Inocybe praetervisa group - A clade of 
four closely related species with partly 
different geographical distribution 
ranges in Europe
Authors: Ellen Larsson, Jukka Vauras, and Cathy L. 
Cripps 
NOTICE: this is the author’s version of a work that was accepted for publication in Mycoscience. 
Changes resulting from the publishing process, such as peer review, editing, corrections, 
structural formatting, and other quality control mechanisms may not be reflected in this 
document. Changes may have been made to this work since it was submitted for publication. A 
definitive version was subsequently published in Mycoscience, vol. 59, iss. 4, (July 2018) 
DOI# 10.1016/j.myc.2017.11.002.
Larsson, Ellen, Jukka Vauras, and Cathy L. Cripps. "Inocybe praetervisa group - A clade of four 
closely related species with partly different geographical distribution ranges in Europe." 
Mycoscience 59, no. 4 (July 2018): 277-287. DOI: 10.1016/j.myc.2017.11.002.
Made available through Montana State University’s ScholarWorks 
scholarworks.montana.edu 
Inocybe praetervisa group e A clade of four closely related species with
partly different geographical distribution ranges in Europe
Ellen Larsson a, b, *, Jukka Vauras c, Cathy L. Cripps d
a University of Gothenburg, Department of Biological and Environmental Sciences, Box 461, SE-40530, G€oteborg, Sweden
b Gothenburg Global Biodiversity Centre, Box 461, SE-40530, G€oteborg, Sweden
c Biological Collections of Åbo Akademi University, Herbarium, Biodiversity Unit, FI-20014, University of Turku, Finland
d Montana State University, Department of Plant Sciences and Plant Pathology, 119 Plant Biosciences Building, Bozeman, MT, 59717, USAbroad
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Sequence data from a
clade is comprised o
by having nodulose
praetervisa occurs in
from Northern Europ
associated with Betul
confirmed as a later
Europe, mainly restr
arctica is here describ
Dryas octopetala, Sali
an intercontinental d
occurrence of one ad
group is provided.geographical region and different habitats show that the Inocybe praetervisa
losely related species. These species of section Marginatae are characterized
and a stipe that is abundantly pruinose only in the upper half. Inocybe
rn Europe in mountainous mixed coniferous forests, and is not confirmed
ybe rivularis occurs in northern boreal forests up to the lower alpine zone,
ist habitats, and is not confirmed from Southern Europe. Inocybe taxocystis is
ym of I. favrei. The species has a wide geographical distribution range in
the alpine zone and moist soils, associated with Salix herbacea. Inocybe
new species. It occurs in the arctic and higher alpine zones, associated with
s, S. reticulata and S. herbacea. All species except I. arctica are shown to have
ion range and are confirmed from North America. Sequence data suggest the
l species in the alpine zone of China. A key to the species in the I. praetervisa1. Introduction
Inocybe section Marginatae Kühner (1933) is known to be
comprised of several species complexes where names have been
interpreted differently and often misapplied (Esteve-Raventos,
Moreno, Bizio, & Alvarado, 2015, 2016; Jacobsson & Larsson,
2012; Vauras & Larsson, 2015).
The species in Marginatae are usually characterized by the
absence of a cortina, having nodulose spores and caulocystidia
descending to the base. However species in the I. praetervisa clade,
as defined by Esteve-Raventos et al. (2016), have abundant caulo-
cystidia in the upper half that are sparse or absent in lower half of
the stipe. In this recent paper dealing with the taxonomy of the
Inocybe praetervisa group (Esteve-Raventos et al., 2016) a lectotype
for I. praetervisa Quel. was selected. In addition, an epitype was
designated, from a representative specimen of the Bresadolaherbarium collected from the type locality in Italy in order to fix the
name also by sequence data of the internal transcribes spacer (ITS)
region (Ariyawansa et al., 2014).
Inmorphology, I. praetervisa is characterized by having a smooth
yellowish-brown pileus that becomes darker brownwith age and a
margin that is rimose. The lamellae are at first pale white with a
greyish tint, and are later more brown. The stipe is pale whitish at
first and later yellowish brown, possesses a more or less marginate
bulbous stipe base, and is densely pruinose in the upper half. Spores
are heterodiametric and distinctly nodulose. Cheilo- and pleuro-
cystidia are utriform to broadly fusiform, and rather thick-walled
with crystals at the apex. Specimens become darker brown upon
drying but are not blackening. The selected epitype, collected by
Bresadola, originates from a coniferous forest. For a more detailed
description see Esteve-Raventos et al. (2016).
In the Esteve-Raventos et al. (2016) study, phylogenetic analyses
based on ITS sequences were performed to infer phylogenetic re-
lationships among the species. In the phylogeny presented, the
sequence of a paratype of I. rivularis Jacobsson & Vauras fell within
the I. praetervisa clade with strong support, suggesting it should be
a later synonym of I. praetervisa. Accordingly, based on the
phylogenetic results and the similarity in morphology I. rivularis
was regarded as a synonym of I. praetervisa (Esteve-Raventos et al.,
2016).
Inocybe rivularis was described by Jacobsson and Vauras (1989)
from moist forest habitats in the boreal zones of Fennoscandia,
likely associated with Betula spp., Picea abies (L.) H. Karst. and Salix
spp. The species is considered to have a rather wide distribution
and is common in suitable habitats, usually close to small spring
brooks. In morphology it is rather similar to the description of
I. praetervisa (Esteve-Raventos et al., 2016) but was characterized as
having a cortina in young specimens that soon disappears leaving
no remnants on the stipe. The stipe base is clavate to bulbous but
not marginately bulbous. As the species was densely pruinose only
in the upper part of the stipe it was at first thought to belong in
section Cortinatae Kühner.
A species with similar morphology is I. taxocystis (J. Favre)
Singer. The species was first described by Favre (1955) as a variety
of I. decipientoides Peck (I. decipientoides var. taxocystis Favre), from
the Swiss alpine zone associated with Salix herbacea L. and Dryas
octopetala L. Favre's variety was characterized by having a stipe that
is pruinose at least to half the stipe, glabrous below, with a bulbous
but not marginate stipe base. It was first raised to species level by
Singer (1986), but without selecting a lectotype. It was also
described as the new species Asterosporina taxocystis J. Favre & E.
