A nomenclatural revision of the genus Alnicola - Fungal diversity

Fungal Diversity 

A nomenclatural revision of the genus Alnicola (Cortinariaceae) 

Pierre-Arthur Moreau * 

Herbarium Z+ZT. Geobotanisches Institut ETH. Zollikerstrasse 107. CH-8008 Zürich. Current 

address : Département de Botanique. Faculté des Sciences Pharmaceutiques et Biologiques. 

B.P. 83. F-59006 Lille Cedex. France 

Moreau, P.-A. (2005). A nomenclatural revision of the genus Alnicola (Cortinariaceae). 

Fungal Diversity 20: 121-155. 

Some nomenclatural points are discussed as a prodrom of a taxonomic revision of the genus 

Alnicola (= Naucoria sensu Moser). Lectotypifications of Alnicola and Agaricus "tribe" 

Naucoria are critically analyzed. Alnicola luteolofibrillosa is proposed as a new lectotype of 

Alnicola; Ag. pannosus is proposed as a new lectotype for Ag. "tribe" Naucoria. A critical 

check-list of taxa published so far in the genus Alnicola is provided. The usual names Alnicola 

(Naucoria) alnetorum and Alnicola (Naucoria) langei are invalid and must be replaced by A. 

inculta and A. macrospora respectively. Alnicola (Naucoria) dasypus is an earlier synonym of 

Pholiotina (Conocybe) subnuda according to type revision. Lectotypes are designed for 

Agaricus melinoides, A. scolecinus, Alnicola macrospora, Naucoria amarescens and Naucoria 

dasypus. Four new combinations are introduced: Alnicola zonata, Galerina albotomentosa, G. 

clavuligera, Pholiotina dasypus. 

Key words: Basidiomycota, Galerina, Naucoria, Pholiotina, taxonomy, typification. 

Introduction 

The genus Alnicola has been introduced by Kühner (1926), in order to 

isolate a homogenous group of species from the nebulous Agaricus "tribe" 

Naucoria of Fries (1821, 1836-1838). All are characterized by warty spores, 

sterile gill edge, differentiated pileipellis and strict association with Alnus spp. 

(alder trees). It was initially made up of 4 species briefly mentioned in a short 

key by Kühner (loc. cit.), Alnicola includes presently 60 specific taxa, all 

mycorrhizal and mainly associated with Betulaceae and Salicaceae, most of 

them originally described from Europe. 

Up to now this genus has never been misinterpreted, but a significant 

number of modern authors (Moser, 1955; Orton, 1960; Reid, 1984; Døssing, 

1992; etc.) prefer to use the name Naucoria (Fr.: Fr.) P. Kumm. It was created 

by Fries (1821: 260) as a tribe of Agaricus, in order to group together 

collybioid, ochre-spored gill fungi which could not be classified in the closely 

* Corresponding author: P.A. Moreau; e-mail: pierre-arthur.moreau@univ-lille2.fr 

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elated tribes Inocybe and Galera. ‘Trib. Naucoria’ (Fries, 1821), which 

initially included 7 species (according to current systematic: 1 Agrocybe, 1 

Alnicola, 2 Flammulaster, 1 Tubaria, 2 not interpreted), is so heterogenous that 

the first modern systematicians (Fayod, 1889; Scherffel, 1897; Earle, 1909; 

Kühner, 1926; Heim, 1931) split it into a number of smaller genera. Only the 

residual species remaining in Naucoria could justify the use of this name, 

which each author used to do according to his own conception. Amongst them, 

Moser (1955) was the first restricting Naucoria to a strict synonym of Alnicola; 

he was followed in this way by Reid (1984), Døssing (1992) and Ludwig 

(2000, 2001), whereas others like Kühner and Romagnesi (1953, 1957), 

Locquin (1956), Orton (1960) or Malençon and Bertault (1970) used Naucoria 

as a "default" genus grouping all brown-spored species of undefined or 

doubtful systematic affinities. 

The typification of Alnicola and Naucoria, already discussed by Donk 

(1962), Singer (1975, 1986), Kühner (1980, 1987) and Moser (1985-2003), is a 

critical point in this debate. Because of the fact that different interpretations of 

Fries’ texts (all more or less equally acceptable according to the value 

attributed to various arguments) lead to opposite conclusions, at this stage a 

somewhat arbitrary choice seems unavoidable in order to solve the recent but 

recurrent dilemma between Alnicola and Naucoria for current use. 

Lectotypifications are clearly proposed here in order to adopt a stable 

nomenclature for these genera, based on an exegesis of Fries’ texts and on 

Moser’s and Kühner’s arguments, but also taking into account traditions, 

usages and comparative disturbances of all alternatives. Finally the name 

Alnicola is adopted here, along the lines of a mainly West-European tradition 

formalized by Kühner (1987) and discussed, according to the arguments 

developed below. 

Even if the generic definition of Alnicola does not constitute a problem, 

the nomenclatural status of a number of taxa at specific or infraspecific rank 

needs to be revised, independently of their taxonomical treatment. The aim of 

this paper is to prepare a taxonomic revision of the genus Alnicola by stating 

on some nomenclatural points, according to the International Code of Botanical 

Nomenclature, Saint-Louis (Greuter et al., 2000). It is assumed that the 

classification of several mentioned taxa in Alnicola is only provisionally 

accepted, and is likely to be modified according to molecular data in a near 

future. 

Typification of Alnicola 

Alnicola Kühner, Botaniste 17 (1-4): 175 (1926). 

≡ Naucoria subg. Alnicola (Kühner) R. Heim, genre Inocybe: 74 (1931). 

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≡ Naucoria sect. Alnicola (‘Alnicolae’) (Kühner) J.E. Lange, Dansk bot. Ark. 9 (6): 21 

(1938). 

≡ Hebeloma subg. Alnicola (Kühner) Kühner, Hyménom. agaricoïdes: 228 (1980). 

= Naucoria ss-g. Cytocutis J.E. Lange, Dansk bot. Ark. 9 (6): 21 (1938) (inval., art. 

36.1). 

?= Cystocybe Velen., České Houby I: 495 (1921) (fide Singer, 1942: 342, 1950a: 412) 

Original diagnosis: ‘Petites espèces grêles, à spores ventrues, en amande ou en fuseau, 

à paroi jaune épaissie et grenelée ponctuée avec le sommet atténué et à membrane souvent 

amincie mais sans pore germinatif.’ (Kühner, 1926: 175). 

Original species: Alnicola submelinoides Kühner, A. luteolofibrillosa Kühner, A. badia 

Kühner, A. melinoides (Bull. : Fr.) Kühner. 

Typifications 

Konrad (1934: 176, as a subgenus of Naucoria): Agaricus escharoides 

Fr. : Fr., an incorrect neotype, out of the original names of species (Art. 10.2) 

(a synonym of A. melinoides Bull. : Fr. ss. Kühner for Konrad, see taxonomic 

discussion below). 

Singer (1936: 342): Alnicola submelinoides Kühner, followed by Singer 

(1949: 538) and Kühner (1980: 228), acceptable. 

Horak (1968: 73): Agaricus melinoides Bull. : Fr. ss. Kühner, already 

suggested by Donk (1962: 19), followed by Singer (1975: 582; 1986: 612) and 

Bon (1992: 8), acceptable. 

Discussion 

Kühner (1926) originally described 4 species in his new genus Alnicola: 

3 were introduced as new, the fourth being A. melinoides, identified following 

Ricken’s flora (1915). Later, Kühner (1931) provided extensive descriptions of 

his taxa, but he finally abandoned the name A. melinoides after 1926, even in 

his personal notes. Kühner never published any description of "his" 

A. melinoides, but one collection (V-85, la Dent-du-Villard, herb. R. Kühner, 

G) is probably (date not indicated) the one used for his key of 1926. This 

collection from Alnus viridis, which is at present difficult to interpret, is not 

likely identical to A. escharoides ss. Lange. 

Before 1926, Kühner collected all his Alnicola species under Alnus 

viridis in the subalpine zone of the Alps (La Dent du Villard, Savoie, F; 

R. Kühner's unpublished notes, G), an habitat where most taxa differ 

consistently from the lowland species described e.g. by Konrad and Maublanc 

(1926), Lange (1938), Romagnesi (1942) or Orton (1960) (Moreau, 

unpublished). Kühner’s unpublished notes (G) prove that the synonymies this 

author admitted to later between lowland and subalpine taxa (e.g. Kühner, 

1942; in Kühner and Romagnesi, 1953) were strongly influenced by 

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M. Josserand and R. Maire (unpublished correspondence, herb. Kühner, G, and 

private coll. D. Lamoure) and by Lange (1938). None of these last authors was 

experienced in orophilic species which the genus Alnicola is originally based 

on. In fact, the synonymies admitted by Kühner (1942, 1980, 1987) were only 

based on the interpretations of his senior correspondents, respectfully trusted, 

but never on comparative studies or critical taxonomic discussion. This has 

important consequences on several usually admitted synonymies, and 

especially on 2 already proposed lectotypes of Alnicola: Ag. escharoides and 

Ag. melinoides. 

Konrad’s (1934) earlier (neo)typification (Ag. escharoides) was justified 

by the synonymy with A. melinoides explicited in Konrad and Maublanc (1926, 

pl. 175 II). But this synonymy is based on a personal interpretation of Kühner’s 

own interpretation of Ag. melinoides (see below, list of species names). A. 

escharoides ss. Konrad (= A. melinoides ss. Ricken) is a common European 

species usually associated with Alnus glutinosa but not confirmed under A. 

viridis, and not susceptible to have been included in Kühner’s original list of 

taxa for his genus Alnicola (contradiction with recomm. 9A3). However, as 

indicated above, Konrad’s neotype must be superseded because of the 

existence of original species. 

Donk (1962) also discussed Ag. melinoides as a possible lectotype for 

Alnicola, but was aware of the taxonomic and nomenclatural problems 

associated with this selection. Consequently he did not recommend it formally 

in order to allow more documented alternatives. This choice has been 

formalized by Horak (1968), who gave as a reference some personal 

collections of "Alnicola melinoides" from Alnus incana (preserved in ZT), 

which undoubtedly belong to Alnicola but which cannot be considered as 

identical with Bulliard’s taxon (most likely a Galerina). 

A. submelinoides, proposed as a lectotype by Singer (1936), is the first 

species mentioned by Kühner (1926), and was the only species found by him to 

have clavate cystidia and hymeniform pileipellis. It cannot be proved that for 

his designation, Singer (1936) used, the "first species method" in an arguable 

way (this method was not constantly used in Singer's work); here, as the first 

species keyed out by Kühner (loc. cit.) was logically the less typical one, this 

method would have been clearly unadequate. Kühner (1980) himself 

recommended A. submelinoides as a lectotype for Alnicola, but for another 

reason: he esteemed Alnicola and Hebeloma to be synonyms and, speculating 

on their phylogenetic relations, stated that A. submelinoides was the most 

distant taxon from Hebeloma because of its hymeniform pileipellis. 

Kühner’s phylogenetic hypothesis is not confirmed by recent molecular 

studies (Moreau et al., 2005), which tend to prove that A. submelinoides and A. 

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inculta form a distinct group not directly related to any other Alnicola and 

closer to (but distinct from) Hebeloma. Because it appears necesary to 

segregate both taxa in an independent genus in a near future (Moreau and 

Peintner, unpublished), in order to keep Alnicola as a monophyletic genus in 

the broadest traditional sense (in order to limit the number of new 

combinations), it is proposed here to reject A. submelinoides as a lectotype for 

Alnicola ss. str. 

