Persoonia 35, 2015: 166 – 229
www.ingentaconnect.com/content/nhn/pimj
RESEARCH ARTICLE
http://dx.doi.org/10.3767/003158515X688000
A monograph of Otidea (Pyronemataceae, Pezizomycetes)
I. Olariaga1, N. Van Vooren 2, ε. Carbone 3, K. Hansen1
Key words
Flavoscypha
ITS
ITS1 minisatellites
LSU
Otideopsis
resinous exudates
Abstract The easily recognised genus Otidea is subjected to numerous problems in species identification. A number
of old names have undergone various interpretations, materials from different continents have not been compared and
misidentifications occur commonly. In this context, Otidea is monographed, based on our multiple gene phylogenies
assessing species boundaries and comparative morphological characters (see Hansen & Olariaga 2015). All names
combined in or synonymised with Otidea are dealt with. Thirty-three species are treated, with full descriptions and
colour illustrations provided for 25 of these. Five new species are described, viz. O. borealis, O. brunneoparva, O. oregonensis, O. pseudoleporina and O. subformicarum. Otidea cantharella var. minor and O. onotica var. brevispora
are elevated to species rank. Otideopsis kaushalii is combined in the genus Otidea. A key to the species of Otidea
is given. An δSU dataset containing 167 sequences (with 44 newly generated in this study) is analysed to place
collections and determine whether the named Otidea sequences in GenBank were identified correctly. Fourty-nine
new ITS sequences were generated in this study. The ITS region is too variable to align across Otidea, but had low
intraspecific variation and it aided in species identifications. Thirty type collections were studied, and ITS and δSU
sequences are provided for 12 of these. A neotype is designated for O. cantharella and epitypes for O. concinna,
O. leporina and O. onotica, along with several lectotypifications. The apothecial colour and shape, and spore characters are important for species identification. We conclude that to distinguish closely related or morphologically
similar species, a combination of additional features are needed, i.e. the shape of the paraphyses, ectal excipulum
structure, types of ectal excipulum resinous exudates and their reactions in εelzer’s reagent and KOH, tomentum
and basal mycelium colours and exudates. The KOH reaction of excipular resinous exudates and basal mycelium
are introduced as novel taxonomic characters.
Article info Received: 26 November 2013; Accepted: 1 February 2015; Published: 10 April 2015.
INTRODUCTION
Species of Otidea produce typically ear-shaped apothecia that
are unique within Pyronemataceae (Pezizomycetes). The genus
is monophyletic based on multilocus phylogenetic analyses
from a few, but broadly sampled, Otidea species (Hansen
et al. 2013). Despite being distinct at the generic level, the
species identification and nomenclature of Otidea are highly
controversial. A few recent typifications have been proposed
(Carbone 2009, 2010a), but many names are still subjected to
different interpretations. Several new species were described
in the last decades from Europe (Harmaja 1976, 2009a) and
Asia (Cao et al. 1990, Zhuang & Yang 2008), with detailed
descriptions and updated identification keys. However, often
no illustrations were presented and colour photographs have
rarely been published when describing new species. εany
names of European species currently used in North America
and Asia are misapplied. εultilocus phylogenetic analyses have
not been previously implemented to critically address species
delimitation issues and material from different continents has
not been compared. A worldwide critical revision of Otidea to
clarify species limits is highly needed. The aims of this study
were: i) to undertake a nomenclatural and taxonomic revision
of Otidea, to clarify misinterpretations and to propose pertinent
typifications to stabilise the use of names; and ii) to provide
detailed species descriptions and colour photographs of both
macro- and microscopic structures, and a key for identification.
1
2
3
Swedish εuseum of Natural History, Department of Botany, P.O. Box 50007,
SE-104 05 Stockholm, Sweden;
corresponding author e-mail: karen.hansen@nrm.se.
36 rue de la Garde, F-69005 δyon, France.
Via Don δuigi Sturzo 173 I-16148 Genova, Italy.
Our multilocus phylogenies and robust hypotheses of species
limits, employing genealogical concordance phylogenetic species recognition (GCPSR; Taylor et al. 2000), which is the basis
for the present work, are given in Hansen & Olariaga (2015).
In the present study we present an δSU rDNA phylogeny to
place a larger number of collections for which we have been
unable to obtain multiple genes, including several sequences
from GenBank, many of which we here re-identify.
Taxonomic history
The first valid publication of Otidea is by Bonorden (1851),
based on Peziza (unranked) Otidea Pers., although it has sometimes been attributed to Fuckel (Kanouse 1949, Nannfeldt
1966, δiu & Zhuang 2006, Smith & Healy 2009). Otidea species
were treated in a broad heterogeneous genus Peziza by early
authors. Persoon (1822) defined Peziza (unranked) Otidea
as producing auriculate apothecia with a split, sometimes
elongated on one side, and included 10 species. Fries (1822)
referred to this group as Peziza (unranked) Cochleatae, but he
included also taxa with non-split apothecia. Bonorden (1851)
elevated Otidea to generic rank with split apothecia as the key
feature, but also referred to Fries’ (1822) Cochleatae. He did
not make any combinations or list any species. Fuckel (1870)
refined the genus using microscopic details, namely uni- or
biguttulate spores and filiform to subclaviform paraphyses, and
included four species: O. abietina, O. cochleata O. leporina
and O. onotica. Boudier (1885) notably contributed to disassemble the large genus Peziza into smaller and more natural
genera. He placed in the genus Otidea species with entire or
split apothecia, biguttulate spores and, importantly non-amyloid
asci and curved paraphyses, which he was the first to introduce. He divided Otidea into two subgenera: Otidea with split
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167
I. Olariaga et al.: A monograph of Otidea
apothecia and Pseudotis with entire apothecia. Interestingly,
Boudier erected the genus Wynnella (Helvellaceae) to accommodate W. silvicola (as P. leporina / P. auricula), a species
with distinctly ear-shaped apothecia, but differing from Otidea
in the uniguttulate spores and tough consistency. In Boudier’s
(1907) subsequent treatment, he elevated Pseudotis to genus
rank and placed here species with entire apothecia, including
O. daliensis (as P. apophysata) and O. propinquata (as P. abietina), both with biguttulate spores and hooked paraphyses.
Saccardo listed O. onotica as an ‘exemplar’ species of Peziza
subg. Otidea in his synopsis of the discomycete genera (1884;
he listed in general only one species per genus / subgenus that
appear to have been selected as typical for the genera) and
it has since been accepted as the type species by most (Rifai
1968, Eckblad 1968, Korf 1972, δiu & Zhuang 2006, Parslow &
Spooner 2013), Index Nominum Genericorum (Eckblad in Farr
et al. 1979), NCU-3 (Greuter et al. 1993) and is prepared to be
adopted on the δist of Protected generic Names for fungi (Kirk
et al. 2013). Clements & Shear (1931) listed O. cochleata as
the type, but this name was not among the original species accepted by Persoon (1822) and has furthermore been considered
an ambiguous name. Kanouse (1949) proposed O. leporina as
the type species, but because of the confusions surrounding
the identity of this species until now, it has been considered an
inappropriate choice.
The genus Scodellina was described by Gray (1821) and involved species attributed today to Otidea, and also Aleuria,
Peziza and Tarzetta. Seaver (1928) refined Scodellina to
species with split to ear-shaped apothecia only and typified it
with Peziza leporina (Seaver 1927), considering Otidea a later
synonym. This was for a period followed by several American
authors (e.g. Korf 1963, Kimbrough 1966). The typification by
Seaver can however, be considered largely mechanical, taken
as the first species listed by Gray, and be superseded under
the ICN (Art. 10.5; εcNeill et al. 2012). Also, even though
Gray (1821) included several species with split apothecia, he
did not mention this feature in the diagnosis of Scodellina, but
emphasised “thallus … hemispherical, spreading” and coined
the vernacular name “spread cup”. Therefore Rifai (1968) designated P. vesiculosa Bull.: Fr. as the type species of Scodellina,
consequently making it a later synonym of the genus Peziza.
Eckblad (1968) came to the same conclusion.
Kanouse (1949) broadened the concept of Otidea and described in detail a number of North American species. She
included species with split apothecia and straight paraphyses
with swollen apices. She also included Wynnella silvicola (as
Otidea auricula), with straight paraphyses and uniguttulate
spores. Nannfeldt (1966) delimited Otidea to species with
non-amyloid asci; smooth, uninucleate, biguttulate spores; a
medullary excipulum of textura intricata; and an ectal excipulum with isodiametric cells, covered by short chains of barrelshaped cells. Nannfeldt’s concept more or less conforms to the
genus Otidea as we recognise it today. He considered Wynnella so distant that it should be treated in a separate tribe of
Pezizaceae. Korf (1963) reviewed the monotypic, sparassoid
genus Ascosparassis, and subsequently (Korf 1973a) assigned
A. shimizuensis to Otidea, based on the hooked paraphyses,
small biguttulate spores and excipulum structure. Pfister (1979)
however, considered Ascosparassis a distinct monotypic genus,
based on “small asci and spores and peculiar growth habit”, and
combined the older name Midotis heinricheri in Ascosparassis. δater Pfister collected A. heinricheri in South America, in
north coastal mountains of Venezuela (Pfister & Halling 1989),
extending its Asian distribution (China, Indonesia and Japan),
still considering the species separate from Otidea.
A new genus Flavoscypha was erected for two species of
Otidea, O. concinna (as Flavoscypha cantharella) and O. phlebophora, with strong emphasis on the ectal excipulum of textura
prismatica (vs textura angularis in Otidea) (Harmaja 1974).
Otidea was further emended to include a species with ornamented spores, O. unicisa, otherwise ‘fitting perfectly’ Otidea
(Harmaja 1986). Otideopsis was published with Otideopsis
yunnanensis as the type species, distinguished from Otidea
by having ornamented spores and paraphyses fused at the
apices (δiu & Cao 1987). Flavoscypha and Otideopsis are now
considered synonyms of Otidea based on molecular phylogenetic analyses (δiu & Zhuang 2006, Hansen & Olariaga 2015).
Recently the circumscription of Otidea was further broadened
when the first hypogeous species, O. subterranea, was discovered using ITS and δSU sequences (Smith & Healy 2009). All
the characters proposed so far as diagnostic for Otidea have exceptions across the genus. Nevertheless, Otidea can be recognised by the non-amyloid asci, in combination with at least
two of these characters (except O. subterranea): a) biguttulate
spores; b) hooked or bent paraphyses; c) medium-large, split
apothecia; and d) a medullary excipulum of textura intricata, and
an ectal excipulum of textura angularis or textura prismatica.
Systematic position and relationships
Nannfeldt (1937, 1938, 1966) suggested a close relationship
between Otidea (incl. Pseudotis), Tarzetta (as Pustularia) and
Helvella (Pezizaceae, tribe Acetabuleae sensu Nannfeldt),
based on similarities in asci, paraphyses and anatomical structures of the apothecia, i.e. a medullary excipulum of dense
textura intricata, ectal excipulum of almost isodiametric large
cells, and an outermost layer of shorter or longer chains of
cells, possible forming distinct clusters or warts (and in Tarzetta
prolonged to cylindrical, hyaline, wavy hairs). δe Gal (1947)
similarly placed Otidea in the tribe Otideeae in her Aleuriaceae
(i.e. a family including taxa with both amyloid and non-amyloid
asci), together with Pseudotis and Tarzetta (as Pustularia), but
placed Helvella in Helvellaceae. Following the ideas of Nannfeldt (1966), Eckblad (1968) erected the family Otideaceae
as a small taxon of closely related genera that produce larger
apothecia, most of which typically lack bright orange to red colours, including in it Tarzetta (as Pustulina) and Otidea, but also
Ascosparassis, Geopyxis and Sowerbyella. Eckblad considered
Helvella (Helvellaceae) to be distant, but having a possible
shared origin with members of Otideaceae (and Morchellaceae
and Rhizinaceae), due to the structure of the excipulum, spores
and asci, especially of Tarzetta. At the same time, he expanded
the concept of Pyronemataceae (to 21 genera) to taxa mostly
characterised by the presence of carotenoid pigments, stating
the inability to satisfactorily subdivide the family on the basis
of common characters. Korf (1972, 1973b) placed Otidea in
the tribe Otideeae (in the subfamily Otideoideae) in an even
more encompassing Pyronemataceae (49 genera), together
with Ascosparassis and Psilopezia. He followed the ideas
of Arpin (1969), in excluding taxa with carotenoids from the
Otideoideae, instead including taxa with prominent hairs such
as Geopora, Humaria and Trichophaea. Otidea has generally
been included in a broadly circumscribed Pyronemataceae
in recent treatments (Dissing 2000, Hansen & Pfister 2006,
Perry et al. 2007, Hansen et al. 2013). εultigene phylogenetic
analyses of Pyronemataceae do not support a close relationship between Otidea and Geopyxis, Psilopezia, Sowerbyella
or Tarzetta (Hansen et al. 2013). Surprisingly, the cleistothecial
Warcupia and the highly reduced (gymnohymenial) Monascella are suggested as the closest relatives of Otidea. Otidea,
Monascella and Warcupia are strongly supported as a distinct
sister group to the rest of the Pyronemataceae in a strict sense
(Hansen et al. 2013).
168
MATERIALS AND METHODS
Material and morphological methods
This study is based on a total of 450 specimens. One hundred
and forty two of these were collected and studied fresh during this project and are deposited in S and a few in TUR-A.
Specimens were studied from the following herbaria: AH, ARAN,
BIO, C, FH, H, HKAS, K, εCVE, εICH, εIN, OSC, PC, PRε,
S, TUR and UPS (Thiers 2014), GεFN (gruppo AεB di Fara
Novarese, Italy), SEST (Sociedad de Ciencias Naturales de
Sestao, Spain), and from the private herbaria of G. Corriol, GC;
C. δavorato, Cδ; N. Van Vooren, NV; and ε. Tabarés, εT. Thirty
type collections were examined, along with other original material. Colour codes are based on Kornerup & Wanscher (1961).
For O. apophysata, O. borealis, O. daliensis, O. oregonensis,
O. phlebophora, O. pseudoleporina, O. smithii and O. unicisa
colour codes for fresh material were taken from photographs.
Apothecial sections, i.e. the thickness of the apothecia including
the hymenium, was measured midway between the apothecial
margin and base. Smell and taste are listed when recorded.
Only discharged, mature spores were measured from living
material. To obtain and ensure mature spores from dried material, a square of c. 3 mm 2 of an apothecium was revived in
a drop of water on a slide, with the hymenium surface facing down and then removed after 1 hour. In this way mature
spores deposited on top of the hymenium were recovered and
measured. Spore measures exclude ornamentation. Spore
statistics are based on measurements of 20 spores from each
collection: Lm = mean length, Wm = mean width and Qm = Lm/
Wm. The number of populations that the statistics are based on
is indicated by ‘n’. Extreme values are given in parentheses.
Hymenial elements were observed by teasing apart a small
piece of hymenium with a needle. To observe the excipulum
structure, sections of apothecia were made by hand prior to
soaking the material in water. Basal mycelium was examined
by mounting clumps of hyphae from the apothecial base (the
tomentum) and from among the substrate particles. Only asci
with mature spores were measured. All measurements were
made in water: in living state whenever possible; only when
not possible, measurements were made on rehydrated (over
2 hours in water) material. εelzer’s reagent (εδZ) and 10 %
KOH were used to observe the reaction of resinous exudates
and other pigmentation. Cotton Blue in lactic acid was used to
observe spore ornamentation. εicroanatomical terminology
follows Korf (1973b). The notation ‘!’ indicates that type or other
original material was examined by us.
If not otherwise indicated in the legends, the photographs presented in this paper were taken by K. Hansen and I. Olariaga.
DNA extraction, PCR amplification, sequencing and
alignment
DNA was extracted from dried material, or from fresh material stored in 1 % SDS DNA extraction buffer. The extraction
method follows Hansen et al. (1999), except dried material was
ground in a εini-Beadbeater Tε (Biospec Products, Bartlesville,
OK, USA) and fresh material using a plastic pestle, in eppendorf tubes. The primer combination ITS5-ITS4, and in a few instances ITS1–ITS4, ITS5–5.8S and ITS3–ITS4, were used to
PCR amplify the ITS region, and δR0R –δR5 the δSU region.
For DNA extracted from fresh material (stored in SDS extraction buffer), the ITS and δSU regions were amplified in a
single piece using the primers ITS1– δR5. The ITS2 region
for O. integra (possible original material from 1892) was
successfully amplified in three pieces, in combination with
newly designed primers for the O. concinna clade: ITS3 –
ConcITS2midR (5´-GCCTGTAAATTTTAAAGACGAA-3´);
ConcITS2midF (5´-CCAGGGTTGCTTTGGTA-3´) – ConcITS4
Persoonia – Volume 35, 2015
intR (5´-CACTGGGTAATTGGAGGTTT-3´); ConcITS2midF
(5´-CCAGGGTTGCTTTGGTA-3´) – ITS4. PCR products were
cleaned using ExoSAP-IT® (USB, Cleveland, OH, USA). The
ITS was sequenced in both directions, using the primers ITS1
and ITS4 (and/or in a few instances ITS5, 5.8S and ITS3) and
the δSU using δR0R and δR5. For O. integra, the same primers as used for PCR, were also used for sequencing. PCR
and sequencing conditions follows Hansen & Olariaga (2015).
Sequences were edited and assembled using Sequencher v. 4.10
(Gene Codes Corporation Ann Arbor, εichigan, USA) and have
been deposited in GenBank (Table 1). The sequences were
aligned manually in Se-Al v. 2.0a11 Carbon (Rambaut 2002).
An all taxa δSU dataset was prepared. Monascella botryosa
and Warcupia terrestris were used as outgroup, based on a
higher level phylogenetic study of Pyronemataceae (Hansen
et al. 2013), which supports these as the closest sister group.
To explore inter- and intraspecific variation of the new species
O. borealis, O. subformicarum and their closest relatives, two
smaller datasets were prepared using ITS and δSU rDNA sequences, based on a more species-inclusive, multi-gene phylogeny of Otidea (Hansen & Olariaga 2015). All three alignments
are available from TreeBASE as accession no. S15887. The
first dataset (the O. borealis dataset) contained O. borealis and
related species in the O. concinna clade. The second dataset
(the O. formicarum dataset) contained specimens of the O. formicarum clade. Otidea caeruleopruinosa and O. nannfeldtii
were used as outgroup for each of these datasets. Nucleotide
diversity (e.g. Nei 1987, equation 10.6), as the average number
of nucleotide differences per sites between two ITS sequences,
was calculated within O. brunneoparva and O. subformicarum.
One insertion or deletion, despite the length, was calculated as
only one basepair difference.
Phylogenetic analyses
εaximum δikelihood (εδ) analyses of the all taxa δSU dataset were performed using the ‘RAxεδ HPC2 on XSEDE’ tool
(Stamatakis 2006) via CIPRES Science Gateway (εiller et al.
2010), employing mixed models of evolution and starting from a
random tree. For the two smaller O. borealis and O. formicarum
datasets, εδ analyses were conducted using RAxεδ v. 7.3.1
(Stamatakis 2006) on the Bioportal, University of Oslo (Kumar
et al. 2009). A GTR-GAεεA model with four rate categories
was assigned and all free model parameters were estimated
by the program. For the εδ bootstrap analyses (εδ-BP) 1 000
rapid bootstrapping replicates from random starting trees were
performed, followed by a subsequent εδ search similarly using
1 000 replicates. As no strongly supported conflict was detected
(εδ-BP ≥ 75 %, PP ≥ 95 %), the ITS and δSU region were concatenated for the O. borealis and O. formicarum datasets. Each
combined dataset was analysed using εδ analyses under the
same settings as specified above. Relationships were likewise
constructed using εetropolis-coupled εarkov Chain εonte
Carlo (εCεCεC) and ‘model-jumping’ as implemented in
εrBayes v. 3.2.1 (Ronquist et al. 2012). The substitution model
was sampled across the GTR space by the εCεC analysis
(Huelsenbeck et al. 2004). Four parallel searches, each with
four chains, were run for ten and three million generations,
respectively, for the all-inclusive LSU dataset and the O. borealis and O. formicarum datasets, initiated with random starting
trees. The chains were sampled every 100 generations from
the posterior distribution. The first 25 % of the trees sampled
was discarded as the ‘burn-in’, and the remaining trees were
used to calculate the posterior probabilities (PP) of the clades.
For the combined εδ and Bayesian analyses the ITS and
δSU regions were specified as distinct partitions. εδ bootstrap
values ≥ 70 % and PP ≥ 95 % were considered to be significant.
169
I. Olariaga et al.: A monograph of Otidea
Table 1 Collections used in the molecular phylogenetic analyses, with voucher information and GenBank accession numbers for ITS and δSU regions. Some
GenBank sequences are re-identified by us and the names originally used in GenBank are listed after the taxon names (‘as’). For type specimens (in bold) the
original names are kept regardless of synonymy. Numbers in parentheses following the species names indicate multiple collections of a species. The GenBank
accessions of sequences generated in this study are in bold.
Taxon
Voucher
δocality / year/ collector
ITS
δSU
Monascella botryosa
Otidea alutacea (1)
O. alutacea (2)
O. alutacea (3)
O. alutacea (4)
O. alutacea (5)
O. alutacea (6)
O. alutacea (7)
O. alutacea (8)
O. alutacea (9)
O. alutacea (10)
O. alutacea (11)
O. alutacea (12)
O. alutacea (13)
O. alutacea (14)
O. alutacea (15)
O. alutacea (16)
O. alutacea (17)
O. alutacea (18)
O. alutacea (19)
O. alutacea (20) as O. umbrina
O. alutacea (21) as O. umbrina
O. alutacea (22)
O. alutacea (23)
O. alutacea (24)
O. alutacea (25)
O. alutacea (26)
O. alutacea (27)
O. alutacea (28)
O. alutacea (29)
O. alutacea (30)
O. angusta
O. apophysata
CBS 233.85
KH.09.170 (S)
KH.10.193 (S)
KH.07.46 (S)
JS.08.81 (S)
OSC 56747
OSC 56770
OSC 56798
OSC 56777
JS.08.43 (S)
KH.09.135 (S)
KH.10.198 (S)
KH.09.178 (S)
KS-94-192 (C)
C-F-48045
HεAS52742
HεAS57844
S-F257085
εoorefun19 (OSC)
OSC 56758
OSC 56813
OSC 56782
KH.09.133 (S)
ARAN A3023204
GC 98092002
HεAS83560
HεAS83563
KS-94-111 (C)
HεAS83559
S-F257084
KH.13.50 (S)
H6010804
S-F257062, dupl. private herb.
Kasparek s.n.
S-F242694
HKAS 43003
Spain, 1985, J. Guarro
Sweden, 2009, K. Hansen & I. Olariaga
Sweden, 2010, K. Hansen, K. Gillen & I. Olariaga
Denmark, 2007, H. Knudsen
Sweden, 2008, J. Santos
USA, 1996, E.T. Peterson
USA, 1997, E.T. Peterson
USA, 1996, E.T. Peterson
USA, 1997, E.T. Peterson
Sweden, 2008, J. Santos
Norway, 2009, V. Kučera & I. Kautmanova
Sweden, 2010, K. Hansen, K. Gillen & I. Olariaga
Sweden, 2009, K. Hansen & I. Olariaga
Denmark, 1994, K. Hansen & S.K. Sandal
Sweden, 1974, D. Paulsen & N. Tams
China
China
Italy, 2010, ε. Carbone
USA, 2010, J. εoore
USA, 1996, E.T. Peterson
USA, 1997, E.T. Peterson
USA, 2010, E.T. Peterson
Norway, 2009, K. Hansen & I. Olariaga
Spain, 2009, J.I. δópez Amiano
France, 1998, G. Corriol
China, 2003, W.Y. Zhuang & Y. Nong
China, 2003, W.Y. Zhuang & Y. Nong
Denmark, 1994, K. Hansen & S.K. Sandal
China, 2003, W.Y. Zhuang & Y. Nong
Italy, 2010, ε. Carbone
Sweden, 2013, K. Hansen & X. Wang
Finland, 1965, H. Harmaja
Germany, 1999, F. Kasparek
–
Kε0100591
Kε0100601
KM010061
Kε0100621
–
–
–
–
KM010063
Kε0100641
KM010065
Kε0100661
KM010067
KM010068
–
–
Kε0100691
Kε0100701
–
–
–
Kε0100711
Kε0100721
KM010073
–
–
KM010074
–
KM010075
KM010076
KF7175741
Kε0100771
KC012688
KC012691
Kε8231881
KM823457
Kε8231871
Kε8231891
AF072073
AF086583
AF086582
KM823458
Kε8231901
KM823459
Kε8231911
KM823460
KM823461
DQ443438
DQ443439
Kε8231921
Kε8231941
Kε8231931
AF086584
AF086586
Kε8231851
Kε8231861
KM823462
DQ443442
DQ443440
KM823463
DQ443441
KM823464
KM823465
Kε8231951
Kε8231961
Finland, 2010, ε. Carbone
China, 2003, Z.δ. Yang
Kε0100231
–
Kε8231971
DQ443450
KH.09.82 (S)
S-F249386 (Ex-H6017193)
S-F257086, dupl. TUR–A 198579
JS.08.66 (S)
KH.08.107 (S)
TUR-A 198582
KH.09.172 (S)
JS.08.55 (S)
KH.07.37 (S)
KH.09.248 (S)
KH.09.249 (S)
NV 2009.11.01 (S)
C-F-94240
H6010805
εT 10082601 (SCε, dupl. S)
KH.13.48 (S)
JS.08.18 (S)
JS.08.47 (S)
KH.09.125 (S)
NV 2008.09.16 (dupl. S)
JS.08.59 (S)
KH.09.183 (S)
KH.09.250 (S)
HMAS583571
HMAS57688
SEST-06081702
H6010806
H6010830
KH.09.153 (S)
H6003350
JS.08.63 (S)
H6003549
S-F244372 (dupl. O)
KH.11.104 (S)
GεFN 2293
S-F108342
AH21147 (εICH)
Sweden, 2009, K. Hansen & I. Olariaga
Finland, 1978, H. Harmaja
Finland, 2009, ε. Carbone
Sweden, 2008, J. Santos
Sweden, 2008, K. Hansen
Finland, 2011, ε. δahti
Sweden, 2009, K. Hansen & I. Olariaga
Sweden, 2008, J. Santos
Denmark, 2007, K. Hansen
Spain, 2009, J.δ. Teres & P.ε. Pasaban
France, 2009, J.δ. Teres
France, 2009, G. εoyne
Denmark, 2011, ε. Sasa
Finland, 1978, H. Harmaja
Spain, 2010, ε. Tabarés & S. Santamaría
Sweden, 2013, I.-δ. Walter
Sweden, 2008, J. Santos
Sweden, 2008, J. Santos
Sweden, 2009, K. Hansen & I. Olariaga
France, 2008, J. Cavet
Sweden, 2008, B. Wasstorp
Sweden, 2009, K. Hansen & I. Olariaga
Spain, 2009, F. Prieto & A. González
China, 2003, W.Y. Zhuang & Y. Nong
China, 1988, S. Wang & W.Y. Zhuang
Spain, 2003, J.δ. Pérez Butrón
Finland, 1978, H. Harmaja
Finland, 1987, P. Askola
Norway, 2009, K. Hansen & I. Olariaga
Finland, 2005, U. Salo & P. Salo
Sweden, 2008, J. Santos
Finland, 1970, δ. Fagerström
Norway, 2009, J. δorås
Sweden, 2011, J.C. Zamora & I. Olariaga
Italy, 2003, G. Jamoni
Italy, 1892, G. Bresadola
USA, 1917, A.H. Smith & R.J. Porter
Kε0100291
KM010024
Kε0100251
KM010028
Kε0100261
KM010027
JN942764
KM010078
JN942767
JN942766
Kε0100791
JN942765
KP119674
KF7175751
Kε0100301
KM010081
KM010082
KM010083
Kε0100841
Kε0100851
KM010031
Kε0100321
JN942775
–
–
Kε0100861
KF7175761
Kε0100871
Kε0100881
KM010036
Kε0100351
KF7175771
Kε0100341
KM010033
KM010037
KP006504
AF072095
Kε8231981
KM823466
Kε8231991
KM823467
Kε8232001
KM823468
JN941097
KM823469
JN941098
JN941084
Kε8232011
JN941085
–
Kε8232021
Kε8232031
KM823470
KM823471
KM823472
Kε8232051
Kε8232041
KM823473
JN941089
JN941095
DQ443444
DQ443445
Kε8232061
Kε8232091
Kε8232081
Kε8232071
KM823474
Kε8232121
Kε8232111
Kε8232101
KM823475
KM823476
–
–
O. borealis
O. brevispora as O. onotica var.
brevispora
O. brunneoparva (1)
O. brunneoparva (2)
O. brunneoparva (3)
O. brunneoparva (4)
O. brunneoparva (5)
O. brunneoparva (6)
O. bufonia (1)
O. bufonia (2)
O. bufonia (3)
O. bufonia (4)
O. bufonia (5)
O. bufonia (6)
O. bufonia (7)
O. caeruleopruinosa (1)
O. caeruleopruinosa (2)
O. caeruleopruinosa (3)
O. cantharella (1)
O. cantharella (2)
O. cantharella (3)
O. cantharella (4)
O. concinna (1)
O. concinna (2)
O. concinna (3)
O. crassa
O. daliensis (1)
O. daliensis (2)
O. flavidobrunneola (1)
O. flavidobrunneola (2)
O. flavidobrunneola (3)
O. formicarum (1)
O. formicarum (2)
O. formicarum (3)
O. formicarum (4)
O. formicarum (5)
O. fusconigra
O. integra
O. kauffmanii (1)
GenBank Accession no 4
170
Persoonia – Volume 35, 2015
Table 1 (cont.)
Taxon
Voucher
δocality / year/ collector
O. kauffmanii (2)
O. kaushalii
O. lactea
O. leporina (1)
O. leporina (2)
O. leporina (3) as O. smithii
O. leporina (4)
O. leporina (5)
O. leporina (6) as O. sp.
O. leporina (7)
O. leporina (8)
O. leporina (9)
O. leporina (10)
O. microspora
O. minor (1)
O. minor (2)
O. minor (3)
O. minor (4)
O. minor (5)
O. minor (6)
O. minor (7)
O. mirabilis (1)
MICH14409
T. δæssøe 6236 (C, dupl. BORH)
HMAS61359 (ex-MHSU 1803)
HεAS83579
HεAS83568
−
OSC 56824
OSC 56784
HεAS583570
JS.08.46 (S)
KH.09.93 (S)
JS.08.92 (S)
NV 2008.09.28 (dupl. S)
AH30502 (εICH)
H6003841
KH.10.311 (S)
H6008618
Tδ-Vorsø-0754 (C)
Cδ 950914-01 (dupl. S)
KH.98.84 (C)
C-F-83445
KH.09.188 (S)
O. mirabilis (2)
O. mirabilis (3) as O. umbrina
O. mirabilis (4)
O. mirabilis (5)
O. mirabilis (6)
O. myosotis
O. nannfeldtii (1)
O. nannfeldtii (2)
O. nannfeldtii (3)
O. nannfeldtii (4) (= O. lohjaënsis nom.
prov. Harmaja)
O. nannfeldtii (5)
O. nannfeldtii (6)
O. nannfeldtii (7)
O. nannfeldtii (8)
O. nannfeldtii (9)
O. onotica (1)
O. onotica (2)
O. onotica (3)
O. onotica (4)
O. onotica (5)
O. onotica (6)
O. onotica (7)
O. onotica (8)
O. onotica (9)
O. onotica (10)
O. oregonensis (1)
O. oregonensis (2)
O. oregonensis (3)
O. oregonensis (4) as O. rainierensis
O. oregonensis (5) as O. rainierensis
O. oregonensis (6) as O. rainierensis
O. oregonensis (7)
O. papillata (1)
O. papillata (2)
O. papillata f. pallidefurfuracea
O. phlebophora (1)
O. phlebophora (2)
O. phlebophora (3)
O. platyspora (1)
O. platyspora (2)
O. platyspora (3)
O. propinquata (1)
O. propinquata (2)
O. propinquata (3)
O. pseudoleporina (1) as O. concinna
O. pseudoleporina (2) as O. concinna
O. pseudoleporina (3) as O. concinna
O. pseudoleporina (4)
O. pseudoleporina (5)
O. pseudoleporina (6)
O. rainierensis
O. sinensis
O. smithii (1)
O. smithii (2)
KH.10.285 (S)
KH.01.09 (C)
S-F257083
NV 2008.09.14 (dupl. S)
S-F256929
H6003548
Cδ 091116-17 (S)
S-F257096
Cδ 091207-01 (S)
S-F249387 (Ex-H6017194)
USA, 1915, C.H. Kauffman
εalaysia, 1999, T. δæssøe
China, 1987, J. Z. Cao
China, 2003, W.Y. Zhuang & Y. Nong
China, 2003, W.Y. Zhuang & Y. Nong
−
USA, 1997, E.T. Peterson
USA, 1997, E.T. Peterson
China, 2003, W.Y. Zhuang & Y. Nong
Sweden, 2008, J. Santos
Sweden, 2009, K. Hansen & I. Olariaga
Sweden, 2008, J. Santos
France, 2008, N. Van Vooren
USA, 1948, A.H. Smith
Finland, 2006, U. Salo & P. Salo
Sweden, 2010, K. Hansen, K. Gillen & I. Olariaga
Finland, 1992, R. Saarenoksa
Denmark, 1982, T. δæssøe
Italy, 1995, C. δavorato
Denmark, 1998, K. Hansen
Denmark, 2007, T. δæssøe
Sweden, 2009, E. Bohus-Jensen, K. Hansen &
I. Olariaga
Sweden, 2010, K. Hansen, K. Gillen & I. Olariaga
Denmark, 2001, C. δange
Finland, 2010, ε. Carbone
France, 2008, J. Cavet
Italy, 1999, D. Bolognini
Finland, 1970, δ. Fagerström
Italy, 2009, C. δavorato
Italy, 2009, B. De Ruvo
Italy, 2009, C. δavorato
Finland, 1978, H. Harmaja
JS.08.103 (S)
NV 2008.10.01 (dupl. S)
H6002902
rh101310 (OSC)
KH.10.302 (S)
OSC 56801
OSC 56734
OSC 56759
C-F-89691
JS.08.48 (S)
KH.10.284 (S)
KH.09.132 (S)
KH.09.136 (S)
εCVE 23277
KH.98.107 (C)
rh139 (S)
Moorefun 58 (OSC, S)
εoorefun 31 (S)
OSC 56829
NSW6354 (OSC)
OSC 56745
EGS2179 (εICH)
H6003547
TUR 102134
NV 2007.09.27 (S)
JV06-385 (C)
S-F108338
K(ε)33068
KH.09.163 (S)
HK0846 (S)
JV06-656 (C-F-75309)
KH.09.99 (S)
JS.08.67 (S)
NV 2008.09.15 (dupl. S)
NSW7574 (OSC)
OSC 56749
OSC 56760
rh101910 (OSC)
εoorefun 14 (S)
OSC 56809
A.H. Smith 30553 (MICH)
HεAS61360
OSC 56799
ecv3345 (S)
Sweden, 2008, J. Santos
France, 2008, N. Van Vooren
Finland, 1972, C.-A. Haeggström
USA, 2010, R. Helliwell
Sweden, 2010, K. Hansen, K. Gillen & I. Olariaga
USA, 1997, E.T. Peterson
USA, 1996, E.T. Peterson
USA, 1996, E.T. Peterson
Denmark, 2008, H. Knudsen
Sweden, 2008, J. Santos
Sweden, 2010, K. Hansen, K. Gillen & I. Olariaga
Norway, 2009, K. Hansen & I. Olariaga
Norway, 2009, K. Hansen & I. Olariaga
Italy, 2008, ε. Carbone
Denmark, 1998, K. Hansen, T. δæssøe & C. δange
USA, 2010, R. Helliwell
USA, 2010, J. εoore
USA, 2010, J. εoore
USA, 1997, ε. Castellano
USA, 1990, D. εcKay
USA, 1996, J. Trappe
USA, 1948, E.G. Simmons
Finland, 1971, H. Harmaja
Finland, 1990, T. δindholm
France, 2007, N. Van Vooren
Denmark, 2006, δ. & J. Vesterholt
Sweden, 1949, G. Haglund & R. Rydberg
UK
Sweden, 2009, K. Hansen & I. Olariaga
Sweden, 2008, H. Kauffman
Denmark, 2006, J. Vesterholt
Sweden, 2009, K. Hansen & I. Olariaga
Sweden, 2008, J. Santos
France, 2008, J. Cavet
USA, N. S. Weber
USA, 1996, E.T. Peterson
USA, 1996, E.T. Peterson
USA, 2010, R. Helliwell
USA, 2010, J. εoore
USA, 1997, J. Spatafora
USA, 1948, A.H. Smith
China
USA, 1997, E.T. Peterson
USA, 2005, E. Vellinga
GenBank Accession no 4
ITS
δSU
KF717579
Kε0101191
–
–
–
–
–
–
–
KM010089
Kε0100901
KM010091
Kε0100921
AF072094
KM010040
Kε0100421
Kε0100391
KM010043
Kε0100441
Kε0100411
KM010038
JN942770
–
AF335111
DQ443447
DQ443448
DQ443449
AF0865732
Kε8232161
Kε8232151
DQ443443
KM823477
Kε8232131
KM823478
Kε8232141
–
KM823479
Kε8232181
Kε8232191
KM823480
Kε8232201
Kε8232171
KM823481
JN941086
Kε0100941
JN942769
KM010095
JN942768
KF717580
KF7175781
KM010096
KM010097
KM010098
Kε0100931
Kε8232211
AY500540
KM823482
JN941094
KM823483
Kε8232221
KM823484
KM823485
KM823486
Kε8232251
Kε0100451
Kε0100991
KF7175811
Kε0101001
Kε0101011
AF072067
AF072066
–
JN942773
KM010102
KP0065051
KM010103
JN942772
KM010104
–
KM010046
Kε0100481
Kε0100471
AF072087
AF072088
AF072089
AF072088
KF7175821
Kε0101051
KF7175841
Kε0100491
KM010050
EU784392
Kε0101061
KM010107
Kε0101081
Kε0101091
KM010110
Kε0101111
AF072083
AF072082
AF072081
Kε0101121
Kε0101131
AF072080
KF7175831
–
AF072063
JN942771
Kε8232241
Kε8232271
Kε8232281
Kε8232261
Kε8232231
AF086578
AF086577
JN941088
JN941090
KM823487
Kε8232291
KC012692
JN941096
KM823488
AF335121
KM823489
Kε8232311
Kε8232301
AF086597
AF086598
Kε8232321
–
Kε8232341
Kε8232331
Kε8232351
Kε8232361
KM823490
–
Kε8232381
KM823491
Kε8232371
Kε8232391
KM823492
Kε8232401
AF086593
AF086592
Kε8232441
Kε8232431
Kε8232421
Kε8232411
Kε8232451
DQ443451
JN941087
JN941093
171
I. Olariaga et al.: A monograph of Otidea
Table 1 (cont.)
Taxon
O. smithii (3)
O. smithii (4)
O. subformicarum (1)
O. subformicarum (2)
O. subformicarum (3)
O. subformicarum (4)
O. aff. subformicarum (1)
O. aff. subformicarum (2)
O. subterranea (1)
O. subterranea (2)
O. tuomikoskii (1)
O. tuomikoskii (2)
O. tuomikoskii (3)
O. tuomikoskii (4)
O. tuomikoskii (5)
O. tuomikoskii (6)
O. tuomikoskii (7)
O. tuomikoskii (8) as O. leporina
O. tuomikoskii (9)
O. tuomikoskii (10)
O. tuomikoskii (11)
O. unicisa (1)
O. unicisa (2) as O. grandis
O. unicisa (3) as O. grandis
O. yunnanensis
O. sp. ‘a’ (1)
O. sp. ‘a’ (2)
O. sp. ‘b’
Warcupia terrestris
1
2
3
4
Voucher
OSC 56753
OSC 56811
S-F242696
S-F256979
Cδ 050928-30, dupl. S-F256978
Private herb. CεP 1179, Rε 1095,
dupl. S-F256980
FH301035
FH301036
RH97 (FH)
RH69 (FH)
JS.08.68 (S)
εK200065 (S)
H6002901
JS.08.100 (S)
NV 2008.09.08 (S)
KH.09.130 (S)
KH.11.77 (S)
−
OSC 56756
OSC 56826
OSC 56761
KH.06.06 (FH)
HεAS51684
ZW Geo65-Clark (S)
HεAS 82166
εK0942 (S)
εK1081 (S)
KH.09.79 (S)
CBS 891.69
δocality / year/ collector
GenBank Accession no 4
ITS
δSU
USA, 1996, E.T. Peterson
USA, 1997, E.T. Peterson
Spain, 2012, J. Herranz & J.C. Campos
Spain, 2008, J. Fernández Vicente et al.
Italy, 2005, C. δavorato
Spain, 2009, C. ε. Pérez del Amo & R. Gil
AF072062
AF072060
KM010054
KM010051
Kε0100521
Kε0100531
AF086574
AF086572
KM823495
KM823494
Kε8232471
Kε8232461
εexico, 2007, ε. Hernández
εexico, 2007, ε.E. Smith
USA, 1997, R. Healy
USA, 1997, R. Healy
Sweden, 2008, J. Santos
Sweden, 2000, ε. Karström
Finland, 1972, R. Tuomikoski
Sweden, 2008, J. Santos
France, 2008, N. Van Vooren
Norway, 2009, K. Hansen & I. Olariaga
Sweden, 2011, ε. Prieto & I. Olariaga
−
USA, 1996, E.T. Peterson
USA, 1996, ε. εadsen & R. Davis
USA, 1996, E.T. Peterson
USA, 2006, δ. εillman
USA, Burdsall
USA, 2003, Z. Wang
China, 2003, Z.δ. Yang
Sweden, 2009, ε. Karström
Sweden, 2010, ε. Karström
Sweden, 2009, K. Hansen & I. Olariaga
Canada, 1966, J.W. Paden
Kε0100551
Kε0100561
FJ404766
FJ404767
KM010114
KM010115
KF7175851
KM010116
JN942777
JN942776
KM010117
–
AF072084
AF072086
AF072085
–
–
KM010118
–
KM010057
KM010058
Kε0101201
–
Kε8232491
Kε8232481
FJ404766
FJ404767
KM823496
KM823497
Kε8232501
KM823498
JN941091
JN941092
KM823499
AF0865883
AF086594
AF086596
Kε8232511
KC012693
DQ443446
AY789369
DQ443452
KM823500
KM823501
Kε8232521
DQ220467
Sequences from Hansen & Olariaga (2015).
The voucher specimen for AF086573 is mistakenly given as OSC 56823 in GenBank. This voucher (OSC 56823) is O. smithii based on morphological re-examination and the ITS sequence
(AF072061) deposited by the same authors. The δSU sequence AF086573 is O. leporina (Fig. 1).
The voucher specimen for AF086588 is mistakenly given as OSC 56825 in GenBank. This voucher (OSC 56825) is O. leporina based on morphological re-examination and the ITS sequence
(AF072078) deposited by the same authors. The δSU sequence AF086588 is O. tuomikoskii (Fig. 1).
ITS: Internal transcribed spacers (ITS1 and ITS2) and the 5.8S gene of the nrDNA; δSU: 28S large subunit of the nrRNA gene.
RESULTS
Alignment and ITS minisatellites
Fourty-nine ITS and 44 δSU sequences were newly generated
in this study (Table 1). In total 146 ITS sequences were utilised,
including 34 obtained from GenBank and 63 from Hansen &
Olariaga (2015). The ITS sequences were too variable to align
across all of Otidea, due to a highly polymorphic part in ITS1
and large length variability (insertions and deletions), and therefore were not included in phylogenetic analyses of the entire
genus. The ITS sequences were aligned among closely related species or species groups, and used as an aid to verify
identifications. The ITS region was especially useful in cases
were the protein-coding genes (RPB1, RPB2 and EF1) failed
to amplify, due to poor quality DNA from old or poorly treated
material. ITS sequences of the type specimens of O. mirabilis
and O. kauffmanii that could not be amplified for the multiple
genes are provided here. The all taxa δSU alignment consisted
of 167 δSU sequences, including 57 from GenBank and 68
from Hansen & Olariaga (2015) (Table 1) and 956 bp including
inserted gaps, of which 263 bp were parsimony informative
characters.
The O. borealis dataset consisted of 31 taxa, represented by
24 complete ITS-δSU and six ITS sequences, and 1 555 bp
including inserted gaps (ITS 614 bp; δSU 941 bp), of which
216 were parsimony informative characters. The O. formicarum
dataset consisted of 14 sequences and 2 531 bp (ITS 1 695 bp;
δSU 836 bp). Part of the ITS1 (805 bp) in the O. formicarum
dataset was omitted from the analyses, due to a long insertion
and tandem repeats in O. subformicarum and the two εexican
specimens, and the combined dataset thus included 1 726 bp,
of which 145 were parsimony informative characters. The insertion was composed of four tandem repeats (minisatellites) in
the four O. subformicarum sequences (212 bp in total), and in
one of them (S-F256979) the repeat was present a fifth time
(275 bp in total). In the εexican FH301036 the insertion was
extremely long (715 bp in total), of variable to random repeats.
The tandem repeats were 43 or 63 nucleotides. They were
composed of three parts (A-B-C) of 30, 20 and 13 nucleotides,
respectively, which were duplicates of the preceding part of
the ITS1 sequence. In the first two repeats the B part was
missing, whereas in the third-fifth all parts were present. The
C part was 100 % identical in all specimens and repeats; the
A part was mostly identical, but showed 3.3–6.7 % variation
in one repeat; and the B part showed 5–15 % variation in the
repeats. To ascertain the correctness of the long insertion in
the εexican specimens, the ITS sequence of FH301036 was
amplified and sequenced twice, using different sets of primers
(in one piece using ITS1-ITS4 and in two pieces using ITS55.8S / ITS3-ITS4). The two sequences were found to be identical. Unfortunately, we were not able to sequence the complete
insertion of FH301035 and the ITS1 was only sequenced in one
direction; both the part of the insertion recovered and the ITS1
were highly different from FH301036.
All taxa LSU phylogeny
The εδ analysis of the all taxa δSU dataset resulted in a single
best εδ tree of -lnδ = 6511.68. Bayesian analyses reached
an average standard deviation of split frequencies of 0.004
after 10 ε generations. A majority rule consensus tree was
constructed from the 300 000 trees sampled from the four
runs, each consisting of 75 000 trees sampled from the station-
172
Persoonia – Volume 35, 2015
O. cantharella (1) SE
100
O. cantharella (2) SE
O. cantharella (3) SE
100
O. cantharella
O. cantharella (4) FR
93
75
100
75 O. brunneoparva (1) SE
- O. brunneoparva (2) FI
O. brunneoparva (3) FI
100
76
99
O. brunneoparva
O. brunneoparva (4) SE
O. brunneoparva (5) SE
O. propinquata (1) SE
100/100
O. propinquata (2) SE
O. propinquata
O. propinquata (3) FR
O. tuomikoskii (1) SE
O. tuomikoskii (2) SE
97
100
O. tuomikoskii (3) FI
O. papillata f. pallidefurfuracea FR
O. tuomikoskii (4) SE
89
100
78
98
100
O. tuomikoskii (5) FR
98
O. tuomikoskii (6) NO
84
96
O. tuomikoskii
O. tuomikoskii (7) SE
91
O. tuomikoskii (8) US
O. tuomikoskii (9) US
100
O. tuomikoskii (10) US
O. tuomikoskii (11) US
O. pseudoleporina (1) US
O. pseudoleporina (2) US
100
O. pseudoleporina (3) US
100
O. pseudoleporina (4) US
O. pseudoleporina
O. pseudoleporina (5) US
93
O. pseudoleporina (6) US
100
100
O. leporina (3) US
O. leporina (4) US
100
O. leporina (5) US
O. crassa CHN
O. leporina (6) CHN
O. leporina
O. leporina (7) SE
100
O. myosotis FI
100
O. leporina (8) SE
O. leporina (9) SE
O. leporina (10) FR
100
O. papillata (1) FI
100
O. papillata (2) FI
O. papillata
O. alutacea (1) SE
99
O. alutacea (2) SE
100
O. alutacea (3) DK
clade 1
O. alutacea (4) SE
O. alutacea (5) US
O. alutacea (6) US
99
100
100
100
O. alutacea (7) US
clade 2
O. alutacea (8) US
O. alutacea (9) SE
98
clade 3a
O. alutacea (10) NO
100
O. alutacea (11, 30) SE
O. alutacea (12) SE
clade 3b
O. alutacea (13) DK
O. alutacea (14) SE
O. alutacea (15) CHN
75
O. alutacea complex
O. alutacea (16) CHN
100
O. alutacea (17) IT
O. alutacea (18) US
74
O. alutacea (19) US
95
-
94
97
96
clade 4
O. alutacea (20) US
O. alutacea (21) US
100
O. alutacea (22) NO
100
O. alutacea (23) ES
O. alutacea (24) FR
71
-
O. alutacea (25) CHN
O. alutacea s.str.
O. alutacea (26) CHN
98
O. alutacea (27) DK
100
O. alutacea (28) CHN
O. alutacea (29) IT
100
O. daliensis (1) CHN
100
O. daliensis (2) ES
100
100
99
O. subterranea (1) US
O. subterranea
O. subterranea (2) US
O. apophysata GE
100
O. daliensis
O. apophysata
O. platyspora (1) SE
-
98
O. platyspora (2) SE
O. platyspora (3) DK
O. platyspora
0.4
Monascella botryosa ES
Warcupia terrestris CA
Fig. 1 Bayesian inference 50 % majority rule consensus phylogram of Otidea from δSU sequence data. εaximum δikelihood bootstrap values (εδ-BP)
≥ 70 % and Bayesian posterior probabilities (PP) ≥ 95 % are shown above and below the branches, respectively. Thickened branches received support by
both εδ-BP ≥ 70 % and PP ≥ 95 %. Type collections are highlighted in bold. Country of origin for each collection is given using ISO country codes. Names of
species recognised are indicated by the vertical bars.
173
I. Olariaga et al.: A monograph of Otidea
O. bufonia (1) SE
100
O. bufonia (2) SE
O. bufonia (3) DK
100
O. bufonia
O. bufonia (4) ES
99
O. bufonia (5) FR
O. bufonia (6) FR
O. mirabilis (1) SE
O. mirabilis (2) SE
O. mirabilis (3) DK
-
O. mirabilis (4) FI
94
O. mirabilis
O. mirabilis (5) FR
O. mirabilis (6) IT
O. leporina (1) CHN
98
97 O. leporina (2) CHN
O. smithii (1) US
O. smithii (2) US
100
O. smithii
O. smithii (3) US
100
O. smithii (4) US
O. flavidobrunneola (1) FI
100
O. flavidobrunneola
O. flavidobrunneola (2) FI
100
O. flavidobrunneola (3) NO
O. brevispora
O. brevispora CHN
O. onotica (1) US
O. onotica (2) US
71
O. onotica (3) US
99
91
O. onotica (4) DK
100
O. onotica
O. onotica (5) SE
82
91
O. onotica (6, 7, 8) SE, NO
O. onotica (9) IT
O. onotica (10) DK
O. unicisa (1) US
100
100
100
100
O. unicisa
O. unicisa (2) US
O. unicisa (3) US
O. kaushalii
O. yunnanensis
O. kaushalii MY
100
O. yunnanensis CHN
100
O. minor (1) FI
100
O. minor (2) SE
100
O. minor (3) FI
O. minor (4) DK
95
71
O. minor
O. minor (5) IT
98
O. minor (6) DK
O. minor (7) DK
99
O. lactea CHN
O. lactea
O. oregonensis (1) US
O. oregonensis (2) US
O. oregonensis (3) US
100
O. oregonensis (4) US
100
O. oregonensis
O. oregonensis (5) US
O. oregonensis (6) US
95
100
100
O. sp. ‘a’ (1) SE
O. sp. ‘a’
O. sp. ‘a’ (2) SE
100
100
O. phlebophora (1) DK
100
100
O. phlebophora (2) SE
O. borealis FI
O. rainierensis US
O. concinna (1) SE
100
O. concinna
O. concinna (2) SE
-
100
O. concinna (3) ES
97
O. caeruleopruinosa (1, 3) FI, SE
100
O. caeruleopruinosa (2) ES
100
99
89
100
98
O. caeruleopruinosa
O. sinensis
O. sinensis CHN
O. subformicarum (1) ES
O. subformicarum (2) ES
O. subformicarum
O. aff. subformicarum (1) ME
O. aff. subformicarum (2) ME
98
O. formicarum (1) FI
100
97
100
O. formicarum (2) SE
O. formicarum (4) NO
O. formicarum (5) SE
91
O. sp. ‘b’ SE
99
-
O. nannfeldtii (1) IT
100
91
O. nannfeldtii (2) IT
99
100
100
99
O. nannfeldtii (6) FR
O. nannfeldtii (7) FI
O. nannfeldtii (8) US
O. nannfeldtii (9) SE
Fig. 1 (cont.)
O. sp. ‘b’
O. nannfeldtii (3) IT
O. angusta FI
O. nannfeldtii (4) FI
99
O. nannfeldtii (5) SE
98
73
-
O. formicarum
O. formicarum (3) FI
O. nannfeldtii
O. phlebophora
O. borealis
O. rainierensis
174
Persoonia – Volume 35, 2015
ary tree distribution (the first 25 % discarded as the burn-in)
(Fig. 1). The εδ and Bayesian tree topologies were congruent and recovered the same moderate to strongly supported
clades (Fig. 1). The terminal clades that constitute species we
recognise (Hansen & Olariaga 2015) have moderate to strong
support in both analyses, except for O. leporina and O. mirabilis.
A few synonymies inferred from sequences of type collections
from which we were not able to obtain the protein-coding
genes, and clear misidentifications of sequences deposited
in GenBank are evidenced by the δSU phylogeny (see Table
1). Based on δSU sequences from GenBank: the holotype of
O. crassa is nested within the O. leporina clade (= O. leporina
and O. pseudoleporina); O. lactea is suggested to be a sister
lineage to O. minor; O. sinensis is resolved as a sister species
to O. caeruleopruinosa, but without support; O. yunnanensis
forms a monophyletic group with O. kaushalii and O. unicisa;
and the type of O. onotica var. brevispora (= O. brevispora) is
supported as a sister lineage to O. onotica. The holotype of
O. mirabilis is nested among other collections of O. mirabilis
and the LSU sequence differs only in 1 bp from sequences of
collections from Scandinavia and France.
The O. formicarum and O. concinna clades in separate
ITS-LSU phylogenies
The εδ analysis of the O. formicarum dataset recovered a single
tree of -lnδ = 3579.93 (Fig. 2). Bayesian analyses reached
an average standard deviation of split frequencies of 0.003
after 3 ε generations. A majority rule consensus tree was
constructed from the 90 004 trees sampled from the four runs,
each consisting of 22 501 trees sampled from the stationary
tree distribution (the first 25 % discarded as the burn-in). The
four collections of O. subformicarum from Spain and Italy form
a distinct, strongly supported monophyletic group (εδ-BP and
PP 100 %). δikewise, the five collections of O. formicarum from
Fennoscandia, and the two collections of O. aff. subformicarum
from εexico, each form separate, strongly supported clades
(εδ-BP and PP 100 %; εδ-BP 84 %, PP 100 %, respectively).
Phylogenetic analyses of the combined ITS-δSU dataset fail
however, to resolve relationships among these three clades
with any certainty.
The combined εδ analysis of the O. borealis dataset resulted in
a single best εδ tree of -lnδ = 4230.82 (Fig. 3). Bayesian analyses reached an average standard deviation of split frequencies
of 0.005 after 3 ε generations. A majority rule consensus tree
was constructed, as for the O. formicarum dataset (above).
The supported topology (PP ≥ 95 %) did not differ from the
supported topology recovered by εδ analysis. The placement
of the single O. borealis collection is unresolved, but the ITSδSU phylogeny confirms it is genetically divergent from its sister
species (Fig. 3). Other species with a yellow outer surface of the
apothecia, O. concinna, O. minor, O. oregonensis and O. phlebophora, are each strongly supported as monophyletic (εδ
97–99 %, PP 100 %). An exception is O. minor (5) from Italy
that is resolved as a sister lineage to the rest of the O. minor
collections and O. integra without support. The ITS and δSU
sequences of O. minor (5) differ by 16 and 5 bp, respectively,
from the rest of the sequences of O. minor, which are 100 %
identical, except for the ITS sequence of O. minor (7) that
differs in 1 bp. Otidea integra is represented only by the ITS2
region (281 bp). The holotype of O. rainierensis is forming a
O. microspora US
O. rainierensis US
97
100
O. kauffmanii (1) US
98
100 O. kauffmanii (2) US
O. integra IT
O. minor (7) DK
O. minor (3) FI
99
O. minor (1) FI
100
O. minor (6) DK
O. formicarum (5) SE
O. minor (2) SE
100
O. formicarum (4) NO
100
100
O. minor (4) DK
100
O. formicarum (3) FI
O. minor (5) IT
O. oregonensis (1) US
O. formicarum (2) SE
O. oregonensis (3) US
O. oregonensis (2) US
O. formicarum (1) FI
84
100
100
100
O. oregonensis (4) US
98 O. oregonensis (5) US
99
O. oregonensis (6) US
O. aff. subformicarum (1) ME
O. aff. subformicarum (2) ME
100
100
O. oregonensis (7) US
100
100
O. subformicarum (2) ES
100
100
O. subformicarum (3) IT
100
100
O. subformicarum (4) ES
O. sp. ‘a’ (1) SE
O. sp. ‘a’ (2) SE
O. phlebophora (3) UK
O. phlebophora (2) SE
O. phlebophora (1) DK
O. subformicarum (1) ES
O. borealis FI
O. concinna (3) ES
97
99 O. concinna (1) SE
O. concinna (2) SE
O. nannfeldtii (7) FI
100
100
O. angusta FI
100
100 O. nannfeldtii (5) SE
0.0080
Fig. 2 The single best tree resulting from the εaximum δikelihood analysis
of the ITS-δSU regions of the O. formicarum clade. εδ bootstrap values
(εδ-BP) are shown above nodes and Bayesian posterior probabilities (PP)
below nodes. Thickened branches are nodes with high support (εδ-BP ≥ 75;
PP ≥ 95). Type collections are in bold.
100
100
O. caeruleopruinosa (2) ES
O. caeruleopruinosa (1) FI
0.02
Fig. 3 The single best tree resulting from the εaximum δikelihood analysis
of the ITS-δSU regions of O. borealis and closely allied species. εδ bootstrap
values (εδ-BP) are shown above nodes and Bayesian posterior probabilities
(PP) below nodes. Thickened branches are nodes with high support (εδ-BP
≥ 75 %; PP ≥ 95 %). Type collections are in bold.
I. Olariaga et al.: A monograph of Otidea
strongly supported clade with the holotype of O. kauffmanii
and a paratype of O. microspora (εδ 97 %, PP 100 %). The
O. kauffmanii and O. microspora types are only represented
by ITS and are therefore not included in any of our previous
analyses. We conclude O. kauffmanii is a synonym of O. rainierensis and O. microspora a doubtful name (see further under
Taxonomy). Otidea concinna is strongly supported as a sister
group to the rest of the ingroup, but otherwise the relationships
among the species are without support.
The phylogenetic results, and the ITS sequence similarity and
divergence (for species identification), will be further discussed
where applicable in the descriptive notes below.
Morphological characters for species delimitation
All Otidea species recognised by concordance of our four genes
phylogenies (Hansen & Olariaga 2015) can be recognised by
a combination of morphological characters. We evaluated the
characters in the context of the phylogeny and discovered
several new characters. The apothecial shape, colours, and
spore characters (size, shape, ornamentation) are important
for species identification, but to distinguish closely related species (or otherwise morphologically similar species) additional
characters are needed. These are the shape of the paraphyses,
ectal excipulum structure, type of exudates on the medullary
excipulum hyphae, resinous exudates on the outer surface of
the ectal excipulum and on the mycelium at the base of the
apothecia, and their possible reactions in εδZ and KOH (see
further in Hansen & Olariaga 2015). Below we provide details
on the resinous exudates, and their reactions in KOH and εδZ,
because they largely have been overlooked.
175
Excipular resinous exudates and reactions in MLZ and KOH
A resinous exudate is here used for a substance that is released
from the cells and in many species is vulnerable to common
mountants, but not water (following Huhtinen 1990). In Otidea
the exudates are concentrated in the excipulum cells, and on
the tomentum and mycelium at the base of the apothecia.
The resinous exudates are deposited on the outside of the
cell or hyphal walls. Harmaja (2009a) introduced the reaction
of coloured resinous exudates on the outermost cells of the
ectal excipulum in εδZ as a taxonomic character. Our study
revealed in addition, different reaction patterns of exudates
on the medullary excipulum cells and on the mycelium at the
base of the apothecia (including the tomentum), extending out
among the soil particles, which turned out to be diagnostic for
some species (see under εycelium at the base of the apothecia). In the medullary excipulum, most frequently scattered,
golden brown, resinous exudates are present at septa, e.g. in
O. alutacea, O. leporina, O. nannfeldtii (Fig. 4a). Otidea bufonia
has unique exudates, wrapping some hyphae and appearing
striate (referred to as ‘fingerprint-like’ by Korf & Zhuang (1991)),
sometimes forming big crystal-like aggregates (Fig. 4b, c). In
contrast, the sibling species, O. mirabilis, has only sometimes
biflabellate crystal-like exudates in the medullary excipulum
(Fig. 4d). Otidea papillata also possesses unique brown exudates, embedding some hyphae of the medullary excipulum
and sometimes appearing rod-like.
Small, resinous drops or amorphous matter are present in variable amounts on the outer surface of the apothecia of nearly
all Otidea species. In most species they are abundant and
Fig. 4 εedullary excipulum resinous exudates in Otidea. a. Hyphae with golden brown resinous exudates at septa in O. leporina (KH.11.02), in water*;
b, c. brown crystal-like exudates in O. bufonia (KH.07.37) in water †: b. overview; c. close-up of hyphae wrapped in striate exudates; d. close-up of biflabellate
crystal-like exudates in O. mirabilis (GεFN 1951, holotype), in water† — Scale bars = 10 µm; * = fresh material; † = dried material.
176
Persoonia – Volume 35, 2015
Fig. 5 Reactions of resinous exudates on the outermost ectal excipulum cells in Otidea. a. O. nannfeldtii † (H6010804, holotype of O. angusta), in water (left)
and converting into amber drops in εelzer’s reagent (right); b. O. bufonia* (KH.09.171), in water (left) and converting into reddish particles in εelzer’s reagent
(right); c. O. borealis† (S-F242694, holotype), in water (left) and turning bright yellow in KOH (right); d. O. nannfeldtii † (H6010804), in water (left) and turning
reddish brown in KOH (right); e. O. pseudoleporina† (rh101910, holotype), in water (left) and converting into reddish grey drops in KOH (right); f. ectal excipulum
cells showing a gelatinous sheath in O. formicarum* (KH.11.104). — Scale bars = 10 µm; * = fresh material; † = dried material.
easy to observe in water, but scarce and difficult to detect in a
few (O. alutacea, O. formicarum). One species, O. kaushalii,
has a unique type of exudate, i.e. crystal-like, oblate spheroid,
striate bodies, with a constricted centre. The possible reaction
of the exudates in εδZ and KOH is useful to separate certain
species or groups. Using these characters requires experience.
The reaction appears to vary depending on the amount of exudate and the concentration of εδZ. As Harmaja (2009a), we
observed two types of reactions in εδZ: i) Resinous exudates
dissolve and coalesce into spheroid drops, ‘amber drops’, that
contain hyaline bubbles (Fig. 5a). The ‘amber drops’ are overlooked if the ectal excipulum is directly mounted in εδZ, since
the exudates coalesce instantly and the drops can be washed
away. The reaction is best observed if εδZ is added to a water
mount. This reaction is present in many species. ii) Resinous
exudates partly convert into small reddish particles (Fig. 5b).
The reaction is often subtle and visible only in certain parts of
a mount. This reaction is typical in O. bufonia, O. mirabilis and
O. smithii. In some species the exudates do not react in εδZ
or they simply dissolve.
I. Olariaga et al.: A monograph of Otidea
177
Fig. 6 εycelium at the base of the apothecia and extending out in the substrate in Otidea†. a, b. O. flavidobrunneola (H6010806, holotype): a. resinous exudates on the hyphal walls in water; b. dissolved in εelzer’s reagent; c, d. O. borealis (S-F242694, holotype): c. pale yellow in water; d. turning bright yellow in
KOH. — Scale bars = 10 µm; † = all dried material.
In this study we detected three discriminative reactions of resinous exudates in 10 % KOH, which we propose as a novel
taxonomic character: i) In water the resinous exudates range
from yellow to dark reddish yellow (Fig. 5c, left) and in KOH
these dissolve completely ± exuding a yellow pigment, or partly
with the rest turning distinctly brighter yellow (Fig. 5c, right). This
reaction occurs in O. concinna and closely related species, viz.
O. borealis, O. caeruleopruinosa, O. flavidobrunneola, O. kaushalii, O. minor and O. oregonensis, and slightly less strikingly in
O. unicisa. ii) The resinous exudates are yellow-brown in water
(Fig. 5d, left) and turn reddish brown in KOH (Fig. 5d, right).
This reaction has been observed in O. nannfeldtii and O. propinquata. iii) The resinous exudates, yellow brown in water
(Fig. 5e, left), partly dissolve in KOH, and partly coalesce into
heterogeneous, pale reddish grey drops, with bubbles inside
(Fig. 5e, right). These drops are similar to the ‘amber drops’
observed in εδZ in many species, except for the pale red colour.
This reaction has been observed in O. leporina and O. pseudoleporina. A number of species have the outermost cells of the
ectal excipulum sometimes covered with a thin gelatinous
sheath (Fig. 5f).
Mycelium at the base of the apothecia
All Otidea species studied showed a conspicuous tomentum
covering the base of the apothecia and spreading out in the
substrate. In the microscopic descriptions we refer to both as
the basal mycelium. The hyphae are septate, straight and frequently branch and anastomose. No rhizomorphs have been
observed, although slender hyphal threads are sometimes
formed. Resinous exudates are often present on the surface,
especially on the mycelia extending out in the substrate, and
can appear like hyphal ornamentation.
Harmaja (2009a) proposed the colour of the basal tomentum
as a taxonomic character, but microscopic features of the basal
mycelium have been largely overlooked and have not been
included in species descriptions. Two types of characters are
useful for species identification: i) Resinous exudates occurring
on the hyphal walls (Fig. 6a). These can be inconspicuous or
nearly absent in some species, such as in O. alutacea s.l., in
which only a few refractive drops or minute hyaline exudates
are sometimes present. Resinous exudates are also scarce in
some species of the O. concinna clade. εany other species
show hyphae densely covered with resinous exudates, such as
O. formicarum, O. propinquata and O. tuomikoskii. The shape of
the exudates is variable, i.e. rod-shaped, hemispherical, conical
or bipyramidal. They dissolve and completely disappear in εδZ
(Fig. 6b). Sometimes doubts can arise about the nature of the
differently shaped exudates, but the facts that they detach from
the hyphal wall when the mount is squashed and dissolve in
εδZ, show these are not true ornamentation, i.e. not part of the
hyphal wall. ii) The hyphal wall turns yellow in KOH (Fig. 6d;
see Fig. 6c in water). We have observed this reaction clearly
in O. borealis and O. onotica. It can be observed in isolated
hyphae, but is more conspicuous when a mass of hyphae is
observed together. It can also be observed macroscopically.
178
TAXONOMY
Otidea (Pers.) Bonord., Handb. εykol.: 205. 1851
≡ Peziza (unranked) Otidea Pers., εycol. Eur. 1: 220. 1822.
≡ Peziza (unranked) Cochleatae Fr., Syst. εycol. 2: 46. 1822: Fr. loc. cit.
Type species. Otidea onotica (Pers. : Fr.) Fuckel, indicated by Saccardo,
Bot. Centralbl. 18: 215. 1884 (‘P. onotica Pers.’).