Horak by Horak (1987), and later moved to Inocybe (Fr.) Fr. by Senn-
Irlet (1992).
Another species with similar morphology and habitat was
published as I. favrei Bon (1985). The species was described with a
stipe that is more or less pruinose particularly in the upper half and
with an obtusely marginate stipe base, and is associated with
S. herbacea, S. reticulata L. and Dryas. The species was described by
Bon to accommodate the description of the species I. decipiens Bres.
sensu Kühner by Favre from the alpine zone (Favre, 1955). Bon
designated a holotype (Bon 84095) from the alpine zone in France
associated with S. herbacea.
Analyses of ITS sequence data have become an important tool
for evaluating the morphological characters used to determine and
diagnose species. This has been especially valuable for groups with
morphologically similar characters and seemingly broad ecology as
often encountered within Inocybe (Cripps, Larsson, & Horak, 2010;
Kokkonen & Vauras, 2012; Larsson, Vauras, & Cripps, 2014; Vauras
& Larsson, 2015). The ITS region is now agreed upon as the bar code
marker for fungi (Schoch et al., 2012) and has been found to be
suitable and to work well for delimitation and segregation of spe-
cies in the majority of the Basidiomycetes (K~oljalg et al., 2013).
In the present study, we compare morphology and sequence
data of a broader sampling of specimens determined as I. rivularis,
I. taxocystis and I. favrei, to get a better picture of I. praetervisa sensu
Esteve-Raventos et al. (2016).
2. Material and methods
2.1. Morphological methods
The majority of the sequenced specimens were collected by the
authors. Additional material received as loans from different
herbaria were studied (AH, G, LIP, O, TUR, S, UPS). Abbreviation of
herbaria follows Index Herbariorum (http://sweetgum.nybg.org/
science/ih/).
Macro-morphological characters were noted and cross-sections
drawn from some fresh basidiomata. The color codes refer to
Cailleux (1981). Micro-morphological characters were measured
and drawn from dried material mounted in 10% NH4OH solution
following the methodology described in Vauras and Kokkonen
(2009).Photos of micro-morphological characters were made using an
Axioskop 2 (Zeiss, Oberkochen, Germany) light microscope and the
AxioVision software (http://www.zeiss.com/microscopy/int/home.
html).
2.2. Molecular methods
Sixty-three specimens from Northern Europe, Svalbard, and
North Americawere targeted for sequencing in this study, including
the holotype of I. favrei (Bon 84095, LIP!). In addition the ITS
sequence of the lectotype of I. taxocystis (JN580884) and paratype
of I. rivularis (JN580885) from the study by Kokkonen and Vauras
(2012), the epitype of I. praetervisa (KT201675), I. tabacina Furrer-
Ziogas and I. phaeosticta Furrer-Ziogas from the study by Esteve-
Raventos et al. (2016) were included. Also a few additional ITS
and LSU sequences of Inocybe section Marginatae published in
Ryberg, Larsson, and Jacobsson (2010) were included. For compar-
ison and rooting of trees the sampling was selected to correspond
with Clade B in Esteve-Raventos et al. (2016). Sequence data of the
ITS and LSU regions of I. fibrosoides Kühner (EL51-14; Genbank
KY033846) and I. decemgibbosa (Kühner) Vauras (JV8115; Genbank
KY033847) were generated. Each target ITS sequence type was also
blasted in GenBank (Clark, Karsch-Mizrachi, Ostell, & Sayers, 2016)
and the UNITE database (K~oljalg et al., 2013) to search for additional
data generated from specimens, environmental samples as soil and
ectomycorrhizae. In all 13 sequences were found and added to the
data set.
Sequences from the complete internal transcribed spacer (ITS)
region and about 1400 or 375 base pairs (bp) of the 50end of the
large subunit (LSU) of the nuclear ribosomal DNA were generated.
DNA extractions were performed using DNeasy Plant Mini kit
(Qiagen, Hilden, Germany), PCR reactions using Illustra PuReTaq™
ReadyToGo PCR beads (GE Healthcare, Buckinghamshire, United
Kingdom). For PCR clean up QIAquick PCR purification kit (Qiagen,
Hilden, Germany) was used. The DNA from the type specimen was
extracted using a CTAB method, and primers for PCR followed
protocols described in Larsson and Jacobsson (2004). Primers used
to amplify the complete ITS region and the 50end of the LSU region
were ITS1F (Gardes& Bruns,1993) and LR21, LR0R, and LR7 (Hopple
& Vilgalys, 1999). Primers used for sequencing were ITS1, ITS4
(White, Bruns, Lee, & Taylor, 1990), Ctb6 (https://nature.berkeley.
edu/brunslab/), LR3R and Lr5 (Hopple & Vilgalys, 1999).
Sequences were edited and assembled using Sequencher 5.1
(Gene Codes, Ann Arbor, Michigan). Sequences generated for this
study have been deposited in GenBank and accession numbers
given (KY033785eKY033847). Specimens sequenced in this study
are indicated with an asterix (*) and the GenBank number is noted
in the list of specimens examined.
Alignment was performed using the L-INS-i strategy as imple-
mented in MAFFT v. 7.017 (Katoh & Standley, 2013). The alignment
was adjusted using Aliview 1.17.1 (Larsson, 2014). For inferring
phylogenetic relationships among species heuristic searches for the
most parsimonious trees were performed using PAUP* (Swofford,
2003). All transformations were considered unordered and
equally weighted. Variable regions with ambiguous alignment
were excluded and gaps were treated as missing data. Heuristic
searches with 1000 random-addition sequence replicates and TBR
branch swapping were performed, saving at most 25 trees in each
replicate. Relative robustness of clades was assessed by the boot-
strap method using 1000 heuristic search replicates with 100
random taxon addition sequence replicates and TBR branch
swapping, the latter saving at most 100 trees in each replicate. In
addition a Bayesian analysis was carried out in MrBayes 3.2.6
(Ronquist&Heulsenbeck, 2012), with a best-fit model of nucleotide
evolution supplied by MrModeltest 2.2 (Nylander, 2004). Eight
default-setting Metropolis-Coupled Markov Chain Monte Carlo
(MCMCMC) chains were run for 10 million generations with trees
sampled every 5000 generations and an initial burn-in of 1000
trees. After discarding the trees prior to the burn-in threshold a 50%
majority-rule consensus phylogram was computed from the
remaining trees.
