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Russula vinosoflavescens sp. nov., from deciduous forests of Northern Alsace, France

Authors:
MYCOTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October–December 2017—Volume 132, pp. 707–721
https://doi.org/10.5248/132.707
Russula vinosoavescens sp. nov.,
from deciduous forests of Northern Alsace, France
Jean Michel Trendel1, Felix Hampe2 & Annemieke Verbeken2*
1 7 rue des Coquilles, 67500 Haguenau, France
2 Department of Biology, Ghent University,
K. L. Ledeganckstraat 35, 9000 Ghent, Belgium
Correspondence to: *2mieke.verbeken@ugent.be 1jmtrendel@free.fr2felix.hampe@email.de
Abstract—Based on morphological, molecular, and ecological data, a new species of
Russula sect. Russula, found on several occasions under deciduous trees in Northern Alsace
is described and illustrated as: Russula vinosoavescens, belonging to R. subsect. Sardoninae.
Key wordsBasidiomycota, Russulaceae, ITS, Russula persicina, phylogeny
Introduction
Russula Pers. (Russulales, Basidiomycota) is an ectomycorrhizal genus with
a world-wide distribution that is particularly well studied in temperate regions,
notably in Europe (Romagnesi 1967; Sarnari 1998, 2005). e ectomycorrhizal
associations, which oen involve a selective fungus–plant partnership, were
sometimes advantageously used to delimit infrageneric taxa. Such has been
the case for R. subsect. Sardoninae Singer (Sarnari 1998, emend.), which is
divided into series—provisionally dened by Sarnari (1998)—comprising, on
one hand, species strictly associated with conifers (R. ser. Sardonia and R. ser.
Sanguinea, both essentially corresponding to R. subsect. Sardoninae as
conceived by Romagnesi 1967, 1987) and, on the other hand, species associated
with deciduous trees, sometimes in a host-specic relationship (R. ser.
Exalbicans = R. subsect. Exalbicantinae Singer as retained by Romagnesi, with
species found only under birches) and sometimes with a broader host-range
(R. ser. Persicina = R. subsect. Persicinae Romagn., species that can be found
under oaks, hornbeams, beeches, birches, chestnuts, poplars, willows).
708 ... Trendel, Hampe & Verbeken
Table 1. Russula, Gymnomyces, and uncultured specimens with GenBank
and UNITE sequence accession numbers used in the molecular analyses.
Species Specimen Country Sequence
G. gilkeyae JT 2572 USA AY239346
G. monosporus OSC 117360 USA EU669222
root sample (uncultured) France JQ890299
— — JN197654
— USA EF434062
— USA GU997950
— USA GU997948
R. americana F 18871 USA HQ604839
R. aquosa TU 101831 Estonia UDB015988
TU 101708 Estonia UDB011290
R. atrorubens TU 101691 Estonia UDB011358
TU 101851 Estonia UDB016006
R. bresadolae
(“R. atropurpurea”) HUE 178 Germany UDB000313
PRM 858109 Czech Republic HG423574
R. cavipes RT 9149 Italy JF908681
HUE 163 Germany AF418623
Russula cf. ammodica F 18874 USA HQ604840
R. citrinochlora TU 101905 Norway UDB016045
20024 Sweden UDB002550
R. consobrina TU 118108 Estonia UDB011223
TU 106965 Estonia UDB011207
R. depallens TU 101829 Estonia UDB015986
TU 101838 Estonia UDB015994
R. emetica F 14309 AY228360
DG 18 United Kingdom JQ888196
DG 44 United Kingdom UDB001628
R. exalbicans 2-1117IS76 — AY061674
R. fageticola (“R. mairei”) FH 12262 Germany KT934013
R. fageticola (“R. nobilis”) HUE 054 Germany UDB000346
R. fellea HUE 218 Germany UDB000345
HUE 177 Germany UDB000314
R. fragilis UE 2006-11-08-22 Italy UDB018436
UE 2004-20-09-04 Sweden UDB018434
R. gracillima TU 101725 Estonia UDB011361
TU 106437 Estonia UDB011176
R. luteotacta TU 106379 Estonia UDB011166
FH 12-187 Germany KT933991
R. ochroleuca TU 118113 Estonia UDB011228
LW 115 Germany UDB000295
R. persicina TU 101826 Estonia UDB015984
TU 106950 Estonia UDB011196
R. pumila F 133 Finland UDB009582
L 3X87 FM993279
R. queletii FH 12-237 Germany KT934007
RT 5140 Ital y JF908668
Russula vinosoavescens sp. nov. (France) ... 709
R. renidens TU 101809 Estonia UDB015975
JR 1758F IT Finland UDB011117
R. rhodomelanea FH 2011-BT307 Germany UDB018429
IB 92/451 HT Italy UDB018435
