoie, Sept. 10, 1964, G. Bruchet 64.21

(LY).

Observations on the material: The material

includes a badly fragmented specimen

which refers to one of the basidiomes illus-

trated by the line drawing of Plate XIX A.

Personal study

Spores (50/1) (8.0)-9.0-10.6-(12) x 5.4-

6.4-(6.8) µm. Q = (1.46)-1.53-1.71-(1.76),

on average (50/1) mL 9.79 µm, mW 6.09

µm, mQ 1.60; morphology mostly amyg-

daliform (77 %), less frequently broadly

amygdaliform (23 %); apex not mucronate

(obovoid in front view); ornamentation

subdistinct to distinct; myxosporium undi-

lating; light-coloured, greenish-grey

[30A/B3]; subdextrinoid [4C4]. Apical

spore wall modified by the occurrence of a

reduced pore.

Basidia 27-34 x (7.0)-8.0-11 µm, four-

spored, clavate.

Lamellar edge sterile. Cheilocystidia

dimorphic, prevalently lageno-clavate,

(35)-50-67-(73) x (4.5)-6.0-8.5-(9.0) x

3.0-5.0 x (5.0)-6.0-10-(12) µm, on average

(27/1) mL 58.1 µm, maB 7.1 µm, mmB

4.0, mbB 7.8 µm, less frequently clavato-

cylindraceous, (60)-70-77-(82) x (6.0)-

7.5-10 x (3.7)-4.0-5.0 µm, on average

(15/1) mL 72.7 µm, maB 8.1 µm, mmB

4.4 µm; congophilous plaque-like thicken-

ings not observed.

Pleurocystidia none.

Comments

The collection exhibits small-sized

spores - but, again, well within the overall

size range reported below and that record-

ed by VESTERHOLT (2005) for H. velutipes

- and apparently lacks cystidia with bifid

157

What is Hebeloma leucosarx ?

Fig. 13 - H. velutipes. Isotype. Spores & cheilocystidia. Bar 10 µm (spores); bar 30 µm

(cheilocystidia.

158 Grilli E.

apex and the typical plaque-like thicken-

ing. However, BRUCHET (1970) describes

also larger spores and in the plate XIX B

illustrates a caulocystidium with a distinct-

ly bifid apex. Also, stature, general facies

and pileal colours of the basidiomes

described are in agreement with H. leu-

cosarx as circumscribed in the present

study.

Further, Bruchet was well aware of

the fuzzy boundary line between H. velu-

tipes and H. crustuliniforme as he inter-

preted it: “Il est par contre beaucoup plus

difficile à distinguer de crustuliniforme

(…) Ses caractères distinctifs seraient:

sont chapeau moins charnu, s’étalant plus

tôt, son stipe plus long, moins épais,

jamais aussi nettement bulbeux, à orne-

mentation différente puisque celui-ci

apparaît velouté sous les lamelles et

éparsément floconneux plus bas et non

densément poudré-sablé à floconneux sur

toute sa surface, ses lamelles ne présen-

tant pas de reflet carné, et ses spores en

moyenne moins ventrues”. Certainly, this

is not H. crustuliniforme ss Vesterholt

with indextrinoid, subfusiform to

fusiform spores and cheilocystidia typi-

cally sinuous-pedicellate, rather similar to

those of H. lutense (see below).

Bruchet’s confusion between H. crus-

tuliniforme and H. leucosarx need not be

necessarily surprising, it was, in fact,

rather common during last century, when

the separating traits had not yet been clear-

ly defined and almost all collections with

beaded lamellae were determined as H.

crustuliniforme.

HEBELOMA FAVREI Romagn. &

Quadr. 1984 (Fig. 14).

Basyonym: Hebeloma favrei Romagn. &

Quadr., Doc. Mycol. 14 (56): 31; 1984

=H. bulbosum Romagn. 1983, Sydowia

36: 263, non H. bulbosum Fayod 1892,

Ann. R. Acad. Agric. Torino: 95

= H. crustuliniforme var. bulbiferum ss

Favre 1961 non Maire 1937,

Holotypus: France, Aureilhan (Landes),

Oct. 13, 1977, H.Romagnesi 77.333 (PC)

Description of type collection (H.

Romagnesi 77.333; PC)

ROMAGNESI, 1983: “Pileo (2)-2,5-5,7

cm, primum e campanulato convexo, dein

e-xpanso, margine angusta involuta, dein

magis minusve undata, e rufulo ochraceo,

in ambitu pallidiore, cute viscidula, laeve

atque paulum micacea, vetustate rugulosa.

Sipite brevi, 2,5-3,5 x 0,5-0,7 cm, sursum

cylindrato, deorsum manifeste bulboso

(usque ad 1 cm), pieno dein medullato,

demum cavo, cortice firmo, albo vel albi-

do, fere toto pruinoso. Carne crassiuscula,

firmula, albida, paulum pallidissimo colo-

re brunneolo in cortice stipitis tincta.

Odore raphanoideo, sapore amarescente.

Lamellis stipatis, triplici serie lemellula-

rum, (2)-3-5 mm latis, rotundatis sublibe-

ris vel adnatis, pallide argillaceis, acie

crenata ac guttulis plerumque limpidis,

interdum lacteis. Sporis late ovoideis-

amygdaliformibus, (8,7)-9-11,7 x 6-6,8

µm, 10-11,7 x 5,7-6,5-(8) µm (NH4OH),

luteolis, sublaevibus vel subtiliter puncta-

tis. Basidiis 20-32 x 8-10 µm. Pilis margi-

nalibus longe clavatis, interdum inferne

ampullaceis, 37-60-(80) x 5-6 x 6-9 µm.

Epicute gelata, pilis erectis (ixotrichoder-

mate) longioribus x 3-4 µm, cylindratis,

haud vel parum summis incrassatis, bre-

vioribus (in strato inferiore) manifeste cla-

vatis (5-7 µm), sparsis; subcute magis

minusve manifeste specie pseudopa-

renchymatica. Fibulis praesentibus. – Sub

Pinus pinastris Quercubusque peduncula-

tis, locis graminosis, fossis luteis, ad

Lacum Aureilhan (Labdes), 13 et 17 X

1977. – Holotypus: 77.333 (Herb.

Romagnesi); 77.348”.

Observations on the material: The type

consists of a legthwise section of a speci-

men and fragments of another. The condi-

159

tions of the material did not allow for an

exhaustive study of the pileipellis, of the

discal zone in particular.

Personal study

Spores (10.4)-10.6-12.2-(12.6) x 6.4-7.2

µm., Q = (1.54)-1.58-1.79-(1.84); on aver-

age [72/2] mL 10.91 µm, mLr 10.64-

11,19 µm., mW 6.58, mWr 6.52-6.65

µm., mQ 1,65, mQr 1.63-1.68; morphol-

ogy mostly amygdaliform (81.9 %), more

rarely broadly amygdaliform (15.3 %),

occasionally subfusiform (2,8 %); not

mucronate, obovoid in front view; orna-

mentation subdistinct to distinct; myx-

osporium undilating; light-coloured,

greenish-grey [29-30A/B2]; subdextrinoid

[4C4-5; 5B/C4]; acyanophilous to

hypocyanophilous. Reduced pore hardly

visibile.

Basidia 28-33 x 8.0-10 µm., four-spored,

some with median constriction..

Lamellar edge sterile. Cheilocystidia

dimorphic, lageno-clavate, (36)-40-56-

(60) x (5.5)-6.5-8.5-(10) x (3.0)-4.0-5.5-

(6.0) x (5.5)-6.0-7.5-(9.0) µm., on average

[36/2] mL 48.15 µm., mLr 47.7-48.6 maB

7.5 µm., maBr 7.2-7.9 µm, mmB 4.7 µm,

mmBr 4.5-4.9 µm, mbB 6.7 µm, mbBr

6.5-7.0 µm, and clavato-cylindraceous

(44)-50-67-(70) x (6.0)-7.0-9.5-(13) x

(3.5)-4.8-6,0 µm., on average [37/2] mL

57.7 µm, mLr 57.5-57.9 µm, maB 8.35

µm., maBr 8.3-8.4 µm, mmB 5.1 µm.,

mmBr 5.0-5.2 µm.; apex with occasional

plaque-like thickenings.

Pleurocystidia none.

Caulocystidia similar to cheilocystidia, but

often much longer, (36)-41-113-(118) µm.

Pileipellis: suprapellis a c. 100-110 µm

deep loosely interwoven ixotomentocutis

of hypongophilous hyphae, 1,6-4,0-(5,0)

µm. wide; pigmentation intracellular and

epiparietal scanty; terminal hyphal cells

(34)-44-88-(96) µm., apex subcylindrical

to clavate (2,5)-3,0-6,0-(7,0). Subpellis

chromatically distinct, brownish-orange

[c. 4B/C5], more or less pseudoparenchy-

matic in structure.

Comments

The microanatomical picture offered

by the study of the type coincides rather

satisfactorily with that from Favre’s

description (FAVRE, 1960), that from the

relative voucher collections previously

studied (see SUPPLEMENTARY TABLE I and

GRILLI, 2004) and the one presented above.

Since Romagnesi in the diagnosis

emphasizes the shortness of the stipe

(“stipite brevi”), it was considered whether

H. favrei could individuate a short-stiped

variant, but a high variability in stipe

length, both within a single collection and

across the very ample supply of material

studied over the years, has been observed

(data not included in this study).

Furthermore, Favre’s material included

also long-stiped specimens (at least GK

9169); accordingly, also H. favrei seems to

fall within the variability range of H. leu-

cosarx.

Given the perfect coincidence of all

the basic qualitative features of spores ( i.e.

morphology, ornamentation pattern, myx-

osporium expansion, colour and dextri-

noidity) and of cystidia (morphology, mor-

phological types and occurrence of plaque-

like thickenings), it can be remarked that

also sporal and cystidial measures, vari-

able as they are across collections, do not

show any real hiatus and can be arranged

in a cline. Therefore, it would seem safe to

conclude that it is impossible to discrimi-

nate any of the taxa presented above on the

basis of both qualitative and quantitative

microanatomical traits.