3. Results
The aligned complete dataset consisted of 95 sequences and
2095 characters. After exclusion of ambiguous regions mainly from
the beginning and the end of the data set 1977 characters remained
for the analysis. Of these, 1670 were constant, 57 were variable but
parsimony uninformative, and 250 (12.6%) were parsimony infor-
mative. The maximum parsimony analysis yielded 18,950 equally
most parsimonious trees (length ¼ 460 steps, CI ¼ 0.8217, and
RI ¼ 0.9359). The 50% majority-rule consensus bootstrap tree is
presented in Fig. 1. The bootstrap analysis recovered the
I. praetervisa complex with 100% support. Five supported clades
with low to strong bootstrap support were found within the
complex corresponding to I. praetervisa (91%), I. favrei (95%),
I. rivularis (54%), Inocybe sp. (100%), I. arctica (see taxonomy section,
83%). The four last clades form a subclade within the complex with
a support value of 71%.
As suggested by MrModeltest, the nucleotide evolution model
HKY þ G was used for the ITS1 spacer; K80 was used for the 5.8S
gene; HKYþ1was used for the ITS2 spacer, and GTRþ I was used for
the LSU in the Bayesian analysis. The MCMC analysis converged
well in advance of the burn-in threshold and chain mixing was
found to be satisfactory, as assessed by using Tracer v1.5
(Drummond, Suchard, Xie, & Rambaut, 2012). Also in the Bayesian
analysis, the I. praetervisa complex was recovered as monophyletic
with strong support (a Bayesian posterior probability (BPP) of 1.00).
The Bayesian tree topology is similar to the MP bootstrap tree. The
same major clades supported in the bootstrap analysis were also
recovered in the Bayesian analysis, and the I. rivularis clade, with
only 54% bootstrap support, received no support. BPP values are
indicated on the corresponding branches in Fig. 1.
4. Taxonomy
Inocybe arctica E. Larss., Vauras & C.L. Cripps, sp. nov. Figs. 2,
3AeF, 4A, B, 5AeC.
MycoBank no.: MB 820654.
Diagnosis: Medium sized to fairly small species with brown to
dark brown pileus, often with frosty remnants of velum, stipe
abundantly pruinose at apex but caulocystidia less frequent to rare
at the base, spores nodulose, (10.3e)11.0e13.1(e14.0)  (6.5e)
6.8e8.7(e9.2) mm, pleurocystidia (50e)56e82(e88)  (11e)
13e22(e28) mm. Closely related to I. rivularis and I. favrei but differs
from these by having on average longer and more narrowly
ventricose pleurocystidia and slightly larger spores, a distribution
range in the arctic and higher alpine zones and with preferences for
more rich and calcareous soils, associatedwithDryas octopetala and
dwarf Salix spp., and by having a different ITS sequence.
Type: NORWAY, Troms, Parasdalen, fjeld Paras, SW slope, alpine
zone, in a snowbed, among Salix polaris Wahlenb. and Bistorta
vivipara (L.) Gray, 69050N, 20100E, 12. VIII. 1986, Jukka Vauras
2238F, (Holotype TUR-A 147440, Isotypes GB, MONT, WTU, Gen-
Bank no. KY033843).
Etymology: Refers to the area of distribution of the species.
Pileus 1e3.5 cm, when young conical to plano-convex, later
more applanate, flat to broadly or indistinctly umbonate, margin
deflexed, often somewhat undulate; dirty brown with yellowishtinge, or dark brown, fibrils often dark brown, subtomentose-
smooth around disc, outwards radially fibrillose-felty, often with
frosty ormore abundant whitish velipellis, sometimes cracking into
squares. Lamellae moderately crowded, up to 6 mm broad,
narrowly adnate to adnexed, initially pale yellowish (71K), then
pale greyish brown with yellow tinge (70L), later yellowish brown
(69N). Stipe 1e3.5  0.3e0.7 cm, often equal, base bulbous but not
marginate, yellowish, apex often with reddish tint, base whitish,
distinct pruinose at apex and in the upper half, but few in the lower
part, longitudinally striate, silky shiny. Cortina fugacious, scarce,
whitish, present in young basidiomata. Fleshwhitish to pale brown,
in stipe also pale yellowish brown, sometimes with reddish tint at
apex, shiny. Smell indistinct to slightly spermatic. Spores nodulose,
heterodiametric, fairly dark yellow-brown, (10.3e)
11.0e13.1(e14.0)  (6.5e)6.8e8.7(e9.2) mm, on average
12.0  7.6 mm, range of mean values 11.5e12.2  7.2e7.9 mm, Q ¼
(1.35e)1.4e1.8(e1.9), on average 1.57, range of mean Q-values
1.49e1.69 (120 spores from 6 collections). Basidia (29e)
30e43(e44) 11e14(e16) mm, on average 35 13 mm, generally 4-
spored (47 basidia from 6 collections). Pleurocystidia with crystals
(50e)56e82(e88)  (11e)13e22(e28) mm, on average 70 17 mm,
range of mean values 65e77  14e20 mm (100 pleurocystidia from
6 collections), cylindrical to narrowly utriform, with up to 4 mm
thick, pale yellow wall. Cheilocystidia with crystals
44e72  13e26 mm, on average 59  19 mm, (n ¼ 26), abundant,
paracystidia 10e32  9e13 mm (n¼ 11). Caulocystidia at stipe apex
thick-walled, abundant, often with crystals (30e)
38e62(e65)  (12e)14e21(e22) mm, on average 49  18 mm,
(n ¼ 21), more rare but present to stipe base. Cauloparacystidia at
stipe apex 17e38  11e16 mm (n ¼ 8), few of them thick-walled.
