ArticlePDF Available

Abstract and Figures

Species of Tarzetta are morphologically well-defined but their taxonomy is confused because of the multiple interpretations of the oldest names, i.e. T. catinus and T. cupularis. The exploration of this genus with molecular and ecological data gave us the opportunity to partially clarify its taxonomy and to propose typifications of names. The type collections of T. pusilla, T. gaillardiana, and T. spurcata in the sense of Harmaja were reviewed. Hypotarzetta insignis, a semi-hypogeous species, is reinstated in Tarzetta based on molecular data. New species are also described and illustrated: T. gregaria sp. nov. is characterized by its small size and gregarious orangish ascocarps, T. alnicola sp. nov. is a small species associated with alders in riparian woodlands, T. alpina sp. nov. grows in boreo-alpine regions, T. sepultarioides sp. nov. is characterized by its Sepultaria-like appearance and ascus base shape, T. melitensis sp. nov. is a small species described from Malta, and T. quercus-ilicis sp. nov. is a small, sessile species, associated with Quercus ilex in Mediterranean regions. T. betulicola is a provisional name proposed for a species from the T. catinus complex, probably associated with birch trees
Content may be subject to copyright.
Preliminary notes on the genus Tarzetta (Pezizales) with typifi-
cations of some species and description of six new species
Arthur C. GRUPE, 11 (6) : 309–334
Mise en ligne le 24/12/2019
Abstract: Species of Tarzetta are morphologically well-defined but their taxonomy is confused because of
the multiple interpretations of the oldest names, i.e. T. catinus and T. cupularis. The exploration of this genus
with molecular and ecological data gave us the opportunity to partially clarify its taxonomy and to propose
typifications of names. The type collections of T. pusilla, T. gaillardiana, and T. spurcata in the sense of Harmaja
were reviewed. Hypotarzetta insignis, a semi-hypogeous species, is reinstated in Tarzetta based on molecular
data. New species are also described and illustrated: T. gregaria sp. nov. is characterized by its small size and
gregarious orangish ascocarps, T. alnicola sp. nov. is a small species associated with alders in riparian wood-
lands, T. alpina sp. nov. grows in boreo-alpine regions, T. sepultarioides sp. nov. is characterized by its Sepul-
taria-like appearance and ascus base shape, T. melitensis sp. nov. is a small species described from Malta, and
T. quercus-ilicis sp. nov. is a small, sessile species, associated with Quercus ilex in Mediterranean regions. T. be-
tulicola is a provisional name proposed for a species from the T. catinus complex, probably associated with
birch trees.
Keywords: Ascomycota, Pyronemataceae, Otideaceae, Tarzettaceae, rDNA analyses, nomenclature, 6 new
The name Tarzetta was established by COOKE (1879) as a subgenus
of Peziza Fr., but no type was designated. It was later combined to
the genus Tarzetta by LAMBOTTE (1888) and typified by ROGERS et al.
(1971) with Peziza tarzetta Cooke as type-species (according to Art.
22 ICN). This name was considered as a later synonym of Peziza
catinus Holmsk. (NANNFELDT, 1938). Consequently, Tarzetta replaced
Pustulina Eckblad, a new name published in replacement of Pustu-
laria Fuckel, nom. illeg. (ECKBLAD, 1968).
The case of orthography of Tarzetta was previously discussed by
VAN VOOREN (2010) where we explain why we do not agree with YAO
& SPOONER (2002) who proposed to correct it as Tazzetta.
The most complete work on Tarzetta was published by HÄFFNER
(1992) but the typification of T. catinus (Holmsk.) Korf & J. K. Rogers
and T. cupularis (L.) Lambotte, two of the oldest names in the genus,
were not treated. Serious confusion exists in the application of these
names, and the precise interpretations differ in the literature. This is
mainly due to the exceedingly few microscopic differences ob-
served between these two species. In its current concept, the genus
Tarzetta contains approximately ten species that are distributed
worldwide (source: MycoBank database, accessed on Sept. 2019).
Species of Tarzetta form ectomycorrhizae (EcM) with trees, e.g. Fagus
sylvatica (TEDERSOO et al., 2006), Quercus douglasii (SMITH et al., 2007),
Alnus spp. (TEDERSOO et al., 2009), Pinus monticola (SOUTHWORTH et al.,
2009) and Corylus avellana (BENUCCI et al., 2014). Our phylogenetic
analyses (Fig. 1 and 2) suggest a higher diversity and a possible host-
specificity for several species based on the EcM sequences available
in public databases (e.g. GenBank).
The goal of this work is to clarify the taxonomy of the species
mainly described and illustrated by BOUDIER (1905-1910) and HARMAJA
(1974), based on type studies, morphological and phylogenetic data
from the internal transcribed spacer region (ITS) and the 28S large-
subunit (LSU) of the rDNA. This work will provide a new framework
for future studies. A type revision of T. gaillardiana (Boud.) Korf & J.K.
Rogers, T. spurcata (Pers.) Harmaja and T. pusilla Harmaja is also pro-
vided. Six new species are described and illustrated based on mor-
phological, ecological, and molecular data: Tarzetta alnicola,
T.alpina, T. gregaria, T. melitensis, T. quercus-ilicis, and T. sepultarioides.
Material and methods
Morphology and cytology. — Observations were made from
both fresh and dried material. For dried samples, parts of the spec-
imens were rehydrated for approximately eight hours in tap water
at room temperature. Observations of microscopic characters were
made in various solutions and stains: tap water, Cotton Blue (CB) in
lactophenol, 5% potassium hydroxide (KOH), and Lugol’s solution
(IKI) to test the amyloid reaction. Measurements were made on 30
ascospores from each collection, mounted in water, under the 100×
oil immersion lens of a transmission light microscope, excluding the
ornamentation. “X” represents the average value of spore dimen-
sions, and “Q” the ratio between spore length and width. Macropho-
tographs were made in situ using digital cameras, while micrographs
were taken using a microscope mounted digital camera. Line draw-
ings were made freehand to scale. The symbol † is used to annotate
observations from rehydrated material.
DNA extraction, amplification and sequencing. DNA was
extracted using the method described in VAN VOOREN et al. (2015).
Loci that were amplified included the ITS (ITS1, 5.8S and ITS2 rDNA)
and LSU (D1, D2 and D3 of the 28S rDNA). Sequences obtained dur-
ing this study were deposited in GenBank under the accession num-
bers listed in Table 1.
Phylogenetic analyses. — BLAST® was used to select the most
highly similar ITS and LSU sequences from the International Nu-
cleotide Sequence Database Collaboration public databases. For the
LSU locus (662 sites), the evolutionary analyses were conducted in
Mega X (KUMAR et al., 2018) with 68 nucleotide sequences. The align-
ment was done with the MUSCLE algorithm (EDGAR, 2004). The evo-
lutionary history was inferred by using the Maximum Likelihood
(ML) method based on the General Time reversible (GTR) model (NEI
& KUMAR, 2000) with 1000 bootstrap iterations. The LSU tree with the
highest log likelihood (-3494.72) is shown in Fig. 1. The percentage
of trees in which the associated taxa clustered together is shown
next to the branches. Initial tree(s) for the heuristic search were ob-
tained automatically by applying Neighbor-Join and BioNJ algo-
rithms to a matrix of pairwise distances estimated using the
Maximum Composite Likelihood (MCL) approach, and then select-
ing the topology with superior log likelihood value. A discrete
Gamma distribution was used to model evolutionary rate differ-
ences among sites [5 categories (+G, parameter = 0.5999)]. The rate
variation model allowed for some sites to be evolutionarily invari-
able ([+I], 42.22% sites). For the ITS locus (515 sites), 72 sequences
were aligned in Mesquite (MADDISON & MADDISON, 2015) using the
MUSCLE algorithm. A maximum likelihood (ML) search was per-
formed using RAxML (STAMATAKIS, 2014) with the GTR GAMMA model
of nucleotide substitution and rapid bootstrapping on the CIPRES
Science Gateway (MILLER et al., 2010). Support for phylogenetic rela-
tionships was assessed based on 1000 bootstrap replicates in
RAxML and trees were viewed using FigTree (RAMBAUT, 2010). The
trees are drawn to scale, with branch lengths measured in the num-
ber of substitutions per site.
Nomenclature. —All references to articles of ICN come from the
Shenzhen Code (TURLAND et al., 2018). Typifications and novelties
were registered in the MycoBank Database (
Definition of the genus Tarzetta
Since its reassessment at the beginning of the 1970s (ROGERS et al.,
1971; KORF, 1972), the genus Tarzetta was considered as a member
of the Pyronemataceae. The phylogenetic work by HANSEN et al.
(2013) — based on a four-gene phylogeny demonstrated that
this family was paraphyletic and placed Tarzetta in a /tarzetta-
geopyxis lineage, a position confirmed by WANG et al. (2016) and
KUMAR et al. (2017) with an enlarged dataset. BARAL (in JAKLITSCH et al.,
2016) resurrected the family Otideaceae erected by ECKBLAD (1968)
and included Tarzetta as a member, while WIJAYAWARDENE et al. (2017)
placed Tarzetta in Pezizales incertae sedis and EKANAYAKA et al. (2018),
with a five-gene phylogeny, published the new family Tarzettaceae
Nom Voucher Number Host(s) GenBank Accession #
Hypotarzetta insignis LY:NV 2014.03.07 Cedrus atlantica MN712290 MN712245
Hypotarzetta insignis F.S. 2013002 Pseudotsuga menziesii MN712291 MN712246
Tarzetta alnicola LY:NV 2017.09.08 Alnus sp. MN712299 MN712254
Tarzetta alnicola LY:NV 2017.08.33 Alnus sp. MN712300 MN712255
Tarzetta alnicola LY:NV 2017.08.36 Alnus incana MN712301 MN712256
Tarzetta alnicola LY:NV 2008.07.06 Alnus glutinosa MN712302 MN712257
Tarzetta alpina LY:NV 2009.08.11 unknown MN712259
Tarzetta betulicola ad int. M.P. 2018-133 Betula, Pinus uncinata – MN712273
Tarzetta bronca SAK-17-0423-5 Populus sp. MN712304 MN712261
Tarzetta catinus V.R. 20190509 Fagus sylvatica, Quercus MN712315 MN712274
Tarzetta catinus GM 20190505 Fagus sylvatica, Quercus MN712316 MN712275
Tarzetta catinus LY:NV 2007.04.18 Fagus sylvatica, Corylus avellana, Picea abies – MN712276
Tarzetta catinus LY:NV 2002.06.05 Fagus sylvatica, conifers MN712277
Tarzetta catinus LY:NV 2010.05.09 Fagus sylvatica MN712317 MN712278
Tarzetta cf. catinus M.C. 18-10-13 Castanea sativa, Quercus sp. MN712303 MN712258
Tarzetta cf. catinus AH:53705 Abies alba, Salix caprea, Corylus avellana MN712325 –
Tarzetta cupularis LY:NV 2019.05.11 Carpinus betulus, Quercus, Castanea sativa MN712319 MN712280
Tarzetta cupularis MC 18-5-26 Quercus pubescens, Castanea sativa MN712320 MN712281
Tarzetta cupularis LY:NV 2013.11.24 hardwoods MN712321 MN712282
Tarzetta cupularis LY:NV 2006.10.31 Quercus, Buxus sempervirens MN712322 MN712283
Tarzetta cupularis LY:NV 2004.10.38 Quercus, Pinus sylvestris MN712323 MN712284
Tarzetta gregaria LY:NV 2017.08.16 unknown MN712288 MN712243
Tarzetta melitensis LY:CS871 Quercus ilex, Laurus nobilis MN712324 MN712285
Tarzetta ochracea LY:NV 2018.06.05 Fagus sylvatica MN712308 MN712266
Tarzetta ochracea M.C. 15-6-20 Abies alba, Fagus sylvatica MN712309 MN712267
Tarzetta ochracea LY:NV 2019.06.01 Quercus, Populus tremula MN712310 MN712268
Tarzetta ochracea LY:NV 2013.06.12 Castanea sativa, Tilia MN712311 MN712269
Tarzetta ochracea TaCu18061801 Abies sp., Fagus sylvatica MN712312 MN712270
Tarzetta ochracea LY:NV 2018.06.11 Fagus sylvatica MN712313 MN712271
Tarzetta ochracea LY:NV 2013.06.43 Fagus sylvatica MN712314 MN712272
Tarzetta pseudocatinus M.C. 18-5-21 Crataegus MN712292 MN712247
Tarzetta pseudocatinus C.A. 20121020.1 Fagus sylvatica MN712293 MN712248
Tarzetta pseudocatinus LY:NV 2019.05.12 Carpinus betulus MN712294 MN712249
Tarzetta pseudocatinus LY:NV 2014.08.19 Corylus avellana MN712295 MN712250
Tarzetta pseudocatinus LY:NV 2013.11.09 Castanea sativa, Carpinus betulus MN712296 MN712251
Tarzetta pseudocatinus LY:NV 2000.05.01 hardwoods MN712297 MN712252
Tarzetta pseudocatinus LY:NV 2006.09.01 Carpinus betulus, Quercus MN712298 MN712253
Tarzetta quercus-ilicis LY:NV 2014.03.20 Quercus ilex MN712306 MN712264
Tarzetta quercus-ilicis M.C. 14-3-15 Quercus ilex MN712307 MN712265
Tarzetta sepultarioides LY:NV 2017.08.03 Picea abies, Alnus incana, Acer sp. MN712289 MN712244
Tarzetta sp. M.C.-Sardinia unknown MN712260
Tarzetta sp. LY:NV 2001.03.01 unknown (Pinus ?) – MN712262
Tarzetta sp. LY:NV 2018.06.12 Corylus avellana MN712305 MN712263
Tarzetta sp. M.C. 08-5-23 Quercus ilex MN712318 MN712279
Tarzetta sp. LY:NV 2014.07.10 Betula, Salix MN712326 MN712286
Table 1 – List of sequenced collections for this study
Fig. 1 –Molecular phylogenetic analysis by Maximum Likelihood method of LSU alignment of Tarzetta, rooted by other species of the /tar-
zetta-geopyxis lineage.
Fig. 2 – Molecular phylogenetic analysis by Maximum Likelihood method of ITS alignment of Tarzetta, rooted by other species of the /tar-
zetta-geopyxis lineage.
corresponding to the lineage /tarzetta-geopyxis as previously de-
fined. This group also contains the apothecial genus Geopyxis (Pers.)
