Preliminary notes on the genus Tarzetta (Pezizales) with typiﬁ-
cations of some species and description of six new species
Nicolas VAN VOOREN
Arthur C. GRUPE
Ascomycete.org, 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
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 (http://www.my-
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-
Fig. 2 – Molecular phylogenetic analysis by Maximum Likelihood method of ITS alignment of Tarzetta, rooted by other species of the /tar-
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.
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
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
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
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
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
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 6 – Tarzetta 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, ⅓unc. 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
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
alba… grisea” 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,
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-
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:
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
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
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
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
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
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
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
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 14.717° W, ~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 “Asci… porus 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-
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,
Ascomycete.org 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.
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2: M. Carbone – Via D on Luigi Sturzo 173, 16148 Genova, Italy – firstname.lastname@example.org
1: N. Van Vooren – 36 rue de la Garde, 69005 Lyon, France – email@example.com
3: C. Sammut – 216/1 St. Joseph Flts., GZR1367 Gzira, Malta – firstname.lastname@example.org
4: A.C. Grupe – Department of Plant Pathology, University of Florida, Gainesville, Florida, U.S.A. – agrupe@uﬂ.edu