Habitat: Growing in small groups but also single, mainly asso-
ciated with Salix polaris, S. reticulata L., Dryas octopetala but also
Bistorta vivipara and S. herbacea. Usually found on somewhat more
rich and calcareous soils. Once found in a mountain birch forest
close to the treeline.
Distribution: So far known from the arctic and higher alpine
regions on Svalbard and Fennoscandia.
Specimens examined: FINLAND, Enonteki€on Lappi, Enonteki€o,
Kilpisj€arvi, Pikku-Malla, forest with Betula pubescens Ehrh. subsp.
czerepanovii (N.I. Orlova) H€amet-Ahti, Betula nana L., and Salix spp.
near timberline, herb-rich site, by path, 19 Aug 2004, J. Vauras
21746* (TUR-A; GenBank KY033844). NORWAY, Svalbard and Jan
Mayen, Svalbard, Nordenski€old Land, Adventsdalen, Endalen, moist
with Salix herbacea and Bistorta vivipara, 10 Aug 2015, E. Larsson 56-
15* (GB; GenBank KY033827); 14 Aug 2009, with Salix polaris, E.
Larsson 77-09* (GB; TUR-A; GenBank KY033828); 14 Aug 2009, in
matt of Dryas octopetala, E. Larsson 80-09* (GB; GenBank
KY033832); 14 Aug 2009, G. Gulden 181/09* (O; GenBank
KY033833); Svalbard, Nordenski€old Land, Adventsdalen, Todalen,
with S. polaris and Dryas octopetala, 21 Aug 2002, leg. C. Cripps,
CLC1923* (MONT; GenBank KY033826); 21 Aug 2002, leg. C. Cripps,
CLC1925 (MONT); Svalbard, Nordenski€old Land, Kapp Linne, 8 Sep
2009, G. Gulden 128/09* (GB; GenBank KY033833); Svalbard,
Nordenski€old Land, Longyearbyen, close to the old churchyard,
with Dryas octopetala and Salix polaris, 7 Aug 2015, E. Larsson 18-
15* (GB; TUR-A; GenBank KY033830); 7 Aug 2015, with Dryas
octopetala and Salix polaris, E. Larsson 16-15* (GB; GenBank
KY033836); Svalbard, Nordenski€old Land, Longyearbyen, with
Dryas octopetala and Salix polaris, 8 Aug 2009, E. Larsson 34-09*
(GB; TUR-A; GenBank KY033837); 12 Aug 2009, with Dryas octo-
petala, Salix polaris and Bistorta vivipara, E. Larsson 50-09* (GB;
GenBank KY033838); 8 Aug 2015, moist area with Dryas octopetala
and Salix polaris, E. Larsson 30-15 (O; NOBAS2288, KY033834);
Svalbard, Nordenski€old Land, Longyearbyen, Nybyen, with Dryas
octopetala and Salix polaris, 17 Aug 2009, E. Larsson 123-09* (GB;
Fig. 1. The 50% majority-rule bootstrap consensus tree from the parsimony analysis. Parsimony bootstrap values and Bayesian posterior probabilities are indicated on branches.
Clades discussed in the text are indicated with bars and species epithets. Sequences originating from type specimens are marked. Clade B refers to Esteve-Raventos et al. (2016).GenBank KY033831); Svalbard, Nordenski€old Land, Longyearbyen.
2002, B. Gahne 02-96 (GB; GenBank AM882980); Svalbard, Nor-
denski€old Land, Bolterdalen, moist with Salix polaris, 13 Aug 2015,
A. Molia 25-2015 (O; NOBAS2299, KY033835). SWEDEN, Torne
Lappmark, Jukkasj€arvi, Latnajaure, moist with Salix reticulata, 10
Aug 2007, E. Larsson 72-07* (GB; TUR-A; GenBank KY033839); 7Aug 2006, with Salix polaris, E. Larsson 71-06* (GB; TUR-A; Gen-
Bank KY033842); Lule lappmark, Jokkmokk, Padjelanta NP,
Vielgggisbakte on NW side of Vastenjaure Lake, near Salix herbacea
and S. reticulata, 12 Aug 2016, J. Vauras 31471* (TUR-A; GB; Gen-
Bank KY033840); Lule lappmark, Jokkmokk, Padjelanta NP, Arra-
noaijvve on N side of the Vastenjaure Lake, near Salix herbacea, 11
Fig. 2. Inocybe arctica. Basidiomata of the holotype growing with Salix polaris and Bistorta vivipara (J. Vauras 2238F).Aug 2016, J. Vauras 31460* (TUR-A, GB; GenBank KY033841); Lule
lappmark, Jukkasj€arvi, Padjelanta NP, Slahpejaure NO, with Salix
herbacea, 14 Aug 2016, E. Larsson 117-16* (GB; GenBank KY033845).
Inocybe praetervisa Quel., in Bresadola, Fung. Trident. 1:35,
1881. Fig. 4E, F.
For the description of I. praetervisa, we refer to Esteve-Raventos
et al. (2016). The species resembles I. rivularis, but differs somewhat
by having smaller spores often with a long apical nodule, on
average smaller nodules, and in the form of the pleurocystidia,
which tend to attenuate at the apex on average more than in
I. rivularis.
Micro-morphological measurements: Spores yellow-brown,
(8.8e)9.1e11.4(e12.7)  (6.0e)6.2e7.4(e8.1) mm, on average
10.2  6.8 mm, range of mean values 9.8e10.4  6.5e6.9 mm, Q ¼
(1.2e)1.3e1.7(e2.0), on average 1.50, range of mean Q-values
1.49e1.52 (80 spores from 3 collections). Basidia
31e38  10e12 mm, on average 34  11 mm, generally 4-spored,
clavate (8 basidia from 1 collection). Pleurocystidia (55e)
59e83(e92) (15e)18e28(e29) mm, on average 69 22 mm, range
of mean values 67e71  21e24 mm (54 pleurocystidia from 3 col-
lections), mainly fusiform, thick-walled, with up to 4 mm pale yel-
low to yellow wall. Caulocystidia abundant at upper part of stipe,
very few or not in the lower half, variable, thick-walled, some
cauloparacystidia thick-walled.