= Flavoscypha Harmaja, Karstenia 14: 107. 1974.
Type species. Peziza phlebophora Berk. & Broome.
= Otideopsis B. δiu & J.Z. Cao, Shanxi Univ. J., Nat. Sci. Ed. 4: 70. 1987.
Type species. Otideopsis yunnanensis B. δiu & J.Z. Cao.
Apothecia small to large, 3–75 mm high, 4–80 mm wide, often
in fascicles or caespitose, epigeous, cup- to ear-shaped and
split to the base on one side, less often entire, stipitate or not;
or hypogeous and enclosed. Hymenium white, yellow, ochre,
brown, almost black, often with pink stains. Receptacle surface
concolorous or with similar colours as hymenium, sometimes
with purplish, greenish or bluish tones, with conical to broadly
conical warts or pustules, less often smooth or furfuraceous,
concolorous or darker than the background. Base of the apothecium tomentose, mycelium white, ochre, yellow, orange or
brown, extending out in the substrate, base ribbed-veined in
a few species. Spores uniseriate, ellipsoid, oblong or fusoid,
typically with 2 guttules, sometimes with a few smaller granules, smooth (or verruculose in SEε), rarely spinose or with
low ridges, with de Bary bubbles in εδZ and Cotton Blue when
dried, thin-walled to slightly thick-walled, hyaline to very pale
brown. Paraphyses typically curved to hooked, rarely straight,
sometimes with notches or swollen at the apices, septate, typically containing refractive small guttules at the apices, fading in
colour and collapsed when dried. Asci cylindrical, operculate,
8-spored, 116–275 × 8–19 µm, with pleurorhynchous base.
Subhymenium c. 100–150 µm thick, of dense textura intricata,
hyphae sometimes swollen, often with scattered pigmented
exudates at septa. Medullary excipulum 400–1500 µm thick,
of textura intricata, hyphae cylindrical to slightly swollen, thinwalled to thick-walled, hyaline to pale brown, often with pigmented resinous exudates at septa. Ectal excipulum 70–150
µm thick, of textura angularis, less often of textura prismatica.
Surface with warts up to 180 µm high, formed by fasciculate, short hyphoid hairs of globose to elongated cells, or of
textura globulosa-angularis with single hyphoid hairs. Resinous
exudates often present on the surface, yellow to dark brown,
sometimes dissolving in εδZ, turning reddish or into brownish
yellow amber drops, sometimes changing colour in KOH. Basal
mycelium of septate, straight hyphae, that frequently branch
and anastomose, turning yellow or not in KOH, often covered
with pigmented, small, resinous exudates.
Ecology & Distribution — see Hansen & Olariaga (2015).
Key to species of Otidea
We were not able to study and interpret the following Chinese
species and these are therefore not treated nor included in the
key: Otidea bicolor W.Y. Zhuang & Zhu δ. Yang, O. kunmingensis W.Y. Zhuang, O. olivaceobrunnea Harmaja, O. sinensis J.Z.
Cao & δ. Fan, O. subpurpurea W.Y. Zhuang and O. tianshuiensis J.Z. Cao, δ. Fan & B. δiu. For O. integra (Bres.) Harmaja
see notes under O. phlebophora.
1. Ascomata hypogeous, globose to subglobose, truffle-like
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. O. subterranea
1. Ascomata epigeous, cup-shaped to ear-shaped, split or
entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Spores ornamented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2. Spores smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3. Spores with fine warts ± ridges . . . . . . . . . . 17. O. unicisa
3. Spores spiny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Persoonia – Volume 35, 2015
4. Spores 14 –17 × 7–9 µm . . . . . . . . . . . . . 16. O. kaushalii
4. Spores 16.5–20 × 7.6–10 µm . . . . . . 18. O. yunnanensis
5. Spores δm > 17 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5. Spores δm < 17 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
6. Apothecia broadly ear-shaped, split; typically ochraceous
yellow to ochre orange; often associated with Cudonia in
mossy Picea forests . . . . . . . . . . . . . . 11. O. cantharella
6. Apothecia cup-shaped, split or entire; brown; under Picea
or other trees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Apothecia entire; reddish brown; basal mycelium normally
with abundant minute resinous exudates; ectal excipulum
hyphoid hairs often with a gelatinous sheath (more easily
seen in εδZ); with Picea . . . . . . . . . 12. O. propinquata
7. Apothecia split or entire; purple or ochre-brown; basal
mycelium without or with sparse resinous exudates; ectal
excipulum hyphoid hairs without a conspicuous sheath;
with angiosperms. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Apothecia split, up to 75 mm wide; spores broadly ellipsoid
to oblong, Qm = 1.7–1.8; associated with Fagaceae . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. O. platyspora
8. Apothecia entire or split, up to 16 mm diam; spores narrowly
ellipsoid to fusoid, Qm = 1.9–2.1; most likely associated
with Betulaceae and Salicaceae . . . . . . . . . . . . . . . . . . 9
9. Apothecia deeply cup-shaped, split; ectal excipulum without
resinous exudates . . . . . . . . . . . . . . . . 1. O. apophysata
9. Apothecia shallowly cup-shaped, usually entire; ectal excipulum surface with abundant resinous exudates . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. O. daliensis
10. Receptacle surface with bright citrine yellow tones in young
apothecia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
10. Receptacle surface without citrine yellow tones . . . . . 15
11. Spores Qm = 1.7– 2; apothecial base at most wrinkled,
without high ribs or veins . . . . . . . . . . . . . . . . . . . . . . . 12
11. Spores Qm = 2–2.3; apothecial base with ribs or strongly
veined at least in some apothecia . . . . . . . . . . . . . . . . 14
12. Receptacle surface ochraceous yellow; basal mycelium
yellow in KOH; spores Wm = 6.5 µm, Qm = 1.7 . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25. O. borealis
12. Receptacle surface citrine yellow; basal mycelium unchanged in KOH; spores Wm = 5.6–6 µm, Qm = 1.8–2 . . . . . 13
13. Apothecia sometimes entire, sometimes with blunt ribs at
the base; North America . . . . . . . . . . 31. O. oregonensis
13. Apothecia split, without ribs at the base; Europe. . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27. O. concinna
14. Apothecia mostly entire, base strongly ribbed-veined and
anastomosing in all the apothecia . . . 32. O. phlebophora
14. Apothecia mostly split, base with a few ribs-veins, smooth
in some apothecia . . . . . . . . . . . . . . . . . . . . 30. O. minor
15. At least some paraphyses straight or curved, claviform
to almost capitate at apices; and/or resinous exudates of
the ectal excipulum yellow or reddish yellow, turning bright
yellow in KOH; apothecia split or not . . . . . . . . . . . . . 16
15. Paraphyses not as such; resinous exudates of the ectal
excipulum, when present, not turning bright yellow in KOH;
apothecia always split . . . . . . . . . . . . . . . . . . . . . . . . . 20
16. Apothecia shallowly cup-shaped and irregular, entire . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29. O. lactea
16. Apothecia ear-shaped or deeply cup-shaped, split . . . 17
17. Paraphyses often claviform or capitate at apices, 3–9 µm
broad; spores Qm = 1.6–1.8 . . . . . . . . . . . . . . . . . . . . 18
17. Paraphyses at most slightly swollen at apices, 2–5 µm
broad; spores Qm = 1.9–2 . . . . . . . . . . . . . . . . . . . . . . 19
18. Basal mycelium turning yellow in KOH; Europe . . . . . . .
. . . . . . . . . . . . . . . . . . . (see under O. borealis) O. sp. ‘a’
179
I. Olariaga et al.: A monograph of Otidea
18. Basal mycelium not yellow in KOH; North America . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33. O. rainierensis
19. Spores δm = 11.2–11.8 µm; sometimes receptacle with a
bluish hue, ochraceous grey when dried; basal tomentum
light ochre in dried specimens . . 26. O. caeruleopruinosa
19. Spores δm = 10–10.6 µm; receptacle without bluish hue
when fresh, reddish brown when dried; basal tomentum
orange-ochre in dried specimens 28. O. flavidobrunneola
20. Spores δm < 12 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
20. Spores δm > 12 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
21. Apothecia dark brown with lilaceous tones; Asia . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23. O. purpurea
21. Apothecia not dark brown, without lilaceous tones; Asia,
Europe or North America . . . . . . . . . . . . . . . . . . . . . . . 22
22. εedullary excipulum with reddish brown resinous exudates
scattered among and covering some hyphae; ectal excipulum of textura prismatica to textura intricata; receptacle surface with contrasting brown warts . . . . . . . 6. O. papillata
22. εedullary excipulum sometimes with yellowish brown resinous exudates at septa, not covering hyphae; ectal excipulum of textura angularis; receptacle surface with contrasting warts or not . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23. At least some warts higher than 85 µm; basal tomentum
orange-ochre when dried; apothecial sections often yellow
in KOH; without resinous exudates at septa in the medullary
excipulum . . . . . . . . . . . . . . . . . . . . . . . 9. O. tuomikoskii
23. Warts up to 85 µm; basal tomentum pale ochre or yellow
when dried; apothecial sections not yellow in KOH; sometimes with resinous exudates at septa in the medullary
excipulum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24. Hymenium with distinct yellow or orange tones, ochreyellow to pinkish orange; resinous exudates on the ectal
excipulum partly dissolving into reddish grey, heterogeneous drops in KOH or basal mycelium yellow in KOH . 25
24. Hymenium without or with weak orange tones; resinous
exudates sometimes turning reddish brown but not dissolving into drops in KOH; basal mycelium not turning yellow
in KOH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
25. Resinous exudates on the ectal excipulum partly dissolving into reddish grey heterogeneous drops in KOH; basal
mycelium not turning yellow in KOH; North America . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. O. pseudoleporina
25. Resinous exudates on the ectal excipulum not dissolving
into reddish grey heterogeneous drops in KOH; basal mycelium turning yellow in KOH; Asia . . . . . 19. O. brevispora
26. Hymenium sometimes with pink tones; apothecia yellowish
ochre to brown; narrowly ear-shaped in the beginning . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14. O. nannfeldtii
26. Hymenium without pink tones; apothecia reddish brown to
orange-brown; broadly ear-shaped . . . . . . . . . . . . . . . 27
27. Spores δm = 10 –10.7 µm; Qm = 1.6 –1.7 . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. O. formicarum
27. Spores δm = 11.1–11.7 µm; Qm = 1.7–1.9 . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . 15. O. subformicarum
28. Spores Qm < 1.8; ear-shaped . . . . . . . . . . . . . . . . . . . 29
28. Spores Qm > 1.8; ear- or cup-shaped . . . . . . . . . . . . . 30
29. Apothecia dark brown, sometimes with olivaceous tint; paraphyses mostly with distinct notches . 10. O. brunneoparva
29. Apothecia cinnamon brown, without olivaceous tint; paraphyses not or slightly notched . . . . . . . . . . 7. O. leporina
30. Apothecia ochraceous yellow, hymenium often with pink
tones; basal mycelium turning yellow in KOH . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22. O. onotica
30. Apothecia pale or dark brown, without pink tones; basal
mycelium not turning yellow in KOH . . . . . . . . . . . . . . 31
31. Receptacle medium brown; ectal excipulum resinous exudates absent or scarce, light yellowish brown; basal mycelium without dark brown resinous exudates . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. O. alutacea s.l.
31. Receptacle dark purple brown; ectal excipulum resinous
exudates abundant, dark brown; basal mycelium with dark
brown resinous exudates . . . . . . . . . . . . . . . . . . . . . . 32
32. Spores Qm < 2, ellipsoid; apothecia mostly ear-shaped;
North America . . . . . . . . . . . . . . . . . . . . . . 24. O. smithii
32. Spores Qm > 2, ellipsoid-fusoid; apothecia mostly cupshaped, split; Eurasia and North America . . . . . . . . . . 33
33. Receptacle strikingly purple-violaceous (fresh); medullary
excipulum without, or rarely with flabellate crystal-like exudates, forming cross-like aggregates; under conifers on calcareous ground . . . . . . . . . . . . . . . . . . . . 21. O. mirabilis
33. Receptacle mostly without purple tones; medullary excipulum with striate exudates covering some hyphae, sometimes forming crystal-like aggregates; often under deciduous trees or on acidic ground . . . . . . . . . . 20. O. bufonia
Based on genealogical concordance phylogenetic species
recognition (GCPSR: Taylor et al. 2000), using the four loci,
RPB1, RPB2, EF1 and δSU rDNA, we delimited 25 species
within Otidea (see Hansen & Olariaga 2015). In addition eight
species were recognised by genetic divergence from their sisters. Twenty-eight of these are treated and discussed below,
along with O. brevispora, O. lactea, O. subterranea and O. yunnanensis included in our LSU phylogeny, and O. purpurea that
has only been studied morphologically. The species are presented following their phylogenetic relationships, inferred from
our combined three- and four-gene analyses (f. 3 in Hansen
& Olariaga 2015).
Otidea platyspora clade
Apothecia disc-shaped, cup-shaped and split, or globose and
hypogeous, brown. Spores large, exceeding 20 µm, except
14–16.5 µm if hypogeous. Basal mycelium smooth or with very
sparse resinous exudates.
Species — Otidea apophysata, O. daliensis, O. platyspora,
O. subterranea.
1. Otidea apophysata (Cooke & W. Phillips) Sacc., Syll. Fung.
8: 96. 1889
Basionym. Peziza apophysata Cooke & W. Phillips in Cooke, Grevillea 5:
60. 1876.
≡ Pseudotis apophysata (Cooke & W. Phillips) Boud., Hist. Classific. Discomyc. Europe: 52. 1907.
Holotype. England, Shrewsbury, in a damp ditch, 1876, W. Phillips (K(ε)
30410 ex Herb. Phillips). Isotype (K(ε) 167215 ex Herb. Cooke) !
Misapplied names
– Otidea felina sensu Boudier, Icon. εycol. livr. 29: n°. 512, pl. 331. 1910
(preliminary text with ‘circulaires’).
Apothecia solitary to caespitose, 8–30 mm high, up to 15 mm
wide, initially ear-shaped, then soon expanding and becoming cup-shaped, split, sessile or stipitate. Hymenium purple
brown (6D3, 6D4), when dried dark orange brown (6E7, 6F7).
Receptacle surface pale greyish orange (6C5), purple brown
(6D4), when dried dark orange brown (6E7, 6F7), furfuraceous.
Warts scarce to absent, low. Stipe absent. Basal tomentum and
mycelium whitish to pale orange grey (5B2). Spores narrowly ellipsoid to fusoid, narrowing toward the poles, sometimes inequilateral, with two large guttules, and often with several smaller
guttules, smooth, hyaline, 20–24.5 × 9–11 µm (δm = 21.6 µm,
Wm = 10 µm, Qm = 2.1; n = 1). Paraphyses curved to hooked,
seldom straight, slightly enlarged at apices, 3–4(–5) µm wide,
180
sometimes with a sinuous underside or with 1–2 notches, frequently branching, entangled and interconnected, when dried
containing small, refractive, hyaline guttules. Asci 172–197 ×
12–13 µm. Apothecial section 700–850 µm thick. Medullary
excipulum of textura intricata, 400 – 500 µm thick, hyphae
slightly thick-walled, 5–9 µm wide, pale brown. Ectal excipulum
of textura angularis, 70–110 µm, cells thin-walled, yellowish
brown, 18–33 × 13–22 µm. Surface with low warts, up to 40
µm high, cells ovoid to globose, constricted at septa, 8–13 µm
wide. Resinous exudates absent. Basal mycelium of 4–5 µm
broad, pale brown hyphae, sometimes with oily refractive drops
on the surface.
Specimens examined. gErmany, Nordrhein, Herten, 1 Sept. 1999, F. Kasparek, private herb. Kasparek s.n. (dupl. S-F257062).
Notes — Otidea apophysata is characterised by deeply cupshaped, split, brown apothecia, and large, ellipsoid to fusoid
spores. Otidea daliensis is a closely related species, distinguished by darker brown, shallowly cup-shaped, usually entire
apothecia, and abundant dark brown resinous exudates on the
outermost cells of the ectal excipulum. Otidea platyspora has
also brown apothecia and large spores, but it differs from O. apophysata in the larger apothecia, partly buried in the substrate,
broadly ellipsoid to oblong spores, and non-entangled paraphyses without notches.
The name O. apophysata has been misapplied twice for O. daliensis (Boudier 1909b, Pérez-Butrón & Fernández-Vicente
2008). The type material of O. apophysata lacks resinous exudates on the ectal excipulum, which clearly distinguishes it
from O. daliensis. An original painting by W. Phillips, based
on the type material of O. apophysata and preserved at RBG
Kew (reproduced in Parslow & Spooner 2013), shows typical
brown, split, deeply cup-shaped apothecia. The illustration by
Cooke (1878, f. 350), based on drawings and (likely dried)
specimens communicated by W. Phillips, shows slightly darker
apothecia than typical.
Otidea apophysata is only known from very few reports from
France (Boudier 1910 as O. felina), Germany (Häffner &
Winterhoff 1989, Kasparek 2000), Belgium and Spain (Van
Vooren 2011a). In εid to South Britain it is widely distributed,
with collections from fourteen different localities (Parslow &
Spooner 2013). Unlike most Otidea species, O. apophysata
shows preference for damp habitats, and might be associated
with Alnus and Populus (Häffner & Winterhoff 1989, Parslow
& Spooner 2013).
2. Otidea daliensis W.Y. Zhuang & Korf, εycotaxon 35: 300.
1989
Holotype. China, Yunnan, Dali, Hudiequan Park, alt. 2100 m, on bare soil
under seedlings of Plantago major, 5 Nov. 1988, R.P. Korf, L.S. Wang & W.Y.
Zhuang (HεAS 57688). Isotype (CUP-CH 2532).
Misapplied names
– Pseudotis apophysata sensu Boudier, Icon. εycol. livr. 24: n°. 471, pl. 332.
1909 (preliminary text with ‘circulaires’).
– Otidea apophysata sensu Pérez-Butrón & Fernández-Vicente, Errotari 5:
37. 2008.
Apothecia gregarious, up to 9 mm high, 3–16 mm wide, initially
cup-shaped, sometimes split, then becoming shallowly cupshaped, sessile or stipitate. Hymenium when dried dark purple
brown (7F5, 7F6) to dark brown (6F5). Receptacle surface
dark purple brown (7F5, 7F6), when dried dark brown (6F5),
furfuraceous. Warts absent or very low. Stipe if present very
short. Basal tomentum and mycelium whitish to pale orange
grey (5B2). Spores broadly ellipsoid to ellipsoid and narrowing
toward the poles, sometimes inequilateral, with two large guttules, and often with several smaller granules, smooth, hyaline,
Persoonia – Volume 35, 2015
(19.5 –)20.5 – 23 × 10.5 –12(–13) µm (δm = 21.2 – 21.5 µm,
Wm = 10.7–11 µm, Qm = 1.9–2.1; n = 2). Paraphyses curved to
hooked, sometimes slightly enlarged at apices, 2.5–3.5(–5) µm
wide, sometimes with slightly swollen areas, apices sometimes
embedded in a brown matter, when dried containing small,
refractive, brown guttules. Asci 199–212 × 15–17 µm. Apothecial section 600–850 µm thick. Subhymenium c. 90–110 µm
thick, of dense textura intricata, visible as a darker zone, cells
cylindrical to swollen, with scattered brown resinous exudates
at septa. Medullary excipulum of textura intricata, 300–400 µm
thick, hyphae 4–6.5 µm wide, slightly thick-walled, pale brown,
with brown resinous exudates at septa. Ectal excipulum of
textura angularis, 90–120 µm thick, cells thin-walled, yellowish
brown, 18–28 × 11–28 µm. Surface with broadly conical warts.
Non-warted parts with 2–5-celled hyphoid hairs, with claviform
uppermost cell, more rarely cylindrical, constricted at septa,
6–9 µm wide. Resinous exudates abundant, dark brown, partly
dissolving in εδZ. Basal mycelium of 3.5–4 µm wide, hyaline
hyphae, with yellowish brown, small, resinous exudates.
Specimens examined. FranCE, 1869, L. Quélet (UPS F-629790). – Spain,
Basque Country, Bizkaia, Galdames, Presa de Aguas Juntas, sandy soil
under Populus nigra, 11 Aug. 2003, J.L. Pérez Butrón, SEST-03071103; 17
Aug. 2006, SEST-06081702.
Notes — Otidea daliensis is recognised by small, usually
entire, dark purple brown, shallowly cup-shaped apothecia,
large ellipsoid spores often narrowing toward the poles, and
abundant brown resinous exudates on the ectal excipulum. This
species has been confused with O. apophysata (see O. apophysata).
The first known report of O. daliensis was by Boudier (1909b),
as Pseudotis apophysata. The plate 332 (n°. 471) shows the
typical dark purple, shallowly cup-shaped apothecia, in contrast
to O. apophysata, depicted in plate 331 (n°. 512) as O. felina
(Boudier 1910). εornand & Courtecuisse (2005) proposed a
provisional name, O. boudieri, for the Boudier P. apophysata
plate (= O. daliensis, n°. 332). Zhuang & Korf (1989) described
O. daliensis without comparing it to O. apophysata. εaterial with
small, shallowly cup-shaped apothecia and darker colour was
reported from the Iberian Peninsula as O. apophysata (PérezButrón & Fernández-Vicente 2008), and thus had similarities
with O. daliensis and Boudier’s plate 332. Van Vooren (2011a)
considered the Iberian O. apophysata material to represent
O. daliensis. After restudying the Iberian material and comparing it to O. apophysata, we agree with that statement. δSU
sequences obtained from the Iberian material and from the
Chinese holotype of O. daliensis are identical.
3. Otidea platyspora Nannf., Ann. Bot. Fenn. 3: 317. 1966.
— Fig. 7
Holotype. SwEdEn, Uppland, Djursholm, Oct. 1951, A. Zander, Fungi Exs.
Suec. 3284 (UPS F-005428). Isotype (S-F88395) !
Misapplied names
– Otidea cochleata sensu Boudier, Icon. εycol. livr. 21: n°. 461, pl. 329. 1908
(preliminary text with ‘circulaires’).
Apothecia caespitose, 60–70 mm high, 40–75 mm wide, initially ear-shaped, then soon expanding and becoming deeply
cup-shaped, split, sessile or shortly stipitate. Hymenium initially
yellowish brown (5C6, 5C7), pale greyish brown (5C3, 5C4) to
dark brown (6F3–6F6), when bruised margin blackish, when
dried brownish ochre (5B4, 5B5). Receptacle surface dark ochre
brown (5D7, 5D8), slightly hygrophanous, in drying pale ochre
brown (5B6), when dried yellowish brown (5C6, 5C7), sometimes wrinkled at the base, finely furfuraceous in the margin.
Warts absent. Stipe not well developed. Basal tomentum and
mycelium whitish to pale brown (5A3). Spores broadly ellipsoid
I. Olariaga et al.: A monograph of Otidea
181
Fig. 7 Otidea platyspora*. a. Apothecia; b. spores; c. paraphyses; d. ectal excipulum (a: KH.10.183; b–d: KH.09.163). — Scale bars = 10 µm; * = all fresh
material.
to oblong, rarely slightly inequilateral, with two large guttules,
often with several smaller guttules, smooth, hyaline, 18–22 ×
(9.5–)10.5–12 µm (δm = 19.8–20.7 µm, Wm = 10.9–11.6 µm,
Qm = 1.7–1.8; n = 6). Paraphyses curved to hooked, of the
same width or slightly enlarged at apices, 2.5 – 3.5(– 5) µm
wide, without notches, rarely with a slightly swollen area on
the underside, when fresh containing small, refractive, light
brownish yellow guttules; when dried tiny, light yellow granules.
Asci 168–213 × 14–19 µm. Apothecial section 700–1700 µm
thick. Subhymenium 80–100 µm thick, visible as a darker zone,
cells cylindrical, densely arranged, with scarce yellowish brown
resinous exudates at septa. Medullary excipulum of textura
intricata, 550–1100 µm thick, hyphae 3.5–10 µm wide, sometimes slightly swollen, thin-walled, light yellowish brown. Ectal
excipulum of textura angularis, 70–90 µm, cells thin-walled,
pale brown, 13–37 × 12–22 µm. Surface with hyphoid hairs,
33–70 µm long, of 4–7 ovoid to subglobose cells, constricted
at septa, 6–9 µm wide, sometimes with a light brown matter.
Resinous exudates absent to scarce, pale brown, dissolving
in εδZ. Basal mycelium of 3.5–4.5 µm wide, hyaline to very
pale brown hyphae, sometimes with oily refractive drops on the
surface, sometimes with minute bipyramidal resinous exudates.
Specimens examined. azErbaijan, εontes Talysh, in silva mixta, 14 Oct.
1962, E. Parmasto (UPS F-629452). – dEnmark, East Jylland, Kammerherrens Ege, εoesgård Skov, S of Århus, under Fagus and Quercus, 11 Sept.
2006, J. Vesterholt, JV06-656 (C). – FranCE, Ain, commune de Saint-Benoit,
forêt d´Évieu, under Quercus robur, Carpinus betulus, Corylus avellana and
Alnus glutinosa, 15 July 2011, F. Armada, NV 2011.07.04 (dupl. S). – SwEdEn, Uppland, Stockholm, Drottningsholm, under Quercus robur, 13 Sept.
2009, K. Hansen & I. Olariaga, KH.09.163 (S); Uppland, Stockholm, Solna,
Karlbergsparken, garden with broadleaf trees, 29 July 2006, H.-G. Toresson
(S-F248339); Uppland, Stockholm, Sånga, Svartsjö slott, under Quercus,
Fagus and Corylus, 31 Aug. 2008, H. Kauffman, HK08046 (S); Uppland,
Uppsala, mixed forest with Pinus sylvestris, Picea abies, Betula pendula
and Quercus robur, 9 Oct. 2011, J.C. Zamora (BIO-Fungi 16391); Uppland,
Uppsala, Hågadalen-Nåsten Nature Reserve, Predikstolen, under Corylus,
Populus, Quercus and Picea, on rich ground, 6 Sept. 2010, K. Hansen &
I. Olariaga, KH.10.183 (S).
Notes — Otidea platyspora is characterised by large, caespitose, brown apothecia, blackening in bruised margins and large
spores. εacroscopically it resembles members of the O. alutacea complex, but these differ in the smaller spores. Otidea apophysata and O. daliensis have likewise brown apothecia and
large spores, but can be distinguished by having even larger
spores and smaller (up to 30 mm) apothecia (see further under
those species).
Otidea platyspora is a striking species with scarce records.
It was described from Sweden and has been reported from
France (Boudier 1908, Nannfeldt 1966, Van Vooren & Armada
2011), the Netherlands (εaas Geesteranus 1967) and recently
from Britain and Denmark (Parslow & Spooner 2013). We report it as new from Azerbaijan and give additional records from
Denmark and Sweden. Otidea platyspora appears to have been
overlooked and is more widespread than thought previously.
4. Otidea subterranea Healy & ε.E. Sm. in Smith & Healy,
εycol. Res. 113: 860. 2009
Holotype. USA, Iowa, δedges State Park, shallowly hypogeous, erumpent
on soil, 30 Aug. 1997, R. Healy RH69 (FH).
Notes — Otidea subterranea is the only known hypogeous
species of Otidea. The pustules (c. 50–100 µm high) on the
182
outer surface of the ptychothecia and the incrusted tomentum
are typical for Otidea. The asci are cylindrical, with 8 uniseriate spores, placed in a defined hymenium. Unlike most Otidea
species the spores are uniguttulate. As an adaptation to a hypogeous habit active spore discharge has been lost. Probably for
the same reason, the paraphyses are aggregated and fused
subapically to form an epithecium of brown thick-walled cells,
and are reminiscent of the frequently branching, entangled and
interconnected paraphyses in O. apophysata. There are no
other morphological features that support the exact placement
of O. subterranea in the O. platyspora clade. The receptacle
surface of O. subterranea ascomata is whitish to cream in
young and peach-cream to buff with tan-brown areas in older
specimens. The gleba is of dark brown fertile veins lined with
a thin, light yellowish brown hypothecium, but the sterile veins
are of whitish hyphae (Smith & Healy 2009).
Otidea alutacea clade
Apothecia cup-shaped and split, brown. Spores typically ellipsoid, with almost parallel sides. Outermost ectal excipulum and
basal mycelium smooth or with very sparse resinous exudates.
Species — Otidea alutacea s.l.
5. Otidea alutacea (Pers.) εassee, Brit. Fungus-Fl. 4: 446.
1895. — Fig. 8
Basionym. Peziza alutacea Pers., Observ. εycol. 2: 78. 1799; non Peziza
alutacea Schumach., Enum. Pl. 2: 431. 1803 (homonym).
≡ Scodellina alutacea (Pers.) Gray, Nat. Arr. Brit. Pl. 1: 668. 1821.
≡ Peziza cochleata var. alutacea (Pers.) Fr., Syst. εycol. 2: 50. 1822:
Fr. loc. cit. (‘ß alutacea’).
≡ Plicaria alutacea (Pers.) Fuckel, Jahrb. Nassauischen Vereins Naturk.
23–24: 327. 1870.
≡ Aleuria alutacea (Pers.) Gillet, Champ. France Discom. 1: 42. 1879.
Lectotype. Bull., Hist. Champ. France 1: t. 154, f. b., designated by Carbone (2010a). Epitype. (δ0111551, Herb. Persoon), designated by Carbone
(2010a).
Apothecia gregarious, rarely caespitose, 15 –75 mm high,
8–48 mm wide, initially ear-shaped, soon expanding, becoming shallowly to deeply cup-shaped, split, sessile or stipitate.
Hymenium initially brown (5D6), then yellowish brown (5D4,
5D5) to dark reddish brown (7D7, 7E7), when dried purple
brown (6D6, 6D7). Receptacle surface slightly hygrophanous, in
drying yellowish brown (5D5, 5D6), or sometimes with purplish
brown (7D5) tones, when dried light ochre (5A4, 5B4) brownish
ochre (5B5, 5B6), finely furfuraceous to slightly warty in the
margin. Warts flattened, gregarious, concolorous, darker when
the outside in drying. Stipe 3 – 6 × 3–4 mm. Basal tomentum
and mycelium white to very light ochre (5A2). Spores ellipsoid
to broadly ellipsoid, oblong ellipsoid, slightly inequilateral,
with two large guttules, often with several smaller granules,
smooth, hyaline, (13.5–)14.5–16.5(–17.5) × 6.5–7.5(–8) µm
(δm = 14.6–16.1 µm, Wm = 6.6–7.5 µm, Qm = 2.1–2.2; n = 5).
Paraphyses curved to hooked, only few straight, of the same
width or slightly enlarged at apices, 2.5–4.5 µm wide, without
notches, sometimes embedded in a brown matter at apices,
when fresh containing small, refractive, light brownish yellow
guttules; when dried brownish yellow. Asci 140–187 × 11–13
µm. Apothecial section 750 –1050 µm thick. Subhymenium
c. 80–100 µm thick, visible as a darker zone, cells cylindrical
to swollen, densely arranged, with scattered yellowish brown
resinous exudates at the septa. Medullary excipulum of textura
intricata, 400–650 µm thick, hyphae thin-walled to slightly thickwalled, 4–9 µm wide, hyaline to light brown, sometimes with
yellowish brown resinous exudates at septa. Ectal excipulum of
textura angularis of 80–100 µm, cells thin-walled, pale brown,
1
1
Description based only on specimens of O. alutacea s.str.
Persoonia – Volume 35, 2015
9–21 × 9–17 µm. Surface with broad conical warts, 35–57
µm high, formed by short, fasciculate, hyphoid hairs, of 6–7
globose to subglobose cells, constricted at septa, 7–10 µm
wide. Resinous exudates absent to scarce, yellowish brown,
dissolving in εδZ. Basal mycelium of 3–4.5 µm wide, hyaline
hyphae, with oily refractive drops on the surface, sometimes
with minute resinous exudates.
Specimens examined. O. alutacea s.str. — dEnmark, S Sjælland, εøn,
Store Klinteskov, by Svantestenen, on calcareous soil (pH 7.0) along forest
road, under deciduous trees, together with Humaria hemisphaerica and
Trichophaea woolhopeia, 11 Sept. 1994, K. Hansen & S.K. Sandal, KS94-111 (C). – FranCE, Puy-de-Dôme, Auvergne, Nadayat, sous feuillus en
terrain neutrocline, 20 Sept. 1998, G. Corriol, GC 98092002 (dupl. S). – italy,
Piemonte, Vignole Borbera (Aδ), Fraz. Variano superiore, under Quercus
pubescens and Castanea sativa, 19 Oct. 2010, M. Carbone (S-F257084). –
norway, Nord-Trøndelag, δeksvik, Gjøråsvika, on rich, bare ground, under
Corylus and Picea, on a steep slope, 3 Sept. 2009, K. Hansen & I. Olariaga,
KH.09.133 (S). – Spain, Gipuzkoa, Tolosa, Elosegi markesaren lorategiak,
under broadleaf trees in a garden, 29 εay 2009, J.I. López-Amiano, JδA
2009052902 (ARAN-Fungi A3023204). – SwEdEn, Gotland, Ollajvs Nature
Reserve, close to δjugarn, under Picea and Pinus on calcareous ground,
27 Sept. 2010, K. Hansen, K. Gillen & I. Olariaga, KH.10.278 (S). Clade 1
— dEnmark, Eastern Falster, Korselitze-forests, 5 Oct. 2007, H. Knudsen,
KH.07.46 (S). – SwEdEn, Uppland, Stockholm, Norra Järvafältet, Hansta
Nature Reserve, on rich ground under Corylus and Quercus, 8 Sept. 2010,
K. Hansen, K. Gillen & I. Olariaga, KH.10.193 (S); Uppland, Stockholm,
N Djurgården, Stora skuggan, on soil in grazed open oak forest, 12 Sept.
2008, J. Santos, JS.08.81 (S); Uppland, Uppsala, Hågadalen-Nåsten Nature
Reserve, Predikstolen, under Quercus robur, Picea abies, Corylus and Salix,
on rich bare ground, 17 Sept. 2009, K. Hansen & I. Olariaga, KH.09.170 (S).
Clade 2 — USA, Oregon, δincoln Co., Devil’s Punchbowl State Park, 13
εar. 1997, E.T. Peterson (OSC 56770); Washington, Pierce Co., εt Rainier
National Park, δower Tahoma Creek, under Pseudotsuga, Tsuga, Picea and
Calocedrus, 29 Oct. 1996, E.T. Peterson (OSC 56747); ibid., 30 Oct. 1996
(OSC 56754); ibid., 18 Oct. 1997 (OSC 56798); Washington, Snohomish
Co., Sloan Creek trail, 24 Sept. 1997, E.T. Peterson (OSC 56777). Clade 3a
— norway, Nord-Trøndelag, δeksvik, Gjøråsvika, on slope under Corylus and
Picea, on rich ground, 3 Sept. 2009, V. Kučera & I. Kautmanová, KH.09.135
(S). – SwEdEn, Södermanland, Nynäshamn, Herrhamra, on soil under Fagus,
in narrow forest area along the road, 19 Sept. 2013, K. Hansen & X.H. Wang,
KH.13.50 (S); Uppland, Norrtälje, δänna, under Corylus, 26 Aug. 2008,
J. Santos, JS.08.43 (S); Uppland, Uppsala, Hågadalen-Nåsten Nature
Reserve, Predikstolen, under Quercus robur, Picea abies, Corylus and
Salix, on rich, bare ground, 6 Sept. 2010, K. Hansen, K. Gillen & I. Olariaga,
KH.10.198 (S). Clade 3b — dEnmark, S Sjælland, εøn, Store Klinteskov,
Vestre Ulvemose, on calcareous soil in deciduous forest, 26 Sept. 1994,
K. Hansen & S.K. Sandal, KS-94-192 (C). – SwEdEn, Uppland, Uppsala,
Hågadalen-Nåsten Nature Reserve, Predikstolen, under Corylus, Populus
and Picea, on rich ground, 19 Sept. 2009, K. Hansen & I. Olariaga, KH.09.178
(S). Clade 4 — USA, Oregon, Benton Co., Corvallis, εcDonald-Dunn Research Forest, 10 Oct. 1997, E.T. Peterson (OSC 56782); ibid., 18 Nov. 1996
(OSC 56758); Oregon, Benton Co., Corvallis, west side of NW Beechwood
Place, scattered to clustered on rotting bark mulch and in thin grass under
Pseudotsuga menziesii, 17 Nov. 2010, N.S. Weber, NSW10200 (OSC
150345); Oregon, Corvallis, Witham Hill, 25 Nov. 1997, E.T. Peterson (OSC
56813); Oregon, Douglas Co., 20 Oct. 2010, J. Moore, εoorefun 19 (OSC);
Oregon, Douglas Co., Bear Gulch, under Pseudotsuga menziesii, 13 Jan.
1999, R. Davidson (OSC 67524); Oregon, Douglas Co., Slimewater, under
Pseudotsuga menziesii, Quercus garryana, Abies grandis, etc., 12 Sept.
1999, Frymire (OSC 72978); Oregon, Douglas Co., Umpqua, under Pseudotsuga menziesii, Abies concolor, Calocedrus decurrens, Corylus cornuta,
etc., 23 εar. 2000, E. Stewart (OSC 72979); under Pseudotsuga menziesii,
Pinus ponderosa, Calocedrus decurrens, Pinus lambertiana, Abies concolor,
etc., 15 Dec. 1999, C. Rusch (OSC 72176); Oregon, δane Co., Willamette
National Forest, εiddle Fork Ranger District, under Tsuga heterophylla, Thuja
plicata and Abies grandis, 18 Nov. 2002, Smith (OSC 119567).
Additional material of O. alutacea s.l. dEnmark, Sjælland, Hareskoven, N of
Copenhagen, 2 Aug. 1961, H. Dissing (C-F-48301). – FranCE, Orliénas, sous
Quercus et Cedrus atlantica, 13 Nov. 2008, B. Rivoire & N. Van Vooren, NV
2008.11.01 (dupl. S-F256976); Rhône, Bron, Parc de Parilly, 24 Sept. 2008,
J. Cavet, NV 2008.09.32 (dupl. S-F256974); Rhône, Courzieu, hameau des
Verchères, sous Pseudotsuga menziesii, 26 Oct. 2008, D. Carbonnel, NV
2008.10.02 (dupl. S-F256975). – italy, Puglia, εesagne (BR), Bosco δucci,
in soil, mainly under Quercus ilex, 12 Oct. 2010, M. Carbone (S-F257085).
– norway, Nord-Trøndelag, δeksvik, Gjøråsvika, mixed forest on rich ground,
I. Olariaga et al.: A monograph of Otidea
183
Fig. 8 Otidea alutacea s.str. (KH.09.133). a. Apothecia; b. spores in water †; c. paraphyses in water †; d. ectal excipulum in KOH†. — Scale bars = 10 µm;
= dried material.
†
3 Sept. 2009, K. Hansen & I. Olariaga, KH.09.137 (S); ibid., R. Braathen,
KH.09.139 (S); Nordland, Rana, Rausandaksla, in salico-betuletum, on limestone, 21 Sept. 1974, S. Sivertsen (C-F-60697). – Spain, εadrid, Arboreto
de ETSI εontes, under Quercus suber, Pinus pinea and Nerium oleander,
21 Nov. 2006, L. Rubio Casas (AH42204). – SwEdEn, Gotland, near Visby,
Värnhem, on rich ground under Fagus and Quercus, with Hepatica nobilis,
22 Sept. 2009, E. Bohus-Jensen, K. Hansen & I. Olariaga, KH.09.187 (S);
Gästrikland, Hofors, Sibbersbovägen, on rich ground under Corylus, 1 Sept.
2010, K. Hansen, K. Gillen & I. Olariaga, KH.10.154 (S); Jämtland, Ändsjön
Nature Reserve, in rich Picea forest, 26 Aug. 2009, H. Lindström, KH.09.97
(S); Närke, Ekeby, Kvarntorp, under a big Quercus by the road, 10 Sept.
2008, J. Santos & K. Hansen, JS.08.57 (S); Närke, Tysslinge, δatorpsbruk,
Grytsätterskogen, grassland with Quercus, 13 Sept. 2008, JS.08.76 (S);
Närke, Örebro, Hästhagen, by Svartån, mixed forest, 13 Sept. 2008, H. Kauffman, JS.08.74 (S); Skåne, Helsingborg, Fredriksdals Friluftmuseum, 16 Sept.
2010, G. Hamilton, KH.10.206 (S); Skåne, Kjugekull Nature Reserve, on bare
ground under Quercus rubra, Corylus and Fagus, 24 Sept. 2010, K. Hansen,
K. Gillen & I. Olariaga, KH.10.262 (S); Skåne, εaltesholm, forest close to the
castle, on the ground under Fagus, close to Alnus, 25 Sept. 2010, K. Hansen,
K. Gillen & I. Olariaga, KH.10.265 (S); Torne δappmark, Abisko, 12 Aug.
1974, M.D. Paulsen & N. Tams (C-F-48045); Uppland, Stockholm, Enebyberg, Rinkebyskogen, on bare soil in a ditch, under Picea, Betula and
Populus, 28 Aug. 2008, J. Santos, JS.08.50 (S); ibid., in deciduous forest
under Corylus, but also Tilia, Quercus and Betula, 1 Sept. 2008, J. Santos,
JS.08.56 (S); Uppland, Stockholm, Norra Järvafältet, Hansta Nature Reserve,
on naked soil among leaves, under large Corylus, also Quercus, 8 Sept.
2010, K. Hansen, K. Gillen & I. Olariaga, KH.10.189 (S); Uppland, Uppsala,
Hågadalen-Nåsten Nature Reserve, Predikstolen, on rich ground under
Quercus, Corylus, Populus tremula and Picea, 17 Sept. 2009, K. Hansen &
I. Olariaga, KH.09.173 (S).
Notes — We consider O. alutacea s.l. to comprise a species
complex. It is recognised by the medium brown, cup-shaped,
split apothecia, an ectal excipulum with only sparse resinous
exudates if any, and predominantly oblong spores. Although
sometimes treated as a well-delimited species (Harmaja
2009a), spore sizes of O. alutacea provided by different authors
vary considerably, e.g. 14–16 × 7–9 µm (Dissing 2000) or 12.5–
14.5 × 6.2–7.3 µm (Harmaja 2009a). In fact, O. cochleata has
been separated from O. alutacea on account of larger spores,
16–18 × 7–8 µm (Dissing 2000), or darker apothecia (εornand
& Courtecuisse 2005, δiu & Zhuang 2006, Zhuang 2006).
εeanwhile, two taxa of the O. alutacea species complex have
been separated in North America, based on apothecia colour,
and spore size and shape (Peterson 1998). Our δSU phylogeny
resolved several clades within O. alutacea s.l. (Fig. 1), which
are strongly supported in our multigene phylogeny (Hansen
& Olariaga 2015). It appears that the spore sizes within each
clade have a fairly narrow range, but overlap exists between
the clades. Patterns of continental speciation are suggested as
well; two clades have North American specimens (clade 2, 4),
and the rest contain samples from Europe and Asia.
Carbone (2010a) selected a lectotype and an epitype for O. alutacea. A spore range of 15.5–17 × 7 µm was given for the epitype specimen (Carbone 2010a), and based on this we assign
O. alutacea s.str. to the clade inferred from the phylogenetic
analyses encompassing this spore size. The description above
is based solely on the specimens of that clade, which are
characterised by the initially shallowly cup-shaped apothecia,
with rather light ochraceous brown hymenium, which later
becomes purple brown and more deeply cup-shaped. The
lectotypification of O. alutacea proposed by Parslow & Spooner
(2013) is superfluous.
Among the Eurasian clades, clade 1 contains North European
specimens characterised by small spores (12–13.5 × 5.5–7
µm, Lm = 12.2 –12.9 µm, Wm = 5.8 – 6.6 µm, Qm = 1.9 – 2.2),
non-overlapping with O. alutacea s.str. and clade 3. Clade 1
184
should be compared to O. kunmingensis, a taxon belonging
to the O. alutacea complex characterised by short spores
(Zhuang & Yang 2008). Clade 3a encompasses three specimens with spores sizes 13.5–15 × 6.5–8 µm (δm = 14.5–14.6
µm, Wm = 6.7–7.3 µm, Qm = 2 – 2.2) overlapping with those
of O. alutacea s.str. as described here. The apothecia differ
slightly macroscopically from O. alutacea s.str. in being deeply
cup-shaped in the beginning, reddish brown when young, later
pale ochre-brown. Clade 3b is composed of two collections
with larger spores, 15.5 –17.5 × 7.5 – 8 µm (δm = 16.1–17.4
µm, Wm = 7.7–8 µm, Qm = 2–2.3), but slightly overlapping with
O. alutacea s.str. and clade 3a. Clade 3b may correspond to
O. cochleata sensu Dissing (2000). The two North American
clades (2 and 4) comprise specimens with clearly non-over-
Persoonia – Volume 35, 2015
lapping spore sizes, 15 –18 × 7– 8 µm (δm = 15.5 –16.9 µm,
Wm = 7.3 –7.9 µm, Qm = 2 – 2.2) and 12 –14.5 × 6.5 – 8.5 µm
(δm = 12.5–14 µm, Wm = 6.6–7.5 µm, Qm = 1.8–2), respectively.
These two clades correspond to the two species distinguished
by Peterson (1998) in Western North America, as O. alutacea
(clade 2) and O. umbrina (clade 4). These species were said to
differ in colour of fresh apothecia. Clades 2 and 4 have spores
that overlap with European clades, and it is so far problematic
to distinguish them using only morphology and disregarding
the geographical origin.
To be able to fully clarify species boundaries within the O. alutacea complex, sampling of additional collections for molecular
study is needed. Distinguishing morphological and ecological
characters should be sought, especially through studying fresh
Fig. 9 Otidea papillata†. a. Apothecia; b. spores in water; c. paraphyses; d. resinous exudates in the medullary excipulum in water; e. ectal excipulum in KOH;
f. basal mycelium in KOH (a–d, f: H6003547, holotype; e: TUR 102134). — Scale bars = 10 µm; † = all dried material. — Photos: a. J. Kearey.
185
I. Olariaga et al.: A monograph of Otidea
material. Several names that belong to this complex, such as
O. alba, O. cinerascens, O. cochleata, O. felina and O. kunmingensis should be considered as this study is undertaken.
Otidea papillata clade
Apothecia cup-shaped, split. Receptacle surface with contrasting warts. Spores small, 9.5–11 µm. εedullary excipulum with
brown resinous exudates embedding some hyphae. Ectal excipulum poorly differentiated, of textura prismatica to textura
intricata. Resinous exudates on the ectal excipulum not dissolving in εδZ.
Species — Otidea papillata.
6. Otidea papillata Harmaja, Karstenia 15: 31. 1976 — Fig. 9
Holotype. Finland, Kainuu, Paltamo, εelalahti, εyllymäki, predominantly
coniferous grass-herb forest on distinctly calcareous soil, in litter mainly
composed of spruce needles, 23 Sept. 1971, H. Harmaja (H6003547) !
Apothecia 23–30 mm high, 7–33 mm wide, initially broadly earshaped, with upper margin rounded, then becoming cup-shaped,
split, stipitate or sessile. Hymenium ochre (5B5, 5B6) to yellowish ochre (4A5) when dried. Receptacle surface yellowish
brown (5C6, 5C7) when dried, warty. Warts conical, angular or
rounded, gregarious, distinctly darker than the background, dark
ochre brown to brown. Stipe 7–10 × 3–4 mm. Basal tomentum
and mycelium abundant, pale brownish ochre (5A3) to orangeochre (6A3). Spores broadly ellipsoid, seldom very slightly
inequilateral, with two large guttules, smooth, hyaline, 9.5 –
11(–11.5) × 5.5–6.5 µm (δm = 10–10.7 µm, Wm = 6.1–6.3 µm,
Qm = 1.6–1.7; n = 2). Paraphyses curved to tightly hooked, usually enlarged at apices, 3–4 µm wide, sometimes with 1–2
shallow notches, sometimes truncate or forked at apices,
when dried containing small, refractive, hyaline granules. Asci
116–165 × 9–10.5 µm. Apothecial section 900–1200 µm thick.
Subhymenium 70 – 90 µm thick, of dense textura intricata,
visible as a pale brown zone. Medullary excipulum of textura
intricata, 600–800 µm thick, hyphae 5.5–10.5 µm wide, thinwalled to slightly thick-walled, hyaline to very pale yellow, with
brown resinous exudates scattered among and covering some
hyphae, sometimes rod-shaped, paler and partially dissolving in KOH. Ectal excipulum of textura prismatica to textura
intricata, 70–100 µm, cells thin-walled, hyaline, 13–35 × 9–18
µm. Surface with conical to rounded warts, 65–100 µm high,
formed by short, fasciculate, hyphoid hairs, sometimes with
a gelatinous sheath. Resinous exudates abundant, yellowish
brown to reddish brown, paler in KOH, not dissolving in εδZ.
Basal mycelium of 2.5 – 4.5(– 6) µm wide, septate, hyaline
to very pale yellow hyphae, unchanged in KOH, smooth or
with regularly arranged, spheroid, yellow, resinous exudates,
sometimes embedded in a yellowish matter, dissolving in εδZ,
partially and more slowly in KOH.
Specimen examined. Finland, Varsinais-Suomi, Parainen, Petteby, Stornäset (Paltbacken), in coniferous forest among mosses, 30 Sept. 1990, T. Lindholm (TUR 102134).
Notes — Otidea papillata is only known from two Finnish
collections and its apothecial colours in fresh state are still
unknown. It is a distinct species, characterised by cup-shaped
apothecia with conspicuous warts on the outside, and small
spores (Harmaja 1976). Two diagnostic characters have been
observed in the two collections examined: 1) a very poorly
differentiated ectal excipulum of textura prismatica to textura
intricata, which is unique within the Otidea species studied by
us; and 2) scattered brown resinous exudates on the hyphae
of the medullary excipulum, that are somewhat reminiscent of
those in O. bufonia.
Harmaja (1976) emphasised the high warts on the outside of
the apothecia as a unique character for O. papillata, but two additional Otidea species, O. tuomikoskii and O. nannfeldtii, have
as high or higher warts. These two species have in fact been
confused with O. papillata (δundell et al. 1985, Van Vooren et
al. 2008). Otidea tuomikoskii is distinguished from O. papillata
by narrowly ear-shaped apothecia, a yellow reaction of the
excipulum in KOH, an ectal excipulum of textura angularis, and
by lacking pigmented exudates on the hyphae of the medullary
excipulum. Highly warted apothecia of O. nannfeldtii probably
resemble O. papillata more. However, O. nannfeldtii possesses
pigmented resinous exudates at the septa in the medullary excipulum, resinous exudates of the ectal excipulum that convert
into amber drops, and most importantly, an ectal excipulum of
textura angularis.
Otidea leporina clade
Apothecia ear-shaped, yellowish ochre to brown. Resinous exudates on the ectal excipulum converting into reddish grey drops
in KOH. Associated with conifers.
Species — Otidea leporina, O. pseudoleporina.
7. Otidea leporina (Batsch) Fuckel, Jahrb. Nassauischen Vereins Naturk. 23 –24: 330. 1870 ‘1869 –1870’ — Fig. 4a, 10
Basionym. Peziza leporina Batsch, Elench. Fung. 1: 117. 1783: Fr., Syst.
εycol. 2: 47. 1822.
≡ Scodellina leporina (Batsch) Gray, Nat. Arr. Brit. Pl. 1: 668. 1821.
≡ Helvella leporina (Batsch) Franchi, δ. δami & ε. εarchetti, Rivista
εicol. 1: 63. 1999.
≡ Helvella auricula Schaeff., Fung. Bavar. Palat. Nasc. 4: 103. 1774
(‘Elvela’).
≡ Wynnella auricula (Schaeff.) Boud., Icon. εycol. list prél. 600 sp.: (2).
1904.
≡ Otidea auricula (Schaeff.) Sacc., Syll. Fung. 8: 95. 1889.
Lectotype designated here: Schaeffer, Fung. Bavar. Palat. Nasc. 2: t. 156.
1763 (‘Elvela decima tertia’). Epitype designated here: SwEdEn, Jämtland,
Östersund, Andersön Nature Reserve, under Picea abies on rich ground, 28
Aug. 2009, K. Hansen & I. Olariaga, KH.09.93 (S); εycoBank εBT178082.
= Otidea leporina f. minor Rehm, Ber. Naturhist. Vereins Augsburg 26:
63: 1881.
≡ Otidea leporina var. minor (Rehm) Sacc., Syll. Fung. 8: 94. 1889.
Lectotype designated here: gErmany, δeipzig, in der Harth, in spruce
forest, Aug. 1873, G. Winter, Rehm Ascomyceten no. 251 (S-F88382) !
Isolectotype (UPS F-641412) !; εycoBank εBT178088.
= Otidea leporina f. major Rehm, Hedwigia 3–4: 2. 1883.
= Otidea leporina var. rubescens Velen., εonograph. Discom. Bohemiae
1: 354. 1934.
Lectotype designated here: CzECh rEpubliC, Kosoř near Prague, Sept.
1920, F. Fechtner (PRε 614790) !; εycoBank εBT200087.
= Otidea myosotis Harmaja, Karstenia 15: 32. 1976.
Holotype. Finland, Etelä-Karjala, Hamina, Vehkalahti, Pyhältö, mixed
forest, 3 Oct. 1970, L. Fagerström (H6003548) !
= Otidea crassa W.Y. Zhuang, εycotaxon 94: 366. 2006 (‘2005’).
= Otidea fuckelii ε. Carbone & Van Vooren, Rivista εicol. 52: 322. 2010
(‘2009’).
Holotype. auStria, Nassau, in pinetis umbrosis, Fungi Rhen. Exs. no.
1233 (G00110768). Isotype (S-F114092) !
Misapplied names
– Non Wynnella auricula sensu Boudier, Icon. εycol. livr. 26: n°. 535, pl. 250.
1909 (preliminary text with ‘circulaires’) (= Wynnella silvicola (Beck) Nannf.).
Apothecia gregarious or caespitose, 17–52 mm high, 4–25
mm wide, narrowly to broadly ear-shaped, split, stipitate or
sessile. Hymenium yellowish brown (5C6), cinnamon brown
(5D6), pale ochre brown (5A5, 5B6) to orange brown (6C7,
6D6), sometimes dark brown (5D8, 5F8, 6E8) when young,
seldom with pale pink stains (6A2), bruised margin orange
brown (6D8), when dried cinnamon brown (5A5, 5B5) to rusty
brown (6D7, 6D8). Receptacle surface ochre brown (5B6),
hygrophanous, in drying paler ochre brown (4A4, 4A5, 4B6),
186
Persoonia – Volume 35, 2015
Fig. 10 Otidea leporina*. a, b. Apothecia; c. spores; d. paraphyses; e. ectal excipulum with resinous exudates, insert shows close-up of amber drops on the
ectal excipulum in εelzer’s reagent; f. basal mycelium pale brown, with very small, regularly arranged, resinous exudates (a: KH.09.93, epitype; b: KH.09.102;
c–g: KH.11.02). — Scale bars = 10 µm; * = all fresh material.
when dried yellowish brown (5C7, 5C8), furfuraceous to finely
warty, seldom wrinkled at the base. Warts conical to flattened,
gregarious, concolorous, sometimes darker than the background, golden brown. Stipe 4–15 × 2–8 mm. Taste slightly
bitter. Basal tomentum and mycelium abundant, white to cream
white (5A3), sometimes very pale brown when dried. Spores
broadly ellipsoid, sometimes inequilateral, with two large
guttules, smooth, hyaline, (12 –)12.5 –14(–15) × 7– 8.5 µm
(δm = 12.8–13.8 µm, Wm = 7.5–8.2 µm, Qm = 1.6–1.8; n = 16).
Paraphyses curved to hooked, of the same width or slightly
enlarged at apices, 2.5–4 µm wide, without notches or with 1–3
low notches, seldom forked at apices, when fresh containing
small, refractive, light yellow guttules; when dried small, refrac-
tive, hyaline granules. Asci 170–215 × 9–10.5 µm. Apothecial
section 700–900 µm thick. Subhymenium c. 80–100 µm thick,
of dense textura intricata, visible as an orange-brown darker
zone, cells cylindrical to swollen. Medullary excipulum of textura
intricata, 300 – 550 µm thick, sometimes differentiated into
two parts: a) textura angularis underneath the subhymenium,
40–50 µm thick, cells 6–12 µm broad; b) textura intricata, hyphae 5–13 µm wide, sometimes slightly swollen, thin-walled
to slightly thick-walled, hyaline to very pale brown, sometimes
with yellow-brown resinous exudates at septa. Ectal excipulum
of textura angularis, 85–110 µm thick, cells thin-walled, pale
brown, 15–48 × 12–31 µm. Surface with broadly conical warts,
50–85 µm high, formed by short, fasciculate, hyphoid hairs,
I. Olariaga et al.: A monograph of Otidea
of 2 – 4 subglobose to elongated cells, constricted at septa,
10–16 µm wide, sometimes with a gelatinous sheath. Resinous
exudates abundant, yellow brown, sometimes dissolving in part
into amber drops or converting into reddish particles in εδZ,
dissolving into yellowish reddish grey heterogeneous drops in
KOH. Basal mycelium of 3–4.5(–6) µm wide, hyaline to pale
brown hyphae, unchanged in KOH, smooth or with very small,
regularly arranged, spheroid, pale brown, resinous exudates,
dissolving in εδZ, and partially in KOH.
Specimens examined. Canada, Québec, δe Verendrye Park, on ground
under spruce, 16 Sept. 1965, M.E. Elliot 65-123 (UPS F-629640). – CzECh
rEpubliC, Central Bohemian region, Zdice, Aug. 1924, F. Fechtner (PRε
614787, as O. felina); in piceto ad aciculos, Sept. 1925 (PRε 148836);
Prague-West district, Kosoř near Prague, Sept. 1920, J. Velenovský (PRε
614792, as O. umbrina). – dEnmark, Bornholm, Rø Plantage, coniferous forest, 29 Sept. 1985, W. Rummel (C-F-47633); N Jylland, δangdal Plantage
(near Tranum), under Juniperus, near Picea, 13 Aug. 2009, T. Læssøe,
Tδ-13769 (C); N Jylland, Rønhøj Plantage, 7 Oct. 1962, A. Hauerbach (C-F86691); Jylland, Virklund, Silkeborg Sønderskov, in moss, coniferous forest,
26 Sept. 1964, H. Dissing (C-F-48298); Sjælland, Bromme Plantage, N of
Sorø, under Picea, 9 Oct. 1965, H. Dissing (C-F-48299). – Finland, EteläHäme, εustiala, in pineto, 29 Aug. 1866, P.A. Karsten (UPS F-146429);
Perä-Pohjanmaa, Rovaniemi, Kaittiainen, acidic Picea forest, 11 Sept. 2011,
T. Kekki, TK407 (TUR); Perä-Pohjanmaa, Rovaniemi, Pisajärvi, old Picea
forest, 2 Sept. 2011, T. Kekki, TK304 (TUR); Perä-Pohjanmaa, Rovaniemi,
Välljoki, calciferous Picea forest, 25 Aug. 2011, T. Kekki, TK231 (TUR). –
FranCE, δoire, δa Chamba, au sol dans la litière d’aiguilles d’épicéa, 27 Sept.
2008, N. Van Vooren, NV 2008.09.28 (dupl. S). – gErmany, Thüringen, in
silvis abiegnis, Kl. & Op. (UPS F-629404, Klotzsch, Herb. Viv. εycol. 143).
– norway, Nord-Trøndelag, Kvam, Noem Northeast, under Picea, among
Rhytidiadelphus squarrosus, 2 Sept. 2009, H. Lindström, KH.09.131 (S);
Nord-Trøndelag, Namdalseid, Flåbekkåsen Nature Reserve, Picea and Pinus
old-growth forest, on acidic soil, among mosses, 4 Sept. 2009, K. Hansen &
I. Olariaga, KH.09.141 (S); Nord-Trøndelag, Steinkjer, Skrattåsen, in rich
Picea abies forest, 5 Sept. 2009, K. Hansen & I. Olariaga, KH.09.145 (S);
ibid., KH.09.147 (S). – SwEdEn, Härjedalen, Torkilstöten, δjungdalen, on
an active anthill in Picea forest, 19 Aug. 2011, J.C. Zamora & I. Olariaga,
KH.11.02 (S); Jämtland, in the surroundings of Sällsjö, in young stand of
Picea abies, with Betula and Salix, 29 Aug. 2009, K. Hansen & I. Olariaga,
KH.09.100 (S); ibid., KH.09.102 (S); Jämtland, SW of εörsil, Sandtjärndalen
Nature Reserve, under Picea abies on rich ground, 7 Sept. 2009, K. Hansen &
I. Olariaga, KH.09.156 (S); δappland, Ekopark Vuollerim, stream from Brännmyran, Picea forest, 28 Aug. 2008, M. Karström, εK0828 (S); δappland,
Jokkmokk, ‘Nornaskogen’ by Ållojaur, mossy Picea forest, on rich ground,
29 Aug. 2011, K. Hansen & I. Olariaga, KH.11.12 (S); ibid., KH.11.14 (S);
δappland, Jokkmokk, Ultevis Fjällurskog Nature Reserve, Sitoätno, near a
Picea, 31 Aug. 2011, K. Hansen & I. Olariaga, KH.11.33 (S); ibid., KH.11.36
(S); δappland, Kuouka, 15 km SE εessaure, herb-rich Picea forest on rich
ground, among mosses, 3 Sept. 2011, K. Hansen & I. Olariaga, KH.11.67
(S); δappland, S of Kvikkjobb-Kabla FUR Nature Reserve, by Kassavare
εt, Köpenhamn, under Picea, mossy place on acidic ground, 1 Sept. 2011,
K. Hansen & I. Olariaga, KH.11.76 (S); Närke, Knista, δekhyttan, Kungshall,
under Picea on calcareous ground, among mosses and litter, 12 Sept. 2008,
J. Santos, JS.08.065 (S); ibid., coniferous forest, with old Picea and Pinus,
K. Hansen, KH.08.108 (S); Skåne, δoshult, δilla δoshult, Picea forest, 5 Sept.
1998, S.-Å. Hanson, SÅH 105838 (C); Uppland, Stockholm, Enebyberg,
Rinkebyskogen, under Picea and Betula on acidic ground, 2 Sept. 2009,
K. Hansen & I. Olariaga, KH.09.169 (S); Uppland, Täby, Rönninge, close to
parking place by Arninge, under Picea on thick litter layer, 21 Sept. 2008,
J. Santos, JS.08.92 (S); Uppland, Uppsala, Ersta Nature Reserve, on soil
under Picea in young plantation, 23 Sept. 2008, J. Santos & K. Hansen,
JS.08.99 (S); Uppland, Uppsala, Sävja, Norra δunsen Nature Reserve, under
Picea, 28 Aug. 2008, J. Santos, JS.08.46 (S); Värmland, Gustav Adolf,
Hagfors, εalmbackarna, on moss, under Picea and Betula, 10 Aug. 2009,
F. Turander s.n. (S). – USA, California, Del Norte Co., δake Earl Wildlife
Area, 14 Dec. 1997, E.T. Peterson (OSC 56824); ibid., 15 Dec. 1997 (OSC
56825); ibid., under Picea sitchensis, Pinus contorta, Abies concolor, 26
Nov. 2001, M. Castellano & E. Cazares (OSC 108820); California, Humboldt
Co., Big δagoon Park, 14 Dec. 1956, A.H. Smith 56668 (UPS F-629302);
ibid., under Picea, 16 Dec. 1956, A.H. Smith 56799 (UPS F-629304); ibid.,
23 Dec. 1956, A.H. Smith 56954 (UPS F-629305); Colorado, Tolland, on
ground in coniferous woods, 28 Aug. 1920, F.B. Cotner (UPS F-629390);
Oregon, δincoln Co., Fogarty Creek State Park, 15 Oct. 1997, E.T. Peterson
187
(OSC 56784); Washington, Okanogan National Forest, Pasayten Wilderness,
under Picea engelmannii, Pseudotsuga menziesii, Abies lasiocarpa, Pinus
contorta, 16 Sept. 1999, R. Davis (OSC 108856).
Notes — Otidea leporina is probably the most common Otidea species in boreal coniferous forests of Europe. In the Alps
it often occurs together with Cudonia circinans, an association
not seen in Fennoscandia. It is characterised by ear-shaped,
brown apothecia, together with relatively broad spores that
are almost unique within Otidea. Otidea brunneoparva shares
similar spores, but differs from O. leporina in the darker brown
apothecia and strongly notched paraphyses. Other species of
Otidea that macroscopically resemble O. leporina are distinguished by different spore size and shape.
Our morphological and molecular study of the holotype of O. myosotis shows it is a synonym of O. leporina (Hansen & Olariaga
2015). The original description of O. myosotis (Harmaja 1976)
pointed out the apothecial shape and colours, and paraphyses
as diagnostic characters, all of which agree with our concept
of O. leporina. Recently, Harmaja (2009a) stated that the excipular resinous exudates in O. leporina convert in εδZ into
reddish particles and show no reaction in O. myosotis. In the
material of O. leporina examined by us, the resinous exudates
dissolve in part, can appear unchanged or can convert into
reddish particles. Sometimes, small amber-drops have also
been observed, though not as strikingly as in other species.
Therefore, it seems that the reaction of the exudates in εδZ
is variable within O. leporina, and cannot be used to separate
O. myosotis from O. leporina. Otidea crassa is a synonym
based primarily on the GenBank δSU sequence of the type
collection (DQ443444).
Nomenclatural notes — When Batsch (1783) described Peziza leporina, he referred to Schaeffer’s plate (1763), being unaware or disregarding Schaeffer’s later description of ‘Elvela’
auricula Schaeff. (1774) based on the same plate. Since Fries
(1822) sanctioned Batsch’s name, referring to H. auricula Schaeff.
(as P. auricula Schaeff.) as a synonym, P. leporina has priority.
While most authors have interpreted the plate by Schaeffer
(1763, t. 156) as a species that belongs to Otidea (e.g. Fuckel
1870, Rehm 1883, Bresadola 1898, Seaver 1904), others have
considered it to represent the monotypic Wynnella (Gonnermann & Rabenhorst 1869, Quélet 1886). Recently, Franchi et
al. (1999) stated that the Schaeffer plate represents W. silvicola
and as they consider Wynnella to be part of the genus Helvella,
they made the combination Helvella leporina. Carbone & Van
Vooren (2010) expressed doubts about how to interpret the protologue by Batsch (1783) and Schaeffer’s plate, and concluded
the name is ambiguous and recommended it not be used. Instead they introduced the new name O. fuckelii for the Otidea
species treated here. Based on our phylogenetic and morphological studies (see also Hansen & Olariaga 2015) this new
name is, however, superfluous, since O. myosotis and O. crassa
are shown to be synonyms of O. leporina.
The original Schaeffer plate shows several, more or less evenly
coloured, light ochraceus brown apothecia, conforming to O. leporina, and not bi-coloured apothecia (dark reddish brown with
a white base) as in W. silvicola. In our opinion, it leaves little
doubt it shows a species of Otidea. To settle the use of the name
O. leporina, and at the same time preserve the use of the wellestablished name, W. silvicola (Beck) Nannf. (Nannfeldt 1966),
we propose a modern epitype for Peziza leporina Batsch: Fr.
(Fig. 10a), which represents the Otidea species for which the
name has most often been used. The epitype is from Sweden
where Fries saw and studied living material, as indicated by
the abbreviation ‘v. v.’ (vidi vivam, seen living).
188
8. Otidea pseudoleporina Olariaga & K. Hansen, sp. nov. —
εycoBank εB808972; ITS barcode GenBank: Kε010112;
Fig. 5e, 11
Etymology. From ancient Greek ψ υ ο-, which means ‘false, fake’, referring to a close relationship with O. leporina.
Holotype. USA, Oregon, Douglas Co., E of εill Creek, under Pseudotsuga
menziesii, Abies concolor, Pinus lambertiana, 19 Oct. 2010, R. Helliwell,
rh101910 (OSC).