R. sanguinea TU 106710 Estonia UDB019728
FH 12240 Germany KT934008
R. sardonia TU 106951 Estonia UDB011197
FH 12215 Germany KT933999
R. silvestris JK 11080504 Sweden UDB018431
JK 11080802 Sweden UDB018430
R. thindii HT BSHC KD11-095 India KM386693
R. torulosa TU 106444 Estonia UDB011177
TU 101626 Italy UDB016261
R. vinosoavescens JMT 05081006 France UDB031189
JMT 08081512 France UDB031188
JMT 14090502 HT France UDB031187
JMT 05081007 France UDB031190
JMT 10080701 France UDB024104
JMT 13090715 France UDB024105
Outgroup
R. puellula (“R. puellaris”) MC 01-502 Denmark UDB000010
R. cuprea FH 12250 Germany KT934010
R. olivobrunnea TU 101883 Finland UDB016034
HT holotype; IT isotype.
e aim of this paper is to introduce a new Russula species occurring in
deciduous forests of Northern Alsace (NE France) and belonging to R. subsect.
Sardoninae. Our initial morphological concept of the species, based on several
collections from various localities, is fully supported by molecular data. ITS
sequence analysis shows that the dierent collections form a well-supported
clade in the Russula phylogenetic tree.
Materials & methods
Sampling
All collections of the new taxon were made by Jean Michel Trendel and are deposited
in GENT Herbarium. ese collections are referred to with an eight-digit code
YYMMDDXX (year; month; day; collection number). All collecting sites are located
in Northern Alsace (NE France); the code of the municipality is that assigned by the
National Institute for statistics and economic studies (INSEE); names (in quotation
marks) of forest or place are those indicated on the topographic maps (1:25000 scale)
from the Institut national de l’information géographique et forestière (IGN); map
coordinates of the collections refer to the Universal Transverse Mercator (UTM)
32N kilometer grid system; geological data (underlying rock) were mainly obtained
710 ... Trendel, Hampe & Verbeken
from geological maps (1:50000) edited by the Bureau de Recherches Géologiques et
Minières (BRGM).
Morphological analysis
Specimens were photographed in situ. e pictures are available at
http://www2.muse.it/russulales-news/.
Microscopic observations were carried out
on fresh material. Pileipellis elements were studied on radial sections made by hand
midway from the cap margin, mounted either in SDS Congo red (Clémençon 1999)
or in distilled water. e search for acido-resistant encrustations was performed by
staining with Ziehl carbol fuchsin followed by a rapid dierentiation using a 1M HCl
solution. Pileocystidia content was revealed using sulfovanillin (SV) freshly prepared
with a 50% or 80% H2SO4 aqueous solution. Hymenial elements were studied in the
above-mentioned observation media. In some cases, additional controls were carried
out on very ne cap sections made from dried material and further rehydrated in a
moisture-saturated closed chamber for 24 h before observation in water. Basidiospores
(from spore deposits) were examined in Melzer’s reagent and spore measurements
(ornamentation excluded) were recorded randomly for 300 spores (6 collections) in
side view. Measurements are given as (MINa) [AVa – 2*SD]–AvaAVb–[AVb + 2*SD]
(MAXb), with AVa = lowest mean value for the measured collections and MINa the
minimum value corresponding with this mean value, AVb = greatest mean value and
MAXb the maximum value corresponding with this mean value, and SD = standard
deviation calculated for the measurements of one collection (minimum and maximum
value are only given if not in the 2*SD-interval). Q stands for spore “length/width ratio
and is given as (MINQa)–QaQb–(MAXQb) with Qa and Qb being the lowest, and the
highest respectively, mean ratio for the measured specimen. e colour of the spore
deposit, referring to Romagnesi’s (1967) scale, was assessed against a personal chart
(JMT) and the Dagrons chart (unpublished). Macrochemical reactions were determined
using FeSO4 in crystalline form, a 4% phenol solution, and a strong Guaiac solution
made of permanently soaked Guaiac wood in 95% ethanol.