Specimens examined: ITALY, Bolzano, NE of CampoTúres, Riva di
Túres, coniferous forest, ca. 1600 m, 23 Jul 2006, G. Veroi (J. Vauras
23918*; TUR-A; GB; GenBank KY033785); Trentino, Rabbi, Jul 1909,
in coniferous forest, G. Bresadola (S-F229598, epitype); SPAIN,
Madrid, Rascafria, Pinus sylvestris and Salix sp., 8 Jul 2013, leg. F.
Pancorbo, det. F. Esteve-Raventos AH44415 (AH, GB).
Inocybe rivularis Jacobsson & J. Vauras, Windahlia18:18, 1990
[1989]. Figs. 4G, H, 5GeI.
For the description of I. rivularis we refer to Jacobsson and
Vauras (1989), repeating here only the sizes of spores and pleuro-
cystidia, and the distribution of caulocystidia. Spores (9e)
10e12.5(e13.5)  6.5e8.5(e9) mm, on average 11.1  7.8 mm, range
of mean values 10.5e12.3  7.1e8.1 mm, Q ¼ 1.2e1.6(e1.8), average
1.43, range of mean Q-values 1.35e1.6 (300 spores from 15collections). Pleurocystidia 50e76(e90)  14e28(e33) mm. With
further experience we complement the description and add that
sometimes the species can have fairly abundant velipellis on the
pileus (Fig. 5C), and with few thick-walled caulocystidia down to
base of stipe. Further, it can be found even in the low-alpine zone
with Betula nana. Even though this quite large species is fairly
common in suitable habitats in northern part of Fennoscandia,
there are rather few reports of it (Alpago Novello, 2002). It has been
selected as one of the indicator fungi for forests of Finland, asso-
ciated with spruce mires, brook-side forests and spring habitats
(Bonsdorff et al., 2014).
Specimens examined: CANADA, Quebec, Schefferville area,
Snowy Channel, alpine region, moist depression, growing by water,
14 Aug 1963, leg. P. Kallio 482*, det. J. Vauras (TUR; GenBank
KY033825); 14 Aug 1963, leg. P. Kallio 489b (TUR); ESTONIA, Hiiu,
K€aina, moist forest with Picea abies, Betula sp., Alnus glutinosa (L.)
Gaertn., Populus tremula L., Pinus sylvestris L. and Fraxinus excelsior
L., 14 Sep 2010, J. Vauras 27165 (TUR-A; TU; TAA101616, UNITE
UDB011734); FINLAND, Oulun Pohjanmaa, Oulu, Sanginsuu, Hakala,
4 Aug 1986, J. Ruotsalainen and J. Vauras 2152F (TUR! Holotype;
GenBank JN580885); Inarin Lappi, Utsjoki, Kevo, around Kevoj€arvi,
subarctic birch forest with Salix spp., close to the lake. 30 Aug 2012,
E. Larsson 178-12*(GB; GenBank KY033823); 27 Aug 2012, subarctic
birch forest with Salix spp., E. Larsson 160-12* (GB; GenBank
KY033824); Inarin Lappi, Utsjoki, Kevo, Kevoj€arvi, W. slope of
Korkea-Jehkas, SE of Jehkasj€arvi, low alpine slope with small
brooks, near Betula nana and Salix spp., amongst moist mosses, 24
Aug 2014, J. Vauras 30708* (TUR-A; GenBank KY033802);
Enonteki€on Lappi, Enonteki€o, Kilpisj€arvi, Malla Strict Reserve,
Mountain Pikku-Malla, alpine site with Salix spp. and Betula nana,
close to snow bed, moist, gently sloping site with mosses and
stones, 7 Aug 1985, J. Ruotsalainen and J. Vauras 1929* (TUR-A;
GenBank KY033804); Enonteki€on Lappi, Enonteki€o, Kilpisj€arvi,
mountain birch forest near small river Tsahkaljohka, rather moist
site by path, 13 Aug 1990, J. Vauras 4744* (TUR-A; GenBank
KY033808); Enonteki€on Lappi, Enonteki€o, Kilpisj€arvi, E side slope of
Jehkas, snow bed area, moist with Salix herbacea, 29 Aug 2013, E.
Larsson 288-13* (GB; GenBank KY033815); Kainuu, Ristij€arvi,
church village, Lukkarila, camping place Ristij€arven Pirtti, open
Fig. 3. Inocybe arctica (Vauras 2238F, holotype). A: Cross-sections of basidiomata. B: Pleurocystidia. C: Cheilocystidia and paracystidia. D: Basidiospores. E: Basidia. F: Caulocystidia
and cauloparacystidia at stipe apex. Bars: A 1 cm; BeF 10 mm.park lawnwith Betula sp., 27 Aug 2016, J. Vauras 31686* (TUR-A; GB,
GenBank KY033810); Pohjois-H€ame, Multia, Lehtom€aki, E of the
farm house, pastured forest on E slope with Picea abies and Betula
sp., onmull soil, 8 Sep 2013, J. Issakainen and J. Vauras 30173* (TUR-
A; GenBank KY033813); 8 Sep 2013, E of the farm house, pastured
forest on E slope with Picea abies and Betula sp., Pinus sylvestris,
Alnus incana (L.) Moench., Populus tremula and Salix caprea L., near
brook, J. Issakainen and J. Vauras 30172* (TUR-A; GenBank
KY033820); Per€a-Pohjanmaa, Rovaniemi, Meltaus, N of the Meltaus
e Marrasj€arvi, SE of Sortovaara, Kenk€akorpi, herb-rich forest with
Picea abies, Populus tremula, Betula sp., Alnus incana, Salix caprea
and Pinus sylvestris, with moist depressions, 23 Aug 1999, J. Vauras
15316*(TUR-A; GenBank KY033822); SWEDEN, Lycksele lappmark,
T€arna, Hemavan, Portbron, Job€acken, subalpine Betula pubescens
forest with Salix spp., 27 Jul 2014, E. Larsson 24-14* (GB; GenBank
KY033803); Lule lappmark, Jukkasj€arvi, Padjelanta NP, Huorso,Vastejaure, among Salix spp., 1 Aug 2001, S. Kuoljok 0125* (GB;
GenBank KY033805); 5 Aug 2000, torvkulle med Betula nana, S.