Sacc. and the truffle-like genera Hydnocystis Tul. & C. Tul., Pauro-
cotylis Berk. and Densocarpa Gilkey (KUMAR et al., 2017).
Macromorphologically Tarzetta is characterised by apothecial as-
comata, sessile or stipitate, deeply cupulate, grey to beige coloured,
sometimes yellowish, ochraceous, rarely orangish, with an external
surface pustulate at least when young; the hymenium is whitish or
concolourous to the outer surface; the margin is often eroded or
denticulate. Microscopically, the asci are operculate, often slightly
curved at the top, inamyloid, without crozier. The paraphyses are
septate, simple or ± diverticulate at the top and contain nuclei stain-
ing red in acetocarmine solution. The ascospores are smooth or
warted in one species, biguttulate, often accompanied by numerous
internal droplets; this content is generally merged into one large oil
drop on rehydrated material. The flesh is composed of two layers: a
thick medullary excipulum of textura intricata and an ectal
excipulum of textura globulosa to angularis; the outer part is often
composed of aggregates of roundish cells which form macroscopic
warts. Marginal cells are generally made of textura prismatica. Finally,
the spore print is white.
Concerning the diagnostic characters for species, the size of
apothecia is an important criterion. The presence of a stipe is also
to be noted but its length can vary within the same collection and
it can be buried in the substrate. Microscopically the size and shape
of mature ascospores are important, but we believe that the spore
content in the living state must be also considered because on re-
hydrated material the oil drops are merged. The top of the paraphy-
ses was often used as an important diagnostic character —
presence of diverticulate items or branchlets or bumps — to sepa-
rate some species, but the situation seems more complex because
we observed that this feature can vary within the same collection,
depending on the age of the specimen or the area of hymenium ob-
served. Thus, it does not appear to be a stable character. Finally, the
ecology appears to be preponderant for many species and should
be noted when collecting samples although it is not always easy to
determine the possible tree (or plant) association.
Phylogenetic results
The analyses done on LSU and ITS sequences, respectively shown
on Fig. 1 and 2, produced similar topologies. The monophyly of the
genus Tarzetta is confirmed, including the monospecific genus Hy-
potarzetta Donadini. The main clades are strongly supported and
give useful information on the hypothesized host-specificity of
several taxa.
The clade /tarz-1 (Fig. 1) is represented by a collection made in
Chile (FH:DHP-CH30, see KUMAR et al., 2017) and an environmental
sequence — with Nothofagus obliqua as host (JX31634) — from Ar-
gentina. On Fig. 2, a similar clade is represented and the collection
FLAS:MES-2162 may represent another species, also associated with
Nothofagus (N. antarctica or N. dombeyi based on the collector data).
These collections should be compared with Tarzetta microspora
(Raithelh.) Raithelh. (= Pustularia microspora Raithelh., 1983), a
species described from Argentina. Tarzetta jafneospora probably
also associated with Nothofagus spp., appears in its own clade.
The clade /tarz-2 (Fig. 1 only) corresponds to T. jafneospora, an en-
demic species from New Zealand. Tarzetta jafneospora could be as-
sociated with Nothofagus spp., a dominant canopy tree. As a “sister
branch, there is a new species, T. gregaria (see under this name in
the chapter on Taxonomy).
The clade /tarz-3 comprises the new species T. alpina (see under
this name), and two North-American collections, one tentatively de-
termined as T. bronca (Peck) Korf & J.K. Rogers and another one
probably new to science, growing in an oak dominated woodland,
with Q. alba and Q. macrocarpa (HEALY, pers. comm.).
The clade /tarz-4 comprises two robust subclades: 1) Hypotarzetta
insignis (Berthet & Riousset) Donadini, a semi-hypogeous species;
this demonstrates that Hypotarzetta is nested within the Tarzetta lin-
eage and the correct name is Tarzetta rather than Hypotarzetta as
proposed by DONADINI (1985). Originally the species was published
in the genus Pustularia, the genus name used by mycologists before
the resurrection of Tarzetta. 2) Two species, i.e. T. pusilla and T. sepul-
tarioides sp. nov., discussed in the next chapters.
The clade /tarz-5 is divided into two subclades: ‘5a’ corresponds
to the sequences identified as Tarzetta ochracea (see next chapter);
‘5b’ contains a group of Mediterranean taxa in which we describe
T. quercus-ilicis, a new species associated with Quercus ilex. See the
comments under this name. Another collection, from Colorado,
USA, determined as T. cupularis, occupies an isolated position within
/tarz-5 and requires further investigation.
The clade /tarz-6 has been chosen to represent Tarzetta cupularis
(see discussion in the next chapter). A collection made in Malta oc-
cupies an isolated position and is described herein as T. melitensis
sp. nov.
The clade /tarz-7 has been chosen to represent Tarzetta catinus
(see discussion in the next chapter). It forms a complex of species
with large apothecia, mainly associated with Fagus spp. More genes
are required to circumscribe the different species. We tentatively de-
limited two collections made under Betula, the first one determined
as T. spurcata (AMNH:44124) and the second one under the provi-
sional name T. betulicola (see under this name).
Based on the hypothesis that collections represented in the /tarz-
7 include the original concept of Tarzetta catinus, the clade /tarz-8
may represent a complex of species often determined as T. catinus.
Within this complex, the subclade 8a is named T. alnicola sp. nov., a
species associated with alders. The host tree for other collections,
provisionally named T. pseudocatinus, is not clearly identified (Carpi-
nus betulus is noted for 3 collections, Corylus avellana = 1, Fagus syl-
vatica = 2, Pinus sp. = 1, Castanea sativa = 2, Quercus sp. = 1,
Crataegus sp. = 1 and “hardwoods” = 1). The sequenced collections
do not differ significantly from T. catinus and we do not find any dif-
ference in spore size or shape. Further investigations are required
to explore this species complex and circumscribe correctly the
species where there is unambiguous evidence to. One of these taxa
appears to be strictly associated with Corylus spp. (represented by
the collection LY:NV 2018.06.12 and an ITS sequence from an envi-
ronmental sample, GenBank JX474834).
Finally, one of our collections (LY:NV 2014.07.10) is included in the
clade /tarz-9 with two environmental ITS sequences (Fig. 2), one iso-
lated from roots of Quercus liaotungensis, in China (GenBank
KX444378), and one isolated from soil in the Arctic part of Alaska
(GenBank KC966151; TIMLING et al., 2014). Our collection comes from
the French Alps, at 1190 m elev., and was growing along a small
brook, close to Betula and Salix trees. Nothing can be here con-
cluded, and more collections are needed.
Review of existing names and typifications
Tarzetta catinus (Holmsk.) Korf & J.K. Rogers, in Korf, Phytologia,
21 (4): 206 (1971). Plates 1–2 and fig. 5A.
Basionym: Peziza catinus Holmsk., Beata ruris otia fungis danicis,
2: 22 (1799); sanctioned by Fries, Syst. mycol., 2: 61 (1822).
Original diagnosis: stipitata, globoso-campanulata; limbo
patente, repando; superficie externa pallide ochracea, punctis fuscis
notata, interna glabra, ex fusco ochracea.
Lectotype here designated: Plate 8, HOLMSKJOLD (1799); MBT
Homotypic synonyms: Peziza sphacelata subsp. catinus (Holmsk.)
Pers., Mycol. Europ., 1: 231 (1822); Pustularia catinus (Holmsk.) Fuckel,
Jahrb. Nass. Verein. Naturk., 23-24: 328 (1870); Aleuria catinus
(Holmsk.) Gillet, Champ. Fr., Discom.: 39 (1879); Geopyxis catinus
(Holmsk.) Sacc., Syll. fung., 8: 71 (1889); Pustulina catinus (Holmsk.)
Eckblad, Nytt Mag. Bot., 15: 84 (1968).
Taxonomical synonyms: Peziza tarzetta Cooke, Mycographia, 1:
166 (1877); Discina pallida Velen., Česke Houby, 4-5: 860 (1922), fide
SVRČEK (1979).
Peziza catinus is lectotypified by the beautiful plate depicted by
HOLMSKJOLD (1799, pl. 8). It is part of the material cited by FRIES (1822)
in his sanctioning publication Systema mycologicum. This plate
shows several specimens at different stages of development. This
species has a rather long stipe compared to the other species and
large apothecia. FRIES (op. cit.) indicated: “Stipes praecedentis longior,
constanter praesens, firmus, 2 lin. fere crassus, subglaber. Cupula [...]
1-2 unc. lata.” This suggests that the stipe usually measures about
4 mm diam. and the apothecium is between 25 to 50 mm1diam.
The plate 2 illustrates an authentic Fries’ collection housed in UPS
which perfectly matches his description. This concept was followed
by BOUDIER (1905-1910, under Pustularia catinus), NANNFELDT (1938,
under P. catinus), DENNIS (1978), DISSING et al. (2000), YAO & SPOONER
(2002) and VAN VOOREN (2014). Our phylogenetic analyses suggest
this concept is not homogeneous regarding the morphology. The
designation of an epitype based on a recent and well-documented
collection coming from Denmark, near Aarhus (where the original
“collection” was found) is urgently needed.
1lin. = linea, which means “line”, an ancient unit of length. The value was not the same in all countries, but in Sweden before 1879 one line
corresponded to about 2 mm after J. Melot (comm. pers.). The same applied to “unc. = uncia” which means “once” (inch), about 25 mm.
Plate 1 – Tarzetta catinus.A: Lectotype of Peziza catinus from HOLMSKJOLD (1799, pl. 8). B: Coll. G.M. 2019 05 05; photo G. Moyne. C: Coll. V.R.
20190509; photo V. Ricard. D: Coll. LY:NV 2007.04.18 (young specimens); photo N. Van Vooren. E: Coll. AH:53705; photo L. Sánchez.
Plate 2 – Tarzetta catinus.Collection of Peziza catinus from Fries herbarium. a: dried specimens. b: close-up view of of a specimen (scale
bar = 20 mm). c: Label. d: Nannfeldt’s label. All photos Åsa Kruys (UPS).
Waiting for the opportunity to study such collection, we select
the collection C:F-55260 ex KS-94-10 [GenBank ITS DQ200833; LSU
DQ062984; RPB2 JX943768] from Denmark (TEDERSOO et al., 2006;
HANSEN et al., 2013) to represent T. catinus. It likely forms ectomycor-
rhizae with beech (Fagus sylvatica). This hypothesis seems to be cor-
rect based on the other collections or EcM data of the /tarz-7 clade
shown in Fig. 1 & 2.
Tarzetta pseudocatinus (provisional name; Plate 3 and fig. 5D) is
morphologically very close to, but molecularly distant from T.
catinus. The morphological criteria to distinguish them could be
hard to establish. The ecological data may prove helpful once the
correct hosts have been identified. It is also possible that this taxon
has developed interactions with several plants.
Studied material: Tarzetta catinus. – FRANCE: Isère, Lans-en-Ver-
cors, Col de la Croix Chabaud, 45.10576° N 5.54601° E, 1215 m a.s.l.,
on soil, under Fagus sylvatica and conifers, 15 Jun. 2002, leg.
M. Poizat, herb. NV 2002.06.05 (LY). Aude, Belcaire, Clos de la Plaine,
42.841328° N 1.954536° E, 980 m a.s.l., on soil, under Picea abies,
Fagus sylvatica and Corylus avellana, 30 Apr. 2007, leg. N. Van Vooren,
herb. NV 2007.04.18 (LY). Loire, Saint-Georges-en-Couzan, pont du
Diable, 45.6954455° N 3.9356588° E, 560 m a.s.l., on soil, under Fagus
sylvatica, 5 May 2010, leg. N. Van Vooren, herb. NV 2010.05.09 (LY).
Doubs, Amancey, le Grand Bois, ~47.03925° N 6.08879° E, 595 m
a.s.l., on soil, under Fagus sylvatica and Quercus sp., 21 May 2019, leg.
G. Moyne, pers. herb. G.M. 2019 05 05. Haute-Marne, Ormancey,
forêt domaniale, 47.919962° N 5.160481° E, 430 m a.s.l., on soil under
Fagus sylvatica, Quercus sp. and Carpinus sp., 9 May 2019, leg. V. Ri-
card, pers. herb. V.R. 20190509-03. SPAIN: Catalonia, Riu de Cerdanya,
Prat de les Cabanes, 42.323675° N 1.8184611° E, 1550 m a.s.l., along
a stream, on soil, under Abies sp., Salix caprea and Corylus avellana,
28 Jun. 2014, leg. L. Sánchez, herb. 53705 (AH). Tarzetta
pseudocatinus agg. – FRANCE: Rhône, Les Haies, Col de Croix-Régis,
45.52189° N 4.74278° E, 500 m a.s.l., on soil, under hardwoods, 1 May
2000, leg. H. Orcel, herb. NV 2000.05.01 (LY). Rhône, La Tour de Sal-
vagny, Cerqueminal, 45.8077383° N 4.7014671° E, 275 m a.s.l., on
soil, under Carpinus betulus and Quercus sp., 4 Sep. 2006, leg. N. Van
Vooren, herb. NV 2006.09.01 (LY). Rhône, Saint-Priest, bois de Feuilly,
45.71302° N 4.92475° E, 210 m a.s.l., on soil, under Castanea sativa
and Carpinus betulus, 7 Nov. 2013, leg. J. Cavet, herb. NV 2013.11.09
(LY). Hautes-Pyrénées, Aulon, Granges de Lurgues, 42.85388° N
0.27167° E, 1445 m a.s.l., on soil, under hardwoods, mainly Corylus
avellana, 26 Aug. 2014, leg. N. Van Vooren, herb. NV 2014.08.19 (LY).
Seine-et-Marne, Vaux-le-Pénil, parc Faucigny-Lucinge, 48.5367° N
2.66905° E, 80 m a.s.l., on soil in an urban park, under Carpinus betu-
lus, 29 May 2019, leg. R. Chalange, herb. NV 2019.05.12 (LY). ITALY:
Caserta, Piano della Corte, ~41.463333° N 14.394166° E, 1400 m a.s.l.,
on soil, under Fagus sylvatica, 20 Oct. 2012, leg. A. Conte, pers. herb.