Consequently, two specific taxa considered by Kühner (1926, 1931) as 

typical of his genus Alnicola (Moreau and Peintner, unpublished) can be 

designated as lectotypes: A. luteolofibrillosa and A. badia. The first is also the 

best documented in literature and is not an ambiguous taxon. If this lectotype 

came to be contested, it would be necessary to propose the conservation of 

Alnicola with a typus conservandus, which could be A. luteolofibrillosa. At 

present the situation of Alnicola does not seem to justify such a procedure. 

Lectotype proposed here: Alnicola luteolofibrillosa Kühner 1926, Botaniste 

17:175. 

Typification of Agaricus "tribe" Naucoria 

Agaricus "tribe" Naucoria Fr., Syst. mycol. 1: 261 (1821): Fr. 

≡ Ag. subgen. Naucoria (Fr. : Fr.) Loud., Encycl. pl.: 1002 (1829). 

≡ Naucoria (Fr. : Fr.) P. Kumm., Führer Pilzk.: 22, 76 (1871). 

Original diagnosis: ”Char.: Velum cum integumento (epidermide) pilei squamuloso 

homogenum, fugacissimum. Stipes subfistulosus, aequalis, tenuis, squamulosus, intus 

flocculosus. Pileus carnoso-membranaceus, e campanulato-planiusculus, squamulis l. fibrillis 

innatis adpressis, raro secedentibus, tectus. Lamellae cinnamomeae. Obs.: Fungi minores, 

gregarii, epiphyti (in stipulis, lignis, foliis, humo limoso etc.), persistentes, fragiles, inodori. 

Statura quidem Collybiae; sed velum ejusdem inodolis ac Lepiotarum, naucum (unde nomen) 

referens.- Spec. 1-3 pileo subcarnoso obtuso, lamellis adfixis secedentibus; 4-6 pileo 

submembranaceo minute umbonato, lamellis adnexis; 7-8 pileo subcarnoso demum umbilicato, 

lamellis subdecurrentibus & c. dignoscuntur.- Accedunt aliae species pileo glabro.” (Fries, 

1821: 260). 

Original species: Ag. escharoides Fr. (currently: Alnicola); Ag. conspersus Fr. 

(currently: Tubaria); Ag. siparius Fr. (currently: Flammulaster); Ag. pannosus Fr. (not 

interpreted); Ag. graminicola Nees [= Crinipellis scabella (Alb. & Schwein. : Fr.) Murrill, after 

Gillot and Lucand, 1891: 238]; Ag. furfuraceus Fr. (currently: lectotype of Tubaria); Ag. 

segestrius Fr. (currently: Tubaria). 

Typifications 

Earle (1909): Ag. melinoides Bull. : Fr. Not acceptable, mentioned by 

Fries (1821) out of the ‘Genuini’ group of his Naucoria "tribus", and 

transferred in his XXIX Galera "tribus" in Elenchus (1828). 

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Singer (1936, 1949), Singer and Smith (1946), Horak (1968): Ag. 

centunculus Fr. : Fr. (Simocybe). Not acceptable, mentioned by Fries (1821: 

262) in the unnamed sect. ††, not in the ‘Genuini’, and in tribe Galera later 

(1828: 35). 

Romagnesi (1950: 78, 1979: 18): Ag. carpophilus Fr. (Flammulaster). 

Not acceptable, not mentioned by Fries (1821). 

Donk (1962: 196), Moser and Jülich (1985-2003): proposal for 

Ag. escharoides Fr.: Fr. (acceptable; now Alnicola in the sense of Konrad and 

Maublanc, 1926; Lange, 1938). 

Kühner (1980: 322, 898, 932), Romagnesi (1950): Ag. siparius Fr.: Fr. 

(acceptable; Flammulaster in the sense of Orton, 1960). 

Discussion 

Fries (1821: 261) created the small "tribus" XXVII Naucoria for 14 taxa, 

considered as the "correspondent" of Collybia in brown-spored groups of 

Agaricus. 7 species constitute the group "Genuini" (considered by Fries as 

typical) and 7 other taxa mentioned in an unnamed section, transient towards 

"tribe" XXVIII Galera (transferred in Galera later, Fries, 1828: 33). Many 

authors have criticized the heterogeneity of the tribe Naucoria according to 

modern knowledge on their anatomy (Kühner, 1926, 1980; Donk, 1962; 

Singer, 1962, 1975, 1986); for this reason all susequent authors have proposed 

to restrict Naucoria to a small part of its initial components (see table 1). This 

point has been extensively discussed by Kühner (1980, 1987), who pointed out 

the necessity of choosing an adequate lectotype of Naucoria in order to 

stabilize its taxonomic situation. 

The lectotype must be chosen amongst the 7 taxa initially mentioned by 

Fries (1821) in his "Genuini" group. It must be noticed that Fries obviously 

classified his species by decreasing importance of veil, the most veiled taxon 

being Ag. escharoides. In his following works, Fries (1836-1838: 201) emends 

Ag. escharoides and seems to transfer the most veiled elements in the 

following taxon, Ag. conspersus. This modified interpretation of Ag. 

escharoides matches the diagnosis of Naucoria and more closely corresponds 

to Konrad and Maublanc (1926), Lange (1939) and nowadays interpretation of 

this taxon. 

Kühner (1942, 1980, 1987) and Singer (1986: 612) refused the 

typification of Naucoria by Ag. escharoides as a doubtful taxon. But Moser 

(1985-2003) defends Ag. escharoides as an interpretable taxon and reliable 

lectotype for Naucoria, according to his own visits in Fries’ localities (Femsjö, 

Sweden); one of his collections from Femsjö (74/264, IB) has been labelled as 

"Naucoria escharoides" by himself. This collection 74/264, studied by the 

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Table 1. Interpretations of Naucoria in literature. 


Authors Modern genera included in Naucoria Name used 

for Alnicola 

Fries (1874) Agrocybe pp., Alnicola, Flammulaster, Galerina pp., Naucoria 

Konrad (1934) Macrocystidia, Phaeocollybia, Phaeomarasmius, 

Lange (1938) 

Pilát (1951) 

Phaeonematoloma, Simocybe 

Heim (1931) Alnicola, Flammulaster, Phaeomarasmius, Simocybe Naucoria 

Kühner and 

Romagnesi (1953, 

1957) 

Locquin (1956) 

Malençon and 

Bertault (1970) 

Singer (1936, 1946, 

1949, 1950a, etc.) 

Horak (1968) 

Romagnesi (1942) 

Favre (1948) 

Alnicola, Flammulaster, Phaeomarasmius, Simocybe, 

Tubaria 

Alnicola, Descolea, Flammulaster, Phaeomarasmius, 

Simocybe, Tubaria 

Naucoria 

Naucoria 

Simocybe Alnicola 

Flammulaster, Phaeomarasmius, Simocybe Alnicola 

Romagnesi (1950) Flammulaster, Phaeomarasmius Alnicola 

Orton (1960) Alnicola, Galerina pp., Phaeogalera, Simocybe Naucoria 

Romagnesi (1977) Flammulaster pp., Tubaria Alnicola 

Kühner (1980) Flammulaster, Phaeogalera, Phaeomarasmius, Tubaria Hebeloma 

Pegler and Young 

(1968) 

Reid (1984) 

Moser (1955) 

Døssing (1992) 

Alnicola Naucoria 

Singer (1962, 1975, 

1986) 

Kühner (1987) 

Rejected Alnicola 

author, seems somewhat different from continental collections usually reported 

as A. escharoides (it gets closer to A. diplocystis Sing.); it is not a 

significatively veiled collection. 

Moser’s interpretation is certainly the interpretation which comes closest 

to the one put forward by Fries (1828) redefinition of Ag. escharoides; but 

originally (1821), Ag. escharoides is a veiled taxon probably based mainly on 

A. luteolofibrillosa (as Moser (1985-2003) and before him Kühner (1942) 

pointed out), and probably partly on collections of Tubaria (Singer, 1986). In 

order to select a lectotype, only the Systema Mycologicum (Fries, 1821) should 

be considered as an original document, and Ag. escharoides should then be 

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taken in his original sense, where it is not the most representative member of 

Naucoria but the first cited. Nevertheless its designation as lectotype cannot be 

considered as an automatic choice and cannot be rejected for this only reason. 

If considered an Alnicola, Ag. escharoides would be the only Alnicola 

species included in Naucoria Genuini by Fries all over his opera, all the others 

belonging to other "tribes". The choice of this lectotype for Naucoria would 

drive to a complete amending of Naucoria according to Fries’ (1821) concept, 

which excludes all initial taxa but one, and moreover not the most 

representative one. Finally, Ag. escharoides has already been proposed as a 

neotype of Alnicola (Konrad, 1934). For all these reasons the author proposes 

to reject Ag. escharoides as a lectotype for Naucoria. 

It should be mentioned that 3 other collections from Femsjö were 

collected by M. Moser, labelled as "Naucoria sp.", and another identified 

"Naucoria sphagneti P.D. Orton" (herb. Moser, IB). This illustrates the 

diversity of Swedish Alnicola, not perceptible in Fries’ opera who probably 

could not separate them reliably on macroscopical characters only. This is also 

illustrated by a plate of Ag. escharoides directed by Fries (S) in 1854, which 

might represent Inocybe calospora Quél. 

Agaricus conspersus Fr.: Fr. is eligible as lectotype of Naucoria, but has 

never been proposed. Unless Fries has modified his acceptance of this taxon 

after 1821 (Fries, 1836-1838: 210; see above, Ag. escharoides), it is now 

considered by all authors as a member of the genus Tubaria (W.G. Sm.) Gillet. 

As Tubaria has been the first genus separated from Naucoria (Gillet, 1876: 

540) and contains presently more than 60 taxa worldwide, a lectotypification of 

Naucoria by Ag. conspersus would uselessly force to introduce a considerable 

number of new combinations, against the recommendations of the Code of 

Nomenclature regarding nomenclatural stability. This possibility is rejected 

here. 

Agaricus siparius Fr.: Fr. has been proposed by Romagnesi (1950), on 

the basis of his own interpretation of this taxon (Romagnesi, 1942; in Kühner 

and Romagnesi, 1957). According to Romagnesi (loc. cit.), N. siparia ss. 

Romagn. might be a synonym of N. ferruginea R. Maire (now Tubaria, 

Phaeomarasmius or Flammulaster: Watling, 1967; Horak and Moreau, 2005). 

According to Orton (1960) and Vellinga (1994), it is an autonomous species of 

Flammulaster. Agaricus siparius is the closest species matching Fries’ 

diagnosis of Naucoria (1821: 261) and would certainly be the best candidate 

for an orthodox lectotypification of Naucoria. But considering the still 

uncertain interpretation and systematic position of Ag. siparius, and the 

nomenclatural consequences of such an uncertain choice, it is proposed not to 

adopt it. 

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Ag. pannosus Fr. : Fr. has not been interpreted in the recent literature, but 

fits in well with the original diagnosis of Naucoria; the only restriction is that 

the cap is described “squamis destituto” and therefore classified in the 4 th 

group of Naucoria sect. Lepidoti, when Ag. escharoides, Ag. siparia and Ag. 

conspersus are classified in the 3 rd section (Fries, 1836-1838, 1874). However, 

this is not contradictory with the protolog and it has always been considered by 

Fries, from 1821 to 1874, as a typical member of Naucoria (sect. Lepidoti). 