Kuoljok 0052* (GB; GenBank KY033809); Lule lappmark, Juk-
kasj€arvi, Padjelanta NP, Njoammeljaure NO, moist with Betula nana
and Salix spp., 18 Aug 2016, E. Larsson 241-16* (GB; GenBank
KY033806); Lule lappmark. Jukkasj€arvi, Padjelanta NP, Njoammel-
jaure, moist depression on S slope, near Salix spp. and Betula nana,
18 Aug 2016, J. Vauras 31568* (TUR-A; GenBank KY033801); Lule
lappmark, Jukkasj€arvi, Padjelanta NP, Slahpejaure NO, moist area
with small brook, with Betula nana and Salix spp., 14 Aug 2016, E.
Larsson 129-16* (GB; GenBank KY033807); Lule lappmark, Juk-
kasj€arvi, Padjelanta NP, Unna Duvgge, moist area with Betula nana
and Salix spp., 15 Aug 2016, E. Larsson 158-16* (GB; GenBank
KY033812); Torne Lappmark, Jukkasj€arvi, Latnajaure, moist with
Salix herbacea and Betula nana, 2 Aug 2006, E. Larsson 23-06* (GB;
GenBank KY033814); H€arjedalen, T€ann€as, Andersborgsv€agen, moist
Fig. 4. Comparison of basidiospores and pleurocystidia of Inocybe spp. A, B: I. arctica (EL 34-09; A basidiospores, B pleurocystidia). C, D: I. favrei (JV 31508F; C basidiospores, D
pleurocystidia). E, F: I. praetervisa (JV 23918; E basidiospores, F pleurocystidia). G, H: I. rivularis (EL & JV 30072F; G basidiospores, H pleurocystidia). Bars: 10 mm.area with brooks, Betula pubescens, B. nana and Salix spp., 15 Aug
2006, E. Larsson 82-06* (GB; GenBank KY033817); 15 Aug 2006, E.
Larsson 82B-16* (GB; GenBank KY033816); V€armland, Sunne,
G€ardesrud, 26 Aug 1990, B. Jansson* (GB; GenBank KY033818);
NORWAY, Troms, Storfjord, Lullesl€attan, mossy forest with Pinus
sylvestris, Betula sp. and scattered Picea abies, on tractor track, 22
Aug 2014, J. Vauras 30676* (TUR-A; GenBank KY033819); Oppland,
Valdres, W of Fisketjernsknausen, fj€allhed, 7 Jul 1995, M. Jeppson
3722* (GB; GenBank FN550887).Inocybe favrei Bon, Bull. trimest. Fed. Mycol. Dauphine-Savoie
25 (no. 97): 28, 1985. Figs. 4C, D, 5DeF, 6AeF.
¼ Inocybe decipientoides var. taxocystis J. Favre, Ergebnisse der
Wissenschaftlichen Untersuchungen des Schweizerischen Natio-
nalparks 5(33): 202, 1955.
¼ Astrosporina taxocystis J. Favre & E. Horak, Arctic and Alpine
Mycology 2: 230, 1987.
¼ Inocybe taxocystis (J. Favre & E. Horak) Senn-Irlet, Bot. Helv.
102(1): 55, 1992.
Fig. 5. Photos of Inocybe spp. AeC: I. arctica (EL123-09); A basidiomata, B pleurocystidia, C basidiospores. DeF: I. favrei (EL78-06; D basidiomata, E pleurocystidia, F basidiospores).
GeI: I. rivularis (JV30676; G basidiomata, H pleurocystidia, I basidiospores). Bars: B, E, H 20 mm; C, F, I 10 mm.¼ Inocybe taxocystis (J. Favre) Singer, The Agaricales in modern
taxonomy: 604, 1986 (nom. inval., lectotype not selected).
For the description of I. favrei we refer to Bon (1985) and Favre
(1955). In macro-morphology, I. favrei resembles I. rivularis and
I. arctica. In micro-morphology, it is characterized by having cys-
tidia (both pleurocystidia, cheilocystidia and caulocystidia) that are
ventricose and are fairly broad at the apex (see Figs. 4e6). Further,
the spores of I. favrei are on average smaller and pleurocystidia
shorter and wider compared with I. arctica. The species is most
common in the low alpine zone, but has been found on few occa-
sions in forests of the northern boreal zone on calcareous soil.Micro-morphological measurements: Spores nodulose, but
variable in form and size, fairly dark yellow-brown, thick-walled,
(9.7e)10.3e12.2(e13.5)  (6.4e)6.6e8.4(e8.7) mm, on average
11.2  7.3 mm, range of mean values 10.9e11.4  7.1e7.5 mm, Q ¼
(1.25e)1.4e1.7(e1.75), on average 1.52, range of mean Q-values
1.50e1.59 (100 spores from 5 collections). Basidia (28e)
29e40  (10e)11e14 mm, on average 33  13 mm, generally 4-
spored, clavate (17 basidia from 4 collections). Pleurocystidia
(54e)58e74(e80) (11e)14e26(e30) mm, on average 66 20 mm,
range of mean values 63e70 16e25 mm (83 pleurocystidia from 5
collections), mainly ventricose, thick-walled, with up to 4 mm pale
Fig. 6. Photos of Inocybe spp. AeC: I. taxocystis (lectotype, J. Favre 122a; A cheilocystidia, B pleurocystidia, C basidiospores). DeF: I. favrei (holotype, Bon 84095; D cheilocystidia, E
pleurocystidia, F basidiospores). Bars: A, D 20 mm; C, F 10 mm.yellow to yellow wall. Cheilocystidia 44e65  14e25 mm, on
average 56  19 mm (n ¼ 15), mainly ventricose, abundant, para-
cystidia pyriform, some thick-walled. Caulocystidia abundant at
stipe apex, (40e)45e66(e68)  (13e)16e24(e27) mm, on average
54  20 mm (n ¼ 20), more rare but present to stipe base, variable,
thick-walled, some of the cauloparacystidia thick-walled.