C. Agnello C.A. 20121020.1. Lombardy, Milan, Paderno Dugnano,
Parco Toti, 45.5715° N 9.175333° E, 170 m a.s.l., on soil, under Cratae-
gus sp., 21 May 2018, leg. E. Sacchi, pers. herb. M.C. 18-5-21 (dupli-
cata in LY). Liguria, Ruta di Camogli, 44.348944° N 9.17775° E, 245
m a.s.l., on soil, under Castanea sativa and Quercus sp., 13 Oct. 2018,
leg. M. Carbone, pers. herb. M.C. 18-10-13 (duplicata in LY).
Tarzetta cupularis (L.) Lambotte, Mém. Soc. roy. sci. Liège, série 2,
14: 325 (1888). Plate 4 and fig. 5B.
Basionym: Peziza cupularis L., Syst. Plant., 2: 1181 (1753); sanc-
tioned by Fries, Syst. mycol., 2: 62 (1822).
Type: “neotype” designated by PANT & TEWARI (1971), collection
Fuckel’s Fungi Rhenani no. 1878, housed in Royal Botanic Gardens,
Kew (K). See comments.
Lectotype here designated: Bot. paris., pl. 11, fig. 1-3, VAILLANT
(1727); MBT 389443.
Original diagnosis: Peziza globoso-campanulata: margine cre-
Peziza calice campanulato globoso: ore crenato. Guett. Stamp. 16.
Dalib. paris. 388.
Fungoides glandis cupulam referens, margine dentato. Vaill. paris.
57. t. 11. f. 1. 2. 3.
Habitat in Gallia.
Homotypic synonyms: Pustularia cupularis (L.: Fr.) Fuckel, Jahrb.
Nass. Verein. Naturk., 23-24: 328 (1870); Aleuria cupularis (L.) Gillet,
Champ. Fr., Discom.: 39 (1879); Geopyxis cupularis (L.) Sacc., Syll. fung.,
8, p. 72 (1889); Pustulina cupularis (L.) Eckblad, Nytt Mag. Bot., 15: 85
Taxonomical synonyms: Peziza crenata Bull., Herb. Champ. Fr., pl.
396, fig. III (1786); Geopyxis grossegranulosa Velen., Novit. Mycol. No-
vissim.: 152 (1947), fide SVRČEK (1979).
PANT & TEWARI (1971) designated a neotype based on Fuckel’s
Fungi Rhenani no. 1878 collection (from Hessen in Germany),
housed in Royal Botanic Gardens, Kew (K). Unfortunately, this des-
ignation is not acceptable because the authors did not respect the
rules for typification, especially Art. 9.7 and 9.10 of ICN. As the
species was originally described by LINNÉ (1753) from France, we des-
ignated a lectotype based on Vaillant’s drawing (1727), listed by
LINNÉ (op. cit.) and by FRIES (1822) in agreement with Art. 9.10 for
sanctioned names. This drawing is here reproduced on the plate 3.
Of course, this lectotype is not sufficient to clearly circumscribe this
name because no microscopical or molecular data are available
from this original material. The species is often considered as a ses-
sile or short-stipitate species (as illustrated by VAILLANT, 1727). FRIES
(1822) indicates the ascocarps are “3–8 lin. lata”, which is approxi-
mately 6.8–18 mm, and is thus considered as a rather small species
for the genus. A French collection made near Versailles2(or in the
south of Paris), growing under hardwoods and possessing such
characters is actively searched to provide a modern definition of this
taxon and designate an epitype. In the meantime, we have selected
a clade containing collections which fit well this definition. In this
clade, the collections are probably associated with oaks (Quercus
spp.) and show a wide distribution (presence in Europe and USA).
Tarzetta scotica (Rea) Y. J. Yao & Spooner is considered by YAO &
SPOONER (2002) as a species close to T. cupularis mainly differing by
smaller ascospores. After the original description (REA, 1928), this
species is sessile. No information is given about the ecology of this
taxon first collected near Perth, on the eastern coast of Scotland. We
did not yet have the opportunity to study T. scotica and evaluate its
characters to make a conclusion.
Studied material: FRANCE: Rhône, La Tour-de-Salvagny, les Engro-
lets, 45.8025517° N 4.7011771° E, 310 m a.s.l., on burnt ground,
under Quercus sp. and Pinus sylvestris, 21 Oct. 2004, leg. N. Van
Vooren, herb. NV 2004.10.38 (LY). Rhône, Saint-Romain-au-Mont-
d’Or, Mont Thoux, 45.83935°N 4.807991° E, 420 m a.s.l., on soil, under
Quercus sp. and Buxus sempervirens, 24 Oct. 2006, leg. N. Van Vooren,
herb. NV 2006.10.31 (LY). Rhône, La Tour-de-Salvagny, Cerqueminal,
45.8077383° N 4.7014671° E, 275 m a.s.l., on soil, under Carpinus be-
tulus, Quercus sp. and Castanea sativa, 31 May 2019, leg. N. Van
Vooren, herb. NV 2019.05.11 (LY). PORTUGAL: Viseu, Parque de Fontelo,
40.659155° N 7.898876° W, 475 m a.s.l., on soil under hardwoods, 22
Nov. 2013, leg. N. Van Vooren, herb. NV 2013.11.24 (LY). ITALY: Pied-
mont, Variano, 44.7202778° N 8.938611° E, 330 m a.s.l., on soil, under
Quercus pubescens and Castanea sativa, 26 May 2018, leg. M. Car-
bone, duplicata herb. M.C. 18-5-26. Liguria, Mezzema, Deiva Marina,
44.252583° N 9.548555° E, 500 m a.s.l., on soil, under Quercus ilex, 23
May 2008, leg. M. Carbone, pers. herb. M.C. 08-5-23.
Tarzetta gaillardiana (Boud.) Korf & J.K. Rogers, in Korf, Phytolo-
gia, 21 (4): 206 (1971). Fig. 3.
Basionym: Pustularia gaillardiana Boud., Bull. Soc. mycol. France,
18: 141 (1902).
Type: holotype in PC, ex herb. Boudier; isotype, no. 28 in herb.
Gaillard (Musée botanique d’Angers, Arboretum).
Original diagnosis: Pustularia gaillardiana Boud. (Pl. 8, fig. 1).
Minuta, 4–8 mm lata, cupularis, cinereo-fulva, brevissime stipitata
aut sessilis, extus parce tomentella margine subcrenulato. Receptacu-
lum cupulare, minute tomentello-granulosum, margine subcrenulato,
brevissime stipitatum, griseo-fuscum, hymenio concolore laeve. Thecae
cylindricae, ad basim sub-attenuatae, hyalinae, operculatae, octospo-
2In the original VAILLANT’s text, this author said: “Il croist dans les bosquets de Versailles vers la fin d’Aoust & le commencement de Septembre.”
which means “It grows in the groves of Versailles from the end of August to the beginning of September.”
Plate 3 Tarzetta pseudocatinus agg.A: Coll. LY:NV 2013.11.09; photo N. Van Vooren. B: Coll. LY:NV 2014.08.19; photo N. Van Vooren. C: Coll.
LY:NV 2019.05.12; photo M.-P. Vigneron. D: Tarzetta cf. catinus - coll. M.C. 18-10-13; photo M. Carbone.
rae, 300–320 µ longae, 15–20 crassae. Paraphyses tenues, hyalinae,
septatae, intus vix granulosae ad apicem paululum incrassatae, 3–4 µ
spissae. Sporae majores, ellipticae, laeves, intus guttulis duobus oleosis,
majoribus et granulis primo sat numerosis dein maturitate rarioribus
aut deficientibus repletae. Long. 25–27 µ. Latitud. 13–14.
Angers, ad terram arenosam in nemoribus legit Novembre 1901
clar. D. Gaillard.
Homotypic synonyms: Geopyxis gaillardiana (Boud.) Sacc. &
D. Sacc., Syll. fung., 18: 15 (1906); Pustulina gaillardiana (Boud.) D.C.
Pant & V.P. Tewari, Mycologia, 62 (6): 1191 (1971).
Taxonomical synonyms: Geopyxis albocinerea Velen., Novit.
Mycol. Novissim.: 152 (1947), fide SVRČEK (1979); Geopyxis cavinae
Velen., Česke Houby, 4-5: 859 (1922), fide SVRČEK (1979); Geopyxis
cupularis Velen., Česke Houby, 4-5: 859 (1922), fide SVRČEK (1979);
Geopyxis patellaris Velen., Monog. Discom. Bohem.: 336 (1934), fide
SVRČEK (1979).
The isotype has been reviewed and possesses ascospores shorter
than those described by BOUDIER (1902), † (20) 21–24 × (11) 11.5–
14 µm vs. 25–27 × 13–14 µm, with a Q value between 1.6 and 2.1
(mean 1.8). PANT & TEWARI (1970) revised the type collection housed
in PC and they found ascospores measuring 13–19 × 6.5–11 µm
(sic!), but they do not discuss this important difference with the pro-
logue. The desiccation process cannot explain such a difference in
spore length. We cannot exclude a difference of maturity between
the specimens examined by Boudier or the fact he measured some
Plate 4 Tarzetta cupularis.A: Lectotype of Peziza cupularis from VAILLANT (1727, pl. 11, fig. 1-3). B: Coll. LY:NV 2019.05.11; photo N. Van
Vooren. C: Coll. M.C. 08-5-23; photo M. Carbone. D: Coll. LY:NV 2004.10.38 (on burnt ground); photo N. Van Vooren. E: Coll. M.C. 18-5-26;
photo M. Carbone. D: Coll. LY:NV 2012.05.43 (on burnt ground).
overmature ascospores. Unfortunately, our attempts to obtain a
DNA sequence from the isotype failed.
SVRČEK (1979) listed several Velenovský’s names as synonyms of
T. gaillardiana but it seems difficult to evaluate their relevance with-
out a reexamination of these collections or a DNA analysis.
Finally, new collections3from the type locality, near Angers
(France), are required to fix a modern interpretation of this species.
Studied material: FRANCE: Maine-et-Loire, Saint-Sylvain-d’Anjou,
“La Baronnerie”, 15 Nov. 1901, leg. D. Gaillard, coll. no. 28 in herb.
Gaillard, isotype (Musée botanique d’Angers, Arboretum).
Tarzetta insignis (Berthet & Riousset) Korf & J.K. Rogers, in Korf,
Phytologia, 21 (4): 206 (1971). Plate 5.
Basionym: Pustularia insignis Berthet & Riousset, Bull. Soc. mycol.
France, 79: 397 (1963).
Original diagnosis:Apotheciis primum cupulatis pedunculatisque,
deinde subsessilibus, late exaplanatis, exterius convolutis, saepe pli-
catis, 6–12 cm maturescentibus. Margine semper reflexa, primum in-
tegra, deinde incisa. Hymenio pallido, albo-flavescente. Facie externa
concolore, sed paulum atriore, dense furfurácea. Carne tenue, firma,
flexile, alba. Ascis longe cylindratis, octosporis, 390–410 × 18–20 µ.
Sporis ellipticis, laevibus, maturitate duabus magnis guttulis fere totis
impletis, 25–27 × 14–15 µ. Paraphysibus filiformibus, linearibus, sep-
tatis, saepe, ramosis, anastomosantibus, ascos longitudine aequan-
tibus, 2,5–3 µ latis. Sparsis in terra argilacea humosa, in cedreto. Typo
in Ly sub n° 346.
Type: holotype in LY, ex herb. Berthet PB 346.
Homotypic synonyms: Pustulina insignis (Berthet & Riousset)
Korf & Berthet, Natur. Canad., 96: 248 (1969); Hypotarzetta insignis
(Berthet & Riousset) Donadini, Doc. mycol., 15 (60): 49 (1985).
We sequenced one of our collections coming from the type local-
ity in the French Massif of Luberon (BERTHET & RIOUSSET, 1963). Our
analysis confirms the placement in Tarzetta and the combination in
the monotypic genus Hypotarzetta by DONADINI (1985) is therefore
superfluous. This species was initially associated with cedar trees,
i.e. Cedrus atlantica, but it was recently discovered in Spain under
pine trees, i.e. Pinus sylvestris (MATEO FERNÁNDEZ, 2011) and in north-
eastern France under Douglas fir, Pseudotsuga menziesii (SARRAILLON
& DOLL, 2013). The latter collection was sequenced and its ITS is
100% identical to the ITS obtained from the “topotype” collection.
This result suggests T. insignis is not exclusively associated with
cedar trees, but also with other Pinaceae species.
T. insignis possesses the largest ascospores of the European
Tarzetta, sometimes reaching 29 µm in length.
Ascomata initially hypogeous, emerging when mature. Apothe-
cia 45–85 mm diam., sessile or very shortly stipitate, irregularly
cupuliform or strongly deformed, with a whitish, creamy or yellow-
ish grey, velvelty hymenium; external surface concolorous, strongly
pustulate. Margin curved, eroded. Flesh thick, brittle.
Medullary excipulum 800–1100 µm thick, of textura intricata,
with hyaline hyphae. Ectal excipulum rather thick, ~250 µm, made
of clavate cells, 17–35 µm wide, mixed with some globose, subglo-
bose or angular elements; presence of emerging hyaline hyphae,
septate, reaching 130 µm in length. Asci (330) 370–400 × 18–20 µm,
cylindrical, long narrowed to the base, without crozier, operculate,
inamyloid, 8-spored. Paraphyses filiform, 3–4 µm diam., not en-
larged at the top, straight or flexuous, simple or sometimes forked,
often with lateral bumps or appendages, hyaline. Ascospores (22)
23–27 (29) × (12.5) 13–17 µm [X = 25.3 × 13.9 µm, n=51], Q=1.6–2.1
(2.3), Qm=1.8, ellipsoid, sometimes slightly tapered at the ends, hya-
line, smooth, rather thin-walled, containing two large oil drops, ac-
companied with numerous small guttules.
Studied material:FRANCE: Vaucluse, Bonnieux, massif des Cèdres,
43.8011115° N 5.2712607° E, 705 m a.s.l., on soil, under Cedrus at-
lantica and Buxus sempervirens, leg. N. Van Vooren, 14 Apr. 2007,
herb. N.V. 2007.04.08 (LY); idem, under C. atlantica and Pinus sp., leg.