The range of measures of all the vari-

ous microanatomical parameters (see

SUPPLEMENTARY TABLE I) is in broad agree-

ment with the ones reported by AANEN &

KUYPER (2004): e.g. spores 10.4-11.9 x

6.3-7.2 µm, Q 1.57-1.80, and VESTERHOLT

What is Hebeloma leucosarx ?

160 Grilli E.

Fig. 14 - H. favrei. Holotype. Spores & cheilocystidia. Bar 10 µm (spores); bar 30 µm

(cheilocystidia).

161

(2005): e.g. spores 9.5-12.1 x 5.6-6.9 µm,

Q 1.67-1.83, for H. velutipes.

Material studied

Hebeloma leucosarx

United Kingdom: Loch Loy, Morayshire

(Grampian), Scotland, Sept. 25,1955, P. D.

Orton 583 holotypus (K); Loch Loy,

Morayshire (Grampian), Scotland, Sept.

25, 1955, P. D. Orton 593 (E00076310)

(E); Bookham Common, Surrey, Oct. 15,

1955, P. D. Orton 596 (E00076300) (E);

Loch Rannoch S., Pertshire, Oct. 25, 1964,

Orton 2611 (E00076306) (E); Thack

Marsh, Surlingham, Norfolk, Sept. 6,

1969, P. D. Orton 1967 (E00076305)(E);

Hothfield Common, Kent, Sept. 22, 1981,

M. Moser 81/424 (IB).

Denmark: Marbaek Plantage N of Esbjerg,

Oct. 6, 1991, J. Vesterholt 91-509 (C).

France: Les Echets (Ain), Sept. 24,

1963, G. Bruchet 63.20 isotypus of H.

velutipes (duplum 00875 RO); ditto, G.

Bruchet 63.21 paratypus (LY); ditto,

Oct. 9, 1964, G. Bruchet 64.31 paraty-

pus (LY); ditto, Oct. 2, 1965, G. Bruchet

65.14 paratypus (LY); Forêt de l’Isertan,

Pralognan, Savoie, Sept. 10, 1964, G.

Bruchet 64.21 (as H. crustuliniforme)

(LY); Aureilhan (Landes), Oct. 13,

1977, H. Romagnesi 77.333 Holotypus

of H. favrei (PC).

Italy: Val Saviore (BS), July 31, 1996, G.

Consiglio 96021 (HG); Civago (RE), Sept.

25, 1998, G. Consiglio 98064 (HG);

Goriano Sicoli (AQ), Oct. 13, 1998, E.

Grilli 981013.1.1 (HG); Val Fondillo, Opi

(AQ), Sept. 20, 2003, E. Grilli 030920.2

(HG).

Hungary: Szakonyfalu, Com. Vas, Oct. 30,

1962 G. Bohus & M. Babos 37.743 (as H.

crustuliniforme var. minus) (BP).

Spain: Lleida, Font Roja Sept. 22, 1993, J.

Llistosella MA 1460; ditto, Sept. 23, 1993,

J. Llistosella MA 1461 and MA 1462 (as

H. helodes) (BBC).

Switzerland: Forêt de Chaubon, Haute

Savoie, Oct. 21, 1951, J. Favre, GK 9169

(as H. crustuliniforme var. bulbiferum)

(G); Forêt de Flin, Engadine, Aug. 28,

1954, J. Favre, GK 14784 (as H. crustulin-

iforme var. bulbiferum) (G); God God,

Engadine, Aug. 30, 1954, J. Favre, GK

14785 (as H. crustuliniforme var. bulbifer-

um) (G).

HEBELOMA LUTENSE Romagn. 1965

(Fig.s 15, 18).

Basyonym: Hebeloma lutense Romagn.,

Bull. Soc. Mycol. Fr. 81: 344; 1965

Holotypus: France. Forêt de Rambouillet,

Etang de la Tour, Oct. 11, 1959, H.

Romagnesi 59.232 (PC)

Description of type collection (H.

Romagnesi 59.232; PC).

ROMAGNESI, 1965: “Pileo 1,8-5,5 cm,

primum convexo, deinde espanso, saepe

esimie irregolari et ad marginem flexuoso

ac lobato, interdum circui albido, medio

castaneo vel saltem e fulgido ochraceo,

viscoso, glabro. Stipite curto, (1,5)-2-5,5

cm x 3,5-7 mm, cilindrato vel inferne levi-

ter incassato, candido, deinde paulum

brunnescente, fibrilloso, atque primum

fere toto farinoso. Carne medio crassa,

alba, paulum in stipite ex ochraceo rufulo

colore tincta, odore levi, potius cacaino

quam rapaceo, sapore amaro. Lamellis

distantibus, praesertim in veteribus speci-

minibus, latis vel atiam latissimis, (3)-4-10

mm, sinuatis, falciformibus vel ventricosis,

deinde segmentiformibus vel fere triangu-

laris, pallidissime brunneolis, deinde

umbrinis, acie esimie serrulata ac punctis

albis praedita, plorante. Sporsi in cumulo

umbrinis, 10-12-(12,5) x 5,2-6-(6,5) µm,

amygdaliformibus, pallide luteis s. m., ver-

ruculosis. Pilis acierum claviformibus,

interdum paulum haud procul a basi

ampullaceis, brevioribus quam apud popu-

linum, 35-57 x 6,5-10,5 µm. Epicute ex

hyphis filiformibus, 1,5-2,8 µm. Praeditis,

What is Hebeloma leucosarx ?

162 Grilli E.

cute hyphis amplioribus, sub cute specie

pseudoparenchymatica manifesta; pig-

mento membranari, interdum etiam incro-

stante, praesertim in hyphis connexivis

cutis subcutisque. – In limo stagni, prope

Salices et Betulas, inter Phragmites,

Carices ac Hydrocotylas. Typus in herb.

Romagnesiano, Mus. Nat. Hist. Nat. Paris,

n° 59.232.”

Observations on the material: The type

consists of at least 6 specimens, all in good

conditions, and allowed for an exhaustive

study of the diagnostic microanatomical

features.

Personal study

Spores (9.6)-10.4-12.4-(13.6) x (5.2)-5.6-

6.3-(7.0) µm., Q = (1.69)-1.79-2.03-

(2.21); on averge [137/4] mL 11.29 µm.,

mLr 10.95-11.93 µm., mW 5.91 µm.,

mWr 5.84-6.06 µm.; mQ 1.91; mQr 1.87-

1.96; morphology mostly subfusiform

(65.7%), less frequently fusiform (21.9%),

rarely amygdaliform (12.4%); ornamenta-

tion subdistinct to distinct; myxosporium

undilating; light-coloured, greenish-white

[29A/B2; 30B3]; subdextrinoid [4B/C4-5;

4C4-5]. Reduced pore more or less visible.

Basidia 28-44 x (6,0)-7,5-8,5 µm., four-

spored.

Subhymenium subcellular; hymenophoral

trama subregular.

Lamellar edge sterile. Cheilocystdia

dimorphic, prevalently (58%) clavato- to

capitato/swollen-pedicellate, (28)-31-64-

(70) x (6.0)-7.0-15-(19.5) x (2.0)-2.5-4.0-

(4.5) µm; on average [70/3] mL 52.8 µm.,

mLr 43.2-59.6 µm., maB 9.6 µm., maBr

7.0-11.9 µm., mmB 3.0 µm., mmBr 2.7-

3.4 µm, and lagenoclavate, (25)-33-50-

(56) x 6.0-8.0-(10) x (1.6)-2.5-4.0-(4.5) x

(4.5)-5.0-6.5-(8.0) µm., on average [50/3]

mL 39.4 µm., mLr 33.5-45.6 µm., maB

7.6 µm., maBr 7.2-7.9 µm., mmB 3.4

µm., mmBr 3.3-3.5 µm., mbB 5.8 µm.,

mbBr 5.5-6.0 µm.; pedicel frequently typ-

ically sinuous; no plaque-like thickenings

observed, apex often hypercongophilous

and slightly thick-walled, at times in the

form of a peanut pod or distinctly swollen.

Pleurocystidia none.

Caulocystidia morphologically similar to

hymenial ones, but not as inflated at the

apex, (32)-38-63-(108) µm long.

Pileocystidia along pileus margin abun-

dant, similar to hymenial ones.

Pileipellis: suprapellis a 100-140 µm deep

ixotomentocutis of hypocongophilous

hyphae, 1.6-4.8 µm wide, embedded in a

complete gel matrix; terminal hyphal cells

not very long, with subcylindrical to sub-

clavate apex; pigment intracellular and

epiparietal scanty; subpellis chromatically

distinct [c. 4C6 slightly more brownish]

from the pileitrama; distinctly pseudo-

parenchymatic in structure at the disc, with

large, subisodiametrical elements, up to

18-20 µm in diameter, but mixed to

hyphoid in the peridiscal zone; epiparietal

pigment incrustation thick and continuous

in the pseudoparenchymatic zone, discon-

tinuous in the hyphoid one (Fig. 15).

Additional collections

To have an idea of the microanatomical

variability pattern, it was deemed useful to

study also some additional material from

Romagnesi’s Herbarium at PC.

France. Exposition du Museum

(Rambouillet?) Oct. 6, 1967, H.

Romagnesi 67.217 (0022095; PC).

Observation on the material: The name

Hebeloma lutense on the herbarium label

is crossed out in pencil. The collection,

made “dans une androit marécageux”,

consists of a single specimen and a spore

print. The crossing out of the name would

imply that Romagnesi had second

thoughts about the correct identification

of the fungus, but the microanatomy of

163

What is Hebeloma leucosarx ?

the specimen is absolutely comparable to

that of the type.

Spores (9.4)-9.8-11.2-(12) x (5.2)-5.6-6.4

µm, Q (1.63)-1.73-1.87-(2.0); on average

(64/2) mLr 10.42-10.71 µm, maWr 5.68-

5.77µm, mQr 1.83-1.85; morphology sub-

fusiform (50%) to amygdaliform (45.3%),

rarely fusiform; ornamentation distinct;

myxosporium undilating or occasionally

slightly vesiculate at the base; light-

coloured, greenish-white [29A/B2]; sub-

desxtrinoid to dextrinoid [5D5; 6C7].