Molecular analysis
Total genomic DNA was extracted from dried material according to Nuytinck &
Verbeken (2003), with modications described in Van de Putte et al. (2010). e ITS
region was amplied using the primers ITS1F–ITS4 (White et al. 1990, Gardes & Bruns
1993) and with polymerase PerfectTAQ (5 PRIME, Hilden, Germany) in accordance
with the manufacturer’s recommendation. PCR amplication followed Eberhardt
(2012), and the PCR products were puried using the Qiaquick PCR Purication
Kit (Qiagen, Hilden, Germany) and directly sequenced with BigDye 3.1 technology
(Applied Biosystems, now ermo Fisher Scientic, Wilmington, USA). Specimens
JMT-05081006, JMT-05081007, and JMT-05081512 were extracted and sequenced in
Muséum National d’Histoire Naturelle de Paris (Marc-André Selosse) following Séne
et al. (2015).
Raw sequences were edited in the BioEdit Sequence Alignment Editor version 7.2.5
(Hall, 2013) or Sequencher version 4.8 (Gene Codes Corporation). Edited sequences
Russula vinosoavescens sp. nov. (France) ... 711
were aligned by MAFFT version 7 using the strategy E-INS-i (Katoh & Standley 2013).
Maximum-likelihood searches for tree building were carried out locally with 100
replicates with the GTR+GAMMA model, selecting the best solution of all replicates
analysis in RaXML 8.1.12 (Stamatakis 2014). Fast bootstrap searches were done locally
or through the CIPRES Science Gateway (Miller et al. 2010) with 10 000 replicates.
e nal alignment included a total number of 67 ITS sequences, with 21 sequences
corresponding to the current concept of R. subsect. Sardoninae (Sarnari 1998). Except
for the R. vinosoavescens collections cited as studied material all sequences were
retrieved from GenBank or UNITE (Table 1). e tree with the highest log likelihood
(–4523.1303) is shown. Initial tree(s) for the heuristic search were obtained automatically
by applying the Maximum Parsimony method.
Taxonomy
Russula vinosoavescens Trendel & F. Hampe, sp. nov. Figures 1, 2
MycoBank MB 819428
Diers from Russula persicina by its colour range, its mild or only slightly acrid taste,
its reticulate spore ornamentation, and the presence of granular pigments in cuticular
hyphae.
Type: France, Alsace, Morsbronn-les-Bains (67303), “Niederwald”, UTM 32N:
0406318/5419158, 213 m alt., on argillaceous soil with eumull humus (underlying
rock: upper Keuper red marls), under Quercus robur and Carpinus betulus with some
Fagus sylvatica nearby, in an area characterized by calcicolous mycoora with numerous
Phlegmacium (Cortinarius) species, 5 Sept. 2014, JMT-14090502 (Holotype, GENT;
UNITE UDB031187).
Etymology: Referring to the purplish-red wine-coloured (vinoso) cap and the
tendency of the cap to become brown-yellowish or ochraceous (avescens) as well as
when handling the stipe.