D. Borgarino, 3 May 2008, herb. N.V. 2008.05.02 (LY); idem, leg. N. Van
Vooren, 2 May 2009, herb. N.V. 2009.05.02 (LY); idem, leg. N. Van
Vooren, 16 May 2010, herb. N.V. 2010.05.23 (LY); idem, leg. N. Van
Vooren, 16 May 2010, herb. N.V. 2010.05.23 (LY); Lacoste, massif des
Cèdres, 43.80149° N 5.26823° E, 710 m a.s.l., on soil, under C. at-
lantica, leg. N. Van Vooren, 21 Mar. 2014, herb. N.V. 2014.03.07 (LY);
Haut-Rhin, Osenbach, Bickenberg, 560 m a.s.l. on soil, under Pseudot-
suga menziesii, leg. D. Doll, 9 Jun. 2013, duplicata herb. F.S. 2013002.
GREECE: West Macedonia, Grevena, Ziakas-Perivolaki, 40.035378° N
21.245003° E, 950 m a.s.l., on soil, under Abies borisii-regis, leg. G. Kon-
stantinidis, 31 Mar. 2007, duplicata herb. G.K. 2094.
Tarzetta ochracea (Boud.) Van Vooren, comb. nov. – MB 833284.
Plate 6 and fig. 5E.
Basionym: Pustularia ochracea Boud. ex Boud., Icon. mycol., sér. I,
livr. 2, pl. 337 (1904), a new name based on Peziza ochracea Boud.,
in Cooke, Mycographia, 5: 225 (1879), illeg.
Type: no type specimen is housed in Cooke’s herbarium at K
(DAVIES, pers. comm.).
Fig. 3 – Tarzetta gaillardiana (isotype). A: Ascospores. B: Paraphyses
and ascus. Scale bars = 10 µm. Drawing N. Van Vooren
3A collection cited in MORNAND & PÉAN (1995) was revised and corresponds to a Sepultaria species (confirmed by a DNA sequence).
Lectotype here designated: Fig. 377, Peziza ochracea, COOKE
(1879); MBT 389444.
Epitype here designated: France, Essonne, forêt de Rougeau, LY:
NV 2019.06.01; MBT 389445.
Original diagnosis: Caespitosa, integra, subflexuosa, laete
ochracea, extus furfurácea. Ascis cylindraceis. Sporidiis ellipticis, binu-
cleatis, hyalinis. Paraphysibus linearibus, supra in processibus brevibus
Although the synonymy between Pustularia ochracea and Tarzetta
spurcata was accepted by authors like NANNFELDT (1938), KORF (1986)
or YAO & SPOONER (2002), we believe they must be separated (see our
discussion under T. spurcata).
This species differs from Tarzetta catinus by the absence of a true
stipe, although some specimens present a furrowed base, some-
times elongated that resembles a pseudostipe (Pl. 5, fig. C and F),
embedded in the substrate. This is a large species reaching 50 mm
in diameter and showing a typical ochraceous colour, especially on
the oldest specimens. The ascospores measure 20–25 × 11–13 µm
after BOUDIER (1904), and 19–24 × 11–13.5 (14) µm after our collec-
tions. This species was originally described from the Montmorency
forest, in the Paris area, which is mainly composed by chestnut trees
(Castanea sativa) at 70%, oaks (Quercus sp.) at 11% and beech (Fagus
sylvatica) at 2% [source: ONF]. COOKE (1879) — who figured the
species from specimens and drawing sent by Boudier — indicates:
“On the ground in beech woods”. Therefore, we hypothesize it may
be associated with Fagaceae.
Plate 5 Tarzetta insignis.A: Coll. LY:NV 2014.03.07. B: Ascospores from coll. LY:NV 2010.05.23. C: Coll. LY:NV 2007.04.08. D: Coll. LY:NV
2009.05.02 (detail of the outer surface). All photos N. Van Vooren.
We designate an epitype with a recent collection made close to
Paris (Pl. 5, fig. B) which fits morphologically well Boudier’s concept
of this species. Based on our data, the species is widely distributed
in Europe.
NANNFELDT (1938) — followed for example by SCHUMACHER (1979)
— suggested Pustularia ochracea to be a synonym of Tarzetta
catinus, considering there exists intermediates between typical
forms of both species. As the two species share similar habitats, this
idea may seem seductive, but it does not reflect the diversity ex-
pressed through molecular data.
BRESADOLA (1892) considered Peziza ochracea and P. spurcata as
later synonyms of his Aleuria pustulata, based on Octospora pustu-
lata Hedw. Although the species described and illustrated by HEDWIG
(1789) can figure a Tarzetta (see Pl. VI, fig. 1–2, showing pustulate
apothecia, and fig. 5 biguttulate ascospores), it seems difficult to de-
fine if this name can apply to T. ochracea or not (or to another known
species) without authentic material.
Studied material: FRANCE: Essonne, Saint-Pierre-du-Perray, forêt
de Rougeau, ~48.587559° N 2.5167034° E, 85 m a.s.l., on soil, under
Quercus sp. and Populus tremula, leg. J.-C. Tinlot, 4 Jun. 2019, herb.
N.V. 2019.06.01 (LY), epitype. Doubs, Labergement-Sainte-Marie, in
a private garden, 46.775665° N 6.280288° E, 867 m a.s.l., on soil,
under Fagus sylvatica, leg. C. Page, 5 Jun. 2018, herb. N.V. 2018.06.11
(LY). Labergement-Sainte-Marie, forêt domaniale du Mont Sainte-
Marie, near the natural reserve of Remoray, ~46.78233° N 6.261407°
E, 960 m a.s.l., in a mixed wood (Picea abies, Fagus sylvatica), leg.
C. Frund, 18 Jun. 2018, pers. herb. TaCu18061801, duplicata N.V.
Plate 6Tarzetta ochracea.A: Coll. LY:NV 2018.06.05; photo B. Capoen. B: Coll. LY:NV 2019.06.01 (epitype); photo J.-C. Tinlot. C: Coll.
TaCu18061801; photo C. Frund. D: Coll. LY:NV 2018.06.11; photo C. Page. E–F: Coll. M.C. 15-6-20; photos M. Carbone.
2018.06.49 (LY). SPAIN: Asturies, Castro, road to Pineda, 43,13624° N
6.26573° W, 665 m a.s.l., on soil, under Castanea sativa and Tilia sp.,
leg. N. Van Vooren, 5 Jun. 2013, herb. N.V. 2013.06.12 (LY). SWITZER-
LAND: Fribourg, Montagny, les Arbognes, forêt de Berley, 46.8124525°
N 7.0054741° E, 525 m a.s.l., on soil, under Fagus sylvatica, leg.
R. Dougoud, 30 Jun. 2013, herb. N.V. 2013.06.43 (LY). Bern, Tramelan,
les Embreux, 47.262777° N 7.128583° E, 1085 m a.s.l., on soil, under
conifers and Fagus sylvatica, leg. B. Capoen, 15 Jun. 2018, herb. N.V.
2018.06.05 (LY). ITALY: Piedmont, Parco natural della Alta Valle Pesio
e Tanaro, Pian delle Gorre, 44.2146667° N 7.6608333° E, 1040 m a.s.l.,
on soil, under Abies alba mixed with Fagus sylvatica, leg. M. Carbone,
20 Jun. 2015, pers. herb. M.C. 15-6-20 (duplicata in LY). FINLAND: Uusi-
maa, Helsinki, Länsi-Pakila, ~60.23764° N 24.92213° E, in a garden,
on soil, leg. S. Manninen, det. H. Harmaja under T. spurcata, 29 Jul.
1970, herb. H6074351.
Tarzetta pusilla Harmaja, Karstenia, 14: 116 (1974).
Type: Finland, prov. Kuusamo, Oulanka National Park, the ravine
of the brook Tulilammenpuro, ~66.36407° N 29.444572° E, 20 Aug.
1971, leg. H. Harmaja (H 6074352, holotype) [studied].
Original diagnosis: A T. spurcata (Pers.) Harmaja apotheciis mi-
noribus et sessilibus, sporis ellipsoides et latioribus et paraphysibus cras-
sioribus, a T. catino (Pers.) Korf & Rogers apotheciis minoribus, sporis
latioribus et apicibus paraphysium valde ramosis differt.
Our revision of the holotype is in conformity with the microscop-
ical description of HARMAJA (1974) except for the size of ascospores
which is a bit smaller, † (17.5) 19–21 (22) × (10.5) 11–12 µm [Qm =
1.8, n=30] vs. 20–23 × 11.5–13 µm and the rarity of digitate para-
physes (as illustrated by the author on the right part of Fig. 2, top
row). We only observed paraphyses with a simple tip or with few
bumps, as represented on the left part of Harmaja’s fig. 2, top row.
We also observed longer ectal hairs, reaching up to 150 µm in length
vs. 100 µm in Harmaja’s description. Nothing is said about the spore
content in the fresh state and it is evident that HARMAJA (op. cit.) stud-
ied his material from exsiccata, judging by the dead ascospores rep-
resented in his Fig. 1a.
Although no associated trees are mentioned in the original de-
scription, the habitat “in moist moss on dolomitic rock and stones
near the water” and its small size suggest affinities with Tarzetta al-
nicola (this paper) but the latter has larger ascospores (especially
the width). Unfortunately, two attempts to obtain rDNA sequences
from the type collection failed due to contamination. We regard the
Hansen collection KH.03.66 (FH), from Norway, as a reference for this
species until a fresh collection from Finland can be examined, se-
quenced and eventually designated as an epitype.
Tarzetta spurcata (Pers.) Harmaja, Karstenia, 14: 119 (1974).
Basionym: Peziza spurcata Pers., Mycologia Europaea, 1: 226
Type: L 0116965, ex herb. Persoon, without mention. This collec-
tion was designated as a neotype by HARMAJA (1974) and not holo-
type, considering that it cannot be proved these specimens were
collected before Persoon’s publication.
Original diagnosis:“17. spurcata, gregaria majuscula sordide alba;
seu subfuliginea, externe farinacea, cupula margine integerrimo. Hab.
in sylvis locis adustis, sestate.
Sessilis, unc. magna, fragilis, grisea, cupulae margo quasi straitus
et magis farinaceus, sed non crenatus, uti in sequente, cui haud dissim-
As the neotype of Peziza spurcata is part of Persoon’s historical
herbarium and unavailable for DNA extraction, we morphologically
and molecularly investigated one of the studied collections cited by
HARMAJA (1974) when he combined this name within Tarzetta (coll.
H 6074351). Our revision of the latter gives the following microscop-
ical characters (in water): Medullary excipulum of textura intricata,
with yellowish hyphae. Ectal excipulum thin, of textura angularis,
with cells up to 48 µm wide; presence of emerging septate hyphae,
hyaline, reaching 170 µm in length and 3–5 µm wide. Paraphyses
not enlarged at the top, ± diverticulate. Ascospores ellipsoid, with
tapered ends, (19.5) 20–23 (23.5) × 11–13 µm, Qm = 1.8 (n=30),
hyaline, smooth, biguttulate.
Unfortunately, our attempt to extract DNA from this sample failed,
but we believe that this collection refers to T. ochracea (see under
this name).
Contrary to the opinion of HARMAJA (1974) who suggests that “the
younger Peziza ochracea Boud. in Cooke is almost surely a synonym,
we do not believe this hypothesis. The Finnish author was probably
influenced by Boudier’s drawing of paraphyses (BOUDIER, 1905-1910,
pl. 337, fig. f ), partly similar to the ones he depicted in fig. 2, lower
row (HARMAJA, op. cit.). But BOUDIER (op. cit.: 188) writes: “Paraphyses
[…] à peine épaissies au sommet qui est souvent simple, mais
quelquefois divisé en deux ou trois petits rameaux courts.” which
means “Paraphyses […] barely thickened at the top which is often
simple, but sometimes divided into two or three small and short
branchlets.Harmaja also describes a large species, reaching 45 mm
in diameter. This is the case for Pustularia ochracea — Boudier gives
a range between 20 to 50 mm — but not for Peziza spurcata in its
original sense, PERSOON (1822) indicating about 8~9 mm in diameter
(confirmed by the dried specimens of neotype, L 0116965, see plate
7). The colour given by PERSOON (op. cit.) seems also different, sordide
albagrisea” versus pale ochraceous in P. ochracea after Boudier’s
description (see also the plate 337). At last P. spurcata is described
as growing on burnt ground in a forest during summer (“in sylvis
Plate 7 – Peziza spurcata.A: Voucher of the authentic material of Peziza spurcata Pers. B: Detail of the specimens. Photos Roxali Bijmoer
(Naturalis Biodiversity Center, Leiden).
Tarzetta jafneospora W.Y. Zhuang & Korf, Mycotaxon, 28 (2): 365
(1987). Plate 8 and fig. 4.
Type: New Zealand, Gisborne, Urewera National Park – PDD
49572 (holotype); CUP 61778 (isotype).
Original diagnosis:Ab Tarzetta speciebus aliis ascosporis fusoideis
cum ornamentation cyanophilica differens.
This species is only known from New Zealand (ZHUANG & KORF,
1987) where it is considered as a rather common species, probably
associated with Nothofagus trees. It occupied an isolated position
in our phylogenetic tree of LSU sequences (Fig. 1). Morphologically
it possesses warted ascospores, a character unique in the genus. The
elongated fusoid shape of ascospores is also characteristic.
Macroscopically it can be confused with Jafnea pallida (Rodway)
G.M. Gates & Van Vooren, a species known from Tasmania, also hav-
ing fusoid and warted ascospores but of different type and asco-
carps being stipitate (GATES & VAN VOOREN, 2016).
Plate 8 – Tarzetta jafneospora.A: Coll. PDD 96264; photo J.A. Cooper (Landcare Research). B: Coll. CSAK 416; photo C. Shirley.
locis adustis, aestate”). We think that HARMAJA (1974) misinterpreted
Peziza spurcata, which must be considered different from Pustularia
ochracea Boud. For these reasons and the impossibility to trace the
original locality of Peziza spurcatawe consider this name as a nomen
Tarzetta velata (Quél.) Svrček, Česká Mykol., 35 (2): 88 (1981).
Basionym: Peziza velata Quél., C. R. Assoc. fr. Avanc. Sci., 24 (2): 621
Type: no type material exists.
Lectotype here designated: Plate VI, fig. 17, QUÉLET (1896); MBT
Original diagnosis: “Peridium cupulaire (0m,01), stipité, avec la
marge crénelée, fragile, translucide, blanc crème grisonnant, cou-
vert de grains pulvérulents brunâtres. Hyménium crème, fermé par
un voile soyeux et blanc qui s’ouvre en étoile par le développement
de la cupule. Spore ellipsoïde (0mm,015), biguttulée, hyaline. (Pl. VI,
fig. 17.)