Lamellar edge sterile. Cheilocystidia

dimorphic, mostly (65.5%) clavato- to

swollen-pedicellate, (40)-48-74-(78) x 6.5-

13-(15) x (2.0)-2.5-3.5-(4.0) µm, on aver-

age (19/1) mL 62.1 µm, maB 10.3 µm,

mmB 3.2 µm, and lageno clavate, (38)-42-

46-(62) x (6.0)-7.0-8.0 x 2.5-3.5-(4.0) x

(4.5)-5.0-7.0-(7.8) µm, on average (10/1)

mL 45.4 µm, maB 7.2 µm, mmB 3.1 µm,

mbB 5.9 µm; pedicel typically sinuous.

Pleurocystidia none.

Caulocystidia and pileipellis not studied.

Comparison of H. lutense with “speci-

men 2” and the voucher material of H.

leucosarx ss Bruchet 1970

“Specimen 2” of the type of Hebeloma leu-

cosarx plus P.D. Orton 596.

Spores (9.6)-10.4-12.2-(13.6) x (5.4)-5.6-

6.4-(7.0) µm, Q (1.55)-1.70-2.10-(2.14);

on average (202/5/2) mLr 10.93-11.53

µm, maWr 6.01-6.07 µm; mQr 1.75-1.90;

morphology often subfusiform (55.4%),

less frequently amygdaliform (36.1%),

rarely fusiform (7.4%), occasionally

broadly amygdaliform; not mucronate;

ornamentation subdistinct to distinct; myx-

osporium undilating to rugulose, occasion-

ally shallowly vesiculate; light-coloured;

subdextrinoid. Reduced pore more or less

visible.

Lamellar edge sterile. Cheilocystidia

dimorphic, mostly (62.7%) clavato- to cap-

itato/swollen-pedicellate, (28)-36-64-(76)

x (5.0)-6.0-10-(15) x (2.0)-2.8-4.0-(5.0)

µm, on average (170/5/2) mLr 42.1-55.8

µm, maBr 7.5-8.3 µm, mmBr 2.8-3.3

µm; less frequently (37.3%) lageno-

clavate, (25)-30-53-(74) x (4.5)-6.0-8.5-

(10) x (2.0)-2.5-4.0-(4.5) x (4.5)-5.0-6.5-

(7.5) µm; on average (101/5/2) mLr 38.5-

46.2 µm, maBr 6.6-8.0 µm, mmBr 2.7-

3.4 µm, mbBr 5.1-5.9 µm; apex not infre-

quently hypercongophilous, pedicel often

typically sinuous to tortuous.

Pleurocystidia none.

Caulocystidia not studied.

Pileipellis: suprapellis a 90-120 µm deep

loosely interwoven ixotomentocutis,

hyphae (1,6). 2.4-4.0-(5.6) µm wide,

embedded in a complete gel matrix; termi-

nal hyphal cells 34-94 x 3.0-5.0-(6.0) µm,

cylindrical to subclavate, occasionally

cystidiform; pigment mostly intracellular,

occasionally epiparietal in the lower lay-

ers; subpellis more or less chromatically

distinct [2A/B3; 3B5] in comparison to the

pileitrama [29A2], pseudoparenchyma-

tous in structure.

Comments

There seems to be good correspon-

dence between the microscopical descrip-

tions of H. lutense and that of “specimen

2” above: the agreement is satisfactory for

what concerns qualitative and quantitative

characters alike.

H. leucosarx ss Bruchet (1970)

(Fig. 18).

France. “Étang du Grand Albert”, Saint-

Jean-de-Bournay (Isere), Oct. 15, 1964,

Bruchet 64.32 (LY).

Observations on the material: the collec-

tion consists of fragments of two speci-

mens which did not allow for an exhaus-

tive study.

164 Grilli E.

Fig. 15 - H. lutense. Holotype, Spores & cheilocystidia. Bar 10 µm (spores); bar 30 µm

(cystidia).

165

What is Hebeloma leucosarx ?

Fig. 17 - Hebeloma lutense Romagn. (M. Contu 981013.1, M. Limbara, SS, Sardinia).

Fig. 16 - Hebeloma lutense Romagn. (M. Contu 970912.2, M. Limbara, SS, Sardinia).

166 Grilli E.

Personal study

Spores (10)-10.8-12.4-(13.6) x (5.4)-5.6-

6.2-(6.6) µm, Q (1.64)-1.82-2.07-(2.31),

on average (62/2) mLr 11.34-11.70 µm;

mWr 5.83-5.96 µm; mQr 1.88-2.00; mor-

phology mostly subfusiform (50%), less

frequently fusiform (40.3%), rarely amyg-

daliform (9.7%); ornamentation subdis-

tinct to distinct; myxosporium mostly

undilating, rarely slightly vesiculate at the

base; light-coloured, greenish-grey

(29A/B2-3; 30A/B2-3); subdextrinoid

(4C4). Reduced pore more or less visible.

Lamellar edge sterile. Cheilocystidia

dimorphic, prevalently (58.7%) clavato- to

capitato-pedicellate, (35)-46-70-(78) x

(5.0)-6.0-9.8-(13) x 2.0-4.0 µm; on aver-

age (44/2) mLr 54.3-57.3 µm, maBr 7.2-

7.3 µm, mmBr 3.0 µm, and lagenoclavate,

(32)-36-48-(50) x (5.0)-5.8-8.0-(9.0) x

(2.0)-3.0-4.0-(5.0) x 5.0-8.0-(9.0) µm, on

average (31/2) mLr 39.4-41.3 µm, maBr

6.2-7.0 µm, mmBr 3.4-3.8 µm, mbBr 5.8-

6.7 µm; pedicel often sinuous to tortuous.

Pleurocystidia none.

Caulocystidia and Pileipellis not studied.

Comments

The micromorphological picture

above is much the same as that of H.

lutense Romagn and that of the second

element of Orton’s type. Therefore, since

also the gross morphological features as

described by Bruchet are perfectly com-

parable, based on the result of this study

H. leucosarx as conceived by BRUCHET

(1970) should now be regarded as a mis-

application. Bruchet must have based his

conception on the study of the second

element of Orton’s type; he, in fact, sur-

mised thart H. leucosarx and H. lutense

might be very close: “L’étude micro-

scopique et anato-mique du type milite

d’ailleurs en faveur du rapprochement

de ces deux espèces”; a surmise that

became certainty in QUADRACCIA (1993):

“Incidentally, our study of the holotype

of H. lutense Romagn. persuaded us to

regard it as a later synonym of H. leu-

cosarx”.

On the basis of the relative collection

(Brenns, Étang du Grand Migner, Sept. 17,

1971, Bon-Delzenne 71091704) also the

H. leucosarx briefly described and com-

mented in BON & al (1972) has to be

assigned to H. lutense. For the sake of

brevity the description is not reported, but

the quantitative data is included in

SUPPLEMENTARY TABLE II.

For the range of measures of all the

microanatomical parameters of H. lutense

see SUPPLEMENTARY TABLE II.

In Index Fungorum (CABI Bioscience

UK), H. lutense is reported as a later syn-

onym of Hebeloma cavipes Huijsman,

most likely on the basis of VESTERHOLT

(1995): “H. lutense is probably identical to

H. cavipes”, but the synonymyzation

would seem impractical from a phenetic

point of view. Based on my unpublished

type revisions, in fact, H. cavipes

Hujisman 1961 belongs in the Hiemale-

complex – another taxonomic-nomenclat-

ural tangle including, apart from H.

hiemale Bres. 1898 (GRILLI, 1997), H.

album Peck 1901, H. fragilipes Romagn.

1965 and a number of other taxa both

European and American - and is morpho-

logically well distinguished from H.

lutense as characterized above.

The same stance has been adopted by

AANEN & KUYPER (2004), again maybe on

the grounds of Vesterholt’s 1995 opinion

and of molecular evidence from a culture

collection of H. cavipes sharing the same

ITS sequence as collections of H. hiemale

and H. lutense, taxa which are all listed as

synonyms of H. helodes J. Favre.

Apparently, such a point of view has

recently received further support from

BOYLE & al (2006); their phylogenetic

inferences, however, are based on the same

GenBank sequences of H. cavipes, H.

hiemale and H. lutense generated by

167

What is Hebeloma leucosarx ?

AANEN & al (2000) which gave molecular

confirmation to the two Dutch authors’

synonymy assumption.

However, VESTERHOLT (2005), with

whom I had an exchange of information on

this very species prior to the publication of

his latest revision of the genus, now con-

sider H. lutense a good species, quite dis-

tinct from both H. hiemale and H. helodes,

even if he does not include or cite H.

cavipes among the species presented. I

have also studied one of his collections

(JV96-231) and have found it congruent

with the circumscription defined above

(data not shown, but see SUPPLEMENTARY

TABLE II).

Hebeloma lutense is rather similar in

gross morphological features to

Hebeloma leucosarx. In the field, diag-

nostic clues may be the stipe base nor-

mally not bulbous or only slightly

enlarged, the less crowded lamellae (a

feature which should be confirmed and

better defined on the basis of further col-

lections) and the generally more colourful

(cinnamon, sienna, rust, brick) or darker

(hazel, at times mimicking H.

mesophaeum) pileal surface. However,

some specimens might prove difficult to

separate with certainty without recourse

to microscopy. In addition, from the liter-

ature it would seem that the former has

more specialized ecological requirements

in comparison with H. leucosarx, which

is associated with a very broad range of

photoautotrophic hosts. In fact, H. lutense

appears to have a preference for damp

sites among “des Phragmites, Carex,

Hydrocotyle et autres plantes des

marécages” (ROMAGNESI, 1965) in associ-

ation especially with Salix (Vesterholt, in

litt.). Nevertheless, Romagnesi’s fungus

has been sparingly recorded in the litera-

ture and its host range might be wider, as

demonstrated by the two Sardinian col-

lections, from a humid, grassy site in a

Pinus laricium stand at an altitude of 700

m., presented in this study. (Fig.s 16, 17).