Pileus 4.0–6.5 cm diam., more or less eshy, rather rm to very rm but with
margin occasionally slightly elastic, oen irregular, at times knotty, convex-
attened with the margin sometimes incurved or even inrolled, then broadly
but shallowly depressed; margin not or shortly sulcate, oen exuose-undulate
or lobed, in some cases showing a very clear, whitish border extending to the
edge of the gills (giving a “festooned” appearance); surface rugulose, more or
less radially veined, frequently uneven-bumpy in the centre, a little shiny in wet
conditions, but soon dry (except sometimes in the centre which remains a little
greasy) and then more mat, purple-red, carmine-red, vinaceous-red, somewhat
violet, rather vivid, but frequently paler, livid pinkish-vinaceous, with mauve
(lilac) greyish shades that can entirely replace the reddish tinges, even more
clearly greyish purple at the margin, becoming yellowish cream-ochraceous
from the centre or showing locally ochre-bistre areas merging in the reddish
and greyish mauve tints, sometimes becoming almost totally beige-brown,
712 ... Trendel, Hampe & Verbeken
ochreous-brown, with livid purple-violet glints mostly near the margin and a
residual purple-brown at centre. Lamellae adnate or attenuated-subdecurrent,
sometimes sub-emarginate, with a variable number of lamellulae, crowded to
rather spaced, interveined, anastomosing and more or less forked at every
level, in some cases more particularly near the stem (exceptionally regular,
without any forking or anastomosing), (sub)acute at the cap margin (rarely
subobtuse), narrow (0.3–0.5 cm, exceptionally broader 0.7 cm), whitish or
very pale cream; edge entire but oen somewhat irregularly wavy. Stipe 2–6 ×
0.9–1.5(–2) cm, cylindrical or slightly enlarged downwards, usually rounded
at the base (in a few cases tapering), sometimes eccentric, rm to almost hard,
with a rm medulla (exceptionally hollow-stulous); surface nearly smooth,
nely wrinkled, white with occasionally a purple tinge (in one case remarkably
grey-purplish, somewhat brownish), distinctly yellowing (dirty, oen with
greyish-brown shades) when handled or bruised. Context white, in the pileus
almost unchanging, in medulla of stipe slightly yellowing, sometimes a little
purple-violet beneath the cuticle in the most coloured forms; taste mild (or
almost mild, occasionally with an unpleasant aertaste, somewhat bitter),
mild to slightly (and sometimes volatile) acrid in the gills, exceptionally more
strongly acrid; smell not distinctive.
Macrochemical reactions FeSO4: rose-orange; Phenol: brown; Guaiac:
in most cases reacting on the stem surface almost immediately (<2 s, in one
case more slowly), but of little intensity at the beginning, developing (10–15 s)
in blue of medium intensity, exceptionally with a stronger reaction, (dark blue);
on gills, the reaction developing slowly and weakly (blue-green to light blue).
Spore print cream, II(a)b–c.
Basidiospores subglobose to broadly ellipsoid, 6.4–7.28.1–9.0 × 5.9–6.5
6.9–7.7 µm (n = 300), Q = 1.06–1.111.20–1.30; ornamentation amyloid,
oen with incompletely amyloid warts; warts conical-obtuse, or more acute,
sometimes somewhat truncated, generally around 0.4–0.8 µm high, but
also frequently reaching 1.0 µm or higher, sometimes locally catenulate,
interconnected by crests or more frequently by (oen very ne) connectives,
forming a well-developed and oen nearly complete (with numerous meshes)
reticulum, which may appear confused in some cases; suprahilar plage
moderately amyloid, more or less bordered by small warts or low crests.
Basidia 4-spored, 42–57 × 9.5–12.5 µm, including sterigmata (about 5 µm
long). Pleuromacrocystidia 60–90 × 10.5–14.5 µm, numerous, fusiform
or more or less clavate, most commonly with a pointed appendage, which
can show constrictions, more rarely ending in a short obtuse protuberance,
Russula vinosoavescens sp. nov. (France) ... 713
Fig. 1. Russula vinosoavescens: colour forms observed. A, B. holotype, collection JMT-14090502;
C. collection JMT-05082042, with marked reddish pigmentation; D. collection JMT-05081006,
with marked reddish pigmentation; E. collection JMT-08081512, with predominating ochraceous
colour; F. collection JMT-10080701, with mauve–lilac colour reminiscent of a Griseinae.