Specimens examined: FINLAND, Enonteki€on Lappi, Enonteki€o,
Kilpisj€arvi, at border of Finland and Norway, W of the main road,
low alpine site at small brook, near Betula nana and Salix spp., 20
Aug 2014, J. Vauras 30653F* (TUR-A; GB; GenBank KY033797);
Koillismaa, Kuusamo, J€ak€al€aniemi, fairly moist forest with Picea
abies, Pinus sylvestris and Betula sp., 28 Aug 2011, J. Vauras 28431*
(TUR-A; GenBank KY033795); FRANCE, Savoie, Haute-Maurienne,
Termignon, Bellecombe, parc national de la Vanoise, 30 Aug 1984,
Bon 84095* (LIP, Holotype; GenBank KY033786); Savoie, Bourg-
Saint-Maurice, Arc 2000, lac Marloup, 21 Aug 1993, with Salix
herbacea onwet soil along a small lake, higher alpine zone, 2500m,
leg. P.-A. Moreau, PAM93082101* (LIP; GenBank KY033799);
NORWAY, Troms, Storfjord, near the border of Finland, Bossovarri,
low alpine heath, margin of dried pond, near Salix spp. and Betula
nana, 21 Aug 2014, J. Vauras 30673F* (TUR-A; GenBank KY033798);
SWEDEN, Torne lappmark, Jukkasj€arvi, Latnajaure, with Salix her-
bacea, 6 Aug 2006, E. Larsson 57-06 (GB; TUR-A; GenBank
FN550886); 7 Aug 2006, moist place with Salix herbacea and Bis-
torta vivipara, E. Larsson 78-06* (GB; GenBank KY033800), 5 Aug2011, with Salix herbacea, E. Larsson 105-11* (GB; GenBank
KY033796); Lule lappmark, Jokkmokk, Padjelanta NP, Arralåbbdå,
fj€allhed med Salix herbacea, 11 Aug 2016, E. Larsson 41-16* (GB;
GenBank KY033794); Lule lappmark, Jokkmokk, Padjelanta NP,
Vielgggisbakte, in snow bed area with Salix herbacea, 12 Aug 2016,
E. Larsson 69B-16* (GB; GenBank KY033793); Lule lappmark, Juk-
kasj€arvi, Padjelanta NP, Slahpejaure NO, in snow bed areawith Salix
herbacea, 14 Aug 2016, E. Larsson 124-16* (GB; GenBank
KY033792), 14 Aug 2016, on S slope, moist boggy meadow near
Salix polaris, S. reticulata, J. Vauras 31508* (TUR-A; GB; GenBank
KY033789); 14 Aug 2016, on S slope near Salix herbacea, J. Vauras
31515* (TUR-A; GB; GenBank KY033788); Lule lappmark, Juk-
kasj€arvi, Padjelanta NP, Njoammeljaure NO, with Salix herbacea and
Bistorta vivipara, 18 Aug 2016, E. Larsson 229-16* (GB; GenBank
KY033791); 16 Aug 2016, snow bed area with Salix herbacea, E.
Larsson 231-16* (GB; GenBank KY033790); 16 Aug 2016, at dried
pond near Salix and Betula nana, J. Vauras 31579 (TUR-A; GB;
GenBank KY033787); SWITZERLAND, Kt. Graubünden, Tarasp,
Munt de la Bescha, Salix herbacea, 8 Sep 1945, J. Favre 122a* (G;
Lectotype; GenBank JN580884); Val S-charl, Mount Plazer, sur tapis
de Salix herbacea, sol gneissique, 20 Aug 1952, J. Favre 122c (G); 26
Aug 1952, J. Favre 122-1 (G).
5. Key to the species in the I. praetervisa clade
1a. Alpine/arctic ecosystems 2.
1b. Forest ecosystems 3.2a. High alpine to arctic, spores on average
11.5e12.2  7.2e7.9 mm, pleurocystidia on average
65e77  14e20 mm I. arctica.
2b. Sub- to low alpine, spores on average
10.9e11.4  7.1e7.5 mm, pleurocystidia on average
63e70  16e25 mm I. favrei.
3a. N. Europe and Canada, with Betula, spores on average
10.5e12.3  7.1e8.1 mm I. rivularis.
3b. S. Europe and N. America, in coniferous forests, spores on
average 9.8e10.4 6.5e6.9 mm I. praetervisa.6. Discussion
With the support from phylogenetic analyses of sequence data
we show that the I. praetervisa clade is comprised of at least four
morphologically and genetically similar but separate species. In
addition, two sequences from ectomycorrhizal (ECM) root tips
isolated from KobresiaWilld. in the alpine zone of China form a sub-
clade within this grouping, indicating the occurrence of one addi-
tional species in Asia. The species show overlapping but partly
different geographical distribution patterns in Europe.