Misapplied names
– Otidea cantharella var. minor sensu Kanouse, εycologia 41: 667. 1949.
Apothecia gregarious, 10–30(–50) mm high, 8–22(–31) mm
wide, initially narrowly to broadly ear-shaped, margin rounded,
then expanding and sometimes becoming irregularly cup-shaped,
Persoonia – Volume 35, 2015
split, stipitate or sessile. Hymenium ochre-orange (4A6, 5A7) to
pinkish orange (5A6, 5A7), sometimes with pink spots or stains
(6A4), when dried orange-ochre (5A5, 5B5) to reddish brown
(6D7). Receptacle surface ochre-brown (5B5), hygrophanous,
in drying yellowish ochre (4A4, 4A5), when dried yellowish ochre
(5A5) to brownish ochre (5B5), furfuraceous to finely warty,
sometimes wrinkled at the base. Warts conical to rounded,
gregarious, concolorous, sometimes distinctly darker than
the background, reddish brown. Stipe 4–11 × 3–5 mm. Basal
tomentum and mycelium abundant, white to pale yellow (4A2)
or ochre (5A2). Spores ellipsoid, sometimes slightly inequilateral, with two large guttules, sometimes with up to 4 smaller
guttules, smooth, hyaline, (9.5–)10–12(–12.5) × 5.5–6.5 µm
(δm = 10.2–11.6 µm, Wm = 5.7–6.4 µm, Qm = 1.7–1.9; n = 6).
Fig. 11 Otidea pseudoleporina. a, b. Apothecia; c. apothecia †; d. spores in water †; e. paraphyses in water †; f. ectal excipulum in water † (a, d–f: rh101910,
holotype; b, c: εoorefun 14). — Scale bars = 10 µm; † = dried material. — Photos: a. R. Helliwell; b. c. J. εoore.
189
I. Olariaga et al.: A monograph of Otidea
Paraphyses curved to hooked, of the same width or slightly
enlarged at apices, 2.5 – 4.5 µm wide, sometimes with 1– 3
notches, apices seldom forked and rarely covered with a hyaline
coating, when dried containing small, refractive, yellow granules. Asci 155–231 × 9–10 µm. Apothecial section 900–1200
µm thick. Subhymenium c. 80–100 µm thick, of dense textura
intricata, visible as an orange-brown darker zone, of cylindrical
to swollen cells. Medullary excipulum 500–750 µm thick, differentiated into two parts: a) textura angularis underneath the
subhymenium, 150–200 µm thick, hyphae 8–18 µm broad;
b) textura intricata, hyphae 5–10(–18) µm wide, sometimes
slightly swollen, thin-walled to slightly thick-walled, very pale
yellow, sometimes with yellow-brown resinous exudates at
septa. Ectal excipulum of textura angularis, sometimes of a
textura prismatica, 80–110(–150) µm thick, cells thin-walled
to slightly thick-walled, pale yellow-brown, 13–37 × 8–27 µm.
Surface with broadly conical warts, 77–115 µm high, formed by
short, fasciculate, hyphoid hairs, of 2–3 subglobose to elongated cells, constricted at septa, 6–12 µm wide, sometimes with
a gelatinous sheath. Resinous exudates abundant, yellowbrown, dissolving into amber drops in εδZ, partially to entirely
dissolving into reddish grey heterogeneous drops in KOH.
Basal mycelium of 3 – 5 µm wide, very pale yellow hyphae,
unchanged in KOH, with very small, yellow resinous exudates,
regularly arranged, spheroid, dissolving in εδZ, partially and
more slowly dissolving in KOH.
Specimens examined. USA, California, Trinidad, under spruce, 30 Nov.
1956, A.H. Smith 56168 (UPS F-629690); Idaho, Idaho Co., Rickliff Creek
Public Camp, on the ground in Thuja-Tsuga woods, 10 Oct. 1947, W.B. Cooke
21227 (UPS F-629388); Idaho, Papoose Creek, Seven Devils εts, on the
ground in Douglas fir association, 3 Sept. 1954, A.H. Smith & H.E. Bigelow,
47346 (UPS F-629331); Oregon, Bear Springs, εt Hood National Forest, 18
Oct. 1947, A.H. Smith 27946 (UPS F-629430); Oregon, Benton Co., Corvallis,
west side of NW Beechwood Place, scattered to clustered on rotting bark
mulch and in thin grass under Pseudotsuga menziesii, 14 Nov. 2010, N.S.
Weber, NSW 10202 (OSC 150347); ibid., 17 Nov. 2010, N.S. Weber, NSW
10200 (OSC 150345); ibid., 27 Nov. 2010, NSW 10201 (OSC 150346);
Oregon, Douglas Co., E of εill Creek, under Pseudotsuga menziesii, Abies
concolor, Pinus lambertiana, 22 Oct. 2010, J. Moore, εoorefun 24 (OSC);
Oregon, Douglas Co., εill Creek, under conifers, 19 Oct. 2010, J. Moore,
εoorefun 14 (S); Oregon, Douglas Co., Roseburg District Bureau of δand
εanagement, under Pseudotsuga menziesii, Arbutus menziesii, Castanopsis
chrysophylla, 11 εay 1997, J. Klein (OSC 66261); Oregon, Douglas Co.,
south of δemolo δake, under conifers, 5 Nov. 2010, R. Heliwell, rh 179 (S);
Oregon, Douglas Co., Thorn Unit I, under Pseudotsuga menziesii, Tsuga
heterophylla, 21 Oct. 2010, C. Durbecq, Durbecq 16 (OSC); Oregon, Jackson
Co., εedford Bureau of δand εanagement, Ashland Resource Area, Beaver
Creek, under Pseudotsuga menziesii, Arbutus menziesii, Toxicodendron
diversilobum, Berberis piperiana, 18 Dec. 2000, R. Brock (OSC 119311);
Oregon, Jackson Co., εedford District Bureau of δand εanagement, Butte
Falls Resource Area, under Pseudotsuga menziesii, Calocedrus decurrens,
Pinus lambertiana, Pinus ponderosa, Quercus kellogii, Rhus diversiloba,
Berberis piperiana, Fragaria vesca, Moehringia macrophylla, grasses, 8 εar.
2000, M. Wineteer (OSC 72956); Oregon, δane Co., Willamette National
Forest, Blue River Ranger District, under Pseudotsuga menziesii, 18 Nov.
1999 (OSC 72296); Oregon, εarion Co., Breitenbuch Hot Springs Community, near Detroit Reservoir, in woods, 8 Nov. 1997, J.W. Spatafora (OSC
56809); Oregon, εt Hood, among moss under conifers, 15 Oct. 1922, L.E.
Wehmeyer (UPS F-629375); Oregon, Warm Springs R., εt Hood National
Forest, Skyline Trail, 29 Sept. 1947, A.H. Smith & W.B. Gruber, 27064 (UPS
F-629689); Washington, Clallam Co., Olympic National Park, Whiskey Bend
trailhead, 26 Nov. 1996, E.T. Peterson (OSC 56760); Washington, Fish Creek
Region, εt Rainier National Park, 25 Aug. 1948, E.G. Simmons 2067 (UPS
F-629332); Washington, lower slopes of Rampart Ridge, W. of δongmire, εt
Rainier National Park, 4 Sept. 1948, E.G. Simmons 2172 (UPS F-629386);
Washington, Park Creek, εt Baker National Forest, 9 Sept. 1941, A.H. Smith
16755 (UPS F-629820); Washington, Pierce Co., εt Rainier National Park,
δower Tahoma Creek, 29 Oct. 1996, E.T. Peterson (OSC 56749).
Notes — Otidea pseudoleporina is recognised by the broadly
ear-shaped apothecia, ochre-orange to pinkish orange hymenium and small spores. Our multi-gene phylogenetic analyses
(Hansen & Olariaga 2015) suggest O. pseudoleporina is the
sister species of O. leporina. They share ear-shaped apothecia and resinous exudates on the outer excipulum that partly
convert into heterogeneous reddish drops in KOH. Otidea
leporina differs in the brown apothecia and larger, broadly ellipsoid spores. Otidea pseudoleporina resembles O. nannfeldtii
and O. formicarum in the general apothecial shape and small
spores. Otidea nannfeldtii is distinguished by most often lacking
orange tones, having narrowly ear-shaped young apothecia,
and ectal excipular resinous exudates turning reddish brown in
KOH. Otidea formicarum is distinguished by having apothecia
devoid of orange tones, and spores with a lower Qm (1.6–1.7)
than O. pseudoleporina (1.7–1.9).
The material cited by Kanouse (1949) under O. cantharella var.
minor most likely represents O. pseudoleporina. Otidea cantharella var. minor as described by Boudier (1909a) has a pale
ochre or grey hymenium, citrine yellow outside, and veins at the
apothecial base, and represents a different species (see under
O. minor). Peterson (1998) treated under O. concinna material
that we refer to O. pseudoleporina and stated that Harmaja
(1974) used the name O. cantharella for the same species.
Otidea concinna is a well-known species in Europe, clearly
distinct from O. pseudoleporina (see O. concinna). As for the
name O. cantharella, the protologue describes a fungus with
the colour of Cantharellus cibarius (Fries 1822), and we typify
it with material of the large-spored species sometimes called
O. caligata (see O. cantharella). So far O. pseudoleporina is
only known from Western North America.
Otidea tuomikoskii clade
Apothecia ear-shaped. Receptacle surface with warts often
more than 100 µm high. Basal tomentum light ochre to orange
ochre. Spores small, 10–11 µm long. Sections of apothecia
turning yellow in KOH, especially the subhymenium and ectal
excipulum. Associated with conifers.
Species — Otidea tuomikoskii.
9. Otidea tuomikoskii Harmaja, Karstenia 15: 30. 1976 — Fig.
12
Holotype. Finland, Etelä-Häme, δammi, Pappilankylä, Koiransuolenoja, in
needles of Picea abies on an anthill, 9 Sept. 1972, R. Tuomikoski (H6002901) !
= Otidea papillata f. pallidefurfuracea Van Vooren & Hairaud, Bull. εycol.
Bot. Dauphiné-Savoie 188: 56. 2008.
Holotype. FranCE, Jura, δes Rousses, tourbière près du lac des Rousses,
au sol dans la litière d’aiguilles, sous épicéas (Picea abies), 19 Sept. 2007,
N. Van Vooren, NV 2007.09.27 (PC). Isotype (S) !
Misapplied names
– Otidea papillata sensu δundell, Nannfeldt & Holm, Fungi Exs. Suec. 66:
3282. 1985.
Apothecia gregarious to caespitose, 17–60 mm high, 7–30 mm
wide, long and narrowly ear-shaped, split, stipitate or sessile.
Hymenium pale whitish ochre (4A2, 4A3) to ochre yellow (4A4–
4A6) rarely with pink stains, when dried light ochre yellow (4A4)
to ochre (5B7, 5B8). Receptacle surface brownish ochre (4B7,
4C7) to yellow brown (5B6–5D6), hygrophanous, ochre yellow
(4A5, 4A6, 4B6) in drying, when dried yellowish brown (5C8,
5D8), warty, rarely wrinkled at the base. Warts conical, gregarious, brown, distinctly darker than the background or rarely lighter. Stipe 3–6 × 2–3 mm. Smell faintly aromatic. Basal tomentum
and mycelium abundant, light ochre (5A2) to orange-ochre
(5A4). Spores ellipsoid, slightly inequilateral, with two large
guttules, and sometimes with 1–4 smaller granules, smooth,
hyaline, (9.5–)10–11(–12) × 5.5–6.5(–7) µm (δm = 10.3–11.4
µm, Wm = (5–)5.5–6.5(–7.5) µm, Qm = 1.7–1.9; n = 15). Paraphyses curved to hooked, often broader at apices, 2.5–5(–6)
µm wide, sometimes with up to two shallow notches or forked at
190
Persoonia – Volume 35, 2015
Fig. 12 Otidea tuomikoskii*. a, b. Apothecia; c. spores; d. paraphyses; e. ectal excipulum; f. basal mycelium (a: JS.08.68; b–f: KH.11.77). — Scale bars = 10 µm;
* = all fresh material. — Photos: a. J. Santos.
apices, when fresh containing small to large, refractive, hyaline
to pale yellow guttules; when dried pale yellow. Asci 113–199
× 9–11.5 µm. Apothecial section 600–700(–1200) µm thick,
pale to bright yellow in KOH. Subhymenium c. 50–90 µm thick,
of dense textura intricata, visible as a yellowish brown zone.
Medullary excipulum of textura intricata, 300–500(–850) µm
thick, hyphae 3–13 µm wide, thin-walled to slightly thick-walled,
hyaline to very pale yellow, without resinous exudates at septa.
Ectal excipulum of textura angularis, 80–120 µm, cells thinwalled, hyaline to light yellow, 13–40 × 8–25 µm. Surface with
conical warts, 55–177 µm high, formed by fasciculate, short,
hyphoid hairs, of 3–8 globose to elongated cells, constricted at
septa, 7–15 µm wide, sometimes with a gelatinous sheath. Resinous exudates abundant, yellow-orange to yellowish brown,
dissolving into amber drops in εδZ, unchanged in KOH. Basal
mycelium of 3.5–6(–7.5) µm wide, often thick-walled, septate,
hyaline to very pale yellow hyphae, unchanged in KOH, with
regularly arranged, spheroid, yellow to orange resinous exudates, dissolving in εδZ, partially and more slowly in KOH.
Specimens examined. dEnmark, NW Jylland, Klim Bjerg, soil along forest road, S.A. Elborne & K. Hansen, 16 Sept. 1998, KH.98.92 (C-F-53155).
– EStonia, Põlvamaa, ad terram in piceto humida, 11 Aug. 1960, A. Elango
(UPS F-629392). – Finland, Perä-Pohjanmaa, Rovaniemi, Pisajärvi, old Picea
forest, 2 Sept. 2011, T. Kekki, TK305 (TUR). – FranCE, Charente-εaritime, île
de Ré, près du camping ‘δa Bonne Étoile’, under Pinus maritima and Quercus
ilex, on leaf litter, 26 Nov. 2006, M. Hairaud, NV 2006.11.05 (dupl. S); Rhône,
δes Halles, col de Croix-Régis, 25 Oct. 2006, J. Cavet, NV 2006.10.33 (dupl.
S); Rhône, Saint-Nizier-d’Azergues, forêt de Pramenoux, sous Picea abies,
20 Sept. 2008, N. Van Vooren, NV 2008.09.08 (dupl. S). – gErmany, Lower
I. Olariaga et al.: A monograph of Otidea
Saxony, δüneburg, Boitze, Pinus and Abies, Oct. 2010, M. Vega private herb.
s.n. (dupl. S-F256977). – norway, Nord-Trøndelag, Snåsa, Bergsåsen Nature
Reserve, under Picea and Pinus, 2 Sept. 2009, K. Hansen & I. Olariaga,
KH.09.130 (S). – Spain, Navarre, Orokieta, Loiandi, Picea abies plantation,
17 Oct. 2008, J.M. Lekuona (ARAN-Fungi A5041195). – SwEdEn, Lappland,
3 miles NW Vuollerim, Bombmurkleskogen, along the Stora δuleälven, herbrich Picea forest, 19 Aug. 2000, M. Karström, εK200065 (S); δappland,
Norrbotten, εessaure, Kaltisbäcken Nature Reserve, herb rich Picea forest,
3 Sept. 2011, M. Karström, KH.11.60 (S); εedelpad, Södra Sillre, Hussborg,
on wood, 22 Aug. 1998, K. Olofsson (S-F256896); Närke, Hidinge, δekhyttan, Katte εajaskogen, coniferous forest on lime rich soil, B. Wasstorp, 13
Sept. 2008, JS.08.77 (S); Närke, Snavlunda, Ö Snavlunda Nature Reserve,
under Picea in a mixed forest, 12 Sept. 2008, L.G. Hellsten & A. Stridvall,
JS.08.68 (S); Närke, Vintrosa, Kanterboda skans Nature Reserve, on soil
under conifers, 10 Sept. 2008, A.B. Nilsson, JS.08.60 (S); Skåne, δoshult,
Lilla Loshult, Picea forest with a few broadleaf trees (Betula, Quercus), 5 Sept.
1998, S.-Å. Hanson, SÅH 105768 (C); Södermanland, Nacka, Kvarnhagen
by Söderbysjön, on soil in shadow, moist area, under Picea, 26 Sept. 2008,
J. Santos, JS.08.100 (S); Uppland, Björklinge, Drälinge, amongst needles
and mosses under pine in coniferous woods, 10 Sept. 1936, H.G. Bruun &
H. Smith (S-F92983, Fungi Exs. Suec. 3282); Uppland, Trehörningsskogen
Nature Reserve, under Picea on rich ground, on needle litter and decayed
wood, 1 Sept. 2011, M. Prieto & I. Olariaga, KH.11.77 (S). – USA, California, Del Norte Co., Earl δake State Park, access by Sand Hill Road, 15
Dec. 1997, M. Madsen & R. Davis (OSC 56826); California, Humboldt Co.,
Trinidad, Nov. 1931, H.E. Parks 3749 (UPS F-629376); Oregon, Benton Co.,
Corvallis, εcDonald-Dunn Research Forest, under conifers, 23 Oct. 1996,
E.T. Peterson (OSC 56761); Oregon, Benton Co., Corvallis, west side of NW
Beechwood Place, scattered clusters of apothecia on duff and adjacent to
rotting wood under Pseudotsuga menziesii, 19 Nov. 1997, N.S. Weber, NSW
8553 (OSC 150344); Oregon, Douglas Co., Bureau of δand εanagement,
Roseburg District, Swiftwater Resource Area, under Tsuga heterophylla,
Pseudotsuga menziesii, Polystichum munitum, Berberis nervosa and Holodiscus tricolor, 8 Nov. 2000, R. Furriel (OSC 105550); Oregon, εarion Co.,
Salem District Bureau of δand εanagement, Cascades Resource Area,
under Pseudotsuga menziesii, Tsuga heterophylla, Gautheria shallon, Polystichum munitum, Berberis nervosa, Oxalis oregana, Acer circinatum, Alnus
rubra and Rhododendron macrophyllum, 12 Nov. 1997, K. Dougan (OSC
66350); Washington, Bremerton, 26 Oct. 1942, J.B. Flett (UPS F-629383);
Washington, Eatonville, 18 Oct. 1954, A.H. Smith 49143 (UPS F-629385);
Washington, δower Nisqually R., εt Rainier National Park, 2 Sept. 1948,
A.H. Smith 30888 (UPS F-629384).
Notes — Otidea tuomikoskii is characterised by the narrowly
ear-shaped apothecia, with high warts on the outside, small
spores, and the excipulum almost always turning yellow in
KOH, together with the ochre to orange-ochre basal tomentum
in dried specimens. Otidea nannfeldtii is probably the species
that resembles O. tuomikoskii most, but O. nannfeldtii has
lower warts, lacks orange tones in the basal tomentum, has
paraphyses only rarely with slightly swollen areas, and resinous exudates on the outer excipulum that turn reddish in KOH.
Otidea papillata shares with O. tuomikoskii conspicuous dark
warts on the outside of the apothecia (see under O. papillata).
The yellow KOH reaction of the excipulum, especially strong in
the subhymenium and ectal excipulum, has been observed to be
constant, although weak in some collections. The KOH reaction
is stronger in recent collections, and can also be macroscopically
observed in fresh apothecia. The holotype of O. tuomikoskii is
from an anthill (Harmaja 1976), but O. tuomikoskii most often
produces apothecia among needle litter or even on very decayed
wood, in coniferous forests. It is widespread in Europe, where it
occurs in coniferous plantations, and in Western North America
(Peterson 1998), and has been found in Asia (Cao et al. 1990).
Otidea cantharella clade
Apothecia ear-shaped, or cup-shaped and entire, usually clearly
stipitate. Spores exceeding 20 µm, biguttulate and with several
additional small guttules (except in O. brunneoparva). Paraphyses often strongly notched. Associated with Picea.
Species — Otidea brunneoparva, O. cantharella, O. propinquata.
191
10. Otidea brunneoparva K. Hansen, ε. Carbone, Olariaga &
Van Vooren, sp. nov. — εycoBank εB537590; ITS barcode
GenBank: Kε010026; Fig. 13, 14
Etymology. Harmaja (2009a) used the epithet brunneoparva to provisionally name this species. The name is validated here and refers to the small
size and brown colour of the apothecia.
Holotype. SwEdEn, Närke, Knista, δekhyttan, Kungshall, calcareous oldgrowth forest, in thick litter layer, with Picea and Pinus, 12 Sept. 2008, K. Hansen,
KH.08.107 (S). Isotype (C).
Apothecia gregarious, 12–35 mm high, 7–25 mm wide, initially
ear-shaped, apex subacute, broadly ear-shaped in the end, seldom almost cup-shaped, split, stipitate. Hymenium dark brown
(6F3–6F7), sometimes olivaceous brown (5D7, 5E7, 5E8) or
reddish brown (7F3–7F5), sometimes paler in the margin, purplish ochre (5B3, 5B4, 6D7), when dried olivaceous brown (5F4,
5F5, 5F7, 5E7) or dark brown (6E6, 7F7, 7F8). Receptacle surface concolorous or slightly lighter, dark brown (5F6, 6F7, 7F8),
slightly hygrophanous, in drying golden brown (5D6, 5D7), when
dried dark brown (6E7, 7E7) to cinnamon brown (5D7, 6D7,
6E7), furfuraceous, sometimes finely warty at the base, often
longitudinally wrinkled at the base, sometimes almost reaching the margin. Warts hemispherical, gregarious, concolorous,
sometimes darker or paler brown than the background. Stipe
4–7 × 2–3 mm. Basal tomentum and mycelium white to very
pale brown, when dried ochre brown. Spores ellipsoid to broadly
ellipsoid, sometimes very slightly inequilateral, with two large
guttules, seldom with 1–2 additional smaller granules, smooth,
hyaline, (11–)11.5–14(–15) × 6.5–8.5 µm (δm = 11.7–13.8 µm,
Wm = 7.1–8.3 µm, Qm = 1.6–1.7; n = 6). Paraphyses hooked,
often inrolled, of the same width or slightly enlarged at the
apices to 3–5.5 µm wide, often with clear notches or forked at
apices, when fresh containing refractive, pale yellowish brown
guttules, restricted to the uppermost part of the paraphyses;
when dried pale yellow. Asci 131–195 × 8–10 µm. Apothecial
section 700–800 µm thick. Subhymenium c. 100–120 µm thick,
visible as a darker brown zone, cells cylindrical to swollen,
densely arranged, with scattered brown resinous exudates at
septa. Medullary excipulum of loosely woven textura intricata,
300–500 µm thick, hyphae cylindrical to slightly swollen, thickwalled, 5–11 µm wide, hyaline to very pale brown, sometimes
with brown resinous exudates at septa. Ectal excipulum of
textura angularis, 80 –100 µm, cells thick-walled, yellowish
brown, 20–47 × 12–25 µm. Surface with broadly conical warts,
25–65 µm high, composed of fasciculate, short hyphoid hairs.
Non-warted parts with scattered hyphoid hairs, of 2–3 subglobose to elongated cells, 7.5–10.5 µm wide, slightly constricted
at septa, sometimes with a thin gelatinous sheath. Resinous
exudates abundant, yellowish to reddish brown, dissolving into
amber drops in εδZ. Basal mycelium of 3–4.5 µm wide, very
pale brown hyphae, unchanged in KOH, smooth or with very
small, resinous exudates, dissolving in εδZ.
Specimens examined. Finland, Etelä-Häme, Hämeen, δammi, Evo,
Kotinen virgin forest, mesic forest of the Myrtillus type, in needle litter of
Pinus sylvestris and Picea abies in basal part of an active anthill, 8 Sept.
1978, H. Harmaja (S-F249386); Kainuu, Paltamo, Saukkovaara, under
Picea abies in moist spring-fed site, nearly in water, 24 Aug. 2011, M. Lahti
(TUR-A 198582); Koillismaa, Kuusamo, Oulanka National Park, first part of
the Kiutaköngäs trail, on rich soil among Picea and Betula leaves, 25 Aug.
2008, M. Carbone (TUR-A 198581); ibid., 16 Aug. 2009 (S-F257086, dupl.
TUR-A 198579); ibid., 14 Aug. 2010 (TUR-A 198580). – SwEdEn, Jämtland,
Östersund, Ändsjön Nature Reserve, on an abandoned anthill in rich Picea
forest, 26 Aug. 2009, K. Hansen & I. Olariaga, KH.09.82 (S); Närke, Hidinge,
δekhyttan, Katte εajaskogen, coniferous lime-rich forest, in litter, 13 Sept.
2008, B. Wasstorp, JS.08.73 (S); Närke, Knista, δekhyttan, Kungshall, calcareous oldgrowth forest, in thick litter layer, with Picea and Pinus, 12 Sept.
2008, J. Santos, JS.08.66 (S); ibid., JS.08.69 (S).
Notes — Otidea brunneoparva is morphologically and genetically a clearly distinct species. It is macroscopically character-
192
Persoonia – Volume 35, 2015
Fig. 13 Otidea brunneoparva apothecia. a. KH.08.107, holotype; b. JS.08.66; c. TUR-A 198579; d. TUR-A 198580. — Photos: b. J. Santos; c, d. ε. Carbone.
ised by stipitate, broadly ear-shaped apothecia with dark brown
colours and sometimes olivaceous shades. εicroscopically, the
spore shape and size are diagnostic (Fig. 14a), although with
wide variation. The strongly inrolled and notched apices of the
paraphyses are very characteristic (Fig. 14b) and otherwise only
found in a few species like O. propinquata or O. daliensis. The
thick-walled, yellowish brown, angular cells forming the outer
excipulum are diagnostic too (Fig. 14c, d); such thick-walled
cells in the outer excipulum are only otherwise found in O. propinquata. The six ITS sequences of O. brunneoparva, from five
different localities in Finland and Sweden, are identical or with
3–8 bp differences. The ITS nucleotide diversity within O. brunneoparva, as sampled here, is 0.74 % per site. Phylogenetic
analyses of four gene-regions (Hansen & Olariaga 2015) show
O. brunneoparva forms a distinct monophyletic group with O. propinquata and O. cantharella. These three species are nevertheless, easily distinguished both macro- and microscopically.
The distinctive spores of O. brunneoparva are only otherwise
found in O. leporina within Otidea. Otidea brunneoparva is
clearly distinguished from O. leporina by the darker coloured
apothecia and notched paraphyses. Otidea bufonia, O. mirabilis
and O. smithii share dark brown colours with O. brunneoparva,
but clearly differ in apothecial shape and stature (O. brunneoparva being more delicate), narrower spores (δm = 6.3–7.3 µm,
Qm = 1.9–2.5) of different shape (fusoid in O. bufonia and O. mirabilis) and the resinous exudates on the ectal excipulum not dissolving into amber drops. Otidea fusconigra was published as a
provisional name too. It also resembles O. brunneoparva in the
dark brown apothecia, and spore size and shape (Jamoni 2004).
Nevertheless, the paler hymenium colour (‘grey caffelatte’),
the paraphyses without notches and the habitat among alpine
dwarf Salix, suggests O. fusconigra is a different species. ITS
and LSU sequences of O. fusconigra (collection GεFN 2293),
obtained by us, confirms O. fusconigra is not conspecific, but
a sister taxon to O. smithii.
Following Cao et al. (1990), O. brunneoparva keys out as
O. olivacea J.Z. Cao & δ. Fan. The cup-shaped, dark brown
apothecia, with olivaceous tinge, suggest these may be closely
related. However, the spores of O. olivacea were described
as considerably longer (14 –17 × 8 – 8.5 µm), almost nonoverlapping with those of O. brunneoparva. Unfortunately, we
were not able to get the type specimen of O. olivacea on loan for
study. It should be noted that O. olivacea is a later homonym of
O. olivacea Bucholtz. Therefore Harmaja (2009b) published the
new name O. olivaceobrunnea for the illegitimate O. olivacea.
Otidea pusilla Rahm might be conspecific with O. brunneoparva, but the name is not validly published since no type was
indicated (Art. 40.1 ICN) and more than one gathering was
cited (Art. 40.2 ICN) (εcNeill et al. 2012), ‘collected over three
weeks in the same site’. The description of O. pusilla agrees with
O. brunneoparva in the dark brown, cup-shaped apothecia and
the relatively broad spores. The spore size given in the protologue was ‘15/6–9 µ’ (Rahm 1958), which we find difficult to
interpret due to the unusually broad width range. Unfortunately,
no material could be traced in ZH (pers. comm. R. Berndt).
A collection named by Harmaja as O. brunneoparva (H6017193)
was found to be morphologically identical to our material. Also
ITS and δSU sequences confirm that Harmaja´s and our material are conspecific. Therefore the name O. brunneoparva is
here adopted and validated as a new species.
Otidea brunneoparva appears to be rather widespread in the
Scandinavian Picea forests. While at least two of our collections, as indicated by Harmaja (2009a) grew on anthills, the
rest were found in places with abundant Picea litter, often with
presence of Betula. Two localities, including the type locality,
were calcareous suggesting O. brunneoparva is calciphilous.
I. Olariaga et al.: A monograph of Otidea
193
Fig. 14 Otidea brunneoparva (KH.08.107, holotype). a. Spores in water †; b. paraphyses*; c. medullary and ectal excipulum in water †; d. wart from the ectal
excipulum in water †; e. ectal excipulum in εelzer’s reagent †; f. basal mycelium in water †. — Scale bars = 10 µm; * = fresh material; † = dried material.
11. Otidea cantharella (Fr.) Quél., Enchir. Fung.: 275. 1886
— Fig. 15
Basionym. Peziza cantharella Fr., Syst. εycol. 2: 48. 1822 : Fr., loc. cit.
≡ Flavoscypha cantharella (Fr.) Harmaja, Karstenia 14: 107. 1974.
Neotype designated here: SwEdEn, Jämtland, Östersund, Ändsjön Nature
Reserve, in rich Picea forest, with Hepatica nobilis and Oxalis acetosella, 31
Aug. 2009, K. Hansen & I. Olariaga, KH.09.125 (S); εycoBank εBT178085.
= Peziza caligata Nyl., Ex Not. Sällsk. Fauna Fl. Fenn. Förh. 10: 8. 1868
‘1869’.
≡ Otidea caligata (Nyl.) Sacc., Syll. Fung. 8: 95. 1889.
≡ Acetabula caligata (Nyl.) Boud., Hist. Classific. Discomyc. Europe: 41.
1907.
Holotype. Finland, Uusimaa, Helsinki, 1850, W. Nylander (H009215) !
Misapplied names
– Otidea abietina sensu Breitenbach & Kränzlin, Fung. Switzerland 1: 82.
1984.
– Peziza propinquata sensu Nannfeldt, Ann. Bot. Fenn. 3: 313. 1966.
Apothecia gregarious or caespitose, 23–68 mm high, 13–43
mm wide, initially broadly ear-shaped, in the end broadly earshaped to almost cup-shaped, split, stipitate. Hymenium initially
light brown (5B5, 5B6, 5C6), then ochraceous yellow (4A5, 4A6,
5A5), ochre-orange (4A7, 5B7), sometimes with pinkish areas or
red dots (6B8), when wounded pinkish and margin brownish red
(6B8), when dried orange brown (6D7, 6D8) to reddish brown
(6C6, 6D6). Receptacle surface concolorous, ochre brown (4B7,
5C7, 5B6), slightly hygrophanous, in drying yellowish ochre (4A6,
4A7), sometimes with pale brown stains, when dried orange
194
brown (6D7, 6D8), furfuraceous to finely warty, sometimes
wrinkled at the base. Warts hemispherical, gregarious, concolorous, sometimes darker than the background, brown. Stipe
4–25 × 3–9 mm. Basal tomentum and mycelium abundant,
white to very pale brown (5A3), very pale brown when dried.
Spores ellipsoid and often narrowing toward poles, sometimes
very slightly inequilateral, with two large and several smaller
guttules, smooth, hyaline, (17–)18–21 × (9–)10–11.5(–12) µm
(δm = 17.7–20 µm, Wm = 10.4–11.4 µm, Qm = 1.7–1.8; n = 13).
Paraphyses hooked, of the same width or slightly enlarged at
apices, 2.5–3.5(–5.5) µm wide, without or with a low notch,
seldom forked at apices, when fresh containing small, refractive,
pale yellow guttules; when dried small, refractive granules. Asci
191– 217 × 11–12 µm. Apothecial section 900–1200 µm thick.
Persoonia – Volume 35, 2015
Subhymenium c. 70–100 µm thick, visible as a darker yellow
zone, of cylindrical to swollen cells, densely arranged. Medullary excipulum of textura intricata, (500–)700 –900 µm thick,
hyphae cylindrical to slightly swollen, thick-walled, 3–12 µm
wide, very pale yellow, sometimes with yellow-brown resinous
exudates at septa. Ectal excipulum of textura angularis 90–120
µm, cells thin-walled, pale yellow, 23–55 × 12–25(–30) µm.
Surface with conical to broadly conical warts, 55–80 µm high,
formed by fasciculate, parallel, short hyphoid hairs, of 3–4 ovoid
cells, constricted at septa, 5–8 µm wide. Non-warted parts with
single hyphoid hairs, of 2–4 subglobose to elongated cells,
slightly constricted at septa, 7–10 µm wide, sometimes with
a gelatinous sheath. Resinous exudates abundant, yellow to
yellow-brown, dissolving into amber drops in εδZ. Basal my-
Fig. 15 Otidea cantharella*. a, b. Apothecia; c. spores; d. paraphyses; e. medullary excipulum showing resinous exudates at septa; f. ectal excipulum (a:
KH.09.125, neotype; b: KH.11.69; c, d: KH.10.152; e: KH.09.144; f: KH.09.155). — Scale bars = 10 µm; * = all fresh material.
I. Olariaga et al.: A monograph of Otidea
celium of 3.5–4.5(–6) µm wide, hyaline to very pale yellowish
brown hyphae, unchanged in KOH, smooth or with very small,
regularly arranged, spheroid, resinous exudates, dissolving in
εδZ and partially in KOH.
Specimens examined. Finland, Etelä-Häme, Hattula, Parola, Alppilankallio, Vaccinium myrtillus-type forest, under Picea abies, 25 Sept. 1967, P. Uotila
618 (H); Etelä-Häme, Janakkala, Tervakoski, 6 Sept. 1970, P. Uotila 6195
(H); Etelä-Häme, εustiala, 5 Sept. 1895, J. Lindroth (H6010923); ibid., in
pineto, 24 Aug. 1866, P.A. Karsten (H6010922); Etelä-Häme, εustiala, Tammela, 2 Sept. 1882, P.A. Karsten (H6010829); Perä-Pohjanmaa, Rovaniemi,
Kalkkinulkki, near old limestone quarry, under Picea, 23 Aug. 2011, T. Kekki,
TK211 (TUR); Perä-Pohjanmaa, Rovaniemi, Ounasvaara, Picea forest, 29
Aug. 2011, T. Kekki, TK279 (TUR); Perä-Pohjanmaa, Rovaniemi, Välijoki, calciferous Picea forest, 25 Aug. 2011, T. Kekki, TK236 (TUR); Perä-Pohjanmaa,
Tervola, Peura, old calciferous Picea forest, 5 Sept. 2011, T. Kekki, TK177
(TUR); Perä-Pohjanmaa, Ylitornio, Kuusikkorommas, calciferous Picea and
Pinus forest, 2 Sept. 2011, T. Kekki, TK301 (TUR); Varsinais-Suomi, Vihti,
Nummela, εetsäkulma, on soil, 16 Sept. 1979, H. Kotiranta (H6010925).
– FranCE, Isère, Villard-de-δans, Bois Barbu, under Picea abies, 20 Sept.
2008, J. Cavet, NV 2008.09.16 (dupl. S). – hungary, Hohe Tatra bei UnterSchmecks, Oct. 1884, Linhart (UPS F-629314, Rehm Ascomyceten 251b,
as O. leporina f. minor). – italy, Trentino-Alto Adige, Cavelonte, in silvis
coniferis, semper socia Cudoniae confusae, Aug. 1898, G. Bresadola (UPS
F-629361). – norway, Nord-Trøndelag, Steinkjer, Noem, under Picea, 2 Sept.
2009, K. Hansen & I. Olariaga, KH.09.129 (S); Nord-Trøndelag, Steinkjer,
Strattåsen, in rich Picea forest, 5 Sept. 2009, K. Hansen & I. Olariaga,
KH.09.144 (S). – SwEdEn, Blekinge, Rödeby, Spjutsbygd, c. 2 km NW from
the train station, on needle litter under a Picea in coniferous forest, 10 Sept.
1946, S. Lundell & S. Wikland (UPS F-146484); Gästrikland, close to Bergby,
mossy Picea forest on acidic ground, 31 Aug. 2010, K. Hansen, K. Gillen &
I. Olariaga, KH.10.152 (S); Hälsingland, Kårböle, St Olofs, 26 Aug. 2001,
H.-G. Toresson s.n. (S); Hälsingland, south to crossing between roads E45
and 310, under Picea abies, among leaf litter, 21 Aug. 2011, J.C. Zamora
& I. Olariaga, KH.11.109 (S); Härjedalen, δinsell, Djursvallen, montane coniferous forest with birch, on very rotten log, 13 Aug. 2000, B. Gahne (UPS
F-125589); Härjedalen, Torkilstöten, under Picea abies among leaf litter,
on acidic ground, 20 Aug. 2011, J.C. Zamora & I. Olariaga, KH.11.106 (S);
Jämtland, Hammarstrand, Picea mossy forest, 26 Aug. 2009, H. Lindström,
KH.09.83 (S); ibid., KH.09.84 (S); Jämtland, SW of εörsil, Sandtjärndalen
Nature Reserve, under Picea abies on rich ground, 7 Sept. 2009, K. Hansen
& I. Olariaga, KH.09.155 (S); Jämtland, Sällsjö surroundings, in young
stand of Picea abies, on rich ground with Betula and Salix, 29 Aug. 2009,
K. Hansen & I. Olariaga, KH.09.104 (S); Jämtland, Östersund, Andersön
Nature Reserve, Picea forest on rich ground, 28 Aug. 2009, K. Hansen &
I. Olariaga, KH.09.95 (S); ibid., under Picea and Pinus, KH.09.96 (S); ibid.,
KH.09.101 (S); Jämtland, Östersund, Fillstabäcken Nature Reserve, on old
anthill under Picea, K. Hansen & I. Olariaga, 30 Aug. 2009, KH.09.111 (S);
ibid., KH.09.117 (S); ibid., K. Hansen & X.H. Wang, 5 Sept. 2012, KH.12.99
(S); Jämtland, Östersund, Ändsjön Nature Reserve, in rich Picea forest, with
Hepatica nobilis and Oxalis acetosella, 26 Aug. 2009, K. Hansen & I. Olariaga,
KH.09.78 (S); ibid., KH.09.80 (S); ibid., KH.09.98 (S); Jämtland, Åre, Kall,
along Stor-Grundsviken, Kallsjön, on old anthill in lime rich Picea forest, 15
Aug. 2008, J. Santos & K. Hansen, JS.08.18 (S); δappland, Jokkmokk, SE
of Vuollerim, Andersviksravinerna (Natura 2000 area), mixed forest, 31 Aug.
2011, A. Stridvall, KH.11.111 (S); ibid., KH.11.112 (S); δappland, Jokkmokk,
by Kassavare mountain, Köpenhamn, under Picea abies, 1 Sept. 2011,
K. Hansen & I. Olariaga, KH.11.110 (S); δappland, Jokkmokk, ‘Nornaskogen’
by Ållojaur, mossy Picea forest on rich ground, 29 Aug. 2011, K. Hansen &
I. Olariaga, KH.11.16 (S); δappland, Kuouka, 15 km SE εessaure, herb-rich
Picea forest on rich ground, among mosses, 3 Sept. 2011, K. Hansen &
I. Olariaga, KH.11.68 (S); ibid., KH.11.69 (S); δappland, 5 km SE Vuollerim,
part of Andersviksravinerna, Rävabacken Nature Reserve, herb-rich Picea
forest, 21 Aug. 2000, M. Karström, εK200061 (S); ibid., under Picea on
mossy ground, together with Cudonia confusa, 2 Sept. 2011, K. Wiking,
KH.11.59 (S); Småland, Ryssby, Gärdsholmen, Björnö, coniferous forest,
6 Sept. 1930, H.G. Bruun (UPS F-146485); ibid., on deep moss in coniferous forest (UPS F-146486); Uppland, Uppsala, Sävja, Norra δunsen Nature
Reserve, Picea forest, on soil, thick layer of litter and mosses, 28 Aug. 2008,
J. Santos, JS.08.47 (S). – SwitzErland, Wallis Canton, δiddes, Palazuit, in
side of brook under Picea abies, 17 Aug. 2008, M. Carbone (εCVE 24217).
Notes — Otidea cantharella can be recognised by broadly
ear-shaped apothecia, with yellow to orange tones, often with
a well-developed stipe, and large spores. No other species of
Otidea have the spore size of O. cantharella, in combination
with those macroscopic characters.
195
Otidea cantharella is associated with Picea abies, and often
produces apothecia on anthills or thick needle layers. Furthermore, we have often found apothecia of O. cantharella and
Cudonia confusa together (Fig. 15a), and once with Spathularia
rufa, both species belonging to the Rhytismatales. An apparent
closer association in the same fairy ring has also been observed
once. Interestingly, Bresadola noted the association with C. confusa in one of his collections (UPS F-629361) studied by us. The
presence of C. confusa in the same collecting spot may give
a first plausible field identification of O. cantharella. εoreover,
it suggests a possible biotrophic association between O. cantharella and C. confusa.
Nomenclatural notes — This species has been referred to
as O. caligata (Nyl.) Sacc. (Nannfeldt 1966, Dissing 2000).
Nevertheless, Harmaja (2009a) came to the conclusion that
the name O. cantharella must refer to the large-spored species
treated here, with which some authors have later disagreed
(Carbone 2010b, Van Vooren 2011b). Harmaja stressed the
following characters from the protologue supporting the usage
of the name as presented here: ear-shaped, stipitate apothecia,
the yellow colour of Cantharellus cibarius and the occurrence
in Picea forests in southern Sweden. We have studied material
from Småland (UPS F-146485, UPS F-146486), from where
Fries described O. cantharella. It is therefore very likely that
Fries had in mind the species described here when he coined
the name O. cantharella, although he had only seen dried
material, as indicated by the abbreviation ‘v. s.’ (vidi siccam,
seen dried). Since no original material is known to exist, we
propose a neotype that will attach the name O. cantharella to
the Otidea species described here. This stabilises the interpretation proposed by Harmaja (2009a), also used by εornand &
Courtecuisse (2005), and long before adopted by Bresadola
(1900: 102), and should serve to settle the interpretation of
O. cantharella. Our ITS-δSU sequences of four O. cantharella
collections from Sweden and France are identical.
12. Otidea propinquata (P. Karst.) Harmaja, Karstenia 15: 32.
1976 — Fig. 16
Basionym. Peziza propinquata P. Karst., Not. Sallsk. Fauna Fl. Fenn.
Forh. 10: 110. 1869.
Lectotype. Finland, Tavastland, εessuby, 7 Oct. 1860, P.A. Karsten
(H6010807) !, selected by Nannfeldt (1966).
= Otidea abietina f. nigra Rick, Oesterr. Bot. Z. 48: 62. 1898.
≡ Otidea abietina var. nigra (Rick) Sacc., Syll. Fung. 14: 746. 1899.
Lectotype. auStria, Vorarlberg, an der Gamp, im Nadelwald, 1700 m,
Sept. 1897, Rick (S-F9962) !, indicated by Nannfeldt (1966).
= Otidea indivisa Velen., εonograph. Discom. Bohemiae 1: 355. 1934.
Lectotype. CzECh rEpubliC, Karlštejn N, Oct. 1922, J. Fechtner (PRε
149147) !, selected by Nannfeldt (1966).
Misapplied names
– Pseudotis abietina sensu Boudier, Icon. εycol. livr. 7: nº. 131, pl. 333.
1906 (preliminary text with ‘circulaires’).
– Otidea cochleata sensu Breitenbach & Kränzlin, Fung. Switzerland 1: 84.
1984.
Apothecia gregarious or caespitose, 7–20 mm high, 15–35
mm wide, obconical to broadly cup-shaped, entire, very rarely
split, regular or sometimes undulate in the margin, stipitate.
Hymenium ochre brown (5C7, 6D5, 6D6) to dark reddish brown
(6F6, 6F7, 7F7, 7F8), when dried ochre brown (6B7, 6C7).
Receptacle surface concolorous, orange brown (6F6, 6F7),
slightly hygrophanous, in drying yellowish brown (5C6, 5C7),
when dried orange brown (6D6, 6D7, 6E7), furfuraceous to
warty, sometimes wrinkled at the base. Warts hemispherical,
gregarious, concolorous, sometimes darker than the background, brown. Stipe 7–16 × 2–7 mm. Basal tomentum and
mycelium abundant, white to very pale brown (5A3), very pale
brown when dried. Spores ellipsoid and often narrowing toward
196
Persoonia – Volume 35, 2015
Fig. 16 Otidea propinquata. a. Apothecia; b. spores in water †; c. paraphyses in KOH †; d. wart of the ectal excipulum in water †; e. ectal excipulum in KOH †;
f. basal mycelium in water† (a: KH.11.21; b–f: KH.09.99). — Scale bars = 10 µm; † = dried material.
the poles, sometimes very slightly inequilateral, with two large
and several smaller guttules, smooth, hyaline, (18–)19–21 ×
10–12.5 µm (δm = 19.3–20 µm, Wm = 10.9–11.6 µm, Qm = 1.6–
1.7; n = 4). Paraphyses hooked, of the same width or often
enlarged at apices, 3–5 µm wide, often with 1–3 notches or
forked at apices, when fresh containing small, refractive, light
yellow guttules; when dried hyaline. Asci 231– 275 × 12–16
µm. Apothecial section 1100–1500 µm thick. Subhymenium
c. 100–120 µm thick, of dense textura intricata, visible as a
darker brown zone. Medullary excipulum of textura intricata,
600–800 µm thick, hyphae 4–9 µm wide, sometimes slightly
swollen, thin- to thick-walled, very pale brown, sometimes with
brown resinous exudates at septa. Ectal excipulum of textura
angularis 100–130 µm, cells rather thick-walled, pale brown,
17–50 × 10–40 µm. Surface with conical to broadly conical
warts, 50–75 µm high, formed by short, fasciculate hyphoid
hairs, of 2 – 3 subglobose to elongated cells, constricted at
septa or not, 8–11 µm wide, sometimes with a pale brown thick
gelatinous sheath. Resinous exudates abundant, yellow-brown
to brown, turning brownish red in KOH, dissolving in εδZ. Basal
mycelium of 3–6 µm wide, hyaline to pale brown hyphae, unchanged in KOH, smooth or normally with regularly arranged,
spheroid to rod-shaped, resinous exudates, dissolving in εδZ
and partially in KOH.
Specimens examined. CzECh rEpubliC, Praha, Karlštejn, 9 Oct. 1922,
F. Fechtner (UPS F-629369, syntype of O. indivisa); Sept. 1924 (UPS F-629367,
syntype of O. indivisa). – dEnmark, NE Sjælland, Asserbo Plantage, under
Pinus and Picea among needles, 7 Oct. 1975, H. Knudsen (C-F-87203).
I. Olariaga et al.: A monograph of Otidea
– FranCE, Isère, δans-en-Vercors, on the ground, on Picea leaf litter, 17
Sept. 2008, J. Cavet, NV 2008.09.15 (dupl. S). – italy, Trentino-Alto Adige,
Sopramonte, ad acus abiegnos in sylvis coniferis, Aug. 1898, G. Bresadola
(UPS F-629357). – SwEdEn, Jämtland, Sällsjö surroundings, in young stand
of Picea abies, on rich ground, 28 Aug. 2009, K. Hansen & I. Olariaga,
KH.09.99 (S); Jämtland, Östersund, Andersön Nature Reserve, under Pinus
and Picea, on rich ground, among mosses, 28 Aug. 2009, K. Hansen &
I. Olariaga, KH.09.94 (S); ibid., KH.09.103 (S); Jämtland, Östersund, Ändsjön
Nature Reserve, in rich Picea forest, 26 Aug. 2009, K. Hansen & I. Olariaga,
KH.09.81 (S); ibid., 31 Aug. 2009, KH.09.123 (S); δappland, Jokkmokk,
‘Nornaskogen’ by Ållojaur, mossy Picea forest on rich ground, 28 Aug.
2011, K. Hansen & I. Olariaga, KH.11.21 (S); δappland, Jokkmokk, Ultevis
Fjällurskog Nature Reserve, Sitoätno, Picea mossy forest on rich ground,
among leaf litter, 31 Aug. 2011, K. Hansen & I. Olariaga, KH.11.30 (S); ibid.,
KH.11.35 (S); δappland, S of Kvikkjobb-Kabla FUR Nature Reserve, by
Kassavare mountain, Köpenhamn, young Picea stand by a road, 1 Sept.
2011, K. Hansen & I. Olariaga, KH.11.53 (S); δappland, εessaure, herbrich Picea forest, 29 Aug. 2008, M. Karström, εK0834 (S); δappland, 17 km
WSW of Vuollerim, Slubbojaureskogen, herb-rich Picea forest, 8 Aug. 2002,
M. Karström, εK0222 (S); Närke, Knista, δekhyttan, under Picea, in thick litter
layer, in lime rich forest, 12 Sept. 2008, J. Santos, JS.08.67 (S); Uppland,
Uppsala, Sätra Nature Reserve, on litter on lime rich soil, under Picea, 23
Sept. 2008, J. Santos & K. Hansen, JS.08.98 (S). – USA, Washington, δake
Crescent, under fir, 28 Oct. 1935, A.H. Smith (UPS F-629351).
Notes — Otidea propinquata is easily recognised by the
stipitate, entire, brown apothecia, large spores, and notched
or forked paraphyses. Otidea daliensis shares brown, entire
apothecia with O. propinquata, but differs in having sessile
apothecia, with purplish brown tones, and basal mycelium that
lacks abundant resinous exudates.
There has been uncertainty surrounding the correct name of
this taxon. A number of early authors used the epithet abietina
to refer to O. propinquata (Boudier 1906, Bresadola 1933). In
agreement with Harmaja (2009a) and Carbone (2010c), we
consider Peziza abietina a nomen confusum. Nannfeldt (1966)
referred to O. propinquata as O. indivisa and argued that the
type of O. propinquata was conspecific with O. cantharella (as
caligata). Harmaja (1976) examined the lectotype of O. propinquata and found it to be clearly distinct from O. cantharella and
O. indivisa, a later synonym. This is confirmed by our study of
the type of O. propinquata. Otidea propinquata occurs in Picea
forests on calcareous ground. It is widespread in Northern Fennoscandia and present in central Europe.
Otidea formicarum clade
Apothecia ear- to cup-shaped, split, ochre to reddish brown.
Spores small, 9.5–12 µm long. Basal mycelium with abundant
yellow resinous exudates. Associated with conifers.
Species — Otidea formicarum, O. nannfeldtii, O. subformicarum, O. aff. subformicarum, Otidea sp. ‘b’.
13. Otidea formicarum Harmaja, Karstenia 15: 31. 1976 — Fig.
5f, 17
Holotype. Finland, Etelä-Karjala, εiehikkäla, Savanjärvi, on anthill in
spruce forest, 26 Sept. 1970, L. Fagerström (H6003549) !
Apothecia gregarious to caespitose, 8–22 mm high, 5–20 mm
wide, broadly ear-shaped, upper margin rounded, then expanding and sometimes in the end becoming cup-shaped and flattened, split, stipitate or sessile. Hymenium yellowish brown
(5C7, 5C8) to reddish brown (6C7, 6C8, 6D8), when dried ochre
(4A5, 4A6) to brownish ochre (5B7). Receptacle surface yellowish brown (5C6) to reddish brown (6C6), hygrophanous,
in drying ochre (5B7), when dried ochre (4A5, 4A6), finely
warty, smooth at the base. Warts broadly conical to rounded,
gregarious, concolorous, sometimes slightly darker than the
background, brown. Stipe 3 – 6 × 2–4 mm. Basal tomentum
and mycelium abundant, light yellow-ochre (5A2). Spores ellip-
197
soid, seldom very slightly inequilateral, with two large guttules,
smooth, hyaline, 9.5–11(–11.5) × (5.5–)6–7 µm (δm = 10–10.7
µm, Wm = 6–6.9 µm, Qm = 1.6–1.7; n = 7). Paraphyses curved
to hooked, of the same width or slightly enlarged at apices, 3–4
µm wide, with up to two slightly swollen areas, occasionally
notched, sometimes when fresh containing small, refractive,
light yellow or green guttules; when dried small, refractive,
hyaline to light yellow granules. Asci 160–232 × 10.5–13 µm.
Apothecial section 700–800 µm thick. Subhymenium c. 80–100
µm thick, of dense textura intricata, visible as a yellowish brown
zone, cells cylindrical to swollen, densely arranged. Medullary
excipulum 400–500 µm thick, differentiated into two parts: a)
textura angularis underneath the subhymenium, 120–200 µm
thick, cells 6–15 µm broad; b) textura intricata, hyphae sometimes slightly swollen, thin-walled to thick-walled, 3–11 µm wide,
hyaline to very pale yellow, sometimes with pale yellow resinous exudates at septa. Ectal excipulum of textura angularis,
70–120 µm, cells thin-walled, light yellow, 17–40 × 11–25 µm.
Surface with broadly conical warts, 45–60 µm high, formed by
short, fasciculate, hyphoid hairs, of 2–3 elongated cells, 7–12
µm wide, not or slightly constricted at septa, sometimes with
a gelatinous sheath. Resinous exudates abundant, brownish
yellow, dissolving into amber drops in εδZ. Basal mycelium of
3–5 µm wide, hyaline to very light yellow hyphae, unchanged
in KOH, with regularly arranged, very small, spheroid resinous
exudates, dissolving in εδZ, partially and more slowly in KOH.
Specimens examined. Finland, Etelä-Häme, δammi, Evo, Vahtervehmas,
Kotinen virgin forest, 6 Sept. 1988, H. Harmaja (H6003551); Etelä-Häme,
δoppi, Topeno, Piimästennummi, on a huge anthill, under Picea, 16 Sept.
2011, S. Huhtinen 11/65 (TUR); Perä-Pohjanmaa, Rovaniemi, Välijoki, calciferous Picea forest, on anthill, 25 Aug. 2011, T. Kekki, TK223 (TUR); Uusimaa,
Elimäki, Villikkala, δääksynmäki, mesic heath spruce forest, on anthill, 22 Oct.
2005, U. Nummela-Salo & P. Salo (H6003550); Varsinais-Suomi, Koski Tl.,
Hongisto, on old anthill under Picea abies, with Betula sp., Salix caprea and
Pinus sylvestris, 8 Aug. 1998, M.-L. & P. Heinonen (TUR 124728); VarsinaisSuomi, δieto, Suopohja, SE of Päivärinne, along the road by the house EskoUkura, on old anthill under Picea abies, 2 Oct. 2009, K. Ruottinen (TUR-A
183242). – FranCE, Haute Savoie, Thorens-Glières, plateau des Glières,
under Picea, on the ground, 22 Sept. 2006, L. Francini, NV 2006.09.11 (S).
– norway, Nordland, Grane, Holmvassdalen Nature Reserve, on old anthill
under Picea, 29 Oct. 2009, J. Lorås (S-F244372). – SwEdEn, Dalarna, Stora
Tuna, between Falubäcken and Övre εorbygge fäbod, abundant in the lower
part of an active anthill, 22 Sept. 1963, R. Morander (UPS F-146725); Närke,
Askersund, Orkarebäckens Nature Reserve, calcareous forest, on an active anthill under Picea, 11 Sept. 2008, J. Santos, JS.08.63 (S); ibid., under
Picea in litter, JS.08.62 (S); Uppland, Bondkyrka, Gottsundabergen, 1 Oct.
1927, J.A. Nannfeldt (UPS F-146706); Uppland, Bondkyrka, Nåsten, S of
δäbyvad station, among coniferous needles, 16 Sept. 1932, J.A. Nannfeldt
(UPS F-146709); Uppland, Älvkarleby, Billuddens Nature Reserve, under
Pinus and Picea on sandy ground, at the base of a Pinus, 15 Sept. 2011,
J.C. Zamora & I. Olariaga, KH.11.104 (S); Ångermanland, Ullanger, Håll, in
the eastern slope of εt εoberget, c. 0.5 km SW of p. 35,29, on the ground,
14 Sept. 1974, R. Moberg (UPS F-146735).
Notes — Otidea formicarum is characterised by relatively
small, reddish brown, broadly ear-shaped apothecia, small
spores, and by its habitat, often occurring on anthills. For a
comparison with the sister species O. subformicarum see under
that species below. Otidea nannfeldtii resembles O. formicarum,
but differs in often having more narrowly ear-shaped and paler
coloured apothecia, sometimes with pink tones in the hymenium. Dried apothecia of O. nannfeldtii have, on the contrary,
darker colours than O. formicarum. Otidea pseudoleporina and
O. formicarum share apothecial shape, but O. pseudoleporina
can be distinguished by the ochre-orange colour, and comparatively narrower spores (Qm = 1.7–1.9).
The original description of O. formicarum was based on several
collections made on anthills in Finland (Harmaja 1976), and
more recent material has been cited from anthills (Harmaja
2009a, b), mainly associated with Picea. εost of the material
198
examined by us was also from anthills, but at least one of our
Swedish finds (KH.11.104) was from Pinus needle litter, which
shows the habitat of O. formicarum partly overlaps with the
habitat of O. subformicarum.
14. Otidea nannfeldtii Harmaja, Karstenia 15: 31. 1976 — Fig.
5a, d, 18
Holotype. Finland, Ahvenanmaa, δemland, Nåtö, spruce forest near Övergård, 17 Sept. 1972, C.-A. Haeggström (H6002902) !
= Otidea angusta Harmaja, Karstenia 48: 35. 2009.
Holotype. Finland, Varsinais-Suomi, δohja, Jalassaari, Ahtiala, E of Heimo
house, mixed somewhat moist rich woods with Picea, Betula, Corylus etc.
on somewhat calcareous soils, 23 Aug. 1965, H. Harmaja (H6010804) !
Persoonia – Volume 35, 2015
Misapplied names
– Otidea papillata sensu Van Vooren et al., Bull. εycol. Bot. Dauphiné-Savoie
188: 52. 2008.
Apothecia gregarious to caespitose, 8–35 mm high, 5–15 mm
wide, initially long, narrowly ear-shaped, sometimes expanding
and becoming broadly ear-shaped, split, stipitate or sessile.
Hymenium ochre (4A5), orangish ochre (5B4, 5C4) or pale
brown (6D6, 6D7), sometimes with pink tones or entirely pinkish (6A4), when dried orange-ochre (5A5) to reddish brown
(6C6). Receptacle surface brown orange (5B4, 5C4) to pale
brown (6D6–7), slightly hygrophanous, in drying ochre (4A5),
when dried reddish brown (7D7, 7E7), finely warty, smooth at
the base. Warts conical, gregarious, concolorous, sometimes
distinctly darker than the background, brown. Stipe 4 – 8 ×
Fig. 17 Otidea formicarum (KH.11.104)*. a. Apothecia; b. spores; c. paraphyses; d. ectal excipulum; e. resinous exudates of the ectal excipulum; f. basal
mycelium. — Scale bars = 10 µm; * = all fresh material.
I. Olariaga et al.: A monograph of Otidea
2–3 mm. Basal tomentum and mycelium abundant, white to
light yellow (5A2) or ochre (5A3). Spores ellipsoid, sometimes
slightly inequilateral, with two large guttules, smooth, hyaline,
(9 –)9.5 –10.5(–11.5) × 5.5 – 6.5(–7) µm (δm = 9.8 –10.9 µm,
Wm = 5.5–6.6 µm, Qm = 1.6–1.9; n = 10). Paraphyses curved
to hooked, of the same width or slightly broader at apices,
2.5–5 µm wide, without notches, rarely with up to two slightly
swollen areas or forked at apices, when fresh containing small,
refractive, light yellow guttules; when dried hyaline to light yellow. Asci 137–190 × 8–10 µm. Apothecial section 650–900 µm
thick. Subhymenium c. 80–100 µm thick, visible as a yellowish
brown zone, of cylindrical to swollen cells, densely arranged.
Medullary excipulum of textura intricata, 400–600 µm thick,
differentiated into two parts: a) textura angularis underneath
the subhymenium, 100–150 µm thick, cells 13–32 × 12–21
199
µm; b) textura intricata, 400–450 µm thick, hyphae 4–9 µm
wide, thin-walled to slightly thick-walled, hyaline to very pale
yellow, sometimes with pale yellow resinous exudates at septa.
Ectal excipulum of textura angularis, sometimes of a textura
prismatica, 80–120 µm, cells thin-walled, hyaline to light brown,
16–40 × 11–22 µm. Outer part of conical to broadly conical
warts, 45–85 µm high, formed by short, fasciculate, hyphoid
hairs, of 2–3(–4) subglobose to elongated cells, constricted
at septa, 5–9 µm wide, sometimes with a gelatinous sheath.
Resinous exudates abundant, yellow brown, dissolving into
amber drops in εδZ, reddish brown in KOH. Basal mycelium
of 3–5.5 µm wide, septate, hyaline to very light yellow hyphae,
unchanged in KOH, with very small, regularly arranged, spheroid, yellow resinous exudates, dissolving in εδZ, partially and
more slowly in KOH.
Fig. 18 Otidea nannfeldtii. a. Apothecia; b. spores*; c. paraphyses*; d. ectal excipulum*; e. hyphoid hairs with gelatinous sheath*; f. amber drops on the
outermost ectal excipulum cells in εelzer’s reagent † (a–e: KH.10.302; f: S-F249387). — Scale bars = 10 µm; * = fresh material; † = dried material.
200
Specimens examined. Finland, Varsinais-Suomi, δohja, Jalassaari, Ahtiala, Alho, very close to the Ahtiala Nature Reserve, below a spruce tree in
mixed forest on fairly calcareous soil, 29 Sept. 1978, H. Harmaja (S-F249387,
ex H6017194 as O. lohjaënsis nom. prov.). – FranCE, Hautes Alpes, ProvenceAlpes-Côte-d’Azur, δa Bâtie-εontsaléon, δes Chariots du Buech, au sol dans
la litière d’aiguilles de pins (P. sylvestris), 26 Oct. 2008, N. Van Vooren, NV
2008.10.01 (dupl. S). – italy, Abruzzo, Pietracamela (TE), Prati di Tivo, on
soil in mixed forest, mainly with Larix but also Picea and Pinus, 2 Oct. 2009,
B. De Ruvo (S-F257096); Calabria, Celico (CS), Contrada Colamauci, under
Pinus and Picea abies, 16 Sept. 2009, C. Lavorato, Cδ 091116-17 (dupl. S);
Calabria, εorano Calabro (CS), Campotenese, under Pinus sylvestris, 7 Dec.
2009, C. Lavorato, Cδ 091207-01 (dupl. S). – SwEdEn, Gotland, Ollajvs Nature
Reserve, close to δjugarn, under Picea and Pinus on calcareous ground,
31 Sept. 2010, K. Hansen, K. Gillen & I. Olariaga, KH.10.302 (S); ibid., under
Picea and Pinus on rich calcareous soil, 27 Sept. 2011, H. Tuovila & S. Huhtinen, KH.11.115 (S); δappland, Jokkmokk, ‘Nornaskogen’ by Ållojaur, under
Picea among mosses, K. Olofsson, 1 Sept. 2011, KH.11.45 (S); δappland,
εessaure, 3 miles NW Vuollerim, herb rich Picea forest, 1 Sept. 2005,
M. Karström, εK0536 (S); δappland, Västra Tjetnekbäcken, 9 km E of
Jokkmokk, under Picea abies, K. Olofsson, 1 Sept. 2011, KH.11.112 (S);
Uppland, Stockholm, Enebyberg, Rinkebyskogen, on soil under Picea and
deciduous trees in side of footpath, 29 Sept. 2008, J. Santos, JS.08.103
(S). – USA, Oregon, Douglas Co., Thielsen Creek, under conifers, 13 Oct.
2010, R. Helliwell, rh101310 (OSC).
Otidea cf. nannfeldtii — dEnmark, δøvenholm Skov, δangsø, 25 km W of
Grenaa, 30 Sept. 1968, H. Folkmar (C-F-48295), (C-F-48296), (C-F-48297).
Notes — Otidea nannfeldtii is characterised by ochre to light
brown, narrowly ear-shaped apothecia, small spores and resinous exudates on the ectal excipulum turning reddish brown in
KOH. It resembles other species with ear-shaped apothecia and
small spores, such as O. formicarum, O. papillata, O. pseudoleporina and O. tuomikoskii. The most similar species is O. formicarum (see under O. formicarum). Otidea tuomikoskii is separated from O. nannfeldtii by higher and more densely placed
warts on the apothecial outer surface, along with the yellow
reaction of the excipulum in KOH. Otidea pseudoleporina has
a brighter ochre-orange hymenium, and the resinous exudates
of the outer excipulum only partly convert into reddish grey
heterogeneous drops in KOH.
Otidea nannfeldtii, as treated here, shows some phylogenetic
structure (Fig. 1; but see Hansen & Olariaga 2015). Recently
O. angusta was described as distinct from O. nannfeldtii,
based on a few subtle characters, i.e. taller and slightly thicker
fleshed apothecia, with very faintly brownish basal mycelium,
paraphyses with shorter and thinner apical cells, and maybe
smaller spores (Harmaja 2009a). At that time only a single
collection of each species was known. Based on studies of
additional collections, we did not find correlations between
any morphological characters and the subgroups supported
by our multiple molecular phylogenies. Therefore, we propose
O. angusta be placed in synonymy with O. nannfeldtii. Supporting this, our study of the holotype of O. angusta revealed larger
spores (9.5–11 × 5–5.8 µm) than cited in the protologue (8–9.8
× 4.5–5.2 µm), and broader paraphyses (2–3 µm) with longer
terminal cells (up to 67 µm). We have also observed both tall,
narrow and rounded apothecia of O. nannfeldtii in the same
spots, and we consider the apothecial shape to vary during
development. Otidea lohjaënsis, proposed by Harmaja (2009a)
as a provisional name, is also suggested to be conspecific
with O. nannfeldtii. A collection identified by Harmaja as
O. lohjaënsis (S-F249387, ex-H6017194) showed excipular
resinous exudates dissolving into amber drops in εδZ (Fig.
18f), contrary to the key character given for O. lohjaënsis (i.e.
not responding to εδZ). The variation in the ITS region within
O. nannfeldtii as recognised here is high, but displays no length
variation (alignment 790 bp long). The ITS sequences of the
Finnish holotypes of O. angusta and O. nannfeldtii show 26 bp
differences. At the same time, however, the ITS sequence of
the Swedish KH.10.302 shows 33 and 32 bp differences from
the O. nannfeldtii and O. angusta holotypes, respectively. The
Persoonia – Volume 35, 2015
North American rh101310 shows 14 and 20 bp differences from
the two holotypes, O. nannfeldtii and O. angusta, respectively.
The collections showing no δSU sequence variation (Fig. 1),
show also no, or only 1–2 bp differences in the ITS region. No
variation was found between the holotype of O. angusta and
the Finnish S-F249387 (identified as the provisional O. lohjaënsis by Harmaja).
Otidea nannfeldtii was previously known only from South West
Finland (Harmaja 1976, 2009a). Here we report it from other
areas in Europe, and for the first time from North America.
15. Otidea subformicarum Olariaga, Van Vooren, ε. Carbone
& K. Hansen, sp. nov. — εycoBank εB809252; ITS barcode GenBank: Kε010054; Fig. 19, 21
Etymology. Referring to its similarity to O. formicarum.
Holotype. Spain, Huesca, Bielsa, Ermita de Nª Señora de Pineta, 1270 m,
42.638039, 0.180532 (decimal format), under Pinus sylvestris and Abies alba
on calcareous ground, 13 Oct. 2012, J.C. Campos & J. Herranz (S-F242696).