714 ... Trendel, Hampe & Verbeken
strongly staining with SV. Pileipellis composed of hyphae containing reddish-
violaceous granular pigments; epicuticular elements (2.0–)2.5–4.0 µm diam.,
sometimes branched, exuose, sometimes inated up to 5.5 µm; terminal cell
obtuse, or slightly thickened with a short appendage, also subcapitulate, or
(exceptionally) pear-shaped 7.0 µm diam., but also conversely more or less
attenuated (sometimes abruptly). Pileocystidia of two types; type 1 usually
unicellular (occasionally with 1 septum), rarely bid, clavate–fusiform, less
frequently cylindric, 4.0–9.5 µm diam., oen appendiculate–capitulate or
showing an obtuse apical protuberance, sometimes slender, (at least 120 µm
long, narrowed at the base to 2.5–3.0 µm diam.), with a content reacting variably
depending on the collection—but also on a cuticular section—when treated
with SV, staining (rather) strongly (grey-blackish) or weakly (grey-pinkish)
or even remaining inert (pinkish); type 2 lactiferoid (some clearly termini of
ascending lactifers), pluriseptate, more or less regularly cylindrical, 6.0–10.5
µm diam., with variably shaped terminal elements that sometimes taper (with
possible constrictions), sometimes are somewhat thickened or subclavate, or
with type 1 type appendages, originating in the deeper pellis, distinctly reacting
with SV (except in one specimen where the granular content of pileocystidia
type 2 is inert or reacts only very weakly in SV); without encrusted elements
aer treatment with Ziehl fuchsin.
Ecology & distribution—Under deciduous trees, associated with
Carpinus betulus, or Quercus (Q. robur or Q. petraea), or both (possibly also
with Fagus sylvatica), in a rather wet environment (possibly periodically drier),
on more or less argillaceous soils, neutral to supercially slightly acidic, usually
nutrient-rich with a rather high base saturation level (calcium-rich), with
mesotrophic to eutrophic mull humus. Phenology: late July to early September
(summer fruiting). Known only from six sites (9 collections) in Northern
Alsace (France).
Additional specimens examinedFRANCE, Alsace. Bas-Rhin, Mattstall,
commune associated with Lembach (67263), ‘Sauerhald’, UTM 32N: 0409918/5426543,
alt. 203 m, underlying rock: Upper Muschelkalk marly calcareous formation, under
Quercus, Carpinus, Fagus (forest locally rich in orchids, e.g., Epipactis microphylla),
JMT-13090715 (GENT, UNITE: UDB024105); Dauendorf (67087), ‘Herrenwald’,
0402753/5410066, 168 m, Loess from the Quaternary (and more ancient alluvial
deposits?), in a wet environment, nitrogen-rich and with a high base saturation level,
under Carpinus, Quercus, Alnus glutinosa, Fraxinus excelsior, Ulmus sp., and with a
herbaceous layer comprising Allium ursinum, Carex sylvatica, Circaea lutetiana, Geum
urbanum, Glechoma hederacea, Paris quadrifolia, Stachys sylvatica, JMT-10080701
(GENT, UDB024104); 0402762/5409965, 171 m, Loess from the Quaternary (and
more ancient alluvial deposits?), under Carpinus, Quercus, JMT-05082042 (GENT);
Russula vinosoavescens sp. nov. (France) ... 715
Fig. 2. Russula vinosoavescens (holotype, collection JMT-14090502). A. fruitbody in sectional
view; B. pileus epicutis: pileocystidia (with content shown schematically) and hyphal terminal
elements; C. hymenial cystidia on gill sides; D. spores as observed in Melzer’s reagent. Scale bars:
A = 1 cm; B = 10 µm; C = 20 µm; D = 5 µm.
716 ... Trendel, Hampe & Verbeken
Bettwiller (67036), ‘Buchwald, 0365394/5416535, 347 m, Upper Muschelkalk marly
calcareous formation, under Quercus, Carpinus, with some Fagus and Prunus avium,
JMT-05081006 (GENT, UDB031189) and JMT-05081007 (GENT, UDB031190), most
likely belonging to the same mycelium; Forstheim (67141), ‘der Wald’, 0405631/5415509,
211 m, Pliocene sands and clays, wet environment, under Quercus, Carpinus, Populus
tremula with Russula lutensis Romagn. and R. rutila Romagn., JMT-08081512 (GENT,
UDB031188); Drusenheim (67106), ‘Barrwald’, approximately 0420900/5401000, 124 m,
Quaternary alluvial deposits, wet environment, on bare blackish ground, with Carpinus,
Quercus, Alnus glutinosa, Fraxinus excelsior, 25/07/1981, JMT-81072501 (GENT, spore
print only available).