Therefore it figures as a perfect residue according to the recomm. 9A5. It is not 

risky to propose Ag. pannosus as a lectotype of Naucoria, since it should not 

destabilize any of the well-defined and currently accepted genera Alnicola, 

Flammulaster or Tubaria. 

Lectotype proposed here: Agaricus pannosus Fr. 1821, Syst. 

mycol. I: 261, : Fr. 

Nomenclature of supraspecific taxa 

Amarescens (stirps) 

Status: invalid (art. 36.1). Original species: Alnicola amarescens (Quél.) R. Heim & 

Romagn., A. chamiteae Kühner, A. tantilla (J. Favre) Romagn., A. cholea Kühner. 

Original name: Alnicola (subgen. Alnicola) stirpe Amarescens Kühner, Trav. sci. Parc 

natl Vanoise 11: 114 (1981) (inval.). 

Melinoideae (sect.) 

Status: valid, not typified. Original species: Alnicola luteolofibrillosa Kühner, 

A. umbrina (Maire) Kühner, A. suavis (Bres.) Kühner, A. melinoides (Bull. : Fr.) Kühner. 

Basionym: Alnicola sect. Melinoideae Singer, Rev. Mycol. (Paris) 4 (1-2): 68 (1939). 

Accepted name: Alnicola sect. Alnicola (art. 22.1) (if A. luteolofibrillosa 

accepted as lectotype of Alnicola). 

Salicicolae (sect.) 

Status: invalid (art. 36.1). Original species: Naucoria scorpioides (Fr.) J.E. Lange ss. 

Lange (= A. bohemica Velen.), N. macrospora J.E. Lange. 

Original name: Naucoria (subgenus Cytocutis) sect. Salicicolae J.E. Lange, Dansk Bot. 

Ark. 6 (9): 21(1938); Fl. Agar. Danica 3: 22 (1939). 

= Alnicola subgen. Salicicolae ("Salicicolae") (J.E. Lange) Kühner, Trav. sci. Parc natl 

Vanoise 11: 129 (1981) (inval.). 

Submelinoideae (sect.) 

Status: valid. Holotype: not designed. Original species: Alnicola submelinoides Kühner, 

A. bohemica (Velen.) Kühner, A. lignicola Singer, A. fulgens (J. Favre & R. Maire) Singer. 

Basionym: Alnicola sect. Submelinoideae Singer, Rev. Mycol. (Paris) 4(1-2): 68 (1939). 

Accepted name: Alnicola sect. Submelinoideae Singer. 

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Nomenclature of specific and infraspecific taxa 

albotomentosa 

Status: valid. Holotype (visited): UK, Oversley Wood, Warwicks, 11-II-1973, leg. 

A.W.Brand (K). Substrate: soil and bark ships, rotten wood. 

Basionym: Naucoria albotomentosa D.A. Reid, Trans. Brit. mycol. Soc. 82 (2): 195 

(1984). 

≡ Alnicola albotomentosa (D.A. Reid) Courtec., Doc. mycol. 16 (61): 48 (1985). 

Notes: This taxon is based on two abundant collections (K) but poorly 

documented concerning its macroscopy. According to the study of these 

collections, E. Horak and the author are convinced that this species belongs in 

the genus Galerina, close to G. clavuligera which also owns cystidioid clavate 

hairs on pileipellis and weakly punctuate, fusiform and undextrinoid spores. It 

cannot be considered as related to Alnicola. 

Accepted name: Galerina albotomentosa (D.A. Reid) E. Horak & P.-A. 

Moreau, comb. nov. 

alnetorum 

Status: invalid (art. 34.1b). Authentic material in herb. R. Kühner (G), M. Josserand (G) 

and R. Maire (MPU) + pl. X fig. 7-16 (Maire 1930). Host: Alnus glutinosa. 

Original names (alternative names): Alnicola alnetorum R. Maire ex Kühner, Bull. Soc. 

mycol. France 47: 243 (1931) (provisional name for A. submelinoides ss. R. Maire, Bull. Soc. 

mycol. France 47: 223 (1930), inval.); Naucoria submelinoides var. alnetorum R. Maire ex 

Kühner, Bull. Soc. mycol. France 47: 243 (1931), inval. (provisional name for 

A. submelinoides ss. R. Maire, Bull. Soc. mycol. France 47: 223 (1930), inval.). 

= Alnicola alnetorum (R. Maire) Romagn., Bull. Soc. mycol. France 58: 122 (1942) 

(inval.). 

= Naucoria alnetorum (R. Maire) Kühner & Romagn., Fl. anal. Champ. sup.: 238 

(1953) (inval.). 

= Alnicola submelinoides var. alnetorum (R. Maire) Konrad & Maubl., Agaricales: 164- 

165 (1948) (inval.). 

Notes: although this name has been used widely in European literature, it 

has never been validly published. The earliest and valid name for the species 

appears to be Alnicola inculta (Singer, 1955: 406). 

Accepted name: Alnicola inculta (Peck) Singer. 

amarescens 

Status: valid, not typified. Automatic lectotype (designed here): Quélet, Champ. Jura 

Vosges I pl. VII fig. 3 (1872) (as "Galera ravida"). Host: not precised (on charcoal). 

Basionym: Naucoria amarescens Quél., Assoc. Fr. Av. Sci. 1882: 8 (1883), based on 

Galera ravida Fr. ss. Quél., Champ. Jura Vosges I: 103 (1872). 

≡ Alnicola amarescens (Quél.) R. Heim & Romagn., Bull. Soc. mycol. France 50 (3): 

179 (1934). 

≡ Hylophila amarescens (Quél.) Quél., Enchir. Fung.: 101 (1886). 

Notes: a well-documented taxon thanks to several extensive descriptions 

130


and good iconography (Quélet, 1872; Boudier, 1905 pl. 127; Heim and 

Romagnesi, 1932; Favre, 1948: 130; Krieglsteiner, 1980; Kühner, 1981), but 

possibly collective according to perceptible discrepancies between some of 

these descriptions, especially on spore size and smell. 

Accepted name: Alnicola amarescens (Quél.) R. Heim & Romagn. 

badia 

Status: valid, not typified. Original material from the type locality (Praz Joseph, Savoie, 

F) in G. Host: Alnus viridis. 

Basionym: Alnicola badia Kühner, Bull. Soc. mycol. France 47 (3-4): 239 (1931). 

≡ Naucoria phaea Kühner � R. Maire & Kühner in Maire, Bull. Inst. bot. Barcelona 3 

(4): 101 (1937) (nomen novum, not Naucoria badia Murrill). 

≡ Alnicola phaea (Kühner � R. Maire & Kühner) Romagn., Bull. Soc. mycol. France 

58: 126 (1942) (comb. superfl.). 

Notes: a good species described from Alnus viridis (well illustrated by 

Breitenbach and Kränzlin, 2000, n° 136 as "A. sphagneti", and n° 138 as 

"A. subconspersa"), but still not very well circumscribed and possibly 

collective. 

Accepted name: Alnicola badia Kühner. 

badiolateritia 

Status: valid. Holotype (not visited): UK, Norfolk, Surlingham, Marsh Cottage Carr, 8 

Sep. 1972 (Orton 4425, E). Host: Salix sp. 

Basionym: Naucoria badiolateritia P.D. Orton, Notes Royal Bot. Gard. Edinburgh 41 

(3): 598 (1984). 

≡ Alnicola badiolateritia (P.D. Orton) Courtec. 1985, Doc. mycol. 16 (61): 48 

Notes: E. Horak has studied a collection labelled "Naucoria 

badiolateritia" at E (Orton n° 4426), but this belongs to A. clavuligeroides, as 

quoted by Orton himself (1984: 600). 

Accepted name: Alnicola badiolateritia (P.D. Orton) Courtec. 

bohemica 

Status: valid, not typified. Original material not found in PRM and Brno. Host: 

unknown. 

Basionym: Naucoria bohemica Velen., Česke Houbý: 527 (1921) (basionyme). 

≡ Alnicola bohemica (Velen.) Kühner, genre Galera: 12 (1935). 

Notes: the combination of this well-known species in the genus Alnicola 

has been reported erroneously in literature, attributed to Kühner & Maire 

(Romagnesi, 1937; Reid, 1984) or Singer (Singer, 1949, 1986). None of them 

has been effectively published. Looking for cryptic combinations, Kühner’s 

(1935) publication is the oldest the author could trace, valid thanks to the 

indication "Alnicola (Naucoria) bohemica (Velen.)", which refers explicitely to 

Naucoria bohemica Velen. 

Accepted name: Alnicola bohemica (Velen.) Kühner. 

131

evicystis 

Status: invalid, art. 36.1. No original material could be found in Métrod’s collections 

(PC). Host: unknown. 

Original name: Alnicola brevicystis Métrod, Bull. Soc. Natur. Oyonnax 14-15 [1962 

(1960-1961)]: 144. 

Notes: According to the description and unpublished notes by Métrod 

(PC), this singular species should be compared with A. zonata, but this last has 

much smaller spores. 

cedriolens 

Status: valid. Holotype (not visited): D, nordöstlich Gotezenalm, 1600 m, 12-9-1979, n° 

175, MB 8444/3 (indicated as preserved in herb. M by Schmid-Heckel, 1985). Host: Alnus 

viridis. 

Basionym: Naucoria cedriolens Bresinsky & Schmid-Heckel in Schmid-Heckel, 

National Park Berchtesgaden 8: 163 (1985). 

≡ Alnicola cedriolens (Bresinsky & Schmid-Heck.) Bon, Doc. mycol. 21 (83): 37 

(1990). 

Notes: the type material could not be traced in the Botanische 

Staatssammlung München, and therefore could not be revised. According to 

the author’s own experience of subalpine alder shrubs, this taxon is nothing but 

a fragrant forma of A. luteolofibrillosa, this last being very common under 

Alnus viridis but not reported by Schmid-Heckel (1985: 163). 

Accepted name: Alnicola cedriolens (Bresinsky & Schmid-Heck.) Bon 

(if different from A. luteolofibrillosa Kühner). 

celluloderma 

Status: valid. Holotype (visited): UK, Downwood, Shobdon, Herefordshire, 31-X-1959 

(K). Host: Alnus sp. 

Basionym: Naucoria celluloderma P.D. Orton, Trans. Brit. mycol. Soc. 43 (2): 314 

(1960). 

≡ Alnicola celluloderma (P.D. Orton) Svrček, Česká Mykol. 20: 41 (1966). 

≡ Alnicola celluloderma (P.D. Orton) Romagn., Bull. Fed. mycol. Dauphiné-Savoie 74: 

19 (1979) (comb. superfl.). 

≡ Alnicola alnetorum var. celluloderma (P.D. Orton) Bon, Doc. mycol. 20 (78): 40 

(1990). 

≡ Alnicola alnetorum var. celluloderma (P.D. Orton) Krieglst., Veirbreitungsatlas 

Grosspilze Deutschl. 1B: 422 (1991) (comb. inval., art. 33.3). 

Notes: the diffences between A. celluloderma and A. inculta are mainly 

organoleptic, as already pointed out by Reid (1984) and Bon (1992: 15). No 

reliable differences could be found so far. 

Accepted name: Alnicola inculta (Peck) Singer, s.l. 

132


chamiteae 

Status: valid. Holotype: F, Col de l’Iseran, le Pays désert, 2650 m, 21-8-1971 (herb. 