Selected illustrations: LUCCHINI

(1997), as H. leucosarx; LACHIUSA

(1999), as H. leucosarx; VESTERHOLT

(2005). For the first two references, my

conclusions are based on the study of the

relative voucher material: G. Lucchini

6084 (LUG) and L. La Chiusa 774

(MCVE), data not shown in full (see

SUPPLEMENTARY TABLE II). The species is

also illustrated in COURTECUISSE & DUHEM

(1994), but the description does not

include any microanatomical details, and

those presented in COURTECUISSE & al

(1986) are not very typical. Regrettably,

despite my repeated requests I have not

been able to study the relative collection

housed at LIP.

Collections studied

France: Forêt de Rambouillet, Etang de la

Tour, Oct. 11, 1959, H. Romagnesi 59.232

holotypus (PC); Exposition du Museum

(Rambouillet ?), Oct. 6, 1967, H.

Romagnesi 67.217 (PC 0022095); Brenns,

Étang du Grand Migner, Sept. 17, 1971,

Bon-Delzenne 71091704 (LIP);

Luzarches, Côte-d’Or., Sept. 27, 1978, H.

Romagnesi 78.179 (PC 0022094), as H.

lutense ? (= H. album Peck); “Étang du

Grand Albert”, Saint-Jean-de-Bournay

(Isere), Oct. 15, 1964, Bruchet 64.32 (as

H. leucosarx) (LY); Forêt de Seillon,

Bourg (Ain), Oct. 19, 1964, Bruchet 64.39

(as H. leucosarx) (LY);

Denmark: Sjapmose, Varming Plantage E

of Ribe, Sept. 28.1996, J. Vesterholt

JV96-231.

Luxembourg: Forêt de Stadt-Predimus,

Sept. 21, 1976, H. Romagnesi 76-93 (PC

0088811) as H. cavipes?.

Italy: Cernusco (MI), Oct. 10, 1996, L. La

Chiusa 774 (as H. leucosarx) (MCVE);

Sardinia, M. Limbara, Tempio Pausania

(SS), Sept. 12, 1997, M. Contu 970912.2

(HG); ditto, Oct. 13, 1999, M. Contu

981013.1 (HG).

168 Grilli E.

Fig. 18 - H. leucosarx ss Bruchet. G. Bruchet 64.32 (LY). Spores & cheilocystidia. Bar 10

µm (spores); bar 30 µm (cheilocystidia).

169

What is Hebeloma leucosarx ?

Switzerland: Tenero (Bolle di Magadino),

bank of the Verzasca river, Sept. 17, 1986,

G. Lucchini 6084 (as H. leucosarx)

(LUG); Locarno, Oct. 6, 1988, G.

Lucchini 6362 (as H. crustuliniforme)

(LUG).

United Kingdom: Scotland, Loch Loy,

Morayshire (Grampian), Sept. 25, 1955, P.

D. Orton 583 (“specimen 2” of the type of

H. leucosarx) (K); Bookham Common,

Surrey, Oct. 15, 1955, P. D. Orton 596 (as.

H. leucosarx) (E).

Conclusions

The revision of the types and addition-

al material presented above confirms that,

within a morphological framework, H.

leucosarx can be distinguished from H.

lutense on account of the bulbous stipe

base, the pileus generally paler, (but in

some cases the colour may be very simi-

lar), crowded lamellae, more densely pru-

inose stipe and, above all, the differently

shaped cheilocystidia, longer caulocys-

tidia and less elongate spore morphology.

It also provides supportive evidence that

Hebeloma bulbosum Bres. (inedit.), H.

velutipes Bruchet and H. favrei Romagn.

ex Quadr. are hardly separable from H.

leucosarx as circumscribed and lectotypi-

fied in this study, and reveals that also H.

crustuliniforme ss Bruchet (1970) and H.

crustuliniforme (Bull.) Quél. var. minus

(sic) Cooke ss Bohus (1972) fall within its

circumscription, and that they have conse-

quently to be regarded as cases of misap-

plications. Likewise, it shows that the

microanatomy of H. lutense is perfectly

comparable to that of the second element

in Orton’s type and to that of the voucher

material of H. leucosarx ss Bruchet,

which, accordingly, is to be considered a

misapplication as well.

The synonymy of H. lutense with H.

helodes proposed by AANEN & KUYPER

(2004) is not accepted on the basis of

present data. In fact, even if the two taxa

may be partly macro- and micromorpho-

logically similar – especially for what

concerns the sinuous morphology of the

pedicels of cheilocystidia – and the “mor-

phological quality” of the separating

traits (smaller, almost indextrinoid spores

and cystidia more regularly capitate with

distinctly hypercongophilous thick-

walled capitulum) is not always unequiv-

ocal, these features do, however, provide

an element of doubt. A doubt that is rein-

forced by the facts that Aanen & Kuyper

include H. helodes ss KEIZER & ARNOLDS

(1995) among the misapplications of H.

helodes as conceived by the latter

authors, because allegedly conspecific

with H. velutipes [while based on the rel-

ative voucher collections (Keizer 86173

and 87126 WAG-W), H. helodes ss

Keizer & Arnolds certainly fall within the

Hiemale-complex: H. hiemale Bres. and

H. fragilipes Romagn respectively], and

that VESTERHOLT (2005), who has refined

his species concepts in light of sequence

data, regards H. lutense as an autonomous

species.

It would seem reasonable to conclude

that placing H. velutipes and H. favrei in

synonymy with H. leucosarx is a viable

option, and that the latter name is the one

to be adopted, at least among the taxa

described from Europe, for this very com-

mon Hebeloma species of the section

Velutipes. However, anticipating the

results of an unpublished companion study

(GRILLI, in prep.), it has to be remarked

that two American taxa: Hebeloma bakeri

Earle (1901), described from Stanford,

California, and Hebeloma exiguifolium

Murrill (1917), described from Pittsford,

New York, are potentially serious earlier

nomenclatural rivals.

Having been treading the thin line

which marks the limit of the taxonomic

possibilities of phenetic resolution, I am

well aware that the morphological

hypotheses suggested in the present study,

170 Grilli E.

as well as in the companion articles in

preparation, demand conclusive evidence

from molecular data.

AKNOWLEDGEMENTS

The directors and the curators of the

herbaria of Kew (K), Edinburgh (E),

Barcelona (BBC), Budapest (BP), Geneve

(G), Innsbruck (IB), Lugano (LUG), Lyon

(LY), Paris (PC), Rome (RO), Trento (TR)

and Venice (MCVE) are thanked for the

loan of the types, original material and/or

other collections analyzed in the study. I

am grateful to J. Vesterholt, G. Consiglio

and M. Contu for providing interesting

collections. G. Consiglio and M. Contu are

also thanked for allowing me to publish

their photographs and J. Vesterholt, for

suggestions and helpful comments on the

manuscript. Finally, I have to express my

gratitude to Th. Kuyper for supplying rele-

vant literature, and to G. Pacioni and G.

Lalli (AQUI) for their steady support and

for arranging loans.

Colour Codes

Flora of British Fungi. Colour Identification Chart. Edinburgh 1969

KORNERUP A. & J.H. WANSCHER,1978: Methuen Handbook of Colour. Eyre Methuen,

London.

REFERENCES

AANEN D. K. & T. KUYPER, 1999: Intercompatibility tests in the Hebeloma crustulini-

forme complex in Northwestern Europe. Mycologia 91: 783-795.

AANEN D. K. & KUYPER TH. W., 2004: A comparison of the application of a biological

and phenetic species concepept in the Hebeloma crustuliniforme complex

within a phylogenetic framework. Persoonia 18: 285-316.

AANEN D. K., T. KUYPER, T. BOEKHOUT & R. F. HOEKSTRA, 2000a: Phylogenetic relation-

ships in the genus Hebeloma based on ITSI and 2 sequences, with special

emphasis on the Hebeloma crustuliniforme complex. Mycologia 92(2):

269-281.

AANEN D. K., T. KUYPER & R. F. HOEKSTRA, 2001: A widely distributed ITS polymor-

phism within a biological species of the ectomycorrhizal fungus Hebeloma

velutipes. Mycol. Res. 105(3): 284-290.

AANEN D. K., T. KUYPER, TED H. M. MES & R. F. HOEKSTRA, 2000b: The evolution of

reproductive isolation in the ectomycorrhizal Hebeloma crustuliniforme

aggregate (Basidiomycetes) in northwester Europe: a phylogenetic

approach. Evolution 54(4): 1192-1206.

AGAPOW P.-M., BININDA-EDMONDS O. R.P., CRANDALL K. A., GITTLEMAN J. G., MACE G.

M., MARSHALL J. C. & PURVIS A., 2004: The impact of species concept on

biodiversity studies. The Quarterly Review of Biology 79 (2): 161-179.

ALBERS J. & B. GRAUWINKEL, 2005: Zur Pilzflora der Weserinsel Strohauser Plate (2).

Beitr. z. Kenntn. Pilze Mitteleuropas XIV: 9-42.

171

What is Hebeloma leucosarx ?

BAS C., 1988: Orders and families in agarics and boleti. In “Flora Agaricina Neerlan-dica

Vol. I” (Bas C., Kuyper Th., Noordeloos M. E. & Vellinga E. eds), pp. 40-

49. A.A.Balkema/Rotterdam/Brookfield.

BASTARDO DEL VAL J., A. GARCIA BLANCO & M. SANZ CARAZO, 2001: Hongos-Setas en

Castilla y León.

BOEKHOUT T. & KUYPER TH. W., 1995: Hebeloma. In “Overzicht van de Paddestoelen in

Nederland” (ARNOLDS E., KUYPER TH. W. & NOORDELOOS M. N. eds), pp.

217-226. Nederlandse Mycol. Vereniging. Wijster,

BASTARDO DEL VAL J., GARCIA BLANCO A.& SANZ CARAZO M., 2001: Hongos-Setas en

Castilla y León.