Apothecia gregarious to caespitose, 13–32 mm high, 10–20 mm
wide, broadly ear-shaped, soon becoming deeply cup-shaped,
split, margin sometimes lobate, sessile or stipitate. Hymenium
orange brown (6C6, 6C8) to dark reddish brown (6D7, 6D8),
slightly hygrophanous, when dried orange brown (7A8). Receptacle surface orange brown (6B7, 6C7) to reddish brown
(6D8), hygrophanous, in drying orange ochre (6A6, 6A7), when
dried ochre brown (5B6), brown (6C8) or reddish brown (7D8),
finely furfuraceous, smooth at the base. Warts present near
the margin, flat and rounded, concolorous, brown. Stipe 3–4
× 2–3 mm. Basal tomentum and mycelium pale yellow (4A3).
Spores ellipsoid, slightly inequilateral, with two large guttules,
rarely with one to several smaller granules, smooth, hyaline,
10.5–12 × 6–6.5 µm (δm = 11.1–11.7 µm, Wm = 6.1–6.7 µm,
Qm = 1.7–1.9; n = 4). Paraphyses broadly hooked, sometimes
curved, sometimes enlarged at the apices to 2.5–3 µm wide,
without notches, sometimes with a brown matter covering the
apices, when fresh containing small, pale yellow guttules; when
dried heterogeneous, pale yellow. Asci 184 – 237 × 11–11.5
µm. Apothecial section 800 –1200 µm thick. Subhymenium
c. 100–150 µm thick, of dense textura intricata, with tendency to
textura angularis towards the medullary excipulum, visible as a
darker orange brown zone, cells 2.5–6(–15) µm wide, with scattered brown resinous exudates. Medullary excipulum 300–700
µm thick, of textura intricata, hyphae thin-walled to slightly
thick-walled, 3–12 µm wide, very pale yellow to pale brownish
yellow, sometimes with brown resinous exudates at septa. Ectal
excipulum of textura angularis, (85–)100 –120 µm, cells thinwalled, very pale brown, (9–)17– 22(–35) × (6–)11.5–23 µm.
Surface with narrowly to broadly conical warts, 45–65 µm high,
formed by short, fasciculate, hyphoid hairs, of 2–3 cylindrical to
ovoid cells, 7–10 µm wide, not or slightly constricted at septa.
Non-warted areas with single hyphoid hairs, of 2–3 cylindrical
to ovoid cells, (5–)7–12 µm wide, not or slightly constricted at
septa, sometimes with a gelatinous sheath. Resinous exudates
abundant, pale yellowish brown, dissolving into amber drops
in εδZ, turning slightly darker and partly dissolving in KOH.
Basal mycelium of 2–5.5 µm wide, hyaline to very light yellow
hyphae, unchanged in KOH, with refractive, pale yellow drops
on the surface, dissolving in εδZ.
Specimens examined. italy, Cosenza, Calabria, Colamauci, Celico, under
Pseudotsuga menziesii, C. Lavorato, 28 Sept. 2005, Cδ 050928-30 (dupl.
S-F256978). – Spain, Canary Islands, La Palma, under Pinus canariensis,
26 Nov. 2008, J. Fernández Vicente, P. Iglesias, F. Hidalgo, J.R. Undagoitia,
S. Lequerica & R. Martínez (S-F256979); δa Rioja, Clavijo, under Pinus
sylvestris, on the ground, C.M. Pérez del Amo & R. Gil, 3 Jan. 2009, private
herb. CεP 1179, Rε 1095 (dupl. S-F256980); εadrid, Bustarviejo-Canencia,
Puerto de Canencia, 3 Oct. 1979, E. Álvarez (AH44526).
I. Olariaga et al.: A monograph of Otidea
201
Fig. 19 Otidea subformicarum apothecia. a. S-F242696, holotype; b. S-F256980; c. S-F256979; d. Cδ 050928-30. — Photos: b. C. Pérez del Amo; c. J.
Fernández Vicente; d. C. δavorato.
Other specimens examined. Otidea aff. subformicarum — mExiCo, Salazar, Parque Nacional εiguel Hidalgo, Abies forest, 23 Sept. 2007, M. Hernández (FH301035); Veracruz, Cofre de Perote, Camino de los Conejos,
Los Lescados, montane forest of Pinus teocote, P. montezumae, Arbutus
jalapensis, 18 Sept. 2007, M.E. Smith (FH301036).
Notes — Otidea subformicarum is closely related to O. formicarum based on both morphological and molecular characters.
Both have broadly ear- to cup-shaped apothecia and small
spores, and are associated with conifers. Diagnostic features
of O. subformicarum are the orange-brown to reddish brown
apothecial colours, and especially the long and narrow spores
compared to related species. Otidea formicarum has shorter
and comparatively more rounded spores (Fig. 20). All O. subformicarum apothecia were collected under Pinus, Pseudotsuga
O. formicarum
O. nannfeldtii
O. pseudoleporina
O. subformicarum
O. aff. subformicarum
O. sp. ‘b’
6.9
Wm (µm)
6.7
6.5
6.3
6.1
5.9
9.7
10.2
10.7
11.2
11.7
Lm (µm)
Fig. 20 εean spore length and width in collections of species in the O. formicarum clade and O. pseudoleporina, based on 20 spores from each
collection.
menziesii or Abies. The ecology of O. formicarum partly overlaps (see further under that species). Otidea subformicarum
appears to have a southern European distribution, whereas
O. formicarum is known only from Fennoscandia and the Alps.
Otidea subformicarum forms a strongly supported clade in
the ITS-δSU phylogeny (Fig. 2). The four ITS sequences of
O. subformicarum, from Spain and Italy, are nearly identical
(only S-F256979 differs in 1 bp), except for a small variation
in the minisatellites. The ITS nucleotide diversity is 0.71 % in
the four minisatellites unique to O. subformicarum. The five
ITS sequences of O. formicarum, from Finland, Norway and
Sweden, are likewise almost identical (only JS.08.63 differs
in 2 bp). The ITS sequences of O. subformicarum and O. formicarum show many nucleotide differences (40 / 747 bp). Thus
the interspecific ITS nucleotide diversity is much higher (5.35 %)
than the intraspecific diversity (in O. subformicarum 0.07 %
without the minisatellites and in O. formicarum 0.10 %).
Two collections from εexico, FH301035 and FH301036 (here
referred to as O. aff. subformicarum), belong to the O. formicarum clade based on our molecular phylogenetic analyses
(Fig. 1, 2). Although analyses of the combined ITS-δSU dataset
did not resolve the relationships among these three lineages
(Fig. 2) and the δSU phylogeny place FH301035 as a strongly
supported sister taxon to two collections of O. subformicarum,
our three- and four-gene phylogenies (Hansen & Olariaga 2015)
suggest the two European species share a most recent common ancestor, and the εexican collections diverged earlier. The
spore size and shape of the εexican collections conform to
those of O. subformicarum (Fig. 20), but the apothecial colours
differ in these collections. Fresh material has not been available
to us, but the photo of fresh apothecia of FH301035 shows eggyellow apothecia, clearly distinct from O. subformicarum. Photos
of apothecia of FH301036 have a dark purplish brown colour,
Persoonia – Volume 35, 2015
202
which has not been observed in O. subformicarum. Based on
these colour differences and molecular characters, we suggest
that those collections belong to an additional undescribed species, possibly endemic to North America. Further collections are
needed to get insights into the variation and delimitation of this
taxon for its formal description.
Outside the O. formicarum clade, a few species have similar
spores to O. subformicarum. These differ by a combination
of other features: Otidea tuomikoskii by narrowly ear-shaped
apothecia, a receptacle surface with high warts (55–177 µm
high), an ectal excipulum that turns yellow in KOH and typically
orange ochre basal tomentum; and O. nannfeldtii by apothecia
often having yellow tones, sometimes pink stains, and above
all by the shorter spores (δm = 10–10.7 µm).
Otidea unicisa clade
Apothecia with ochre yellow tones. Basal tomentum with ochre
tones. Spores ellipsoid, with warts ± ridges or spinose. Resinous
exudates on the outermost ectal excipulum cells dissolve and
exude bright yellow pigment in KOH.
Species — Otidea kaushalii, O. unicisa, O. yunnanensis.
16. Otidea kaushalii (J. εoravec) K. Hansen & Olariaga, comb.
nov. — εycoBank εB810994; Fig. 22
Basionym. Sowerbyella kaushalii J. εoravec, εycol. Helv. 2: 94. 1986.
≡ Aleurina kaushalii (J. εoravec) W.Y. Zhuang & Korf, εycotaxon 29:
312. 1987.
≡ Otideopsis kaushalii (J. εoravec) J. εoravec, εycol. Helv. 3: 138.
1988.
Fig. 21 Otidea subformicarum (S-F242696, holotype). a. Spores*; b. paraphyses and asci*; c. ectal excipulum in water showing reddish brown resinous exudates†; d. ectal excipulum in εelzer’s reagent showing amber drops †; e. basal mycelium in water †; f. basal mycelium in εelzer’s reagent†. — Scale bars = 10
µm; * = fresh material; † = dried material.
I. Olariaga et al.: A monograph of Otidea
Holotype. india, West Bengal, Darjeeling, Batasi, on soil and decayed
wood in angiosperm forest, alt. 7600 f, 6 Sept. 1979, R. Kaushal, PAN 18169.
Isotypes herb. J. εoravec (CUP 61814, C-F-60847 !).
Apothecia 16–65 mm high, 12–62 mm wide, broadly ear-shaped,
split, stipitate. Hymenium dirty grey to very faintly incarnate,
greyish yellow, orange-ochre (5A6) when dried. Receptacle surface dark orange brown (6E7, 6E8) when dried, densely warty.
Warts conical, acute or blunt, densely gregarious, darker than
the background, dark reddish brown. Stipe 3 – 32 mm long,
3–10 mm wide. Basal tomentum and mycelium ochre (5A4).
Spores ellipsoid to slightly subfusoid, inequilateral, with one or
two large and/or a few small guttules, with thin, often curved,
spines, up to 1(–1.5) µm high, denser at the poles, sometimes
203
united in short ridges, hyaline, 14–17 × 7–9 µm (δm = 14.9–
15.7 µm, Wm = 7.7–7.9 µm, Qm = 1.9–2; n = 2). Paraphyses
curved to hooked, sometimes enlarged at apices, 2.5–5 µm
wide, when dried containing small, yellowish refractive granules. Apothecial section 700–800 µm thick. Asci 188–213 ×
11–13 µm. Medullary excipulum of textura intricata, hyphae
thick-walled, hyaline to very pale yellow. Ectal excipulum of
textura globulosa-angularis, 70–90 µm thick, of 3–4 cell layers, cells thin-walled, pale yellowish brown, 12–46 × 13–33
µm. Surface with conical warts, densely placed, 40–140 µm
high, 50–137 µm wide, formed by globose to elongated cells,
8–11.5 µm broad. Resinous exudates abundant, reddish brown,
amorphous, and/or many yellowish brown, crystal-like, oblate
Fig. 22 Otidea kaushalii. a. Apothecia; b. spores in water †; c. spores in Cotton Blue†; d. paraphyses in KOH †; e. ectal excipulum and warts with abundant
crystal-like, resinous exudates in water†; f. close-up of crystal-like, oblate spheroid exudates on outermost ectal excipulum cells in water † (a–d, insertion on f:
T. δæssøe 6236; e, f: PAN 18169, isotype). — Scale bars = 10 µm; † = dried material. — Photos: a. T. δæssøe.
204
spheroid, 6–8 × 7.5–11 µm, striate bodies, with a constricted
centre, not dissolving in εδZ, dissolving and exuding bright
yellow pigment in KOH. Basal mycelium of 4.5–6 µm wide,
thin-walled to slightly thick-walled, very pale yellow hyphae,
unchanged in KOH, with regularly arranged, spheroid, yellow
resinous exudates, dissolving in εδZ, unchanged in KOH.
Specimens examined. malaySia, Sabah Kinabalu, on rotten wood, 2 εar.
1999, T. Læssøe 6236 (C, dupl. BORH).
Notes — Otidea kaushalii is closely related to O. yunnanensis based on our analyses of the δSU (Fig. 1) and morphology.
Already εoravec (1988) stated this, when he combined O. kaushalii in Otideopsis. The distinctly spiny spore ornamentation in
these two species appears identical; see SEε photographs of
the spores of the holotype of O. kaushalii in εoravec (1986)
Persoonia – Volume 35, 2015
and of O. yunnanensis in δiu & Zhuang (2006). Both species
have a receptacle surface with densely placed, high, dark brown
warts and a lighter coloured hymenium, i.e. greyish yellow,
cream to yellow. Otidea kaushalii is distinguished by the smaller
spores, and possibly by the ectal excipulum of large, yellowish
brown globose to angular cells (Fig. 22e). We report here for
the first time a so far unique type of crystal-like, oblate spheroid, striate exudates on the outermost ectal excipulum cells
of O. kaushalii (Fig. 22e, f) that might be confined to O. kaushalii within Otidea. These are abundant in both the isotype and
the εalaysian material. A third collection of O. kaushalii has
been reported by Zhuang & Korf (1987) from Xizang, China. The
εalaysian and Chinese collections have smaller apothecia with
shorter or almost no stipe (apothecia up to 19 × 18 mm; stipe
3 × 3 mm in the εalaysian material, Fig. 22a) as compared to
Fig. 23 Otidea unicisa. a. Apothecium; b. apothecia†; c. spores in water †; d. spores in Cotton Blue †; e. paraphyses in water †; f. ectal excipulum in water†
(a: JK12082101; b: H7003343; c–f: KH.06.06). — Scale bars = 10 µm; † = dried material. — Photos: a. J. Karakehian.
I. Olariaga et al.: A monograph of Otidea
the holotype (apothecia up to 65 × 62 mm; stipe 32 × 10 mm).
The apothecia of O. yunnanensis have also been described
with varying stipe length (6 –25 × 4–6 mm).
17. Otidea unicisa (Peck) Harmaja, Karstenia 26: 44. 1986. —
Fig. 23
Basionym. Peziza unicisa Peck, Rep. (Annual) New York State εus.
Nat. Hist. 26: 81. 1874.
≡ Sowerbyella unicisa (Peck) J. εoravec, Czech εycol. 47: 266. 1994.
Holotype. USA, New York, δewis County, Croghan, Felt House, ground
in woods, Sept. (NYSf 3283).
Misapplied names
– Otidea grandis sensu Kanouse, εycologia 41: 672. 1949; δiu & Zhuang,
Fung. Diversity 23: 188. 2006.
Apothecia gregarious to caespitose, 12–25 mm high, 12–35 mm
wide, initially broadly ear-shaped, then becoming cup-shaped,
split, stipitate or sessile. Hymenium ochraceous yellow, sometimes with pink tinges, yellowish ochre (5A5, 5B5), orange-ochre
(5A6) when dried. Receptacle surface ochraceous yellow, when
dried dark orange brown (6E7, 6F7), ochre-brown (5A4) towards
the base, warty, occasionally with shallow ribs at the base.
Warts rounded, gregarious, concolorous or slightly darker than
the background, reddish brown. Stipe 4 – 8 × 2–4 mm. Basal
tomentum and mycelium abundant, yellowish ochre (5A4).
Spores ellipsoid to slightly fusoid, inequilateral, with two large
guttules, often with a few smaller guttules, with small, low warts,
often irregular ridges, denser at the poles, hyaline, (13–)14–
15.5(–16.5) × 6.5–8.5 µm (δm = 14.6–15.2 µm, Wm = 7.1–8 µm,
Qm = 1.8–2.1; n = 3). Paraphyses curved to hooked, sometimes
enlarged at apices, 2.5–4 µm wide, sometimes with 1–2 low
notches, when dried containing refractive, pale yellow granules.
Asci 181–197 × 8–11 µm. Apothecial section 750–900 µm thick.
Subhymenium c. 70–120 µm thick, of cylindrical cells, densely
arranged. Medullary excipulum of textura intricata, 500–700 µm
thick, hyphae thin-walled, 3.5–9 µm wide, hyaline to very pale
yellow, without resinous exudates at septa. Ectal excipulum of
textura angularis, 70–95 µm thick, cells thin-walled, hyaline to
light yellow, 11–30 × 6.5–18 µm. Surface with conical warts,
35–85 µm high, formed by fasciculate, short, hyphoid hairs,
of globose to elongated cells, constricted at septa, 6–11 µm
wide. Resinous exudates abundant, dark yellowish to reddish
brown, dissolving in part and converting into reddish particles
in εδZ, turning brighter yellow in KOH. Basal mycelium of
4.5–6 µm wide, thin-walled to slightly thick-walled, hyaline to
very pale yellow hyphae, unchanged in KOH, with regularly
arranged, spheroid, yellow resinous exudates, dissolving in
εδZ, unchanged in KOH.
Specimens examined. USA, εassachusetts, Carlisle, Great Farm, on
woody debris, 8 July 2006, L. Millman, KH.06.06 (FH); εassachusetts, Carlisle, Towle Conservation δand, 15 July 2006, L. Millman (FH301030); εassachusetts, Purgatory Charm, 9 July 2003, Z. Wang, ZW Geo 65-Clark (S);
εichigan, Cut R., εackinac Co., in frondose woods, 10 Aug. 1949, A.H. Smith
33020 (UPS F-629380); εichigan, N of Hessel, εackinac Co., in frondose
woods, 15 Aug. 1949, H. Imshaug 3458 (UPS F-629377); εichigan, δuce
Co., Tahquamenon Falls State Park, 21 Aug. 1951, A.H. Smith 39070 (UPS
F-629382); εichigan, W of Detour on ε134, Chippewa Co., in beech-maple
woods, 14 Aug. 1949, H. Imshaug 3348 (UPS F-629379); New Hampshire,
Shelburne (UPS F-629427); North Carolina, εacon Co., near summit of
Standing Indian εountain, on duff and buried wood of Betula, 1 Aug. 1969,
H.H. Burdsall 2605 (dupl. H7003343); West Virginia, on light loam and leaf
mold, 15 July 1896, L.W. Nuttall, Ellis 868 (UPS F-630023); West Virginia,
εonongahela National Forest, Dolly Sods Wilderness, Wildlife Trail (TR 560),
on soil among leaf litter under Fagus grandifolia, Acer pennsylvanicum, and
Betula sp., 21 Aug. 2012, J. Karakehian, JK12082101 (FH, dupl. S).
Notes — Otidea unicisa is easily recognised by the spore ornamentation of low, delicate warts and short ridges. The dried
specimens are typically bicoloured, with dark brown outside
and orange-ochre hymenium.
205
The spore ornamentation of O. unicisa was for a long time overlooked (Harmaja 1986). The ornamentation is clearly visible at
1 000×, especially at the poles where it is more prominent. We
agree with Harmaja (2009a) that what Kanouse (1949) described under the name O. grandis is O. unicisa. The bicoloured
apothecia with brown outside and orange hymenium, along
with the ornamented spores, and spore sizes in the range of
O. unicisa, support this view. δiu & Zhuang (2006) also gave a
collection with the typical spore ornament of O. unicisa under
the name O. grandis (for SEε of the ornament see their f. 8).
Both studies mention the Boudier plate 328 (n°. 134, 1905; as
O. grandis), which shows bicoloured apothecia. Otidea unicisa
has not been recorded outside Eastern North America, and
in our opinion the Boudier plate 328 shows typical O. bufonia
apothecia. Corroborating this, two Boudier specimens labelled
O. grandis (UPS F-629342 and likely the collection used for
plate 328: PC0093644), studied by us, are in fact O. bufonia.
For further comments on O. grandis see Excluded, dubious
and imperfectly known taxa.
18. Otidea yunnanensis (B. δiu & J.Z. Cao) W.Y. Zhuang &
C.Y. δiu, Fung. Diversity 23: 188. 2006
Basionym. Otideopsis yunnanensis B. δiu & J.Z. Cao, Shanxi Univ. J.,
Nat. Sci. Ed. 4: 70. 1987.
Holotype. China, Yunnan, Dulong River, δang Tuan, on ground in forest,
30 Aug. 1982, D.C. Zhang (HKAS 12150).
Otidea yunnanensis was described when the monotypic genus
Otideopsis was erected. It was separated from Otidea because
of the ornamented spores and paraphyses with fused apical
portion (δiu & Cao 1987). Upon re-examination of the type
material, δiu & Zhuang (2006) discovered that the paraphyses
had free apices. They also concluded that the spores were with
two large and several small guttules, and not multi-guttulate as
originally described by δiu & Cao (1987). Our δSU phylogeny
confirms that O. yunnanensis is deeply nested within Otidea
(Fig. 1). The species is easily recognised within Otidea by the
large spores with fine, curved spines. Different spore sizes have
however, been reported from the holotype: 18–20 × 8–10 µm
(including ornament, δiu & Cao 1987) and 15–21 × 9.7–10.5
µm (excluding ornament, εoravec 1988), and including one
additional collection: 16.5 – 20 × 7.6 –10 µm (δiu & Zhuang
2006). The very wide spore measurements given by εoravec
overlap somewhat with the spore length of the closely related
O. kaushalii (for further comparisons see under O. kaushalii).
Otidea bufonia-onotica clade
Apothecia dark brown or ochre yellow. Basal tomentum with
brown or yellow tones, especially when dried. Spores fusoid or
ellipsoid, smooth. Resinous exudates on the ectal excipulum
converting into reddish particles or melting into amber drops.
Species — Otidea brevispora, O. bufonia, O. fusconigra ad
interim, O. mirabilis, O. onotica, O. purpurea, O. smithii.
19. Otidea brevispora (W.Y. Zhuang) Olariaga & K. Hansen,
comb. & stat. nov. — εycoBank εB808974
Basionym. Otidea onotica var. brevispora W.Y. Zhuang, εycotaxon 94:
368. 2006 ‘2005’.
Holotype. China, Yunnan, Baoshan, 24 July 2003, Z.L. Yang (HKAS
43003) !
Otidea onotica var. brevispora was distinguished from O. onotica based on shorter spores (Zhuang 2006). Our study of the
O. brevispora holotype confirmed the smaller spores (9.5–10.5
× 5.5–6 µm; δm = 9.9 µm, Wm = 5.6 µm, Qm = 1.75). Otherwise
O. brevispora shares all macro- and microscopic features with
206
O. onotica, including the yellow reaction of the basal mycelium
in KOH. Our εδ analyses, with an δSU GenBank sequence of
the Chinese holotype, show O. onotica var. brevispora is a sister
taxon to a clade of O. onotica specimens (Fig. 1). Based on this
and the smaller spores we consider it to be a distinct species.
20. Otidea bufonia (Pers.) Boud., Hist. Classif. Discomyc.
Europe: 52. 1907. — Fig. 4b, c, 5b, 24
Basionym. Peziza bufonia Pers., εycol. Eur. 1: 225. 1822: Fr., Syst.
εycol. 2: 54. 1822.
≡ Geopyxis bufonia (Pers.) Sacc., Syll. Fung. 8: 73. 1889.
Lectotype designated here: FranCE, in sylvula Vincennes, Aug. 1816 (δ
0116690 / 911.81.97, Persoon herbarium) !; εycoBank εBT178089.
= Peziza umbrina Pers., Observ. εycol. 2: 77. 1799.
≡ Scodellina umbrina (Pers.) Gray, Nat. Arr. Brit. Pl. 1: 668. 1821.
Persoonia – Volume 35, 2015
≡ Peziza cochleata var. umbrina (Pers.) Fr., Syst. εycol. 2: 50. 1822:
Fr. loc. cit (‘a umbrina’).
≡ Otidea umbrina (Pers.) Bres., Fungi Trident. Ser. 2, fasc. 11–13: 68.
1898.
Lectotype designated here: Sowerby, Col. Fig. Engl. Fung. 1: t. 5. 1797
(as Peziza cochleata); εycoBank εBT200088.
= Peziza pseudobadia Cooke, εycographia part 4: 176. 1877.
≡ Aleuria pseudobadia (Cooke) Gillet, Champ. France Discomycetes:
38. 1879 (‘pseudo-badia’).
≡ Geopyxis pseudobadia (Cooke) Sacc., Syll. Fung. 8: 69. 1889 (‘pseudobadia’).
Holotype. FranCE, εérignac, sur un mur, 1814 (K(ε) 195314, ex herb.
Cooke).
= Otidea pedunculata Velen., εonogr. Discomyc. Bohemiae 1: 354.
1934.
Lectotype designated here: CzECh rEpubliC, Hrusice, Aug. 1924, J. Velenovský (PRε 147622) !; εycoBank εBT200086.
Fig. 24 Otidea bufonia*. a, b. Apothecia; c. spores; d. paraphyses; e. ectal excipulum; f. basal mycelium (a: KH.09.172; b: JS.08.55; c–f: KH.09.171) — Scale
bars = 10 µm; * = all fresh material. — Photos: b. J. Santos.
I. Olariaga et al.: A monograph of Otidea
Misapplied names
– Otidea grandis sensu Boudier, Icon. εycol. livr. 6: n°. 134, pl. 328. 1905
(preliminary text with ‘circulaires’).
– Peziza cochleata sensu Sowerby, Col. Fig. Engl. Fung.1: t. 5. 1797.
Apothecia gregarious, rarely caespitose, 15–45 mm high, 17–
32 mm wide, initially ear-shaped, then soon expanding and becoming deeply cup-shaped, split, stipitate or sessile. Hymenium
initially orange brown (6C6) sometimes olivaceous brown (4D6),
then dark orange brown (6F6, 7E8), when dried greyish brown
(5E3, 5E4), slightly purple. Receptacle surface dark brown
(6E4–6E7), sometimes pale rusty brown (6D8, 6E8) or purplish
brown (6E3) or with olivaceous tones, slightly hygrophanous,
in drying slightly paler, when dried dark orange brown (6E5,
6F4), warty, seldom slightly wrinkled at the base. Warts conical to flattened, gregarious, dark brown, sometimes distinctly
darker than the background. Stipe 5–14 × 7–10 mm, often hollow and felty inside. Smell weak; taste mild. Basal tomentum
and mycelium abundant, brownish white to light brown (6B3).
Spores narrowly fusoid, rarely ovoid, inequilateral, with two
large guttules, very rarely with a third small guttule, smooth,
hyaline to pale yellowish, (12–)13–16.5(–18) × 6–7.5(–8) µm
(δm = 12.4–16.1 µm, Wm = 6.3–7.3 µm, Qm = 1.9–2.5; n = 8).
Paraphyses hooked, a few curved, of the same width or slightly
enlarged at apices, 3.5–5(–7) µm wide, without notches or
rarely with a notch on the underside, when fresh containing
small, refractive, light brownish yellow guttules; when dried
light brownish yellow. Asci 143 –172 × 10 –12 µm. Apothecial section 1000 –1300 µm thick. Subhymenium c. 70 –100
µm thick, visible as a darker zone, composed of cylindrical
to swollen cells, densely arranged, with scattered brownish
resinous exudates at septa. Medullary excipulum of textura
intricata, 500–700 µm thick, hyphae thick-walled, 3.5–9 µm
wide, hyaline to light brown, some covered with strikingly dark
brown, striate resinous exudates. Big crystal-like aggregates
sometimes present among the hyphae, dissolving in KOH,
visible in εδZ. Ectal excipulum of textura angularis, 80–100
µm thick, cells thin-walled, brownish, 16–31 × 11–25(–30) µm.
Surface with broadly conical warts, 40–80 µm high, formed by
short, fasiculate hyphoid hairs, of 3–4 subglobose to elongated
cells, constricted at septa, 6–11 µm wide, sometimes with a
gelatinous sheath. Resinous exudates abundant, dark brown,
partly dissolving and converting into small reddish particles in
εδZ, partly dissolving in KOH. Basal mycelium of 3.5–4.5 µm
wide, hyaline to brown hyphae, with oily, light brown drops on
the surface, sometimes crystalloid and rod-shaped.
Specimens examined. CzECh rEpubliC, Prague-East district, infra Klokočná, in carpineto, 5 Oct. 1931, J. Velenovský (PRε 150074, syntype of
O. pedunculata). – dEnmark, Fyn, Juelsberg Skov, N of Nyborg, along roadside in Fagus forest, 28 Sept. 1986, D. Boertman (C-F-47790); E Jylland,
δøvenholm Skovene, Eldrup Skov, in grass along road, 23 Sept. 1979, E. Andersen (C-F-86688); E Jylland, Vejle Nørreskov, at roadside on rich soil
under Populus tremula, Alnus, Corylus, Betula, Fraxinus, Fagus, etc., 13
Sept. 1993, J. Vesterholt (C-F-20453); NE Jylland, Rubjerg Knude, under
Abies, 18 Sept. 1989, C. Lange & J. Vesterholt (C-F-25955); NW Jylland,
Nystrup Plantage near Thisted, 1 Sept. 1972, K. Toft (C-F-48018); W Jylland, Bordrup Plantage, in Picea and Pinus plantation along forest road on
sandy ground, 17 Sept. 2011, M. Sasa (C-F-94240); δolland, Favrsted Skov,
among mosses, on calcareous soil, by forest-road under Fagus, 4 Oct. 2007,
K. Hansen & B. Kullman, KH.07.37 (S); Sjælland, Feldskoven near Sorø,
14 Aug. 1974, M. Lange (C-F-47995); N Sjælland, Gribskov, under Fagus
in the forest, 19 Sept. 1971, H. Dissing (C-F-48019); ibid., 8 Oct. 1973, P.M.
Pedersen (C-F-48015); Sjælland, Horserød Hegn, 6 km W of Helsingør, along
roadside, 17 Sept. 1964, H. Dissing (C-F-48000). – Finland, Varsinais-suomi,
δohja, Virkkala, Pähkinäniemi, herb-rich deciduous forest on calcareous, mull
soil, with Corylus avellana, Betula pendula, Populus tremula, Picea abies
and Pinus sylvestris, 28 Aug. 1997, J. Vauras 12503F (TUR-A, dupl. S). –
FranCE, Fontainebleau, on the ground under Quercus and Abies, Oct. 1876,
E. Boudier (PC0093644, as O. grandis); Forêt de Châtellerault (δ 0111782,
Herb. Persoon, as Peziza umbrina); δandes, Contis, under Pinus pinaster
on sandy soil, 6 Dec. 2009, J.L. Teres, KH.09.249 (S); δoire-Atlantique,
207
Bourgneuf-en-Retz, 5 Nov. 2009, G. Moyne, NV 2009.11.01 (S); Picardie,
Compiègne, Oct. 1892, E. Boudier (UPS F-629342, as O. grandis); Saône-etδoire, Robin, on the ground under Corylus and Quercus, J.-P. Dechaume, NV
2008.09.12 (dupl. S); Vendée, Jard-sur-εer, in dune forest with dominance
of Pinus pinaster and Quercus ilex, G. Ouvrard, NV 2006.11.07 (S); Vosges,
Rambervillers, forêt de Saint-Hélène, under Alnus in a peatbog, 5 Oct. 2006,
M. Hurtu, NV 2006.10.12 (S). – italy, Caorle (VE), Brussa, Vallevecchia,
under Pinus pinea close to the sea, 4 Nov. 2001, E. Campo (S-F257089);
Ceva (CN), 9 km from the city on the road to Viola, on the side of a track
under Quercus pubescens, Castanea sativa and Corylus avellanea, 16 Sept.
2005, M. Carbone (S-F257088); Deiva εarina (SP), under Quercus ilex,
5 Nov. 2008, M. Carbone (S-F257090); Piemonte, Vignole Borbera (Aδ), Fraz.
Variano superiore, in soil under Quercus pubescens and Castanea sativa, 19
Oct. 2010, M. Carbone (S-F257087). – Spain, Barcelona, entre Sant Feliu de
Codines y Castellterçol, forest with Quercus ilex and Pinus sylvestris, on calcareous ground, 14 Oct. 1976, C. Montoliu (AH44527); Huesca, Javierregay,
under Quercus rotundifolia and Q. humilis humus, 5 Dec. 2009, F. Prieto &
A. González s.n. (S); Navarre, Arbizu, in mixed forest, 14 Nov. 2009, J. Martín
(ARAN-Fungi A5048005); Navarre, Etxauri mendatea, under Quercus ilex,
2 Dec. 2006, J.M. Lekuona (ARAN-Fungi A5053019); Navarre, δete, under
Quercus faginea and Q. rotundifolia on calcareous ground, 19 Dec. 2009,
J.L. Teres & P.M. Pasaban, KH.09.248 (S); Valladolid, Tudela de Duero,
Puente Hierro, under Quercus ilex, J. Santos (S-F22110). – SwEdEn, Gotland,
near Visby, Värnhem, under Fagus and Quercus robur on rich ground, with
Hepatica nobilis, 22 Sept. 2009, E. Bohus-Jensen, K. Hansen & I. Olariaga,
KH.09.189 (S); Skåne, Degeberga, εörkavad, broadleaf forest, 27 Sept.
2001, S.-Å. Hanson, SÅH 2001-253 (C); Skåne, Helsinborg, Gyhult, broadleaf
forest dominated by Fagus and Quercus, on the ground, 13 Oct. 1994, S.-Å.
Hanson, SÅH 16457 (C); Uppland, Stockholm, Enebyberg, Rinkebyskogen,
on rich soil in deciduous forest, under Corylus, but also Tilia, Quercus and
Betula, 1 Sept. 2008, J. Santos, JS.08.55 (S); ibid., 17 Sept. 2008, JS.08.79
(S); Uppland, Uppsala, Hågadalen-Nåsten Nature Reserve, Predikstolen,
under Quercus robur, Picea abies, Corylus and Salix, on rich bare ground,
17 Sept. 2009, K. Hansen & I. Olariaga, KH.09.171 (S); ibid., KH.09.172 (S);
Uppland, Vaksala, near Törnby, Ekbacken, grassy ground with bare patches
of soil, close to a Quercus robur tree, 13 Sept. 2010, K. Gillen & I. Olariaga,
KH.10.199 (S); Öland, Algustrum, Hönstorp, 500 m SO of the village, grazed
mixed deciduous forest, 19 Sept. 1993, T. Knutsson, TK93-209 (S); Öland,
δånglöt, Åstad, c. 75 m from Björkerumsvägen, c. 550 S of the T-crossing
in Åstad, under Corylus, 6 Sept. 1998, T. Knutsson, TK98-208 (S). – USA,
εichigan, Cheboybogan Co., Colonial Pt, Burt δ., in beech-maple woods,
9 Aug. 1951, A.H. Smith 37560 (UPS F-629510); εichigan, Emmet Co.,
Pellston, in beech-maple woods, 11 Aug. 1951, A.H. Smith 37654 (UPS
F-629511); εinnesota, Itasca State Park, on soil in mesic deciduous woods of
red oak, bur oak, birch, hazelnut, Ostrya, 28 July 2010, R.A. Healy, RH1218
(εIN 933332); εinnesota, δake Alexander SNA, on soil in mixed woods with
canopy or red oak, Ostrya, white birch, 16 Aug. 2011, R.A. Healy, RH1393
(εIN 9333323). – without loCality (δ 0111780, Herb. Persoon, as Peziza
umbrina); (δ 0111781, Herb. Persoon, as Peziza umbrina).
Notes — Otidea bufonia is macroscopically characterised
by the cup-shaped apothecia, with a dark brown outside and a
brown basal tomentum, especially in dried specimens. εicroscopically, the narrowly fusoid spores and the presence of hyphae with striate resinous exudates in the medullary excipulum
are important diagnostic characters. Otidea mirabilis is very
similar to O. bufonia (see under O. mirabilis).
The hymenium colour shows a high variability in O. bufonia (Fig.
24a, b); we have observed orange brown, olivaceous or dark
brown tones within a single collection. Also the colour of the
receptacle surface varies and can be with rusty to purplish
brown or olivaceous tones. If purplish tones are present it may
be confused with O. mirabilis. The pigmented, striate, crystallike exudates on the hyphae of the medullary excipulum are
considered a unique feature for O. bufonia (Korf & Zhuang
1991, Harmaja 2009a, Parslow & Spooner 2013), and have
been used to separate O. bufonia from O. mirabilis (Carbone
et al. 2010, Van Vooren 2010). Our results strengthen this view.
We have observed striate exudates in all the material examined
(Fig. 4b, c). The size and abundance of the striate exudates
are variable, and often restricted to the outermost part of the
medullary excipulum (Fig. 4b). In O. mirabilis we have only
observed biflabellate, crystal-like exudates (Fig. 4d).
208
An exceptional O. bufonia specimen with deviant spore characters was discovered (NV 2009.11.01), in which the spores
were ovoid and considerably shorter than in the rest of the
material. This collection is nested within the O. bufonia clade,
along with typical collections.
Nomenclatural notes — A specimen from the Persoon herbarium (911.81.97, δ0116690) labelled as O. bufonia by Persoon can be considered type material (Harmaja 2009a, b), being
collected prior to the publication of O. bufonia. We designate it
here as the lectotype to stabilise the name. It includes a single
dark brown apothecium, with uniseriate, smooth, subfusoid
spores, 14 –16 × 5.8 –7 µm, curved paraphyses, numerous
dark brown, striate exudates on the hyphae of the medullary
Persoonia – Volume 35, 2015
excipulum, and outermost ectal excipular cells with dark brown
resinous exudates.
Otidea umbrina has been recognised as a separate species
(εedardi 1995, Dissing 2000), based mainly on different hymenium colour. The original description of O. umbrina and the
Sowerby plate to which Persoon (1799) referred, agree with
our concept of O. bufonia. Confirming Harmaja (2009a), three
of the four collections of O. umbrina studied by us and kept in
Persoon´s herbarium in δ belong to O. bufonia. Thus we regard
O. umbrina as a synonym of O. bufonia in accordance with other
authors (Dennis 1978). The name O. bufonia is sanctioned at
specific rank and thus has priority over O. umbrina, which is
sanctioned only at variety rank. The name O. grandis has been
used for O. bufonia collections with bicoloured hymenium, i.e.
Fig. 25 Otidea mirabilis*. a, b. Apothecia; c. spores; d. paraphyses; e. ectal excipulum; f. ectal excipulum wart in εelzer’s reagent, showing reddish reaction
of resinous exudates (a: KH.09.188; b: KH.10.294; c–f: KH.10.308). — Scale bars = 10 µm; * = all fresh material.
I. Olariaga et al.: A monograph of Otidea
orange-brown or olivaceous hymenium in contrast to a dark
brown outside (Boudier 1905, Van Vooren 2010). For the interpretation of O. grandis see Excluded, dubious and imperfectly
known taxa. We synonymise O. pedunculata with O. bufonia
based on the study of two syntypes, and lectotypify it with
PRε 147622. The lectotype shows dark brown, split apothecia, fusoid spores (13.5–15.5 × 5.5–6.5 µm, from 8 spores)
and importantly striate exudates on some of the hyphae of the
medullary excipulum. Harmaja (2009b) studied the holotype of
P. pseudobadia and found it is a later synonym of O. bufonia.
Only one collection of P. pseudobadia could be located in Kew
(K(ε) 195314); it is from εérignac as given in the protologue,
but from a wall, whereas Cooke wrote ‘on the ground’. In any
case, one of the apothecia corresponds well to the illustration
in the protologue, and we consider this the holotype (and the
material studied by Harmaja, although no annotation by him
could be found; pers. comm. B. Aguirre-Hudson).
21. Otidea mirabilis Bolognini & Jamoni in Jamoni, Funghi e
Ambiente 85 –86: 56. 2001. — Fig. 4d, 25
Holotype. italy, Piemonte, Alagna valsesia (VC), Val d’Otro, on soil under
Picea and Larix, alt. c. 1500 m, 3 Sept. 1999, D. Bolognini, GεFN 1951.
Isotype (S-F256929) !
Misapplied names
– Otidea leporina sensu Zhuang pro parte, εycotaxon 96: 367. 2005.
Apothecia gregarious or caespitose, 18–62 mm high, 9–52 mm
wide, initially ear-shaped, short or elongated, soon expanding
and becoming deeply cup-shaped, split, stipitate or sessile.
Hymenium initially reddish ochre (5B7, 6B7) or sometimes olivaceous brown (5F5, 5F6), then dark reddish ochraceous brown
(5D8) to dark purple brown (6F4–6F7), when dried ochraceous
brown (5D6) to purplish brown (6E5, 6E6). Receptacle surface
dark purple brown with bluish lilaceous shades (7F2, 7F3),
fading away with time to ochraceous dark-brown (5D7), then
dark purple brown (6F4–6F7), sometimes with greyish ochre
(4A3, 4B3) patches in unexposed parts, hygrophanous, in drying light purple brown (5D2, 6D2) to ochre brown (5B4), when
dried purple brown (6D4, 6F3, 6F4), warty, often wrinkled at the
base. Warts broadly conical to hemispherical, obtuse, gregarious, dark brown, sometimes distinctly darker than the background. Stipe 7–17 × 3–7 mm. Smell weak, vaguely resin-like;
taste mild, slightly bitter in the end. Basal tomentum and mycelium abundant, whitish to light brown (5A3). Spores narrowly
fusoid, inequilateral, with two large guttules, very rarely with
additional 1–3 small guttules, smooth, hyaline to pale yellowish, (13–)13.5–16(–17) × 6–7(–7.5) µm (δm = 14.1–15.4 µm,
Wm = 6.3–6.9 µm, Qm = 2.1–2.3; n = 9). Paraphyses curved to
hooked, of the same width to slightly enlarged at apices, 2.5–5
µm wide, without or occasionally with a few low notches towards
the apex, when fresh containing small, refractive, light brownish yellow guttules; when dried refractive, hyaline guttules.
Asci 178–204 × 9–11 µm. Apothecial section 1000 –1300 µm
thick. Subhymenium c. 100–150 µm thick, visible as a darker
zone, composed of irregularly, densely arranged, globose cells,
with scattered brownish resinous exudates at septa. Medullary excipulum of textura intricata, 600–900 µm thick, hyphae
4–11(–15) µm wide, sometimes slightly swollen, thin to thickwalled, hyaline to light brown, seldom with brown resinous
exudates at septa or biflabellate exudates, sometimes forming
cross-like aggregates. Ectal excipulum of textura angularis,
80 –100 µm thick, cells thin-walled to slightly thick-walled,
brownish, 8–30 × 8–18 µm. Surface with broadly conical warts,
35–55 µm high, formed by short, fasciculate, hyphoid hairs,
of 2 – 4 subglobose to elongated cells, constricted at septa,
6.5–10(–17) µm wide, sometimes with a gelatinous sheath.
Resinous exudates abundant, dark brown, partially dissolving
209
and turning reddish in εδZ. Basal mycelium of 3–6 µm wide,
light yellow to light brown hyphae, seldom with oily brownish
drops on the surface, with abundant small dark brown resinous
exudates, rounded to irregular.
Specimens examined. dEnmark, Bornholm, Rø Plantage, on needle layer
under Picea, 30 Sept. 2001, C. Lange, KH.01.09 (C). – Finland, Koillismaa,
Kuusamo, Oulanka National Park, Ampumavaara, south of the main road
to δiikanen, 14 Aug. 2010, M. Carbone (S-F257083). – FranCE, Isère, δansen-Vercors, 1300 m, on the ground, on litter, under Larix, 20 Sept. 2008,
J. Cavet, NV 2008.09.14 (dupl. S). – india, Uttarakhand, Kalika, Nainital, on
loose humus soil, 22 Sept. 1973, S. Chandes (UPS F-630072). – italy, Friuli
Venezia Giulia, Tarvisio (UD), Val Saisera-Valbruna, on calcareous soil under
Picea abies, Larix decidua and Fagus sylvatica, 27 Sept. 2010, G. Dose
(S-F257092). – norway, Rana, St. Alteren, 7 km W of εo i Rana, 4 Sept.
1972, H. Dissing (C-F-87187). – SwEdEn, Gotland, Ala, Näsmyr, under Picea
and Pinus sylvestris on calcareous ground, 30 Sept. 2010, E. Bohus-Jensen,
K. Hansen, K. Gillen & I. Olariaga, KH.10.294 (S); Gotland, δjugarn, forest
close to Kaupungs Fridhem, under Picea abies on calcareous ground, under
a cliff, 27 Sept. 2010, K. Hansen, K. Gillen & I. Olariaga, KH.10.279 (S);
Gotland, δojsta hed, Russpark, grazed forest with Pinus sylvestris, on calcareous ground, 2 Oct. 2010, K. Hansen, K. Gillen & I. Olariaga, KH.10.308 (S);
Gotland, near Visby, Rävhagen, under Pinus sylvestris, with small Quercus
robur, Helianthemum sp., Melampyrum pratense, Gallium verum, on sandy
and acidified soil, 22 Sept. 2009, E. Bohus-Jensen, K. Hansen & I. Olariaga,
KH.09.188 (S); Gotland, Tofta, Smågårde naturskog Nature Reserve, Tofta
strand, under Pinus sylvestris on calcareous ground, 28 Sept. 2010, K. Hansen,
K. Gillen & I. Olariaga, KH.10.285 (S); ibid., KH.10.288 (S).
Otidea cf. mirabilis — SwEdEn, Öland, Gräsgård, Solberga, Stora Alvaret,
1300 m S of the village, 17 Oct. 1993, T. Knutsson, TK93-276 (S).
Notes — Otidea mirabilis is characterised by dark brown
apothecia, purple to lilaceous-bluish shades in the receptacle
surface of especially young apothecia, fusoid spores, and when
present, biflabellate, crystal-like exudates in the medullary excipulum. Otidea mirabilis and O. bufonia are strongly supported
as sister species in our multigene phylogeny (Hansen & Olariaga 2015). Thorough morphological comparisons show that
O. bufonia differs from O. mirabilis in lacking lilaceous-bluish
shades, and having brown striate exudates on some hyphae of
the medullary excipulum. Otidea smithii is distinguished from
O. mirabilis by the shorter spores with lower Qm (1.9–2.0).
Otidea mirabilis was described with emphasis on the lilaceous
tones of the outside, in contrast to the paler ochre to olivaceous
hymenium (Jamoni 2001). Part of our material shows this colour pattern, but collections with dark brown hymenium have
also been observed. Some authors have noted the absence
of pigmented resinous exudates in the medullary excipulum of
O. mirabilis (Carbone et al. 2010, Van Vooren 2010). We have,
however, observed brown crystal-like exudates in KH.10.308
and the holotype. They are flabellate and forming cross-like aggregates (Fig. 4d), clearly different from those present in O. bufonia (Fig. 4b, c).
Coniferous forests on calcareous ground are the typical habitat
of O. mirabilis. Two Chinese collections assigned to O. leporina
by Zhuang (2006) and sequenced by δiu & Zhuang (2006) are
resolved in the O. mirabilis clade in our δSU tree (Fig. 1). Based
on morphology we report it also from India (UPS F-630072).
22. Otidea onotica (Pers.) Fuckel, Jahrb. Nassauischen Vereins Naturk. 23–24: 330. 1870. — Fig. 26
Basionym. Peziza onotica Pers., Syn. εeth. Fung.: 637. 1801: Fr., Syst.
εycol. 2: 48. 1822.
≡ Scodellina onotica (Pers.) Gray, Nat. Arr. Brit. Pl. 1: 668. 1821.
≡ Peziza leporina var. onotica (Pers.) P. Karst., Bidrag Kannedom Finlands Natur Folk 19: 41. 1871 (‘P. leporina * P. onotica’).
Lectotype designated here: Sowerby, Col. Fig. Engl. Fung. 1: t. 79. 1797
(as Peziza leporina). Epitype designated here: SwEdEn, Gotland, Ollajvs
Nature Reserve, close to δjugarn, calcareous ground in mossy Picea forest,
27 Sept. 2010, K. Hansen, K. Gillen & I. Olariaga, KH.10.284 (S); εycoBank
εBT178083.
= Peziza rosea Schumach., Enum. Pl. 2: 416. 1803.
210
Misapplied names
– Otidea concinna sensu Boudier, Icon. εycol. livr. 26: n°. 552, pl. 325. 1909
(preliminary text with ‘circulaires’).
– Peziza leporina sensu Sowerby, Col. Fig. Engl. Fung.1: t. 79. 1797.
Apothecia gregarious to caespitose, 25–100 mm high, 14–80
mm wide, initially long and narrowly ear-shaped, then soon
expanding, and becoming deeply cup-shaped, split, stipitate
or sessile. Hymenium initially light yellow (4A3 – 4A5), then
ochraceous yellow (4A6, 4A7, 5A6, 5A7, 5B6, 5B7), in some
parts light orange (5A3, 5A4), often with pink tones or entirely
pinkish (6A4), sometimes with red dots, when dried orangeochre (5A5, 5A6) to reddish ochre (6B7, 6B8). Receptacle
surface ochraceous yellow (4A6, 4A7), slightly hygrophanous,
in drying slightly paler, when bruised sometimes brownish ochre
(5B7) in the margin, when dried brownish ochre with orange
Persoonia – Volume 35, 2015
tinge (5B7, 5B8), slightly warty, sometimes wrinkled at the base
when old. Warts broadly conical, gregarious, concolorous or
sometimes distinctly darker than the background, brown. Stipe
13–27 × 4–14 mm. Smell weak; taste mild. Basal tomentum
and mycelium abundant, whitish to light yellow (4A2, 5A2).
Spores ellipsoid to broadly ellipsoid, inequilateral, with two large
guttules, smooth, hyaline, (11–)12–13.5(–14) × (5.5–)6–7 µm
(δm = 12.1–13.3 µm, Wm = 6.2–6.8 µm, Qm = 1.8–2; n = 10).
Paraphyses curved to hooked, of the same width or slightly
broader at apices, 2.5–4.5 µm wide, without notches or rarely
with a slightly swollen area on the underside, when fresh containing small, refractive, light yellow guttules; when dried hyaline
to pale yellow guttules. Asci 138–233 × 9.5–12 µm. Apothecial
section 850 –1400 µm thick. Subhymenium c. 100 –120 µm
thick, of dense textura intricata, visible as a darker zone, cells
Fig. 26 Otidea onotica*. a, b. Apothecia; c. spores; d. paraphyses; e. ectal excipulum; f. basal mycelium (a: KH.10.284, epitype; b: KH.09.132; c – f:
KH.09.165). — Scale bars = 10 µm; * = all fresh material.
I. Olariaga et al.: A monograph of Otidea
cylindrical to swollen, densely arranged. Medullary excipulum
of textura intricata, 400–600 µm thick, hyphae 5–7(–12) µm
wide, thick-walled, hyaline. Ectal excipulum of textura angularis,
80–110 µm thick, cells 13–55 × 11–28 µm, thin-walled, light
yellow. Surface with conical warts, 85–105 µm high, formed by
short, fasciculate hyphoid hairs, of 2–3 subglobose to elongated
cells, constricted at septa, 11–14 µm wide. Resinous exudates
abundant, yellow, dissolving into amber drops in εδZ. Basal
mycelium of 3.5–6 µm wide, hyaline to very light yellow hyphae,
turning yellow in KOH, with very small, regularly arranged,
spheroid, resinous exudates on the surface, dissolving in εδZ,
partially and more slowly in KOH.
Specimens examined. CzECh rEpubliC, South Bohemia, Netolice, Sept.
1922, Hampl (PRε 148341). – dEnmark, Amager, Kongelunden, soil along
forest path, 21 July 1998, K. Hansen, T. Læssøe & C. Lange, KH.98.107 (C);
εøn, Fanefjord Skov, under conifers on calcareous influenced soil, 30 Sept.
2008, H. Knudsen (C-F-89691); Sjælland, Geelskov, 12 km N of Copenhagen,
under Fagus, 18 Sept. 1963, L. Hansen & A. Kjøller (C-F-47985). – FranCE,
Landes, Contis phare, under Pinus pinaster on sandy soil, 9 Sept. 2009,
J.L. Teres (ARAN-Fungi A8200131C). – italy, Piemonte, Vinadio (CN), San
Bernolfo, Picea abies and Abies alba forest, with presence of Fagus sylvatica,
29 Sept. 2008, M. Carbone (εCVE 23277). – norway, Nord-Trøndelag,
Gjøråsvika, δeksvik, under Picea on rich ground, among mosses, 3 Sept.
2009, K. Hansen & I. Olariaga, KH.09.132 (S); under Corylus and Picea on
rich ground, KH.09.136 (S). – Spain, Gipuzkoa, Aia, Amezketalardiko bidegurutzea, under Abies alba, 3 Oct. 2009, J.L. Teres (ARAN-Fungi A3033701A);
εadrid, Sierra de Guadarrama, deciduous forest, among Quercus pyrenaica
leaves, 4 Oct. 1981, G. Moreno (AH2528); Navarre, Arbizu, mixed forest, 14
Nov. 2009, J. Martín (ARAN-Fungi A5048003); Navarre, Orokieta, δoiandi, in
Picea abies plantation, 10 Oct. 2009, Aranzadi ZE (ARAN-Fungi A50411742). – SwEdEn, Skåne, Vittskövle, Segesholm, Herremöllan, broadleaf forest,
27 Sept. 2001, S.-Å. Hanson, SÅH 2001-266 (C); Uppland, Stockholm,
Norra Järvafältet, Hansta Nature Reserve, on rich ground under Picea abies
with Corylus avellana, 15 Sept. 2009, K. Hansen & I. Olariaga, KH.09.164
(S); ibid., KH.09.165 (S), KH.09.166 (S); Uppland, Uppsala, forest-covered
hill NNE of Naturicum, coniferous forest with Quercus and Corylus, 4 Oct.
2011, R. Sundin, KH.11.108 (S); Uppland, Uppsala, Norra δunsen Nature
Reserve, δunsentorpet, on rich organic soil, under dead standing Picea, 28
Aug. 2008, J. Santos, JS.08.48 (S); Uppland, Uppsala, Vänge, Fiby urskog
Nature Reserve, old Picea forest, in the middle of a path, 17 Sept. 2009,
K. Hansen & I. Olariaga, KH.09.175 (S). – USA, εinnesota, Cedar Creek
ESR, on sandy soil in oak savannah, canopy of pin oak, hazelnut, 18 July
2011, R.A. Healy, RH1121 (εIN 933307); εinnesota, δake Alexander SNA,
on soil, mixed woods, canopy of red oak, paper birch, Ostrya, 29 July 2010,
R.A. Healy, RH1222 (εIN 933311); εinnesota, Wild River State Park εany,
on moist, sandy soil in mesic deciduous woods with oak, 23 July 2010,
R.A. Healy, RH1199 (εIN 933309); Oregon, Benton Co., Philomath, Wood
Creek Road, 18 Nov. 1996, E.T. Peterson (OSC 56759); Washington, δower
Tahoma Creek, εt Rainier National Park, 27 Aug. 1948, A.H. Smith 30764
(UPS F-629424); Washington, Nisqually R., εt Rainier National Park, 30
Aug. 1948, H.A. Imshaug 2116 (UPS F-629438).
Notes — Otidea onotica is one of the most common and wellknown species of the genus. It is characterised by rather large,
ochraceous yellow apothecia, often with a pinkish tinge, dots
or stains in the hymenium, and spores of unusual size within
Otidea. The presence of pinkish tones varies considerably.
Apothecia ranging from entirely pink to completely devoid
of pinkish tones have been observed in the same locality, in
KH.09.132 and KH.09.136, respectively. These two collections
have identical ITS sequences (GenBank accessions Kε010103
and JN942772). A consistent character, here reported for the
first time, is the yellow reaction of the basal mycelium in KOH.
Otidea unicisa resembles O. onotica macroscopically, but is
distinguished by ornamented spores.
Nomenclatural notes — Carbone (2009) revised the nomenclature of O. onotica. He considered the plate by Sowerby (1797,
as Peziza leporina) the only element seen by Persoon, and
thus, the holotype. However, Persoon did indirectly refer to
two elements by giving the habitat in the protologue as beech
forests (faginetis) and citing Sowerby’s plate, which shows an
oak leaf. Thus, the plate is not the holotype; i.e. the one element
on which the author based the name (Art. 9.1 ICN; εcNeill et
211
al. 2012). The use of the term holotype could be corrected to
lectotype according to Art. 9.9, but this article is not applicable
since Carbone’s holotype indication does not fulfil Art. 7.10 of
the ICN (it does not include the phrase ‘designated here’ or
equivalent) (εcNeill et al. 2012). Carbone (2009) selected in
addition an epitype from Persoon´s herbarium. But as an epitype must refer to the type it interprets and there was no validly
selected type in 2009, the epitypification by Carbone is also
not valid. We therefore typify O. onotica here, by designating
as lectotype the Sowerby plate, supported by a newly collected
epitype with multiple gene sequences and colour photographs.
The selection of an epitype from Sweden is justified by the
name being sanctioned (Fries saw and studied living material
as indicated by ‘v. v.’), and our ITS sequences of O. onotica
(4, 5, 6, 7, 8, 9) from Denmark, Italy, Norway and Sweden
(including the epitype) being identical. The δSU sequences of
O. onotica (4, 5) from Denmark and Sweden differ only 2 bp
from the other collections.
23. Otidea purpurea (ε. Zang) Korf & W.Y. Zhuang, εycotaxon
22: 507. 1985
Basionym. Acetabula purpurea ε. Zang, Acta Bot. Yunnan. 1: 101. 1979.
Holotype. China, Tibet, Zayu, on the ground in forest with Pinus yunnanensis, 1 Sept. 1976, M. Zang 670 (HKAS 5670) !
Notes — Otidea purpurea is probably closely related to
O. mirabilis. It is also characterised by dark brown apothecia
with lilaceous tones, but it clearly differs in the smaller spores
(8.8–10 × 4.5–5.2 µm). Two additional taxa, O. subpurpurea
and O. bicolor, should be compared with O. purpurea. Otidea
subpurpurea likewise has violaceous tones on the outside,
but has larger spores and asci (Zhuang & Yang 2008). Otidea
bicolor has dark brown apothecia and small spores (Zhuang
2010), and is probably also closely related or conspecific with
O. purpurea, based on the spore size. We treat here O. purpurea in the O. bufonia clade, because of the dark brown apothecia
and brown basal tomentum.
24. Otidea smithii Kanouse, Pap. εichigan Acad. Sci., Part 1.
24: 28. 1939 ‘1938’ — Fig. 27
Holotype. USA, California, Crescent City, 18 Nov. 1937, A.H. Smith 8843
(εICH 14408) !
Apothecia gregarious or caespitose, 32–70 mm high, 13–40
mm wide, initially narrowly ear-shaped, short or elongated,
finally expanding, at late stages becoming deeply cup-shaped,
split, normally shortly stipitate. Hymenium purple-brown (6D5)
to dark purple-brown (7E5), then ochraceous brown (5D8),
when dried very pale purplish brown (5A2). Receptacle surface
dark purple-brown (7E5), sometimes with lilaceous shades or
lighter ochre-brown (6C5) patches in unexposed parts, slightly
hygrophanous, in drying dark ochraceous brown (6D7), when
dried reddish brown (6E5, 6E6), warty, sometimes wrinkled
at the base. Warts conical, obtuse or acute, gregarious, dark
brown, sometimes slightly darker than the background. Stipe
12–26 × 5–12 mm. Basal tomentum and mycelium abundant,
whitish to brown (5B2, 5C2). Spores ellipsoid, sometimes fusoid, inequilateral, with two large guttules, smooth, hyaline,
12–14(–14.5) × 6–7.5 µm (δm = 12.5–13.6 µm, Wm = 6.4–7.1
µm, Qm = 1.9–2.0; n = 5). Paraphyses curved to hooked, of the
same width to slightly enlarged at apices, 3–5 µm wide, sometimes with 1–3 notches or with a low notch near the apex, when
dried containing refractive, pale yellow guttules. Asci 175–251 ×
9–11 µm. Apothecial section 800–1300 µm thick. Subhymenium
c. 140–180 µm thick, of dense textura intricata, visible as a
darker zone, hyphae intermixed with a few subglobose cells,
with scattered brownish resinous exudates at septa. Medullary excipulum of textura intricata, 700–950 µm thick, hyphae
212
Persoonia – Volume 35, 2015
Fig. 27 Otidea smithii (ECV3345). a. Apothecia; b. spores in water †; c. paraphyses in water †; d. ectal excipulum in water †. — Scale bars = 10 µm; † = dried
material. — Photos: a. E. Vellinga.
Fig. 28 Otidea borealis (S-F242694, holotype). a, b. Apothecia; c. close-up of apothecia showing a warty external receptacle and dentate margin; d. Young
narrowly cup-shaped apothecium. — Photos: ε. Carbone.
213
I. Olariaga et al.: A monograph of Otidea
4–14(–21) µm wide, sometimes slightly swollen, thin-walled to
slightly thick-walled, light brown, seldom with light brown resinous exudates at septa. Ectal excipulum of textura angularis,
70–100 µm thick, cells thin-walled to slightly thick-walled, pale
brown, 13–30 × 8–15 µm. Surface with conical warts, 34–71
(–102) µm high, formed by short, fasciculate, hyphoid hairs,
of 2 – 5 subglobose to elongated cells, constricted at septa,
6–14 µm wide, sometimes with a gelatinous sheath. Resinous
exudates rather abundant on the outside, dark brown, partly
dissolving and turning reddish in εδZ. Basal mycelium of 3–5
µm wide, light to darker brown hyphae, sometimes with pale
brown drops on the surface, with abundant small, rod-shaped
to irregular, brown resinous exudates.
Specimens examined. Canada, British Columbia, Goldstream Park, Victoria, on decayed wood under mature Douglas fir, 21 Sept. 1968, J. Ginns
1212 (UPS F-629486). – USA, California, Alameda Co., Berkeley, δe Conte
Avenue, in lawn under Betula and Cedrus, 25 Oct. 2005, E. Vellinga,
ECV3345 (S); California, Del Norte Co., California State Park, δake Earl
Wildlife Area, access by Sand Hill Road, with Abies grandis, Picea sitchensis, Polystichum munitum, Rubus spectabilis, Vaccinum ovatum, 15 Dec.
1997, M. Madsen & R. Davis (OSC 56823); Oregon, Benton Co., Philomath,
Woods Creek Road, 15 Nov. 1997, E.T. Peterson (OSC 56811); Washington,
Pierce Co., εt Rainier National Park, δower Tahoma Creek, 30 Oct. 1996,
E.T. Peterson (OSC 56753); ibid., 18 Oct. 1997 (OSC 56799); Washington,
δewis Co., Gifford-Pinchot National Forest, Camp Creek Falls Trail, on litter
with Acer circinatum, A. macrophyllum, Tsuga heterophylla, Berberis nervosa,
Polystichum munitum, Linnaea borealis, Rubus sp., Goodyera oblongifolia,
6 Nov. 1997, E. Hathaway & E. Millian (OSC 56830).
Notes — Otidea smithii is closely related to O. bufonia and
O. mirabilis, as shown by morphological and molecular characters. These three species share dark brown apothecia, brown
basal tomentum and dark brown resinous exudates in the ectal
excipulum. Otidea smithii is distinguished by shorter spores with
a lower Qm value, and typically narrower, ear-shaped apothecia.
The unique, biflabellate or striate exudates in the medullary
excipulum, as seen in O. bufonia and O. mirabilis (Fig. 4c, d),
respectively, are not present in O. smithii. Only pigmented resinous exudates at the hyphal septa in the medullary excipulum
are rarely found in O. smithii (as in Fig. 4a). Otidea smithii is so
far only known from Western North America (Kanouse 1949,
Peterson 1998). Our ITS sequence of O. smithii (JN942771)
is identical to the ITS sequence of the holotype (AF072065).
Otidea concinna clade
Apothecia cup-shaped, split or sometimes entire, often with bright
yellow tones. Spores small, 10–12.5 µm long, smooth. Paraphyses typically straight and claviform at apices. Basal mycelium with scattered reddish or yellow resinous exudates.
Species — Otidea borealis, O. caeruleopruinosa, O. concinna,
O. flavidobrunneola, O. lactea, O. minor, O. oregonensis, O. phlebophora, O. rainierensis, O. sinensis, O. tianshuiensis, O. sp. ‘a’.
brown (5D7) in the margin, hygrophanous, when dried reddish
brown (6D5, 6D6), warty, seldom slightly wrinkled at the base.
εargin sometimes finely dentate and darker. Warts broadly
conical, gregarious, light to dark brown. Stipe well developed,
3–12 × 2–6 mm. Basal tomentum and mycelium white to very
pale yellow (3A2), when dried very pale yellow (3A2). Spores
broadly ellipsoid, symmetrical, with two large guttules, smooth,
hyaline, (10–)10.5–11.5 × 6–7 µm (δm = 10.5 µm, Wm = 6.5
µm, Qm = 1.7; n = 1). Paraphyses straight to curved, very few
hooked, 2–2.5 µm wide, at apices up to 3–4.5(–5) µm wide,
without notches, terminal segment 33–47 µm long, narrowly
claviform to claviform, when dried containing slightly refractive,
hyaline granules. Asci 167–198 × 9–10.5 µm. Apothecial section 1 000 µm thick. Subhymenium c. 100 µm thick, of dense
textura intricata, cells 2–4 µm thick. Medullary excipulum 600
µm thick, of textura intricata, hyphae thin-walled to slightly
thick-walled, 4–7 µm wide, hyaline, without resinous exudates.
Ectal excipulum of textura prismatica, 80 µm thick, cells thinwalled, hyaline, unchanged to very pale yellow in KOH, 10–21
× 8.5–11 µm. Surface with broadly conical warts, 63–90 µm
high, formed by fasciculate, short, hyphoid hairs, of 3–6 subglobose to ovoid cells, constricted at septa, 6–12 µm wide.
Non-warted parts with single, 2–5-celled hyphoid hairs, with
cylindrical to claviform upper cell, 45–73 × 8–10 µm. Resinous
exudates golden brown, partly dissolving in εδZ (amber drops
not observed), bright yellow and partly dissolving in KOH. Basal
mycelium of 3–4.5 µm wide, very pale yellow hyphae, bright
yellow in KOH, with spheroid to rod-shaped, reddish yellow
resinous exudates, dissolving in εδZ, partially dissolving and
turning bright yellow in KOH.
Other specimens examined. Otidea sp. ‘a’ — SwEdEn, Lappland, Arvidsjaurs, δillån Allmänningreservat, 8 km SV Järvträsk, herb-rich Picea forest
on calcareous ground, 12 Sept. 2009, M. Karström, εK0942 (S); ibid., 10
Sept. 2010, εK1081 (S).
Notes — Within the O. concinna clade, O. borealis is distinguished by broadly ear-shaped apothecia, with a pale yellowish
ochre hymenium and a darker ochre yellow outer surface with
small brown warts. εicroscopically, the spores are proportionally broader than in closely related species, and the mycelium
at the base of the apothecia turns bright yellow in KOH (tested
in dried specimen only; Fig. 30f).
At least four species of the O. concinna clade have bright
yellow outer apothecial colours, similar to O. borealis. Otidea
concinna appears morphologically most similar, but besides the
citrine-yellow outer receptacle, it differs in the narrower spores
(Fig. 29) with higher Qm value. Otidea phlebophora and O. minor
are distinguished by the often entire apothecia with ribs or
anastomosing veins at the base and distinctly narrower spores.
Otidea oregonensis differs from O. borealis in typically having
O. borealis
O. concinna
7
25. Otidea borealis ε. Carbone, Olariaga, K. Hansen & Van
Vooren, sp. nov. — εycoBank εB809250; ITS barcode
GenBank Kε010023; Fig. 28, 30
Holotype. Finland, Koillismaa, Kuusamo, Juuma, Jäkälävuoma, western
part of Jäkälävuoma gorges, many apothecia in moist soil among mosses
at brookside, under Picea abies, 16 Aug. 2010, M. Carbone (S-F242694).
Isotype (TUR-A 198578).
O. kauffmanii type
6.6
O. minor
6.4
Wm (µm)
Etymology. Referring to a supposedly boreal distribution.
6.8
O. oregonensis
6.2
O. phlebophora
6
O. rainierensis type
5.8
O. sp. ‘a’
5.6
5.4
Apothecia gregarious to caespitose, 12–22 mm high, 8–15 mm
wide, initially narrowly ear-shaped, upper margin subacute to
rounded, then broadly ear-shaped, split, stipitate. Hymenium
pale yellowish ochre (4A2, 5A2), when dried greyish ochre
(4A2, 5A2). Receptacle surface ochraceous yellow (4A5,
5A5), yellowish brown (5B5, 5B6) in some parts, sometimes
5.2
5
10.1
10.6
11.1
11.6
12.1
12.6
Lm (µm)
Fig. 29 εean spore length and width in collections of species in the core of
the O. concinna clade, based on 20 spores from each collection.