Kühner n° K.71.70, G). Host: Salix herbacea. 

Basionym: Alnicola chamiteae Kühner, Trav. Sci. Parc natl Vanoise 11: 133 (1981). 

≡ Naucoria chamiteae (Kühner) Senn-Irlet, Mycol. helvetica 2 (1): 47 (1986). 

Notes: Kühner described A. chamiteae by misinterpreting A. tantilla, as 

already suggested by De Haan (2000). Revision of original collections of both 

taxa reveal a complete identity (Moreau, unpublished). 

Accepted name: Alnicola tantilla (J. Favre) Romagn. 

cholea 

Status: valid. Holotype (visited): F, Pralognan-la-Vanoise, cirque du Génépy, (herb. 

Kühner n° G K.62.36). Host: Salix spp. 

Basionym: Alnicola cholea Kühner, Trav. Sci. Parc natl Vanoise 11: 133 (1981). 

Note: a rare but very distinctive species of arctic-alpine distribution. 

Accepted name: Alnicola cholea Kühner. 

clavuligera 

Status: valid. Holotype: F, Sens, Yonne, 10-IX-1941 (PC). Substrate: Alnus glutinosa or 

Salix sp., on wood. 

Basionym: Alnicola clavuligera Romagn., Bull. Soc. mycol. France 58 (2): 148 (1942). 

≡ Naucoria clavuligera (Romagn.) Pilát, Klíč: 271 (1951). 

≡ Naucoria clavuligera (Romagn.) Kühner & Romagn., Fl. Anal. Champ. Sup.: 236 

(1953) (comb. inval., art. 33.3). 

≡ Hebeloma clavuligerum (Romagn.) P. Collin, Doc. mycol. 19 (74): 61 (1988). 

Notes: according to the study of holotype (absence of differenciated 

subpellis and thick-walled, undextrinoid spores), this species most likely 

belongs to Galerina. The suprapellis with typically capitate cystidia-like 

terminal hyphae is analogous to G. albotomentosa (spores punctuate) and 

G. permixta (spores smooth). It is probably identical to Naucoria salicetorum 

D.A. Reid (see under this name). 

Accepted name: Galerina clavuligera (Romagn.) P.-A. Moreau, comb. 

nov. 

clavuligeroides 

Status: valid. Holotype (visited by E. Horak): UK, Norfolk, Surlingham, Parish Marsh - 

9 Oct. 1971, Orton 4186 (E). A spore print (visited) is deposited in K. Host: not precised 

(Corylus, Alnus, Salix). 

Basionym: Naucoria clavuligeroides P.D. Orton, Notes Royal Bot. Gard. Edinburgh 41 

(3): 599 (1984) (based on N. clavuligera ss. Orton in sched. (K); ss. Reid 1984: 199). 

≡ Alnicola clavuligeroides (P.D. Orton) Courtec., Doc. mycol. 16 (61): 48 (1985). 

Notes: according to E. Horak’s personal notes on the holotype, this 

species is clamped with hymeniform pileipellis; but another collection (Orton 

4426, Parish Marsh, 6-IX-1972, E), also studied by E. Horak, has a pileipellis 

of A. salicis-type, more conform to Orton’s (1984: 600) description. A. 

133

clavuligeroides still has an ambiguous systematic position, possibly close to A. 

erebia (Huijsm.) Romagn. It may also be an Hebeloma, if clamp connections 

are confirmed. So far only known from the type locality. 

Accepted name: Alnicola clavuligeroides (P.D. Orton) Courtec. 

dasypus 

Status: valid, not typified. Lectotype (designed here): F, Paris, jardin du Muséum de 

Paris, novembre 1934 (herb. H. Romagnesi, PC; ½ specimen). Host: unknown (amongst dead 

leaves), probably not mycorrhizal. 

Basionym: Naucoria dasypus Romagn., Bull. Soc. mycol. France 53: 121 (1937). 

≡ Alnicola dasypus (Romagn.) Romagn., Bull. Soc. mycol. France 58: 126 (1942). 

≡ Hebeloma dasypus (Romagn.) Singer, Lilloa 23: 536, 538 (1949 [1952]). 

≡ Hebeloma dasypus (Romagn.) Bon, Docum. Mycol. 21 (83): 37 (1991) (comb. 

superfl.). 

Description (lectotype): spores 9-13,5 × 5,5-7 µm, ochre yellow, 

distinctly verruculose with large supra-apicular plage, with distinct pore 0,8-1 

µm large. Basidia (2-)4-spored, 22-35 × 8-10 µm, shortly clavate or with more 

or less tapering base. Subhymenium pseudoparenchymatous. Cheilocystidia 

not observed (due to strongly parasited gill edge), clavate-capitate 27-40 × 10- 

14 µm according to Romagnesi (1937: 122). Suprapellis an (ixo?)hymenoderm 

of cylindro-clavate to pyriform articles, 20-32 × 8-17 µm, thin- to slightly 

thick-walled, smooth. Subpellis filamentous, thin, smooth. Caulocystidia 

numerous all over the stipe, often in fascicles, cylindro-clavate, sometimes 

septate, 35-75 × 9-17 µm. Pileal and hymenial trama heteromerous, with 

strongly inflated to almost globulose hyphae × 30-45 µm, mixed with 

cylindrical, very long hyphae up to 1000 × 8-10 µm, and inconspicuous 

generative hyphae × 2,5-4 µm. Clamp connections small and not conspicuous 

but present at all septa. 

Notes: only one collection, here designed as lectotype, could be found in 

Romagnesi’s personal herbarium (PC). Gill edge and pileipellis are parasited 

and can hardly be observed, and Romagnesi’s description of pileipellis is 

erroneous due to the parasit (“laticifères ramifiés, sinueux”). In fact intact parts 

of the pileipellis can be found and are typically hymeniform. Romagnesi 

himself wrote later on his pack: “Serait Conocybe subnuda R. Maire selon 

Kühner”, was is confirmed here. This collection presents particularly 

ornamented spores, but falls into the high variability of Pholiotina subnuda as 

reevaluated by Hausknecht (1993) and Meusers (1996); Romagnesi’s name 

becomes the older valid name for this species. In a more restricted sense 

(Kühner, 1935; Bon, 1992), Naucoria dasypus comes closer to P. subnuda ss. 

str. by cystidial shape (according to Romagnesi, 1937: 122), but sporal 

ornamentation is as conspicuous as in P. subverrucispora, which should have 

partly fusiform cystidia. 

134


Accepted name: Pholiotina dasypus (Romagn.) P.-A. Moreau, comb. 

nov. 

Taxonomic synonym : Conocybe subnuda R. Maire ex Kühner & Watling in Watling 

1983, Roy. bot. Gard. Edinburgh 40 (3): 553, ≡ Pholiotina subnuda (Kühner ex Kühner & 

Watling) Bon 1991, Doc. mycol. 21 (83): 39. 

(for exhaustive synonymy see Hausknecht, 1993) 

devia 

Status: valid. Holotype (not visited): ARG, prov. Buenos Aires, Punta Lara, 9-VI-1949 

(R. Singer n° S111, LIL). Host: Salix humboldtiana. 

Basionym: Alnicola devia Singer, Sydowia 4: 150 (1950). 

≡ Naucoria devia (Sing.) Raithelh., Fl. Mycol. Argentina 2: 121 (1988). 

Notes: only known from the type locality. The author has no opinion 

about this species. 

Accepted name: Alnicola devia Singer. 

diplocystis 

Status: valid. Holotype (not visited): ARG, prov. Tucumán, Supra Tafi del Valle, 1-V- 

1949 (R. Singer n° T503, LIL). A part of the holotype (3 specimens, visited) sent by R. Singer 

to H. Romagnesi has been found in Romagnesi’s herbarium (PC). Host: Alnus jorullensis. 

Basionym: Alnicola diplocystis Singer, Sydowia 4: 151 (1950). 

≡ Naucoria diplocystis (Singer) Raithelh., Fl. Mycol. Argentina 2: 120 (1988). 

Notes: this is the most common Alnicola species growing under Alnus 

jorullensis, and apparently also A. acuminata, all along the Cordillera (Singer, 

1950b: 152). Its distribution and affinities with the European species are still 

unclear. 

Accepted name: Alnicola diplocystis Singer. 

dubis 

Status: valid. Holotype: F, Doubs, Deluz, 22 novembre 1953, herb. G. Métrod (PC). 

Host: Alnus glutinosa. 

Basionym: Alnicola dubis Métrod ex P.-A. Moreau & Vidonne, Doc. mycol. 34 (133- 

134), 2005, in press. 

Origianl name: Alnicola dubis (‘dubii’) Métrod, Bull. Soc. Natur. Oyonnax 14-15: 143 

(1962 [1960-1961]) (inval., art. 36.1). 

Notes: this taxon is validated and compared to A. scolecina ss. Lange 

(1938) by Moreau and Vidonne (2005). 

Accepted name: Alnicola dubis Métrod ex P.-A. Moreau & Vidonne. 

erebia 

Status: valid. Holotype (not visited): NL, prope lacum ‘Quackjeswater’, insluar Voorne, 

12 jul. 1972 (not found in Huijsman’s herbarium, L). Host: Betula or Salix. 

Basionym: Naucoria erebia Huijsman, Kew Bull. 31 (3): 585 (1976). 

≡ Alnicola erebia (Huijsm.) Romagn., Bull. Fed. mycol. Dauphiné-Savoie 74: 19 

(1979). 

135

Notes: the author cannot interpret this species, with abundant clamp 

connections according to Huijsman (1976) but otherwise very similar to some 

four-spored collections of A. salicis. To be compared with A. clavuligeroides 

(P.D. Orton) Courtec.; possibly also an Hebeloma. 

escharoides 

Status: valid, not typified. Original material: none; authentic material: 1 unpublished 

plate directed by Fries, Femsjö, 26-IX-1854, Akerlund pinx. (S). Host: undefined. 

Basionym: Agaricus escharoides Fr., Obs. Mycol. 2: 131 (1818): Fr, Syst. mycol. I: 260 

(1821) ("escharioides", corrected by Fries, 1857-1863: 478). 

≡ Naucoria escharoides (Fr. : Fr.) P. Kumm., Führer Pilzk.: 76 (1871). 

≡ Hylophila escharoides (Fr. : Fr.) Quél., Enchir. Fung.: 104 (1886). 

≡ Alnicola escharoides (Fr. : Fr.) Romagn., Bull. Soc. mycol. France 58: 126 (1942). 

Notes: Ag. escharoides is a critical taxon, which nevertheless has been 

proposed as a neotype for Alnicola (Konrad, 1934) and as a lectotype for 

Naucoria (Horak, 1968). Whether Ag. escharoides can be considered as a 

member of the genus Alnicola or not is debatable. Fries (1818: 131; 1821: 260) 

originally describes a heavily veiled species, including probably 

A. luteolofibrillosa (Kühner, 1942: 7) but also Tubaria conspersa and related. 

Only Ag. escharoides β naucosus Fr. : Fr. (Fries, 1818: 132) is in agreement 

with the most usual interpretation of Ag. escharoides (Konrad and Maublanc, 

1926, pl. 175 II; Lange, 1939, pl. 125E), as an alnicolous species with 

darkening stipe ("l. c emarginatae, stipite fuscescente"), but it contradicts the 

definition of Ag. escharoides typical: "stipes (...) ut totus fungus pallidus, (...) 

lamellae primo subdecurrentes". If this variety matches the current concept of 

Alnicola escharoides (defined by Konrad and Maublanc, loc. cit.) and can be 

considered as a typical Alnicola, it is certainly not representative of Fries’ 

initial concept of Ag. escharoides. 