Inocybe praetervisa is confirmed to have an intercontinental
distribution range. According to our data it occurs in the mountains
of Southern Europe then associated with coniferous mixed forests
and in Canada associated with Pseudotsuga Carriere. Also, seven
sequences deposited in GenBank under various names for collec-
tions from British Columbia, lacking information on host or forest
type, are 100% identical with the epitype ITS sequence. So far, we
have no specimens or evidence from sequence data of environ-
mental samples that confirm the occurrence of the species in
Northern Europe, although it has frequently been reported. How-
ever, these reports are based on misinterpretations of the species
concept (Esteve-Raventos et al. (2016)). It likely reflects the fact that
there are rather many species with similar morphology and tradi-
tionally I. praetervisa has been treated as a species with a broad
species concept in the Nordic countries (Jacobsson & Larsson,
2012). It was thought to be a species with a more or less
completely pruinose stipe, a marginate bulbous stipe base and a
pileus and stipe which darken upon drying. All four species treated
in this article have abundant caulocystidia at the apex of the stipe,
which are more rare but present to the stipe base. In addition, at
least I. arctica, I. favrei and I. rivularis have a more or less bulbous,
not marginate stipe base. This set of morphological characteristics
was observed by Kobayashi (1995) when he described the new
species I. fastuosa Tak. Kobay., from Tokyo, associated with Pasania
Oerst. and Cedrus Duhamel. The species is likely to belong to the
same group and fall within the same clade as the species treated
here but no sequence data is yet available.
Kokkonen and Vauras (2012) compared sequence data and
morphology of the lectotype of I. taxocystis (G13208, J. Favre 122a)
with I. rivularis (JV3610, paratype). They found that the two differ in
the ITS sequence data by 8 base pairs and 5 insertion deletion
events (1e4 bp in length) and somewhat in spore morphology,
color of pileus and habitat preferences. They concluded that they
should be regarded as separate species: a result that is confirmed
by this study.
In our analyses of ITS sequence data from the type specimens of
both I. taxocystis and I. favrei fall within the same clade and haveidentical ITS sequence data (Fig. 1), and according to the rules of
nomenclature, the oldest name, I. favrei should be used. This result
is also in agreement with the morphology which is very similar
(Fig. 6).
Horak (1987) described Astrosporina taxocystis as having a
brown to dark brown pileus, occasionally with a reddish tint, an
absence of veil, and a pruinose stipe with a marginate bulb at the
base. Based on thematerial he had at hand it should be restricted to
acid soils in association with S. herbacea, rarely S. retusa L. Bon
(1985) described I. favrei to have a pale yellowish brown pileus
with cortina (araneoso-velatus), a stipe that is pruinose in the up-
per part and a bulbous base that is obtusely marginate. This
contradiction suggests that determining the presence or absence of
a cortina in this group of species can be problematic, or that it is
variable. Bon found the species to be indifferent to pH and occur-
ring in association with S. herbacea, S. reticulata and Dryas octope-
tala. In our experience I. favrei has a brown color of the pileus, often
with a reddish tint, and often also with remnants of velum that
form patches (Fig. 5B). Because of the uneven distribution of the
caulocystidia on the stipe there has been confusion with these two
names. Inocybe favrei has been keyed out to the sectionMarginatae
(Bon, 1997, 1998), and I. taxocystis to the section Cortinatae (Singer,
1986; Bon, 1997, 1998). Also Ferrari (2006) follows these ideas e his
I. taxocystis is the same taxon as presented here (I. favrei).
From our molecular data we confirm that also I. favrei has an
inter-continental distribution range and seems to be rather ho-
mogenous in the molecular data (Fig. 1). In Europe it has a broad
geographic distribution but is likely restricted mainly to the alpine
zone and especially to the lower alpine zone, associated with Betula
nana, various Salix shrubs, S. herbacea and S. reticulata on moist
places as close to small brooks and in snow bed areas. The three
sequences from Alaska, downloaded from Genbank, are isolated
from soil and from ECM of Picea mariana, and suggest that it also
occurs in North America with a northern distribution in the taiga
and boreal forest ecosystems.
In the molecular data, I. rivularis shows a much larger sequence
variation than I. favrei. The herein recognized I. rivularis clade re-
ceives a week bootstrap support and eight of the sequences form a
sub-cladewith 84% support value (Fig.1). The collections within the
sub-clade are mainly collected in boreal to subalpine regions, likely
associated with Betula pendula Roth., B. pubescens or B. nana. One of
the specimens originates from Estonia and was collected in moist
mixed forest. The residual specimens originate from northern
boreal forests to the alpine zone, in associationwith B. pubescens, B.
nana, various Salix shrubs and S. herbacea in moist habitats. The
species seems to have a northern distribution within Europe, with
the most southern confirmed specimens from Estonia. Sequence
data of one specimen collected in Canada (Quebec) was included
and confirms the species also from North America. With the data at
hand the genetic differences cannot be correlated with differences
in ecology, geographic distribution ormorphology. So, we choose to
recognize the major clade despite the low support value.
Inocybe arctica is here described and represents the recovered
clade within the complex without a name. Our specimens originate
from Svalbard and Fennoscandia, mainly from the arctic and higher
alpine zones. On Svalbard it is one of the most common Inocybe
species found in mats of Dryas octopetala and with Salix polaris.
Specimens from the higher alpine zone in Fennoscandia are also
found with D. octopetala and S. polaris but also with S. reticulata and
S. herbacea. In comparison to I. rivularis and I. favrei, seems to have
preferences for more rich and calcareous soils. Inocybe arctica
seems to occur in higher alpine regions in Northern Fennoscandia
than I. favrei. Inocybe favrei is not confirmed from Svalbard and
I. arctica has not yet been confirmed from North America.
Disclosure
The authors declare no conflict of interest.
Acknowledgments
Curators of herbaria G, LIP, AH, O are gratefully acknowledged
for arranging loans, and Pierre-Arthur Moreau, Gro Gulden, Fer-
nando Esteve-Raventos and Fermin Pancorbo for sharing inter-
esting collections and photos, Seppo Huhtinen and reviewers for
valuable comments on the ms. Financial support was received from
The Swedish Taxonomy Initiative, ArtDatabanken SLU Uppsala,
Anna och Gunnar Vidfelts Fond, Stiftelsen Wilhelm och Martina
Lundgrens Vetenskapsfond, and the FinBOL-project funded by the
Finnish Cultural Foundation and Kone Foundation.
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