BOHUS G.,1972: Hebeloma studies I. Annls. hist. nat. Mus. natn. Hung.64:71-78.

BON M., 1972: Itineraire mycologique dans le pays de Brenne (Suite). Doc. Mycol. 1(4):

29-42.

BON M., 1993: Espèces rares, intéressantes, ou nouvelles étudiées aux journées

mycologiques Phoecennes de 1992. Bull. Féd. Ass. Mycol. Médit. 2(4):

4-14.

BREITEMBACH J. & F. KRÄNZLIN, 2000: Champignons de Suisse. Tome 5 Agaricales

(Champignons à lames) 3éme partie. Cortinariaceae. Lucerne. Édition

Mycologia

BRIDGE P. D., SPOONER B. M., ROBERTS P. J. & PANCHAL G., 2003: On the unreliability of

published DNA sequences. New Phytologist 160 (1): 43-48.

BRIDGE P. D., SPOONER B. M. & ROBERTS P. J., 2004: Reliability and use of published

sequence data. New Phytologist 16 (1): 15-17.

BRUCHET G.,1970: Contribution a l’étude du genre Hebeloma (Fr.) Kummer; Partie spe-

ciale. Supp. Bull. Soc. Linn. Lyon 39(6):1-132.

COURTECUISSE R. & B. DUHEM, 1994: Les Champignons de France. Guide encyclopé-

dique. Eclectis.

COURTECUISSE R., J. P. PRIOU & P. BOISSELET,1986: Contribution à la connaissance de la

flore fongique du Morbihan et de quelques départements voisins - I. Doc.

Mycol.16(62): 1-22.

ENDERLE M., 2004: Die Pilzflora des Ulmer Raumes. Verein für Naturwissenschaft und

Mathematik in Ulm e. V.

FAVRE J.,1948: Les associations fongiques des hautes marais jurassiens et de quelques

régions voisines.Matériaux Flore Cryptog.-suisse,10(3). Berne.

FAYOD V., 1892: Censimento dei funghi osservati nella valli valdesi del Piemonte duran-

te i mesi di Agosto-Ottobre del 1885-87. Annali R. Accad. d’Agric. 35(6):

81-114.

FRANCIS AA & BOUGHER NL, 2002 [2003]: Historical and current perspectives in the sys-

tematics of Australian cortinarioid sequestrate (truffle-like) fungi.

Australasian Mycologist 21(3): 81-93.

172 Grilli E.

GERHARDT E., VILA J. & LLIMONA X., 2000: Hongos de España y de Europa. Ediciones

Omega. Barcellona.

GRILLI E.,1994: Specie interessanti o rare della micoflora abruzzese. Micol. e Veget.

Medit. 9(1): 3-12.

GRILLI E, 2004 : Type studies in Hebeloma. On Hebeloma bulbiferum and H. colossus.

Micol. e Veget. Medit. 19(2): 83-107.

GRILLI E., 2006: Type studies in Hebeloma. H. angustispermum, an earlier synonym of H.

cylindrosporum. Micol. e Veget. Medit. 21(1): 3-34.

GRILLI E., (in prep.): Type studies in Hebeloma. On some little-known North-American

species described by S.F. Earle and W. A. Murrill.

HAWKSWORTH D.L., 2004: “Misidentifications” in fungal DNA sequence databanks. New

Phytologist 161(1): 13-15.

HIBBET D.S., BINDER M., BISCHOFF J.F., BLACKWELL M., CANNON P. F., ERIKSSON O. E.,

HUHNDORF S., JAMES T., KIRK P. M., LÜCKING R., THORSTEN LUMBSCH H.,

LUTZONI F., MATHENY P. B., MCLAUGHLIN D.J., POWELL M. J., REDHEAD S.,

SCHOCH C. L., SPATAFORA J. W., STALPERS J. A., VILGALYS R., AIME M. C.,

APTROOT A., BAUER R., BEGEROW D., BENNY G. L., CASTLEBURY L. A.,

CROUS P. W., DAI Y.-C., GAMS W., GEISER D. M., GRIFFITH G. W., GUEIDAN

C., HAWKSWORTH D. L., HESTMARK G., HOSAKO K., HUMBER R. A., HYDE

K. D., IRONSIDE J. E., KÖLJAG U., KURTZMAN C.P., LARSSON K.-H.,

LICHTWARDT R., LONGCORE J., MIADLIKOWSKA J., MILLER A., MONCALVO J.-

M., MOZLEY-STANDRIDGE S., OBERWINKLER F., PARMASTO E., REEB V.,

ROGERS J. D., ROUX C., RYVARDEN L., SAMPAIO J. P., SCHÜ‚LER A.,

SUGIYAMA J., THORN R. G., TIBELL L., UNTEREINER W. A., WALKER K., YAO

Y.-J. & ZHANG N., 2007: A higher-level phylogenetic classification of the

Fungi. Mycol. Res. 111: 509-547.

HOLST-JENSEN A., VRÅLSTÅD T. & SCHUMACHER T., 2004: On reliability. New Phytologist

161 (1): 11-13.

KEIZER P.J. & E. ARNOLDS, 1996: Taxonomical notes on macrofungi on roadside verges

planted with trees in Drenthe (2). Persoonia 16(1): 81-122.

KELLER G. & M. MOSER, 2001: Die Cortinariaceae Österreichs. Catalogus Florae

Austriae, III. Teil, Pilze, Heft 2, Agaricales: Cortinariaceae. Österreichi-

sche Akademie der Wissenschaften. Wien.

KIRK P. M., P. F. CANNON, J. C. DAVID & J. A. STALPERS, 2001: Ainsworth & Bixby’s

Dictionary of the Fungi. CABI Bioscience, U. K.

KONRAD P. & A. MAUBLANC,1924-1937: Icones selectae fungorum.Voll.I-VI. Paris.

KORF R. P., 2005: Reinventing taxonomy: a curmadgeon’s view of 250 years of fungal tax-

onomy, the crisis in biodiversity, and the pitfalls of the phylogenetic age.

Mycotaxon 93: 407-415.

KREISEL H., 1987: Pilzflora der Deutschen Demokratischen Republik. Basidiomycetes

(Gallert-, Hut- und Bauchpilze). Jena. Gustav Fischer Verlag.

173

What is Hebeloma leucosarx ?

LACHIUSA L., 1999: Contributo allo studio del genere Hebeloma. RdM. 42 (3): 221-240.

LANGE J.E.,1938: Flora agaricina danica.Vol.3.København.

LANZONI G., 1989: Hebeloma longicaudum (Pers.: Fr.) P. Kumm. RdM 31(5-6): 352 (back

cover).

LEGON, N.W. & A. HENRICI, (with ROBERTS P.J., SPOONER B.M. & WATLING R.). 2005.

Checklist of the British & Irish Basidiomycota. Royal Botanic Gardens,

Kew. 517 pgs.

LLISTOSELLA J., M. AGUASCA & J. VILA, 1996: Basidiomycets de les torberes dels Pirineus

catalans. Primera contribució. Rev. Catalana Micol. 19 (1/3): 67-72.

LOCQUIN M., 1977 ( ?1979): Flore Mycologiques, 3. Cortinariales-A. Le Guyot, Paris.

LUCCHINI G., 1997: I funghi del Cantone Ticino e di altre regioni svizzere ed estere con-

servati al Museo di Storia Naturale. Ed. E. Lucchini-Balmelli, CH-

Gentilino.

MACNEIL JF, BARRIE F, BURDET HM, DEMOULIN V, H AWKSWORTH DL, MARHOLD K,

NICOLSON DH, PRADO J, SILVA PC, SKEG JE, WIERSEMA J & TURLAND NJ

(eds), 2006: International Code of Botanical Nomenclature (Vienna code).

ARG Gauter Verlag, Ruggel.

MATHENY B. P, CURTIS J. M., HOFSTETTER V., AIME M. C., MONCALVO J.-M., YANG Z.-W.

G. Z.-L.; SLOT J. C., AMMIRATI J. F., BARONI T. J., BOUGHER N. L., HUGHES

K. W., LODGE D. J., KERRIGAN R. W., SEIDL M. T., AANEN D. K., DENITIS

M., DANIELE G. M., DESJARDIN D. E., KROPP B. R., NORVELL L. L., PARKER

A., VELLINGA E. C., VILGALYS R. & HIBBETT D. S., 2006: Major clades

of Agaricales: a multilocus phylogenetic overview. Mycologia 98 (6):

982-995.

MONCALVO J. M., LUTZONI. F. M., REHNER S. A., JOHNSON J. & VILGALYS R., 2000:

Phylogenetics relationships of agaric fungi based on nuclear large subunit

ribosomal DNA sequences. Syst. Biol. 49 (2): 278-305.

MONCALVO J. M., VILGALYS R., REDHEAD S. A., JOHNSON J. E., JAMES T. Y., AIME M. C.,

HOFSTETTER V., VERDUIN S. J. W., LARSSON E., BARONI T. J., THORN R. G.,

JACOBSSON S., CLEMENÇON H. & MILLER O. K., 2002: One hundred and

seventeen clades of euagarics. Mol. Phylogenet. Evol. 23: 357-400.

MOSER M. & U. PEINTNER,1985-2007: Farbatlas der Basidiomyceten (1-24). Spektrum

Akademische Verlag. G. Fischer. Stuttgart - New York.

ORTON P.D.,1960: New check list of British agarics and boleti 3. Notes on genera and

species in the list. Trans. Brit. mycol. Soc. 43: 159-439.

PEINTNER U., BOUGHER N. L., CASTELLANO M. A., MONCALVO J-M., MOSER M., TRAPPE J.

M. & VYLGALYS R., 2001: Multiple origins of sequestrate fungi related to

Cortinarius (Cortinariaceae). Am. Journ. Bot. 88: 2168-2179.

PEINTNER U., LADURNER H. & SIMONINI G., 2003: Xerocomus cisalpinus sp. nov., and the

delimitation of species in the X. chrysenteron complex based on morphol-

ogy and rDNA-LSU sequences. Mycol. Res. 107 (6): 659-679.