Été. — Sur l’humus formé par les souches de hêtre, Jura. Cette
jolie peziza ne paraît pas spécifiquement différente de cupularis et
pourrait être rapportée à catinus.
Homotypic synonyms: Geopyxis velata (Quél.) Sacc. & P. Syd., Syll.
fung., 14: 744 (1899); Pustularia catinus var. velata (Quél.) Boud., Hist.
class. Discom. Eur.: 53 (1907); Pustularia velata (Quél.) Le Gal, Bull. Soc.
mycol. France, 70 (3): 200 (1955); Pustulina velata (Quél.) Svrček, Česká
Mykol., 31 (2): 70 (1977); Tarzetta cupularis var. velata (Quél.) Häffner,
Rheinl.-Pfälz. Pilzj., 2 (1): 43 (1992).
Taxonomical synonyms: Geopyxis alba Velen., Novit. Mycol. No-
vissim.: 152 (1947), fide SVRČEK (1979); Geopyxis radicans Velen.,
Monog. Discom. Bohem.: 337 (1934), fide SVRČEK (1979).
The abundant veil at the margin is the main feature that justified
the creation of this taxon by QUÉLET (1896), but several species of
Tarzetta present such a character in the young state (e.g. Tarzetta al-
nicola – see under this name), so this cannot be considered as tax-
onomically valuable.
The main work done on this species came from LEGAL (1955) who
recognized T. velata in a collection made in Switzerland in a beech
forest. Although she gave a detailed description and illustrations, it
is hard to determine which species could be applied to her collec-
tion. An examination of this collection with our modern techniques
is required to fix this name, among the taxa known to be associated
with Fagaceae. The synonyms proposed by SVRČEK (1979) — based
on Velenovský’s collections — also require investigation in the light
of our results and the heterogeneity of spore size given by the Czech
author (see Table 2).
Table 2 – Comparison of some characters of T. velata in the literature
Author Apothecia size (in mm) Spore size (in µm)
QUÉLET (1896) 10 15
LEGAL (1955) up to 8 mm (17.25) 19–22 × (11) 12–14
SVRČEK (1979), sub Geopyxis alba 10 19–22 × 11.5–13
SVRČEK (1979), sub Geopyxis radicans 7–15 20–22–25.5 × 12–14–15.5
HÄFFNER (1992), sub T. cupularis var. velata up to 27.5 mm 18–22 × 11–14
Apothecia 5–8 mm diam., sessile, cupuliform, with a greyish to
livid grey hymenium; external surface beige or brownish grey, pus-
tulate. Margin denticulate.
Medullary excipulum of textura intricata, with hyaline hyphae.
Ectal excipulum rather thin, of textura suglobulosa/angularis, with
hyaline cells, † 10–35 µm, forming in the outer part some pyramidal
cells aggregates. Marginal cells of textura prismatica. Asci † 300–
320 × 12–14 µm, cylindrical, narrowed to the base, without crozier,
operculate, inamyloid, 8-spored. Paraphyses filiform, † 2.5–3 µm
diam., not enlarged at the top, straight, simple, hyaline. Ascospores
† (20.5) 23–26 (27) × (8) 10–11.5 µm [X = 24.6 × 10.6 µm, n=30],
Q=2.1–2.7, Qm=2.3, fusoid, hyaline, rather thick-walled, containing
two large oil drops, ornamented with cyanophilic low warts, dense,
isolated or coalescent, ± angular, up to 0.5 µm wide, often concen-
trated at the poles and looking like caps.
Studied material: NEW ZEALAND: Auckland district, Hunua, Work-
man Road, Workman Track, ~37.122333° S 175.233742 E, on mossy
soil, in a Leptospermum and Nothofagus mixed forest, 14 Jun. 2015,
leg. C. Shirley, det. N. Van Vooren, duplicata of pers. herb. CSAK 416.
Tarzetta bronca (Peck) Korf & J.K. Rogers, in Korf, Phytologia, 21
(4): 206 (1971). Plate 9 and fig. 5F.
Basionym: Peziza bronca Peck, Ann. Rep. N.Y. St. Mus. Nat. Hist., 29:
54 (1878).
Type: USA, New York State – NYS F-000544 (syntype).
Original diagnosis: “Cups gregarious or crowded, sessile, sub-
hemispherical, four to nine lines broad, whitish or very pale-buff, ex-
ternally roughened by small crowded whitish warts; asci cylindrical;
spores elliptical, one to two-nucleate, .0008-.0009’ long, .0005’ broad.
Ground. Knowersville and Sandlake. July and August. (Plate 2, figs.
Homotypic synonyms: Geopyxis bronca (Peck) Seaver, N. Amer.
Cup-fungi (Operculates): 213 (1928); Pustularia bronca (Peck)
Kanouse, Mycologia, 42 (4): 497 (1950); Pustulina bronca (Peck) Korf
& Berthet, Natural. Can., 96 (2): 248 (1969).
Fig. 4 – Ascospores of Tarzetta jafneospora in Cotton Blue (coll. CSAK
416). Scale bar = 10 µm. Drawing N. Van Vooren
Plate 9 – Tarzetta bronca.Coll. SAK-17-0423-5; photo S. Krstic.
Shortly described by PECK (1878) from the state of New York, the
species was redescribed by KANOUSE (1950) based on numerous col-
lections coming from different North American localities. KANOUSE
(op. cit.), in her microscopical description, gave two different sizes
of asci and ascopores, and evoked that “heterosporous condition
was found in all of the 29 collections” and also in the type collec-
Macroscopically the species seems characterized by its yellowish
tinges of ascocarps, especially visible on outer surface of dried spec-
imens (KANOUSE, 1950: 499) but also on fresh specimens (SEAVER, 1928:
213). We selected a collection that may represent T. bronca. We
measured ascospores † (19) 20.5–22 × (10) 10.5–12 µm [X = 21.2 ×
11.2 µm, Qm=1.9, n=30] vs. 20–24 × 12–14 µm in KANOUSE (op. cit.)
and also noted the presence of numerous immature asci (so with
smaller ascospores) that may correspond to the “heterosporous con-
dition” indicated by Kanouse.
The taxonomic diversity of Tarzetta in North America requires ex-
ploration in the light of this preliminary work.
Studied material:USA, Oregon, Multnomah Co., Portland, Kelley
Point Park, ~45.6463889° N 122.7625° W, 35 m a.s.l., on soil, under
cottonwoods (Populus sp.), leg. S. Krstic, 23 Apr. 2017, duplicata herb.
SAK-17-0423-5 (as T. cupularis).
Following the results of molecular, morphological and ecological
data, we describe and illustrate here six new species, and propose
another one ad interim.
Tarzetta gregaria Van Vooren, sp. nov. – MB 833286 – Pl. 10
Diagnosis: Differs from the other Tarzetta spp. by its gregarious
habit of growth, its orangish colour, and its genetic profile.
Holotype: France, coll. NV 2017.08.16 (LY).
Etymology: From Latin “gregarius” which means “related to the
herds”, as in very numerous.
Ascomata gregarious. Apothecia 1–3 mm diam., sessile, cupuli-
form, with a beige orangish hymenium; external surface concolor-
ous, finely white-powdery. Margin eroded. Flesh thin.
Subhymenium and medullary excipulum of textura intricata,
with hyaline hyphae. Ectal excipulum of textura globulosa/subglob-
ulosa, with cells 10–32 µm diam., mixed with some clavate elements.
Marginal cells of textura subprismatica, composed of hyaline hy-
phae, 4–7 µm wide, with their top slightly emerging. Asci 310–360
× 15–17 µm, cylindrical, long narrowed to the base, without crozier,
operculate, inamyloid, 8-spored. Paraphyses filiform, 3–3.5 µm
diam., not enlarged at the top, straight or slightly curved at the top,
simple, hyaline. Ascospores (20) 21–24.5 (25.5) × (11.5) 12.5–15 µm
Fig. 5 – Ascospores of Tarzetta species. A: T. catinus (coll. LY:NV 207.04.18). B: T. cupularis (coll. LY:NV 2019.05.11). C: T. betulicola ad int. (coll.
M.P. 2018-133). D: T. pseudocatinus agg. (coll. LY:NV 2019.05.12). E: T. ochracea (coll. LY:NV 2019.06.01, epitype). F: T. bronca (coll. SAK-17-
0423-5). Scale bar = 10 µm. Drawings N. Van Vooren.
Plate 10 – Tarzetta gregaria.A: Coll. LY:NV 2017.08.16 (holotype); photo N. Van Vooren. B: idem; photo R. Dougoud.
[X = 21 × 13 µm, n=30], Q=1.1–1.3, Qm=1.2, ellipsoid, sometimes
slightly tapered at the ends or subfusoid, hyaline, smooth, rather
thin-walled, containing two large oil drops, rarely only one, with nu-
merous small guttules.
Studied material: FRANCE: Savoie, Beaufort, lac de la Gittaz, above
the dam, 45.71546° N 6.66664° E, 1675 m a.s.l., on soil, in the border
of a wood of green alders (Alnus alnobetula), leg. N. Robert, 29 Aug.
2017, herb. LY NV 2017.08.16 (holotype) and pers. herb. B. Capoen
BC250817 (isotype).
Comments: The great number of apothecia (about one hundred
specimens on a small surface) with a light orangish colour was rem-
iniscent of a Sepultariella species (VAN VOOREN et al., 2017), but the
examination of microscopical characters changed this initial super-
ficial identification. The phylogenetic analyses (Fig. 1 & 2) confirmed
the placement in Tarzetta in an isolated position. Its specific charac-
ters and the molecular data justify its publication as a new species.
ITS and LSU sequences of T. gregaria are identical to an environ-
mental sequence isolated from soil in the Arctic part of Canada
(GenBank KC966365), in a site characterized by “prostrate dwarf
shrub, herb tundra (Dryas integrifolia-Salix arctica)” (TIMLING et al.,
2014). Since no Dryas sp. or dwarf willows were growing in the site
where T. gregaria was collected, we speculate that it could be asso-
ciated with another boreo-alpine plant.
Tarzetta sepultarioides Van Vooren, sp. nov. – MB 833297 – Pl. 11
Diagnosis: Differs from other sessile Tarzetta spp. by its ascus
base often enlarged or with a protuberance (as if it was pleurorhyn-
Holotype: France, coll. NV 2017.08.03 (LY).
Etymology: From “Sepultaria, the genus name in which I first
placed the type collection, and -oides meaning “resembling”.
Apothecia 4–6 mm diam., sessile, cupuliform, with a pale beige
to pale greyish hymenium; external surface finely furfuraceous or
low-warted, pale beige to light ochraceous. Margin eroded or ±
postulate, concolorous or ochraceous.
Medullary excipulum of textura intricata, with hyaline hyphae.
Ectal excipulum of textura subglobulosa/angularis, with cells 10–
35 µm diam., hyaline to light yellow, ± organized in small pyramidal
brownish warts; presence of some emerging hyphae, septate, rather
short, 40–155 (260) × 7–8 µm. Asci 260–280× 18–21 µm, cylindrical,
long narrowed to the base, without crozier but often enlarged (as if
they were pleurorhynchous), operculate, inamyloid, 8-spored. Pa-
raphyses filiform, 5–8 µm diam., not enlarged at the top, sometimes
sublanceolate, simple, hyaline. Ascospores 22–25 (26.5) × (11.5) 12–
13 µm [X = 23.9 × 12.4 µm, n=20], Q=1.8–2.1, Qm=1.9, ellipsoid ta-
pered at the ends to fusoid, hyaline, smooth, thick-walled,
containing two large oil drops, with small guttules, the latter lacking
in mature spores.
Studied material: FRANCE: Savoie, Beaufort, la Tetaz, 45.7073° N
6.61507° E, 1000 m a.s.l., along the river Doron, on wet naked soil,
under Petasites sp., in a riparian woodland (Alnus incana, Acer sp. and
Picea abies), leg. N. Van Vooren, 27 Aug. 2017, herb. NV 2017.08.03
(LY), holotype.
Comments: On the field, Tarzetta sepultariodes was first “deter-
mined” as a Sepultaria species. A first microscopical examination
confirmed partially this feeling because of the ascospores and the
enlarged ascus base, but at the same time, the absence of true hairs
emerging from the ectal excipulum and an outer surface marked
with clustered cells excluded this hypothesis. The rDNA analysis of
the collection demonstrated this species was a member of Tarzetta.
Based on its LSU sequence our collection is 99% identical to a
Danish collection (C:F-55462) identified as T. gaillardiana. As de-
scribed by BOUDIER (1902; 1905-1910), the microscopical characters
of T. sepultarioides seem very close, but as we discussed above, the
review of the isotype of T. gaillardiana is not fully in conformity with
the protologue (especially the spore size). Moreover the species de-
picted by BOUDIER (op. cit.) is shortly stipitate and mature ascospores
content is different. For these reasons we do not believe that our
collection is T. gaillardiana.
Tarzetta alnicolaVan Vooren, sp. nov. – MB 833287 – Pl. 12
Diagnosis: Differs from other Tarzetta spp. by its small size, asso-
ciation with a habitat in riparian woodland, under alder trees (Alnus
spp.), and its genetic profile.
Holotype: France, coll. NV 2017.08.36 (LY).
Etymology: From Latin alnus, meaning alder, and cola meaning
“linked to”.
Misapplication: Tarzetta velata in VAN VOOREN (2017).
Apothecia 2.5–9 (18) mm diam., sessile or shortly stipitate, cupuli-
form, with a whitish to pale cream hymenium; external surface
beige to buff, finely pustulate with scattered small reddish-brown
tufts. Margin denticulate and a bit darker than the outer surface,
with an abundant veil in young state. Flesh thin. Stipe buried in the
substrate, sometimes with a white mycelial tomentum.
Subhymenium and medullary excipulum thick, of textura intri-
cata, with hyaline hyphae. Ectal excipulum of textura globulosa/
subglobulosa, with cells 7–30 µm diam., mixed with some clavate el-
Plate 11 – Tarzetta sepultarioides.A: Coll. LY:NV 2017.08.03 (holotype). B: Ascus base in CR. All photos N. Van Vooren.
Plate 12 Tarzetta alnicola.A: Coll. LY:NV 2017.08.36 (holotype). B: Ascospores in living state, in water (scale bar = 10 µm). C: Coll.
LY:NV 2017.08.33. D: Coll. LY:NV 2013.08.26. All photos N. Van Vooren.