Fries evolved in his concept of Ag. escharoides in the ‘Monographia’ 

(1857-1863: 383): this later description excludes collections with abundant veil 

(and consequently A. escharoides is cited below Ag. conspersus after this date). 

In fact he probably restricts it mainly to his former "var. naucosus” : the same 

indications, e.g. ‘Supra Hallandsås Scaniae’ are given for naucosus (1918: 132) 

and escharoides later (1836-1838: 201; 1857-1863: 383). By the way Fries 

emends Ag. conspersus, in which a wide part of his original typical 

Ag. escharoides is obviously included. 

Amongst the aquarelles directed by E. Fries preserved at S, only one is 

labelled "Agaricus (Naucoria) escharoides" (n° 645: Akerlund pinx., Femsjö, 

26.IX.1854, 5 specimens). These specimens are dark brown with pale gills, do 

not show any veil, and Fries has written and erased several names before 

choosing this one at last. The author rather tend to refer them to Inocybe 

calospora Quél. 

136


The wide and somewhat unstable conception Fries had of Ag. 

escharoides (and of its limits with Ag. conspersus) is also illustrated by his 

reference (as “var." sine nomine; Fries, 1874: 264) to Ag. pulverulentus 

Schaeff., t. 226 (fig. II), a distant-gilled, white-spored fungi comparable to a 

Laccaria, and somewhat reminiscent of a Tubaria species, not of any Alnicola. 

Konrad and Maublanc (1926) have interpreted A. escharoides as closely 

as possible to Fries’ later descriptions (Fries, 1836-1838, 1857-1863, 1874); 

but not according to the original sense (Fries, 1818, 1821), which is 

significantly different. However it is premature to propose a name in 

replacement of Ag. escharoides without a deep taxonomic revision of all the 

taxa involved and a revision of Swedish collections. 

In Moser’s herbarium (IB) a collection labeled "Naucoria escharoides" 

from Femsjö (SW) might be considered as a reference, but corresponds to an 

unveiled taxon. At this early stage of the author’s revision this collection 

cannot surely be identified to any currently well-circumscribed European 

taxon. 

Ag. escharoides var. naucosus Fr. : Fr. seems to be the most correct name 

for Ag. escharoides ss. auct. pl. 

fellea 

Status: valid. Holotype (visited): CH, Le Locle, tourbière du Grand Cachot (herb. 

J. Favre n° GK 9457, G). Host: Betula pendula or Pinus uncinata according to Favre (1948: 

133). 

Basionym: Alnicola scolecina var. fellea J. Favre, Beitr. Kryptogamenfl. Schweiz 10 

(3): 214 (1948). 

≡ Alnicola fellea (J. Favre) Courtec., Doc. mycol. 16 (61): 48 (1985). 

≡ Naucoria fellea (J. Favre) Raithelh., Metrodiana 7 (2-3): 53 (1978). 

Notes: quite an enigmatic species, so far only known from the type 

collection. Favre compared it to A. amarescens, from which it differs 

consistently by spore shape and ornamentation. Possibly illustrated by 

Breitenbach and Kränzlin (2000, pl. 128) as A. amarescens. More 

investigations on this interesting taxon are required. 

Accepted name: Alnicola fellea (J. Favre) Courtec. 

fulgens 

Status: valid, not typified. Original material in coll. J. Favre (G) and R. Maire (MPU); 

additional topotypic collections in R. Kühner (G). Substrate: manured peaty ground. 

Basionym: Naucoria fulgens J. Favre & Maire, Bull. Soc. mycol. France 53: 267 (1937). 

≡ Fulvidula fulgens (J. Favre & Maire) Kühner, Bull. Soc. linn. Lyon 8 (2): 44 (1939). 

≡ Alnicola fulgens (J. Favre & Maire) Singer, Rev. mycol. (Paris) 4 (1-2): 68 (1939) 

(inval., art. 33.3); Not. Sect. crypt. Inst. bot. Acad. Sci. U.S.S.R. t.5, N: 7-9: 93 (1941). 

≡ Gymnopilus fulgens (J. Favre & Maire) Singer, Lilloa 22: 561 (1949 [1951]). 

137

≡ Flammula fulgens (J. Favre & Maire) S. Lundell, Fungi Exsiccati Suecici 55: 9 

(1960). 

Notes: this well-known taxon was at first temptatively classified in 

Alnicola sect. Submelinoideae by Singer (1939: 68; 1950a: 413), before being 

recognized as a Gymnopilus. 

Accepted name: Gymnopilus fulgens (J. Favre & Maire) Singer. 

fusispora 

Status: valid, not typified. Original material not found in PRM and Brno. Host: 

undefined. 

Basionym: Galera fusispora Velen., Novitates I: 128 (1939). 

≡ Alnicola fusispora (Velen.) Singer, Fieldiana Bot. N.S. 21: 116 (1989). 

Notes: Singer (1989) gave no justification of his combination in Alnicola. 

According to Velenovsky’s (1939: 128) description, the author feels unable to 

interpret precisely this species, apparently belonging to the group of A. 

macrospora, and prefers considering it as a nomen dubium. 

geraniolens 

Status: valid. Holotype (visited): F, Santes, près de Lille (Nord), 20-X-1982, herb. 

R. Courtecuisse n° 82.10.20.03 (LIP). Host: Salix viminalis, S. caprea. 

Basionym: Alnicola geraniolens Courtec., Doc. mycol. 13 (50): 55 (1983). 

≡ Naucoria geraniolens (Courtec.) G. Keller in Keller & Moser, Biosystem. Ecol. 19: 

177 (2001). 

≡ Naucoria amarescens var. geraniolens (Courtec.) E. Ludwig, Pilzkompendium 1: 421 

(2001). 

Accepted name: Alnicola geraniolens Courtec. 

gracillima 

Status: invalid, art. 36.1. Original material (icon): pl. 125F in Lange (1939). Host: Alnus 

glutinosa. 

Original name: Naucoria scolecina f. gracillima J.E. Lange, Dansk Bot. Arkiv 9 (5): 20 

(1938) (inval.); Fl. agar. danica V: 21 (1939) (inval.). 

Notes: no validating latin diagnosis was published by Lange (1940) for 

this taxon. It is interpreted by Moreau (2004) as a septentrional variant of 

Alnicola umbrina. Its taxonomic level and the opportunity of a validation 

requires further observations. 

Accepted name: Alnicola umbrina (R. Maire) Kühner s.l. (see Moreau, 

2004). 

inculta 

Status: valid. Holotype (visited by E. Horak): USA, Catskill mountains, Sept. 1887 

(MICH). Host: willows and alders (most probably Alnus sp.). 

Basionym: Galera inculta Peck, Ann. Rep. N.Y. St. Mus. 41: 69 (1888). 

≡ Galerula inculta (Peck) G.F. Atk., Proc. Amer. philos. Soc. 48: 362 (1918). 

≡ Alnicola inculta (Peck) Singer, Sydowia 9: 406 (1955) (comb. inval., art. 33.3). 

138


≡ Alnicola inculta (Peck) Singer, Sydowia 15 (1-6): 71 (1961 [1962]). 

Notes: already described by Singer (1955) to be an earlier synonym of 

A. alnetorum, this is confirmed by E. Horak’s observations (personal notes, 

ZT). Peck (1888: 69) describes it correctly as looking like "Clitocybe laccata 

in its small glabrous striatulate form". 

Accepted name: Alnicola inculta (Peck) Singer. 

iodiolens 

Status: valid. Holotype (visited): Germany: Berlin-Wannsee (Heckensorn), on the edge 

of "Grosser Wannsee", 1-12-86 (Hort. Bot. Berlin-Dahlem, 1 specimen in poor condition). 

Host: Alnus glutinosa. 

Basionym: Naucoria scolecina var. iodiolens E. Ludwig, Pilzkompendium 1: 417 

(2001). 

Notes: although known only from one collection, this taxon close to 

A. scolecina ss. Lange (1938, 1939) is clearly defined by its very long spores 

and, according to Ludwig (2001), by its peculiar smell (inde nomen). This 

smell is distinctive in the genus Alnicola, if not due to the advanced age of the 

specimens; according to most other features it also comes close to A. 

subconspersa and A. dubis, the only significant difference being the very long 

spores. For redescription and further discussion see Moreau and Vidonne 

(2005). 

Accepted name: ‘Naucoria’ scolecina var. iodiolens E. Ludwig. 

lactariolens 

Status: valid. Holotype: JAP, Tomikawa, Otsu-shi, Shiga-ken, 15-VIII-1988, leg. 

T. Hongo & H. Clémençon (HC 88/95, TNS); isotype (visited) in LAU. 

Basionym: Alnicola lactariolens Clémençon & T. Hongo, Mycoscience 35 (1): 25 

(1994). 

Notes: this species seems better placed in the genus Anamika, but the 

purplish spore print is unusual for this genus (B. Matheny, comm. pers.). 

langei 

Status: invalid, art. 32.3 and 34.2; as nomen novum for Naucoria macrospora 

J.E. Lange, illegit. 

Original names: Alnicola langei J.E. Lange → Kühner in Kühner & Romagnesi, Bull. 

Soc. Natur. Oyonnax 10-11: 3 (1957); Naucoria langei J.E. Lange → Kühner in Kühner & 

Romagnesi, Bull. Soc. Natur. Oyonnax 10-11: 15 (1957). 

= Alnicola langei (Kühner) Singer, Sydowia 30 (1-6): 207 (1977 [1978]) (inval.). 

= Alnicola langei (Kühner) Romagn., Bull. Fed. mycol. Dauphiné-Savoie 74: 19 (1979) 

(inval.). 

Notes: In Kühner’s mind (in Kühner and Romagnesi, 1957: 11; 

unpublished notes, herb. G), N. langei is only a nomen novum for N. 

macrospora J.E. Lange (illegit.), including a 2-spored (typical, not seen by 

Kühner) and a 4-spored forma (described by Kühner, loc. cit. p. 9). If no 

139

taxonomic value is given to the number of sterigmata in this group (Moreau 

and Deïana, 2004), the collection described by Kühner (in Kühner and 

Romagnesi, 1957) can be reported to Alnicola macrospora J.E. Lange → J. 

Favre (= A. salicis), or in a restricted sense to A. saliceti (see ‘tetraspora’ 

below). Kühner (1957) retrospectively admits that A. bohemica "4-sp." of 

Maire and Kühner (1934) also corresponds to this taxon. See also Horak and 

Moreau (2005). 

Accepted name: Alnicola macrospora J.E. Lange → J. Favre. 

leucocnemis 

Status: valid. Holotype (visited): F, La Chapelle-en-Serval, près Fosses, Oise (PC). 

Host: undefined. 

Basionym: Alnicola leucocnemis Romagn., Bull. Soc. mycol. France 102 (2): 129 

(1986). 

≡ Naucoria leucocnemis (Romagn.) E. Ludwig, Pilzkomp. I Beischreib.: 53 (2001). 

Notes: Related to A. amarescens, but the limit between both taxa needs to 

be precised (Moreau and Garcia, 2005; see also Hausknecht and Forstinger, 

2004). 

Accepted name: Alnicola leucocnemis Romagn. (if different from 

A. amarescens). 

lignicola 

Status: valid, not typified (original material not traced). 