174 Grilli E.

ROMAGNESI H.,1965: Étude sur le genre Hebeloma. Bull. Soc. Mycol. Fr.81(3): 321-344.

ROMAGNESI I H.,1983: Études sur le genre Hebeloma. II. Sydowia 36: 255-268.

SMITH A.H.,1984: Studies of species of Hebeloma (Fr.) Kummer from the Great Lakes

Region of North America, I. Sydowia 37: 271-283.

TAYLOR J.W., JACOBSON D.J., KROKEN S., KASUGA T., GEISER D.M., HIBBETT D.S. &

FISHER M.C., 2000: Phylogenetic species recognition and species concepts

in fungi. Fungal Genetics and Biology 31: 21-32.

THOMAS K. A., PEINTNER U., MOSER M. M. & MANIMOHAN P., 2002: Anamika, a new myc-

orrhizal genus of Cortinariaceae from India and its phylogenetic position

on ITS and LSU sequences. Mycol. Res. 106 (2): 245-251.

VESTERHOLT J.,1992: Almindelig Tåreblad (Hebeloma crustuliniforme) og dens almin-

delige dobbeltgængere. Svampe 25: 15-22.

VESTERHOLT J.,1995: Hebeloma crustuliniforme and related taxa - notes on some charac-

ters of taxonomic importance. Acta Univ. Ups. Symb. Bot. Ups. 30(3): 129-

137.

VESTERHOLT J, 2000: Hebeloma crustuliniforme and related species. Field Mycology 1(2):

58-68.

VESTERHOLT J., 2004: The identity of Hebeloma fastibile, the type species of Hebeloma.

Ann. Micol. A.G.M.T. 1: 53-63.

VESTERHOLT J., 2004a: Danmarks Svampe. Gyldendal. Københaven.

VESTERHOLT J., 2005: The genus Hebeloma. Fungi of Northern Europe – Vol. 3.

Svampetryk.

VILGALYS R., 2003: Taxonomic misidentification in public DNA databases. New

Phytologist 160 (1): 4-5.

WHEELER Q. D., 2004: Taxonomic triage and the poverty of phylogeny. Phil. Trans. R.

Soc. London B 359: 571-583.

WHEELER Q. D., RAVEN P.H. & WILSON E.O., 2004: Taxonomy: Impediment or Expedient?

Science 303 (5656): 285.

YANG Z. L., MATHENY P. B., GEZ-W., SLOT J. C. & HIBBETT D., 2005: New Asian species

of the genus Anamika (euagarics, hebelomatoid clade) based on morpholo-

gy and ribosomal DNA sequences. Mycol. Res. 109 (11): 1259-1267.

175

What is Hebeloma leucosarx ?

(*&&#!$!%) '+ ) #! " $"$'*(! '$.#*,!+/ 8$ 1$ 7OQML!

;NMO@J @LC BVPQHCH@J PHWD @BOMPP QGD BMJJDBQHMLP NODPDLQDC HL QGD PQRCV" NJRP QGD SMRBGDO

K@QDOH@J ME #+.,-.'&)&%*+($ S@O$ (&)., 0MMID DK$ :D@ PP /MGRP /75=;" &.,'!$ <GD

C@Q@ ODF@OCHLF /MGRPX @LC /ODP@CMJ@XP K@QDOH@J HP LMQ PGMTL HL ERJJ HL QGD PQRCV$

;NMODP

0MJJDBQHMLP PQRCHDC K6O K?O K9O

5$ JDRBMP@OU QVND N@O@QVNDP &%$-#&&$. +$)#+$* &$+-#&$-(

5$ JDRBMP@OU PP >DPQDOGMJQ &%$,#&%$- +$&#+$( &$,%#&$,,

5$ ARJAHEDORK PP 3@SOD &%$'#&&$( +$(#+$- &$+%#&$++

5$ BORPQ$ S@O$ KHLRP PP /MGRP &&$( +$) &$,+

5$ E@SODH QVND &%$+#&&$' +$*#+$+ &$+(#&$+-

5$ SDJRQHNDP HPM# N@O@QVNDP &%$'#&&$' +$&#+$) &$+(#&$,*

5$ BORPQRJHLHEMOKD PP /ORBGDQ .$- +$& &$+%

5$ ARJAMPRK /ODP$ &%$&#&&$) +$'#+$- &$+'#&$,(

24#%(%.'%$' &%$,#&&$' +$)#+$+ &$+-#&$,%

4$ 0MLPHFJHM .+%'& &&#&&$( +$'#+$) &$,)#&$-'

7SDO@JJ O@LFD .$-#&&$. +$&#+$- &$+%#&$-(

6@FDLMBJ@S@QD BVPQHCH@

0MJJDBQHMLP PQRCHDC K6O K@/O KK/O KA/O

5$ JDRBMP@OU QVND N@O@QVNDP *&$+#*)$+ ,$&#,$- )$+#)$- ,$&#,$+

5$ JDRBMP@OU PP >DPQDOGMJQ )'$%#)+$* +$+#,$) )$'#)$+ +$-#,$%

5$ ARJAHEDORK PP 3@SOD )+$-#+)$. -$&#.$) )$)#)$- +$-#,$'

5$ P@BB$ S@O$ KHLRP PP /MGRP *($' ,$( )$( +$)

5$ E@SODH QVND ),$,#)-$+ ,$'#,$. )$*#)$. +$*#,$%

5$ SDJRQHNDP HPM# N@O@QVNDP (-$&#++$) +$&#-$- )$(#)$, +$*#,$(

5$ BORPQRJHLHEMOKD PP /ORBGDQ *-$& ,$& )$% ,$-

5$ ARJAMPRK /ODP$ )+$*#*.$) ,$*#-$& )$%#)$. +$)#,$(

24#%(%.'%$' *'$-#*)$( -$+#.$' *$' ,$,#,$.

4$ 0MLPHFJHM .+%'& *-$-#+* -$-#.$' )$%#*$& +$-#+$.

7SDO@JJ :@LFD (-$&#++$) +$&#.$) )$%#*$' +$)#,$.

0J@S@QMBVJHLCO@BDMRP BVPQHCH@

0MJJDBQHMLP PQRCHDC K6O K@/O KK/O

5$ JDRBMP@OU QVND N@O@QVNDP +'$,#+.$( -$'#.$& )$*#)$.

5$ JDRBMP@OU PP >DPQDOGMJQ *($*#*-$) ,$-#-$( )$,#*$(

5$ ARJAHEDORK PP 3@SOD ).$-#,'$) -$'#.$) *$&#*$'

5$BORPQ$ S@O$ KHLRP PP /MGRP +-$+ -$* )$-

5$ E@SODH QVND *,$*#*,$. -$(#-$) *$%#*$'

5$ SDJRQHNDP HPM# N@O@QVNDP )-$)#,($' +$,#-$- )$)#)$-

5$ BORPQRJHLHEMOKD PP /ORBGDQ ,'$, -$& )$)

5$ ARJAMPRK /ODP$ *-#+- -$&#.$( )$)#*$)

24#%(%.'%$' +($*#+)$) .$+#&%$* *$,#+$)

4$ 0MLPHFJHM .+%'& +-$-#,*$( .$+#&%$- *$'#*$,

7SDO@JJ :@LFD )-$)#,*$( +$,#&%$- )$)#+$)

SUPPLEMENTARY TABLE I(Hebeloma leucosarx P. D. Orton)

Sporal and cystidial size across the collections presented in the study, plus the voucher

material of H. crustuliniforme var. minus Cooke em. Rea ss Bohus (BOHUS, 1972). The

data regarding Bohus’ and Bresadola’s material is not shown in full in the study.

(*&&#!$!%) '+ ) #! " $"$'*(! '$.#*,!+/ 8$ 1$ 7OQML!

;NMO@J @LC BVPQHCH@J PHWD @BOMPP QGD BMJJDBQHMLP NODPDLQDC HL QGD PQRCV" NJRP QGD SMRBGDO

K@QDOH@J ME #+.,-.'&)&%*+($ S@O$ (&)., 0MMID DK$ :D@ PP /MGRP /75=;" &.,'!$ <GD

C@Q@ ODF@OCHLF /MGRPX @LC /ODP@CMJ@XP K@QDOH@J HP LMQ PGMTL HL ERJJ HL QGD PQRCV$

;NMODP

0MJJDBQHMLP PQRCHDC K6O K?O K9O

5$ JDRBMP@OU QVND N@O@QVNDP &%$-#&&$. +$)#+$* &$+-#&$-(

5$ JDRBMP@OU PP >DPQDOGMJQ &%$,#&%$- +$&#+$( &$,%#&$,,

5$ ARJAHEDORK PP 3@SOD &%$'#&&$( +$(#+$- &$+%#&$++

5$ BORPQ$ S@O$ KHLRP PP /MGRP &&$( +$) &$,+

5$ E@SODH QVND &%$+#&&$' +$*#+$+ &$+(#&$+-

5$ SDJRQHNDP HPM# N@O@QVNDP &%$'#&&$' +$&#+$) &$+(#&$,*

5$ BORPQRJHLHEMOKD PP /ORBGDQ .$- +$& &$+%

5$ ARJAMPRK /ODP$ &%$&#&&$) +$'#+$- &$+'#&$,(

24#%(%.'%$' &%$,#&&$' +$)#+$+ &$+-#&$,%

4$ 0MLPHFJHM .+%'& &&#&&$( +$'#+$) &$,)#&$-'

7SDO@JJ O@LFD .$-#&&$. +$&#+$- &$+%#&$-(

6@FDLMBJ@S@QD BVPQHCH@

0MJJDBQHMLP PQRCHDC K6O K@/O KK/O KA/O

5$ JDRBMP@OU QVND N@O@QVNDP *&$+#*)$+ ,$&#,$- )$+#)$- ,$&#,$+

5$ JDRBMP@OU PP >DPQDOGMJQ )'$%#)+$* +$+#,$) )$'#)$+ +$-#,$%

5$ ARJAHEDORK PP 3@SOD )+$-#+)$. -$&#.$) )$)#)$- +$-#,$'

5$ P@BB$ S@O$ KHLRP PP /MGRP *($' ,$( )$( +$)

5$ E@SODH QVND ),$,#)-$+ ,$'#,$. )$*#)$. +$*#,$%

5$ SDJRQHNDP HPM# N@O@QVNDP (-$&#++$) +$&#-$- )$(#)$, +$*#,$(

5$ BORPQRJHLHEMOKD PP /ORBGDQ *-$& ,$& )$% ,$-

5$ ARJAMPRK /ODP$ )+$*#*.$) ,$*#-$& )$%#)$. +$)#,$(

24#%(%.'%$' *'$-#*)$( -$+#.$' *$' ,$,#,$.