Plate 13 Tarzetta alpina.A: Coll. LY:NV 2009.08.11 (holotype); photo J.-L. Cheype. B: Ascospores in living state, in water (scale bar = 10 µm);
photo N. Van Vooren.
ements, up to 65 µm wide, forming in the outer part some cells ag-
gregates, brownish-coloured. Marginal cells of textura subprismat-
ica, composed of hyaline hyphae, 6–13 µm wide, with some
marginal “hairs”, 120–180 × 6–12 µm, composed of cell chains, pale
ochre. Asci (250) 280–340 × 14–17 (20) µm, cylindrical, long nar-
rowed to the base, without crozier, operculate, inamyloid, 8-spored.
Paraphyses filiform, 3–4 (6) µm diam., straight or slightly enlarged
at the top, sometimes sublanceolate, more rarely subclavate, simple,
hyaline. Ascospores (20) 21–24.5 (25.5) × (12) 13–15 µm [X = 22.8 ×
13.9 µm, n=108], Q=1.5–1.8, Qm=1.6, widely ellipsoid, slightly ta-
pered at the ends, hyaline, smooth, rather thick-walled, containing
two large oil drops, and numerous small guttules which fill all the
space, especially in asci.
Studied material: FRANCE: Savoie, Hauteluce, l’Infernet, 45.76657°
N 6.62345° E, 1160 m a.s.l., on wet soil, along the brook Dorinet,
under Petasites hybridus, Alnus sp. and Fraxinus excelsior, 30 Aug.
2017, leg. N. Van Vooren, NV 2017.08.33 (LY). Savoie, Beaufort, la
Tetaz, 45.7073° N 6.61507° E, 1000 m a.s.l., along the river Doron, on
wet naked soil, under Alnus incana, Acer sp. and Picea abies, 31 Aug.
2017, leg. N. Van Vooren, NV 2017.08.36 (LY) [holotype]; same place,
28 Aug. 2013, leg. N. Van Vooren, NV 2013.08.26 (LY). Savoie, Beau-
fort, près du lac de la Gittaz, 45.719212° N 6.667183° E, 1620 m a.s.l.,
along a rivulet, with Alnus alnobetula in the vicinity, 24 Aug. 2013,
leg. B. Jeannerot, NV 2013.08.06 (LY). Alpes-de-Haute-Provence,
Castellane, near the camping “la Ferme”, 43.8376° N 6.54244° E,
760 m a.s.l., along the brook of Destourbes, on wet naked soil, under
Alnus sp., 9 Sept. 2017, leg. M. Vega & N. Van Vooren, NV 2017.09.08
(LY). Rhône, La Tour-de-Salvagny, Cerqueminal, 45.808192° N
4.699982° E, 280 m a.s.l., along the brook of “la Grande Rivière”, on
naked soil, under Alnus glutinosa and Fraxinus excelsior, 14 Jul. 2008,
leg. N. Van Vooren, NV 2008.07.06 (LY).
Comments: Initially we used the name T. velata for these collec-
tions made along small rivers (VAN VOOREN, 2017), partly based on
the concept of HÄFFNER (1992). The DNA analyses revealed the orig-
inality of this taxon, inside the /tarz-8 clade (Fig. 1 and 2), based on
its habitat and association with alder trees, confirmed by environ-
mental ITS sequences present in public databases (e.g. GenBank
KX897769, with Alnus glutinosa subsp. barbata; FM993170, with
Alnus sp.).
Macroscopically it is characterized by small apothecia, less than
10 mm diam. in our collections (up to 18 mm in one specimen), and
an outer surface finely marked by darker pustules. Microscopically,
it is characterized by its wide ascospores, fully filled by the two oils
drops and small guttules.
Tarzetta alpina Van Vooren & Cheype, sp. nov. – MB 833290 – Pl.
Diagnosis: Differs from other Tarzetta spp. by its non-pustulate
outer surface, becoming dark brown at the margin, its boreo-alpine
habitat and its genetic profile.
Holotype: France, coll. NV 2009.08.11 (LY).
Etymology: From Latin alpinus, the Alps, referring to its growth
at the alpine zone.
Apothecia 5–11 mm diam., sessile or substipitate, deeply cupuli-
form, with a beige to yellow beige hymenium, becoming darker
with age; external surface finely furfuraceous but not pustulate,
beige, caramel, becoming dark brown near the margin. Margin
whitish, teeth-like when young, then eroded or jagged.
Medullary excipulum of textura intricata, with hyaline hyphae.
Ectal excipulum mainly of textura globulosa, with cells hyaline to
yellowish, mixed with some clavate or subglobose elements. Asci
250–300 × 14–16 µm, cylindrical, long narrowed to the base, with-
out crozier, operculate, inamyloid, 8-spored. Paraphyses filiform, 4–
6 µm diam., not enlarged at the top, straight, simple or forked at the
middle, rarely diverticulate, hyaline. Ascospores 20.5–23 × 11–13.5
(14) µm [X = 22.1 × 12.5 µm, n=21], Q=(1.5) 1.6–2.0, Qm=1.8, ellip-
soid, tapered at the ends to subfusoid, hyaline, smooth, rather thin-
walled, containing two large oil drops, with numerous small
Studied material: FRANCE: Savoie, Aime-la-Plagne, Cormet
d’Arêches, 45.617421° N 6.608059° E, 2100 m a.s.l., on wet soil,
among alpine plants, leg. J.-L. Cheype, 27 Aug. 2009, herb. NV
2009.08.11 (LY, holotype) and pers. herb. JLC (isotype).
Comments: In the field, Tarzetta alpina can easily be confused
with a Sepultaria species. This impression is consolidated by the ses-
sile ascomata, the absence of a pustulate outer surface and the gen-
eral colour. A microscopical examination — especially on fresh
material — allows observation of the usual features of a Tarzetta.
We are not able to determine the plant association and new collec-
tions are required to propose hypotheses on this ecological charac-
T. alpina is phylogenetically identical to an unnamed Norwegian
collection (FH:KH.03.102) but we could not obtain information on
the latter to compare its characters and ecology with our own col-
lection. Another collection from Norway, sharing a similar habitat,
collected by one of us (M. Carbone) — tentatively named ‘cf. gaillar-
diana — was examined. Macroscopically it differs by the outer sur-
face covered by numerous white pustules. Microscopically it also
showed some differences: smaller ascospores, (18) 19–20.5 × 10–
11 µm, and paraphyses strongly digitate. Unfortunately, our attempt
to extract DNA failed. Finally, another collection — from Sardinia
(Italy) — is nested in the same clade of T. alpina (Fig. 1). The asco-
pores measure 21–25 × 12–13.5 µm [X = 23 ×12.8 µm, n=22],
Qm=1.8, on rehydrated material. We hesitate to place it under the
same epithet because we do not have any information on its ecol-
ogy (and biotopes in Sardinia are very different from those of boreo-
alpine regions). More genes are required to better evaluate its
taxonomical position.
Fig. 6 – Microscopic characters. A: Tarzetta gregaria (coll. LY:NV 2017.08.16, holotype); 1: Ascospores; 2: Paraphyses and ascus. B: Tarzetta
alnicola (coll. LY:NV 2013.08.26); 1: Ascospores; 2: Paraphyses and ascus. C: Tarzetta sepultarioides (coll. LY:NV 2017.08.03, holotype); 1: Pa-
raphyses and ascus; 2: Ascospores. Scale bars = 10 µm. Drawings N. Van Vooren.
Tarzetta melitensis Sammut & Van Vooren, sp. nov. – MB 833298
– Pl. 14
Diagnosis: Differs from Tarzetta cupularis and T. quercus-ilicis by
longer ascospores, a different spore shape and its genetic profile.
Holotype: Malta, coll. CS871 (LY ).
Etymology: From “Melita, the Latin name of the island of Malta
where the species was collected.
Apothecia 7–11 mm diam., sessile or shortly stipitate, deeply
cupuliform, goblet-shaped; hymenium cream-whitish; external sur-
face pustulate, grey-brown, a bit more beige near the margin, with
paler pustules. Margin distinctly crenate, with reddish brown tooth.
Medullary excipulum of textura intricata, ~350–450 µm thick,
with hyaline hyphae, 3.5–7 µm diam. Ectal excipulum of textura an-
gularis/subglobulosa, ~100–150 µm thick, with cells 11–30 × 12–
20 µm, and aggregates organized into pyramidal warts, pale
orange-brown. External hyphae scattered, 4.5–7 µm diam., origi-
nating from the ectal excipulum, simple or branched, thin-walled,
septate, hyaline. Asci 280–325 × 14–19 µm, cylindrical, long nar-
rowed to the base, without crozier, operculate, inamyloid, 8-spored.
Paraphyses filiform, slightly inflated at the top, 3–7 µm diam.,
straight, simple or occasionally forked at the base, hyaline. As-
cospores (21.3) 22.9–26.5 × 12.2–13.4 µm [X = 24.6 × 12.9 µm,
n=45], Q = 1.7–2.0, Qm = 1.9, ellipsoid with tapered ends to subfu-
soid, hyaline, smooth, rather thick-walled, containing two large oil
drops, with numerous small guttules.
Studied material: MALTA: Buskett, 35.856111° N 14.398778° E,
200 m a.s.l., on soil, under broadleaved trees, one specimen, leg.
C. Sammut, 18 Apr. 2012, pers. herb. CS368. Same location, close to
Quercus ilex and Laurus nobilis, several scattered specimens, 18 Apr.
2015, leg. C. Sammut, herb. CS871 (LY, holotype).
Comments: Initially determined as T. gaillardiana due to its small
size and colour, the molecular data confirmed the originality of this
taxon. Based on our revision of type-material of T. gaillardiana,
T. melitensis has longer ascospores, 22.9–26.5 × 12.2–13.4 µm vs. 21–
24 × (11) 11.5–14 µm.
The closest ITS sequence comes from an EcM isolate (GenBank
JF927143, 97.8% similar) from roots of Quercus pubescens in a Tuber
melanosporum truffle ground in Italy (BELFIORI et al., 2012). We hy-
pothesize that T. melitensis makes ectomycorrhizae with Mediter-
ranean oak trees like Q. ilex, Q. pubescens, etc. and being endemic
to Malta.
Tarzetta quercus-ilicis Van Vooren & M. Carbone, sp. nov. – MB
833292 – Pl. 15
Diagnosis: Differs from the Tarzetta cupularis by its sessile asco-
carp, its greyish hymenium, its probable strict association with Quer-
cus ilex and its genetic profile.
Holotype: France, coll. NV 2014.03.20 (LY).
Etymology: From Quercus ilex, the Latin botanical name of the
host tree.
Apothecia 3–15 mm diam., sessile, deeply cupuliform, spreading
at maturity; hymenium pale grey, a bit darker with age; external sur-
face postulate, especially at the margin, ochraceous grey. Margin
Plate 14 Tarzetta melitensis.A–B: Coll. LY:CS871 (holotype). C: Cells of ectal excipulum, in water (scale bar = 25 µm). D: Top of paraphyses,
in water (scale bar = 10 µm). E: Ascospores in living state, in water (scale bar = 20 µm). All photos C. Sammut.
Plate 15 – Tarzetta quercus-ilicis.A: Coll. LY:NV 2014.03.20 (holotype); photo N. Van Vooren. B: Top of ascus and paraphyses (scale bar =
10 µm), drawn from the holotype. C: Coll. M.C. 14-3-15; photo M. Carbone.
eroded or denticulate, concolorous or darker but often with whitish
Medullary excipulum of textura intricata, with hyaline hyphae.
Ectal excipulum of textura globulosa/angularis, with cells 10–30 µm
diam. or 18–60 × 11–23 µm. Asci 250–300 × 13–18 µm, cylindrical,
long narrowed to the base, without crozier, operculate, inamyloid,
8-spored. Paraphyses filiform, 4–5 µm diam., straight, hyaline, not
enlarged but irregular at the top, ± digitate or with lateral bumps.
Ascospores 20–23 (24) × (11.5) 12–13 µm [X = 21.7 × 12.1 µm,
n=30], Q= 1.6–2.0, Qm=1.8, † 20–22 (23) × 11–12.5 (13) µm [X = 21.2
× 11.9 µm, n=20], Q= 1.7–1.9, Qm=1.8, ellipsoid, sometimes a bit ta-
pered at the poles, hyaline, smooth, thick-walled, containing two
large oil drops, sometimes with small guttules.
Studied material: FRANCE: Var, La Seyne-sur-Mer, Jarnas forest,
43.07192° N 5.8536568° E, 77 m a.s.l., on soil, under Quercus ilex, leg.
F. Fouchier & N. Van Vooren, 22 Mar. 2014, herb. NV 2014.03.20 (LY,
holotype). ITALY: Apulia, Brindisi, Tuturano, Bosco Colemi,
40.5362222° N 17.93875° E, 50 m a.s.l., on soil, under Quercus ilex,
leg. M. Carbone & C. Agnello, 15 Mar. 2014, pers. herb. M.C. 14-3-15
(duplicata in LY ).
Comments: This species belongs to a small clade, named /tarz-
5b (Fig. 1 and 2), containing taxa from the Mediterranean area. The
second taxon was growing in an urban park containing typical
Mediterranean vegetation and trees (coll. LY:NV 2001.03.01, France,
Fréjus, March 2001). Its LSU sequence is close or identical to an en-
vironmental sequence (partial LSU) extracted from roots of Pinus
pinaster in Spain (GenBank JQ976016). Taking account of the de-
scription of T. melitensis, it proves that there exists a specific diversity
of Tarzetta in the Mediterranean area, adapted to the local vegeta-
T. quercus-ilicis differs from T. melitensis, probably associated with
Mediterranean oak trees, by the shape of ascospores, ellipsoid vs.
subfusoid, and their size, shorter in T. quercus-ilicis. It could also be
confused with T. cupularis which can be found under Quercus ilex,
but the latter is often short-stalked and its ascospores have a mean
size that is shorter, with a Q ratio also shorter.
Tarzetta betulicola ad int. - Pl. 16 and fig. 5C.
Etymology: From Latin betula, meaning birch, and cola mean-
ing “linked to.
Apothecia 10–40 (55) mm diam., substipitate or shortly stipitate,
deeply cupuliform, spreading at maturity; hymenium pallid beige;
external surface pustulate, greyish to beige. Margin paler, eroded
or denticulate. Stipe reaching 11 mm in length and 5.5 mm in diam.