214
Persoonia – Volume 35, 2015
Fig. 30 Otidea borealis (S-F242694, holotype)†. a. Spores in water; b. paraphyses and asci in KOH; c. ectal excipulum in water, with resinous exudates on the
outside; d. ectal excipulum turning bright yellow in KOH, resinous exudates partly dissolving; e. basal mycelium in water; f. basal mycelium in KOH. — Scale
bars = 10 µm; † = all dried material.
obconical apothecia, sometimes with strongly rugulose base,
citrine-yellow colour in the outer receptacle when young, narrower spores (Fig. 29), and the mycelium at the base of the
apothecia does not turn yellow in KOH. The North American
O. rainierensis (= O. kauffmanii) appears to have some yellowish
apothecial pigment (see further under O. rainierensis). Nevertheless, the spores of O. rainierensis are longer than the spores
of O. borealis (Fig. 29). Furthermore, our ITS-δSU phylogenies
(Fig. 3) suggest that these species are distinct from O. borealis.
Two Swedish collections, εK0942 and εK1081, constitute another undescribed species (Otidea sp. ‘a’) nested within the
O. concinna group. It resembles O. borealis in spore shape
(only slightly longer in Otidea sp. ‘a’: 11–12(–13) × 6–7 µm;
Lm = 11.2–11.6 µm, Wm = 6.6–6.8 µm, Qm = 1.6–1.7; n = 2) and
yellow reaction of the basal mycelium in KOH. The two Otidea
sp. ‘a’ collections differ from O. borealis in the brownish ochre
receptacle surface, without yellow tones, and ochre to pinkish
ochre hymenium. Also our analyses of the ITS-δSU regions
support O. sp. ‘a’ and O. borealis as distinct species (Fig. 3).
Since only dried material has been available to us, we postpone
a formal description of O. sp. ‘a’ until fresh and more abundant
material, with good colour photographs, becomes available.
I. Olariaga et al.: A monograph of Otidea
26. Otidea caeruleopruinosa Harmaja, Karstenia 48: 37. 2009
— Fig. 31
Holotype. Finland, Varsinais-Suomi, δohja, Jalassaari, 20 Sept. 1978,
H. Harmaja (H6010805) !
Apothecia gregarious to caespitose, 31– 60 mm high, 17–50
mm wide, initially ear-shaped, upper margin rounded, then cupshaped, split, margin sometimes lobulate, stipitate or sessile.
Hymenium yellowish brown (5C6), when dried yellowish brown
with a faint olivaceous tint (4D6) or orange-ochre (6B6, 6C6).
Receptacle surface upper half concolorous with hymenium or
slightly duller (5C5, 5C6), lower half more whitish (pubescent
pruinose), slightly hygrophanous, when dried concolorous with
hymenium towards the margin, below cream, or pale reddish
brown (6C6), furfuraceous to very finely warty, seldom slightly
ribbed at the base. Warts broadly conical, gregarious, paler
or bluish. Stipe not well developed, rooting, up to 11 × 5 mm.
Basal tomentum and mycelium abundant, when fresh ochraceous white, when dried light ochre (5A2). Spores narrowly
ellipsoid to ellipsoid, often inequilateral, with two large guttules,
smooth, hyaline, (10.5–)11–12.5 × 5.5–6.5 µm (δm = 11.2–11.8
µm, Wm = 5.9–6.1 µm, Qm = 1.9; n = 3). Paraphyses straight
or curved, or broadly hooked, of the same width or broader at
apices, 2–4(–5) µm wide, without notches, terminal segment
26–78 µm long, when fresh containing few, small, weakly refractive, pale guttules; when dried refractive, hyaline granules. Asci
144–200 × 9–11 µm. Apothecial section 700–950 µm thick.
Subhymenium c. 70–80 µm thick, of 2.5–5 µm wide cylindrical
cells, with scattered swollen, 12–18 µm wide cells, densely arranged. Medullary excipulum of textura intricata, 550–700 µm
thick, hyphae thin to thick-walled, 3–9 µm wide, intercalated with
215
swollen cells, 12–18 µm diam, hyaline, without resinous exudates. Ectal excipulum of textura prismatica-angularis, 100–120
µm, cells thin-walled, pale brown, sometimes yellow in KOH,
10–32 × 10–22 µm. Surface with conical to broadly conical
warts, 60–80 µm high, formed by short, fasciculate hyphoid
hairs, of 2 – 4 ovoid to cylindrical cells, 6 –10 µm wide, not
constricted at septa. Non-warted parts with single 2–7-celled
hyphoid hairs, of globose to cylindrical cells, 20–33 × 9–17
µm. Resinous exudates abundant, yellow to reddish brown,
dissolving in εδZ, bright yellow and partly dissolving in KOH
and the outer excipulum turning bright yellow. Basal mycelium
of 3.5–6(–8) µm wide hyphae, with spheroid to rod-shaped,
reddish yellow resinous exudates, dissolving in εδZ, more
slowly and partially dissolving in KOH, turning yellow.
Specimens examined. Spain, Girona, Ripollés, Setcases, under Corylus
avellana, Betula verrucosa and Buxus sempervirens, on calcareous soil, 26
Aug. 2010, M. Tabarés & S. Santamaría, εT 10082601 (dupl. S). – SwEdEn,
Södermanland, Nynäshamn, Herrhamra, on soil under Fagus, in narrow forest area along the road, 18 Sept. 2013, I.-L. Walter, KH.13.48 (S); Uppland,
Älvkarleby, V from the train station, 24 Sept. 1950, G. Fåhraeus & G. Stenlid
(UPS F-146664).
Notes — The main diagnostic characters of O. caeruleopruinosa are cup-shaped, split apothecia, whitish to pale cream hymenium, grey outside and rather small spores. Harmaja (2009a)
described O. caeruleopruinosa with strong emphasis on the
bluish shades of the warts on the apothecial outside, a character that has not been observed in our Swedish, or the Iberian
material (Van Vooren et al. 2011). A third gathering from the
Iberian site does not show bluish tones either (ε. Tabarés, pers.
comm.).
Fig. 31 Otidea caeruleopruinosa. a. Apothecia; b. apothecium showing basal tomentum†; c. spores in water †; d. paraphyses in water †; e. ectal excipulum in
water † (a: εT 11080205; b–e: H6010805, holotype). — Scale bars = 10 µm; † = dried material. — Photos: a. ε. Tabarés; b. J. Kearey.
216
Otidea caeruleopruinosa belongs to the O. concinna clade, and
it shares several characters with the other members of the
clade, such as cup-shaped apothecia, small spores, and an
ectal excipulum of textura prismatica to textura angularis. It is
morphologically most similar to O. flavidobrunneola (see Notes
under that species).
Persoonia – Volume 35, 2015
holm, Naturhistoriska riksmuseet, in front of the Botany building, under a
big Quercus robur tree, by a row of Populus, 20 Sept. 2009, K. Hansen &
I. Olariaga, KH.09.183 (S); εycoBank εBT178084.
= Helvella pyxidata Schaeff., Fung. Bavar. Palat. Nasc. 4: 111. 1774
(‘Elvela’).
≡ Peziza marsupium var. pyxidata (Schaeff.) Pers., Syn. εeth. Fung. 2:
640. 1801 (‘ß pyxidata’).
≡ Scodellina pyxidata (Schaeff.) Gray, Nat. Arr. Brit. Pl. 1: 669. 1821.
27. Otidea concinna (Pers.) Sacc., Syll. Fung. 8: 96. 1889 —
Fig. 32
Misapplied names
– Flavoscypha cantharella sensu Harmaja, Karstenia 14: 107. 1974.
Basionym. Peziza concinna Pers., εycol. Eur. 1: 221. 1822: Fr., Syst.
εycol. 2: 49. 1822.
≡ Helvella scutellata Schaeff., Fung. Bavar. Palat. Nasc. 4: 101. 1774
(‘Elvela’).
Lectotype designated here: Schaeffer, Fung. Bavar. Palat. Nasc. 2: t. 150,
f. 1. 1763 (‘Elvela tertia’). Epitype designated here: SwEdEn, Uppland, Stock-
Apothecia gregarious to caespitose, 17–71 mm high, 12–66 mm
wide, initially broadly ear-shaped, then soon expanding and becoming deeply cup-shaped, split, often broader above, very
rarely entire when young, stipitate or broadly sessile. Hymenium
pale yellowish ochre (3A3, 4A5) to pale greyish ochre (5A2),
Fig. 32 Otidea concinna*. a, b. Apothecia; c. spores; d. paraphyses; e. ectal excipulum; f. basal mycelium (a, c, e, f: KH.09.183, epitype; b: KH.09.217;
d: KH.09.176). — Scale bars = 10 µm; * = all fresh material.
I. Olariaga et al.: A monograph of Otidea
sometimes with rose stains or spots, when dried light yellowish
grey (2A2) to very pale yellowish ochre (3A2, 3A3). Receptacle
surface bright citrine yellow (3A5–3A7), sometimes with a greenish hue (2A6), slightly hygrophanous, in drying slightly paler,
fading to brownish ochre (4B5) in age, when dried yellow (3A7),
furfuraceous, sometimes warty in young apothecia, sometimes
shallowly wrinkled at the base. Warts flat to broadly conical,
leaving a reticulum among them, concolorous. Stipe not well
developed, rooting, 7–8 × 4–5 mm. Smell sweet, fruity, weak;
taste mild. Basal tomentum and mycelium abundant, white to
very pale cream (4A2), sometimes with very pale purplish tones,
when dried white to very pale yellow (4A2). Spores narrowly ellipsoid to ellipsoid, inequilateral, with two large guttules, smooth,
hyaline, (10–)10.5–12 × (5–)5.5–6.5 µm (δm = 10.8–11.2 µm,
Wm = 5.6–6 µm, Qm = 1.8–2; n = 8). Paraphyses straight to
bent, seldom broadly hooked, at apices claviform to abruptly
capitate, 2.5–6(–8.5) µm wide, without notches, terminal segment 19–66 µm long, when fresh content hyaline homogenous
or of small, refractive, hyaline to pale yellow granules; when
dried refractive, hyaline granules. Asci 176 –196 × 10 –11.5
µm. Apothecial section 1200 –2100 µm thick. Subhymenium
c. 60–90 µm thick, composed of cylindrical cells, 2–4 µm wide,
with scattered swollen cells, 9–13 µm wide, densely arranged.
Medullary excipulum 600 –1000(–1500) µm thick, of textura
intricata, hyphae cylindrical to slightly swollen, 3–11(–15) µm
wide, thin to thick-walled, hyaline, without resinous exudates.
Ectal excipulum of textura prismatica-angularis, 80–100 µm
thick, cells thin-walled, hyaline to very pale yellow, sometimes
light reddish in KOH, 12 – 35 × 6 – 22 µm. Surface with low
broadly conical warts, 60–70 µm high, formed by fasciculate,
parallel, short, hyphoid hairs, of 4–6 ovoid cells, constricted
at septa, 5–9.5 µm wide. Non-warted parts with single, 2–6celled, hyphoid hairs, uppermost cell narrowly claviform to
subcapitate, 20–45 × 4.5–7 µm, sometimes with a gelatinous
sheath. Resinous exudates abundant, yellow or reddish, dissolving into amber drops in εδZ, turning bright yellow in KOH.
Basal mycelium of cylindrical to slightly swollen, hyaline to very
pale yellow hyphae, 3–4.5(–8.5) µm wide, yellow in KOH, with
scattered, spheroid to rod-shaped, yellow or reddish resinous
exudates, dissolving in εδZ, partially in KOH.
Specimens examined. dEnmark, NE Sjælland, Jægersborg Dyrehave,
8 Sept. 1979, H. Knudsen (C-F-87186); NW Jylland, Nystrup Plantage, ‘Kridtstien’, on calcareous soil with Abies, 14 Sept. 1985, T. Læssøe (C-F-81617).
– EStonia, Saaremaa, Abruka, ad terram, 16 Sept. 1966, K. Kalamees (UPS
F-629562). – Finland, Varsinais-Suomi, Turku, Ispoinen Katariinanlaakso,
W part of the reserve, in rich, essentially deciduous woods with Corylus, Quercus, Tilia, etc. on bare clayey mull soil, 22 Aug. 1977, H. Harmaja (S-F249360,
ex-H6015773). – FranCE, Saône-et-δoire, δa Grande-Verrière, Senavelle, on
the ground under Pseudotsuga menziesii, 12 Sept. 2008, J.-P. Dechaume, NV
2008.09.13 (dupl. S). – Spain, Huesca, Javierregay, humus of Quercus rotundifolia and Q. humilis, 5 Dec. 2009, F. Prieto & A. González, KH.09.250 (S);
Huesca, Sallent de Gállego, δanuza, 14 Oct. 2006, I. Olariaga (BIO-Fungi
13002); Huesca, Yebra de Basa, close to Sta Orosia chapel, in Fagus sylvatica forest, 11 Oct. 2009, P. Siljeström (ARAN-Fungi A8700091). – SwEdEn,
Gotland, Ala, Stenstugu, near a big Quercus robur tree in meadow, on rich
ground, among grass, 26 Sept. 2009, E. Bohus-Jensen, K. Hansen & I. Olariaga, KH.09.217 (S); Gotland, δojsta, δojsta slot, under Corylus on a chalkrich slope, 19 Sept. 2000, T. Knutsson, TK2000-078 (S); Närke, Havsta, Bruntorpskärret, under conifers, 10 Aug. 2008, B. Wasstorp, JS.08.59 (S); Skåne,
Degeberga, Segesholm, on a steep slope on bare ground, under Fagus and
Ulmus, 23 Sept. 2010, K. Hansen, K. Gillen, I. Olariaga, KH.10.256 (S);
Skåne, Fjälkstad, Balsberget, on rich ground among leaf litter under Fagus,
20 Sept. 2010, K. Hansen, K. Gillen & I. Olariaga, KH.10.212 (S); Skåne,
Kristianstad, N. δingenäset, Storskogen, on ground in broadleaf forest,
18 Sept. 1996, S.-Å. Hanson s.n. (C); Uppland, Uppsala, Carolinaparken,
under Corylus avellana, 7 Sept. 2010, S. Ryman, K. Hansen, K. Gillen &
I. Olariaga, KH.10.180 (S); Uppland, Uppsala, Hågadalen-Nåsten Nature
Reserve, Predikstolen, under Quercus robur, Ulmus glabra, Sorbus aucuparia, with Hepatica nobilis, Geum urbanum, Convallaria majalis, on rich
ground, 19 Sept. 2009, K. Hansen & I. Olariaga, KH.09.176 (S); ibid., 6 Oct.
2010, KH.10.182 (S).
217
Notes — Otidea concinna is characterised by cup-shaped,
split apothecia, citrine yellow receptacle surface, along with paraphyses predominantly straight and claviform at apices. Otidea
minor and O. phlebophora share with O. concinna a yellow
receptacle surface, but they have anastomosing ribs at the
apothecial base and narrower spores (Fig. 29). Otidea oregonensis is distinguished from O. concinna by the often obconical
apothecia, sometimes with blunt ribs at the base.
The bright yellow receptacle surface is one of the main diagnostic characters of O. concinna, but we have observed the yellow
colour can disappear with age, and the outer surface becomes
brownish ochre (Fig. 32b) thus making it difficult to recognise
the species. Otidea rainierensis is a North American taxon that
shows an apothecial shape similar to O. concinna. Nevertheless, bright citrine yellow tones are absent in O. rainierensis,
and our molecular data support it as distinct from O. concinna
(Fig. 1, 3). For further comments see under O. rainierensis.
Nomenclatural notes — No authentic material has been
located in Persoon’s herbarium in δ. We therefore select one
element in Schaeffer’s plate (t. 150, f. 1) as the lectotype, because it most closely resemble Persoon’s description, showing
two or more large, convolute, caespitose, sessile, cup-shaped
apothecia with a citrine (although light) outer surface. Persoon (1822) collected the species in ‘sylvula Vincennes prope
Parisios’ and Schaeffer’s plate is from Bavaria, Germany. Fries
sanctioned the name, but did not see material (‘v. ic.’, seen from
icones). Nevertheless, we select a newly collected epitype from
Sweden (Fig. 32a, c, e, f), associated with Quercus (the habitat
described by Persoon) and backed by sequences of multiple
gene regions (ITS, δSU, PRB1, RPB2 and EF1) of O. concinna
collections from southern Europe (Spain) and from the epitype
being identical (Hansen & Olariaga 2015), acknowledging the
species is widely distributed in Europe.
28. Otidea flavidobrunneola Harmaja, Karstenia 48: 38. 2009
— Fig. 33
Holotype. Finland, Varsinais-Suomi, δohja, Jalassaari, Ahtiala, Alho,
Ahtiala Nature Reserve, rich, predominantly deciduous (Quercus, Corylus
etc.) woods on calcareous soils, 20 Sept. 1978, H. Harmaja (H6010806) !
Apothecia gregarious to caespitose, 15–70 mm high, 8–50 mm
wide, initially narrowly to broadly ear-shaped, then cup-shaped,
split, margin sometimes lobulate, shortly stipitate or sessile.
Hymenium cream or pale yellowish (4A2, 4A3), when dried
orange-ochre (5B6). Receptacle surface pale brownish ochre
(5A3, 5A4), slightly hygrophanous, when dried dark reddish
brown (5E5), furfuraceous to very finely warty, seldom shallowly
wrinkled at the base. Warts flattened, gregarious, concolorous. Stipe not well developed. Basal tomentum and mycelium
abundant, white to pale cream (4A2) when fresh, ochre (5A4)
to orange-ochre (6A4) when dried. Spores narrowly ellipsoid
to ellipsoid, inequilateral, with two large guttules, very rarely
with a third small guttule, smooth, hyaline, 9.5–11 × (4.5–)5–6
µm (δm = 10–10.6 µm, Wm = 5.1–5.5 µm, Qm = 1.9–2; n = 5).
Paraphyses curved, a few straight or broadly hooked, of the
same width or slightly broader at apices, 3–4 µm wide, without
notches or slightly sinuous underside, uppermost cell 40–76 µm
long, when fresh containing yellow guttules; when dried hyaline
guttules or granules. Asci 127–170 × 9 –10 µm. Apothecial
section 900–1900 µm thick. Subhymenium c. 100 µm thick, of
dense textura intricata, visible as a yellowish zone, cells cylindrical to slightly swollen. Medullary excipulum 600–1300 µm
thick, of textura intricata, hyphae 5–13 µm wide, thick-walled,
sometimes with a thinner outer wall, hyaline, without resinous
exudates. Ectal excipulum of textura prismatica, sometimes of
218
Persoonia – Volume 35, 2015
Fig. 33 Otidea flavidobrunneola. a. Apothecia; b. apothecium showing basal tomentum†; c. spores in water †; d. paraphyses in water †; e. ectal excipulum in
water †; f. ectal excipulum in KOH†; g. basal mycelium in water † (a, c: KH.09.153; b, d–g: H6010806, holotype). — Scale bars = 10 µm; † = dried material. —
Photos: b. J. Kearey.
a textura angularis, 100–120 µm, cells thin-walled, yellowish,
distinctly citrine yellow in KOH, 18–38 × 11–13 µm. Surface with
broadly conical warts, 40–70 µm high, formed by fasciculate,
short hyphoid hairs, of 3–4 ovoid to elongated cells, constricted
at septa, 5–9 µm wide, sometimes with a gelatinous sheath.
Non-warted parts with 2–3-celled hyphoid hairs, with claviform
uppermost cell, more rarely cylindrical, 33 – 55 × 8 –13 µm.
Resinous exudates abundant, yellowish brown, partially dissolving and turning slightly reddish in εδZ, partly dissolving,
bright yellow and appearing gelatinous in KOH. Basal mycelium of 4–6 µm wide, hyaline to very pale yellow hyphae, with
scattered swollen septa, with rounded to rod-shaped, yellow
resinous exudates, slowly dissolving in KOH, quickly in εδZ.
Specimens examined. Finland, Uusimaa, Nurmijärvi, parish centre, under
Quercus, 16 Sept. 1987, P. Askola 2220 (H); Uusimaa, Nurmijärvi, parish
centre, clayey soil under Quercus, 11 Aug. 1988, P. Askola 2334 (H); Uusimaa, Nurmijärvi, parish centre, the margin of the park by the vicarage, under
Quercus, 18 Aug. 1988, P. Askola 2360 (H). – norway, Nord-Trøndelag, δeksvik, Gjøråsvika, on rich ground, on slope under Corylus and Picea, 3 Sept.
2009, K. Hansen & I. Olariaga, KH.09.153 (S). – SwEdEn, Uppland, Bondkyrka,
Vårdsätra naturpark, 17 Aug. 1927, H. Svensson (UPS F-146554); Uppland,
Uppsala, in front of the prison, on bare ground under deciduous trees, 14 Sept.
1938, R. Gustafsson (UPS F-146718).
Notes — Otidea flavidobrunneola is macroscopically characterised by cup-shaped, split apothecia, especially with age very
pale yellowish brown outside and cream-white to pale yellow
hymenium. εicroscopically, the small spores and the yellow
I. Olariaga et al.: A monograph of Otidea
ectal excipulum turning brighter yellow in KOH are diagnostic.
The apothecia and basal tomentum become characteristically
darker upon drying: the outside turns dark brown, the hymenium and the basal tomentum orange ochre (Fig. 33b). Otidea
caeruleopruinosa and O. flavidobrunneola are morphologically
very similar, but differ in the spore sizes (δm = 11.2–11.8 µm
vs 10.1–10.6 µm) and colours of the basal tomentum in dried
specimens. According to Harmaja (2009a), O. flavidobrunneola
is reminiscent of O. bufonia, but the latter has a darker brown
basal tomentum and larger, narrowly fusoid spores. We report
here the first finds outside Finland.
29. Otidea lactea J.Z. Cao & δ. Fan in Cao et al., εycologia
82: 735. 1990
Holotype. China, Heilongjiang province, Yichun City, on ground (rotten
wood?) under broadleaf trees, 6 Sept. 1987, J.Z. Cao (HεAS 61359, exεHSU 1803).
Notes — We were not able to get the material of O. lactea
on loan, but it appears to be a distinct species based on the
entire, cup-shaped, cream white apothecia and paraphyses
with broadly clavate to subglobose apices (Cao et al. 1990).
Also, in our δSU phylogeny (including a GenBank sequence of
the holotype, DQ443447 from δiu & Zhuang (2006)), O. lactea
forms a distinct sister lineage (εδ 71 %, PP 99 %) to a clade of
seven collections of O. minor from Europe (εδ 95 %, PP 98 %,
Fig. 1). Otidea minor differs from O. lactea in the (most often)
split apothecia, with a yellow outer surface and often more
narrow, straight to bent, subclaviform to claviform paraphyses
apices. Otidea lactea has the characteristic broad apices of
the paraphyses of the O. concinna clade, which however, in
O. lactea becomes hooked with age (Cao et al. 1990). The holotype was originally deposited in εHSU (Cao et al. 1990), but
later transferred to HεAS (confirmed by Hong-εei δu, HεAS).
30. Otidea minor (Boud.) Olariaga & K. Hansen, comb. & stat.
nov. — εycoBank εB808975; Fig. 34
Basionym. Otidea cantharella var. minor Boud., Icon. εycol. livr. 23: n°.
411. 1909 (preliminary text with ‘circulaires’).
≡ Flavoscypha cantharella var. minor (Boud.) Häffner, Rheinland Pfäl.
Pilzj. 4: 36. 1994.
Lectotype designated here: Boud., Icon. εycol. livr. 23: n°. 411, pl. 326.
1909; εycoBank εBT178086.
Misapplied names
– Otidea cantharella sensu δundell & Nannfeldt, Fungi Exs. Suec. 1–2: 93.
1934.
– Flavoscypha cantharella sensu Dennis, Brit. Ascomyc.: pl. 8 D. 1978.
Apothecia gregarious to caespitose, 8 – 33 mm high, 4 – 41
mm wide, broadly ear-shaped or shallowly cup-shaped, often
elongated on one side, normally split, often stipitate. Hymenium
light ochre (3A2, 3A3) to yellowish grey (4A3), sometimes with
weak rose stains or spots, when dried yellowish ochre (4A4,
4A5) to ochre (5A4). Receptacle surface citrine yellow (2A6,
2A7), slightly hygrophanous, in drying slightly paler, fading to
brownish ochre (4B5, 4B6) in age, when dried reddish ochre
(6B6–6D6) to reddish brown (5B4, 5B5), slightly furfuraceous
to slightly warty, some apothecia with low ridges coming from
the stipe, restricted to the base or reaching 1/2–1/3 of the apothecium, sometimes interconnected by low veins. Warts minute,
conical, concolorous to brownish. Stipe often well developed,
cylindrical, 2–12 × 2–5 mm. Basal tomentum and mycelium
abundant, white, seldom with purplish tones, very pale yellow
(4A2) when dried. Spores narrowly ellipsoid, often inequilateral,
with two large guttules, smooth, hyaline, (10–)10.5–12.5(–13)
× (4.5–)5–5.5(–6.2) µm (δm = 10.7–12.3 µm, Wm = 5–5.6 µm,
Qm = 2.1– 2.3; n = 10). Paraphyses straight to bent, seldom
curved, subclaviform to claviform at apices, 3–8 µm wide, with-
219
out notches, uppermost cell 20–71 µm long, when fresh with
homogeneous content or seldom containing slightly refractive,
light yellow granules at apices, often with a yellow refractive
body (Fig. 34d); when dried small, slightly refractive, hyaline
granules. Asci 153 –169 × 9 –10.5 µm. Apothecial section
750–1000 µm thick. Subhymenium c. 100 µm thick, of dense
textura intricata, hyphae 3–6 µm wide. Medullary excipulum
of textura intricata, 500–700 µm thick, hyphae 4–9(–13) µm
wide, sometimes slightly swollen, thin to thick-walled, hyaline
to very pale yellow, without resinous exudates. Ectal excipulum
of textura prismatica-angularis, 80–150 µm, cells thin-walled,
very pale yellow, unchanged in KOH, 10–33 × 6–15 µm. Surface with low flattened warts, 35–53 µm high, formed by short,
fasciculate, hyphoid hairs, of 2–3 ovoid cells, constricted at
septa, 6–9 µm wide. Non-warted parts with 2–4-celled hyphoid
hairs, with subclaviform to claviform uppermost cell, 43–58 ×
7.5–13 µm. Resinous exudates abundant, yellow or reddish
to brownish yellow, dissolving into amber drops in εδZ. Basal
mycelium of 3–7 µm wide, very pale yellow, slightly swollen
hyphae, not changing in KOH, with very scattered amorphous
or rod-shaped, yellow resinous exudates, dissolving in εδZ.
Specimens examined. dEnmark, Jylland, Nystrup Plantage, Kridtstien, on
calcareous soil under Abies, 1 Nov. 2007, T. Læssøe, Tδ-13332 (C); N Jylland,
Rold Skov, Buderupholm Bjergskov, by fence of Cypripedium, calcareous soil
along roadside, deciduous forest, 14 Sept. 1998, K. Hansen, KH.98.84 (C);
Jylland, Vorsø, under Salix caprea, 25 Sept. 1981, T. Læssøe, Tδ-0684 (C);
ibid., on base of Salix caprea, 8 Sept. 1982, T. Læssøe, Tδ-0754 (C); Sjælland, Geelskov, 10 km N of Copenhagen, 3 Aug. 1950, M. Lange (C-F-47967).
– Finland, Helsinki, Toukola, Koreankatu, Acer, Betula, Populus tremula, Salix
caprea, Sambucus, Sorbus, Aegopodium podagraria, Urtica dioica, 8 Sept.
1992, R. Saarenoksa 24592 (H); Varsinais-Suomi, δohja, Pähkinäniemi, very
rich, somewhat dry grass-herb forest with calcareous ground, 1 Aug. 1997,
U. Nummela-Salo & P. Salo 4051 (H); Varsinais-Suomi, δohja, Virkkala,
NE-slope of Pähkinäniemi, very rich, somewhat dry grass-herb forest with
calcareous soil, nearby Corylus avellana, Populus tremula, Betula pendula,
12 Sept. 2006, U. Nummela-Salo & P. Salo 10724 (H). – italy, Calabria, Acri
(CS), Croce di Greca, 14 Sept. 1995, C. Lavorato, Cδ 950914-01 (dupl. S).
– SwEdEn, Gotland, Fårö, Avanäset, under Pinus sylvestris, on humus,
sandy soil, 27 Sept. 2011, J.C. Zamora, KH.11.103 (S); Gotland, δojsta hed,
Russpark, grazed forest with Pinus sylvestris, on calcareous ground, 2 Oct.
2010, K. Hansen, K. Gillen & I. Olariaga, KH.10.311 (S); Småland, Stenbrohult, Stockanäs SSV of Stenbrohults kyrka, under Pyrus, Salix and Prunus
domestica, 9 Aug. 2011, G. Aronsson (UPS F-548414); Södermanland,
Södertalje, εörkö, Oaxen, in rich soil in deciduous forest, Corylus avellana,
Salix caprea, with Lactarius citriolens, 10 Sept. 1994, P. Höljer (H7003652);
Uppland, Uppsala, the park in front of the prison, on bare soil, among needles
etc. under Abies, 16 Aug. 1932, S. Lundell (S-F108335, Fungi Exs. Suec.
93); Öland, Högby, Horns kungsgårds Nature Reserve, under Corylus,
6 Aug. 2000, T. Knutsson, TK2000-057 (S).
Notes — Otidea minor is recognised macroscopically by
apothecia with a yellow outside, and shallow ribs and veins
at the base of at least some apothecia. Otidea concinna and
O. oregonensis can be distinguished by the broader spores
with a lower Qm (Fig. 29). In fresh material of O. minor, some
paraphyses had a strikingly, yellowish refractive body in the
upper part, a so far unique feature within Otidea. To assess its
taxonomic value, this feature has to be checked in additional
fresh material and in closely related species.
Otidea minor has been confused with O. phlebophora due to the
presence of ribs or veins at the base of the apothecia (δundell &
Nannfeldt 1934, Dennis 1978). Harmaja (2009a) proposed the
provisional name O. subconcinna for O. minor as circumscribed
here. After examining two collections annotated by him, we
consider the Finnish material to be conspecific with our Swedish finds, based on both morphological and molecular data.
Although Harmaja (2009a) did not directly compare O. minor
(as O. subconcinna) and O. phlebophora, he stated in the key
that O. phlebophora has ‘other tinges of yellow’ and mostly
straight paraphyses. We could not confirm these differences
in the material we examined. Instead, O. minor can be distin-
220
guished from O. phlebophora by the presence of at least some
apothecia without ribs, and ribs when present shallower and
less anastomosing, as well as normally split apothecia. In spite
of their morphological similarity, O. minor and O. phlebophora
are not sister species in our four-gene phylogeny, but both
are deeply nested within the O. concinna clade (Hansen &
Olariaga 2015; see also Fig. 3). Boudier´s plate under Otidea
cantharella var. minor shows shallow ribs at the base of some
apothecia, agreeing with our material, and as considered by
Harmaja (2009a). No original material could be located in PC,
and we therefore designate Boudier’s plate as the lectotype.
Persoonia – Volume 35, 2015
31. Otidea oregonensis K. Hansen & Olariaga, sp. nov. —
εycoBank εB808973; ITS barcode GenBank Kε010048;
Fig. 35
Etymology. Named after Oregon, the area where most of the specimens
of this species have been collected.
Holotype. USA, Oregon, Douglas Co., Umpqua National Forest, Diamond
δake Ranger District, under Pseudotsuga menziesii and Abies concolor, 7 Nov.
2010, J. Moore, εoorefun 58 (OSC). Isotype (S).
Apothecia gregarious to caespitose, 12–48 mm high, 23–80
mm wide, shallowly to deeply cup-shaped, sometimes elongated on one side or obconical, split, seldom entire, sometimes
stipitate. Hymenium greyish white (2A2) to pale ochre (4A2–
4A4), sometimes with rose stains or spots, when dried pale
ochre (4A2 – 4A4). Receptacle surface bright citrine yellow
Fig. 34 Otidea minor (KH.10.311)*. a. Apothecia; b. apothecium showing veined outer receptacle; c. spores; d. paraphyses showing a crystallized body; e. ectal
excipulum; f. ectal excipulum in εelzer’s reagent, resinous exudates mostly washed away, showing free hypoid hairs. — Scale bars = 10 µm; * = fresh material.
I. Olariaga et al.: A monograph of Otidea
(2A6–2A8), slightly hygrophanous, fading to brownish ochre
(4B5, 4B6) in age, when dried brownish ochre (4A4, 4B4, 5B4),
furfuraceous, sometimes some apothecia wrinkled-veined at
the base, seldom with short ribs reaching up 1/3 to the margin, partly covered by white tomentum from the base. Warts
sometimes present toward the base, minute, flat, concolorous.
Stipe sometimes well developed, cylindrical, 3–18 × 3–11 mm.
Smell mild. Basal tomentum and mycelium abundant, white,
when dried very pale yellow (4A2). Spores narrowly ellipsoid
to ellipsoid, sometimes inequilateral, with two large guttules,
rarely with a few small granules, smooth, hyaline, 10 –11.5
× 5.5 – 6(– 6.5) µm (δm = 10.4 –11.3 µm, Wm = 5.6 – 5.9 µm,
Qm = 1.8 – 2; n = 7). Paraphyses bent to curved, sometimes
straight, subclaviform to capitate at apices, 3–6(–8) µm wide,
221
without notches, uppermost cell 28–70 µm long, when dried
containing small, weakly refractive, hyaline granules. Asci
171–203 × 9–10.5 µm. Apothecial section 1000–1400(–2000)
µm thick. Subhymenium 100–140 µm thick, of dense textura
intricata, hyphae 2–4.5 µm wide, with scattered swollen, up to
12 µm wide cells. Medullary excipulum 650–850(–1400) µm
thick, of textura intricata, hyphae 5–9(–17) µm wide, sometimes swollen, thin- to thick-walled, hyaline, without resinous
exudates. Ectal excipulum of textura prismatica to textura
angularis, 70–100 µm thick, cells thin-walled, very pale yellow,
brighter yellow in KOH, 16–37 × 8–14 µm. Surface with low
warts, 35–70 µm high, formed by short, fasciculate hyphoid
hairs, of 2–4 ovoid cells, constricted at septa, 5–9 µm wide.
Non-warted parts with 2–4-celled, hyphoid hairs, with claviform
Fig. 35 Otidea oregonensis. a. Apothecia; b. apothecia showing faded colour in the outer receptacle; c. spores in water †; d. paraphyses in water †; e. ectal
excipulum in water †; f. ectal excipulum in KOH† (a, c–f: εoorefun 58, holotype; b: rh139). — Scale bars = 10 µm; † = dried material. — Photos: a. J. εoore;
b. R. Helliwell.
222
uppermost cell, 33–55 × 7–12 µm. Resinous exudates often
abundant, yellow, dissolving into amber drops in εδZ, brighter
yellow in KOH. Basal mycelium of 3–5(–8) µm wide, sometimes
slightly swollen, very pale yellow hyphae, not changing in KOH,
with very scattered, spheroid to rod-shaped, yellow resinous
exudates, dissolving in εδZ.
Specimens examined. USA, Oregon, Jackson Co., Rogue River National
Forest, under Pseudotsuga menziesii, Abies concolor, Pinus ponderosa,
2 Dec. 1999, B. Schroeter (OSC 72950); ibid., 27 Oct. 1990, D. McKay,
NSW6354 (OSC 132740, dupl. S); Oregon, Josephine Co., Bureau of δand
εanagement, εedford District, Grants Pass Resource Area, δittle Pickett
Creek, under Pseudotsuga menziesii, Calocedrus decurrens, Lithocarpus
densiflorus, Quercus chrysolepis, 19 Dec. 2000, R. Meyer (OSC 108041);
Oregon, εarion Co., near Breitenbush Hot Springs Community, 27 Oct.
1996, J. Trappe (OSC 56745); Oregon, Umpqua NF, Diamond δake RD,
under Pseudotsuga menziesii, 2 Nov. 2010, J. Moore, εoorefun 31 (S); ibid.,
old growth forest with Abies concolor, Pseudotsuga menziesii, 2 Nov. 2010,
R. Helliwell, rh139 (OSC); Washington, δewis Co., Gifford Pinchot National
Forest, Camp Creeks Falls Trail, 6 Nov. 1997, M. Castellano (OSC 56829).
Notes — Otidea oregonensis is characterised by a citrine
yellow receptacle surface, often a wrinkled to veined, or shallowly ribbed apothecial base, and relatively broad spores. The
apothecia are typically obconical cup-shaped, with a narrow
base. Otidea oregonensis was treated as O. rainierensis Kanouse by Peterson (1998), primarily based on the presence of
swollen apices of the paraphyses. Also, one of the paratypes
of O. rainierensis (EGS2179), sequenced by Peterson (1998),
is nested within the O. oregonensis lineage in our ITS-δSU
phylogeny (Fig. 3). Nevertheless, our sequences of the holotype
of O. rainieriensis, from four different gene regions, show it
belongs to another lineage, well separated from O. oregonensis
(Hansen & Olariaga 2015). Kanouse´s concept of O. rainier-
Persoonia – Volume 35, 2015
ensis was therefore mixed. The ITS-δSU phylogeny, including
a larger sampling of the O. concinna clade, likewise resolves
O. oregonensis as a distinct species.
Otidea phlebophora and O. minor, so far only known from
Europe, resemble O. oregonensis in the cup-shaped apothecia with yellow outside and swollen paraphyses. Otidea
phlebophora differs macroscopically from O. oregonensis in
predominantly entire apothecia, with always a strongly veinedribbed base reaching up 1/2 to the margin, while only small
veins are present in some apothecia of O. oregonensis. Based
on the material examined here, the slightly narrower spores in
O. phlebophora (Qm = 2–2.1 vs Qm = 1.8–2 in O. oregonensis)
are a constant and reliable character to separate these species
(Fig. 29). Otidea minor tends to have smaller, shallower and
more broadly cup-shaped apothecia than O. oregonensis. The
spores of O. minor have a higher Qm value (2.1–2.3) that does
not overlap with the Qm of O. oregonensis.
32. Otidea phlebophora (Berk. & Broome) Sacc., Syll. Fung.
8: 97. 1889 — Fig. 36
Basionym. Peziza phlebophora Berk. & Broome, Ann. εag. Nat. Hist.,
ser. III, 18: 122. 1866.
≡ Flavoscypha phlebophora (Berk. & Broome) Harmaja, Karstenia 14:
107. 1974.
Lectotype designated here: England, North Somerset, Brislington, 16
Sept. 1853 (K(ε) 144045, ex Herb. C.E. Broome) !; εycoBank εBT178087.
Apothecia gregarious, 8–12 mm high, 3–40 mm wide, shallowly to deeply cup-shaped, sometimes elongated on one side,
entire, seldom split, often stipitate. Hymenium pale ochre (4A2,
4A3), sometimes with rose stains or spots, when dried yellowish
Fig. 36 Otidea phlebophora (JV06-385). a. Apothecia; b. spores in water †; c. paraphyses in water †; d. ectal excipulum in water †. — Scale bars = 10 µm;
= dried material. — Photos: a. J. Vesterholt.
†
223
I. Olariaga et al.: A monograph of Otidea
ochre (4B4, 5B6, 5C6) to reddish brown (6C5, 6C6). Receptacle surface bright citrine yellow (3A7, 2A7), sometimes with a
greenish hue (2A6), slightly hygrophanous, fading to brownish
ochre (4B5) in age, when dried yellowish brown (5B6, 5C6) to
reddish brown (6C6, 6D6), furfuraceous, with high ridges in the
basal part of all apothecia reaching up 1/2–1/3 to the margin,
interconnected by veins, partly covered by white basal tomentum. Warts minute, flat to rounded, appressed, concolorous.
Stipe often well developed, cylindrical, 1–10 × 1.5–3 mm. Basal
tomentum and mycelium abundant, white, very pale ochre (5A2)
when dried. Spores narrowly ellipsoid, sometimes inequilateral,
with two large guttules, smooth, hyaline, (9.5–)10–11.5(–12)
× (4.5 –)5 – 5.5(– 6.2) µm (δm = 10.2 –11 µm, Wm = 4.9 – 5.1
µm, Qm = 2–2.1; n = 5). Paraphyses straight to bent, cylindrical to claviform at apices, 2–3.5 µm wide, without notches,
uppermost cell 23–46 µm long, when dried containing small,
rather refractive, hyaline guttules. Asci 130–153 × 8–9 µm.
Apothecial section 650–1000 µm thick. Subhymenium 80–100
µm thick, of dense textura intricata, hyphae 2 – 3 µm wide.
Medullary excipulum of textura intricata, 450–650 µm thick,
hyphae 4–7(–14) µm wide, sometimes swollen, thick-walled,
hyaline to very pale yellow, without resinous exudates. Ectal
excipulum of textura prismatica-angularis, 80–100 µm thick,
cells thin-walled, hyaline to very pale yellow, unchanged in KOH,
15–21 × 6–11 µm. Surface with low flattened warts, 40–60 µm
high, formed by short, fasciculate hyphoid hairs, of 3–4 ovoid
cells, constricted at septa, 8–12 µm wide. Non-warted parts
with 2–3-celled hyphoid hairs, with claviform uppermost cell,
30–60 × 7–13 µm. Resinous exudates abundant, yellow, dissolving into amber drops in εδZ. Basal mycelium of 3.5–7 µm
wide, slightly swollen, very pale yellow hyphae, not changing
in KOH, with very scattered, spheroid to rod-shaped, yellow
resinous exudates, dissolving in εδZ.
Specimens examined. dEnmark, NE Jylland, Rubjerg Knude Plantage,
under Abies in large fairy ring, 28 Aug. 2006, J. Vesterholt & L. Vesterholt,
JV06-385 (C); NE Sjælland, Ravnsholts Hegn, under Picea abies, 30 Aug.
1999, B.W. Pedersen (C-F-71506). – Finland, Varsinais-Suomi, δohja,
Jalassaari, Alho, by the Ahtiala manor, in rich forest with Quercus robur,
Corylus avellana, Prunus padus, cultivated Larix sibirica and Betula, 30 Aug.
1967, H. Harmaja (H6010675). – SwEdEn, Dalarna, Husby, Husby värdhus,
on lawn under Quercus, Tilia, Larix and Acer, 31 Aug. 1958, R. Morander
(UPS F-144691); Skåne, Helsinborg, Jordbodalen by Harlyckan, on sandy
ground in deciduous forest under Quercus, 13 July 1995, S.-Å Hanson, SÅH
30601 (C); Uppland, Djurö, Runmarö, Södersunda, in the city, on the ground
under Syringa, 18 Sept. 1949, G. Haglund & R. Rydberg (S-F108338, UPS
F-144689); Uppland, Uppsala, Sunnersta, Almlund, calcareous, humus-rich
clay, 23 Aug. 1986, J. Nitare (UPS F-119845). – UK, Northampshire, King´s
Cliffe, on soil, 1853, M.J. Berkeley (K(ε) 144046, syntype of P. phlebophora);
North Somerset, Brislington, The Beeches, Sept. 1853, C.E. Broome (K(ε)
194582, syntype of P. phlebophora).
Other specimen examined. Otidea integra — italy, Sopramonte, 1892,
G. Bresadola (S-F108342). δocality not specified, in silvis mixtis, 1892, G. Bresadola (PC 124965).
Notes — Otidea phlebophora is primarily characterised by
high anastomosing ribs and veins towards the apothecial base.
Diagnostic characters are in addition, the predominantly entire
apothecia, with a citrine yellow outside. For a comparison with
O. concinna, O. minor and O. oregonensis see Notes under
those species. Harmaja (1986) elevated O. integra to species
rank based on smaller apothecia and broader, curved paraphyses. The likely original specimen of O. integra collected in 1892
and kept in Bresadola’s herbarium (S-F108342), shows curved
paraphyses up to 5 µm broad, the same as Harmaja (1986,
2009a) gave for O. phlebophora. The paraphyses in the O. phlebophora material examined by us were straight to bent, and
otherwise similar to the original material of O. integra. In spite of
the morphological similarity, the ITS2 sequence of O. integra (281
bp obtained) is different from the ITS sequences of the material
assigned to O. phlebophora by us, and we preliminary accept
O. integra as a separate species. The position of O. integra is
without support in our ITS-δSU phylogeny (Fig. 3).
Nomenclatural notes — Harmaja (1974: 107) indicated a lectotype of O. phlebophora at K, but he gave no collection number
and the typification was not achieved. We have studied three
of four syntypes at Kew and select here the richest collection
containing ten apothecia as the lectotype. It conforms to the
current interpretation of the name, with several entire apothecia
(i.e. without a split), a ribbed-veined base seen on two of the
apothecia (the others with the base glued to the cardboard),
spores in the range 9.5 –11.7 × 4.7– 5.9 µm (δm = 10.6 µm,
Wm = 5.1 µm), paraphyses straight and enlarged at apices, and
abundant yellow resinous exudates in the ectal excipulum that
dissolve into amber drops in εδZ.
33. Otidea rainierensis Kanouse, εycologia 41: 674. 1949
Holotype. USA, Washington, Pierce County, Lower Tahoma Creek, εT
Rainier National Park, 23 Aug. 1948, A.H. Smith 30553 (εICH 14410) !
= Otidea kauffmanii Kanouse, εycologia 41, 6: 673. 1949.
Holotype. USA, εichigan, δakeland, 18 July 1915, C.H. Kauffman (εICH
14409) !
Kanouse (1949) only had dried material of O. rainierensis and
described the outside of the apothecia as ‘ochraceous buff ’,
‘cinnamon buff ’ to ‘wood brown’ and the hymenium as ‘avellaneous’, ‘vinaceous buff’ to ‘drab grey’. For O. kauffmanii she
had notes on fresh material and she separated it from O. rainierensis based on the presence of yellow tones in the apothecia
(outside ‘chamois’ to ‘ochraceous’, hymenium ‘cream buff ’). We
believe O. rainierensis does have yellow apothecial tones, as
we observed small yellow resinous exudates (in water) in the
type collection of O. rainierensis, and as observed in closely
allied species the yellow colour can fade and almost disappear
with age (Fig. 32b). Different spore sizes were also used to
distinguish the two species. The spores of the holotype of O.
kauffmanii are however, larger than noted in the protologue,
11.5–12.5 × 5.5–6.5 µm, δm = 11.9 µm, Wm = 6 µm, Qm = 2
(spores 8–10(–12) × 5–6(–7) µm in the protologue), and thus
overlapping with the spores of the holotype of O. rainierensis,
11–12 × 6.5–7 µm, δm = 11.7, Wm = 6.7 µm, Qm = 1.7. We suggest O. kauffmanii and O. rainierensis constitute a single species, based on molecular and morphological study of the type
material (Fig. 3). Our ITS sequences of the holotypes differ by
4 bp. See also comments on O. microspora under Excluded,
dubious and imperfectly known taxa. Otidea rainierensis is
characterised by a smooth apothecial base, long and relatively
broad spores, compared to the rest of the species in the O. concinna clade, and by paraphyses with abruptly enlarged, broadly
clavate to globose apices.
EXCLUDED, DUBIOUS AND IMPERFECTLY KNOwN
TAXA
Cochlearia (Cooke) δambotte, εém. Soc. Roy. Sci. δiege, ser.
2. 14: 323. 1888
Nom. illegit. Art. 53.1, non Cochlearia δ., Sp. Pl. 2: 647. 1753 (Cruciferae).
Basionym. Peziza subg. Cochlearia Cooke, εycographia part 6: 252
(‘Index Systematicus’). 1879.
Notes — Eckblad (1968) selected Peziza cochleata as the
type species for the genus Cochlearia considering it a synonym
of Otidea. However, Rifai (1968) had already selected Peziza
aurantia as the type species, herewith making Cochlearia a
synonym of Aleuria. He felt too much doubt had surrounded
the identity of P. cochleata and it better not be selected as the
type species.
224
Otidea abietina (Pers.) Fuckel, Jahrb. Nassauischen Vereins
Naturk. 23–24: 330. 1870 ‘1869 –1870’
Basionym. Peziza abietina Pers., Neues εag. Bot. 1: 113. 1794: Fr., Syst.
εycol. 2: 47. 1822.
≡ Discina abietina (Pers.) Rehm, Rabenh. Krypt.-Fl., ed. 2, 3: 977. 1896.
≡ Pseudotis abietina (Pers.) Boud., Icon. εycol. list prél. 600 sp.: 3 (unnumbered page). 1904.
Notes — Nannfeldt (1966) stated that the original material
belongs to Peziza badia, but Harmaja (2009a) and Carbone
(2010c) reported two collections in Persoon’s herbarium under
O. abietina representing O. propinquata and a third collection
O. bufonia. If typified with elements belonging to either O. propinquata or O. bufonia, the name O. abietina would take priority
over any of those. Otidea abietina is the type species of the
genus Pseudotis (Boud.) Boud. The identity of Pseudotis will
thus remain open until O. abietina is clarified or typified. Another
way to typify O. abietina would be to select an element belonging to Peziza badia. However, this choice should be studied
more thoroughly, since it would make Pseudotis available as
a genus name for the Peziza depressa-Ruhlandiella lineage, if
Peziza is split into smaller genera in the future (Hansen et al.
2005). For the time being we regard O. abietina as a nomen
ambiguum, as several others (Harmaja 2009a, Carbone 2010c,
Parslow & Spooner 2013).
Otidea alba Velen., εonograph. Discom. Bohemiae 1: 354.
1934
Holotype. CzECh rEpubliC, Karlštejn, Sept. 1924, Fechtner (PRε 149788) !
Notes — The holotype specimen has the typical oblong
spores of the O. alutacea complex, (13.5–)14.5–16.5(–17.5)
× 6.5–7.5 µm (δm = 14.9 µm, Wm = 6.9, Qm = 2.1). Judging from
the spore size, O. alba might represent either O. alutacea s.str.
or the O. alutacea clade 3b. It should be considered in future
studies of the complex.
Otidea aurantia (Pers.) εassee, Brit. Fungus-Fl. 4: 448. 1895
Basionym. Peziza aurantia Pers., Observ. εycol. 2: 76. 1800: Fr., Syst.
εycol. 2: 49. 1822.
Notes — This is the type species of Aleuria, A. aurantia
(Pers.: Fr.) Fuckel.
Otidea aurantia var. atromarginata (W. Phillips & Plowr.)
εassee, Brit. Fungus-Fl. 4: 449. 1895
Basionym. Peziza aurantia var. atromarginata W. Phillips & Plowr., Gard.
Chron. 17: 191. 1882.
Notes — The blood red disc with short, obtuse, dark brown,
3–4 septate hairs, giving the margin a dark appearance suggest
this may be a species of Melastiza. The spores are tuberculate
with thread-like appendages.
Otidea aurantia var. stipitata (W. Phillips) εassee, Brit. FungusFl. 4: 448. 1895
Basionym. Peziza aurantia var. stipitata W. Phillips, εan. Brit. Discom.:
57. 1887.
Notes — The small, bright scarlet apothecia with a ‘stem
equalling the height of the cup, 4 mm’ and the ornamented
spores suggest this is a species of Aleuria or Sowerbyella. It was
described as a variety of Aleuria aurantia (as Peziza aurantia
Oed.). Ramsbottom (1914) cited O. aurantia var. stipitata as a
synonym of Sowerbyella rhenana (as Aleuria rhenana Fuckel),
but the type material is presumably lost (Spooner & Yao 1995)
and no modern interpretation can be provided.
Persoonia – Volume 35, 2015
Otidea auriculariiformis Henn., Hedwigia 36: 232. 1897
Holotype. brazil, A. Glaziou no. 20181 (S-F9965, ex Herb. Sydow) !
Notes — This species belongs to the genus Phillipsia, Sarcoscyphaceae. The large spores, (30.5 –)31.5 – 36.5(– 37) ×
12.5–14.0 µm (δm = 33.2 µm, Wm = 13.1 µm, from 13 spores),
are ellipsoid, inequilateral in profile view, smooth or with faint
cyanophobic, parallel, longitudinal ridges. Asci seem thickwalled, with an internal eccentric thickened apical pad. The
medullary excipulum is of interwoven hyphae, running mostly
parallel with the outer surface and the ectal excipulum is a
narrow band of textura prismatica, with the long axes of the
cells parallel to the exterior. The tropical distribution and the
substrate, suggested in the diagnosis to be wood, are typical
for Phillipsia.
Otidea cinerascens Velen., Novit. εycol.: 152 .1947
Holotype. CzECh rEpubliC, εoravia, Žarošice, Aug. 1940, V. Vacek (PRε
151779).
Notes — The grey-ochraceous apothecia and spores with
parallel sides suggest O. cinerascens belongs to the O. alutacea complex. Type not studied by us, but annotated in 2009 by
B. Spooner as O. alutacea (a photograph of the collection and
annotation provided by Jan Holec, PRε).
Otidea cochleata (δ.) Fuckel, Jahrb. Nassauischen Vereins
Naturk. 23–24: 329. 1870. ‘1869 –1870’
Basionym. Peziza cochleata δ., Sp. Pl. 4: 183. 1753: Fr., Syst. εycol.,
Index: 129. 1832.
≡ Cochlearia cochleata (δ.) δambotte, Fl. εycol. Belgique 1: 323. 1880.
Notes — The interpretation of the original description is
difficult, but O. cochleata has been treated as a taxon in the O.
alutacea group (e.g. δundell & Nannfeldt 1938, Dissing 2000,
εornand & Courtecuisse 2005, Zhuang 2006), probably following Bulliard (1791: plate 154 as Peziza cochleata). Part of the
Bulliard plate (f. b) has now been selected as the lectotype for
O. alutacea (Carbone 2010a). The name O. cochleata should
be considered in future revisions of the O. alutacea complex.
For a review of the nomenclatural history of O. cochleata see
Carbone (2010a) and Parslow & Spooner (2013).
Otidea darjeelensis (Berk.) Sacc., Syll. Fung. 10: 4. 1892
Basionym. Peziza darjeelensis Berk., Hooker’s J. Bot. Kew Gard. εisc.
3: 202. 1851.
Notes — Rifai (1968) stated that the type specimen of P. darjeelensis at Kew has iodine positive asci and echinulate spores,
and does not belong to Otidea. Two collections of P. darjeelensis
are present in Kew that may represent original material: india,
Sikkim, J.D. Hooker (K(ε) 177412, ex Herb. Berkeley); and
india, Sikkim (K(ε) 177413, ex Herb. Cooke). No annotation
label by Rifai was found (B. Aguirre-Hudson, pers. comm. ), but
he most likely studied the collection from Berkeley’s herbarium,
because he listed J.D. Hooker as the collector. The collection
K(ε) 177413 might be a part of K(ε) 177412, because Cooke
(1876, f. 215) illustrated P. darjeelensis from specimens in
Berkeley’s herbarium. Both collections should be studied and
a lectotype selected.
Otidea dochmia (Berk. & ε.A. Curtis) Sacc., Syll. Fung. 8:
95. 1889
Basionym. Peziza dochmia Berk. & ε.A. Curtis, J. δinn. Soc. Bot. 10:
364. 1869.
≡ Phillipsia dochmia (Berk. & ε.A. Curtis) Seaver, N. Amer. Cup-fungi,
Operc.: 184. 1928.
I. Olariaga et al.: A monograph of Otidea
≡ Aurophora dochmia (Berk. & ε.A. Curtis) Rifai, Verh. Kon. Ned. Akad.
Wetensch., Afd. Natuurk., sect. 2, 57: 52. 1968.
Notes — This is the type species of the genus Aurophora.
Rifai (1968) distinguished Aurophora from Phillipsia by its fanshaped apothecia and the presence of a gelatinous matrix in
the medullary excipulum.
Otidea domingensis (Berk.) Sacc., Syll. Fung. 8: 97. 1889
Basionym. Peziza domingensis Berk., Ann. εag. Nat. Hist., ser. II, 9: 201.
1852.
Notes — This is the type species of the genus Phillipsia,
P. domingensis (Berk.) Berk. (see Hansen et al. 1999).
Otidea doratophora (Ellis & Everh.) Sacc., Syll. Fung. 8: 96.
1888
225
Notes — The original sense of O. grandis corresponds most
likely to a species of the Peziza depressa-Ruhlandiella lineage
(Hansen et al. 2005) due to the lack of a split. No original material of O. grandis seems to be kept in Persoon´s herbarium (δ).
The name O. grandis has been used for O. bufonia (Boudier
1905; specimen PC0093644 is O. bufonia) or O. unicisa (Kanouse 1949, δiu & Zhuang 2006). We regard it here as a nomen
dubium and confusum in agreement with Harmaja (2009a).
Otidea grandis var. scheremetjeffii Henn., Hedwigia 42, 3:
(116). 1903
Notes — Hennings (1903) described this taxon based on
specimens kept in formalin and the colours provided in the
protologue are probably imprecise. The spore size can fit
O. bufonia or O. onotica, but there is no type material extant in
B or S, and a precise interpretation cannot be proposed here.
Basionym. Peziza doratophora Ellis & Everh., J. εycol. 1: 90. 1885.
Notes — The small spores and asci, along with the pointed
paraphyses, suggest this taxon does not belong to Otidea. Cash
(1953) proposed O. doratophora is a synonym of Ionomidotis
irregularis (Schwein.) E.J. Durand (as Midotis irregularis).
Otidea euplecta (Cooke) Sacc., Syll. Fung. 8: 97. 1889
Basionym. Peziza euplecta Cooke, εycographia part 3: 125. 1876.
Holotype. USA, Alabama, Peters 4560 (K(ε) 161851, ex Herb. Berkeley
as Peziza phlebophora var.) !
Notes — The type is in a poor condition: one apothecium immature; the other infected. But the species likely belongs to
Sarcoscypha, Sarcoscyphaceae. The asci are inamyloid, thickwalled and with an eccentrically placed, thickened operculum,
spores are ellipsoid, slightly inequilateral, 19–21 × 10.5–11 µm,
smooth, and paraphyses straight, filiform, branching above. The
excipulum is composed of interwoven hyphae that give rise on
the outside to shallow pustules.
Otidea felina (Pers.) Bres., Fungi Trident. ser. 2, fasc. 14:
103. 1900
Basionym. Peziza felina Pers., εycol. Eur. 1: 223. 1822.
Holotype. FranCE, prope Pariseos, sylvula εeudon (δ0116774, Herb.
Persoon).
Notes — Van Vooren & Carbone (2012) revised the holotype
and demonstrated that it belongs to the O. alutacea group.
Parslow & Spooner (2013) considered O. felina a synonym of
O. alutacea. Further studies on the O. alutacea group should
consider the name O. felina, which might be epitypified for an
unequivocal interpretation.
Otidea fibrillosa εassee, Brit. Fungus-Fl. 4: 449. 1895
≡ Pseudaleuria fibrillosa (εassee) J. εoravec (‘Pseudoaleuria’), Acta
εus. εorav. Sci. Biol. 88: 51. 2003.
Otidea harperiana Rehm, Ann. εycol. 2: 34. 1904
Holotype. USA, Ohio, Blue εountains, on ground, 6 June 1903, Harper
333 (S-F9961, ex Herb. Rehm, ‘Herb. R.A. and A.ε. Harper 333’) !
Notes — This species is closely related or conspecific with
Peziza phyllogena Cooke. The asci are strongly amyloid in
εδZ with a general bluing over the apex and the spores are
ornamented with irregular, low, separate warts that are higher
and more densely placed at the poles (forming ‘pole caps’).
Our spore measurements from the holotype, 18–19.5 × 8–9
µm (δm = 18.6 µm, Wm = 8.5 µm, from 15 spores) are larger
than those given in the protologue, 15 –17 × 5–7 µm.
Otidea hirneoloides (Berk.) Sacc., Syll. Fung. 8: 96. 1889
Basionym. Peziza hirneoloides Berk. in Berkeley & Curtis, J. δinn. Soc.
Bot. 10: 365. 1869.
≡ Phillipsia hirneoloides (Berk.) Berk., J. δinn. Soc. Bot. 18: 388. 1881.
Notes — The wood-inhabiting, ear-shaped apothecia, and
especially, the cymbiform spores suggest this name is to be
referred to Phillipsia. Hansen et al. (1999) suggested P. hirneoloides belongs to the Phillipsia domingensis complex.
Otidea lechria (Berk. & Broome) Sacc., Syll. Fung. 8: 97. 1889
Basionym. Peziza lechria Berk. & Broome, J. δinn. Soc. Bot. 14: 103.
1875.
Holotype. Sri lanka, on rotten wood, Nov. 1867, G.H.K. Thwaites
(K(ε)161847, ex Herb. Berkeley).
Notes — B. Spooner annotated the holotype in 2008, and
noted it has amyloid asci and belongs to Peziza.
Otidea lilacina R. Heim & δ. Remy, Bull. Soc. εycol. France
48: 65. 1932
Notes — According to εoravec (2005) this is a species of
Pseudaleuria, P. fibrillosa.
Notes — The ornamented, multi-guttulate spores and straight
paraphyses suggest that this taxon does not belong to Otidea.
No original material could be traced in PC (B. Buyck, pers.
comm.).
Otidea grandis (Pers.) Boud., Bull. Soc. εycol. France 9:
10. 1893
Otidea lobata Rodway, Pap. & Proc. Roy. Soc. Tasmania: 116.
1925 ‘1924’
Basionym. Peziza grandis Pers., Ann. Bot. Usteri 15: 27. 1795.
≡ Peziza abietina var. grandis (Pers.) Pers., εycol. Eur. 1: 233. 1822.
≡ Aleuria grandis (Pers.) Gillet, Champ. France Discomyc.: 42. 1879.
≡ Scodellina grandis (Pers.) Seaver, N. Amer. Cup-fung., Operc.: 186.
1928.
Notes — Rifai (1968) stated that the type specimen appears
to represent the inoperculate genus Discinella.
226
Otidea luculenta (Cooke) εassee, Brit. Fungus-Fl., 4: 450.
1895 (‘leuculenta’)
Basionym. Peziza luculenta Cooke, εycographia part 3: 121. 1876.
Notes — The entire, orange apothecia and straight paraphyses with orange granules suggest this name does not belong
to Otidea. Nannfeldt (1966) noted that O. luculenta has ‘other
affinities’ than Otidea, but did not provide a generic placement.
Otidea luteonitens (Berk. & Broome) εassee, Brit. FungusFl. 4: 449. 1895
Basionym. Peziza luteonitens Berk. & Broome, Ann. εag. Nat. Hist., ser.
II, 7: 180. 1851.
Notes — This name is currently placed in Aleuria as Aleuria
luteonitens (Berk. & Broome) Gillet.
Otidea micropus (Pers.) Sacc., Syll. Fung. 8: 98. 1889
Basionym. Peziza micropus Pers., Icon. Desc. Fung. 2: 30. 1800: Fr.,
Syst. εycol. 2: 54. 1822.
Notes — This name is a synonym of Peziza varia (Hedw.:
Fr.) Fr. sensu Hansen et al. (2002).
Otidea microspora (Kanouse) Harmaja, Karstenia 15: 32. 1976
Basionym. Otidea alutacea var. microspora Kanouse, εycologia 41: 668.
1949.
Notes — Kanouse (1949) described this taxon as a variety
of O. alutacea. She indicated two different collections as the
type; A.H. Smith 9351 after the diagnosis and A.H. Smith 17699
in the material examined. We have studied A.H. Smith 9351
(εICH 14406, dupl. UPS F-629985 !) and it has far larger spores
than stated in the protologue (13–15.5 × 7–8 µm vs 9–10 ×
5.5–6.5 µm). The oblong spores, an ectal excipulum of textura
angularis, and the absence of yellow pigment in KOH indicate
it belongs to the O. alutacea complex. As for A.H. Smith 17699
(UPS F-629996 !), the spores match the original description.
The apothecial shape is similar to that of O. rainierensis. A GenBank ITS sequence of a paratype of O. microspora (A.H. Smith
30502) differs only 1 bp from the ITS sequence of the holotype
of O. rainierensis (Fig. 3), but we prefer not to select a lectotype
until all the original material has been examined. We therefore
treat the name as doubtful for the time being.
Otidea neglecta εassee, Grevillea 22: 66. 1894
Persoonia – Volume 35, 2015
species of Otidea. The Bucholtz herbarium was bought by the
FH, but no authentic material of O. olivacea could be located
there.
Otidea onotica var. ochracea (Fr.) Sacc., Syll. Fung. 8: 95.
1889
Basionym. Peziza onotica var. ochracea Fr., Syst. εycol. 2: 48. 1822: Fr.
loc. cit. (‘ß ochracea’).
≡ Peziza ochracea (Fr.) P. Karst., Not. Sallsk. Fauna Fl. Fenn. Forh. 10:
110. 1869.
≡ Otidea ochracea (Fr.) Seaver, Bull. δab. Nat. Hist. Iowa State Univ. 5:
45. 1904.
Notes — Fries (1822) stated that this variety is close to
O. onotica. It was elevated to species rank by Karsten (1869), who
later considered it a synonym of O. onotica (Karsten 1871). As
no original material exists and the original description is meagre,
we are not able to provide a good interpretation of this taxon.
Otidea pleurota (W. Phillips) Sacc., Syll. Fung. 8: 97. 1889
Basionym. Peziza pleurota W. Phillips in Cooke, εycographia part 5:
208. 1878.
≡ Iotidea pleurota (W. Phillips) Clem., Gen. Fungi: 175. 1909.
Holotype. England, 1877, W. Phillips (K(ε)29973).
Notes — Eckblad (1968) observed strongly amyloid asci in
the type and concluded it belongs to Peziza. B. Spooner annotated the type as ‘Peziza badiofusca ?’.
Otidea radiculata (Sowerby) Bres., Fungi Trident. ser. 2, fasc.
11–13: 72. 1898
Basionym. Peziza radiculata Sowerby, Col. Fig. Engl. Fung. 1: 46 (unnumbered page), t. 114. 1797: Fr., Syst. εycol. 2: 81. 1822.
Notes — This name is placed in Sowerbyella, as S. radiculata (Sowerby) Nannf. Yao & Spooner (2006) examined the
type at K and confirmed its placement.
Otidea reisneri Velen., České Houby 4 –5: 872. 1922
Notes — Svrček (1976) studied the type material and concluded it is a synonym of Sowerbyella radiculata (Sowerby: Fr.)
Nannf.
Otidea schulzeri Quél. in Schulzer, Hedwigia 24, 4: 150. 1885
Notes — This name was erected as a new name for O. auricula in the sense of Rehm (1883), Saccardo (1889) and Bresadola (1884, as Peziza). The species that these three authors
treated under the epithet auricula is Wynnella silvicola (Beck)
Nannf. in its current sense.
Notes — The apothecia of O. schulzeri were described as
elongated on one side, split, pale yellow-grey and pseudostipitate. However, the very thick flesh (3–4 mm), the straight
paraphyses and the spore size (20–28 µm) suggest this taxon
does not represent Otidea. Also Nannfeldt (1966) stated that
O. schulzeri could hardly belong to Otidea.
Otidea obtecta (Schwein.) Sacc., Syll. Fung. 8: 98. 1889
Otidea silvicola Beck in Sacc., Syll. Fung. 8: 97. 1889
Basionym. Peziza obtecta Schwein., Trans. Amer. Philos. Soc. ser. 2, 4:
170. 1832 ‘1834’.
Notes — This is Wynnella silvicola. The name was created
for Peziza atrofusca Beck, a later homonym for P. atrofusca
Berk. & ε.A. Curtis.
Notes — The original description gives stipitate, 1–1.5 cm
wide, cinnamon-coloured apothecia with a split, growing among
leaves. Seaver (1928) stated that the identity of O. obtecta is
uncertain. No original material could be located in PH and the
identification of this species cannot be inferred.
Otidea sparassis Quél., Rev. εycol. (Toulouse) 54: 65. 1892
Notes — The uniguttulate spores do not suggest an Otidea.