4$ 0MLPHFJHM .+%'& *-$-#+* -$-#.$' )$%#*$& +$-#+$.

7SDO@JJ :@LFD (-$&#++$) +$&#.$) )$%#*$' +$)#,$.

0J@S@QMBVJHLCO@BDMRP BVPQHCH@

0MJJDBQHMLP PQRCHDC K6O K@/O KK/O

5$ JDRBMP@OU QVND N@O@QVNDP +'$,#+.$( -$'#.$& )$*#)$.

5$ JDRBMP@OU PP >DPQDOGMJQ *($*#*-$) ,$-#-$( )$,#*$(

5$ ARJAHEDORK PP 3@SOD ).$-#,'$) -$'#.$) *$&#*$'

5$BORPQ$ S@O$ KHLRP PP /MGRP +-$+ -$* )$-

5$ E@SODH QVND *,$*#*,$. -$(#-$) *$%#*$'

5$ SDJRQHNDP HPM# N@O@QVNDP )-$)#,($' +$,#-$- )$)#)$-

5$ BORPQRJHLHEMOKD PP /ORBGDQ ,'$, -$& )$)

5$ ARJAMPRK /ODP$ *-#+- -$&#.$( )$)#*$)

24#%(%.'%$' +($*#+)$) .$+#&%$* *$,#+$)

4$ 0MLPHFJHM .+%'& +-$-#,*$( .$+#&%$- *$'#*$,

7SDO@JJ :@LFD )-$)#,*$( +$,#&%$- )$)#+$)

(*&&#!$!%) '+ ) #! " $"$'*(! '$.#*,!+/ 8$ 1$ 7OQML!

;NMO@J @LC BVPQHCH@J PHWD @BOMPP QGD BMJJDBQHMLP NODPDLQDC HL QGD PQRCV" NJRP QGD SMRBGDO

K@QDOH@J ME #+.,-.'&)&%*+($ S@O$ (&)., 0MMID DK$ :D@ PP /MGRP /75=;" &.,'!$ <GD

C@Q@ ODF@OCHLF /MGRPX @LC /ODP@CMJ@XP K@QDOH@J HP LMQ PGMTL HL ERJJ HL QGD PQRCV$

;NMODP

0MJJDBQHMLP PQRCHDC K6O K?O K9O

5$ JDRBMP@OU QVND N@O@QVNDP &%$-#&&$. +$)#+$* &$+-#&$-(

5$ JDRBMP@OU PP >DPQDOGMJQ &%$,#&%$- +$&#+$( &$,%#&$,,

5$ ARJAHEDORK PP 3@SOD &%$'#&&$( +$(#+$- &$+%#&$++

5$ BORPQ$ S@O$ KHLRP PP /MGRP &&$( +$) &$,+

5$ E@SODH QVND &%$+#&&$' +$*#+$+ &$+(#&$+-

5$ SDJRQHNDP HPM# N@O@QVNDP &%$'#&&$' +$&#+$) &$+(#&$,*

5$ BORPQRJHLHEMOKD PP /ORBGDQ .$- +$& &$+%

5$ ARJAMPRK /ODP$ &%$&#&&$) +$'#+$- &$+'#&$,(

24#%(%.'%$' &%$,#&&$' +$)#+$+ &$+-#&$,%

4$ 0MLPHFJHM .+%'& &&#&&$( +$'#+$) &$,)#&$-'

7SDO@JJ O@LFD .$-#&&$. +$&#+$- &$+%#&$-(

6@FDLMBJ@S@QD BVPQHCH@

0MJJDBQHMLP PQRCHDC K6O K@/O KK/O KA/O

5$ JDRBMP@OU QVND N@O@QVNDP *&$+#*)$+ ,$&#,$- )$+#)$- ,$&#,$+

5$ JDRBMP@OU PP >DPQDOGMJQ )'$%#)+$* +$+#,$) )$'#)$+ +$-#,$%

5$ ARJAHEDORK PP 3@SOD )+$-#+)$. -$&#.$) )$)#)$- +$-#,$'

5$ P@BB$ S@O$ KHLRP PP /MGRP *($' ,$( )$( +$)

5$ E@SODH QVND ),$,#)-$+ ,$'#,$. )$*#)$. +$*#,$%

5$ SDJRQHNDP HPM# N@O@QVNDP (-$&#++$) +$&#-$- )$(#)$, +$*#,$(

5$ BORPQRJHLHEMOKD PP /ORBGDQ *-$& ,$& )$% ,$-

5$ ARJAMPRK /ODP$ )+$*#*.$) ,$*#-$& )$%#)$. +$)#,$(

24#%(%.'%$' *'$-#*)$( -$+#.$' *$' ,$,#,$.

4$ 0MLPHFJHM .+%'& *-$-#+* -$-#.$' )$%#*$& +$-#+$.

7SDO@JJ :@LFD (-$&#++$) +$&#.$) )$%#*$' +$)#,$.

0J@S@QMBVJHLCO@BDMRP BVPQHCH@

0MJJDBQHMLP PQRCHDC K6O K@/O KK/O

5$ JDRBMP@OU QVND N@O@QVNDP +'$,#+.$( -$'#.$& )$*#)$.

5$ JDRBMP@OU PP >DPQDOGMJQ *($*#*-$) ,$-#-$( )$,#*$(

5$ ARJAHEDORK PP 3@SOD ).$-#,'$) -$'#.$) *$&#*$'

5$BORPQ$ S@O$ KHLRP PP /MGRP +-$+ -$* )$-

5$ E@SODH QVND *,$*#*,$. -$(#-$) *$%#*$'

5$ SDJRQHNDP HPM# N@O@QVNDP )-$)#,($' +$,#-$- )$)#)$-

5$ BORPQRJHLHEMOKD PP /ORBGDQ ,'$, -$& )$)

5$ ARJAMPRK /ODP$ *-#+- -$&#.$( )$)#*$)

24#%(%.'%$' +($*#+)$) .$+#&%$* *$,#+$)

4$ 0MLPHFJHM .+%'& +-$-#,*$( .$+#&%$- *$'#*$,

7SDO@JJ :@LFD )-$)#,*$( +$,#&%$- )$)#+$)

176 Grilli E.

SUPPLEMENTARY TABLE II (Hebeloma lutense Romagn.)

Sporal and cystidial size across the collections presented in the study, plus additional

unpublished material. The data regarding LaChiusa 774 (MCVE), Lucchini 6084 (LUG)

and Contu 970912.2, 981013.1 (HG) is not shown in full in the study.

(*&&#!$!%) '+ ) #! "" "!"#&$ #)("%'" 9KI?EJ$!

:LKM?H ?JB ATNOGBG?H NGUC ?AMKNN OFC AKHHCAOGKJN LMCNCJOCB GJ OFC NOPBT" LHPN ?BBGOGKJ?H

PJLP@HGNFCB I?OCMG?H$ ;FC B?O? MCE?MBGJE 5?0FGPN? ,,) 60=1!" 5PAAFGJG +%-) 5<2!

?JB 0KJOP .,%.&'$'" .-&%&($& 32! GN JKO NFKRJ GJ DPHH GJ OFC NOPBT$

:LKMCN

0KHHCAOGKJN NOPBGCB I5M I>M I8M

3$ HPOCJNC OTLC &%$.*#&&$.( *$-)#+$%+ &$-,#&$.+

9KI?EJCNG +,$'&, &%$)'#&%$,& *$+-#*$,, &$-(#&$-*

3$ HCPAKN?MS VNLCAGICJ 'W &&$%+ +$%, &$-'

7MOKJ *-+ &%$.(#&&$*( +$%&#+$'& &$,*#&$.%

3$ HCPAKN?MS NN /MPAFCO &&$()#&&$,% *$-(#*$.+ &$--#'$%%

/KJ ,&%.&,%) &%$&,#&%$), *$(,#*$*+ &$-,#&$.'

5? 0FGPN? ,,% &&$&(#&&$), *$,.#+$%. &$-*#&$.'

5PAAFGJG +%-) &%$%.#&%$,% *$*(#*$*- &$-(#&$.)

0KJOP .,%.&'$' .-&%&($& &&$%*#&&$** *$,)#+$%( &$--#&$.-

4=.+#'(& &%$,, *$-% &$-*

7QCM?HH M?JEC &%$%.#&&$.( *$(,#+$'& &$,*#'$%%

0H?Q?OKLCBGACHH?OC ATNOGBG?