Medullary excipulum ~700–800 µm thick, of textura intricata,
with hyaline hyphae. Ectal excipulum ~150 µm thick, of textura
globulosa/subglobulosa, with cells 13–44 µm diam., mixed with
some clavate or subglobose elements. Asci 250–300 × 14–16 µm,
cylindrical, long narrowed to the base, without crozier, operculate,
inamyloid, 8-spored. Paraphyses filiform, (3) 4–6 µm diam., not en-
larged at the top, straight, hyaline, simple or digitate, often with lat-
eral bumps. Ascospores (17) 18–21 (22) × (9) 10–11.5 (12) µm, X =
19.2 × 10.5 µm (n=54), Q= (1.6) 1.7–2.1, Qm=1.8, ellipsoid but rather
elongated to subfusoid, hyaline, smooth, thick-walled, containing
two large oil drops, with numerous small guttules, merged into one
large guttule on rehydrated material.
Studied material: ANDORRA: Soldeu, 42.575475° N 1.657945° E,
1758 m a.s.l., on soil, under Betula sp. and Pinus uncinata, leg.
M. Pélissier, 8 Jun. 2018, herb. M.P. 2018-133 (duplicata LY:NV
2018.06.49). ICELAND: Austurland, Hólarnir, Hallormsstaður, ~65.094°
N 14.71W, ~130 m a.s.l., on soil, in a birch forest, leg. Guðríður Gyða
Eyjólfsdóttir, 2 Aug. 1993, herb. AMNH FA-14527 under T. spurcata.
Comments: Based on the ecology of the two studied collections,
we think it is highly probable that this species grows in association
with Betula trees. Macroscopically it looks like T. catinus or
T. ochracea, but the ascospores are shorter and their shape is more
elongated. Molecularly the LSU sequences of the examined collec-
tions fall within the same clade as T. catinus, along with T. spurcata
suggesting that our current concept of T. catinus is a complex of
species. Other genes may be necessary to resolve the taxonomy of
this group. This is why we present this taxon ad interim.
Geopyxis albocinerea Velen. could be an older name for this
species because its lectotype (PRM 150774) was collected under Be-
tula in Czech Republic and synonymized with T. gaillardiana by
SVRČEK (1979). Unfortunately, the Czech author indicated that the
collection was fully immature and “partially destroyed by moulds”,
so no comparison can be done with the studied material.
Excluded or doubtful species
Here is a list of species placed in Tarzetta that we exclude from
this genus. The correct name is in bold.
Tarzetta ammophila (Sacc.) Theodor., Badania Przyrodnicze Po-
morskie, Tow. nauk., Torun: 11 (1936).
Peziza ammophila Sacc.
Tarzetta brasiliensis Rick, Brotéria, sér. bot., 25 (2-3): 80 (1931).
The short diagnosis of RICK (1931) indicates “sporis […] eguttu-
latis. This feature excludes it from the genus Tarzetta.
Tarzetta cinerascens Rehm, Ann. mycol., 2 (4): 352 (1904).
The diagnosis of REHM (1904) indicates “Asciporus J+”. This fea-
ture excludes it from the genus Tarzetta.
Tarzetta rapulum (Bull.) Rehm, Rabenh. Kryptog.-Fl., Pilze – Ascom.,
1 (3): 1021 (1894).
Stromatinia rapulum (Bull.) Boud.
Tarzetta rapuloides Rehm, Hedwigia, 38 (Beibl.): (243) (1899).
The short size of ascospores, 8–10 × 5 µm, given by REHM (1899)
seems doubtful for a Tarzetta and suggests a member of inopercu-
late discomycetes.
Tarzetta rosea (Rea) Dennis, Brit. Ascom.: 30 (1978).
Rhodotarzetta rosea (Rea) Dissing & Sivertsen
Plate 16 Tarzetta betulicola ad int. Coll. MP 2018-133 (in situ).
Photo M. Pélissier.
We thank the following people who helped us in this study with
their collections, data, photographs or literature: Carlo Agnello, Mar-
tin Bemmann, Brigitte Capoen, Jean-Louis Cheype, Jerry Cooper,
Jean-Marie Cugnot, René Dougoud, Guillaume Eyssartier, Francis
Fouchier, Christian Frund, Beñat Jeannerot, Sava Krstic, Jean Mor-
nand, Gilbert Moyne, Jean-Luc Muller, Claude Page, Maurice
Pélissier, Donald Pfister, Vincent Ricard, Nicole Robert, Leandro
Sánchez, Clive Shirley, Jean-Claude Tinlot et Marie-Paule Vigneron.
We also thank Jacques Melot for his nomenclatural advices. The fol-
lowing curator are also acknowledged for the loans of material or
photographs: Guðríður Gyða Eyjólfsdóttir (AMNH, Iceland), Lee
Davies (K, United Kingdom), Perti Salo (H, Finland), Roxali Bijmoer
(L, The Netherlands), Åsa Kruys (UPS, Sweden) and Javier Rejos (AH,
Spain). is thanked for having financially supported a
large part of the sequencing made by the laboratories Alvalab
(Spain) and Mycea (France).
Finally, Rosanne Healy and Matthew E. Smith (University of
Florida, USA) are warmly thanked for the pre-submission review.
2012. — Comparison of ectomycorrhizal communities in natural
and cultivated Tuber melanosporum truffle grounds. FEMS Micro-
biol Ecology, 81 (3): 547–561. doi: 10.1111/j.1574-6941.2012.01379.x
2014. — Assessment of ectomycorrhizal biodiversity in Tuber
macrosporum productive sites. Mycorrhiza, 24: 281–292. doi:
BERTHET P. & RIOUSSET L. 1964. — Un Pustularia nouveau : P. insignis.
Description de l’espèce et remarque sur un caractère cytologique
du genre Pustularia. Bulletin de la Société mycologique de France,
79: 392–398.
BOUDIER E. 1902. — Champignons nouveaux de France. Bulletin de la
Société mycologique de France, 18: 137–146.
BOUDIER E. 1905-1910. Icones mycologicæ ou Iconographie des
champignons de France. Paris, Paul Klincksieck, 4 vol.
BRESADOLA G. 1892. — Fungi tridentini novi, vel nondum delineati, de-
scripti, et iconibus illustrati. Vol. 2. Trentin, J. Zippel. 118 pp.
COOKE M.C. 1879. Mycographia, seu icones fungorum. Vol. I. Dis-
comycetes. London, Williams and Norgates, 267 pp. + 113 pl.
DENNIS R.W.G. 1978. — British Ascomycetes. 2nd ed. Vaduz, J. Cramer,
486 pp., 31 figures, pl. 1–44.
DISSING H., ECKBLAD F.-E. & LANGE M. 2000. — Pezizales Bessey. In:
HANSEN L. & KNUDSEN H. (éd.) Nordic Macromycetes. Vol. 1. As-
comycetes. Copenhagen, Nordsvamp, 309 pp.
DONADINI J.-C. 1985. — Discomycetes - 2 - Cytologie, taxonomie, et
phylogénie (applications à Hypotarzetta gen. nov., H. insignis c.
nov., Humariaceae et Pyronemataceae). Documents mycologiques,
15 (60): 47–57.
ECKBLAD F.-E. 1968. — The genera of the operculate discomycetes. A
re-evaluation of their taxonomy, phylogeny and nomenclature.
Nytt Magasin for Botanikk, 15 (1): 1–191.
EDGAR R.C.2004. — MUSCLE: a multiple sequence alignment method
with reduced time and space complexity. BMC Bioinformatics, 5
(1): 113. doi: 10.1186/1471-2105-5-113
EKANAYAKA A.H., HYDE K.D., JONES E.B.G. & ZHAO Q. 2018. — Taxonomy
and phylogeny of operculate discomycetes: Pezizomycetes. Fun-
gal Diversity, 90: 161–243. doi: 10.1007/s13225-018-0402-z
GATES G. & VAN VOOREN N. 2016. Jafnea pallida (Pezizales), a new
name for a cup fungus rediscovered in Tasmania.,
8 (3): 101–104. doi: 10.25664/art-0178
HÄFFNER J. 1992. — Die Gattung Tarzetta (Cooke) Lambotte und Be-
nachbarte Keingattungen. Rheinland-Pfälzisches Pilzjournal, 2 (1):
logeny of the highly diverse cup-fungus family Pyronemataceae
(Pezizomycetes, Ascomycota) clarifies relationships and evolution
of selected life history traits. Molecular Phylogenetics and Evolution,
67 (2): 311–335. doi: 10.1016/j.ympev.2013.01.014
HARMAJA H. 1974. — Tarzetta pusilla n. sp. and T. spurcata (Pers.) n.
comb. from Finland. Karstenia, 14: 116–120.
HEDWIG J. 1789. Descriptio et adumbratio microscopico-analytica
muscorum frondosorum. Vol. 2. Lipsiae, J.G. Müller.
HOLMSKJOLD T. 1799. — Beata ruris otia fungis danicis. Vol. 2. Copen-
hagen, F. Brummer, 70 pp.
cota. In: FREY W. (ed). Syllabus of plant families — Adolf Engler’s Syl-
labus der Pflanzenfamilien. Stuttgart, Borntraeger, 322 pp.
KANOUSE B.B. 1950. — A study of Peziza bronca Peck. Mycologia, 42
(4): 497–502. doi: 10.1080/00275514.1950.12017854
KORF R.P. 1972. — Synoptic key to the genera of the Pezizales. My-
cologia, 64 (5): 937–994. doi: 10.1080/00275514.1972.12019349
KORF R.P. 1986. — Révision des espèces illustrées par Boudier dans
les tomes II et III des Icones Mycologicæ. In: BRUMMELEN J. (VAN), KORF
R.P., CLÉMENÇON H., JÜLICH W. & DEMOULIN V. (eds.) Icones Mycologicæ
par Émile Boudier. Volume V: liste préliminaire et explication des
planches. Lausanne, Piantanida, 300 pp.
MCLAUGHLIN D.J., TRAPPE J.M. & HEALY R.A. 2017. — A molecular and
morphological re-examination of the generic limits of truffles in
the tarzetta-geopyxis lineage Densocarpa, Hydnocystis, and Pau-
rocotylis. Fungal Biology, 121 (3): 264–284. doi: 10.1016/j.funbio
lecular Evolutionary Genetics Analysis across computing plat-
forms. Molecular Biology and Evolution, 35 (6): 1547–1549. doi:
LAMBOTTE E. 1888. — La flore mycologique de la Belgique. Premier
supplément comprenant les hyménomycètes - Pyrénomycètes -
Discomycètes. Mémoires de la Société royale des sciences de Liège,
2esérie, XIV: 1–350.
LEGAL M. 1955. — Étude critique sur les Discomycètes récoltés en
Suisse au cours de la session de 1953. Bulletin de la Société my-
cologique de France, 70 (3): 185–218.
LINNÉ C. (VON) 1753. — Systema plantarum. Vol. 2. Holmiae, Impensis
Laurentii Salvii. doi: 10.5962/bhl.title.669
MADDISON W.P. & MADDISON D.R. 2015. — Mesquite: a modular system
for evolutionary analysis. Version 3.51. Available from:
MATEO FERNÁNDEZ J.F. 2011. — Hypotarzetta insignis una especie muy
rara recolectada en Cuenca. Boletín Sociedad Micológica de Madrid,
35: 25–29.
MILLER M.A., PFEIFFER W. & SCHWARTZ T. 2010. — Creating the CIPRES
Science Gateway for inference of large phylogenetic trees. In: Pro-
ceedings of the Gateway Computing Environments Workshop
(GCE), 14 Nov. 2010, New Orleans, LA: 1–8. doi: 10.1109/
MORNAND J. & PÉAN R. 1995.Contribution à la connaissance des
champignons de Maine-et-Loire. 4. Discomycètes (= Pezizomyce-
tideae). Première partie : Operculés. Bulletin de la Société my-
cologique de France, 111 (1): 3–32.
NANNFELDT J.A. 1938. — Contributions to the mycoflora of Sweden.
5. On Peziza catinus Holmskj. ex Fr. and P. radiculata Sow. ex Fr.
with a discussion of the genera Pustularia Fuck. emend. Boud. and
Sowerbyella Nannf. n. gen. Svensk Botanisk Tidskrift, 32: 108–120.
NEI M. & KUMAR S. 2000. Molecular Evolution and Phylogenetics.
New York, Oxford University Press, 333 pp.
PANT D.C. & TEWARI V.P. 1970. — Observations on two species of the
genus Pustulina. Mycologia, 62 (6): 1187–1194. doi: 10.1080/
PECK C.H. 1878. — Report of the Botanist (1875). Annual Report on
the New York State Museum of Natural History, 29: 29–82.
QUÉLET L. 1895. — Quelques espèces critiques ou nouvelles de la
flore mycologique de France. Comptes rendus de l’Association
française pour l’avancement des sciences, 24 (2): 616–622.
RAITHELHUBER J. 1983. — Über die Nomenklatur einiger Argentinischer
Blätterpilze. Metrodiana Sonderheft, 2: 1–24.
RAMBAUT A. 2010. — FigTree v1.3.1. Edinburgh, Institute of Evolution-
ary Biology, University of Edinburgh. Available from:
REA C. 1928. — New or rare British Discomycetae. Transactions of the
British mycological Society, 13 (3-4): 253–260. doi: 10.1016/S0007-
REHM H. 1899. — Ascomycetes exs. fasc. 26. Beiblatt zur Hedwigia, 38
(1): (242)–(246).
REHM H. 1904. — Ascomycetes Americae borealis. II. Annales myco-
logici, 2 (4): 351–354.
RICK J. 1931. Monographia Pezizinearum Riograndensium.
Brotéria, série botânica, 25 (2–3): 77–122.
ROGERS J.K., DUMONT K.P. & KORF R.P. 1971. — Nomenclatural notes. VIII.
Stromatinia, an available generic name (not to be replaced by
Tarzetta, the correct name for Pustulina). Mycologia, 63 (5): 1084–
1086. doi: 10.1080/00275514.1971.12019208
SARRAILLON F. & DOLL D. 2013. Première récolte en Alsace : Hy-
potarzetta insignis (Berthet & Riousset) Donadini. Bulletin de la
Fédération mycologique de l’Est, 11: 50–53.
SCHUMACHER T. 1979. — Notes on taxonomy, ecology, and distribution
of operculate discomycetes (Pezizales) from river banks in Norway.
Norwegian Journal of Botany, 26: 53–83.