Basionym: Alnicola lignicola Singer, Rev. Mycol. (Paris) 4 (2): 67 (1939). Substrate: 

lignicolous on Picea schrenckiana. 

Notes: according to Singer’s description, this is a typical Gymnopilus, 

possibly G. picreus (Pers. : Fr.) P. Karst. ss. str. (Bon and Roux, 2002: 41). 

Alnicolus fulgens (currently a Gymnopilus) is mentioned by Singer (1939: 66) 

as a comparison. 

Accepted name: Gymnopilus aff. picreus (Pers. : Fr.) P. Karst. 

luteolofibrillosa 

Status: valid, not typified (several original collections in herb. Kühner, G). Host: Alnus 

viridis. 

Basionym: Alnicola luteolofibrillosa Kühner, Botaniste 17 (1-4): 175 (1926). 

≡ Naucoria luteolofibrillosa (Kühner) Pilát, Klič: 271 (1951). 

≡ Naucoria luteolofibrillosa (Kühner) Kühner & Romagn., Fl. anal. Champ. Sup.: 237 

(1953) (comb. inval., art. 33.3). 

Accepted name: Alnicola luteolofibrillosa Kühner. 

macrospora 

Status: valid, not typified. Lectotype (obligatory, designed here): Lange 1939, Fl. agar. 

danica 4 pl. 125B. 

140


Basionym: Alnicola macrospora J.E. Lange→ J. Favre, Beitr. Kryptogamenfl. Schweiz 

10 (3): 131 (1948). 

= Naucoria macrospora J.E. Lange, Dansk bot. Ark. 9 (5): 21 (1938) (inval., art. 37.1). 

= Naucoria macrospora J.E. Lange, Fl. agar. danica 5 p. VI (1940) (illegit., not 

N. macrospora Pat. & Doassans→ Bigeard & Guillemin, Fl. Champ. Fr. II p. 248 (1913). 

= Alnicola langei J.E. Lange→ Kühner in Kühner & Romagnesi, Bull. Soc. Natur. 

Oyonnax 10-11: 3 (1957) (inval.). 

= Naucoria langei J.E. Lange→ Kühner in Kühner & Romagnesi, Bull. Soc. Natur. 

Oyonnax 10-11: 15 (1957) (inval.). 

= Naucoria salicis P.D. Orton, Trans. brit. mycol. Soc. 43 (2): 318 (1960). 

Notes: Moreau and Deïana (2004) have redescribed this taxon focusing 

on its microscopical variability, but unfortunately had not noticed the 

haphazard validation of A. macrospora by Favre (1948: 130) and therefore 

used the name A. salicis, a posterior synonym (see under this name). The 

identity of this taxon with A. mirabilis is not excluded (Orton, 1960: 319). 

Accepted name: Alnicola macrospora J.E. Lange → J. Favre. 

melinoides 

Status: valid, not typified. Lectotype (automatic, designed here): Bulliard 1792, Herb. 

Fr., pl. 560, fig. 1 (plate without text). Substrate: undefined (grasslands). 

Basionym: Agaricus melinoides Bull., Herb. Fr., pl. 560, fig. 1 (1792): Fr. (latin name in 

“Table annuelle” of this publication). 

≡ Agaricus melinoides Bull. : Fr., Syst. Mycol. 1: 266 (1821). 

≡ Naucoria melinoides (Bull. : Fr.) P. Kumm., Führer Pilzk.: 77 (1871). 

≡ Alnicola melinoides (Bull. : Fr.) Kühner, Botaniste 17 (1-4): 176 (1926). 

Notes: The interpretation of Ag. melinoides as an Alnicola with necked 

cheilocystidia is due to Ricken (1915), and followed by a part of European 

authors (Kühner, 1926; Bon, 1979, 1992; Runge, 1990). But according to 

Watling and Gregory (1981: 167), Ag. melinoides has also been interpreted 

formerly as Galera mniophila (Lasch) Kühner by various authors, and as a 

Conocybe species by Fayod (1889) according to Singer (1978: 55, as Conocybe 

mesospora) and Watling and Gregory (loc. cit.: 167, as C. tenera). Cooke 

(1885, pl. 457) has made a clear illustration of Galerina clavata (Velen.) 

Kühner under this name. 

Bulliard’s plate cannot (intuitively) be interpreted as an Alnicola, nor the 

associated description (in Bulliard and Ventenat, 1792-1793: 244) but ratheras 

an undefined or collective species of Galerina. The author considers this name 

as a nomen dubium. 

mirabilis 

Status: valid. Holotype (visited by E. Horak): USA, NY, McLean, 19-VI-1903, leg. 

Whetzel, herb. G. Atkinson n° 15117 (CUP). Host: Salix sp. 

Basionym: Galerula mirabilis G.F. Atk., Proc. Amer. philos. Soc. 58: 367 (1918). 

141

≡ Pholiotina mirabilis (G.F. Atk.) Singer, Acta Inst. bot. Kamarov Acad. Sci. URSS, 

ser.2; 6: 425 (1950). 

≡ Alnicola mirabilis (G.F. Atk.) Singer, Sydowia 9: 406 (1955) (comb. inval., art. 33.3). 

≡ Alnicola mirabilis (G.F. Atk.) Singer, Sydowia 15: 70 (1961 [1962]). 

≡ Naucoria mirabilis (Atk.) Watling in Watling & Gregory, Bibl. mycol. 82: 121 (1981) 

[illegit., non N. mirabilis Velen., Česke Houby 1: 522 (1921)]. 

Notes: the holotype also visited by R. Singer (January 1952), annoted : 

"= Alnicola bohemica (Velen.) Kühner = N. hamadryas ss. A.H. Sm. = 

N. scorpioides ss. Lange". Singer published this first synonymy (1955: 406), 

but revised it later (1961: 70) following Orton’s opinion (1960: 319), as a 

possible synonym of A. salicis (= A. macrospora). 

Accepted name: Alnicola mirabilis (G.F. Atk.) Singer. 

paludosa Peck 

Status: valid. Holotype (not visited): USA, Catskill mountains, sept. 1887 (MICH). 

Host: willows and alders (most probably Alnus sp.). 

Basionym: Naucoria paludosa Peck, Ann. Rep. N.Y. St. Mus. 41: 68 (1888). 

≡ Alnicola paludosa (Peck) Singer, Sydowia 30 (1-6): 210 (1977 [1978]). 

Notes: Singer (1977) considered this taxon as a synonym of A. striatula 

(P.D. Orton) Romagn. The material has not been revised by the author yet. 

Jugding from Singer’s and Peck’s notes A. paludosa might be an American 

vicariant of A. umbrina. According to Murrill (1917: 178), 

Ag. pallidomarginatus Peck and Naucoria uliginosa Peck are also synonym; 

types have not been revised either. 

paludosa Velen. 

Status: valid, illegit. (non N. paludosa Peck, 1888). No type could be found in BRN or 

PRM. Host: willows and alders (most probably Alnus sp.). 

Basionym: Naucoria paludosa Velen., Novit. Mycol.: 124 (1939). 

Notes: according to Singer (1977: 208), this taxon is a synonym of 

Alnicola scolecina (consequently in the sense of Singer, loc. cit.: 210; see 

under this name). 

phaea 

Status : valid. Type = type of A. badia Kühner, not typified, see badia. 

Basionym: Alnicola badia Kühner, Bull. Soc. mycol. France 47 (3-4): 239 (1931). 

≡ Naucoria phaea Kühner � R. Maire & Kühner in Maire, Bull. Inst. bot. Barcelona 3 

(4): 101 (1937) (nomen novum for Alnicola badia in the genus Naucoria, non Naucoria badia 

Murrill, 1917). 

≡ Alnicola phaea (Kühner � R. Maire & Kühner) Romagn., Bull. Soc. mycol. France 

58: 126 (1942) (comb. superfl., = A. badia Kühner). 

Notes: see under BADIA. 

Accepted name: Alnicola badia Kühner. 

142


pseudoamarescens 

Status: valid, not typified (original material in herb. R. Kühner, G., and H. Romagnesi, 

PC, visited). Substrate: charcoal, but probably ectomycorrhizal. 

Basionym: Alnicola pseudoamarescens Kühner & Romagn. in Kühner, Ann. Sci. 

Franche-Comté 2: 17 (1947). 

≡ Naucoria pseudoamarescens (Kühner & Romagn.) Kühner & Romagn., Fl. anal. 

Champ. sup.: 236 (1953) (comb. inval., art. 33.3). 

≡ Hebeloma pseudoamarescens (Kühner & Romagn.) P. Collin, Doc. mycol. 19 (74): 

61 (1988). 

≡ Hebeloma pseudoamarescens (Kühner & Romagn.) Kühner & Romagn. ex A. Runge, 

Mitt. Arbeitsgemeinsch. Pilzk. Niederrhein 8 (2): 94 (1990) (comb. superfl.). 

Notes: this is a typical member of the genus Hebeloma, because of its 

gelified pileipellis and narrowly elliptic spores, as recognized by Romagnesi 

(1986) and Kühner (1987) themselves. Romagnesi (1989) postulated a 

synonymy with Hebeloma funariophilum M. Moser, not supported by type 

studies. 

Accepted name: Hebeloma pseudoamarescens (Kühner & Romagn.) 

P. Collin. 

pseudoscolecina 

Status: valid. Holotype (visited): UK, Ripley, Surrey, 12-X-1952 (K). Host: Alnus sp. 

Basionym: Naucoria pseudoscolecina D.A. Reid, Trans. Brit. mycol. Soc. 82 (2): 202 

(1984). 

≡ Alnicola pseudoscolecina (D.A. Reid) Courtec., Doc. mycol. 13 (50): 55 (1983). 

Accepted name: Alnicola pseudoscolecina (D.A. Reid) Courtec. (if 

distinct from A. umbrina (R. Maire) Kühner). 

rubriceps 

Status: valid. Holotype (visited by E. Horak): UK, Norfolk, Surlingham, Tucks East, 16 

oct. 1972 (Orton 4428, E). Host: not precised (deciduous trees). 

Basionym: Naucoria rubriceps P.D. Orton, Notes Royal Bot. Gard. Edinburgh 41 (3): 

600 (1984). 

≡ Alnicola rubriceps (P.D. Orton) Courtec., Doc. mycol. 16 (61): 48 (1985). 

Notes: E. Horak reported the type collection to have unclamped hyphae 

(not specified by Orton, 1984) and strictly bisporical basidia. It comes close to 

A. macrospora. 

Accepted name: Alnicola rubriceps (P.D. Orton) Courtec. 

salabertii 

Status: valid. Holotype (visited): F, Hérault, Graissessac, terril de Garella, 29 oct. 2003 

(P.-A. Moreau 03102993, LIP). Host: Alnus cordata. 

Basionym: Alnicola salabertii Bon & Boutev. ex P.-A. Moreau & Guy Garcia, Bull. 

Soc. mycol. France 120 (1-4): 275 (2005). 

= Alnicola leucocnemis f. salabertii Bon & Bouteville ad int., Bull. Fed. Assoc. mycol. 

médit. 18: 38 (2000). 

143

Notes: a very distinct species, hitherto only reported under Alnus cordata 

(Bon and Bouteville, 2000; Moreau and Garcia, 2005). Possibly related to A. 

diplocystis Singer, but not comparable to any European taxon. 