0KHHCAOGKJN NOPBGCB I5M I?/M II/M

3$ HPOCJNC OTLC )($'#*.$+ ,$%#&&$. ($%

3$ HPOCJNC +,$'&, +'$& &%$( ($'

3$ HCPAKN?MS VNLCA$ 'W *&$( ,$( ($(

7MOKJ *-+ )'$&#),$+ ,$*#-$& '$-#($'

3$ HCPAKN?MS NN /MPAFCO *)$(#*,$( ,$'#,$( ($%

/KJ ,&%.&,%) *&#**$* +$,#-$& ($&#($)

5?0FGPN? ,,% +($)#+*$( +$+#,$( '$*#($%

5PAAFGJG +%-) *'$-#*+$' ,$.#-$* ($%#($&

0KJOP .,%.&'$' .-&%&($& *&$,#*+$. +$,#&&$* ($%#($*

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Decrypting the Hebeloma crustuliniforme complex: European species of Hebeloma section Denudata subsect. Denudata (Agaricales)

Article

Full-text available

Jan 2016

Hebeloma subsection Denudata includes the type of H. section Denudata, Hebeloma crustuliniforme, as well as the majority of the taxa commonly included in the Hebeloma crustuliniforme complex. Complementing the work of D.K. Aanen and co-workers, and using refined morphological and molecular methods we were able to recognize further individual taxa within the section. Fifteen species occurring in Europe are assigned to H. subsect. Denudata. Of these, we describe eight species as new, namely H. aanenii, H. aurantioumbrinum, H. geminatum, H. louiseae, H. luteicystidiatum, H. pallidolabiatum, H. perexiguum and H. salicicola. Naucoria bellotiana, a species very similar to H. alpinum is recombined into Hebeloma. A key to Hebeloma subsect. Denudata is provided. We demonstrate that within this subsection there is good overall consistency between morphological, phylogenetic and biological species concepts. In contrast to current opinion, in this group there is little species overlap, particularly when also considering species frequencies, between arctic and alpine floras on one hand and temperate on the other.

Unexpected species diversity and contrasting evolutionary hypotheses in Hebeloma (Agaricales) sections Sinapizantia and Velutipes in Europe

Article

Full-text available

Dec 2016

Hebeloma velutipes is one of the most common and abundant members of the ectomycorrhizal basidiomycete genus and H. sinapizans is one of its oldest and most commonly recorded species. Using large sample sizes, several loci and the analysis of types, we explored the taxonomy, species limits, distribution and the infrageneric classification of these two species and their relatives. By relying almost exclusively on sequenced material, we were able to attain a marked refinement of species descriptions. Phylogenetic results are congruent with respect to the delimitation of species, but suggest conflicting evolutionary histories of the species phylogeny. Using multi-species coalescent analysis, phylogenetic support for H. sects. Velutipes and Sinapizantia was assessed, finding clear support for H. sect. Sinapizantia but ambiguous results for H. sect. Velutipes. One species, H. subconcolor, previously accommodated in H. sect. Denudata, is placed in H. sect. Velutipes. Hebeloma bulbiferum, so far not considered in systematic treatments, is shown to belong to H. sect. Sinapizantia. Unexpectedly, H. velutipes turned out to be distinct from H. leucosarx. Hebeloma erebium comb. nov., H. celatum sp. nov. and H. quercetorum, formerly treated as a single species (H. quercetorum), are demonstrated to be three taxa that are clearly distinct in molecular terms, even though, morphologically, they can be deceptively similar; H. erebium and H. quercetorum are, moreover, geographically distinct. The morphological characters used to distinguish the ten recognised European species are outlined. Finally, a lectotype and an epitype are designated for H. sinapizans and a lectotype for H. quercetorum.

Epitypification of Hebeloma crustuliniforme

Article

Aug 2013

It is widely recognised that there is no consistent use of the name H. crustuliniforme. The name has been used to represent a number of different taxa and indeed taxa from sections of Hebeloma that are morphologically and molecularly well separated. The holotype of H. crustuliniforme is plate 308 Bulliard from 1787 and while it can be interpreted, no such interpretation can be without ambiguity. It is clear from existing literature that modern authors have applied numerous different interpretations to this name and no real consensus exists. Indeed it appears that at various times most of the medium to large species within both sections Denudata and Velutipes Vesterh. have been referred to as H. crustuliniforme. Within this paper an epitype is selected for H. crustuliniforme in order to give the taxon a precise meaning and a detailed species description is given. Molecular data combined with the results of intercompatibility tests of Aanen and Kuyper published earlier support the definition of H. crustuliniforme adopted in this paper as a distinct taxon and as a biological species. We strongly recommend that this taxon be referred to as Hebeloma crustuliniforme (Bull.) Quél. emend. Vesterh., U. Eberh. & Beker in order to emphasise that it is the specific taxon rather than the complex to which it is being referred.

A comparison of the application of a biological and phenetic species concept in the Hebeloma crustuliniforme complex within a phylogenetic framework

Article

Full-text available

Jan 2004

A method is presented to derive an operational phenetic species concept for the Hebeloma crustuliniforme complex in northwestern Europe. The complex was found to consist of at least 22 biological species (intercompatibility groups; ICGs). Almost none of these biological species could be recognised unambiguously by morphological criteria. It is therefore necessary to base a phenetic species concept on combinations of biological species. However, such species delimitation must be performed within an explicitly phylogenetic context. It is crucial therefore to have a reliable estimate of the phylogeny of 22 biological species in that complex. Based on two nuclear sequences, we present a best estimate of the phylogeny of biological species within the complex. Using this phylogeny, on the basis of strict monophyly only two species can be morphologically recognised among 22 biological species. Relaxing the criterion of monophyly and allowing paraphyletic groupings of biological species as phenetic species would result in the recognition of three phenetic species. A tree, with the five ICGs of the previously defined morphospecies H. crustuliniforme (1, 2, 3, 4 and 5) constrained as a monophyletic group, can not be rejected. This constrained tree, together with the relaxed criterioon that allows for paraphyletic groupings of biological species, leads to the recognition of four phenetic species, viz. H. crustuliniforme, H. helodes, H. incarnatulum and H. velutipes. These phenetic species are described and a key is provided. Other taxon names are briefly discussed. The very limited ability to translate a biological species concept into an operational phenetic species concept is explained by the lack of qualitative characters and the plasticity of quantitative characters. Recency of common evolutionary history is also a major factor. Intercompatibility tests and DNA based phylogenies indicate that most biological species are very closely related and hence provide support for the claim that correspondence between a biological species concept and a phenetic species concept in the H. crustuliniforme complex is not likely to be forthcoming. In an Appendix morphological descriptions are provided of the 22 ICGs.

Intercompatibility Tests in the Hebeloma crustuliniforme Complex in Northwestern Europe

Article

Full-text available

Sep 1999

Sexual intercompatibility has been tested for members of the ectomycorrhizal Hebeloma crustuliniforme complex. Intercompatibility tests have been applied to 110 collections from northwestern Europe, for which monospore cultures were obtained. Crossing experiments indicated the existence of at least 20 intercompatibility groups (ICGs). All ICGs were bifactorial heterothallic with multiple mating type alleles. A low level of partial intercompatibility between ICGs was observed in two species clusters. One isolate was completely compatible with members of two ICGs (and partly compatible with one member of a third ICG) and could therefore not be assigned to an ICG. A large majority of these 20 ICGs could not be recognized by morphological characters, supporting claims that many basidiomycete morphospecies contain cryptic biological species. Possible explanations for this large number of ICGs that are not morphologically, ecologically or geographically delineated are discussed. A simplified strategy for executing intercompatibility tests with a high number of isolates is described. This procedure could substantially reduce the number of necessary matings.

Ainsworth & Bisby's Dictionary of the Fungi

Article

Full-text available

Jan 2001

BASIDIOMICETS DE LES TORBERES DELS PIRINEUS CATALANS. PRIMERA CONTRIBUCIÓ

Article

Full-text available

May 1996

Basidiomycetes from the peat-bogs of the Pyrenees of Catatonia. First contribution. First floristic results of the studies deroted to the Basidiomycetes that grows in the sphagni bogs of Catalan slopes of the Pyrenean range. In all, 9 species, new or rarely recorded in the Iberian Peninsule are recorded and discussed: Hygrocybe coccineocrenata (Ort.) Mos., H. lepida Arnolds, Hypholoma elongatum (Pers.: Fr.) Ricken, H. udum (Pers.: Fr.) Kühner, Omphalina oniscus (Fr.: Fr.) Quél., Hebeloma helodes Favre, Cortinarius palustris Moser, C. sphagneti P.D. Orton i C. uliginosus Berk.

Phylogenetic relationships in the genus Hebeloma based on ITS1 and 2 sequences, with special emphasis on the Hebeloma crustuliniforme complex

Article

Mar 2000

Phylogenetic relationships within the genus Hebeloma (Cortinariaceae, Agaricales) were determined, based on nuclear ribosomal ITS sequences, using cladistic methods. Special emphasis was on phylogenetic relationships within the H. crustuliniforme complex. In total 52 sequences were analysed, representing 51 collections and 39 taxa. Agrocybe praecox and two species of Alnicola were used as out-groups. The genus Hebeloma appears to be monophyletic. Several well supported clades could be recognized. However, many of the basal relationships are unresolved or only weakly supported. Alternative topologies could not be rejected. It is therefore impossible to derive a revised infrageneric classification of Hebeloma. The H. crustuliniforme complex appears paraphyletic, consisting of two clades with three and 17 intercompatibility groups respectively. In the second clade many of the phylogenetic relationships are also unresolved, reflecting a high rate of recent speciation events. Most of the species in this clade form ectomycorrhizae with members of the Salicaceae. The taxon that is basal to this clade, however, is not associated with these hosts. The host tree switch to Salicaceae has been followed by extensive and rapid speciation.

Contribution a l'etude du genre Hebeloma (Fr.) Rummer; partie speciale

Article

G. Bruchet

Reinventing taxonomy: A curmudgeon's view of 250 years of fungal taxonomy, the crisis in biodiversity, and the pitfalls of the phylogenetic age

Article

Jul 2005

R.P. Korf

Biological taxonomy is bleeding. The fungi are but one example. There is an almost ignored crisis of impending loss in biodiversity, while the funding - necessary to provide basic inventories and monographic studies - is simultaneously siphoned away by the misuse of the great potentials of molecular biology. One disturbing result has been frequent support of needless, repetitive phylogenetic studies. A seminal paper, "Taxonomic triage and the poverty of phylogeny," by entomologist Quentin D. Wheeler, is cited as required reading for all biologists. His theses are that "Taxonomy, already weakened by decades of neglect, now suffers the loss of positions and funding," and that "Considering what is at stake for human and environmental welfare in the biodiversity crisis, it is time to triage and move descriptive taxonomy to the forefront of science funding priorities." Reinventing taxonomy may provide answers.

New check list of British Agarics and Boleti: Part III. Notes on genera and species in the list

Article