This taxon might refer to a sparassoid Helvella species.
Otidea olivacea Bucholtz, Bull. Soc. Imp. Naturalists εoscou
2: 325. 1897
Otidea subonotica Henn., Hedwigia 36: 232. 1897
Notes — The curved, uniguttulate spores and the occurrence
on a rotten trunk suggest that this taxon does not refer to a
Notes — The original description does not provide any discordant feature for Otidea, except it was reported from Brazil.
I. Olariaga et al.: A monograph of Otidea
No type material of O. subonotica exists in Hennings herbarium
in B (Carbone 2009) or in S, and its correct placement cannot
be inferred with certainty here.
Otidea succosa (Berk.) Thüm., εycoth. Univ. 15: no. 1411.
1879
Basionym. Peziza succosa Berk., Ann. εag. Nat. Hist., ser. I, 6: 358.
1841.
Notes — This is currently considered to be a species of
Peziza s.l. (Hansen et al. 2005).
Otidea tasmanica Rodway, Pap. & Proc. Roy. Soc. Tasmania:
116. 1925 ‘1924’
Notes — Rifai (1968) studied the type specimen of O. tasmanica in Kew and synonymised it with Peziza praetervisa
Bres. (sensu Dennis, as Rifai depicted ornamented spores
from the type specimen). Thus, O. tasmanica might be close
to, or a synonym of, Peziza subviolacea Svrček.
Otidea violacea A.δ. Sm. & Ramsb., Trans. Brit. εycol. Soc.
5: 237. 1916
Holotype. England, Warwickshire, W.B. Grove 1915 (K(ε) 30407, ex Herb.
W.B. Grove).
Notes — Parslow & Spooner (2013) examined the holotype
and concluded it is a species of Peziza (cf. azureoides Donadini).
Acknowledgements This monograph became more comprehensive thanks
to the great efforts of numerous mycologists who provided us with interesting
material. All our colleagues who contributed collections, discussed issues
or helped us in other ways are warmly thanked. Especially we thank Nancy
S. Weber, Rosanne Healy, εats Karström and Tapio Kekki for putting at our
disposal valuable material accompanied by colour photographs. We also
thank the curators and staff in the herbaria mentioned in the taxonomic
part, H.J.ε. Sipman (B), ε. Herrera and I. Salcedo (BIO), H. Knudsen (C),
P.G. Jamoni and D. Bolognini (GεFN), D.H. Pfister and G. δewis-Gentry
(FH), P. Salo and S. Stenroos (H), E.D. δiu (HKAS), Hong-εei δu (HεAS),
B. Aguirre-Hudson, B. Dentinger and B. Spooner (K), R. Rabeler and
P. Rogers (εICH), D. εcδaughlin (εIN), A.B. εujic and J. Spatafora (OSC),
B. Buyck (PC), A. Freire-Fierro (PH), J. Holec (PRε), S. Huhtinen (TUR,
TUR-A) and R. Berndt (ZH), for searching material and arranging loans. We
appreciate Kerstin Gillen´s assistance during fieldwork in 2010. Juan Santos
and Xiang-Hua Wang collected part of the material during their participation in the Swedish Pezizomycetes project. We are indebted to Xiang-Hua
Wang for sequencing a few important Otidea collections, for discussions and
help in editing the manuscript, and Seppo Huhtinen for instructive discussions on resinous exudates. We thank D.H. Hawksworth and δ.A. Parra for
nomenclatural advice. Donald H. Pfister and Trond Schumacher reviewed
the manuscript and we are grateful for their valuable comments and corrections. Funding for this research was provided by a grant from the Swedish
Taxonomy Initiative to K.H. (grant no. 143/07 1.4).
REFERENCES
Arpin N. 1969. δes caroténoïdes des Discomycètes: essai chimotaxinomique.
Bulletin εensuel de la Société δinnéenne de δyon 38 (suppl.): 1–169.
Batsch AJGK. 1783. Elenchus Fungorum. Halle, Germany.
Bonorden HF. 1851. Handbuch der allgemeinen mykologie als anleitung zum
studium derselben, nebst speciellen beiträgen zur vervolkommung dieses
zweiges der naturkunde. Schweizerbart, Stuttgart, Germany.
Boudier JδÉ. 1885. Nouvelle classification naturelle des Discomycètes charnus connus généralement sous le nom de Pezizes. Bulletin de la Société
εycologique de France 1: 91–120.
Boudier JδÉ. 1905. Icones εycologicae, livr. 6. Klincksieck, Paris, France.
Boudier JδÉ. 1906. Icones εycologicae, livr. 7. Klincksieck, Paris, France.
Boudier JδÉ. 1907. Histoire et classification des Discomycètes d’Europe.
Klincksieck, Paris, France.
227
Boudier JδÉ. 1908. Icones εycologicae, livr. 21. Klincksieck, Paris, France.
Boudier JδÉ. 1909a. Icones εycologicae, livr. 23. Klincksieck, Paris, France.
Boudier JδÉ. 1909b. Icones εycologicae, livr. 24. Klincksieck, Paris, France.
Boudier JδÉ. 1910. Icones εycologicae, livr. 29. Klincksieck, Paris, France.
Bresadola G. 1884. Fungi Tridentini novi, vel nondum delineati, descripti, et
iconibus illustrati. Ser. 1, fasc. 4–5: 43–70, pl. 46–75. εonauni, Trento, Italy.
Bresadola G. 1898. Fungi Tridentini novi, vel nondum delineati, descripti, et
iconibus illustrati. Ser. 2, fasc. 11–13: 47–81, pl. 151–195. Zippel, Trento,
Italy
Bresadola G. 1900. Fungi Tridentini novi, vel nondum delineati, descripti, et
iconibus illustrati. Ser. 2, fasc. 14: 83–118, pl. 196–217. Zippel, Trento, Italy.
Bresadola G. 1933. Iconographia mycologica. Vol. XXV. Trento, Italy.
Bulliard JBF. 1791. Herbier de la France. Tome 11. Paris, France.
Cao JZ, Fan δ, δiu B. 1990. Some species of Otidea from China. εycologia
82: 734–741.
Carbone ε. 2009. Il genere Otidea I. Sull’identità di Peziza onotica. Rivista
di εicologia 52: 11–28.
Carbone ε. 2010a. Il genere Otidea III. Identità e tipifizione di Peziza
alutacea. Bollettino dell’Associazione εicologica ed Ecologica Romana
80–81: 22–38.
Carbone ε. 2010b. Il genere Otidea IV. Prima parte. Otidea caligata, l’attuale
nome di Otidea abietina sensu Breitenbach & Kränzlin, con discussione
sull’abbandono dell’epiteto abietina. Schweizerische Zeitschrift für Pilzkunde 88: 14–17.
Carbone ε. 2010c. Il genere Otidea IV. Seconda parte. Otidea caligata,
l’attuale nome di Otidea abietina sensu Breitenbach & Kränzlin, con discussione sull’abbandono dell’epiteto abietina. Schweizerische Zeitschrift
für Pilzkunde 88: 64–66.
Carbone ε, Campo E, Vauras J. 2010. Records on Otidea mirabilis and
O. tuomikoskii from Finland. Karstenia 50: 25–34.
Carbone ε, Van Vooren N. 2010 ‘2009’. Il genere Otidea – II. Otidea fuckelii,
una nuova specie pubblicata per chiarire le differenti interpretazioni di
O. leporina. Rivista di εicologia 52: 313–330.
Cash EK. 1953. A record of the fungi named by J.B. Ellis. Part. II. The division
of mycology and disease survey, special publication 2: 167–345.
Clements FE, Shear Cδ. 1931. The genera of fungi. Wilson, New York, USA.
Cooke εC. 1876. εycographia seu Icones Fungorum. Part. 3. Williams &
Norgate, δondon, Great Britain.
Cooke εC. 1878. εycographia seu Icones Fungorum. Part. 5. Williams &
Norgate, δondon, Great Britain.
Dennis RWG. 1978. British Ascomycetes. 2nd edn. Cramer, Vaduz, Germany.
Dissing H. 2000. Pezizales Bessey. In: Hansen δ, Knudsen H (eds), Nordic
εacromycetes. Vol. 1. Ascomycetes: 55–127. Nordsvamp, Copenhagen,
Denmark.
Eckblad F-E. 1968. The genera of operculate Discomycetes. A reevaluation
of their taxonomy, phylogeny and nomenclature. Norwegian Journal of
Botany 15: 1–191.
Farr εδ, δeeusink JA, Stafleu FA. 1979. Index Nominum Genericorum
(Plantarum). Regnum Vegetabile vols. 100 –102. Bohn, Scheltema &
Holkema, Utrecht.
Franchi P, δami δ, εarchetti ε. 1999. Helvella leporina, nome corretto per
Helvella silvicola. Rivista di εicologia 42: 63–72.
Fries Eε. 1822. Systema mycologicum. II. Officina Berlingiana, Sweden.
Fuckel δ. 1870 ‘1869–1870’. Symbolae mycologicae. Jahrbücher des Nassauischen Vereins für Naturkunde 23–24: 1–459.
Gal ε le. 1947. Recherches sur les ornamentations sporales des Discomycètes operculés. Bibliotheca εycologica 28: 73 – 297. Reprint 1970.
Cramer, New York, USA.
Gonnermann W, Rabenhorst δ. 1869. εycologia Europaea. Abbildungen
sämmtlicher Schwämme Europas. 3. Dresden, Germany.
Gray SF. 1821. A natural arrangement of British plants. Baldwin, Cradock &
Joy, δondon, United Kingdom.
Greuter W, Brummitt RK, Farr E, et al. (eds). 1993. Names in current use
for extant plant genera. Regnum Vegetabile vol. 129. Koeltz Scientific
Books, Koenigstein.
Häffner J, Winterhoff W. 1989. Rezente Ascomycetenfunde VI. Otidea apophysata (Cooke & Phill.) Sacc. ein extrem seltener Öhrling. Beiträge zur
Kenntnis der Pilze εitteleuropas 5: 175–184.
Hansen K, δæssøe T, Pfister DH. 2002. Phylogenetic diversity in the core
group of Peziza inferred from ITS sequences and morphology. εycological
Research 106: 879–902.
Hansen K, δoBuglio KF, Pfister DH. 2005. Evolutionary relationships of the
cup-fungus genus Peziza and Pezizaceae inferred from multiple nuclear
genes: RPB2, ß-tubulin, and δSU rDNA. εolecular Phylogenetics and Evolution 36: 1–23.
Hansen K, Olariaga I. 2015. Species limits and relationships within Otidea
inferred from multiple gene phylogenies. Persoonia 35: 148–165.
228
Hansen K, Perry BA, Dranginis AW, et al. 2013. A phylogeny of the highly
diverse cup-fungus family Pyronemataceae (Pezizomycetes, Ascomycota)
clarifies relationships and evolution of selected life history traits. εolecular
Phylogenetics and Evolution 67: 311–335.
Hansen K, Pfister DH. 2006. Systematics of the Pezizomycetes - the operculate discomycetes. εycologia 98: 1029–1040.
Hansen K, Pfister DH, Hibbett DS. 1999. Phylogenetic relationships among
species of Phillipsia inferred from molecular and morphological data.
εycologia 91: 299–314.
Harmaja H. 1974. Flavoscypha, a new genus of the Pezizales for Otidea
cantharella and O. phlebophora. Karstenia 14: 105–108.
Harmaja H. 1976. New species and combinations in the genera Gyromitra,
Helvella and Otidea. Karstenia 15: 29–32.
Harmaja H. 1986. Studies on Pezizales. Karstenia 26: 41–48.
Harmaja H. 2009a. Studies in Otidea (Pezizales). Karstenia 48: 33–48.
Harmaja H. 2009b. A note on Otidea (Pezizales, Fungi). Phytotaxa 2: 49–50.
Hennings P. 1903. Beiträge zur Pilzflora Südamerikas. II. Hedwigia 36:
190–246.
Huelsenbeck JP, δarget B, Alfaro εE. 2004. Bayesian phylogenetic model
selection using reversible jump εarkov chain εonte Carlo. εolecular Biology and Evolution 21: 1123–1133.
Huhtinen S. 1990 ‘1989’. A monograph of Hyaloscypha and allied genera.
Karstenia 29: 45–252.
Jamoni PG. 2001. Validazione di nuovi taxa. Funghi e Ambiente 85–86: 56.
Jamoni PG. 2004. I funghi dell’ambiente alpino – XVIII. Funghi e Ambiente
94–95: 5–19.
Kanouse B. 1949. Studies in the genus Otidea. εycologia 41: 660–677.
Karsten PA. 1869. εonographia Pezizarum fennicarum. Notiser ur Sallskapets pro Fauna et Flora Fennica Forhandlingar 10: 99–206.
Karsten PA. 1871. εycologia Fennica. Pars prima. Discomycetes. Bidrag till
Kannedom of Finlands Natur och Folk 19: 1–263.
Kasparek F. 2000. Über einige bemerkenswerte Schlauchpilze. Der Tintling
19: 7–15.
Kimbrough JW. 1966. Studies in the Pseudoascoboleae. Canadian Journal
of Botany 44: 685–704.
Kirk Pε, Stalpers JA, Braun U, et al. 2013. A without-prejudice list of generic
names of fungi for protection under the International Code of Nomenclature
for algae, fungi, and plants. IεA Fungus 4: 381–443.
Korf RP. 1963. Discomycete flora of Asia, precursor II: A revision of the genera Acervus and Ascosparassis and their new positions in the Pezizales.
δloydia 26: 21–26.
Korf RP. 1972. Synoptic key to the genera of the Pezizales. εycologia 64:
937–994.
Korf RP. 1973a. Sparassoid ascocarps in the Pezizales and Tuberales.
Reports of the Tottori εycological Institute 10: 389–403.
Korf RP. 1973b. Discomycetes and Tuberales. In: Ainsworth GC, Sparrow
FK, Sussman S (eds), The fungi: An advanced treatise. Vol. 4A: 249–319.
Academic Press, New York, USA.
Korf RP, Zhuang WY. 1991. A preliminary discomycete flora of εacaronesia:
part 15, Terfeziaceae, and Otideaceae, Otideoideae. εycotaxon 40: 413–
433.
Kornerup A, Wanscher JH. 1961. Farver i farver. Politikens forlag, København, 1st edn.
Kumar S, Skjæveland Å, Orr RJS, et al. 2009. AIR: A batch-oriented web
program package for construction of supermatrices ready for phylogenomic
analyses. BεC Bioinformatics 10: 357.
δiu B, Cao JZ. 1987. Otideopsis yunnanensis gen. et sp. nov. of Pezizales
from China and its position in Pezizales system. Journal of Shanxi University, Natural Science edition 4: 70–73.
δiu CY, Zhuang WY. 2006. Relationships among some members of the genus Otidea (Pezizales, Pyronemataceae). Fungal Diversity 23: 181–192.
δundell S, Nannfeldt JA. 1934. Fungi Exsiccati Suecici, praesertim upsalienses Fasc. 1–2. Almqvist & Wiksells Boktryckeri, Sweden.
δundell S, Nannfeldt JA. 1938. Fungi Exsiccati Suecici, praesertim upsalienses. Fasc. 11–12. Almqvist & Wiksells Boktryckeri, Sweden.
δundell S, Nannfeldt JA, Holm δ. 1985. Fungi Exsiccati Suecici, prasertim
upsalienses. Fasc. 66 (No 3251–3300). Publications from the Herbarium,
University of Uppsala, Sweden 18: 1–18.
εaas Geesteranus RA. 1967. De fungi van Nederland 2a. Pezizales – deel
1, Discinaceae, Helvellaceae, εorchellaceae, Pezizaceae, Rhizinaceae.
Wetenschappelijke mededelingen van de Koninklijke Nederlandse Natuurhistorische Vereniging 69: 1–72.
εcNeill J, Barrie FR, Buck WR, et al. (eds). 2012. International Code of
Nomenclature for algae, fungi, and plants (εelbourne Code) adopted by
the Eighteenth International Botanical Congress εelbourne, Australia, July
2011. [Regnum Vegetabile No. 154.] Koeltz Scientific Books, Königstein,
Germany.
Persoonia – Volume 35, 2015
εedardi G. 1995. Considerazioni sul genere Otidea Fuck., 1870. Bolletino
del Circolo εicologico ‘G. Carini’ 29–30: 23–32.
εiller εA, Pfeiffer W, Schwartz T. 2010. Creating the CIPRES Science
Gateway for inference of large phylogenetic trees. In: Proceedings of the
Gateway Computing Environments Workshop (GCE), 14 Nov.: 1–8. New
Orleans, δA, USA.
εoravec J. 1986. A new species and two new combinations in the genus
Sowerbyella. εycologia Helvetica 2: 93–102.
εoravec J. 1988. Sowerbyella angustispora spec. nov. and Otideopsis kaushalii comb. nov. (Discomycetes, Pezizales, Pyronemataceae). εycologia
Helvetica 3: 135–142.
εoravec J. 2005. A world monograph of the genus Cheilymenia (Discomycetes, Pezizales, Pyronemataceae). δibri Botanici 21: 3–256.
εornand J, Courtecuisse R. 2005. δe genre Otidea et espèces affines en
France. Bulletin mensuel de la Société linnéenne de δyon 74, numero
spécial: 65–84.
Nannfeldt JA. 1937. Contributions to the mycoflora of Sweden. 4. On some
species of Helvella, together with a discussion of the natural affinities within
the Helvellaceae and Pezizaceae trib. Acetabuleae. Svensk Botanisk
Tidskrift 31: 47–66.
Nannfeldt JA. 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 Fuckel emend. Boud. and Sowerbyella Nannf. n.
gen. Svensk Botanisk Tidskrift 32: 108–120.
Nannfeldt JA. 1966. On Otidea caligata, O. indivisa and O. platyspora (Discomycetes, Operculatae). Annales Botanici Fennici 3: 309–318.
Nei ε. 1987. εolecular evolutionary genetics. Columbia University Press, USA.
Parslow ε, Spooner B. 2013. The British species of Otidea: overview and
the large spored species. εycosystema 32: 347–365.
Pérez-Butrón Jδ, Fernández-Vicente J. 2008. Otidea apophysata, en la
Península Ibérica. Errotari 5: 36–43.
Perry BA, Hansen K, Pfister DH. 2007. A phylogenetic overview of the
family Pyronemataceae (Ascomycota, Pezizales). εycological Research
111: 549–571.
Persoon CH. 1799. Observationes εycologicae. Pars secunda. δeipzig &
δuzern.
Persoon CH. 1822. εycologia europaea. I. Erlangae, Germany.
Peterson ET. 1998. Systematics of the genus Otidea in the Pacific Northwest.
Unpublished thesis. Oregon State University.
Pfister DH. 1979. A monograph of the genus Wynnea (Pezizales, Sarcoscyphacea). εycologia 71: 144–159.
Pfister DH, Halling RE. 1989. Ascosparassis heinricheri from Venezuela: an
extended distribution. εycotaxon 35: 283–285.
Quélet δ. 1886. Enchiridion fungorum in Europa media et praesertim in Gallia
vigentium. Douin, Paris, France.
Rahm E. 1958. Otidea pusilla nov. spec. Zwerg-Öhrling. Schweizerische Zeitschrift für Pilzkunde 36: 33–35.
Rambaut A. 2002. Se-Al. Sequence Alignment Editor. Version 2.0 alpha 11.
University of Oxford, Oxford. From http://tree.bio.ed.ac.uk/software/seal/.
Ramsbottom J. 1914. A list of the British species of Discomycetes arranged
according to Boudier’s system, with a key to the genera. Transactions of
the British εycological Society 4: 343–381.
Rehm H. 1883. Ascomyceten XIV. Hedwigia 22: 1–18 (reprint).
Rifai εA. 1968. The Australasian Pezizales in the herbarium of the Royal
Botanic Gardens Kew. Verhandelingen der Koninklijke Nederlandse Akademie van Wetenschappen, Afd. Natuurkunde, Tweede Reeks 57: 1–295.
Ronquist F, Teslenko ε, εark P van der, et al. 2012. εrBayes 3.2: Efficient
Bayesian phylogenetic inference and model choice across a large model
space. Systematic Biology 61: 539–542.
Saccardo PA. 1884. Conspectus generum discomycetum hucusque cognitorum. Botanisches Centralblatt 18: 213–256.
Saccardo PA. 1889. Sylloge Fungorum omnium hucusque cognitorum 8.
Patavii.
Schaeffer JC. 1763. Fungorum qui in Bavaria et Palatinatu circa Ratisbonam
nascuntur icons natives coloribus expressae. Tomus II. Regensburg, Germany.
Schaeffer JC. 1774. Fungorum qui in Bavaria et Palatinatu circa Ratisbonam
nascuntur icons natives coloribus expressae. Tomus IV. Regensburg, Germany.
Seaver FJ. 1904. The discomycetes of eastern Iowa. Bulletin from the laboratories of natural history of the State University of Iowa 5: 230–297.
Seaver FJ. 1927. A tentative scheme for the treatment of the genera of the
Pezizaceae. εycologia 19: 86–89.
Seaver FJ. 1928. The North American cup-fungi (operculates). Reprint 1961.
Hafner Publishing Company, New York, USA.
Smith εE, Healy RA. 2009. Otidea subterranea sp. nov.: Otidea goes below
ground. εycological Research 113: 858–866.
229
I. Olariaga et al.: A monograph of Otidea
Sowerby J. 1797. Coloured figures of English fungi or mushrooms. Vol. I.
δondon, England.
Spooner Bε, Yao Y-J. 1995. Notes on British taxa referred to Aleuria. εycological Research 99: 1515–1518.
Stamatakis A. 2006. Raxml-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics
22: 2688–2690.
Svrček ε. 1976. A taxonomic revision of Velenoský’s types of operculate
discomycetes (Pezizales) preserved in national museum, Prague. Sborník
národního muzea v Praze. Acta εusei Nationalis Pragae 32 B: 115–194.
Taylor JW, Jacobson DJ, Kroken S, et al. 2000. Phylogenetic species recognition and species concepts in fungi. Fungal Genetics and Biology 31: 21–32.
Thiers B (ed). 2014. Index Herbariorum. A global directory of public herbaria
and associated staff. New York Botanical Garden´s Virtual Herbarium. http://
sweetgum.nybg.org/ih/.
Van Vooren N. 2010. Note sur Otidea mirabilis (Pezizales). Ascomycete.
org 2: 33–35.
Van Vooren N. 2011a. δe genre Otidea V. Otidea apophysata et ses interprétations. Bulletin mycologique et botanique Dauphiné-Savoie: 165–174.
Van Vooren N. 2011b. Premiers signalements d’Otidea caligata (Nyl.) Sacc.
(Ascomycota, Pezizales) en France. Ascomycete.org 3: 61–64.
Van Vooren N, Armada F. 2011. Redécouverte d’Otidea platyspora Nannf.
(Ascomycota, Pezizales) en France. Bulletin mycologique et botanique
Dauphiné-Savoie 203: 57–62.
Van Vooren N, Carbone ε. 2012. The genus Otidea. VI. Otidea felina and its
interpretations. Ascomycete.org 4: 29–34.
Van Vooren N, Hairaud ε, Jindřich O. 2008. Otidea tuomikoskii, Otidea papillata et Otidea papillata f. pallidefurfuracea f. nov. trois taxons remarquables
appartenant au genre Otidea (Pezizales, Pyronemataceae). Bulletin mycologique et botanique Dauphiné-Savoie 188: 47–57.
Van Vooren N, Olariaga I, Tabarés ε. 2011. First record of Otidea caeruleopruinosa Harmaja (Ascomycota, Pezizales) in the Iberian Peninsula.
Ascomycete.org 3: 43–46.
Yao Y-J, Spooner Bε. 2006. Species of Sowerbyella in the British Isles, with
validation of Pseudoombrophila sect. Nannfeldtiella (Pezizales). Fungal
Diversity 22: 267–279.
Zhuang WY. 2006 ‘2005’. Notes on Otidea from Xinjiang, China. εycotaxon
94: 365–370.
Zhuang WY. 2010. Taxonomic assessment of some pyronemataceous fungi
from China. εycotaxon 112: 31–46.
Zhuang WY, Korf RP. 1987. Some new species and new records of discomycetes in China. II. εycotaxon 29: 309–314.
Zhuang WY, Korf RP. 1989. Some new species and new records of discomycetes in China. III. εycotaxon 35: 297–312.
Zhuang WY, Yang Zδ. 2008 ‘2007’. Some pezizalean fungi from alpine areas
of southwestern China. εycologia εontenegrina 10: 235–249.
INDEX to species, varieties and forms
Accepted names treated in the monograph are in bolditalic.
abietina 166, 167, 193, 195, 197, 224
abietina var. grandis 225
abietina f. / var. nigra 195
alba 185, 224
alutacea 175–177, 179, 181, 182, 224, 225
alutacea var. microspora 226
angusta 176, 198, 200
apophysata 167, 168, 178, 179, 180–182
atrofusca 226
aurantia 223, 224
aurantia var. atromarginata 224
aurantia var. stipitata 224
auricula 167, 185, 187, 226
auriculariiformis 224
azureoides 227
badiofusca 226
bicolor 178, 211
borealis 168, 171, 174, 176–178, 212, 213
boudieri 180
brevispora 174, 179, 205
brunneoparva 168, 179, 187, 191
bufonia 175, 176, 179, 185, 192, 205, 206, 209,
211, 213, 219, 224, 225
caeruleopruinosa 168, 174, 177, 179, 213,
215, 219
caligata 189, 193, 195, 197
cantharella 167, 178, 189, 191, 192, 193, 197,
216, 219
cantharella var. minor 188, 189, 219, 220
cinerascens 185, 224
cochleata 166, 167, 180, 183–185, 195, 206,
207, 223, 224
cochleata var. alutacea 182
cochleata var. umbrina 206, 175
concinna 167, 168, 174, 175, 177, 178, 189,
210, 213, 216, 219, 223
crassa 174, 185, 187
daliensis 167, 168, 178–180, 181, 192, 197
darjeelensis 224, 225
dochmia 224
domingensis 225
doratophora 225
euplecta 225
felina 179, 180, 185, 187, 225
fibrillosa 225
flavidobrunneola 177, 179, 213, 216, 217
formicarum 168, 171, 174, 176, 177, 179, 189,
197, 200, 201
fuckelii 185, 187
fusconigra 192, 205
grandis 205, 207, 208, 209, 225
grandis var. scheremetjeffii 225
harperiana 225
hirneoloides 225
indivisa 195–197
integra 168, 174, 178, 223
kauffmanii 171, 175, 213, 214, 223
kaushalii 174, 176, 177, 178, 202, 205
kunmingensis 178, 184, 185
lactea 174, 178, 179, 213, 219
lechria 225
leporina 166, 167, 171, 174, 175, 177, 179, 185,
188, 189, 192, 209–211
leporina f. major 185
leporina f. / var. minor 185, 195
leporina var. onotica 209
leporina var. rubescens 185
lilacina 225
lobata 225
lohjaënsis 200
luculenta 226
luteonitens 226
marsupium var. pyxidata 216
micropus 226
microspora 226
minor 174, 177, 178, 189, 213, 217, 219, 222
mirabilis 171, 174–176, 179, 192, 205, 207,
209, 211, 213
myosotis 185, 187
nannfeldtii 168, 175–177, 179, 185, 189, 191,
197, 198, 201, 202
neglecta 226
obtecta 226
ochracea 226
olivacea 192, 226
olivaceobrunnea 178, 192
onotica 166, 167, 174, 177–179, 205, 209,
225, 226
onotica var. brevispora 174, 205, 206
onotica var. ochracea 226
oregonensis 168, 174, 177, 178, 213, 217,
219, 220, 223
papillata 175, 179, 185, 189, 191, 198, 200
papillata f. pallidefurfuracea 189
pedunculata 206, 207, 209
phlebophora 167, 168, 174, 178, 213, 217, 219,
220, 222, 225
phyllogena 225
platyspora 178, 179, 180, 182
pleurota 226
praetervisa 227
propinquata 167, 177, 178, 191–193, 195, 224
pseudobadia 206, 209
pseudoleporina 168, 174, 176, 177, 179, 185,
188, 197, 200, 201
purpurea 179, 205, 211
pusilla 192
pyxidata 216
radiculata 226
rainierensis 174, 175, 179, 213, 214, 217, 222,
223, 226
reisneri 226
rhenana 224
rosea 209
schulzeri 226
scutellata 216
shimizuensis 167
silvicola 167, 185, 187, 226
sinensis 174, 178, 213
smithii 168, 171, 176, 179, 192, 205, 209, 211
sparassis 226
subconcinna 219
subformicarum 168, 171, 174, 179, 197, 198,
200
subonotica 226
subpurpurea 178, 211
subterranea 167, 178, 179, 181
subviolacea 227
succosa 227
tasmanica 227
tianshuiensis 178, 213
tuomikoskii 171, 177, 179, 185, 189, 200, 202
umbrina 184, 187, 206–208
unicisa 167, 168, 174, 177, 178, 202, 205,
211, 225
varia 226
violacea 227
yunnanensis 167, 174, 178, 179, 202, 204, 205
Persoonia 35, 2015: 166 – 229
www.ingentaconnect.com/content/nhn/pimj
RESEARCH ARTICLE
http://dx.doi.org/10.3767/003158515X688000
A monograph of Otidea (Pyronemataceae, Pezizomycetes)
I. Olariaga1, N. Van Vooren 2, ε. Carbone 3, K. Hansen1
Key words
Flavoscypha
ITS
ITS1 minisatellites
LSU
Otideopsis
resinous exudates
Abstract The easily recognised genus Otidea is subjected to numerous problems in species identification. A number
of old names have undergone various interpretations, materials from different continents have not been compared and
misidentifications occur commonly. In this context, Otidea is monographed, based on our multiple gene phylogenies
assessing species boundaries and comparative morphological characters (see Hansen & Olariaga 2015). All names
combined in or synonymised with Otidea are dealt with. Thirty-three species are treated, with full descriptions and
colour illustrations provided for 25 of these. Five new species are described, viz. O. borealis, O. brunneoparva, O. oregonensis, O. pseudoleporina and O. subformicarum. Otidea cantharella var. minor and O. onotica var. brevispora
are elevated to species rank. Otideopsis kaushalii is combined in the genus Otidea. A key to the species of Otidea
is given. An δSU dataset containing 167 sequences (with 44 newly generated in this study) is analysed to place
collections and determine whether the named Otidea sequences in GenBank were identified correctly. Fourty-nine
new ITS sequences were generated in this study. The ITS region is too variable to align across Otidea, but had low
intraspecific variation and it aided in species identifications. Thirty type collections were studied, and ITS and δSU
sequences are provided for 12 of these. A neotype is designated for O. cantharella and epitypes for O. concinna,
O. leporina and O. onotica, along with several lectotypifications. The apothecial colour and shape, and spore characters are important for species identification. We conclude that to distinguish closely related or morphologically
similar species, a combination of additional features are needed, i.e. the shape of the paraphyses, ectal excipulum
structure, types of ectal excipulum resinous exudates and their reactions in εelzer’s reagent and KOH, tomentum
and basal mycelium colours and exudates. The KOH reaction of excipular resinous exudates and basal mycelium
are introduced as novel taxonomic characters.
Article info Received: 26 November 2013; Accepted: 1 February 2015; Published: 10 April 2015.
INTRODUCTION
Species of Otidea produce typically ear-shaped apothecia that
are unique within Pyronemataceae (Pezizomycetes). The genus
is monophyletic based on multilocus phylogenetic analyses
from a few, but broadly sampled, Otidea species (Hansen
et al. 2013). Despite being distinct at the generic level, the
species identification and nomenclature of Otidea are highly
controversial. A few recent typifications have been proposed
(Carbone 2009, 2010a), but many names are still subjected to
different interpretations. Several new species were described
in the last decades from Europe (Harmaja 1976, 2009a) and
Asia (Cao et al. 1990, Zhuang & Yang 2008), with detailed
descriptions and updated identification keys. However, often
no illustrations were presented and colour photographs have
rarely been published when describing new species. εany
names of European species currently used in North America
and Asia are misapplied. εultilocus phylogenetic analyses have
not been previously implemented to critically address species
delimitation issues and material from different continents has
not been compared. A worldwide critical revision of Otidea to
clarify species limits is highly needed. The aims of this study
were: i) to undertake a nomenclatural and taxonomic revision
of Otidea, to clarify misinterpretations and to propose pertinent
typifications to stabilise the use of names; and ii) to provide
detailed species descriptions and colour photographs of both
macro- and microscopic structures, and a key for identification.
1
2
3
Swedish εuseum of Natural History, Department of Botany, P.O. Box 50007,
SE-104 05 Stockholm, Sweden;
corresponding author e-mail: karen.hansen@nrm.se.
36 rue de la Garde, F-69005 δyon, France.
Via Don δuigi Sturzo 173 I-16148 Genova, Italy.
Our multilocus phylogenies and robust hypotheses of species
limits, employing genealogical concordance phylogenetic species recognition (GCPSR; Taylor et al. 2000), which is the basis
for the present work, are given in Hansen & Olariaga (2015).
In the present study we present an δSU rDNA phylogeny to
place a larger number of collections for which we have been
unable to obtain multiple genes, including several sequences
from GenBank, many of which we here re-identify.
Taxonomic history
The first valid publication of Otidea is by Bonorden (1851),
based on Peziza (unranked) Otidea Pers., although it has sometimes been attributed to Fuckel (Kanouse 1949, Nannfeldt
1966, δiu & Zhuang 2006, Smith & Healy 2009). Otidea species
were treated in a broad heterogeneous genus Peziza by early
authors. Persoon (1822) defined Peziza (unranked) Otidea
as producing auriculate apothecia with a split, sometimes
elongated on one side, and included 10 species. Fries (1822)
referred to this group as Peziza (unranked) Cochleatae, but he
included also taxa with non-split apothecia. Bonorden (1851)
elevated Otidea to generic rank with split apothecia as the key
feature, but also referred to Fries’ (1822) Cochleatae. He did
not make any combinations or list any species. Fuckel (1870)
refined the genus using microscopic details, namely uni- or
biguttulate spores and filiform to subclaviform paraphyses, and
included four species: O. abietina, O. cochleata O. leporina
and O. onotica. Boudier (1885) notably contributed to disassemble the large genus Peziza into smaller and more natural
genera. He placed in the genus Otidea species with entire or
split apothecia, biguttulate spores and, importantly non-amyloid
asci and curved paraphyses, which he was the first to introduce. He divided Otidea into two subgenera: Otidea with split
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167
I. Olariaga et al.: A monograph of Otidea
apothecia and Pseudotis with entire apothecia. Interestingly,
Boudier erected the genus Wynnella (Helvellaceae) to accommodate W. silvicola (as P. leporina / P. auricula), a species
with distinctly ear-shaped apothecia, but differing from Otidea
in the uniguttulate spores and tough consistency. In Boudier’s
(1907) subsequent treatment, he elevated Pseudotis to genus
rank and placed here species with entire apothecia, including
O. daliensis (as P. apophysata) and O. propinquata (as P. abietina), both with biguttulate spores and hooked paraphyses.
Saccardo listed O. onotica as an ‘exemplar’ species of Peziza
subg. Otidea in his synopsis of the discomycete genera (1884;
he listed in general only one species per genus / subgenus that
appear to have been selected as typical for the genera) and
it has since been accepted as the type species by most (Rifai
1968, Eckblad 1968, Korf 1972, δiu & Zhuang 2006, Parslow &
Spooner 2013), Index Nominum Genericorum (Eckblad in Farr
et al. 1979), NCU-3 (Greuter et al. 1993) and is prepared to be
adopted on the δist of Protected generic Names for fungi (Kirk
et al. 2013). Clements & Shear (1931) listed O. cochleata as
the type, but this name was not among the original species accepted by Persoon (1822) and has furthermore been considered
an ambiguous name. Kanouse (1949) proposed O. leporina as
the type species, but because of the confusions surrounding
the identity of this species until now, it has been considered an
inappropriate choice.
The genus Scodellina was described by Gray (1821) and involved species attributed today to Otidea, and also Aleuria,
Peziza and Tarzetta. Seaver (1928) refined Scodellina to
species with split to ear-shaped apothecia only and typified it
with Peziza leporina (Seaver 1927), considering Otidea a later
synonym. This was for a period followed by several American
authors (e.g. Korf 1963, Kimbrough 1966). The typification by
Seaver can however, be considered largely mechanical, taken
as the first species listed by Gray, and be superseded under
the ICN (Art. 10.5; εcNeill et al. 2012). Also, even though
Gray (1821) included several species with split apothecia, he
did not mention this feature in the diagnosis of Scodellina, but
emphasised “thallus … hemispherical, spreading” and coined
the vernacular name “spread cup”. Therefore Rifai (1968) designated P. vesiculosa Bull.: Fr. as the type species of Scodellina,
consequently making it a later synonym of the genus Peziza.
Eckblad (1968) came to the same conclusion.
Kanouse (1949) broadened the concept of Otidea and described in detail a number of North American species. She
included species with split apothecia and straight paraphyses
with swollen apices. She also included Wynnella silvicola (as
Otidea auricula), with straight paraphyses and uniguttulate
spores. Nannfeldt (1966) delimited Otidea to species with
non-amyloid asci; smooth, uninucleate, biguttulate spores; a
medullary excipulum of textura intricata; and an ectal excipulum with isodiametric cells, covered by short chains of barrelshaped cells. Nannfeldt’s concept more or less conforms to the
genus Otidea as we recognise it today. He considered Wynnella so distant that it should be treated in a separate tribe of
Pezizaceae. Korf (1963) reviewed the monotypic, sparassoid
genus Ascosparassis, and subsequently (Korf 1973a) assigned
A. shimizuensis to Otidea, based on the hooked paraphyses,
small biguttulate spores and excipulum structure. Pfister (1979)
however, considered Ascosparassis a distinct monotypic genus,
based on “small asci and spores and peculiar growth habit”, and
combined the older name Midotis heinricheri in Ascosparassis. δater Pfister collected A. heinricheri in South America, in
north coastal mountains of Venezuela (Pfister & Halling 1989),
extending its Asian distribution (China, Indonesia and Japan),
still considering the species separate from Otidea.
A new genus Flavoscypha was erected for two species of
Otidea, O. concinna (as Flavoscypha cantharella) and O. phlebophora, with strong emphasis on the ectal excipulum of textura
prismatica (vs textura angularis in Otidea) (Harmaja 1974).
Otidea was further emended to include a species with ornamented spores, O. unicisa, otherwise ‘fitting perfectly’ Otidea
(Harmaja 1986). Otideopsis was published with Otideopsis
yunnanensis as the type species, distinguished from Otidea
by having ornamented spores and paraphyses fused at the
apices (δiu & Cao 1987). Flavoscypha and Otideopsis are now
considered synonyms of Otidea based on molecular phylogenetic analyses (δiu & Zhuang 2006, Hansen & Olariaga 2015).
Recently the circumscription of Otidea was further broadened
when the first hypogeous species, O. subterranea, was discovered using ITS and δSU sequences (Smith & Healy 2009). All
the characters proposed so far as diagnostic for Otidea have exceptions across the genus. Nevertheless, Otidea can be recognised by the non-amyloid asci, in combination with at least
two of these characters (except O. subterranea): a) biguttulate
spores; b) hooked or bent paraphyses; c) medium-large, split
apothecia; and d) a medullary excipulum of textura intricata, and
an ectal excipulum of textura angularis or textura prismatica.
Systematic position and relationships
Nannfeldt (1937, 1938, 1966) suggested a close relationship
between Otidea (incl. Pseudotis), Tarzetta (as Pustularia) and
Helvella (Pezizaceae, tribe Acetabuleae sensu Nannfeldt),
based on similarities in asci, paraphyses and anatomical structures of the apothecia, i.e. a medullary excipulum of dense
textura intricata, ectal excipulum of almost isodiametric large
cells, and an outermost layer of shorter or longer chains of
cells, possible forming distinct clusters or warts (and in Tarzetta
prolonged to cylindrical, hyaline, wavy hairs). δe Gal (1947)
similarly placed Otidea in the tribe Otideeae in her Aleuriaceae
(i.e. a family including taxa with both amyloid and non-amyloid
asci), together with Pseudotis and Tarzetta (as Pustularia), but
placed Helvella in Helvellaceae. Following the ideas of Nannfeldt (1966), Eckblad (1968) erected the family Otideaceae
as a small taxon of closely related genera that produce larger
apothecia, most of which typically lack bright orange to red colours, including in it Tarzetta (as Pustulina) and Otidea, but also
Ascosparassis, Geopyxis and Sowerbyella. Eckblad considered
Helvella (Helvellaceae) to be distant, but having a possible
shared origin with members of Otideaceae (and Morchellaceae
and Rhizinaceae), due to the structure of the excipulum, spores
and asci, especially of Tarzetta. At the same time, he expanded
the concept of Pyronemataceae (to 21 genera) to taxa mostly
characterised by the presence of carotenoid pigments, stating
the inability to satisfactorily subdivide the family on the basis
of common characters. Korf (1972, 1973b) placed Otidea in
the tribe Otideeae (in the subfamily Otideoideae) in an even
more encompassing Pyronemataceae (49 genera), together
with Ascosparassis and Psilopezia. He followed the ideas
of Arpin (1969), in excluding taxa with carotenoids from the
Otideoideae, instead including taxa with prominent hairs such
as Geopora, Humaria and Trichophaea. Otidea has generally
been included in a broadly circumscribed Pyronemataceae
in recent treatments (Dissing 2000, Hansen & Pfister 2006,
Perry et al. 2007, Hansen et al. 2013). εultigene phylogenetic
analyses of Pyronemataceae do not support a close relationship between Otidea and Geopyxis, Psilopezia, Sowerbyella
or Tarzetta (Hansen et al. 2013). Surprisingly, the cleistothecial
Warcupia and the highly reduced (gymnohymenial) Monascella are suggested as the closest relatives of Otidea. Otidea,
Monascella and Warcupia are strongly supported as a distinct
sister group to the rest of the Pyronemataceae in a strict sense
(Hansen et al. 2013).
168
MATERIALS AND METHODS
Material and morphological methods
This study is based on a total of 450 specimens. One hundred
and forty two of these were collected and studied fresh during this project and are deposited in S and a few in TUR-A.
Specimens were studied from the following herbaria: AH, ARAN,
BIO, C, FH, H, HKAS, K, εCVE, εICH, εIN, OSC, PC, PRε,
S, TUR and UPS (Thiers 2014), GεFN (gruppo AεB di Fara
Novarese, Italy), SEST (Sociedad de Ciencias Naturales de
Sestao, Spain), and from the private herbaria of G. Corriol, GC;
C. δavorato, Cδ; N. Van Vooren, NV; and ε. Tabarés, εT. Thirty
type collections were examined, along with other original material. Colour codes are based on Kornerup & Wanscher (1961).
For O. apophysata, O. borealis, O. daliensis, O. oregonensis,
O. phlebophora, O. pseudoleporina, O. smithii and O. unicisa
colour codes for fresh material were taken from photographs.
Apothecial sections, i.e. the thickness of the apothecia including
the hymenium, was measured midway between the apothecial
margin and base. Smell and taste are listed when recorded.
Only discharged, mature spores were measured from living
material. To obtain and ensure mature spores from dried material, a square of c. 3 mm 2 of an apothecium was revived in
a drop of water on a slide, with the hymenium surface facing down and then removed after 1 hour. In this way mature
spores deposited on top of the hymenium were recovered and
measured. Spore measures exclude ornamentation. Spore
statistics are based on measurements of 20 spores from each
collection: Lm = mean length, Wm = mean width and Qm = Lm/
Wm. The number of populations that the statistics are based on
is indicated by ‘n’. Extreme values are given in parentheses.
Hymenial elements were observed by teasing apart a small
piece of hymenium with a needle. To observe the excipulum
structure, sections of apothecia were made by hand prior to
soaking the material in water. Basal mycelium was examined
by mounting clumps of hyphae from the apothecial base (the
tomentum) and from among the substrate particles. Only asci
with mature spores were measured. All measurements were
made in water: in living state whenever possible; only when
not possible, measurements were made on rehydrated (over
2 hours in water) material. εelzer’s reagent (εδZ) and 10 %
KOH were used to observe the reaction of resinous exudates
and other pigmentation. Cotton Blue in lactic acid was used to
observe spore ornamentation. εicroanatomical terminology
follows Korf (1973b). The notation ‘!’ indicates that type or other
original material was examined by us.
If not otherwise indicated in the legends, the photographs presented in this paper were taken by K. Hansen and I. Olariaga.
DNA extraction, PCR amplification, sequencing and
alignment
DNA was extracted from dried material, or from fresh material stored in 1 % SDS DNA extraction buffer. The extraction
method follows Hansen et al. (1999), except dried material was
ground in a εini-Beadbeater Tε (Biospec Products, Bartlesville,
OK, USA) and fresh material using a plastic pestle, in eppendorf tubes. The primer combination ITS5-ITS4, and in a few instances ITS1–ITS4, ITS5–5.8S and ITS3–ITS4, were used to
PCR amplify the ITS region, and δR0R –δR5 the δSU region.
For DNA extracted from fresh material (stored in SDS extraction buffer), the ITS and δSU regions were amplified in a
single piece using the primers ITS1– δR5. The ITS2 region
for O. integra (possible original material from 1892) was
successfully amplified in three pieces, in combination with
newly designed primers for the O. concinna clade: ITS3 –
ConcITS2midR (5´-GCCTGTAAATTTTAAAGACGAA-3´);
ConcITS2midF (5´-CCAGGGTTGCTTTGGTA-3´) – ConcITS4
Persoonia – Volume 35, 2015
intR (5´-CACTGGGTAATTGGAGGTTT-3´); ConcITS2midF
(5´-CCAGGGTTGCTTTGGTA-3´) – ITS4. PCR products were
cleaned using ExoSAP-IT® (USB, Cleveland, OH, USA). The
ITS was sequenced in both directions, using the primers ITS1
and ITS4 (and/or in a few instances ITS5, 5.8S and ITS3) and
the δSU using δR0R and δR5. For O. integra, the same primers as used for PCR, were also used for sequencing. PCR
and sequencing conditions follows Hansen & Olariaga (2015).
Sequences were edited and assembled using Sequencher v. 4.10
(Gene Codes Corporation Ann Arbor, εichigan, USA) and have
been deposited in GenBank (Table 1). The sequences were
aligned manually in Se-Al v. 2.0a11 Carbon (Rambaut 2002).
An all taxa δSU dataset was prepared. Monascella botryosa
and Warcupia terrestris were used as outgroup, based on a
higher level phylogenetic study of Pyronemataceae (Hansen
et al. 2013), which supports these as the closest sister group.
To explore inter- and intraspecific variation of the new species
O. borealis, O. subformicarum and their closest relatives, two
smaller datasets were prepared using ITS and δSU rDNA sequences, based on a more species-inclusive, multi-gene phylogeny of Otidea (Hansen & Olariaga 2015). All three alignments
are available from TreeBASE as accession no. S15887. The
first dataset (the O. borealis dataset) contained O. borealis and
related species in the O. concinna clade. The second dataset
(the O. formicarum dataset) contained specimens of the O. formicarum clade. Otidea caeruleopruinosa and O. nannfeldtii
were used as outgroup for each of these datasets. Nucleotide
diversity (e.g. Nei 1987, equation 10.6), as the average number
of nucleotide differences per sites between two ITS sequences,
was calculated within O. brunneoparva and O. subformicarum.
One insertion or deletion, despite the length, was calculated as
only one basepair difference.
Phylogenetic analyses
εaximum δikelihood (εδ) analyses of the all taxa δSU dataset were performed using the ‘RAxεδ HPC2 on XSEDE’ tool
(Stamatakis 2006) via CIPRES Science Gateway (εiller et al.
2010), employing mixed models of evolution and starting from a
random tree. For the two smaller O. borealis and O. formicarum
datasets, εδ analyses were conducted using RAxεδ v. 7.3.1
(Stamatakis 2006) on the Bioportal, University of Oslo (Kumar
et al. 2009). A GTR-GAεεA model with four rate categories
was assigned and all free model parameters were estimated
by the program. For the εδ bootstrap analyses (εδ-BP) 1 000
rapid bootstrapping replicates from random starting trees were
performed, followed by a subsequent εδ search similarly using
1 000 replicates. As no strongly supported conflict was detected
(εδ-BP ≥ 75 %, PP ≥ 95 %), the ITS and δSU region were concatenated for the O. borealis and O. formicarum datasets. Each
combined dataset was analysed using εδ analyses under the
same settings as specified above. Relationships were likewise
constructed using εetropolis-coupled εarkov Chain εonte
Carlo (εCεCεC) and ‘model-jumping’ as implemented in
εrBayes v. 3.2.1 (Ronquist et al. 2012). The substitution model
was sampled across the GTR space by the εCεC analysis
(Huelsenbeck et al. 2004). Four parallel searches, each with
four chains, were run for ten and three million generations,
respectively, for the all-inclusive LSU dataset and the O. borealis and O. formicarum datasets, initiated with random starting
trees. The chains were sampled every 100 generations from
the posterior distribution. The first 25 % of the trees sampled
was discarded as the ‘burn-in’, and the remaining trees were
used to calculate the posterior probabilities (PP) of the clades.
For the combined εδ and Bayesian analyses the ITS and
δSU regions were specified as distinct partitions. εδ bootstrap
values ≥ 70 % and PP ≥ 95 % were considered to be significant.
169
I. Olariaga et al.: A monograph of Otidea
Table 1 Collections used in the molecular phylogenetic analyses, with voucher information and GenBank accession numbers for ITS and δSU regions. Some
GenBank sequences are re-identified by us and the names originally used in GenBank are listed after the taxon names (‘as’). For type specimens (in bold) the
original names are kept regardless of synonymy. Numbers in parentheses following the species names indicate multiple collections of a species. The GenBank
accessions of sequences generated in this study are in bold.
Taxon
Voucher
δocality / year/ collector
ITS
δSU
Monascella botryosa
Otidea alutacea (1)
O. alutacea (2)
O. alutacea (3)
O. alutacea (4)
O. alutacea (5)
O. alutacea (6)
O. alutacea (7)
O. alutacea (8)
O. alutacea (9)
O. alutacea (10)
O. alutacea (11)
O. alutacea (12)
O. alutacea (13)
O. alutacea (14)
O. alutacea (15)
O. alutacea (16)
O. alutacea (17)
O. alutacea (18)
O. alutacea (19)
O. alutacea (20) as O. umbrina
O. alutacea (21) as O. umbrina
O. alutacea (22)
O. alutacea (23)
O. alutacea (24)
O. alutacea (25)
O. alutacea (26)
O. alutacea (27)
O. alutacea (28)
O. alutacea (29)
O. alutacea (30)
O. angusta
O. apophysata
CBS 233.85
KH.09.170 (S)
KH.10.193 (S)
KH.07.46 (S)
JS.08.81 (S)
OSC 56747
OSC 56770
OSC 56798
OSC 56777
JS.08.43 (S)
KH.09.135 (S)
KH.10.198 (S)
KH.09.178 (S)
KS-94-192 (C)
C-F-48045
HεAS52742
HεAS57844
S-F257085
εoorefun19 (OSC)
OSC 56758
OSC 56813
OSC 56782
KH.09.133 (S)
ARAN A3023204
GC 98092002
HεAS83560
HεAS83563
KS-94-111 (C)
HεAS83559
S-F257084
KH.13.50 (S)
H6010804
S-F257062, dupl. private herb.
Kasparek s.n.
S-F242694
HKAS 43003
Spain, 1985, J. Guarro
Sweden, 2009, K. Hansen & I. Olariaga
Sweden, 2010, K. Hansen, K. Gillen & I. Olariaga
Denmark, 2007, H. Knudsen
Sweden, 2008, J. Santos
USA, 1996, E.T. Peterson
USA, 1997, E.T. Peterson
USA, 1996, E.T. Peterson
USA, 1997, E.T. Peterson
Sweden, 2008, J. Santos
Norway, 2009, V. Kučera & I. Kautmanova
Sweden, 2010, K. Hansen, K. Gillen & I. Olariaga
Sweden, 2009, K. Hansen & I. Olariaga
Denmark, 1994, K. Hansen & S.K. Sandal
Sweden, 1974, D. Paulsen & N. Tams
China
China
Italy, 2010, ε. Carbone
USA, 2010, J. εoore
USA, 1996, E.T. Peterson
USA, 1997, E.T. Peterson
USA, 2010, E.T. Peterson
Norway, 2009, K. Hansen & I. Olariaga
Spain, 2009, J.I. δópez Amiano
France, 1998, G. Corriol
China, 2003, W.Y. Zhuang & Y. Nong
China, 2003, W.Y. Zhuang & Y. Nong
Denmark, 1994, K. Hansen & S.K. Sandal
China, 2003, W.Y. Zhuang & Y. Nong
Italy, 2010, ε. Carbone
Sweden, 2013, K. Hansen & X. Wang
Finland, 1965, H. Harmaja
Germany, 1999, F. Kasparek
–
Kε0100591
Kε0100601
KM010061
Kε0100621
–
–
–
–
KM010063
Kε0100641
KM010065
Kε0100661
KM010067
KM010068
–
–
Kε0100691
Kε0100701
–
–
–
Kε0100711
Kε0100721
KM010073
–
–
KM010074
–
KM010075
KM010076
KF7175741
Kε0100771
KC012688
KC012691
Kε8231881
KM823457
Kε8231871
Kε8231891
AF072073
AF086583
AF086582
KM823458
Kε8231901
KM823459
Kε8231911
KM823460
KM823461
DQ443438
DQ443439
Kε8231921
Kε8231941
Kε8231931
AF086584
AF086586
Kε8231851
Kε8231861
KM823462
DQ443442
DQ443440
KM823463
DQ443441
KM823464
KM823465
Kε8231951
Kε8231961
Finland, 2010, ε. Carbone
China, 2003, Z.δ. Yang
Kε0100231
–
Kε8231971
DQ443450
KH.09.82 (S)
S-F249386 (Ex-H6017193)
S-F257086, dupl. TUR–A 198579
JS.08.66 (S)
KH.08.107 (S)
TUR-A 198582
KH.09.172 (S)
JS.08.55 (S)
KH.07.37 (S)
KH.09.248 (S)
KH.09.249 (S)
NV 2009.11.01 (S)
C-F-94240
H6010805
εT 10082601 (SCε, dupl. S)
KH.13.48 (S)
JS.08.18 (S)
JS.08.47 (S)
KH.09.125 (S)
NV 2008.09.16 (dupl. S)
JS.08.59 (S)
KH.09.183 (S)
KH.09.250 (S)
HMAS583571
HMAS57688
SEST-06081702
H6010806
H6010830
KH.09.153 (S)
H6003350
JS.08.63 (S)
H6003549
S-F244372 (dupl. O)
KH.11.104 (S)
GεFN 2293
S-F108342
AH21147 (εICH)
Sweden, 2009, K. Hansen & I. Olariaga
Finland, 1978, H. Harmaja
Finland, 2009, ε. Carbone
Sweden, 2008, J. Santos
Sweden, 2008, K. Hansen
Finland, 2011, ε. δahti
Sweden, 2009, K. Hansen & I. Olariaga
Sweden, 2008, J. Santos
Denmark, 2007, K. Hansen
Spain, 2009, J.δ. Teres & P.ε. Pasaban
France, 2009, J.δ. Teres
France, 2009, G. εoyne
Denmark, 2011, ε. Sasa
Finland, 1978, H. Harmaja
Spain, 2010, ε. Tabarés & S. Santamaría
Sweden, 2013, I.-δ. Walter
Sweden, 2008, J. Santos
Sweden, 2008, J. Santos
Sweden, 2009, K. Hansen & I. Olariaga
France, 2008, J. Cavet
Sweden, 2008, B. Wasstorp
Sweden, 2009, K. Hansen & I. Olariaga
Spain, 2009, F. Prieto & A. González
China, 2003, W.Y. Zhuang & Y. Nong
China, 1988, S. Wang & W.Y. Zhuang
Spain, 2003, J.δ. Pérez Butrón
Finland, 1978, H. Harmaja
Finland, 1987, P. Askola
Norway, 2009, K. Hansen & I. Olariaga
Finland, 2005, U. Salo & P. Salo
Sweden, 2008, J. Santos
Finland, 1970, δ. Fagerström
Norway, 2009, J. δorås
Sweden, 2011, J.C. Zamora & I. Olariaga
Italy, 2003, G. Jamoni
Italy, 1892, G. Bresadola
USA, 1917, A.H. Smith & R.J. Porter
Kε0100291
KM010024
Kε0100251
KM010028
Kε0100261
KM010027
JN942764
KM010078
JN942767
JN942766
Kε0100791
JN942765
KP119674
KF7175751
Kε0100301
KM010081
KM010082
KM010083
Kε0100841
Kε0100851
KM010031
Kε0100321
JN942775
–
–
Kε0100861
KF7175761
Kε0100871
Kε0100881
KM010036
Kε0100351
KF7175771
Kε0100341
KM010033
KM010037
KP006504
AF072095
Kε8231981
KM823466
Kε8231991
KM823467
Kε8232001
KM823468
JN941097
KM823469
JN941098
JN941084
Kε8232011
JN941085
–
Kε8232021
Kε8232031
KM823470
KM823471
KM823472
Kε8232051
Kε8232041
KM823473
JN941089
JN941095
DQ443444
DQ443445
Kε8232061
Kε8232091
Kε8232081
Kε8232071
KM823474
Kε8232121
Kε8232111
Kε8232101
KM823475
KM823476
–
–
O. borealis
O. brevispora as O. onotica var.
brevispora
O. brunneoparva (1)
O. brunneoparva (2)
O. brunneoparva (3)
O. brunneoparva (4)
O. brunneoparva (5)
O. brunneoparva (6)
O. bufonia (1)
O. bufonia (2)
O. bufonia (3)
O. bufonia (4)
O. bufonia (5)
O. bufonia (6)
O. bufonia (7)
O. caeruleopruinosa (1)
O. caeruleopruinosa (2)
O. caeruleopruinosa (3)
O. cantharella (1)
O. cantharella (2)
O. cantharella (3)
O. cantharella (4)
O. concinna (1)
O. concinna (2)
O. concinna (3)
O. crassa
O. daliensis (1)
O. daliensis (2)
O. flavidobrunneola (1)
O. flavidobrunneola (2)
O. flavidobrunneola (3)
O. formicarum (1)
O. formicarum (2)
O. formicarum (3)
O. formicarum (4)
O. formicarum (5)
O. fusconigra
O. integra
O. kauffmanii (1)
GenBank Accession no 4
170
Persoonia – Volume 35, 2015
Table 1 (cont.)
Taxon
Voucher
δocality / year/ collector
O. kauffmanii (2)
O. kaushalii
O. lactea
O. leporina (1)
O. leporina (2)
O. leporina (3) as O. smithii
O. leporina (4)
O. leporina (5)
O. leporina (6) as O. sp.
O. leporina (7)
O. leporina (8)
O. leporina (9)
O. leporina (10)
O. microspora
O. minor (1)
O. minor (2)
O. minor (3)
O. minor (4)
O. minor (5)
O. minor (6)
O. minor (7)
O. mirabilis (1)
MICH14409
T. δæssøe 6236 (C, dupl. BORH)
HMAS61359 (ex-MHSU 1803)
HεAS83579
HεAS83568
−
OSC 56824
OSC 56784
HεAS583570
JS.08.46 (S)
KH.09.93 (S)
JS.08.92 (S)
NV 2008.09.28 (dupl. S)
AH30502 (εICH)
H6003841
KH.10.311 (S)
H6008618
Tδ-Vorsø-0754 (C)
Cδ 950914-01 (dupl. S)
KH.98.84 (C)
C-F-83445
KH.09.188 (S)
O. mirabilis (2)
O. mirabilis (3) as O. umbrina
O. mirabilis (4)
O. mirabilis (5)
O. mirabilis (6)
O. myosotis
O. nannfeldtii (1)
O. nannfeldtii (2)
O. nannfeldtii (3)
O. nannfeldtii (4) (= O. lohjaënsis nom.
prov. Harmaja)
O. nannfeldtii (5)
O. nannfeldtii (6)
O. nannfeldtii (7)
O. nannfeldtii (8)
O. nannfeldtii (9)
O. onotica (1)
O. onotica (2)
O. onotica (3)
O. onotica (4)
O. onotica (5)
O. onotica (6)
O. onotica (7)
O. onotica (8)
O. onotica (9)
O. onotica (10)
O. oregonensis (1)
O. oregonensis (2)
O. oregonensis (3)
O. oregonensis (4) as O. rainierensis
O. oregonensis (5) as O. rainierensis
O. oregonensis (6) as O. rainierensis
O. oregonensis (7)
O. papillata (1)
O. papillata (2)
O. papillata f. pallidefurfuracea
O. phlebophora (1)
O. phlebophora (2)
O. phlebophora (3)
O. platyspora (1)
O. platyspora (2)
O. platyspora (3)
O. propinquata (1)
O. propinquata (2)
O. propinquata (3)
O. pseudoleporina (1) as O. concinna
O. pseudoleporina (2) as O. concinna
O. pseudoleporina (3) as O. concinna
O. pseudoleporina (4)
O. pseudoleporina (5)
O. pseudoleporina (6)
O. rainierensis
O. sinensis
O. smithii (1)
O. smithii (2)
KH.10.285 (S)
KH.01.09 (C)
S-F257083
NV 2008.09.14 (dupl. S)
S-F256929
H6003548
Cδ 091116-17 (S)
S-F257096
Cδ 091207-01 (S)
S-F249387 (Ex-H6017194)
USA, 1915, C.H. Kauffman
εalaysia, 1999, T. δæssøe
China, 1987, J. Z. Cao
China, 2003, W.Y. Zhuang & Y. Nong
China, 2003, W.Y. Zhuang & Y. Nong
−
USA, 1997, E.T. Peterson
USA, 1997, E.T. Peterson
China, 2003, W.Y. Zhuang & Y. Nong
Sweden, 2008, J. Santos
Sweden, 2009, K. Hansen & I. Olariaga
Sweden, 2008, J. Santos
France, 2008, N. Van Vooren
USA, 1948, A.H. Smith
Finland, 2006, U. Salo & P. Salo
Sweden, 2010, K. Hansen, K. Gillen & I. Olariaga
Finland, 1992, R. Saarenoksa
Denmark, 1982, T. δæssøe
Italy, 1995, C. δavorato
Denmark, 1998, K. Hansen
Denmark, 2007, T. δæssøe
Sweden, 2009, E. Bohus-Jensen, K. Hansen &
I. Olariaga
Sweden, 2010, K. Hansen, K. Gillen & I. Olariaga
Denmark, 2001, C. δange
Finland, 2010, ε. Carbone
France, 2008, J. Cavet
Italy, 1999, D. Bolognini
Finland, 1970, δ. Fagerström
Italy, 2009, C. δavorato
Italy, 2009, B. De Ruvo
Italy, 2009, C. δavorato
Finland, 1978, H. Harmaja
JS.08.103 (S)
NV 2008.10.01 (dupl. S)
H6002902
rh101310 (OSC)
KH.10.302 (S)
OSC 56801
OSC 56734
OSC 56759
C-F-89691
JS.08.48 (S)
KH.10.284 (S)
KH.09.132 (S)
KH.09.136 (S)
εCVE 23277
KH.98.107 (C)
rh139 (S)
Moorefun 58 (OSC, S)
εoorefun 31 (S)
OSC 56829
NSW6354 (OSC)
OSC 56745
EGS2179 (εICH)
H6003547
TUR 102134
NV 2007.09.27 (S)
JV06-385 (C)
S-F108338
K(ε)33068
KH.09.163 (S)
HK0846 (S)
JV06-656 (C-F-75309)
KH.09.99 (S)
JS.08.67 (S)
NV 2008.09.15 (dupl. S)
NSW7574 (OSC)
OSC 56749
OSC 56760
rh101910 (OSC)
εoorefun 14 (S)
OSC 56809
A.H. Smith 30553 (MICH)
HεAS61360
OSC 56799
ecv3345 (S)
Sweden, 2008, J. Santos
France, 2008, N. Van Vooren
Finland, 1972, C.-A. Haeggström
USA, 2010, R. Helliwell
Sweden, 2010, K. Hansen, K. Gillen & I. Olariaga
USA, 1997, E.T. Peterson
USA, 1996, E.T. Peterson
USA, 1996, E.T. Peterson
Denmark, 2008, H. Knudsen
Sweden, 2008, J. Santos
Sweden, 2010, K. Hansen, K. Gillen & I. Olariaga
Norway, 2009, K. Hansen & I. Olariaga
Norway, 2009, K. Hansen & I. Olariaga
Italy, 2008, ε. Carbone
Denmark, 1998, K. Hansen, T. δæssøe & C. δange
USA, 2010, R. Helliwell
USA, 2010, J. εoore
USA, 2010, J. εoore
USA, 1997, ε. Castellano
USA, 1990, D. εcKay
USA, 1996, J. Trappe
USA, 1948, E.G. Simmons
Finland, 1971, H. Harmaja
Finland, 1990, T. δindholm
France, 2007, N. Van Vooren
Denmark, 2006, δ. & J. Vesterholt
Sweden, 1949, G. Haglund & R. Rydberg
UK
Sweden, 2009, K. Hansen & I. Olariaga
Sweden, 2008, H. Kauffman
Denmark, 2006, J. Vesterholt
Sweden, 2009, K. Hansen & I. Olariaga
Sweden, 2008, J. Santos
France, 2008, J. Cavet
USA, N. S. Weber
USA, 1996, E.T. Peterson
USA, 1996, E.T. Peterson
USA, 2010, R. Helliwell
USA, 2010, J. εoore
USA, 1997, J. Spatafora
USA, 1948, A.H. Smith
China
USA, 1997, E.T. Peterson
USA, 2005, E. Vellinga
GenBank Accession no 4
ITS
δSU
KF717579
Kε0101191
–
–
–
–
–
–
–
KM010089
Kε0100901
KM010091
Kε0100921
AF072094
KM010040
Kε0100421
Kε0100391
KM010043
Kε0100441
Kε0100411
KM010038
JN942770
–
AF335111
DQ443447
DQ443448
DQ443449
AF0865732
Kε8232161
Kε8232151
DQ443443
KM823477
Kε8232131
KM823478
Kε8232141
–
KM823479
Kε8232181
Kε8232191
KM823480
Kε8232201
Kε8232171
KM823481
JN941086
Kε0100941
JN942769
KM010095
JN942768
KF717580
KF7175781
KM010096
KM010097
KM010098
Kε0100931
Kε8232211
AY500540
KM823482
JN941094
KM823483
Kε8232221
KM823484
KM823485
KM823486
Kε8232251
Kε0100451
Kε0100991
KF7175811
Kε0101001
Kε0101011
AF072067
AF072066
–
JN942773
KM010102
KP0065051
KM010103
JN942772
KM010104
–
KM010046
Kε0100481
Kε0100471
AF072087
AF072088
AF072089
AF072088
KF7175821
Kε0101051
KF7175841
Kε0100491
KM010050
EU784392
Kε0101061
KM010107
Kε0101081
Kε0101091
KM010110
Kε0101111
AF072083
AF072082
AF072081
Kε0101121
Kε0101131
AF072080
KF7175831
–
AF072063
JN942771
Kε8232241
Kε8232271
Kε8232281
Kε8232261
Kε8232231
AF086578
AF086577
JN941088
JN941090
KM823487
Kε8232291
KC012692
JN941096
KM823488
AF335121
KM823489
Kε8232311
Kε8232301
AF086597
AF086598
Kε8232321
–
Kε8232341
Kε8232331
Kε8232351
Kε8232361
KM823490
–
Kε8232381
KM823491
Kε8232371
Kε8232391
KM823492
Kε8232401
AF086593
AF086592
Kε8232441
Kε8232431
Kε8232421
Kε8232411
Kε8232451
DQ443451
JN941087
JN941093
171
I. Olariaga et al.: A monograph of Otidea
Table 1 (cont.)