Discussion
Molecular analysis
e six ITS sequences of the newly described R. vinosoavescens form
an independent clade, which receives high bootstrap support (Fig. 3). e
clade corresponding to R. vinosoavescens is nested within a large, supported
(bootstrap value 75) clade that includes the typical European representatives of
R. subsect. Sardoninae (R. sardonia Fr., R. queletii Fr., R. torulosa Bres.). Beside
these, it contains a number of species (e.g., R. cavipes Britzelm., R. exalbicans
(Pers.) Melzer & Zvára, R. gracillima Jul. Schä., R. luteotacta Rea, R. persicina
Krombh., R. renidens Ruots. et al., R. sanguinea Fr.) included by Sarnari (1998)
in his emended concept of R. subsect. Sardoninae but attributed to dierent
(although closely related) infrageneric taxa by previous authors (Romagnesi
1967, Singer 1986, Bon 1988). In addition to the abovementioned species,
which fairly well represent R. subsect. Sardoninae sensu Sarnari, the clade
includes extra-European Sardoninae species such as R. thindii K. Das & S.L.
Miller and some poorly known species with an uncertain systematic position
(e.g., R. americana Singer, R. citrinochlora Singer).
e deeper phylogenetic relationships within the Sardoninae clade as
well as its external relationships with R. fellea (Fr.) Fr., R. consobrina (Fr.) Fr.,
R. ochroleuca Fr., and the large clade representing R. subsect. Russula
s.l. (including a sequence of the generic type R. emetica (Schae.) Pers.)
remain poorly resolved in the ITS analysis. Within the Sardoninae clade,
R. vinosoavescens occupies a rather isolated position in a weakly supported
subclade containing R. renidens and samples identied as R. citrinochlora.
Morphological analysis
Despite its mild taste (in most cases only weakly acrid in gills),
R. vinosoavescens shows a combination of characters—a more or less eshy
rm pileus, cuticle oen rugulose with polychromic colours, cream spore
Russula vinosoavescens sp. nov. (France) ... 717
Fig. 3. Molecular phylogenetic analysis by Maximum Likelihood method-based tree with the
highest log likelihood (–4523.1303), based on ITS sequences.
718 ... Trendel, Hampe & Verbeken
print, well-characterised and non-encrusted pileocystidia—that is consistent
with its position in R. subsect. Sardoninae sensu Sarnari. Moreover, by its
habitus, somewhat decurrent narrow gills, and clear yellowing of its stipe when
bruised, this species is very reminiscent of certain forms of R. persicina. Russula
vinosoavescens diers, however, unambiguously from the latter by its colours,
which are never ‘purered. Indeed, even in its reddest forms, the cap always
displays some purple-vinaceous tinges, presumably reecting the existence of
one or several blue pigments, occurring in sucient amount to signicantly
shade the red ones. e presence of these blue pigments becomes even more
apparent when grey-mauve (lilac) tints replace—sometimes completely—the
reddish colouring. In addition, this russula appears to fade readily to yellowish
ochre, or even brown-beige, and therefore colour forms devoid of any reddish
or bluish shades might be expected.
Other striking features are the oen rugulose surface of the cuticle and the
strong tendency of the lamellae to become anastomosed-forked, though it
should be kept in mind that these characters are variable. e same is true for
the variability of the speed and intensity of the Guaiac reaction, which is mostly
in an average position on the reactivity scale.