Accepted name: Alnicola salabertii Bon & Boutev. ex P.-A. Moreau & 

Guy Garcia. 

saliceti 

Status: valid. Holotype (visited by E. Horak): UK, Dorset, Studland, 11 Aug. 1971 

(Orton 4187, E). Host: Salix. 

Basionym: Naucoria saliceti P.D. Orton, Notes Royal Bot. Gard. Edinburgh 41 (3): 601 

(1984). 

≡ Alnicola saliceti (P.D. Orton) Courtec., Doc. mycol. 16 (61): 48 (1985). 

Notes: Orton (1984) published this name in order to rename N. langei 

Kühner (4-spored form), invalid and ambiguous (Kühner and Romagnesi, 

1957). According to Horak (pers. notes, ZT), the specimens constituting the 

holotype are variable concerning the number of sterigmata, some of them 

predominantly 2-sp. Therefore A. saliceti can also be interpreted as a single 

synonym of A. macrospora (= A. salicis), the differences between both species 

being very weak. 

Accepted name: Alnicola macrospora J.E. Lange → J. Favre s.l. 

salicetorum 

Status: valid. Holotype (visited): UK, Warwicks, Bickenhill, leg. A.W. Brand, 10-X- 

1971 (n° AWB 184, K). Host: Salix spp. 

Basionym: Naucoria salicetorum D.A. Reid, Trans. Brit. mycol. Soc. 82 (2): 202 

(1984). 

≡ Alnicola salicetorum (D.A. Reid) Courtec., Doc. mycol. 16 (61): 48 (1985). 

Notes: the holotype and an additional collection show a pileic structure 

analogous to that of Galerina permixta (P.D. Orton) Pegler & Young. No 

subpellis can be distinguished. Therefore the author consider that this species 

belongs to the genus Galerina, close to G. permixta and G. clavus Romagn. 

IThe author provisionally considers it as a synonym of A. clavuligera Romagn. 

according to the study of holotypes and descriptions; the only discrepancy is 

the absence of veil in A. clavuligera according to Romagnesi (1942). 

Accepted name: Galerina clavuligera (Romagn.) P.-A. Moreau. 

salicis 

Status: valid. Holotype (visited): UK, Thack Marsh, Surlingham, Norfolk, 5-IX-1959- 

Dorset (K). Host: Salix spp. 

Basionym: Naucoria salicis P.D. Orton, Trans. Brit. mycol. Soc. 43 (2): 318 (1960). 

≡ Alnicola salicis (P.D. Orton) Bon, Doc. mycol. 9 (35): 41, 49 (1979, March). 

≡ Alnicola salicis (P.D. Orton) Romagn., Bull. Fed. mycol. Dauphiné-Savoie 74: 19 

(1979, July), (comb. superfl.). 

Notes: first described as a collective taxon, as a new name for 

144


N. macrospora J.E. Lange. Agaricus salicis has been splitted by Orton (1984), 

the holotype of A. salicis being redescribed more correctly by Reid (1984: 

203). see under MACROSPORA. 

Accepted name: A. macrospora J.E. Lange → J. Favre. 

scolecina 

Status: valid, not typified. Lectotype (automatic, designed here): original plate by E. 

Fries (n° 665, S), published by Fries (1877-1884, pl. 124-1), mentioned in the protolog. Host: 

Alnus sp. (probably A. incana). 

Basionym: Agaricus scolecinus Fr., Epicr.: 194 (1838). 

≡ Naucoria scolecina (Fr.) Quél., Champ. Jura Vosges: 438 (1875). 

≡ Hylophila scolecina (Fr.) Quél., Ench. Fung.: 101 (1886). 

≡ Hylophila sideroides var. scolecina (Fr.) Quél., Fl. Mycol.: 87 (1888). 

≡ Alnicola scolecina (Fr.) Romagn., Bull. Soc. mycol. France 58 (4): 122 (1942). 

Notes: this taxon is interpreted in modern literature following Lange 

(1938) and Romagnesi (1942), as a bitter, not striate species, often growing 

cespitose. This is certainly not in conformity with Fries’ definition (Fries, 

1838), which concerns an originally very pale and glabrous species. Kühner 

(1980: 933, 967) suggested that A. scolecina ss. Fries might be identical to A. 

inculta (Peck) Singer (= A. alnetorum, inval.), what is acceptable according to 

Fries’ (1877-1884, pl. 124-1) plate; moreover Fries specifies (loc. cit.: 22) that 

this plate represents a more coloured forma. At the opposite Singer (1977: 

210), on the basis of personal collections near Femsjö, interprets A. scolecina 

as a small-spored species with acute cystidia. This taxon should be redefined 

and neotypified after a complete revision of Scandinavian species of Alnicola, 

and if revealed to be identical to A. inculta, asked for a rejection in favour of 

the last. 

Provisionally Lange’s (1938, 1939) interpretation (certainly inadequate 

but usual) is adopted by Moreau and Vidonne (2005). 

Accepted name: Alnicola scolecina (Fr.) Romagn. 

silvae-novae 

Status: valid. Holotype (visited): UK, Denny, Hants, sept.1982 (K). Host: Alnus sp. 

Basionym: Naucoria silvae-novae D.A. Reid, Trans. Brit. mycol. Soc. 82 (2): 206 

(1984). 

≡ Alnicola silvae-novae (D.A. Reid) Courtec., Doc. mycol. 16 (61): 48 (1985). 

Notes: the bispory, probably accidental (and only partial) mentioned by 

Reid (1984) as a specific character of A. silvae-novae, cannot be considered as 

such, since it can occur potentially in all 4-spored species in Alnicola. By all 

other features this taxon is referable to A. subconspersa (because of veil and 

ground colour). 

Accepted name: Alnicola silvae-novae (D.A. Reid) Courtec. (if different 

from A. subconspersa). 

145

sobria 

Status: valid, not typified. Original material : not located. Host: undefined. 

Basionym: Agaricus furfuraceus β sobrius Fr., Obs. mycol. 2: 25 (1818). 

≡ Agaricus sobrius (Fr.) Fr., Epicr.: 200 (1838). 

≡ Naucoria sobria (Fr.) P. Kumm., Führer Pilzk.: 77 (1871). 

≡ Alnicola sobria (Fr.) Kühner ex Konrad & Maublanc, Agaricales: 165 (1948). 

Notes: this uncertain taxon is not described precisely enough by Fries 

(1818: 25, 1836-1838: 200, 1874: 263), Gillet (1876: 541, pl. 368) or Konrad 

and Maublanc (1948) to allow for a reliable interpretation. It might belong to 

A. amarescens group, but is better being abandoned. 

Interpretation: N. sobria ss. Cooke = Conocybe vestita fide Kühner 

(1935: 157). 

spadicea 

Status: valid. Holotype (visited): GB, Monks Woods, Hunts., 28-VII-1965, leg. P.J. 

Houlton (K). Host: undefined. 

Basionym: Naucoria spadicea D.A. Reid, Trans. Brit. mycol. Soc. 82 (2): 206 (1984). 

≡ Alnicola spadicea (D.A. Reid) Bon, Doc. mycol. 20 (78): 40 (1990). 

≡ Alnicola spadicea (D.A. Reid) Bon, Doc. mycol. 21 (83): 37 (1990) (comb. superfl.). 

≡ Naucoria salicis var. spadicea (D.A. Reid) Ludwig, Pilzkomp. I: 407 (2001). 

Notes: published a few months before N. saliceti P D. Orton, both names 

based on Naucoria macrospora f. tetraspora J.E. Lange (Lange, 1939, 1940). 

However it is not clear whether the holotype designed by Reid (1984) matches 

Orton’s (1984) salicicolous taxon, here reported to A. macrospora (see under 

SALICETI). 

Accepted name: Alnicola spadicea (D.A. Reid) Bon (if different from 

A. macrospora J.E. Lange → J. Favre). 

sphagneti 

Status: valid. Holotype (visited): UK, Loch-an-Eilean, Rothiemurchus, Inverness-Shire, 

27-IX-1955 (K). Isotype (visited by E. Horak) at E. Host: undefined. 

Basionym: Naucoria sphagneti P.D. Orton, Trans. brit. mycol. Soc. 43 (2): 320 (1960). 

≡ Alnicola sphagneti (P.D. Orton) Romagn., Bull. Fed. mycol. Dauphiné-Savoie 74: 19 

(1979). 

Notes: Orton (1960) reported A. sphagneti to Agaricus conspersus b 

uliginosus Fr.: Fr. (Fries, 1821: 261), a shortly described taxon which is 

difficult to interpret. Orton does not provide specific information about the 

host-tree; the presence of Sphagnum can only help to interpret N. sphagneti as 

a acidophilic, but certainly not bryotrophic species. Moser (1991 in Moser and 

Jülich, 1985-2003, Agaricales III Leif. 9, only left picture) report under 

"Naucoria sphagneti" a very singular taxon, which could match Orton’s 

description. This possibly septentrional taxon of uncertain affinities requires a 

complete revision. 

Accepted name: Alnicola sphagneti (P.D. Orton) Romagn. 

146


striatula 

Status: valid. Holotype (visited): UK, Kingthorpe, Yorks, 14-IX-1959 (Orton n° 1943, 

K). Host: Alnus sp. 

Basionym: Naucoria striatula P.D. Orton, Trans. brit. mycol. Soc. 43 (2): 322 (1960). 

≡ Alnicola striatula (P.D. Orton) Romagn., Bull. Fed. mycol. Dauphiné-Savoie 74: 18- 

19 (1979). 

Notes: in the original sense of Orton (1960), this is a pale species devoid 

of reddish tones. Reid (1984) and Bon (1992) interpret it in a wider sense 

which virtually includes A. umbrina (Moreau, 2004). It might represent only a 

pale variant of A. umbrina with Atlantic distribution. 

Accepted name: Alnicola striatula (P.D. Orton) Romagn. (if different 

from A. umbrina (R. Maire) Singer). 

suavis 

Status: valid, not typified. Original material : only 1 colour plate, published by 

Bresadola (1884 pl. 59). Other material identified by Bresadola (visited by E. Horak): IX-1882, 

leg. G. Bresadola (S): Host: Alnus sp. (probably A. incana). 

Basionym: Naucoria suavis Bres., Fung. trident. I (4/5): 53 (1884). 

≡ Alnicola suavis (Bres.) Kühner, Bull. Soc. mycol. France 47 (3-4): 242 (1931). 

Notes: taxon not critical, but rare and possibly confined to Alnus incana. 

E. Horak’s notes on Bresadola’s material are conform to the literature (Kühner, 

1931: 238; Horak, 1963: 78, pl. 4 fig. 24; Kränzlin and Breitenbach, 1997: 

274). 

Accepted name: Alnicola suavis (Bres.) Kühner. 

subconspersa Kühner 

Status: invalid, art. 36.1. Original material in R. Kühner (G) and H. Romagnesi (PC) 

collections, visited. Host: Alnus glutinosa. 

Original name: Naucoria subconspersa Kühner in Kühner and Romagn., Fl. Anal. 

Champ. Sup.: 237 (1953) (inval., art. 36.1; sp. nov. for Naucoria conspersa ss. Lange 1939, 

fig. 125G) 

= Alnicola subconspersa (Kühner) M.M. Moser in Gams, Kleine Kryptogamenfl. IIb(2) 

(1955) (basion. inval.). 

= Alnicola subconspersa (Kühner) Romagn., Bull. Fed. mycol. Dauphiné-Savoie 74: 19 

(1979) (basion. inval.). 