Taxon
O. smithii (3)
O. smithii (4)
O. subformicarum (1)
O. subformicarum (2)
O. subformicarum (3)
O. subformicarum (4)
O. aff. subformicarum (1)
O. aff. subformicarum (2)
O. subterranea (1)
O. subterranea (2)
O. tuomikoskii (1)
O. tuomikoskii (2)
O. tuomikoskii (3)
O. tuomikoskii (4)
O. tuomikoskii (5)
O. tuomikoskii (6)
O. tuomikoskii (7)
O. tuomikoskii (8) as O. leporina
O. tuomikoskii (9)
O. tuomikoskii (10)
O. tuomikoskii (11)
O. unicisa (1)
O. unicisa (2) as O. grandis
O. unicisa (3) as O. grandis
O. yunnanensis
O. sp. ‘a’ (1)
O. sp. ‘a’ (2)
O. sp. ‘b’
Warcupia terrestris
1
2
3
4
Voucher
OSC 56753
OSC 56811
S-F242696
S-F256979
Cδ 050928-30, dupl. S-F256978
Private herb. CεP 1179, Rε 1095,
dupl. S-F256980
FH301035
FH301036
RH97 (FH)
RH69 (FH)
JS.08.68 (S)
εK200065 (S)
H6002901
JS.08.100 (S)
NV 2008.09.08 (S)
KH.09.130 (S)
KH.11.77 (S)
−
OSC 56756
OSC 56826
OSC 56761
KH.06.06 (FH)
HεAS51684
ZW Geo65-Clark (S)
HεAS 82166
εK0942 (S)
εK1081 (S)
KH.09.79 (S)
CBS 891.69
δocality / year/ collector
GenBank Accession no 4
ITS
δSU
USA, 1996, E.T. Peterson
USA, 1997, E.T. Peterson
Spain, 2012, J. Herranz & J.C. Campos
Spain, 2008, J. Fernández Vicente et al.
Italy, 2005, C. δavorato
Spain, 2009, C. ε. Pérez del Amo & R. Gil
AF072062
AF072060
KM010054
KM010051
Kε0100521
Kε0100531
AF086574
AF086572
KM823495
KM823494
Kε8232471
Kε8232461
εexico, 2007, ε. Hernández
εexico, 2007, ε.E. Smith
USA, 1997, R. Healy
USA, 1997, R. Healy
Sweden, 2008, J. Santos
Sweden, 2000, ε. Karström
Finland, 1972, R. Tuomikoski
Sweden, 2008, J. Santos
France, 2008, N. Van Vooren
Norway, 2009, K. Hansen & I. Olariaga
Sweden, 2011, ε. Prieto & I. Olariaga
−
USA, 1996, E.T. Peterson
USA, 1996, ε. εadsen & R. Davis
USA, 1996, E.T. Peterson
USA, 2006, δ. εillman
USA, Burdsall
USA, 2003, Z. Wang
China, 2003, Z.δ. Yang
Sweden, 2009, ε. Karström
Sweden, 2010, ε. Karström
Sweden, 2009, K. Hansen & I. Olariaga
Canada, 1966, J.W. Paden
Kε0100551
Kε0100561
FJ404766
FJ404767
KM010114
KM010115
KF7175851
KM010116
JN942777
JN942776
KM010117
–
AF072084
AF072086
AF072085
–
–
KM010118
–
KM010057
KM010058
Kε0101201
–
Kε8232491
Kε8232481
FJ404766
FJ404767
KM823496
KM823497
Kε8232501
KM823498
JN941091
JN941092
KM823499
AF0865883
AF086594
AF086596
Kε8232511
KC012693
DQ443446
AY789369
DQ443452
KM823500
KM823501
Kε8232521
DQ220467
Sequences from Hansen & Olariaga (2015).
The voucher specimen for AF086573 is mistakenly given as OSC 56823 in GenBank. This voucher (OSC 56823) is O. smithii based on morphological re-examination and the ITS sequence
(AF072061) deposited by the same authors. The δSU sequence AF086573 is O. leporina (Fig. 1).
The voucher specimen for AF086588 is mistakenly given as OSC 56825 in GenBank. This voucher (OSC 56825) is O. leporina based on morphological re-examination and the ITS sequence
(AF072078) deposited by the same authors. The δSU sequence AF086588 is O. tuomikoskii (Fig. 1).
ITS: Internal transcribed spacers (ITS1 and ITS2) and the 5.8S gene of the nrDNA; δSU: 28S large subunit of the nrRNA gene.
RESULTS
Alignment and ITS minisatellites
Fourty-nine ITS and 44 δSU sequences were newly generated
in this study (Table 1). In total 146 ITS sequences were utilised,
including 34 obtained from GenBank and 63 from Hansen &
Olariaga (2015). The ITS sequences were too variable to align
across all of Otidea, due to a highly polymorphic part in ITS1
and large length variability (insertions and deletions), and therefore were not included in phylogenetic analyses of the entire
genus. The ITS sequences were aligned among closely related species or species groups, and used as an aid to verify
identifications. The ITS region was especially useful in cases
were the protein-coding genes (RPB1, RPB2 and EF1) failed
to amplify, due to poor quality DNA from old or poorly treated
material. ITS sequences of the type specimens of O. mirabilis
and O. kauffmanii that could not be amplified for the multiple
genes are provided here. The all taxa δSU alignment consisted
of 167 δSU sequences, including 57 from GenBank and 68
from Hansen & Olariaga (2015) (Table 1) and 956 bp including
inserted gaps, of which 263 bp were parsimony informative
characters.
The O. borealis dataset consisted of 31 taxa, represented by
24 complete ITS-δSU and six ITS sequences, and 1 555 bp
including inserted gaps (ITS 614 bp; δSU 941 bp), of which
216 were parsimony informative characters. The O. formicarum
dataset consisted of 14 sequences and 2 531 bp (ITS 1 695 bp;
δSU 836 bp). Part of the ITS1 (805 bp) in the O. formicarum
dataset was omitted from the analyses, due to a long insertion
and tandem repeats in O. subformicarum and the two εexican
specimens, and the combined dataset thus included 1 726 bp,
of which 145 were parsimony informative characters. The insertion was composed of four tandem repeats (minisatellites) in
the four O. subformicarum sequences (212 bp in total), and in
one of them (S-F256979) the repeat was present a fifth time
(275 bp in total). In the εexican FH301036 the insertion was
extremely long (715 bp in total), of variable to random repeats.
The tandem repeats were 43 or 63 nucleotides. They were
composed of three parts (A-B-C) of 30, 20 and 13 nucleotides,
respectively, which were duplicates of the preceding part of
the ITS1 sequence. In the first two repeats the B part was
missing, whereas in the third-fifth all parts were present. The
C part was 100 % identical in all specimens and repeats; the
A part was mostly identical, but showed 3.3–6.7 % variation
in one repeat; and the B part showed 5–15 % variation in the
repeats. To ascertain the correctness of the long insertion in
the εexican specimens, the ITS sequence of FH301036 was
amplified and sequenced twice, using different sets of primers
(in one piece using ITS1-ITS4 and in two pieces using ITS55.8S / ITS3-ITS4). The two sequences were found to be identical. Unfortunately, we were not able to sequence the complete
insertion of FH301035 and the ITS1 was only sequenced in one
direction; both the part of the insertion recovered and the ITS1
were highly different from FH301036.
All taxa LSU phylogeny
The εδ analysis of the all taxa δSU dataset resulted in a single
best εδ tree of -lnδ = 6511.68. Bayesian analyses reached
an average standard deviation of split frequencies of 0.004
after 10 ε generations. A majority rule consensus tree was
constructed from the 300 000 trees sampled from the four
runs, each consisting of 75 000 trees sampled from the station-
172
Persoonia – Volume 35, 2015
O. cantharella (1) SE
100
O. cantharella (2) SE
O. cantharella (3) SE
100
O. cantharella
O. cantharella (4) FR
93
75
100
75 O. brunneoparva (1) SE
- O. brunneoparva (2) FI
O. brunneoparva (3) FI
100
76
99
O. brunneoparva
O. brunneoparva (4) SE
O. brunneoparva (5) SE
O. propinquata (1) SE
100/100
O. propinquata (2) SE
O. propinquata
O. propinquata (3) FR
O. tuomikoskii (1) SE
O. tuomikoskii (2) SE
97
100
O. tuomikoskii (3) FI
O. papillata f. pallidefurfuracea FR
O. tuomikoskii (4) SE
89
100
78
98
100
O. tuomikoskii (5) FR
98
O. tuomikoskii (6) NO
84
96
O. tuomikoskii
O. tuomikoskii (7) SE
91
O. tuomikoskii (8) US
O. tuomikoskii (9) US
100
O. tuomikoskii (10) US
O. tuomikoskii (11) US
O. pseudoleporina (1) US
O. pseudoleporina (2) US
100
O. pseudoleporina (3) US
100
O. pseudoleporina (4) US
O. pseudoleporina
O. pseudoleporina (5) US
93
O. pseudoleporina (6) US
100
100
O. leporina (3) US
O. leporina (4) US
100
O. leporina (5) US
O. crassa CHN
O. leporina (6) CHN
O. leporina
O. leporina (7) SE
100
O. myosotis FI
100
O. leporina (8) SE
O. leporina (9) SE
O. leporina (10) FR
100
O. papillata (1) FI
100
O. papillata (2) FI
O. papillata
O. alutacea (1) SE
99
O. alutacea (2) SE
100
O. alutacea (3) DK
clade 1
O. alutacea (4) SE
O. alutacea (5) US
O. alutacea (6) US
99
100
100
100
O. alutacea (7) US
clade 2
O. alutacea (8) US
O. alutacea (9) SE
98
clade 3a
O. alutacea (10) NO
100
O. alutacea (11, 30) SE
O. alutacea (12) SE
clade 3b
O. alutacea (13) DK
O. alutacea (14) SE
O. alutacea (15) CHN
75
O. alutacea complex
O. alutacea (16) CHN
100
O. alutacea (17) IT
O. alutacea (18) US
74
O. alutacea (19) US
95
-
94
97
96
clade 4
O. alutacea (20) US
O. alutacea (21) US
100
O. alutacea (22) NO
100
O. alutacea (23) ES
O. alutacea (24) FR
71
-
O. alutacea (25) CHN
O. alutacea s.str.
O. alutacea (26) CHN
98
O. alutacea (27) DK
100
O. alutacea (28) CHN
O. alutacea (29) IT
100
O. daliensis (1) CHN
100
O. daliensis (2) ES
100
100
99
O. subterranea (1) US
O. subterranea
O. subterranea (2) US
O. apophysata GE
100
O. daliensis
O. apophysata
O. platyspora (1) SE
-
98
O. platyspora (2) SE
O. platyspora (3) DK
O. platyspora
0.4
Monascella botryosa ES
Warcupia terrestris CA
Fig. 1 Bayesian inference 50 % majority rule consensus phylogram of Otidea from δSU sequence data. εaximum δikelihood bootstrap values (εδ-BP)
≥ 70 % and Bayesian posterior probabilities (PP) ≥ 95 % are shown above and below the branches, respectively. Thickened branches received support by
both εδ-BP ≥ 70 % and PP ≥ 95 %. Type collections are highlighted in bold. Country of origin for each collection is given using ISO country codes. Names of
species recognised are indicated by the vertical bars.
173
I. Olariaga et al.: A monograph of Otidea
O. bufonia (1) SE
100
O. bufonia (2) SE
O. bufonia (3) DK
100
O. bufonia
O. bufonia (4) ES
99
O. bufonia (5) FR
O. bufonia (6) FR
O. mirabilis (1) SE
O. mirabilis (2) SE
O. mirabilis (3) DK
-
O. mirabilis (4) FI
94
O. mirabilis
O. mirabilis (5) FR
O. mirabilis (6) IT
O. leporina (1) CHN
98
97 O. leporina (2) CHN
O. smithii (1) US
O. smithii (2) US
100
O. smithii
O. smithii (3) US
100
O. smithii (4) US
O. flavidobrunneola (1) FI
100
O. flavidobrunneola
O. flavidobrunneola (2) FI
100
O. flavidobrunneola (3) NO
O. brevispora
O. brevispora CHN
O. onotica (1) US
O. onotica (2) US
71
O. onotica (3) US
99
91
O. onotica (4) DK
100
O. onotica
O. onotica (5) SE
82
91
O. onotica (6, 7, 8) SE, NO
O. onotica (9) IT
O. onotica (10) DK
O. unicisa (1) US
100
100
100
100
O. unicisa
O. unicisa (2) US
O. unicisa (3) US
O. kaushalii
O. yunnanensis
O. kaushalii MY
100
O. yunnanensis CHN
100
O. minor (1) FI
100
O. minor (2) SE
100
O. minor (3) FI
O. minor (4) DK
95
71
O. minor
O. minor (5) IT
98
O. minor (6) DK
O. minor (7) DK
99
O. lactea CHN
O. lactea
O. oregonensis (1) US
O. oregonensis (2) US
O. oregonensis (3) US
100
O. oregonensis (4) US
100
O. oregonensis
O. oregonensis (5) US
O. oregonensis (6) US
95
100
100
O. sp. ‘a’ (1) SE
O. sp. ‘a’
O. sp. ‘a’ (2) SE
100
100
O. phlebophora (1) DK
100
100
O. phlebophora (2) SE
O. borealis FI
O. rainierensis US
O. concinna (1) SE
100
O. concinna
O. concinna (2) SE
-
100
O. concinna (3) ES
97
O. caeruleopruinosa (1, 3) FI, SE
100
O. caeruleopruinosa (2) ES
100
99
89
100
98
O. caeruleopruinosa
O. sinensis
O. sinensis CHN
O. subformicarum (1) ES
O. subformicarum (2) ES
O. subformicarum
O. aff. subformicarum (1) ME
O. aff. subformicarum (2) ME
98
O. formicarum (1) FI
100
97
100
O. formicarum (2) SE
O. formicarum (4) NO
O. formicarum (5) SE
91
O. sp. ‘b’ SE
99
-
O. nannfeldtii (1) IT
100
91
O. nannfeldtii (2) IT
99
100
100
99
O. nannfeldtii (6) FR
O. nannfeldtii (7) FI
O. nannfeldtii (8) US
O. nannfeldtii (9) SE
Fig. 1 (cont.)
O. sp. ‘b’
O. nannfeldtii (3) IT
O. angusta FI
O. nannfeldtii (4) FI
99
O. nannfeldtii (5) SE
98
73
-
O. formicarum
O. formicarum (3) FI
O. nannfeldtii
O. phlebophora
O. borealis
O. rainierensis
174
Persoonia – Volume 35, 2015
ary tree distribution (the first 25 % discarded as the burn-in)
(Fig. 1). The εδ and Bayesian tree topologies were congruent and recovered the same moderate to strongly supported
clades (Fig. 1). The terminal clades that constitute species we
recognise (Hansen & Olariaga 2015) have moderate to strong
support in both analyses, except for O. leporina and O. mirabilis.
A few synonymies inferred from sequences of type collections
from which we were not able to obtain the protein-coding
genes, and clear misidentifications of sequences deposited
in GenBank are evidenced by the δSU phylogeny (see Table
1). Based on δSU sequences from GenBank: the holotype of
O. crassa is nested within the O. leporina clade (= O. leporina
and O. pseudoleporina); O. lactea is suggested to be a sister
lineage to O. minor; O. sinensis is resolved as a sister species
to O. caeruleopruinosa, but without support; O. yunnanensis
forms a monophyletic group with O. kaushalii and O. unicisa;
and the type of O. onotica var. brevispora (= O. brevispora) is
supported as a sister lineage to O. onotica. The holotype of
O. mirabilis is nested among other collections of O. mirabilis
and the LSU sequence differs only in 1 bp from sequences of
collections from Scandinavia and France.
The O. formicarum and O. concinna clades in separate
ITS-LSU phylogenies
The εδ analysis of the O. formicarum dataset recovered a single
tree of -lnδ = 3579.93 (Fig. 2). Bayesian analyses reached
an average standard deviation of split frequencies of 0.003
after 3 ε generations. A majority rule consensus tree was
constructed from the 90 004 trees sampled from the four runs,
each consisting of 22 501 trees sampled from the stationary
tree distribution (the first 25 % discarded as the burn-in). The
four collections of O. subformicarum from Spain and Italy form
a distinct, strongly supported monophyletic group (εδ-BP and
PP 100 %). δikewise, the five collections of O. formicarum from
Fennoscandia, and the two collections of O. aff. subformicarum
from εexico, each form separate, strongly supported clades
(εδ-BP and PP 100 %; εδ-BP 84 %, PP 100 %, respectively).
Phylogenetic analyses of the combined ITS-δSU dataset fail
however, to resolve relationships among these three clades
with any certainty.
The combined εδ analysis of the O. borealis dataset resulted in
a single best εδ tree of -lnδ = 4230.82 (Fig. 3). Bayesian analyses reached an average standard deviation of split frequencies
of 0.005 after 3 ε generations. A majority rule consensus tree
was constructed, as for the O. formicarum dataset (above).
The supported topology (PP ≥ 95 %) did not differ from the
supported topology recovered by εδ analysis. The placement
of the single O. borealis collection is unresolved, but the ITSδSU phylogeny confirms it is genetically divergent from its sister
species (Fig. 3). Other species with a yellow outer surface of the
apothecia, O. concinna, O. minor, O. oregonensis and O. phlebophora, are each strongly supported as monophyletic (εδ
97–99 %, PP 100 %). An exception is O. minor (5) from Italy
that is resolved as a sister lineage to the rest of the O. minor
collections and O. integra without support. The ITS and δSU
sequences of O. minor (5) differ by 16 and 5 bp, respectively,
from the rest of the sequences of O. minor, which are 100 %
identical, except for the ITS sequence of O. minor (7) that
differs in 1 bp. Otidea integra is represented only by the ITS2
region (281 bp). The holotype of O. rainierensis is forming a
O. microspora US
O. rainierensis US
97
100
O. kauffmanii (1) US
98
100 O. kauffmanii (2) US
O. integra IT
O. minor (7) DK
O. minor (3) FI
99
O. minor (1) FI
100
O. minor (6) DK
O. formicarum (5) SE
O. minor (2) SE
100
O. formicarum (4) NO
100
100
O. minor (4) DK
100
O. formicarum (3) FI
O. minor (5) IT
O. oregonensis (1) US
O. formicarum (2) SE
O. oregonensis (3) US
O. oregonensis (2) US
O. formicarum (1) FI
84
100
100
100
O. oregonensis (4) US
98 O. oregonensis (5) US
99
O. oregonensis (6) US
O. aff. subformicarum (1) ME
O. aff. subformicarum (2) ME
100
100
O. oregonensis (7) US
100
100
O. subformicarum (2) ES
100
100
O. subformicarum (3) IT
100
100
O. subformicarum (4) ES
O. sp. ‘a’ (1) SE
O. sp. ‘a’ (2) SE
O. phlebophora (3) UK
O. phlebophora (2) SE
O. phlebophora (1) DK
O. subformicarum (1) ES
O. borealis FI
O. concinna (3) ES
97
99 O. concinna (1) SE
O. concinna (2) SE
O. nannfeldtii (7) FI
100
100
O. angusta FI
100
100 O. nannfeldtii (5) SE
0.0080
Fig. 2 The single best tree resulting from the εaximum δikelihood analysis
of the ITS-δSU regions of the O. formicarum clade. εδ bootstrap values
(εδ-BP) are shown above nodes and Bayesian posterior probabilities (PP)
below nodes. Thickened branches are nodes with high support (εδ-BP ≥ 75;
PP ≥ 95). Type collections are in bold.
100
100
O. caeruleopruinosa (2) ES
O. caeruleopruinosa (1) FI
0.02
Fig. 3 The single best tree resulting from the εaximum δikelihood analysis
of the ITS-δSU regions of O. borealis and closely allied species. εδ bootstrap
values (εδ-BP) are shown above nodes and Bayesian posterior probabilities
(PP) below nodes. Thickened branches are nodes with high support (εδ-BP
≥ 75 %; PP ≥ 95 %). Type collections are in bold.
I. Olariaga et al.: A monograph of Otidea
strongly supported clade with the holotype of O. kauffmanii
and a paratype of O. microspora (εδ 97 %, PP 100 %). The
O. kauffmanii and O. microspora types are only represented
by ITS and are therefore not included in any of our previous
analyses. We conclude O. kauffmanii is a synonym of O. rainierensis and O. microspora a doubtful name (see further under
Taxonomy). Otidea concinna is strongly supported as a sister
group to the rest of the ingroup, but otherwise the relationships
among the species are without support.
The phylogenetic results, and the ITS sequence similarity and
divergence (for species identification), will be further discussed
where applicable in the descriptive notes below.
Morphological characters for species delimitation
All Otidea species recognised by concordance of our four genes
phylogenies (Hansen & Olariaga 2015) can be recognised by
a combination of morphological characters. We evaluated the
characters in the context of the phylogeny and discovered
several new characters. The apothecial shape, colours, and
spore characters (size, shape, ornamentation) are important
for species identification, but to distinguish closely related species (or otherwise morphologically similar species) additional
characters are needed. These are the shape of the paraphyses,
ectal excipulum structure, type of exudates on the medullary
excipulum hyphae, resinous exudates on the outer surface of
the ectal excipulum and on the mycelium at the base of the
apothecia, and their possible reactions in εδZ and KOH (see
further in Hansen & Olariaga 2015). Below we provide details
on the resinous exudates, and their reactions in KOH and εδZ,
because they largely have been overlooked.
175
Excipular resinous exudates and reactions in MLZ and KOH
A resinous exudate is here used for a substance that is released
from the cells and in many species is vulnerable to common
mountants, but not water (following Huhtinen 1990). In Otidea
the exudates are concentrated in the excipulum cells, and on
the tomentum and mycelium at the base of the apothecia.
The resinous exudates are deposited on the outside of the
cell or hyphal walls. Harmaja (2009a) introduced the reaction
of coloured resinous exudates on the outermost cells of the
ectal excipulum in εδZ as a taxonomic character. Our study
revealed in addition, different reaction patterns of exudates
on the medullary excipulum cells and on the mycelium at the
base of the apothecia (including the tomentum), extending out
among the soil particles, which turned out to be diagnostic for
some species (see under εycelium at the base of the apothecia). In the medullary excipulum, most frequently scattered,
golden brown, resinous exudates are present at septa, e.g. in
O. alutacea, O. leporina, O. nannfeldtii (Fig. 4a). Otidea bufonia
has unique exudates, wrapping some hyphae and appearing
striate (referred to as ‘fingerprint-like’ by Korf & Zhuang (1991)),
sometimes forming big crystal-like aggregates (Fig. 4b, c). In
contrast, the sibling species, O. mirabilis, has only sometimes
biflabellate crystal-like exudates in the medullary excipulum
(Fig. 4d). Otidea papillata also possesses unique brown exudates, embedding some hyphae of the medullary excipulum
and sometimes appearing rod-like.
Small, resinous drops or amorphous matter are present in variable amounts on the outer surface of the apothecia of nearly
all Otidea species. In most species they are abundant and
Fig. 4 εedullary excipulum resinous exudates in Otidea. a. Hyphae with golden brown resinous exudates at septa in O. leporina (KH.11.02), in water*;
b, c. brown crystal-like exudates in O. bufonia (KH.07.37) in water †: b. overview; c. close-up of hyphae wrapped in striate exudates; d. close-up of biflabellate
crystal-like exudates in O. mirabilis (GεFN 1951, holotype), in water† — Scale bars = 10 µm; * = fresh material; † = dried material.
176
Persoonia – Volume 35, 2015
Fig. 5 Reactions of resinous exudates on the outermost ectal excipulum cells in Otidea. a. O. nannfeldtii † (H6010804, holotype of O. angusta), in water (left)
and converting into amber drops in εelzer’s reagent (right); b. O. bufonia* (KH.09.171), in water (left) and converting into reddish particles in εelzer’s reagent
(right); c. O. borealis† (S-F242694, holotype), in water (left) and turning bright yellow in KOH (right); d. O. nannfeldtii † (H6010804), in water (left) and turning
reddish brown in KOH (right); e. O. pseudoleporina† (rh101910, holotype), in water (left) and converting into reddish grey drops in KOH (right); f. ectal excipulum
cells showing a gelatinous sheath in O. formicarum* (KH.11.104). — Scale bars = 10 µm; * = fresh material; † = dried material.
easy to observe in water, but scarce and difficult to detect in a
few (O. alutacea, O. formicarum). One species, O. kaushalii,
has a unique type of exudate, i.e. crystal-like, oblate spheroid,
striate bodies, with a constricted centre. The possible reaction
of the exudates in εδZ and KOH is useful to separate certain
species or groups. Using these characters requires experience.
The reaction appears to vary depending on the amount of exudate and the concentration of εδZ. As Harmaja (2009a), we
observed two types of reactions in εδZ: i) Resinous exudates
dissolve and coalesce into spheroid drops, ‘amber drops’, that
contain hyaline bubbles (Fig. 5a). The ‘amber drops’ are overlooked if the ectal excipulum is directly mounted in εδZ, since
the exudates coalesce instantly and the drops can be washed
away. The reaction is best observed if εδZ is added to a water
mount. This reaction is present in many species. ii) Resinous
exudates partly convert into small reddish particles (Fig. 5b).
The reaction is often subtle and visible only in certain parts of
a mount. This reaction is typical in O. bufonia, O. mirabilis and
O. smithii. In some species the exudates do not react in εδZ
or they simply dissolve.
I. Olariaga et al.: A monograph of Otidea
177
Fig. 6 εycelium at the base of the apothecia and extending out in the substrate in Otidea†. a, b. O. flavidobrunneola (H6010806, holotype): a. resinous exudates on the hyphal walls in water; b. dissolved in εelzer’s reagent; c, d. O. borealis (S-F242694, holotype): c. pale yellow in water; d. turning bright yellow in
KOH. — Scale bars = 10 µm; † = all dried material.
In this study we detected three discriminative reactions of resinous exudates in 10 % KOH, which we propose as a novel
taxonomic character: i) In water the resinous exudates range
from yellow to dark reddish yellow (Fig. 5c, left) and in KOH
these dissolve completely ± exuding a yellow pigment, or partly
with the rest turning distinctly brighter yellow (Fig. 5c, right). This
reaction occurs in O. concinna and closely related species, viz.
O. borealis, O. caeruleopruinosa, O. flavidobrunneola, O. kaushalii, O. minor and O. oregonensis, and slightly less strikingly in
O. unicisa. ii) The resinous exudates are yellow-brown in water
(Fig. 5d, left) and turn reddish brown in KOH (Fig. 5d, right).
This reaction has been observed in O. nannfeldtii and O. propinquata. iii) The resinous exudates, yellow brown in water
(Fig. 5e, left), partly dissolve in KOH, and partly coalesce into
heterogeneous, pale reddish grey drops, with bubbles inside
(Fig. 5e, right). These drops are similar to the ‘amber drops’
observed in εδZ in many species, except for the pale red colour.
This reaction has been observed in O. leporina and O. pseudoleporina. A number of species have the outermost cells of the
ectal excipulum sometimes covered with a thin gelatinous
sheath (Fig. 5f).
Mycelium at the base of the apothecia
All Otidea species studied showed a conspicuous tomentum
covering the base of the apothecia and spreading out in the
substrate. In the microscopic descriptions we refer to both as
the basal mycelium. The hyphae are septate, straight and frequently branch and anastomose. No rhizomorphs have been
observed, although slender hyphal threads are sometimes
formed. Resinous exudates are often present on the surface,
especially on the mycelia extending out in the substrate, and
can appear like hyphal ornamentation.
Harmaja (2009a) proposed the colour of the basal tomentum
as a taxonomic character, but microscopic features of the basal
mycelium have been largely overlooked and have not been
included in species descriptions. Two types of characters are
useful for species identification: i) Resinous exudates occurring
on the hyphal walls (Fig. 6a). These can be inconspicuous or
nearly absent in some species, such as in O. alutacea s.l., in
which only a few refractive drops or minute hyaline exudates
are sometimes present. Resinous exudates are also scarce in
some species of the O. concinna clade. εany other species
show hyphae densely covered with resinous exudates, such as
O. formicarum, O. propinquata and O. tuomikoskii. The shape of
the exudates is variable, i.e. rod-shaped, hemispherical, conical
or bipyramidal. They dissolve and completely disappear in εδZ
(Fig. 6b). Sometimes doubts can arise about the nature of the
differently shaped exudates, but the facts that they detach from
the hyphal wall when the mount is squashed and dissolve in
εδZ, show these are not true ornamentation, i.e. not part of the
hyphal wall. ii) The hyphal wall turns yellow in KOH (Fig. 6d;
see Fig. 6c in water). We have observed this reaction clearly
in O. borealis and O. onotica. It can be observed in isolated
hyphae, but is more conspicuous when a mass of hyphae is
observed together. It can also be observed macroscopically.
178
TAXONOMY
Otidea (Pers.) Bonord., Handb. εykol.: 205. 1851
≡ Peziza (unranked) Otidea Pers., εycol. Eur. 1: 220. 1822.
≡ Peziza (unranked) Cochleatae Fr., Syst. εycol. 2: 46. 1822: Fr. loc. cit.
Type species. Otidea onotica (Pers. : Fr.) Fuckel, indicated by Saccardo,
Bot. Centralbl. 18: 215. 1884 (‘P. onotica Pers.’).
= Flavoscypha Harmaja, Karstenia 14: 107. 1974.
Type species. Peziza phlebophora Berk. & Broome.
= Otideopsis B. δiu & J.Z. Cao, Shanxi Univ. J., Nat. Sci. Ed. 4: 70. 1987.
Type species. Otideopsis yunnanensis B. δiu & J.Z. Cao.
Apothecia small to large, 3–75 mm high, 4–80 mm wide, often
in fascicles or caespitose, epigeous, cup- to ear-shaped and
split to the base on one side, less often entire, stipitate or not;
or hypogeous and enclosed. Hymenium white, yellow, ochre,
brown, almost black, often with pink stains. Receptacle surface
concolorous or with similar colours as hymenium, sometimes
with purplish, greenish or bluish tones, with conical to broadly
conical warts or pustules, less often smooth or furfuraceous,
concolorous or darker than the background. Base of the apothecium tomentose, mycelium white, ochre, yellow, orange or
brown, extending out in the substrate, base ribbed-veined in
a few species. Spores uniseriate, ellipsoid, oblong or fusoid,
typically with 2 guttules, sometimes with a few smaller granules, smooth (or verruculose in SEε), rarely spinose or with
low ridges, with de Bary bubbles in εδZ and Cotton Blue when
dried, thin-walled to slightly thick-walled, hyaline to very pale
brown. Paraphyses typically curved to hooked, rarely straight,
sometimes with notches or swollen at the apices, septate, typically containing refractive small guttules at the apices, fading in
colour and collapsed when dried. Asci cylindrical, operculate,
8-spored, 116–275 × 8–19 µm, with pleurorhynchous base.
Subhymenium c. 100–150 µm thick, of dense textura intricata,
hyphae sometimes swollen, often with scattered pigmented
exudates at septa. Medullary excipulum 400–1500 µm thick,
of textura intricata, hyphae cylindrical to slightly swollen, thinwalled to thick-walled, hyaline to pale brown, often with pigmented resinous exudates at septa. Ectal excipulum 70–150
µm thick, of textura angularis, less often of textura prismatica.
Surface with warts up to 180 µm high, formed by fasciculate, short hyphoid hairs of globose to elongated cells, or of
textura globulosa-angularis with single hyphoid hairs. Resinous
exudates often present on the surface, yellow to dark brown,
sometimes dissolving in εδZ, turning reddish or into brownish
yellow amber drops, sometimes changing colour in KOH. Basal
mycelium of septate, straight hyphae, that frequently branch
and anastomose, turning yellow or not in KOH, often covered
with pigmented, small, resinous exudates.
Ecology & Distribution — see Hansen & Olariaga (2015).
Key to species of Otidea
We were not able to study and interpret the following Chinese
species and these are therefore not treated nor included in the
key: Otidea bicolor W.Y. Zhuang & Zhu δ. Yang, O. kunmingensis W.Y. Zhuang, O. olivaceobrunnea Harmaja, O. sinensis J.Z.
Cao & δ. Fan, O. subpurpurea W.Y. Zhuang and O. tianshuiensis J.Z. Cao, δ. Fan & B. δiu. For O. integra (Bres.) Harmaja
see notes under O. phlebophora.
1. Ascomata hypogeous, globose to subglobose, truffle-like
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. O. subterranea
1. Ascomata epigeous, cup-shaped to ear-shaped, split or
entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Spores ornamented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2. Spores smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3. Spores with fine warts ± ridges . . . . . . . . . . 17. O. unicisa
3. Spores spiny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Persoonia – Volume 35, 2015
4. Spores 14 –17 × 7–9 µm . . . . . . . . . . . . . 16. O. kaushalii
4. Spores 16.5–20 × 7.6–10 µm . . . . . . 18. O. yunnanensis
5. Spores δm > 17 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5. Spores δm < 17 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
6. Apothecia broadly ear-shaped, split; typically ochraceous
yellow to ochre orange; often associated with Cudonia in
mossy Picea forests . . . . . . . . . . . . . . 11. O. cantharella
6. Apothecia cup-shaped, split or entire; brown; under Picea
or other trees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Apothecia entire; reddish brown; basal mycelium normally
with abundant minute resinous exudates; ectal excipulum
hyphoid hairs often with a gelatinous sheath (more easily
seen in εδZ); with Picea . . . . . . . . . 12. O. propinquata
7. Apothecia split or entire; purple or ochre-brown; basal
mycelium without or with sparse resinous exudates; ectal
excipulum hyphoid hairs without a conspicuous sheath;
with angiosperms. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Apothecia split, up to 75 mm wide; spores broadly ellipsoid
to oblong, Qm = 1.7–1.8; associated with Fagaceae . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. O. platyspora
8. Apothecia entire or split, up to 16 mm diam; spores narrowly
ellipsoid to fusoid, Qm = 1.9–2.1; most likely associated
with Betulaceae and Salicaceae . . . . . . . . . . . . . . . . . . 9
9. Apothecia deeply cup-shaped, split; ectal excipulum without
resinous exudates . . . . . . . . . . . . . . . . 1. O. apophysata
9. Apothecia shallowly cup-shaped, usually entire; ectal excipulum surface with abundant resinous exudates . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. O. daliensis
10. Receptacle surface with bright citrine yellow tones in young
apothecia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
10. Receptacle surface without citrine yellow tones . . . . . 15
11. Spores Qm = 1.7– 2; apothecial base at most wrinkled,
without high ribs or veins . . . . . . . . . . . . . . . . . . . . . . . 12
11. Spores Qm = 2–2.3; apothecial base with ribs or strongly
veined at least in some apothecia . . . . . . . . . . . . . . . . 14
12. Receptacle surface ochraceous yellow; basal mycelium
yellow in KOH; spores Wm = 6.5 µm, Qm = 1.7 . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25. O. borealis
12. Receptacle surface citrine yellow; basal mycelium unchanged in KOH; spores Wm = 5.6–6 µm, Qm = 1.8–2 . . . . . 13
13. Apothecia sometimes entire, sometimes with blunt ribs at
the base; North America . . . . . . . . . . 31. O. oregonensis
13. Apothecia split, without ribs at the base; Europe. . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27. O. concinna
14. Apothecia mostly entire, base strongly ribbed-veined and
anastomosing in all the apothecia . . . 32. O. phlebophora
14. Apothecia mostly split, base with a few ribs-veins, smooth
in some apothecia . . . . . . . . . . . . . . . . . . . . 30. O. minor
15. At least some paraphyses straight or curved, claviform
to almost capitate at apices; and/or resinous exudates of
the ectal excipulum yellow or reddish yellow, turning bright
yellow in KOH; apothecia split or not . . . . . . . . . . . . . 16
15. Paraphyses not as such; resinous exudates of the ectal
excipulum, when present, not turning bright yellow in KOH;
apothecia always split . . . . . . . . . . . . . . . . . . . . . . . . . 20
16. Apothecia shallowly cup-shaped and irregular, entire . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29. O. lactea
16. Apothecia ear-shaped or deeply cup-shaped, split . . . 17
17. Paraphyses often claviform or capitate at apices, 3–9 µm
broad; spores Qm = 1.6–1.8 . . . . . . . . . . . . . . . . . . . . 18
17. Paraphyses at most slightly swollen at apices, 2–5 µm
broad; spores Qm = 1.9–2 . . . . . . . . . . . . . . . . . . . . . . 19
18. Basal mycelium turning yellow in KOH; Europe . . . . . . .
. . . . . . . . . . . . . . . . . . . (see under O. borealis) O. sp. ‘a’
179
I. Olariaga et al.: A monograph of Otidea
18. Basal mycelium not yellow in KOH; North America . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33. O. rainierensis
19. Spores δm = 11.2–11.8 µm; sometimes receptacle with a
bluish hue, ochraceous grey when dried; basal tomentum
light ochre in dried specimens . . 26. O. caeruleopruinosa
19. Spores δm = 10–10.6 µm; receptacle without bluish hue
when fresh, reddish brown when dried; basal tomentum
orange-ochre in dried specimens 28. O. flavidobrunneola
20. Spores δm < 12 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
20. Spores δm > 12 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
21. Apothecia dark brown with lilaceous tones; Asia . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23. O. purpurea
21. Apothecia not dark brown, without lilaceous tones; Asia,
Europe or North America . . . . . . . . . . . . . . . . . . . . . . . 22
22. εedullary excipulum with reddish brown resinous exudates
scattered among and covering some hyphae; ectal excipulum of textura prismatica to textura intricata; receptacle surface with contrasting brown warts . . . . . . . 6. O. papillata
22. εedullary excipulum sometimes with yellowish brown resinous exudates at septa, not covering hyphae; ectal excipulum of textura angularis; receptacle surface with contrasting warts or not . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23. At least some warts higher than 85 µm; basal tomentum
orange-ochre when dried; apothecial sections often yellow
in KOH; without resinous exudates at septa in the medullary
excipulum . . . . . . . . . . . . . . . . . . . . . . . 9. O. tuomikoskii
23. Warts up to 85 µm; basal tomentum pale ochre or yellow
when dried; apothecial sections not yellow in KOH; sometimes with resinous exudates at septa in the medullary
excipulum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24. Hymenium with distinct yellow or orange tones, ochreyellow to pinkish orange; resinous exudates on the ectal
excipulum partly dissolving into reddish grey, heterogeneous drops in KOH or basal mycelium yellow in KOH . 25
24. Hymenium without or with weak orange tones; resinous
exudates sometimes turning reddish brown but not dissolving into drops in KOH; basal mycelium not turning yellow
in KOH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
25. Resinous exudates on the ectal excipulum partly dissolving into reddish grey heterogeneous drops in KOH; basal
mycelium not turning yellow in KOH; North America . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. O. pseudoleporina
25. Resinous exudates on the ectal excipulum not dissolving
into reddish grey heterogeneous drops in KOH; basal mycelium turning yellow in KOH; Asia . . . . . 19. O. brevispora
26. Hymenium sometimes with pink tones; apothecia yellowish
ochre to brown; narrowly ear-shaped in the beginning . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14. O. nannfeldtii
26. Hymenium without pink tones; apothecia reddish brown to
orange-brown; broadly ear-shaped . . . . . . . . . . . . . . . 27
27. Spores δm = 10 –10.7 µm; Qm = 1.6 –1.7 . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. O. formicarum
27. Spores δm = 11.1–11.7 µm; Qm = 1.7–1.9 . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . 15. O. subformicarum
28. Spores Qm < 1.8; ear-shaped . . . . . . . . . . . . . . . . . . . 29
28. Spores Qm > 1.8; ear- or cup-shaped . . . . . . . . . . . . . 30
29. Apothecia dark brown, sometimes with olivaceous tint; paraphyses mostly with distinct notches . 10. O. brunneoparva
29. Apothecia cinnamon brown, without olivaceous tint; paraphyses not or slightly notched . . . . . . . . . . 7. O. leporina
30. Apothecia ochraceous yellow, hymenium often with pink
tones; basal mycelium turning yellow in KOH . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22. O. onotica
30. Apothecia pale or dark brown, without pink tones; basal
mycelium not turning yellow in KOH . . . . . . . . . . . . . . 31
31. Receptacle medium brown; ectal excipulum resinous exudates absent or scarce, light yellowish brown; basal mycelium without dark brown resinous exudates . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. O. alutacea s.l.
31. Receptacle dark purple brown; ectal excipulum resinous
exudates abundant, dark brown; basal mycelium with dark
brown resinous exudates . . . . . . . . . . . . . . . . . . . . . . 32
32. Spores Qm < 2, ellipsoid; apothecia mostly ear-shaped;
North America . . . . . . . . . . . . . . . . . . . . . . 24. O. smithii
32. Spores Qm > 2, ellipsoid-fusoid; apothecia mostly cupshaped, split; Eurasia and North America . . . . . . . . . . 33
33. Receptacle strikingly purple-violaceous (fresh); medullary
excipulum without, or rarely with flabellate crystal-like exudates, forming cross-like aggregates; under conifers on calcareous ground . . . . . . . . . . . . . . . . . . . . 21. O. mirabilis
33. Receptacle mostly without purple tones; medullary excipulum with striate exudates covering some hyphae, sometimes forming crystal-like aggregates; often under deciduous trees or on acidic ground . . . . . . . . . . 20. O. bufonia
Based on genealogical concordance phylogenetic species
recognition (GCPSR: Taylor et al. 2000), using the four loci,
RPB1, RPB2, EF1 and δSU rDNA, we delimited 25 species
within Otidea (see Hansen & Olariaga 2015). In addition eight
species were recognised by genetic divergence from their sisters. Twenty-eight of these are treated and discussed below,
along with O. brevispora, O. lactea, O. subterranea and O. yunnanensis included in our LSU phylogeny, and O. purpurea that
has only been studied morphologically. The species are presented following their phylogenetic relationships, inferred from
our combined three- and four-gene analyses (f. 3 in Hansen
& Olariaga 2015).
Otidea platyspora clade
Apothecia disc-shaped, cup-shaped and split, or globose and
hypogeous, brown. Spores large, exceeding 20 µm, except
14–16.5 µm if hypogeous. Basal mycelium smooth or with very
sparse resinous exudates.
Species — Otidea apophysata, O. daliensis, O. platyspora,
O. subterranea.
1. Otidea apophysata (Cooke & W. Phillips) Sacc., Syll. Fung.
8: 96. 1889
Basionym. Peziza apophysata Cooke & W. Phillips in Cooke, Grevillea 5:
60. 1876.
≡ Pseudotis apophysata (Cooke & W. Phillips) Boud., Hist. Classific. Discomyc. Europe: 52. 1907.
Holotype. England, Shrewsbury, in a damp ditch, 1876, W. Phillips (K(ε)
30410 ex Herb. Phillips). Isotype (K(ε) 167215 ex Herb. Cooke) !
Misapplied names
– Otidea felina sensu Boudier, Icon. εycol. livr. 29: n°. 512, pl. 331. 1910
(preliminary text with ‘circulaires’).
Apothecia solitary to caespitose, 8–30 mm high, up to 15 mm
wide, initially ear-shaped, then soon expanding and becoming cup-shaped, split, sessile or stipitate. Hymenium purple
brown (6D3, 6D4), when dried dark orange brown (6E7, 6F7).
Receptacle surface pale greyish orange (6C5), purple brown
(6D4), when dried dark orange brown (6E7, 6F7), furfuraceous.
Warts scarce to absent, low. Stipe absent. Basal tomentum and
mycelium whitish to pale orange grey (5B2). Spores narrowly ellipsoid to fusoid, narrowing toward the poles, sometimes inequilateral, with two large guttules, and often with several smaller
guttules, smooth, hyaline, 20–24.5 × 9–11 µm (δm = 21.6 µm,
Wm = 10 µm, Qm = 2.1; n = 1). Paraphyses curved to hooked,
seldom straight, slightly enlarged at apices, 3–4(–5) µm wide,
180
sometimes with a sinuous underside or with 1–2 notches, frequently branching, entangled and interconnected, when dried
containing small, refractive, hyaline guttules. Asci 172–197 ×
12–13 µm. Apothecial section 700–850 µm thick. Medullary
excipulum of textura intricata, 400 – 500 µm thick, hyphae
slightly thick-walled, 5–9 µm wide, pale brown. Ectal excipulum
of textura angularis, 70–110 µm, cells thin-walled, yellowish
brown, 18–33 × 13–22 µm. Surface with low warts, up to 40
µm high, cells ovoid to globose, constricted at septa, 8–13 µm
wide. Resinous exudates absent. Basal mycelium of 4–5 µm
broad, pale brown hyphae, sometimes with oily refractive drops
on the surface.
Specimens examined. gErmany, Nordrhein, Herten, 1 Sept. 1999, F. Kasparek, private herb. Kasparek s.n. (dupl. S-F257062).
Notes — Otidea apophysata is characterised by deeply cupshaped, split, brown apothecia, and large, ellipsoid to fusoid
spores. Otidea daliensis is a closely related species, distinguished by darker brown, shallowly cup-shaped, usually entire
apothecia, and abundant dark brown resinous exudates on the
outermost cells of the ectal excipulum. Otidea platyspora has
also brown apothecia and large spores, but it differs from O. apophysata in the larger apothecia, partly buried in the substrate,
broadly ellipsoid to oblong spores, and non-entangled paraphyses without notches.
The name O. apophysata has been misapplied twice for O. daliensis (Boudier 1909b, Pérez-Butrón & Fernández-Vicente
2008). The type material of O. apophysata lacks resinous exudates on the ectal excipulum, which clearly distinguishes it
from O. daliensis. An original painting by W. Phillips, based
on the type material of O. apophysata and preserved at RBG
Kew (reproduced in Parslow & Spooner 2013), shows typical
brown, split, deeply cup-shaped apothecia. The illustration by
Cooke (1878, f. 350), based on drawings and (likely dried)
specimens communicated by W. Phillips, shows slightly darker
apothecia than typical.
Otidea apophysata is only known from very few reports from
France (Boudier 1910 as O. felina), Germany (Häffner &
Winterhoff 1989, Kasparek 2000), Belgium and Spain (Van
Vooren 2011a). In εid to South Britain it is widely distributed,
with collections from fourteen different localities (Parslow &
Spooner 2013). Unlike most Otidea species, O. apophysata
shows preference for damp habitats, and might be associated
with Alnus and Populus (Häffner & Winterhoff 1989, Parslow
& Spooner 2013).
2. Otidea daliensis W.Y. Zhuang & Korf, εycotaxon 35: 300.
1989
Holotype. China, Yunnan, Dali, Hudiequan Park, alt. 2100 m, on bare soil
under seedlings of Plantago major, 5 Nov. 1988, R.P. Korf, L.S. Wang & W.Y.
Zhuang (HεAS 57688). Isotype (CUP-CH 2532).
Misapplied names
– Pseudotis apophysata sensu Boudier, Icon. εycol. livr. 24: n°. 471, pl. 332.
1909 (preliminary text with ‘circulaires’).
– Otidea apophysata sensu Pérez-Butrón & Fernández-Vicente, Errotari 5:
37. 2008.
Apothecia gregarious, up to 9 mm high, 3–16 mm wide, initially
cup-shaped, sometimes split, then becoming shallowly cupshaped, sessile or stipitate. Hymenium when dried dark purple
brown (7F5, 7F6) to dark brown (6F5). Receptacle surface
dark purple brown (7F5, 7F6), when dried dark brown (6F5),
furfuraceous. Warts absent or very low. Stipe if present very
short. Basal tomentum and mycelium whitish to pale orange
grey (5B2). Spores broadly ellipsoid to ellipsoid and narrowing
toward the poles, sometimes inequilateral, with two large guttules, and often with several smaller granules, smooth, hyaline,
Persoonia – Volume 35, 2015
(19.5 –)20.5 – 23 × 10.5 –12(–13) µm (δm = 21.2 – 21.5 µm,
Wm = 10.7–11 µm, Qm = 1.9–2.1; n = 2). Paraphyses curved to
hooked, sometimes slightly enlarged at apices, 2.5–3.5(–5) µm
wide, sometimes with slightly swollen areas, apices sometimes
embedded in a brown matter, when dried containing small,
refractive, brown guttules. Asci 199–212 × 15–17 µm. Apothecial section 600–850 µm thick. Subhymenium c. 90–110 µm
thick, of dense textura intricata, visible as a darker zone, cells
cylindrical to swollen, with scattered brown resinous exudates
at septa. Medullary excipulum of textura intricata, 300–400 µm
thick, hyphae 4–6.5 µm wide, slightly thick-walled, pale brown,
with brown resinous exudates at septa. Ectal excipulum of
textura angularis, 90–120 µm thick, cells thin-walled, yellowish
brown, 18–28 × 11–28 µm. Surface with broadly conical warts.
Non-warted parts with 2–5-celled hyphoid hairs, with claviform
uppermost cell, more rarely cylindrical, constricted at septa,
6–9 µm wide. Resinous exudates abundant, dark brown, partly
dissolving in εδZ. Basal mycelium of 3.5–4 µm wide, hyaline
hyphae, with yellowish brown, small, resinous exudates.
Specimens examined. FranCE, 1869, L. Quélet (UPS F-629790). – Spain,
Basque Country, Bizkaia, Galdames, Presa de Aguas Juntas, sandy soil
under Populus nigra, 11 Aug. 2003, J.L. Pérez Butrón, SEST-03071103; 17
Aug. 2006, SEST-06081702.
Notes — Otidea daliensis is recognised by small, usually
entire, dark purple brown, shallowly cup-shaped apothecia,
large ellipsoid spores often narrowing toward the poles, and
abundant brown resinous exudates on the ectal excipulum. This
species has been confused with O. apophysata (see O. apophysata).
The first known report of O. daliensis was by Boudier (1909b),
as Pseudotis apophysata. The plate 332 (n°. 471) shows the
typical dark purple, shallowly cup-shaped apothecia, in contrast
to O. apophysata, depicted in plate 331 (n°. 512) as O. felina
(Boudier 1910). εornand & Courtecuisse (2005) proposed a
provisional name, O. boudieri, for the Boudier P. apophysata
plate (= O. daliensis, n°. 332). Zhuang & Korf (1989) described
O. daliensis without comparing it to O. apophysata. εaterial with
small, shallowly cup-shaped apothecia and darker colour was
reported from the Iberian Peninsula as O. apophysata (PérezButrón & Fernández-Vicente 2008), and thus had similarities
with O. daliensis and Boudier’s plate 332. Van Vooren (2011a)
considered the Iberian O. apophysata material to represent
O. daliensis. After restudying the Iberian material and comparing it to O. apophysata, we agree with that statement. δSU
sequences obtained from the Iberian material and from the
Chinese holotype of O. daliensis are identical.
3. Otidea platyspora Nannf., Ann. Bot. Fenn. 3: 317. 1966.
— Fig. 7
Holotype. SwEdEn, Uppland, Djursholm, Oct. 1951, A. Zander, Fungi Exs.
Suec. 3284 (UPS F-005428). Isotype (S-F88395) !
Misapplied names
– Otidea cochleata sensu Boudier, Icon. εycol. livr. 21: n°. 461, pl. 329. 1908
(preliminary text with ‘circulaires’).
Apothecia caespitose, 60–70 mm high, 40–75 mm wide, initially ear-shaped, then soon expanding and becoming deeply
cup-shaped, split, sessile or shortly stipitate. Hymenium initially
yellowish brown (5C6, 5C7), pale greyish brown (5C3, 5C4) to
dark brown (6F3–6F6), when bruised margin blackish, when
dried brownish ochre (5B4, 5B5). Receptacle surface dark ochre
brown (5D7, 5D8), slightly hygrophanous, in drying pale ochre
brown (5B6), when dried yellowish brown (5C6, 5C7), sometimes wrinkled at the base, finely furfuraceous in the margin.
Warts absent. Stipe not well developed. Basal tomentum and
mycelium whitish to pale brown (5A3). Spores broadly ellipsoid
I. Olariaga et al.: A monograph of Otidea
181
Fig. 7 Otidea platyspora*. a. Apothecia; b. spores; c. paraphyses; d. ectal excipulum (a: KH.10.183; b–d: KH.09.163). — Scale bars = 10 µm; * = all fresh
material.
to oblong, rarely slightly inequilateral, with two large guttules,
often with several smaller guttules, smooth, hyaline, 18–22 ×
(9.5–)10.5–12 µm (δm = 19.8–20.7 µm, Wm = 10.9–11.6 µm,
Qm = 1.7–1.8; n = 6). Paraphyses curved to hooked, of the
same width or slightly enlarged at apices, 2.5 – 3.5(– 5) µm
wide, without notches, rarely with a slightly swollen area on
the underside, when fresh containing small, refractive, light
brownish yellow guttules; when dried tiny, light yellow granules.
Asci 168–213 × 14–19 µm. Apothecial section 700–1700 µm
thick. Subhymenium 80–100 µm thick, visible as a darker zone,
cells cylindrical, densely arranged, with scarce yellowish brown
resinous exudates at septa. Medullary excipulum of textura
intricata, 550–1100 µm thick, hyphae 3.5–10 µm wide, sometimes slightly swollen, thin-walled, light yellowish brown. Ectal
excipulum of textura angularis, 70–90 µm, cells thin-walled,
pale brown, 13–37 × 12–22 µm. Surface with hyphoid hairs,
33–70 µm long, of 4–7 ovoid to subglobose cells, constricted
at septa, 6–9 µm wide, sometimes with a light brown matter.
Resinous exudates absent to scarce, pale brown, dissolving
in εδZ. Basal mycelium of 3.5–4.5 µm wide, hyaline to very
pale brown hyphae, sometimes with oily refractive drops on the
surface, sometimes with minute bipyramidal resinous exudates.
Specimens examined. azErbaijan, εontes Talysh, in silva mixta, 14 Oct.
1962, E. Parmasto (UPS F-629452). – dEnmark, East Jylland, Kammerherrens Ege, εoesgård Skov, S of Århus, under Fagus and Quercus, 11 Sept.
2006, J. Vesterholt, JV06-656 (C). – FranCE, Ain, commune de Saint-Benoit,
forêt d´Évieu, under Quercus robur, Carpinus betulus, Corylus avellana and
Alnus glutinosa, 15 July 2011, F. Armada, NV 2011.07.04 (dupl. S). – SwEdEn, Uppland, Stockholm, Drottningsholm, under Quercus robur, 13 Sept.
2009, K. Hansen & I. Olariaga, KH.09.163 (S); Uppland, Stockholm, Solna,
Karlbergsparken, garden with broadleaf trees, 29 July 2006, H.-G. Toresson
(S-F248339); Uppland, Stockholm, Sånga, Svartsjö slott, under Quercus,
Fagus and Corylus, 31 Aug. 2008, H. Kauffman, HK08046 (S); Uppland,
Uppsala, mixed forest with Pinus sylvestris, Picea abies, Betula pendula
and Quercus robur, 9 Oct. 2011, J.C. Zamora (BIO-Fungi 16391); Uppland,
Uppsala, Hågadalen-Nåsten Nature Reserve, Predikstolen, under Corylus,
Populus, Quercus and Picea, on rich ground, 6 Sept. 2010, K. Hansen &
I. Olariaga, KH.10.183 (S).
Notes — Otidea platyspora is characterised by large, caespitose, brown apothecia, blackening in bruised margins and large
spores. εacroscopically it resembles members of the O. alutacea complex, but these differ in the smaller spores. Otidea apophysata and O. daliensis have likewise brown apothecia and
large spores, but can be distinguished by having even larger
spores and smaller (up to 30 mm) apothecia (see further under
those species).
Otidea platyspora is a striking species with scarce records.
It was described from Sweden and has been reported from
France (Boudier 1908, Nannfeldt 1966, Van Vooren & Armada
2011), the Netherlands (εaas Geesteranus 1967) and recently
from Britain and Denmark (Parslow & Spooner 2013). We report it as new from Azerbaijan and give additional records from
Denmark and Sweden. Otidea platyspora appears to have been
overlooked and is more widespread than thought previously.
4. Otidea subterranea Healy & ε.E. Sm. in Smith & Healy,
εycol. Res. 113: 860. 2009
Holotype. USA, Iowa, δedges State Park, shallowly hypogeous, erumpent
on soil, 30 Aug. 1997, R. Healy RH69 (FH).
Notes — Otidea subterranea is the only known hypogeous
species of Otidea. The pustules (c. 50–100 µm high) on the
182
outer surface of the ptychothecia and the incrusted tomentum
are typical for Otidea. The asci are cylindrical, with 8 uniseriate spores, placed in a defined hymenium. Unlike most Otidea
species the spores are uniguttulate. As an adaptation to a hypogeous habit active spore discharge has been lost. Probably for
the same reason, the paraphyses are aggregated and fused
subapically to form an epithecium of brown thick-walled cells,
and are reminiscent of the frequently branching, entangled and
interconnected paraphyses in O. apophysata. There are no
other morphological features that support the exact placement
of O. subterranea in the O. platyspora clade. The receptacle
surface of O. subterranea ascomata is whitish to cream in
young and peach-cream to buff with tan-brown areas in older
specimens. The gleba is of dark brown fertile veins lined with
a thin, light yellowish brown hypothecium, but the sterile veins
are of whitish hyphae (Smith & Healy 2009).
Otidea alutacea clade
Apothecia cup-shaped and split, brown. Spores typically ellipsoid, with almost parallel sides. Outermost ectal excipulum and
basal mycelium smooth or with very sparse resinous exudates.
Species — Otidea alutacea s.l.
5. Otidea alutacea (Pers.) εassee, Brit. Fungus-Fl. 4: 446.
1895. — Fig. 8
Basionym. Peziza alutacea Pers., Observ. εycol. 2: 78. 1799; non Peziza
alutacea Schumach., Enum. Pl. 2: 431. 1803 (homonym).
≡ Scodellina alutacea (Pers.) Gray, Nat. Arr. Brit. Pl. 1: 668. 1821.
≡ Peziza cochleata var. alutacea (Pers.) Fr., Syst. εycol. 2: 50. 1822:
Fr. loc. cit. (‘ß alutacea’).
≡ Plicaria alutacea (Pers.) Fuckel, Jahrb. Nassauischen Vereins Naturk.
23–24: 327. 1870.
≡ Aleuria alutacea (Pers.) Gillet, Champ. France Discom. 1: 42. 1879.
Lectotype. Bull., Hist. Champ. France 1: t. 154, f. b., designated by Carbone (2010a). Epitype. (δ0111551, Herb. Persoon), designated by Carbone
(2010a).
Apothecia gregarious, rarely caespitose, 15 –75 mm high,
8–48 mm wide, initially ear-shaped, soon expanding, becoming shallowly to deeply cup-shaped, split, sessile or stipitate.
Hymenium initially brown (5D6), then yellowish brown (5D4,
5D5) to dark reddish brown (7D7, 7E7), when dried purple
brown (6D6, 6D7). Receptacle surface slightly hygrophanous, in
drying yellowish brown (5D5, 5D6), or sometimes with purplish
brown (7D5) tones, when dried light ochre (5A4, 5B4) brownish
ochre (5B5, 5B6), finely furfuraceous to slightly warty in the
margin. Warts flattened, gregarious, concolorous, darker when
the outside in drying. Stipe 3 – 6 × 3–4 mm. Basal tomentum
and mycelium white to very light ochre (5A2). Spores ellipsoid
to broadly ellipsoid, oblong ellipsoid, slightly inequilateral,
with two large guttules, often with several smaller granules,
smooth, hyaline, (13.5–)14.5–16.5(–17.5) × 6.5–7.5(–8) µm
(δm = 14.6–16.1 µm, Wm = 6.6–7.5 µm, Qm = 2.1–2.2; n = 5).
Paraphyses curved to hooked, only few straight, of the same
width or slightly enlarged at apices, 2.5–4.5 µm wide, without
notches, sometimes embedded in a brown matter at apices,
when fresh containing small, refractive, light brownish yellow
guttules; when dried brownish yellow. Asci 140–187 × 11–13
µm. Apothecial section 750 –1050 µm thick. Subhymenium
c. 80–100 µm thick, visible as a darker zone, cells cylindrical
to swollen, densely arranged, with scattered yellowish brown
resinous exudates at the septa. Medullary excipulum of textura
intricata, 400–650 µm thick, hyphae thin-walled to slightly thickwalled, 4–9 µm wide, hyaline to light brown, sometimes with
yellowish brown resinous exudates at septa. Ectal excipulum of
textura angularis of 80–100 µm, cells thin-walled, pale brown,
1
1
Description based only on specimens of O. alutacea s.str.
Persoonia – Volume 35, 2015
9–21 × 9–17 µm. Surface with broad conical warts, 35–57
µm high, formed by short, fasciculate, hyphoid hairs, of 6–7
globose to subglobose cells, constricted at septa, 7–10 µm
wide. Resinous exudates absent to scarce, yellowish brown,
dissolving in εδZ. Basal mycelium of 3–4.5 µm wide, hyaline
hyphae, with oily refractive drops on the surface, sometimes
with minute resinous exudates.
Specimens examined. O. alutacea s.str. — dEnmark, S Sjælland, εøn,
Store Klinteskov, by Svantestenen, on calcareous soil (pH 7.0) along forest
road, under deciduous trees, together with Humaria hemisphaerica and
Trichophaea woolhopeia, 11 Sept. 1994, K. Hansen & S.K. Sandal, KS94-111 (C). – FranCE, Puy-de-Dôme, Auvergne, Nadayat, sous feuillus en
terrain neutrocline, 20 Sept. 1998, G. Corriol, GC 98092002 (dupl. S). – italy,
Piemonte, Vignole Borbera (Aδ), Fraz. Variano superiore, under Quercus
pubescens and Castanea sativa, 19 Oct. 2010, M. Carbone (S-F257084). –
norway, Nord-Trøndelag, δeksvik, Gjøråsvika, on rich, bare ground, under
Corylus and Picea, on a steep slope, 3 Sept. 2009, K. Hansen & I. Olariaga,
KH.09.133 (S). – Spain, Gipuzkoa, Tolosa, Elosegi markesaren lorategiak,
under broadleaf trees in a garden, 29 εay 2009, J.I. López-Amiano, JδA
2009052902 (ARAN-Fungi A3023204). – SwEdEn, Gotland, Ollajvs Nature
Reserve, close to δjugarn, under Picea and Pinus on calcareous ground,
27 Sept. 2010, K. Hansen, K. Gillen & I. Olariaga, KH.10.278 (S). Clade 1
— dEnmark, Eastern Falster, Korselitze-forests, 5 Oct. 2007, H. Knudsen,
KH.07.46 (S). – SwEdEn, Uppland, Stockholm, Norra Järvafältet, Hansta
Nature Reserve, on rich ground under Corylus and Quercus, 8 Sept. 2010,
K. Hansen, K. Gillen & I. Olariaga, KH.10.193 (S); Uppland, Stockholm,
N Djurgården, Stora skuggan, on soil in grazed open oak forest, 12 Sept.
2008, J. Santos, JS.08.81 (S); Uppland, Uppsala, Hågadalen-Nåsten Nature
Reserve, Predikstolen, under Quercus robur, Picea abies, Corylus and Salix,
on rich bare ground, 17 Sept. 2009, K. Hansen & I. Olariaga, KH.09.170 (S).
Clade 2 — USA, Oregon, δincoln Co., Devil’s Punchbowl State Park, 13
εar. 1997, E.T. Peterson (OSC 56770); Washington, Pierce Co., εt Rainier
National Park, δower Tahoma Creek, under Pseudotsuga, Tsuga, Picea and
Calocedrus, 29 Oct. 1996, E.T. Peterson (OSC 56747); ibid., 30 Oct. 1996
(OSC 56754); ibid., 18 Oct. 1997 (OSC 56798); Washington, Snohomish
Co., Sloan Creek trail, 24 Sept. 1997, E.T. Peterson (OSC 56777). Clade 3a
— norway, Nord-Trøndelag, δeksvik, Gjøråsvika, on slope under Corylus and
Picea, on rich ground, 3 Sept. 2009, V. Kučera & I. Kautmanová, KH.09.135
(S). – SwEdEn, Södermanland, Nynäshamn, Herrhamra, on soil under Fagus,
in narrow forest area along the road, 19 Sept. 2013, K. Hansen & X.H. Wang,
KH.13.50 (S); Uppland, Norrtälje, δänna, under Corylus, 26 Aug. 2008,
J. Santos, JS.08.43 (S); Uppland, Uppsala, Hågadalen-Nåsten Nature
Reserve, Predikstolen, under Quercus robur, Picea abies, Corylus and
Salix, on rich, bare ground, 6 Sept. 2010, K. Hansen, K. Gillen & I. Olariaga,
KH.10.198 (S). Clade 3b — dEnmark, S Sjælland, εøn, Store Klinteskov,
Vestre Ulvemose, on calcareous soil in deciduous forest, 26 Sept. 1994,
K. Hansen & S.K. Sandal, KS-94-192 (C). – SwEdEn, Uppland, Uppsala,
Hågadalen-Nåsten Nature Reserve, Predikstolen, under Corylus, Populus
and Picea, on rich ground, 19 Sept. 2009, K. Hansen & I. Olariaga, KH.09.178
(S). Clade 4 — USA, Oregon, Benton Co., Corvallis, εcDonald-Dunn Research Forest, 10 Oct. 1997, E.T. Peterson (OSC 56782); ibid., 18 Nov. 1996
(OSC 56758); Oregon, Benton Co., Corvallis, west side of NW Beechwood
Place, scattered to clustered on rotting bark mulch and in thin grass under
Pseudotsuga menziesii, 17 Nov. 2010, N.S. Weber, NSW10200 (OSC
150345); Oregon, Corvallis, Witham Hill, 25 Nov. 1997, E.T. Peterson (OSC
56813); Oregon, Douglas Co., 20 Oct. 2010, J. Moore, εoorefun 19 (OSC);
Oregon, Douglas Co., Bear Gulch, under Pseudotsuga menziesii, 13 Jan.
1999, R. Davidson (OSC 67524); Oregon, Douglas Co., Slimewater, under
Pseudotsuga menziesii, Quercus garryana, Abies grandis, etc., 12 Sept.
1999, Frymire (OSC 72978); Oregon, Douglas Co., Umpqua, under Pseudotsuga menziesii, Abies concolor, Calocedrus decurrens, Corylus cornuta,
etc., 23 εar. 2000, E. Stewart (OSC 72979); under Pseudotsuga menziesii,
Pinus ponderosa, Calocedrus decurrens, Pinus lambertiana, Abies concolor,
etc., 15 Dec. 1999, C. Rusch (OSC 72176); Oregon, δane Co., Willamette
National Forest, εiddle Fork Ranger District, under Tsuga heterophylla, Thuja
plicata and Abies grandis, 18 Nov. 2002, Smith (OSC 119567).
Additional material of O. alutacea s.l. dEnmark, Sjælland, Hareskoven, N of
Copenhagen, 2 Aug. 1961, H. Dissing (C-F-48301). – FranCE, Orliénas, sous
Quercus et Cedrus atlantica, 13 Nov. 2008, B. Rivoire & N. Van Vooren, NV
2008.11.01 (dupl. S-F256976); Rhône, Bron, Parc de Parilly, 24 Sept. 2008,
J. Cavet, NV 2008.09.32 (dupl. S-F256974); Rhône, Courzieu, hameau des
Verchères, sous Pseudotsuga menziesii, 26 Oct. 2008, D. Carbonnel, NV
2008.10.02 (dupl. S-F256975). – italy, Puglia, εesagne (BR), Bosco δucci,
in soil, mainly under Quercus ilex, 12 Oct. 2010, M. Carbone (S-F257085).
– norway, Nord-Trøndelag, δeksvik, Gjøråsvika, mixed forest on rich ground,
I. Olariaga et al.: A monograph of Otidea
183
Fig. 8 Otidea alutacea s.str. (KH.09.133). a. Apothecia; b. spores in water †; c. paraphyses in water †; d. ectal excipulum in KOH†. — Scale bars = 10 µm;
= dried material.