One of the most striking microscopical features is the aggregation of
pigments into granules (sometimes distinctly combining red and blue
pigments) that are generally easily observed in the cap cuticular hyphae. e
cuticular structure of specimens for which we were unable to examine in fresh
condition was re-examined in dried material. Concerning the least pigmented
collection JMT-08081512 (Fig. 1-E), cuticular sections from the central part
of the cap—apparently the richest in residual red and blue pigments—revealed
the presence of pigmented grains. Our search was much less conclusive for the
vividly coloured specimens JMT-05081006 (Fig. 1-D), and even if we were able
to show the presence of sparse coloured granules, it appears that a signicant
amount of pigment is either vacuolar (soluble) or at least not distinctly granular.
In addition, examination of the cuticle of collection JMT-05082042 (Fig. 1-C),
not studied in a fresh state, clearly revealed the presence of pigmented granules.
As reported by Romagnesi (1967) and Sarnari (1998), granular pigments are
highly characteristic of R. subg. Heterophyllidia but are also found occasionally
in some species belonging to other groups. In this respect, one should especially
mention R. renidens, a Nordic russula growing under birches, which shows
abundant granular red-purplish pigments, the latter probably co-occurring
in a soluble form accounting for the intensity of its colours (Sarnari 1998).
A similar situation may also exist for some of our vividly coloured collections
Russula vinosoavescens sp. nov. (France) ... 719
(notably JMT-05081006 and JMT-05081007). Indeed, it seems likely that
pigment aggregates contribute less eciently to global colouring compared to
those in solution, and that a strong predominance of granular pigments over
their soluble counterparts may explain the relative paleness of some of our
collections.
e morphology of the terminal hyphal cells of the epicuticular elements
appears too variable to be usable for species recognition. On the other hand,
pileocystidial shape seems informative at species level. It is also signicant that
this characteristic can be found in almost all members of R. subsect. Sardoninae
as dened by Sarnari, where the new species belongs.
Another important diagnostic feature is the strongly reticulate spore
ornamentation that distinguishes R. vinosoavescens from similar species
sharing the same cream spore print and also growing under deciduous trees,
such as R. persicina or the birch associates, R. exalbicans and R. renidens.
Ecology
Within Russula subsect. Sardoninae, partitioned by Sarnari (1998) into
ecological series based on Russula/tree partnerships, R. vinosoavescens ts in
R. ser. Persicina, devoted to species associated with broadleaf trees (but not
strictly with birches); the other series cover species linked strictly to either
conifers (R. ser. Sardonia and R. ser. Sanguinea) or birches (R. ser. Exalbicans).
Indeed, R. vinosoavescens appears associated with oak (Quercus robur or
Q. petraea) or hornbeam (Carpinus betulus) or both, the only trees present
on all collecting sites. A wider host association might include beech (Fagus
sylvatica, generally well represented in the forests of interest but sometimes
rather far away from the fruiting bodies) or, less likely, aspen (Populus tremula,
which can occur in more complex environments). Birch (Betula spp.), absent
from all the prospected sites, should not be considered a potential associate.
However, regarding Romagnesis (1967) initial concept of Persicinae, which
refers to “intermediate forms between Emeticinae and Sardoninae [both sensu
Romagnesi!], with less reticulate spores and gills more decurrent or weeping
than the former, and more purely red in colour than the latter” [a concept at
least partly shared by Sarnari (1998)—“pileus pure red”], it becomes necessary
to restrict Sarnari’s R. ser. Persicina denition exclusively to ecology, as in
R. vinosoavescens colouration is not restricted to pure red and its spores are
rather strongly reticulate. In his classication, Bon (1988) does not retain
Romagnesi’s Persicinae group but treats the dierent taxa of R. persicina in his
emended R. subsect. Exalbicantinae. In this context, R. vinosoavescens would
take its place beside R. exalbicans, which also oen displays washed-out greyish
720 ... Trendel, Hampe & Verbeken
colours, and R. renidens, genetically a very closely related species sharing some
outstanding morphological features such as pigments that are at least partly
granular or lamellae that are more or less forked.
Acknowledgments
We are extremely thankful to Josie Lambourdière and Prof. Marc-André Selosse
(Muséum national d’Histoire Naturelle, Paris) for some additional molecular analyses,
and to Dr. Philippe Schaeer for helpful discussions. Dr. Ursula Eberhardt and Georey
Kibby are acknowledged for useful comments and reviewing the paper.
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