Notes: first considered by Kühner (1942: 2) as a synonym of 

A. luteolofibrillosa, A. subconspersa is separated and published as a species 

nova ("nomen novum") (Kühner and Romagnesi, 1953) but not validated later 

(Kühner and Romagnesi, 1957). According to all studied material, 

A. subconspersa ss. Kühner represents lowland collections of 

A. luteolofibrillosa with longer spores and well-developed ring-like zone on the 

stipe. Orton (1960) used this name for another, very distinct species, probably 

closer from Lange’s plate (1939: pl. 125G). 

Accepted name: Alnicola luteolofibrillosa Kühner. 

147

subconspersa P.D. Orton 

Status: valid. Holotype (visited): UK, Badger Falls, Glen Affric, Inverness-Shire, 20- 

VIII-1955 (K). Host: Alnus sp. (probably A. glutinosa). 

Basionym: Naucoria subconspersa P.D. Orton, Trans. Brit. mycol. Soc. 43: 323 (1960). 

≡ Alnicola subconspersa (P.D. Orton) Bon, Doc. mycol. 9 (35): 41 (1979). 

Notes: new name for Naucoria conspersa ss. J.E. Lange (1939) and, 

according to Orton (1960: 324), based on Naucoria subconspersa Kühner 

(1953: 237, inval.) but typified on British material and not in conformity with 

Kühner’s intentions (see above). Nevertheless Orton’s taxon seems to agree 

with Lange’s plate and short description. Macroscopical description and 

variation range need to be completed for this apparently common and possibly 

early-fruiting taxon. 

Accepted name: Alnicola subconspersa (P.D. Orton) Bon. 

submelinoides 

Status: valid, not typified. Original material in R. Kühner collections (G), visited. Host: 

Alnus viridis. 

Basionym: Alnicola submelinoides Kühner, Botaniste 17 (1-4): 175 (1926) 

("submelinoïdes"). 

≡ Naucoria submelinoides (Kühner) R. Maire, Bull. Soc. mycol. France 47: 223 (1930). 

≡ Alnicola submelinoides (Kühner) Bon (as "(Kühner ex Maire) Bon"), Doc. mycol. 9 

(35): 41 (1979) (comb. inval., art. 33.6a). 

≡ Hebeloma submelinoides (Kühner) Kühner, Hyménom. agaricoïdes: 235, 898 (1980). 

Notes: Accepted as validly published in 1926, by a short line in a key. 

Detailed descriptions in Kühner (1931: 242; 1942: 5). 

Accepted name: Alnicola submelinoides Kühner. 

tantilla 

Status: valid. Holotype (visited): CH, Val Nüglia, 2550 m, 15-VIII-1950, herb. J. Favre 

n° GK13461 (G). Host: Salix herbacea. 

Basionym: Naucoria tantilla J. Favre, Ergebn. wiss. Unters. schweiz. Natl Parkes 5: 202 

(1955). 

≡ Alnicola tantilla (J. Favre) Romagn., Bull. Fed. mycol. Dauphiné-Savoie 74: 19 

(1979). 

≡ Alnicola tantilla (J. Favre) Kühner, Trav. sci. Parc natl Vanoise 11: 124 (1981) 

(comb. superfl.). 

≡ Alnicola tantilla (J. Favre) G. Gulden, Arct. Alp. Fungi 2: 43 (1988) (comb. superfl.). 

Notes: a very widespread, arctico-alpine species perfectly described by 

Kühner (1981) as "Alnicola chamitae" (Moreau, unpublished). Collections 

from Salix repens (Reid, 1984: 212; Orton, 1984: 603; De Haan, 1999) seem to 

differ mainly in spore shape, slightly depressed above apiculus. 

Accepted name: Alnicola tantilla (J. Favre) Romagn. 

148


tetraspora 

Status: invalid, art. 36.1. Original material: only Lange (1939) pl. 125D. Host: Salix 

cinerea. 

Original name: Naucoria macrospora f. tetraspora J.E. Lange, Dansk Bot. Ark. 9 (6): 

21 (1938) (inval.). 

= Naucoria langei f. tetraspora (J.E. Lange) Kühner in Kühner and Romagnesi, Bull. 

Soc. Natur. Oyonnax 10-11: 9 (1957) (basion. inval.). 

= Naucoria alnetorum f. tetraspora (J.E. Lange) G. Keller & M. Moser, Biosystem. 

Ecol. 19: 176 (2001) (basion. inval.). 

Notes: renamed "Naucoria spadicea" by Reid (1984) and "Naucoria 

saliceti" by Orton (1984). Lange’s description agrees with A. salicis, but 

Kühner’s (1957: 9) description with pear-shaped superficial articles in 

pileipellis can also be refered to A. saliceti P.D. Orton, if proved to be distinct 

from A. salicis. 

Accepted name: Alnicola macrospora J.E. Lange → J. Favre (see 

Moreau and Deïana, 2004). 

umbrina R. Maire 

Status: valid. Lectotype (Moreau, 2004): Algeria, Azazga, 20-XII-1913. Herbier 

Champignons d’Afrique du Nord n° 2030, R. Maire (MPU). Host: Alnus glutinosa (handwriten 

on unpublished notes of R. Maire, MPU). 

Basionym: Tubaria umbrina Maire, Bull. Soc. mycol. France 44 (1): 48 (1928). 

≡ Naucoria umbrina (Maire) Maire, Bull. Soc Mycol. France 46: 225 (1930) (illegit., 

non N. umbrina Bres.). 

≡ Alnicola umbrina (Maire) Kühner, Bull. Soc. mycol. France 47: 241 (1931). 

Notes: the combination of this taxon in the genus Alnicola is sometimes 

attributed to Singer (Singer, 1947: 246, 1949: 540, etc.). An earlier 

combination by Kühner (1931: 241) has been found, formally written in the 

legend of fig. B (art. 33.1) with explicit reference to the basionym p. 240. 

Alnicola umbrina has been revised by Moreau (2004). 

Accepted name: Alnicola umbrina (R. Maire) Kühner. 

umbrina Singer 

Status: valid. Holotype: (not revised): CHILE, Salto de Laja near San Antonio, 2-IV- 

1967 (R. Singer n° M 6540, BAFC). Host: Alnus sp. (plantation). 

Basionym: Alnicola scolecina var. umbrina Singer, Sydowia 30 (1-6): 276 (1978 

[1977]). 

Notes: it is unclear what Singer (1977: 202) had in mind when he created 

this name, almost homonym and possibly synonym of A. umbrina (Maire) 

Kühner. He had formerly used the latter name for Catalan (Singer, 1947: 246) 

and Caucasian (Singer, 1930: 98; 1939: 68) collections, but later doubted of 

Maire’s (1928) taxon and choosed to rename it (Singer, 1977). However, so 

few is known about South American species of Alnicola that this taxon needs a 

149

deep reevaluation before possibly being renamed at a specific rank. Singer’s 

taxon is certainly close to Maire’s Tubaria umbrina (Moreau, 2004). 

Accepted name: to be precised. 

velutina 

Status: valid. Holotype (visited): USA, Connecticut, Redding, 27-VIII-1902 

(F.S. Earle n° 1246, NYC). 

Basionym: Naucoria velutina Murrill, North Amer. Fl. 10 (3): 177 (1917). Host: 

undefined (mosses in a swamp). 

≡ Alnicola velutina (Murrill) Singer, Sydowia 15: 71 (1962). 

Notes: Singer does not explicit his combination, probably based on a 

revision of the type; some scarce descriptive elements are in Singer (1977: 

209). According to type revision this species is closely related to A. badia. 

Accepted name: Alnicola velutina (Murrill) Singer. 

zonata 

Status: valid. Holotype (visited): Sweden, Medelpad, near Borgsjö, 4-9-1997 

(Berlin-Dahlem). Host: undefined (not Alnus). 

Basionym: Naucoria zonata E. Ludwig & Reil in Ludwig, 2001, Pilzkompendium 1: 

422. 

Notes: this extremely distinct species is isolated amongst European taxa 

by its small spores and cystidia, but seems to have relatives in North America. 

More macroscopical informations are required for this species, so far only 

known from the type collection (Moreau and Garcia, 2005). Also compare with 

A. brevicystis Métrod, inval., with distinctly smaller spores. 

Accepted name: Alnicola zonata (E. Ludwig & Reil) P.-A. Moreau, 

comb. nov. 

Acknowledgements 

This revision has been made possible thanks to the kind collaboration of the curators 

and mycologists of the following institutions, who could provide informations on collections 

and kindly allowed me to borrow many type collections : BD (Dr Burghard Hein, Berlin), G 

(Dr Philippe Clerc, Geneva), IB (Pr Ursula Peintner, Innsbrück), K (Dr Brian Spooner, Kew), 

L (Dr Machiel E. Noordeloos), LAU (Pr H. Clémençon, Lausanne), LIP (Pr Régis 

Courtecuisse, Lille), M (Dr Dietmar Triebel, Münich), MPU (Dr Joël Mathez, Mrs Marie- 

Josèphe Mauruc, Montpellier), NYC (Dr Ellen Bloch), PC (Dr Bart Buyck, Mr Bernard 

Duhem, Paris) and PRM (Dr Jan Holec, Praha). 

I am especially grateful to Pr E. Horak (Zürich), who has accepted to lend me his 

precious notes on many collections, especially on the type material he studied in many herbaria 

during his long career at ETH, and for his valuable comments and suggestions all along my 

work. I also thank all the mycologists who provided me information, bibliography and 

additional collections : Pr Denise Dailly-Lamoure (France), Dr Anne-Marie Fiore (Geneva), Dr 

Béatrice Senn-Irlet (Switzerland), Dr Else C. Vellinga (USA), Pr Joe F. Ammirati (USA), 

François Ayer (Suisse), Miroslav Beran (Czech Republic), André Bidaud (France), Dr Ivano 

150


Brunner (France), Dr Gabriele Cacciali (Italy), Emanuele Campo (Italy), Pierre Collin (Suisse), 

Pr Giovanni Consiglio (Italy), Jean-Marie Cugnot (France), Jean-Claude Deïana (France), 

Yves Deneyer (Belgium), Laurent Deparis (France), Roberto Fernández Sasia (Spain), Francis 

Fouchier (France), Guy Garcia (France), Jacques Guinberteau (France), Anton Hausknecht 

(Austria), Pascal Hériveau (France), Dr Brandon P. Matheny (USA), Paul Pirot (Belgium), 

Gregor Podgornik (Slovenija), Pr Jean-Philippe Rioult (France), Pierre Roux (France), Pr Scott 

A. Redhead (Canada), Jean-Pierre Vidonne (France), Ruben Walleyn (Belgium) and Jean- 

Jacques Wuilbaut (Belgium). 

I warmly thank Jacques Melot (Reykjavik, Iceland) for having provided me with rare 

documents, copies of Fries' unpublished plates preserved in Uppsala (S) and for his precious 

help in nomenclatural problems, Pr Ursula Peintner (IB, Austria) for having provided 

important information from Moser’s archives and literature and for having accepted to revise 

critically this paper, and Ana and Olivier Daillant (Belgium) for having kindly revised English. 

This paper is dedicated to the memory of Guy Redeuilh (ex-president of the Société 

Mycologique de France), who spent a long time revising all nomenclatural points treated in this 

article, and passed away shortly after its submission, in October 2004. 

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(Received 20 November 2004; accepted 15 September 2005) 

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