SEAVER F.J. 1928. — The North American Cup-fungi (Operculates). New
York, Seaver, 265 pp.
SMITH M.E., DOUHAN G.W. & RIZZO D.M. 2007. — Ectomycorrhizal com-
munity structure in a xeric Quercus woodland based on rDNA se-
quence analysis of sporocarps and pooled roots. New Phytologist,
174 (4): 847–863. doi: 10.1111/j.1469-8137.2007.02040.x
& DEVINE W.D. 2009. — Mycorrhizas on nursery and field seedlings
of Quercus garryana. Mycorrhiza, 19 (3): 149–158. doi: 10.1007/s00572-
STAMATAKIS A. 2014. — RAxML version 8: a tool for phylogenetic analy-
sis and post-analysis of large phylogenies. Bioinformatics, 30 (9):
1312–1313. doi: 10.1093/bioinformatics/btu033
SVRČEK M. 1979 [1976]. — A taxonomic revision of Velenovský’s types
of operculate discomycetes (Pezizales) preserved in National Mu-
seum, Prague. Acta Musei Nationalis Pragae, 32B (2-4): 115–194.
TEDERSOO L., HANSEN K., PERRY B. & KJØLLER R. 2006. — Molecular and
morphological diversity of pezizalean ectomycorrhiza. New Phy-
tologist, 170 (3): 581–596. doi: 10.1111/j.1469-8137.2006.01678.x
TEDERSOO L., SUVI T., JAIRUS T., OSTONEN I. & PÕLME S. 2009. — Revisiting
ectomycorrhizal fungi of the genus Alnus: differential host speci-
ficity, diversity and determinants of the fungal community. New
Phytologist, 182 (3): 727–735. doi: 10.1111/j.1469-8137.2009.02792.x
Rich and cold: Diversity, distribution and drivers of fungal com-
munities in patterned-ground ecosystems of the North American
Arctic. Molecular Ecology, 23 (13): 3258–3272. doi: 10.1111/mec.12743
ternational Code of Nomenclature for algae, fungi, and plants (Shen-
zhen Code) adopted by the Nineteenth International Botanical
Congress Shenzhen, China, July 2017. Regnum Vegetabile 159.
Glashütten, Koeltz Scientific Books, 254 pp.
VAILLANT S. 1727. Botanicon parisiense ou dénombrement par ordre
alphabétique des plantes qui se trouvent aux environs de Paris. Leide
et Amsterdam, J. & H. Verbeek et B. Lakeman.
VAN VOOREN N. 2010. — Tarzetta versus Tazzetta. Des difficultés d’in-
terpréter l’article 60.1 du Code de nomenclature.,
2 (3): 15–18. doi: 10.25664/art-0027
VAN VOOREN N. 2014. — Contribution à la connaissance des Pézizales
de Rhône-Alpes. 2nd part. Cahiers de la FMBDS, 4: 1–172.
VAN VOOREN N. 2017. — Contribution à la connaissance des Pézizales
en Auvergne-Rhône-Alpes. 3rd part. Cahiers de la FMBDS, 5: 1–126.
& KUŠAN I. 2015. — Lachnea poiraultii (Pezizales), rediscovered after
more than one hundred years., 7 (3): 105–116. doi:
VAN VOOREN N., LINDEMANN U. & HEALY R. 2017. — Emendation of the
genus Tricharina (Pezizales) based on phylogenetic, morphologi-
cal and ecological data., 9 (4): 101–123. doi:
WANG X.H., HUHTINEN S. & HANSEN K. 2016. — Multilocus phylogenetic
and coalescent-based methods reveal dilemma in generic limits,
cryptic species, and a prevalent intercontinental disjunct distri-
bution in Geopyxis (Pyronemataceae s. l., Pezizomycetes). Mycolo-
gia, 108 (6): 1189–1215. doi: 10.3852/16-100
al. 2017. — Notes for genera: Ascomycota. Fungal Diversity, 86: 1–
594. doi: 10.1007/s13225-017-0386-0
YAO Y.J. & SPOONER B.M. 2002. — Notes on British species of Tazzetta
(Pezizales). Mycological Research, 106 (10): 1243–1246. doi:
ZHUANG W.Y. & KORF R.P. 1987. — A new species of Tarzetta (Pezizales)
from New Zealand. Mycotaxon, 28 (2): 365–368.
2: M. Carbone – Via D on Luigi Sturzo 173, 16148 Genova, Italy –
1: N. Van Vooren – 36 rue de la Garde, 69005 Lyon, France –
3: C. Sammut – 216/1 St. Joseph Flts., GZR1367 Gzira, Malta –
4: A.C. Grupe – Department of Plant Pathology, University of Florida, Gainesville, Florida, U.S.A. –
... Other differences include shape of the cheilocystidia but unfortunately none were observed on the partially dissolved collections. Spore size (7.5-9.9 x 4.7-6.6µm) is within the range published for this species (Uljé, 2005). ...
... also Wächter & Melzer, 2020 (Uljé, 2005). The applied name to the taxon was C. mitraespora and the synonymy with C. spelaiophila has been established in Nagy et al. (2013). ...
... near tree stumps whereas C. subdisseminatus is found in small scattered numbers on dead grass residues. Microscopically they are easily distinguished by the larger pileocystidia and shorter spores in C. disseminatus (Uljé, 2005). The collected specimen was probably old and overripe whilst another specimen close by was stunted in its growth probably due to dry and warm conditions. ...
Fourty new records of macro fungi are reported for the first time from Malta. Some members have been confirmed molecularly using ITS sequence data: Cortinarius castaneus, Crepidotus neotrichocystis, Entoloma graphitipes, Entoloma rusticoides, Lepiota lilacea, Leucoagaricus barssii, Lyophyllum littorale, Mycena olivaceomarginata, Pseudosperma squamatum, Volvariella taylorii and Trichoglossum variabile. A checklist of fungi reported to date from the Maltese Islands is provided.
... Although radical views to bypass some of these difficulties have occasionally been put forward, such as the controversial PhyloCode advocating for a system of rankless phylotaxonomy (de Queiroz and Gauthier 1990, 1992, 1994, or calls to abandon binomial nomenclature altogether (Money 2013), these have been widely rejected by the academic community and Linnaean binomial nomenclature continues to form the basis of scientific communication (e.g., Nixon and Carpenter 2000;Carpenter 2003;Wheeler 2004;Will et al. 2005;Korf 2005;Schoch et al. 2014;Minnis 2015;Dayarathne et al. 2016;Zamora et al. 2018). Therefore, genetic characterization of early-described taxa through sequencing of original material and/or designation of sequenced epitypes remains the most cautious, widely accepted, and least disruptive method of solving complex taxonomic problems and stabilizing taxonomy and nomenclature within critical genera (Hyde and Zhang 2008;Ariyawansa et al. 2014;Liimatainen et al. 2014b;Vesterholt et al. 2014;Borovička et al. 2015;Olariaga et al. 2015;Vizzini et al. 2016Vizzini et al. , 2020Richard et al. 2015;Dima et al. 2016;Skrede et al. 2017;Moreau et al. 2018;Lombard et al. 2018;Turland et al.2018;Van Vooren et al. 2019;Loizides et al. 2020;Van Vooren 2020). Powerful new technologies such as next-generation sequencing, able to produce useful DNA sequences from old and contaminated material, are expected to be decisive in decrypting the genetic identity of early-described taxa in the years to come Bellanger et al. 2021;Bidaud et al. 2021). ...
The genus Morchella has gone through turbulent taxonomic treatments. Although significant progress in Morchella systematics has been achieved in the past decade, several problems remain unresolved and taxonomy in the genus is still in flux. In late 2019, a paper published in the open-access journal Scientific Reports raised serious concerns about the taxonomic stability of the genus, but also about the future of academic publishing. The paper, entitled “High diversity of Morchella and a novel lineage of the esculenta clade from the north Qinling Mountains revealed by GCPSR-based study” by Phanpadith and colleagues, suffered from gross methodological errors, included false results and artifactual phylogenies, had misapplied citations throughout, and proposed a new species name invalidly. Although the paper was eventually retracted by Scientific Reports in 2021, the fact that such an overtly flawed and scientifically unsound paper was published in a high-ranked Q1 journal raises alarming questions about quality controls and safekeeping procedures in scholarly publishing. Using this paper as a case study, we provide a critical review on the pitfalls of Morchella systematics followed by a series of recommendations for the delimitation of species, description of taxa, and ultimately for a sustainable taxonomy in Morchella. Problems and loopholes in the academic publishing system are also identified and discussed, and additional quality controls in the pre- and post-publication stages are proposed.
Based on recent taxonomic publications on Pezizales, an argued update of the names of species presented in the three volumes of the series "Contribution à la connaissance des Pézizales (Ascomycota) en Auvergne-Rhône-Alpes", published between 2014 and 2017, is proposed.
Full-text available
Knjiga „Diverzitet gljiva Zaštićenog pejzaža "Konjuh" - Inventarizacija, zaštita i promocija” bavi se prezentacijom preliminarnih rezultata mikoloških istraživanja realizovanih tokom 2019. i 2020. godine na teritoriji Zaštićenog pejzaža "Konjuh", trenutno jedinog protektorata na području Tuzlanskog kantona. U okviru knjige izdvojeno je i predstavljeno 114 vrsta gljiva iz odjeljaka Ascomycota i Basidiomycota te predstavljen popis gljiva ZP "Konjuh" sa ukupno 399 različitih vrsta. Za manji broj registrovanih vrsta predloženi su odgovarajući IUCN statusi i kategorije, a u skladu sa provedenom evaluacijom i relevantnim konzervacijskim kriterijima. Na osnovu predloženih kategorija i procjene ugroženosti pojedinih staništa, upravi zaštićenog područja su predložene i odgovarajuće mjere i aktivnosti s ciljem održavanja, ali i unapređenja postojećeg stanja i diverziteta gljiva. Autori unutar knjige nastavljaju i s prijedlogom implementacije koncepta IFA područja, u koja se, uzimajući u obzir realizovana istraživanja, ubraja i ZP "Konjuh".
Full-text available
The molecular evolutionary genetics analysis (Mega) software implements many analytical methods and tools for phylogenomics and phylomedicine. Here, we report a transformation of Mega to enable cross-platform use on Microsoft Windows and Linux operating systems. Mega X does not require virtualization or emulation software and provides a uniform user experience across platforms. Mega X has additionally been upgraded to use multiple computing cores for many molecular evolutionary analyses. Mega X is available in two interfaces (graphical and command line) and can be downloaded from free of charge.
Full-text available
Knowledge of the relationships and thus the classification of fungi, has developed rapidly with increasingly widespread use of molecular techniques, over the past 10–15 years, and continues to accelerate. Several genera have been found to be polyphyletic, and their generic concepts have subsequently been emended. New names have thus been introduced for species which are phylogenetically distinct from the type species of particular genera. The ending of the separate naming of morphs of the same species in 2011, has also caused changes in fungal generic names. In order to facilitate access to all important changes, it was desirable to compile these in a single document. The present article provides a list of generic names of Ascomycota (approximately 6500 accepted names published to the end of 2016), including those which are lichen-forming. Notes and summaries of the changes since the last edition of ‘Ainsworth & Bisby’s Dictionary of the Fungi’ in 2008 are provided. The notes include the number of accepted species, classification, type species (with location of the type material), culture availability, life-styles, distribution, and selected publications that have appeared since 2008. This work is intended to provide the foundation for updating the ascomycete component of the “Without prejudice list of generic names of Fungi” published in 2013, which will be developed into a list of protected generic names. This will be subjected to the XIXth International Botanical Congress in Shenzhen in July 2017 agreeing to a modification in the rules relating to protected lists, and scrutiny by procedures determined by the Nomenclature Committee for Fungi (NCF). The previously invalidly published generic names Barriopsis, Collophora (as Collophorina), Cryomyces, Dematiopleospora, Heterospora (as Heterosporicola), Lithophila, Palmomyces (as Palmaria) and Saxomyces are validated, as are two previously invalid family names, Bartaliniaceae and Wiesneriomycetaceae. Four species of Lalaria, which were invalidly published are transferred to Taphrina and validated as new combinations. Catenomycopsis Tibell & Constant. is reduced under Chaenothecopsis Vain., while Dichomera Cooke is reduced under Botryosphaeria Ces. & De Not. (Art. 59).
Full-text available
Tricharina is one of the most difficult genera of Pezizales because it is hard to distinguish morphologically among species. To provide a more robust taxonomy, new investigations on the genus were conducted, both morphologically and phylogenetically. This study focuses on the four key species of the genus: T. ascophanoides, T. gilva, T. ochroleuca and T. praecox. The type material of these species was reviewed. In the literature, T. gilva is considered to be close or even identical to T. ochroleuca. The phylogenetic analysis shows that all recent collections identified as T. ochroleuca appear to be T. gilva. Furthermore, the morphological study of the type material of T. ochroleuca leads to the conclusion that the name is a nomen dubium. The presence of several endophytes sequences in the Tricharina-core clade suggests that this genus has an endophytic lifecycle. The confusion between T. gilva and T. praecox, both considered as pyrophilous species, is now resolved thanks to phylogenetic analyses and new data based on vital taxonomy. In contrast to T. gilva, which can occasionally grow on burnt places, T. praecox is a strictly pyrophilous taxon and belongs genetically to a different clade than T. gilva. In agreement with art. 59 of ICN, the anamorphic genus Ascorhizoctonia is used to accommodate the species belonging to the “T. praecox clade”. T. ascophanoides, an extra-limital species, is excluded from the genus Tricharina and combined in the genus Cupulina based on the phylogenetic results. The genetic exploration of some other tricharinoid species, i.e. Leucoscypha semi-immersa and L. patavina, confirms they do not belong to the genus Leucoscypha in its original sense, and deserve their own genus. The name Sepultariella is validated to accommodate the latter
Two species of the genus Pustulina, P. cupularis and P. gaillardiana comb, nov., have been described and illustrated. The description of P. cupularis is based on the study of Fung. rhen. No. 1878, Peziza cupularis L., in FH which has been designated the neotype specimen of this species. Three Indian collections of P. cupularis, made by the authors, have also been compared with the type. Pustularia patavina, on the basis of the study of type specimen No. 1777 Peziza patavina from PAD, has been shown to belong to the genus Leucoscypha.
Our catalogue of Pezizales species, registered in the Auvergne-Rhône-Alpes region (France), continues through the detailed presentation and illustration of new species. This third part includes 78 species and varieties