†
3 Sept. 2009, K. Hansen & I. Olariaga, KH.09.137 (S); ibid., R. Braathen,
KH.09.139 (S); Nordland, Rana, Rausandaksla, in salico-betuletum, on limestone, 21 Sept. 1974, S. Sivertsen (C-F-60697). – Spain, εadrid, Arboreto
de ETSI εontes, under Quercus suber, Pinus pinea and Nerium oleander,
21 Nov. 2006, L. Rubio Casas (AH42204). – SwEdEn, Gotland, near Visby,
Värnhem, on rich ground under Fagus and Quercus, with Hepatica nobilis,
22 Sept. 2009, E. Bohus-Jensen, K. Hansen & I. Olariaga, KH.09.187 (S);
Gästrikland, Hofors, Sibbersbovägen, on rich ground under Corylus, 1 Sept.
2010, K. Hansen, K. Gillen & I. Olariaga, KH.10.154 (S); Jämtland, Ändsjön
Nature Reserve, in rich Picea forest, 26 Aug. 2009, H. Lindström, KH.09.97
(S); Närke, Ekeby, Kvarntorp, under a big Quercus by the road, 10 Sept.
2008, J. Santos & K. Hansen, JS.08.57 (S); Närke, Tysslinge, δatorpsbruk,
Grytsätterskogen, grassland with Quercus, 13 Sept. 2008, JS.08.76 (S);
Närke, Örebro, Hästhagen, by Svartån, mixed forest, 13 Sept. 2008, H. Kauffman, JS.08.74 (S); Skåne, Helsingborg, Fredriksdals Friluftmuseum, 16 Sept.
2010, G. Hamilton, KH.10.206 (S); Skåne, Kjugekull Nature Reserve, on bare
ground under Quercus rubra, Corylus and Fagus, 24 Sept. 2010, K. Hansen,
K. Gillen & I. Olariaga, KH.10.262 (S); Skåne, εaltesholm, forest close to the
castle, on the ground under Fagus, close to Alnus, 25 Sept. 2010, K. Hansen,
K. Gillen & I. Olariaga, KH.10.265 (S); Torne δappmark, Abisko, 12 Aug.
1974, M.D. Paulsen & N. Tams (C-F-48045); Uppland, Stockholm, Enebyberg, Rinkebyskogen, on bare soil in a ditch, under Picea, Betula and
Populus, 28 Aug. 2008, J. Santos, JS.08.50 (S); ibid., in deciduous forest
under Corylus, but also Tilia, Quercus and Betula, 1 Sept. 2008, J. Santos,
JS.08.56 (S); Uppland, Stockholm, Norra Järvafältet, Hansta Nature Reserve,
on naked soil among leaves, under large Corylus, also Quercus, 8 Sept.
2010, K. Hansen, K. Gillen & I. Olariaga, KH.10.189 (S); Uppland, Uppsala,
Hågadalen-Nåsten Nature Reserve, Predikstolen, on rich ground under
Quercus, Corylus, Populus tremula and Picea, 17 Sept. 2009, K. Hansen &
I. Olariaga, KH.09.173 (S).
Notes — We consider O. alutacea s.l. to comprise a species
complex. It is recognised by the medium brown, cup-shaped,
split apothecia, an ectal excipulum with only sparse resinous
exudates if any, and predominantly oblong spores. Although
sometimes treated as a well-delimited species (Harmaja
2009a), spore sizes of O. alutacea provided by different authors
vary considerably, e.g. 14–16 × 7–9 µm (Dissing 2000) or 12.5–
14.5 × 6.2–7.3 µm (Harmaja 2009a). In fact, O. cochleata has
been separated from O. alutacea on account of larger spores,
16–18 × 7–8 µm (Dissing 2000), or darker apothecia (εornand
& Courtecuisse 2005, δiu & Zhuang 2006, Zhuang 2006).
εeanwhile, two taxa of the O. alutacea species complex have
been separated in North America, based on apothecia colour,
and spore size and shape (Peterson 1998). Our δSU phylogeny
resolved several clades within O. alutacea s.l. (Fig. 1), which
are strongly supported in our multigene phylogeny (Hansen
& Olariaga 2015). It appears that the spore sizes within each
clade have a fairly narrow range, but overlap exists between
the clades. Patterns of continental speciation are suggested as
well; two clades have North American specimens (clade 2, 4),
and the rest contain samples from Europe and Asia.
Carbone (2010a) selected a lectotype and an epitype for O. alutacea. A spore range of 15.5–17 × 7 µm was given for the epitype specimen (Carbone 2010a), and based on this we assign
O. alutacea s.str. to the clade inferred from the phylogenetic
analyses encompassing this spore size. The description above
is based solely on the specimens of that clade, which are
characterised by the initially shallowly cup-shaped apothecia,
with rather light ochraceous brown hymenium, which later
becomes purple brown and more deeply cup-shaped. The
lectotypification of O. alutacea proposed by Parslow & Spooner
(2013) is superfluous.
Among the Eurasian clades, clade 1 contains North European
specimens characterised by small spores (12–13.5 × 5.5–7
µm, Lm = 12.2 –12.9 µm, Wm = 5.8 – 6.6 µm, Qm = 1.9 – 2.2),
non-overlapping with O. alutacea s.str. and clade 3. Clade 1
184
should be compared to O. kunmingensis, a taxon belonging
to the O. alutacea complex characterised by short spores
(Zhuang & Yang 2008). Clade 3a encompasses three specimens with spores sizes 13.5–15 × 6.5–8 µm (δm = 14.5–14.6
µm, Wm = 6.7–7.3 µm, Qm = 2 – 2.2) overlapping with those
of O. alutacea s.str. as described here. The apothecia differ
slightly macroscopically from O. alutacea s.str. in being deeply
cup-shaped in the beginning, reddish brown when young, later
pale ochre-brown. Clade 3b is composed of two collections
with larger spores, 15.5 –17.5 × 7.5 – 8 µm (δm = 16.1–17.4
µm, Wm = 7.7–8 µm, Qm = 2–2.3), but slightly overlapping with
O. alutacea s.str. and clade 3a. Clade 3b may correspond to
O. cochleata sensu Dissing (2000). The two North American
clades (2 and 4) comprise specimens with clearly non-over-
Persoonia – Volume 35, 2015
lapping spore sizes, 15 –18 × 7– 8 µm (δm = 15.5 –16.9 µm,
Wm = 7.3 –7.9 µm, Qm = 2 – 2.2) and 12 –14.5 × 6.5 – 8.5 µm
(δm = 12.5–14 µm, Wm = 6.6–7.5 µm, Qm = 1.8–2), respectively.
These two clades correspond to the two species distinguished
by Peterson (1998) in Western North America, as O. alutacea
(clade 2) and O. umbrina (clade 4). These species were said to
differ in colour of fresh apothecia. Clades 2 and 4 have spores
that overlap with European clades, and it is so far problematic
to distinguish them using only morphology and disregarding
the geographical origin.
To be able to fully clarify species boundaries within the O. alutacea complex, sampling of additional collections for molecular
study is needed. Distinguishing morphological and ecological
characters should be sought, especially through studying fresh
Fig. 9 Otidea papillata†. a. Apothecia; b. spores in water; c. paraphyses; d. resinous exudates in the medullary excipulum in water; e. ectal excipulum in KOH;
f. basal mycelium in KOH (a–d, f: H6003547, holotype; e: TUR 102134). — Scale bars = 10 µm; † = all dried material. — Photos: a. J. Kearey.
185
I. Olariaga et al.: A monograph of Otidea
material. Several names that belong to this complex, such as
O. alba, O. cinerascens, O. cochleata, O. felina and O. kunmingensis should be considered as this study is undertaken.
Otidea papillata clade
Apothecia cup-shaped, split. Receptacle surface with contrasting warts. Spores small, 9.5–11 µm. εedullary excipulum with
brown resinous exudates embedding some hyphae. Ectal excipulum poorly differentiated, of textura prismatica to textura
intricata. Resinous exudates on the ectal excipulum not dissolving in εδZ.
Species — Otidea papillata.
6. Otidea papillata Harmaja, Karstenia 15: 31. 1976 — Fig. 9
Holotype. Finland, Kainuu, Paltamo, εelalahti, εyllymäki, predominantly
coniferous grass-herb forest on distinctly calcareous soil, in litter mainly
composed of spruce needles, 23 Sept. 1971, H. Harmaja (H6003547) !
Apothecia 23–30 mm high, 7–33 mm wide, initially broadly earshaped, with upper margin rounded, then becoming cup-shaped,
split, stipitate or sessile. Hymenium ochre (5B5, 5B6) to yellowish ochre (4A5) when dried. Receptacle surface yellowish
brown (5C6, 5C7) when dried, warty. Warts conical, angular or
rounded, gregarious, distinctly darker than the background, dark
ochre brown to brown. Stipe 7–10 × 3–4 mm. Basal tomentum
and mycelium abundant, pale brownish ochre (5A3) to orangeochre (6A3). Spores broadly ellipsoid, seldom very slightly
inequilateral, with two large guttules, smooth, hyaline, 9.5 –
11(–11.5) × 5.5–6.5 µm (δm = 10–10.7 µm, Wm = 6.1–6.3 µm,
Qm = 1.6–1.7; n = 2). Paraphyses curved to tightly hooked, usually enlarged at apices, 3–4 µm wide, sometimes with 1–2
shallow notches, sometimes truncate or forked at apices,
when dried containing small, refractive, hyaline granules. Asci
116–165 × 9–10.5 µm. Apothecial section 900–1200 µm thick.
Subhymenium 70 – 90 µm thick, of dense textura intricata,
visible as a pale brown zone. Medullary excipulum of textura
intricata, 600–800 µm thick, hyphae 5.5–10.5 µm wide, thinwalled to slightly thick-walled, hyaline to very pale yellow, with
brown resinous exudates scattered among and covering some
hyphae, sometimes rod-shaped, paler and partially dissolving in KOH. Ectal excipulum of textura prismatica to textura
intricata, 70–100 µm, cells thin-walled, hyaline, 13–35 × 9–18
µm. Surface with conical to rounded warts, 65–100 µm high,
formed by short, fasciculate, hyphoid hairs, sometimes with
a gelatinous sheath. Resinous exudates abundant, yellowish
brown to reddish brown, paler in KOH, not dissolving in εδZ.
Basal mycelium of 2.5 – 4.5(– 6) µm wide, septate, hyaline
to very pale yellow hyphae, unchanged in KOH, smooth or
with regularly arranged, spheroid, yellow, resinous exudates,
sometimes embedded in a yellowish matter, dissolving in εδZ,
partially and more slowly in KOH.
Specimen examined. Finland, Varsinais-Suomi, Parainen, Petteby, Stornäset (Paltbacken), in coniferous forest among mosses, 30 Sept. 1990, T. Lindholm (TUR 102134).
Notes — Otidea papillata is only known from two Finnish
collections and its apothecial colours in fresh state are still
unknown. It is a distinct species, characterised by cup-shaped
apothecia with conspicuous warts on the outside, and small
spores (Harmaja 1976). Two diagnostic characters have been
observed in the two collections examined: 1) a very poorly
differentiated ectal excipulum of textura prismatica to textura
intricata, which is unique within the Otidea species studied by
us; and 2) scattered brown resinous exudates on the hyphae
of the medullary excipulum, that are somewhat reminiscent of
those in O. bufonia.
Harmaja (1976) emphasised the high warts on the outside of
the apothecia as a unique character for O. papillata, but two additional Otidea species, O. tuomikoskii and O. nannfeldtii, have
as high or higher warts. These two species have in fact been
confused with O. papillata (δundell et al. 1985, Van Vooren et
al. 2008). Otidea tuomikoskii is distinguished from O. papillata
by narrowly ear-shaped apothecia, a yellow reaction of the
excipulum in KOH, an ectal excipulum of textura angularis, and
by lacking pigmented exudates on the hyphae of the medullary
excipulum. Highly warted apothecia of O. nannfeldtii probably
resemble O. papillata more. However, O. nannfeldtii possesses
pigmented resinous exudates at the septa in the medullary excipulum, resinous exudates of the ectal excipulum that convert
into amber drops, and most importantly, an ectal excipulum of
textura angularis.
Otidea leporina clade
Apothecia ear-shaped, yellowish ochre to brown. Resinous exudates on the ectal excipulum converting into reddish grey drops
in KOH. Associated with conifers.
Species — Otidea leporina, O. pseudoleporina.
7. Otidea leporina (Batsch) Fuckel, Jahrb. Nassauischen Vereins Naturk. 23 –24: 330. 1870 ‘1869 –1870’ — Fig. 4a, 10
Basionym. Peziza leporina Batsch, Elench. Fung. 1: 117. 1783: Fr., Syst.
εycol. 2: 47. 1822.
≡ Scodellina leporina (Batsch) Gray, Nat. Arr. Brit. Pl. 1: 668. 1821.
≡ Helvella leporina (Batsch) Franchi, δ. δami & ε. εarchetti, Rivista
εicol. 1: 63. 1999.
≡ Helvella auricula Schaeff., Fung. Bavar. Palat. Nasc. 4: 103. 1774
(‘Elvela’).
≡ Wynnella auricula (Schaeff.) Boud., Icon. εycol. list prél. 600 sp.: (2).
1904.
≡ Otidea auricula (Schaeff.) Sacc., Syll. Fung. 8: 95. 1889.
Lectotype designated here: Schaeffer, Fung. Bavar. Palat. Nasc. 2: t. 156.
1763 (‘Elvela decima tertia’). Epitype designated here: SwEdEn, Jämtland,
Östersund, Andersön Nature Reserve, under Picea abies on rich ground, 28
Aug. 2009, K. Hansen & I. Olariaga, KH.09.93 (S); εycoBank εBT178082.
= Otidea leporina f. minor Rehm, Ber. Naturhist. Vereins Augsburg 26:
63: 1881.
≡ Otidea leporina var. minor (Rehm) Sacc., Syll. Fung. 8: 94. 1889.
Lectotype designated here: gErmany, δeipzig, in der Harth, in spruce
forest, Aug. 1873, G. Winter, Rehm Ascomyceten no. 251 (S-F88382) !
Isolectotype (UPS F-641412) !; εycoBank εBT178088.
= Otidea leporina f. major Rehm, Hedwigia 3–4: 2. 1883.
= Otidea leporina var. rubescens Velen., εonograph. Discom. Bohemiae
1: 354. 1934.
Lectotype designated here: CzECh rEpubliC, Kosoř near Prague, Sept.
1920, F. Fechtner (PRε 614790) !; εycoBank εBT200087.
= Otidea myosotis Harmaja, Karstenia 15: 32. 1976.
Holotype. Finland, Etelä-Karjala, Hamina, Vehkalahti, Pyhältö, mixed
forest, 3 Oct. 1970, L. Fagerström (H6003548) !
= Otidea crassa W.Y. Zhuang, εycotaxon 94: 366. 2006 (‘2005’).
= Otidea fuckelii ε. Carbone & Van Vooren, Rivista εicol. 52: 322. 2010
(‘2009’).
Holotype. auStria, Nassau, in pinetis umbrosis, Fungi Rhen. Exs. no.
1233 (G00110768). Isotype (S-F114092) !
Misapplied names
– Non Wynnella auricula sensu Boudier, Icon. εycol. livr. 26: n°. 535, pl. 250.
1909 (preliminary text with ‘circulaires’) (= Wynnella silvicola (Beck) Nannf.).
Apothecia gregarious or caespitose, 17–52 mm high, 4–25
mm wide, narrowly to broadly ear-shaped, split, stipitate or
sessile. Hymenium yellowish brown (5C6), cinnamon brown
(5D6), pale ochre brown (5A5, 5B6) to orange brown (6C7,
6D6), sometimes dark brown (5D8, 5F8, 6E8) when young,
seldom with pale pink stains (6A2), bruised margin orange
brown (6D8), when dried cinnamon brown (5A5, 5B5) to rusty
brown (6D7, 6D8). Receptacle surface ochre brown (5B6),
hygrophanous, in drying paler ochre brown (4A4, 4A5, 4B6),
186
Persoonia – Volume 35, 2015
Fig. 10 Otidea leporina*. a, b. Apothecia; c. spores; d. paraphyses; e. ectal excipulum with resinous exudates, insert shows close-up of amber drops on the
ectal excipulum in εelzer’s reagent; f. basal mycelium pale brown, with very small, regularly arranged, resinous exudates (a: KH.09.93, epitype; b: KH.09.102;
c–g: KH.11.02). — Scale bars = 10 µm; * = all fresh material.
when dried yellowish brown (5C7, 5C8), furfuraceous to finely
warty, seldom wrinkled at the base. Warts conical to flattened,
gregarious, concolorous, sometimes darker than the background, golden brown. Stipe 4–15 × 2–8 mm. Taste slightly
bitter. Basal tomentum and mycelium abundant, white to cream
white (5A3), sometimes very pale brown when dried. Spores
broadly ellipsoid, sometimes inequilateral, with two large
guttules, smooth, hyaline, (12 –)12.5 –14(–15) × 7– 8.5 µm
(δm = 12.8–13.8 µm, Wm = 7.5–8.2 µm, Qm = 1.6–1.8; n = 16).
Paraphyses curved to hooked, of the same width or slightly
enlarged at apices, 2.5–4 µm wide, without notches or with 1–3
low notches, seldom forked at apices, when fresh containing
small, refractive, light yellow guttules; when dried small, refrac-
tive, hyaline granules. Asci 170–215 × 9–10.5 µm. Apothecial
section 700–900 µm thick. Subhymenium c. 80–100 µm thick,
of dense textura intricata, visible as an orange-brown darker
zone, cells cylindrical to swollen. Medullary excipulum of textura
intricata, 300 – 550 µm thick, sometimes differentiated into
two parts: a) textura angularis underneath the subhymenium,
40–50 µm thick, cells 6–12 µm broad; b) textura intricata, hyphae 5–13 µm wide, sometimes slightly swollen, thin-walled
to slightly thick-walled, hyaline to very pale brown, sometimes
with yellow-brown resinous exudates at septa. Ectal excipulum
of textura angularis, 85–110 µm thick, cells thin-walled, pale
brown, 15–48 × 12–31 µm. Surface with broadly conical warts,
50–85 µm high, formed by short, fasciculate, hyphoid hairs,
I. Olariaga et al.: A monograph of Otidea
of 2 – 4 subglobose to elongated cells, constricted at septa,
10–16 µm wide, sometimes with a gelatinous sheath. Resinous
exudates abundant, yellow brown, sometimes dissolving in part
into amber drops or converting into reddish particles in εδZ,
dissolving into yellowish reddish grey heterogeneous drops in
KOH. Basal mycelium of 3–4.5(–6) µm wide, hyaline to pale
brown hyphae, unchanged in KOH, smooth or with very small,
regularly arranged, spheroid, pale brown, resinous exudates,
dissolving in εδZ, and partially in KOH.
Specimens examined. Canada, Québec, δe Verendrye Park, on ground
under spruce, 16 Sept. 1965, M.E. Elliot 65-123 (UPS F-629640). – CzECh
rEpubliC, Central Bohemian region, Zdice, Aug. 1924, F. Fechtner (PRε
614787, as O. felina); in piceto ad aciculos, Sept. 1925 (PRε 148836);
Prague-West district, Kosoř near Prague, Sept. 1920, J. Velenovský (PRε
614792, as O. umbrina). – dEnmark, Bornholm, Rø Plantage, coniferous forest, 29 Sept. 1985, W. Rummel (C-F-47633); N Jylland, δangdal Plantage
(near Tranum), under Juniperus, near Picea, 13 Aug. 2009, T. Læssøe,
Tδ-13769 (C); N Jylland, Rønhøj Plantage, 7 Oct. 1962, A. Hauerbach (C-F86691); Jylland, Virklund, Silkeborg Sønderskov, in moss, coniferous forest,
26 Sept. 1964, H. Dissing (C-F-48298); Sjælland, Bromme Plantage, N of
Sorø, under Picea, 9 Oct. 1965, H. Dissing (C-F-48299). – Finland, EteläHäme, εustiala, in pineto, 29 Aug. 1866, P.A. Karsten (UPS F-146429);
Perä-Pohjanmaa, Rovaniemi, Kaittiainen, acidic Picea forest, 11 Sept. 2011,
T. Kekki, TK407 (TUR); Perä-Pohjanmaa, Rovaniemi, Pisajärvi, old Picea
forest, 2 Sept. 2011, T. Kekki, TK304 (TUR); Perä-Pohjanmaa, Rovaniemi,
Välljoki, calciferous Picea forest, 25 Aug. 2011, T. Kekki, TK231 (TUR). –
FranCE, δoire, δa Chamba, au sol dans la litière d’aiguilles d’épicéa, 27 Sept.
2008, N. Van Vooren, NV 2008.09.28 (dupl. S). – gErmany, Thüringen, in
silvis abiegnis, Kl. & Op. (UPS F-629404, Klotzsch, Herb. Viv. εycol. 143).
– norway, Nord-Trøndelag, Kvam, Noem Northeast, under Picea, among
Rhytidiadelphus squarrosus, 2 Sept. 2009, H. Lindström, KH.09.131 (S);
Nord-Trøndelag, Namdalseid, Flåbekkåsen Nature Reserve, Picea and Pinus
old-growth forest, on acidic soil, among mosses, 4 Sept. 2009, K. Hansen &
I. Olariaga, KH.09.141 (S); Nord-Trøndelag, Steinkjer, Skrattåsen, in rich
Picea abies forest, 5 Sept. 2009, K. Hansen & I. Olariaga, KH.09.145 (S);
ibid., KH.09.147 (S). – SwEdEn, Härjedalen, Torkilstöten, δjungdalen, on
an active anthill in Picea forest, 19 Aug. 2011, J.C. Zamora & I. Olariaga,
KH.11.02 (S); Jämtland, in the surroundings of Sällsjö, in young stand of
Picea abies, with Betula and Salix, 29 Aug. 2009, K. Hansen & I. Olariaga,
KH.09.100 (S); ibid., KH.09.102 (S); Jämtland, SW of εörsil, Sandtjärndalen
Nature Reserve, under Picea abies on rich ground, 7 Sept. 2009, K. Hansen &
I. Olariaga, KH.09.156 (S); δappland, Ekopark Vuollerim, stream from Brännmyran, Picea forest, 28 Aug. 2008, M. Karström, εK0828 (S); δappland,
Jokkmokk, ‘Nornaskogen’ by Ållojaur, mossy Picea forest, on rich ground,
29 Aug. 2011, K. Hansen & I. Olariaga, KH.11.12 (S); ibid., KH.11.14 (S);
δappland, Jokkmokk, Ultevis Fjällurskog Nature Reserve, Sitoätno, near a
Picea, 31 Aug. 2011, K. Hansen & I. Olariaga, KH.11.33 (S); ibid., KH.11.36
(S); δappland, Kuouka, 15 km SE εessaure, herb-rich Picea forest on rich
ground, among mosses, 3 Sept. 2011, K. Hansen & I. Olariaga, KH.11.67
(S); δappland, S of Kvikkjobb-Kabla FUR Nature Reserve, by Kassavare
εt, Köpenhamn, under Picea, mossy place on acidic ground, 1 Sept. 2011,
K. Hansen & I. Olariaga, KH.11.76 (S); Närke, Knista, δekhyttan, Kungshall,
under Picea on calcareous ground, among mosses and litter, 12 Sept. 2008,
J. Santos, JS.08.065 (S); ibid., coniferous forest, with old Picea and Pinus,
K. Hansen, KH.08.108 (S); Skåne, δoshult, δilla δoshult, Picea forest, 5 Sept.
1998, S.-Å. Hanson, SÅH 105838 (C); Uppland, Stockholm, Enebyberg,
Rinkebyskogen, under Picea and Betula on acidic ground, 2 Sept. 2009,
K. Hansen & I. Olariaga, KH.09.169 (S); Uppland, Täby, Rönninge, close to
parking place by Arninge, under Picea on thick litter layer, 21 Sept. 2008,
J. Santos, JS.08.92 (S); Uppland, Uppsala, Ersta Nature Reserve, on soil
under Picea in young plantation, 23 Sept. 2008, J. Santos & K. Hansen,
JS.08.99 (S); Uppland, Uppsala, Sävja, Norra δunsen Nature Reserve, under
Picea, 28 Aug. 2008, J. Santos, JS.08.46 (S); Värmland, Gustav Adolf,
Hagfors, εalmbackarna, on moss, under Picea and Betula, 10 Aug. 2009,
F. Turander s.n. (S). – USA, California, Del Norte Co., δake Earl Wildlife
Area, 14 Dec. 1997, E.T. Peterson (OSC 56824); ibid., 15 Dec. 1997 (OSC
56825); ibid., under Picea sitchensis, Pinus contorta, Abies concolor, 26
Nov. 2001, M. Castellano & E. Cazares (OSC 108820); California, Humboldt
Co., Big δagoon Park, 14 Dec. 1956, A.H. Smith 56668 (UPS F-629302);
ibid., under Picea, 16 Dec. 1956, A.H. Smith 56799 (UPS F-629304); ibid.,
23 Dec. 1956, A.H. Smith 56954 (UPS F-629305); Colorado, Tolland, on
ground in coniferous woods, 28 Aug. 1920, F.B. Cotner (UPS F-629390);
Oregon, δincoln Co., Fogarty Creek State Park, 15 Oct. 1997, E.T. Peterson
187
(OSC 56784); Washington, Okanogan National Forest, Pasayten Wilderness,
under Picea engelmannii, Pseudotsuga menziesii, Abies lasiocarpa, Pinus
contorta, 16 Sept. 1999, R. Davis (OSC 108856).
Notes — Otidea leporina is probably the most common Otidea species in boreal coniferous forests of Europe. In the Alps
it often occurs together with Cudonia circinans, an association
not seen in Fennoscandia. It is characterised by ear-shaped,
brown apothecia, together with relatively broad spores that
are almost unique within Otidea. Otidea brunneoparva shares
similar spores, but differs from O. leporina in the darker brown
apothecia and strongly notched paraphyses. Other species of
Otidea that macroscopically resemble O. leporina are distinguished by different spore size and shape.
Our morphological and molecular study of the holotype of O. myosotis shows it is a synonym of O. leporina (Hansen & Olariaga
2015). The original description of O. myosotis (Harmaja 1976)
pointed out the apothecial shape and colours, and paraphyses
as diagnostic characters, all of which agree with our concept
of O. leporina. Recently, Harmaja (2009a) stated that the excipular resinous exudates in O. leporina convert in εδZ into
reddish particles and show no reaction in O. myosotis. In the
material of O. leporina examined by us, the resinous exudates
dissolve in part, can appear unchanged or can convert into
reddish particles. Sometimes, small amber-drops have also
been observed, though not as strikingly as in other species.
Therefore, it seems that the reaction of the exudates in εδZ
is variable within O. leporina, and cannot be used to separate
O. myosotis from O. leporina. Otidea crassa is a synonym
based primarily on the GenBank δSU sequence of the type
collection (DQ443444).
Nomenclatural notes — When Batsch (1783) described Peziza leporina, he referred to Schaeffer’s plate (1763), being unaware or disregarding Schaeffer’s later description of ‘Elvela’
auricula Schaeff. (1774) based on the same plate. Since Fries
(1822) sanctioned Batsch’s name, referring to H. auricula Schaeff.
(as P. auricula Schaeff.) as a synonym, P. leporina has priority.
While most authors have interpreted the plate by Schaeffer
(1763, t. 156) as a species that belongs to Otidea (e.g. Fuckel
1870, Rehm 1883, Bresadola 1898, Seaver 1904), others have
considered it to represent the monotypic Wynnella (Gonnermann & Rabenhorst 1869, Quélet 1886). Recently, Franchi et
al. (1999) stated that the Schaeffer plate represents W. silvicola
and as they consider Wynnella to be part of the genus Helvella,
they made the combination Helvella leporina. Carbone & Van
Vooren (2010) expressed doubts about how to interpret the protologue by Batsch (1783) and Schaeffer’s plate, and concluded
the name is ambiguous and recommended it not be used. Instead they introduced the new name O. fuckelii for the Otidea
species treated here. Based on our phylogenetic and morphological studies (see also Hansen & Olariaga 2015) this new
name is, however, superfluous, since O. myosotis and O. crassa
are shown to be synonyms of O. leporina.
The original Schaeffer plate shows several, more or less evenly
coloured, light ochraceus brown apothecia, conforming to O. leporina, and not bi-coloured apothecia (dark reddish brown with
a white base) as in W. silvicola. In our opinion, it leaves little
doubt it shows a species of Otidea. To settle the use of the name
O. leporina, and at the same time preserve the use of the wellestablished name, W. silvicola (Beck) Nannf. (Nannfeldt 1966),
we propose a modern epitype for Peziza leporina Batsch: Fr.
(Fig. 10a), which represents the Otidea species for which the
name has most often been used. The epitype is from Sweden
where Fries saw and studied living material, as indicated by
the abbreviation ‘v. v.’ (vidi vivam, seen living).
188
8. Otidea pseudoleporina Olariaga & K. Hansen, sp. nov. —
εycoBank εB808972; ITS barcode GenBank: Kε010112;
Fig. 5e, 11
Etymology. From ancient Greek ψ υ ο-, which means ‘false, fake’, referring to a close relationship with O. leporina.
Holotype. USA, Oregon, Douglas Co., E of εill Creek, under Pseudotsuga
menziesii, Abies concolor, Pinus lambertiana, 19 Oct. 2010, R. Helliwell,
rh101910 (OSC).
Misapplied names
– Otidea cantharella var. minor sensu Kanouse, εycologia 41: 667. 1949.
Apothecia gregarious, 10–30(–50) mm high, 8–22(–31) mm
wide, initially narrowly to broadly ear-shaped, margin rounded,
then expanding and sometimes becoming irregularly cup-shaped,
Persoonia – Volume 35, 2015
split, stipitate or sessile. Hymenium ochre-orange (4A6, 5A7) to
pinkish orange (5A6, 5A7), sometimes with pink spots or stains
(6A4), when dried orange-ochre (5A5, 5B5) to reddish brown
(6D7). Receptacle surface ochre-brown (5B5), hygrophanous,
in drying yellowish ochre (4A4, 4A5), when dried yellowish ochre
(5A5) to brownish ochre (5B5), furfuraceous to finely warty,
sometimes wrinkled at the base. Warts conical to rounded,
gregarious, concolorous, sometimes distinctly darker than
the background, reddish brown. Stipe 4–11 × 3–5 mm. Basal
tomentum and mycelium abundant, white to pale yellow (4A2)
or ochre (5A2). Spores ellipsoid, sometimes slightly inequilateral, with two large guttules, sometimes with up to 4 smaller
guttules, smooth, hyaline, (9.5–)10–12(–12.5) × 5.5–6.5 µm
(δm = 10.2–11.6 µm, Wm = 5.7–6.4 µm, Qm = 1.7–1.9; n = 6).
Fig. 11 Otidea pseudoleporina. a, b. Apothecia; c. apothecia †; d. spores in water †; e. paraphyses in water †; f. ectal excipulum in water † (a, d–f: rh101910,
holotype; b, c: εoorefun 14). — Scale bars = 10 µm; † = dried material. — Photos: a. R. Helliwell; b. c. J. εoore.
189
I. Olariaga et al.: A monograph of Otidea
Paraphyses curved to hooked, of the same width or slightly
enlarged at apices, 2.5 – 4.5 µm wide, sometimes with 1– 3
notches, apices seldom forked and rarely covered with a hyaline
coating, when dried containing small, refractive, yellow granules. Asci 155–231 × 9–10 µm. Apothecial section 900–1200
µm thick. Subhymenium c. 80–100 µm thick, of dense textura
intricata, visible as an orange-brown darker zone, of cylindrical
to swollen cells. Medullary excipulum 500–750 µm thick, differentiated into two parts: a) textura angularis underneath the
subhymenium, 150–200 µm thick, hyphae 8–18 µm broad;
b) textura intricata, hyphae 5–10(–18) µm wide, sometimes
slightly swollen, thin-walled to slightly thick-walled, very pale
yellow, sometimes with yellow-brown resinous exudates at
septa. Ectal excipulum of textura angularis, sometimes of a
textura prismatica, 80–110(–150) µm thick, cells thin-walled
to slightly thick-walled, pale yellow-brown, 13–37 × 8–27 µm.
Surface with broadly conical warts, 77–115 µm high, formed by
short, fasciculate, hyphoid hairs, of 2–3 subglobose to elongated cells, constricted at septa, 6–12 µm wide, sometimes with
a gelatinous sheath. Resinous exudates abundant, yellowbrown, dissolving into amber drops in εδZ, partially to entirely
dissolving into reddish grey heterogeneous drops in KOH.
Basal mycelium of 3 – 5 µm wide, very pale yellow hyphae,
unchanged in KOH, with very small, yellow resinous exudates,
regularly arranged, spheroid, dissolving in εδZ, partially and
more slowly dissolving in KOH.
Specimens examined. USA, California, Trinidad, under spruce, 30 Nov.
1956, A.H. Smith 56168 (UPS F-629690); Idaho, Idaho Co., Rickliff Creek
Public Camp, on the ground in Thuja-Tsuga woods, 10 Oct. 1947, W.B. Cooke
21227 (UPS F-629388); Idaho, Papoose Creek, Seven Devils εts, on the
ground in Douglas fir association, 3 Sept. 1954, A.H. Smith & H.E. Bigelow,
47346 (UPS F-629331); Oregon, Bear Springs, εt Hood National Forest, 18
Oct. 1947, A.H. Smith 27946 (UPS F-629430); Oregon, Benton Co., Corvallis,
west side of NW Beechwood Place, scattered to clustered on rotting bark
mulch and in thin grass under Pseudotsuga menziesii, 14 Nov. 2010, N.S.
Weber, NSW 10202 (OSC 150347); ibid., 17 Nov. 2010, N.S. Weber, NSW
10200 (OSC 150345); ibid., 27 Nov. 2010, NSW 10201 (OSC 150346);
Oregon, Douglas Co., E of εill Creek, under Pseudotsuga menziesii, Abies
concolor, Pinus lambertiana, 22 Oct. 2010, J. Moore, εoorefun 24 (OSC);
Oregon, Douglas Co., εill Creek, under conifers, 19 Oct. 2010, J. Moore,
εoorefun 14 (S); Oregon, Douglas Co., Roseburg District Bureau of δand
εanagement, under Pseudotsuga menziesii, Arbutus menziesii, Castanopsis
chrysophylla, 11 εay 1997, J. Klein (OSC 66261); Oregon, Douglas Co.,
south of δemolo δake, under conifers, 5 Nov. 2010, R. Heliwell, rh 179 (S);
Oregon, Douglas Co., Thorn Unit I, under Pseudotsuga menziesii, Tsuga
heterophylla, 21 Oct. 2010, C. Durbecq, Durbecq 16 (OSC); Oregon, Jackson
Co., εedford Bureau of δand εanagement, Ashland Resource Area, Beaver
Creek, under Pseudotsuga menziesii, Arbutus menziesii, Toxicodendron
diversilobum, Berberis piperiana, 18 Dec. 2000, R. Brock (OSC 119311);
Oregon, Jackson Co., εedford District Bureau of δand εanagement, Butte
Falls Resource Area, under Pseudotsuga menziesii, Calocedrus decurrens,
Pinus lambertiana, Pinus ponderosa, Quercus kellogii, Rhus diversiloba,
Berberis piperiana, Fragaria vesca, Moehringia macrophylla, grasses, 8 εar.
2000, M. Wineteer (OSC 72956); Oregon, δane Co., Willamette National
Forest, Blue River Ranger District, under Pseudotsuga menziesii, 18 Nov.
1999 (OSC 72296); Oregon, εarion Co., Breitenbuch Hot Springs Community, near Detroit Reservoir, in woods, 8 Nov. 1997, J.W. Spatafora (OSC
56809); Oregon, εt Hood, among moss under conifers, 15 Oct. 1922, L.E.
Wehmeyer (UPS F-629375); Oregon, Warm Springs R., εt Hood National
Forest, Skyline Trail, 29 Sept. 1947, A.H. Smith & W.B. Gruber, 27064 (UPS
F-629689); Washington, Clallam Co., Olympic National Park, Whiskey Bend
trailhead, 26 Nov. 1996, E.T. Peterson (OSC 56760); Washington, Fish Creek
Region, εt Rainier National Park, 25 Aug. 1948, E.G. Simmons 2067 (UPS
F-629332); Washington, lower slopes of Rampart Ridge, W. of δongmire, εt
Rainier National Park, 4 Sept. 1948, E.G. Simmons 2172 (UPS F-629386);
Washington, Park Creek, εt Baker National Forest, 9 Sept. 1941, A.H. Smith
16755 (UPS F-629820); Washington, Pierce Co., εt Rainier National Park,
δower Tahoma Creek, 29 Oct. 1996, E.T. Peterson (OSC 56749).
Notes — Otidea pseudoleporina is recognised by the broadly
ear-shaped apothecia, ochre-orange to pinkish orange hymenium and small spores. Our multi-gene phylogenetic analyses
(Hansen & Olariaga 2015) suggest O. pseudoleporina is the
sister species of O. leporina. They share ear-shaped apothecia and resinous exudates on the outer excipulum that partly
convert into heterogeneous reddish drops in KOH. Otidea
leporina differs in the brown apothecia and larger, broadly ellipsoid spores. Otidea pseudoleporina resembles O. nannfeldtii
and O. formicarum in the general apothecial shape and small
spores. Otidea nannfeldtii is distinguished by most often lacking
orange tones, having narrowly ear-shaped young apothecia,
and ectal excipular resinous exudates turning reddish brown in
KOH. Otidea formicarum is distinguished by having apothecia
devoid of orange tones, and spores with a lower Qm (1.6–1.7)
than O. pseudoleporina (1.7–1.9).
The material cited by Kanouse (1949) under O. cantharella var.
minor most likely represents O. pseudoleporina. Otidea cantharella var. minor as described by Boudier (1909a) has a pale
ochre or grey hymenium, citrine yellow outside, and veins at the
apothecial base, and represents a different species (see under
O. minor). Peterson (1998) treated under O. concinna material
that we refer to O. pseudoleporina and stated that Harmaja
(1974) used the name O. cantharella for the same species.
Otidea concinna is a well-known species in Europe, clearly
distinct from O. pseudoleporina (see O. concinna). As for the
name O. cantharella, the protologue describes a fungus with
the colour of Cantharellus cibarius (Fries 1822), and we typify
it with material of the large-spored species sometimes called
O. caligata (see O. cantharella). So far O. pseudoleporina is
only known from Western North America.
Otidea tuomikoskii clade
Apothecia ear-shaped. Receptacle surface with warts often
more than 100 µm high. Basal tomentum light ochre to orange
ochre. Spores small, 10–11 µm long. Sections of apothecia
turning yellow in KOH, especially the subhymenium and ectal
excipulum. Associated with conifers.
Species — Otidea tuomikoskii.
9. Otidea tuomikoskii Harmaja, Karstenia 15: 30. 1976 — Fig.
12
Holotype. Finland, Etelä-Häme, δammi, Pappilankylä, Koiransuolenoja, in
needles of Picea abies on an anthill, 9 Sept. 1972, R. Tuomikoski (H6002901) !
= Otidea papillata f. pallidefurfuracea Van Vooren & Hairaud, Bull. εycol.
Bot. Dauphiné-Savoie 188: 56. 2008.
Holotype. FranCE, Jura, δes Rousses, tourbière près du lac des Rousses,
au sol dans la litière d’aiguilles, sous épicéas (Picea abies), 19 Sept. 2007,
N. Van Vooren, NV 2007.09.27 (PC). Isotype (S) !
Misapplied names
– Otidea papillata sensu δundell, Nannfeldt & Holm, Fungi Exs. Suec. 66:
3282. 1985.
Apothecia gregarious to caespitose, 17–60 mm high, 7–30 mm
wide, long and narrowly ear-shaped, split, stipitate or sessile.
Hymenium pale whitish ochre (4A2, 4A3) to ochre yellow (4A4–
4A6) rarely with pink stains, when dried light ochre yellow (4A4)
to ochre (5B7, 5B8). Receptacle surface brownish ochre (4B7,
4C7) to yellow brown (5B6–5D6), hygrophanous, ochre yellow
(4A5, 4A6, 4B6) in drying, when dried yellowish brown (5C8,
5D8), warty, rarely wrinkled at the base. Warts conical, gregarious, brown, distinctly darker than the background or rarely lighter. Stipe 3–6 × 2–3 mm. Smell faintly aromatic. Basal tomentum
and mycelium abundant, light ochre (5A2) to orange-ochre
(5A4). Spores ellipsoid, slightly inequilateral, with two large
guttules, and sometimes with 1–4 smaller granules, smooth,
hyaline, (9.5–)10–11(–12) × 5.5–6.5(–7) µm (δm = 10.3–11.4
µm, Wm = (5–)5.5–6.5(–7.5) µm, Qm = 1.7–1.9; n = 15). Paraphyses curved to hooked, often broader at apices, 2.5–5(–6)
µm wide, sometimes with up to two shallow notches or forked at
190
Persoonia – Volume 35, 2015
Fig. 12 Otidea tuomikoskii*. a, b. Apothecia; c. spores; d. paraphyses; e. ectal excipulum; f. basal mycelium (a: JS.08.68; b–f: KH.11.77). — Scale bars = 10 µm;
* = all fresh material. — Photos: a. J. Santos.
apices, when fresh containing small to large, refractive, hyaline
to pale yellow guttules; when dried pale yellow. Asci 113–199
× 9–11.5 µm. Apothecial section 600–700(–1200) µm thick,
pale to bright yellow in KOH. Subhymenium c. 50–90 µm thick,
of dense textura intricata, visible as a yellowish brown zone.
Medullary excipulum of textura intricata, 300–500(–850) µm
thick, hyphae 3–13 µm wide, thin-walled to slightly thick-walled,
hyaline to very pale yellow, without resinous exudates at septa.
Ectal excipulum of textura angularis, 80–120 µm, cells thinwalled, hyaline to light yellow, 13–40 × 8–25 µm. Surface with
conical warts, 55–177 µm high, formed by fasciculate, short,
hyphoid hairs, of 3–8 globose to elongated cells, constricted at
septa, 7–15 µm wide, sometimes with a gelatinous sheath. Resinous exudates abundant, yellow-orange to yellowish brown,
dissolving into amber drops in εδZ, unchanged in KOH. Basal
mycelium of 3.5–6(–7.5) µm wide, often thick-walled, septate,
hyaline to very pale yellow hyphae, unchanged in KOH, with
regularly arranged, spheroid, yellow to orange resinous exudates, dissolving in εδZ, partially and more slowly in KOH.
Specimens examined. dEnmark, NW Jylland, Klim Bjerg, soil along forest road, S.A. Elborne & K. Hansen, 16 Sept. 1998, KH.98.92 (C-F-53155).
– EStonia, Põlvamaa, ad terram in piceto humida, 11 Aug. 1960, A. Elango
(UPS F-629392). – Finland, Perä-Pohjanmaa, Rovaniemi, Pisajärvi, old Picea
forest, 2 Sept. 2011, T. Kekki, TK305 (TUR). – FranCE, Charente-εaritime, île
de Ré, près du camping ‘δa Bonne Étoile’, under Pinus maritima and Quercus
ilex, on leaf litter, 26 Nov. 2006, M. Hairaud, NV 2006.11.05 (dupl. S); Rhône,
δes Halles, col de Croix-Régis, 25 Oct. 2006, J. Cavet, NV 2006.10.33 (dupl.
S); Rhône, Saint-Nizier-d’Azergues, forêt de Pramenoux, sous Picea abies,
20 Sept. 2008, N. Van Vooren, NV 2008.09.08 (dupl. S). – gErmany, Lower
I. Olariaga et al.: A monograph of Otidea
Saxony, δüneburg, Boitze, Pinus and Abies, Oct. 2010, M. Vega private herb.
s.n. (dupl. S-F256977). – norway, Nord-Trøndelag, Snåsa, Bergsåsen Nature
Reserve, under Picea and Pinus, 2 Sept. 2009, K. Hansen & I. Olariaga,
KH.09.130 (S). – Spain, Navarre, Orokieta, Loiandi, Picea abies plantation,
17 Oct. 2008, J.M. Lekuona (ARAN-Fungi A5041195). – SwEdEn, Lappland,
3 miles NW Vuollerim, Bombmurkleskogen, along the Stora δuleälven, herbrich Picea forest, 19 Aug. 2000, M. Karström, εK200065 (S); δappland,
Norrbotten, εessaure, Kaltisbäcken Nature Reserve, herb rich Picea forest,
3 Sept. 2011, M. Karström, KH.11.60 (S); εedelpad, Södra Sillre, Hussborg,
on wood, 22 Aug. 1998, K. Olofsson (S-F256896); Närke, Hidinge, δekhyttan, Katte εajaskogen, coniferous forest on lime rich soil, B. Wasstorp, 13
Sept. 2008, JS.08.77 (S); Närke, Snavlunda, Ö Snavlunda Nature Reserve,
under Picea in a mixed forest, 12 Sept. 2008, L.G. Hellsten & A. Stridvall,
JS.08.68 (S); Närke, Vintrosa, Kanterboda skans Nature Reserve, on soil
under conifers, 10 Sept. 2008, A.B. Nilsson, JS.08.60 (S); Skåne, δoshult,
Lilla Loshult, Picea forest with a few broadleaf trees (Betula, Quercus), 5 Sept.
1998, S.-Å. Hanson, SÅH 105768 (C); Södermanland, Nacka, Kvarnhagen
by Söderbysjön, on soil in shadow, moist area, under Picea, 26 Sept. 2008,
J. Santos, JS.08.100 (S); Uppland, Björklinge, Drälinge, amongst needles
and mosses under pine in coniferous woods, 10 Sept. 1936, H.G. Bruun &
H. Smith (S-F92983, Fungi Exs. Suec. 3282); Uppland, Trehörningsskogen
Nature Reserve, under Picea on rich ground, on needle litter and decayed
wood, 1 Sept. 2011, M. Prieto & I. Olariaga, KH.11.77 (S). – USA, California, Del Norte Co., Earl δake State Park, access by Sand Hill Road, 15
Dec. 1997, M. Madsen & R. Davis (OSC 56826); California, Humboldt Co.,
Trinidad, Nov. 1931, H.E. Parks 3749 (UPS F-629376); Oregon, Benton Co.,
Corvallis, εcDonald-Dunn Research Forest, under conifers, 23 Oct. 1996,
E.T. Peterson (OSC 56761); Oregon, Benton Co., Corvallis, west side of NW
Beechwood Place, scattered clusters of apothecia on duff and adjacent to
rotting wood under Pseudotsuga menziesii, 19 Nov. 1997, N.S. Weber, NSW
8553 (OSC 150344); Oregon, Douglas Co., Bureau of δand εanagement,
Roseburg District, Swiftwater Resource Area, under Tsuga heterophylla,
Pseudotsuga menziesii, Polystichum munitum, Berberis nervosa and Holodiscus tricolor, 8 Nov. 2000, R. Furriel (OSC 105550); Oregon, εarion Co.,
Salem District Bureau of δand εanagement, Cascades Resource Area,
under Pseudotsuga menziesii, Tsuga heterophylla, Gautheria shallon, Polystichum munitum, Berberis nervosa, Oxalis oregana, Acer circinatum, Alnus
rubra and Rhododendron macrophyllum, 12 Nov. 1997, K. Dougan (OSC
66350); Washington, Bremerton, 26 Oct. 1942, J.B. Flett (UPS F-629383);
Washington, Eatonville, 18 Oct. 1954, A.H. Smith 49143 (UPS F-629385);
Washington, δower Nisqually R., εt Rainier National Park, 2 Sept. 1948,
A.H. Smith 30888 (UPS F-629384).
Notes — Otidea tuomikoskii is characterised by the narrowly
ear-shaped apothecia, with high warts on the outside, small
spores, and the excipulum almost always turning yellow in
KOH, together with the ochre to orange-ochre basal tomentum
in dried specimens. Otidea nannfeldtii is probably the species
that resembles O. tuomikoskii most, but O. nannfeldtii has
lower warts, lacks orange tones in the basal tomentum, has
paraphyses only rarely with slightly swollen areas, and resinous exudates on the outer excipulum that turn reddish in KOH.
Otidea papillata shares with O. tuomikoskii conspicuous dark
warts on the outside of the apothecia (see under O. papillata).
The yellow KOH reaction of the excipulum, especially strong in
the subhymenium and ectal excipulum, has been observed to be
constant, although weak in some collections. The KOH reaction
is stronger in recent collections, and can also be macroscopically
observed in fresh apothecia. The holotype of O. tuomikoskii is
from an anthill (Harmaja 1976), but O. tuomikoskii most often
produces apothecia among needle litter or even on very decayed
wood, in coniferous forests. It is widespread in Europe, where it
occurs in coniferous plantations, and in Western North America
(Peterson 1998), and has been found in Asia (Cao et al. 1990).
Otidea cantharella clade
Apothecia ear-shaped, or cup-shaped and entire, usually clearly
stipitate. Spores exceeding 20 µm, biguttulate and with several
additional small guttules (except in O. brunneoparva). Paraphyses often strongly notched. Associated with Picea.
Species — Otidea brunneoparva, O. cantharella, O. propinquata.
191
10. Otidea brunneoparva K. Hansen, ε. Carbone, Olariaga &
Van Vooren, sp. nov. — εycoBank εB537590; ITS barcode
GenBank: Kε010026; Fig. 13, 14
Etymology. Harmaja (2009a) used the epithet brunneoparva to provisionally name this species. The name is validated here and refers to the small
size and brown colour of the apothecia.
Holotype. SwEdEn, Närke, Knista, δekhyttan, Kungshall, calcareous oldgrowth forest, in thick litter layer, with Picea and Pinus, 12 Sept. 2008, K. Hansen,
KH.08.107 (S). Isotype (C).
Apothecia gregarious, 12–35 mm high, 7–25 mm wide, initially
ear-shaped, apex subacute, broadly ear-shaped in the end, seldom almost cup-shaped, split, stipitate. Hymenium dark brown
(6F3–6F7), sometimes olivaceous brown (5D7, 5E7, 5E8) or
reddish brown (7F3–7F5), sometimes paler in the margin, purplish ochre (5B3, 5B4, 6D7), when dried olivaceous brown (5F4,
5F5, 5F7, 5E7) or dark brown (6E6, 7F7, 7F8). Receptacle surface concolorous or slightly lighter, dark brown (5F6, 6F7, 7F8),
slightly hygrophanous, in drying golden brown (5D6, 5D7), when
dried dark brown (6E7, 7E7) to cinnamon brown (5D7, 6D7,
6E7), furfuraceous, sometimes finely warty at the base, often
longitudinally wrinkled at the base, sometimes almost reaching the margin. Warts hemispherical, gregarious, concolorous,
sometimes darker or paler brown than the background. Stipe
4–7 × 2–3 mm. Basal tomentum and mycelium white to very
pale brown, when dried ochre brown. Spores ellipsoid to broadly
ellipsoid, sometimes very slightly inequilateral, with two large
guttules, seldom with 1–2 additional smaller granules, smooth,
hyaline, (11–)11.5–14(–15) × 6.5–8.5 µm (δm = 11.7–13.8 µm,
Wm = 7.1–8.3 µm, Qm = 1.6–1.7; n = 6). Paraphyses hooked,
often inrolled, of the same width or slightly enlarged at the
apices to 3–5.5 µm wide, often with clear notches or forked at
apices, when fresh containing refractive, pale yellowish brown
guttules, restricted to the uppermost part of the paraphyses;
when dried pale yellow. Asci 131–195 × 8–10 µm. Apothecial
section 700–800 µm thick. Subhymenium c. 100–120 µm thick,
visible as a darker brown zone, cells cylindrical to swollen,
densely arranged, with scattered brown resinous exudates at
septa. Medullary excipulum of loosely woven textura intricata,
300–500 µm thick, hyphae cylindrical to slightly swollen, thickwalled, 5–11 µm wide, hyaline to very pale brown, sometimes
with brown resinous exudates at septa. Ectal excipulum of
textura angularis, 80 –100 µm, cells thick-walled, yellowish
brown, 20–47 × 12–25 µm. Surface with broadly conical warts,
25–65 µm high, composed of fasciculate, short hyphoid hairs.
Non-warted parts with scattered hyphoid hairs, of 2–3 subglobose to elongated cells, 7.5–10.5 µm wide, slightly constricted
at septa, sometimes with a thin gelatinous sheath. Resinous
exudates abundant, yellowish to reddish brown, dissolving into
amber drops in εδZ. Basal mycelium of 3–4.5 µm wide, very
pale brown hyphae, unchanged in KOH, smooth or with very
small, resinous exudates, dissolving in εδZ.
Specimens examined. Finland, Etelä-Häme, Hämeen, δammi, Evo,
Kotinen virgin forest, mesic forest of the Myrtillus type, in needle litter of
Pinus sylvestris and Picea abies in basal part of an active anthill, 8 Sept.
1978, H. Harmaja (S-F249386); Kainuu, Paltamo, Saukkovaara, under
Picea abies in moist spring-fed site, nearly in water, 24 Aug. 2011, M. Lahti
(TUR-A 198582); Koillismaa, Kuusamo, Oulanka National Park, first part of
the Kiutaköngäs trail, on rich soil among Picea and Betula leaves, 25 Aug.
2008, M. Carbone (TUR-A 198581); ibid., 16 Aug. 2009 (S-F257086, dupl.
TUR-A 198579); ibid., 14 Aug. 2010 (TUR-A 198580). – SwEdEn, Jämtland,
Östersund, Ändsjön Nature Reserve, on an abandoned anthill in rich Picea
forest, 26 Aug. 2009, K. Hansen & I. Olariaga, KH.09.82 (S); Närke, Hidinge,
δekhyttan, Katte εajaskogen, coniferous lime-rich forest, in litter, 13 Sept.
2008, B. Wasstorp, JS.08.73 (S); Närke, Knista, δekhyttan, Kungshall, calcareous oldgrowth forest, in thick litter layer, with Picea and Pinus, 12 Sept.
2008, J. Santos, JS.08.66 (S); ibid., JS.08.69 (S).
Notes — Otidea brunneoparva is morphologically and genetically a clearly distinct species. It is macroscopically character-
192
Persoonia – Volume 35, 2015
Fig. 13 Otidea brunneoparva apothecia. a. KH.08.107, holotype; b. JS.08.66; c. TUR-A 198579; d. TUR-A 198580. — Photos: b. J. Santos; c, d. ε. Carbone.
ised by stipitate, broadly ear-shaped apothecia with dark brown
colours and sometimes olivaceous shades. εicroscopically, the
spore shape and size are diagnostic (Fig. 14a), although with
wide variation. The strongly inrolled and notched apices of the
paraphyses are very characteristic (Fig. 14b) and otherwise only
found in a few species like O. propinquata or O. daliensis. The
thick-walled, yellowish brown, angular cells forming the outer
excipulum are diagnostic too (Fig. 14c, d); such thick-walled
cells in the outer excipulum are only otherwise found in O. propinquata. The six ITS sequences of O. brunneoparva, from five
different localities in Finland and Sweden, are identical or with
3–8 bp differences. The ITS nucleotide diversity within O. brunneoparva, as sampled here, is 0.74 % per site. Phylogenetic
analyses of four gene-regions (Hansen & Olariaga 2015) show
O. brunneoparva forms a distinct monophyletic group with O. propinquata and O. cantharella. These three species are nevertheless, easily distinguished both macro- and microscopically.
The distinctive spores of O. brunneoparva are only otherwise
found in O. leporina within Otidea. Otidea brunneoparva is
clearly distinguished from O. leporina by the darker coloured
apothecia and notched paraphyses. Otidea bufonia, O. mirabilis
and O. smithii share dark brown colours with O. brunneoparva,
but clearly differ in apothecial shape and stature (O. brunneoparva being more delicate), narrower spores (δm = 6.3–7.3 µm,
Qm = 1.9–2.5) of different shape (fusoid in O. bufonia and O. mirabilis) and the resinous exudates on the ectal excipulum not dissolving into amber drops. Otidea fusconigra was published as a
provisional name too. It also resembles O. brunneoparva in the
dark brown apothecia, and spore size and shape (Jamoni 2004).
Nevertheless, the paler hymenium colour (‘grey caffelatte’),
the paraphyses without notches and the habitat among alpine
dwarf Salix, suggests O. fusconigra is a different species. ITS
and LSU sequences of O. fusconigra (collection GεFN 2293),
obtained by us, confirms O. fusconigra is not conspecific, but
a sister taxon to O. smithii.
Following Cao et al. (1990), O. brunneoparva keys out as
O. olivacea J.Z. Cao & δ. Fan. The cup-shaped, dark brown
apothecia, with olivaceous tinge, suggest these may be closely
related. However, the spores of O. olivacea were described
as considerably longer (14 –17 × 8 – 8.5 µm), almost nonoverlapping with those of O. brunneoparva. Unfortunately, we
were not able to get the type specimen of O. olivacea on loan for
study. It should be noted that O. olivacea is a later homonym of
O. olivacea Bucholtz. Therefore Harmaja (2009b) published the
new name O. olivaceobrunnea for the illegitimate O. olivacea.
Otidea pusilla Rahm might be conspecific with O. brunneoparva, but the name is not validly published since no type was
indicated (Art. 40.1 ICN) and more than one gathering was
cited (Art. 40.2 ICN) (εcNeill et al. 2012), ‘collected over three
weeks in the same site’. The description of O. pusilla agrees with
O. brunneoparva in the dark brown, cup-shaped apothecia and
the relatively broad spores. The spore size given in the protologue was ‘15/6–9 µ’ (Rahm 1958), which we find difficult to
interpret due to the unusually broad width range. Unfortunately,
no material could be traced in ZH (pers. comm. R. Berndt).
A collection named by Harmaja as O. brunneoparva (H6017193)
was found to be morphologically identical to our material. Also
ITS and δSU sequences confirm that Harmaja´s and our material are conspecific. Therefore the name O. brunneoparva is
here adopted and validated as a new species.
Otidea brunneoparva appears to be rather widespread in the
Scandinavian Picea forests. While at least two of our collections, as indicated by Harmaja (2009a) grew on anthills, the
rest were found in places with abundant Picea litter, often with
presence of Betula. Two localities, including the type locality,
were calcareous suggesting O. brunneoparva is calciphilous.
I. Olariaga et al.: A monograph of Otidea
193
Fig. 14 Otidea brunneoparva (KH.08.107, holotype). a. Spores in water †; b. paraphyses*; c. medullary and ectal excipulum in water †; d. wart from the ectal
excipulum in water †; e. ectal excipulum in εelzer’s reagent †; f. basal mycelium in water †. — Scale bars = 10 µm; * = fresh material; † = dried material.
11. Otidea cantharella (Fr.) Quél., Enchir. Fung.: 275. 1886
— Fig. 15
Basionym. Peziza cantharella Fr., Syst. εycol. 2: 48. 1822 : Fr., loc. cit.
≡ Flavoscypha cantharella (Fr.) Harmaja, Karstenia 14: 107. 1974.
Neotype designated here: SwEdEn, Jämtland, Östersund, Ändsjön Nature
Reserve, in rich Picea forest, with Hepatica nobilis and Oxalis acetosella, 31
Aug. 2009, K. Hansen & I. Olariaga, KH.09.125 (S); εycoBank εBT178085.
= Peziza caligata Nyl., Ex Not. Sällsk. Fauna Fl. Fenn. Förh. 10: 8. 1868
‘1869’.
≡ Otidea caligata (Nyl.) Sacc., Syll. Fung. 8: 95. 1889.
≡ Acetabula caligata (Nyl.) Boud., Hist. Classific. Discomyc. Europe: 41.
1907.
Holotype. Finland, Uusimaa, Helsinki, 1850, W. Nylander (H009215) !
Misapplied names
– Otidea abietina sensu Breitenbach & Kränzlin, Fung. Switzerland 1: 82.
1984.
– Peziza propinquata sensu Nannfeldt, Ann. Bot. Fenn. 3: 313. 1966.
Apothecia gregarious or caespitose, 23–68 mm high, 13–43
mm wide, initially broadly ear-shaped, in the end broadly earshaped to almost cup-shaped, split, stipitate. Hymenium initially
light brown (5B5, 5B6, 5C6), then ochraceous yellow (4A5, 4A6,
5A5), ochre-orange (4A7, 5B7), sometimes with pinkish areas or
red dots (6B8), when wounded pinkish and margin brownish red
(6B8), when dried orange brown (6D7, 6D8) to reddish brown
(6C6, 6D6). Receptacle surface concolorous, ochre brown (4B7,
5C7, 5B6), slightly hygrophanous, in drying yellowish ochre (4A6,
4A7), sometimes with pale brown stains, when dried orange
194
brown (6D7, 6D8), furfuraceous to finely warty, sometimes
wrinkled at the base. Warts hemispherical, gregarious, concolorous, sometimes darker than the background, brown. Stipe
4–25 × 3–9 mm. Basal tomentum and mycelium abundant,
white to very pale brown (5A3), very pale brown when dried.
Spores ellipsoid and often narrowing toward poles, sometimes
very slightly inequilateral, with two large and several smaller
guttules, smooth, hyaline, (17–)18–21 × (9–)10–11.5(–12) µm
(δm = 17.7–20 µm, Wm = 10.4–11.4 µm, Qm = 1.7–1.8; n = 13).
Paraphyses hooked, of the same width or slightly enlarged at
apices, 2.5–3.5(–5.5) µm wide, without or with a low notch,
seldom forked at apices, when fresh containing small, refractive,
pale yellow guttules; when dried small, refractive granules. Asci
191– 217 × 11–12 µm. Apothecial section 900–1200 µm thick.
Persoonia – Volume 35, 2015
Subhymenium c. 70–100 µm thick, visible as a darker yellow
zone, of cylindrical to swollen cells, densely arranged. Medullary excipulum of textura intricata, (500–)700 –900 µm thick,
hyphae cylindrical to slightly swollen, thick-walled, 3–12 µm
wide, very pale yellow, sometimes with yellow-brown resinous
exudates at septa. Ectal excipulum of textura angularis 90–120
µm, cells thin-walled, pale yellow, 23–55 × 12–25(–30) µm.
Surface with conical to broadly conical warts, 55–80 µm high,
formed by fasciculate, parallel, short hyphoid hairs, of 3–4 ovoid
cells, constricted at septa, 5–8 µm wide. Non-warted parts with
single hyphoid hairs, of 2–4 subglobose to elongated cells,
slightly constricted at septa, 7–10 µm wide, sometimes with
a gelatinous sheath. Resinous exudates abundant, yellow to
yellow-brown, dissolving into amber drops in εδZ. Basal my-
Fig. 15 Otidea cantharella*. a, b. Apothecia; c. spores; d. paraphyses; e. medullary excipulum showing resinous exudates at septa; f. ectal excipulum (a:
KH.09.125, neotype; b: KH.11.69; c, d: KH.10.152; e: KH.09.144; f: KH.09.155). — Scale bars = 10 µm; * = all fresh material.
I. Olariaga et al.: A monograph of Otidea
celium of 3.5–4.5(–6) µm wide, hyaline to very pale yellowish
brown hyphae, unchanged in KOH, smooth or with very small,
regularly arranged, spheroid, resinous exudates, dissolving in
εδZ and partially in KOH.
Specimens examined. Finland, Etelä-Häme, Hattula, Parola, Alppilankallio, Vaccinium myrtillus-type forest, under Picea abies, 25 Sept. 1967, P. Uotila
618 (H); Etelä-Häme, Janakkala, Tervakoski, 6 Sept. 1970, P. Uotila 6195
(H); Etelä-Häme, εustiala, 5 Sept. 1895, J. Lindroth (H6010923); ibid., in
pineto, 24 Aug. 1866, P.A. Karsten (H6010922); Etelä-Häme, εustiala, Tammela, 2 Sept. 1882, P.A. Karsten (H6010829); Perä-Pohjanmaa, Rovaniemi,
Kalkkinulkki, near old limestone quarry, under Picea, 23 Aug. 2011, T. Kekki,
TK211 (TUR); Perä-Pohjanmaa, Rovaniemi, Ounasvaara, Picea forest, 29
Aug. 2011, T. Kekki, TK279 (TUR); Perä-Pohjanmaa, Rovaniemi, Välijoki, calciferous Picea forest, 25 Aug. 2011, T. Kekki, TK236 (TUR); Perä-Pohjanmaa,
Tervola, Peura, old calciferous Picea forest, 5 Sept. 2011, T. Kekki, TK177
(TUR); Perä-Pohjanmaa, Ylitornio, Kuusikkorommas, calciferous Picea and
Pinus forest, 2 Sept. 2011, T. Kekki, TK301 (TUR); Varsinais-Suomi, Vihti,
Nummela, εetsäkulma, on soil, 16 Sept. 1979, H. Kotiranta (H6010925).
– FranCE, Isère, Villard-de-δans, Bois Barbu, under Picea abies, 20 Sept.
2008, J. Cavet, NV 2008.09.16 (dupl. S). – hungary, Hohe Tatra bei UnterSchmecks, Oct. 1884, Linhart (UPS F-629314, Rehm Ascomyceten 251b,
as O. leporina f. minor). – italy, Trentino-Alto Adige, Cavelonte, in silvis
coniferis, semper socia Cudoniae confusae, Aug. 1898, G. Bresadola (UPS
F-629361). – norway, Nord-Trøndelag, Steinkjer, Noem, under Picea, 2 Sept.
2009, K. Hansen & I. Olariaga, KH.09.129 (S); Nord-Trøndelag, Steinkjer,
Strattåsen, in rich Picea forest, 5 Sept. 2009, K. Hansen & I. Olariaga,
KH.09.144 (S). – SwEdEn, Blekinge, Rödeby, Spjutsbygd, c. 2 km NW from
the train station, on needle litter under a Picea in coniferous forest, 10 Sept.
1946, S. Lundell & S. Wikland (UPS F-146484); Gästrikland, close to Bergby,
mossy Picea forest on acidic ground, 31 Aug. 2010, K. Hansen, K. Gillen &
I. Olariaga, KH.10.152 (S); Hälsingland, Kårböle, St Olofs, 26 Aug. 2001,
H.-G. Toresson s.n. (S); Hälsingland, south to crossing between roads E45
and 310, under Picea abies, among leaf litter, 21 Aug. 2011, J.C. Zamora
& I. Olariaga, KH.11.109 (S); Härjedalen, δinsell, Djursvallen, montane coniferous forest with birch, on very rotten log, 13 Aug. 2000, B. Gahne (UPS
F-125589); Härjedalen, Torkilstöten, under Picea abies among leaf litter,
on acidic ground, 20 Aug. 2011, J.C. Zamora & I. Olariaga, KH.11.106 (S);
Jämtland, Hammarstrand, Picea mossy forest, 26 Aug. 2009, H. Lindström,
KH.09.83 (S); ibid., KH.09.84 (S); Jämtland, SW of εörsil, Sandtjärndalen
Nature Reserve, under Picea abies on rich ground, 7 Sept. 2009, K. Hansen
& I. Olariaga, KH.09.155 (S); Jämtland, Sällsjö surroundings, in young
stand of Picea abies, on rich ground with Betula and Salix, 29 Aug. 2009,
K. Hansen & I. Olariaga, KH.09.104 (S); Jämtland, Östersund, Andersön
Nature Reserve, Picea forest on rich ground, 28 Aug. 2009, K. Hansen &
I. Olariaga, KH.09.95 (S); ibid., under Picea and Pinus, KH.09.96 (S); ibid.,
KH.09.101 (S); Jämtland, Östersund, Fillstabäcken Nature Reserve, on old
anthill under Picea, K. Hansen & I. Olariaga, 30 Aug. 2009, KH.09.111 (S);
ibid., KH.09.117 (S); ibid., K. Hansen & X.H. Wang, 5 Sept. 2012, KH.12.99
(S); Jämtland, Östersund, Ändsjön Nature Reserve, in rich Picea forest, with
Hepatica nobilis and Oxalis acetosella, 26 Aug. 2009, K. Hansen & I. Olariaga,
KH.09.78 (S); ibid., KH.09.80 (S); ibid., KH.09.98 (S); Jämtland, Åre, Kall,
along Stor-Grundsviken, Kallsjön, on old anthill in lime rich Picea forest, 15
Aug. 2008, J. Santos & K. Hansen, JS.08.18 (S); δappland, Jokkmokk, SE
of Vuollerim, Andersviksravinerna (Natura 2000 area), mixed forest, 31 Aug.
2011, A. Stridvall, KH.11.111 (S); ibid., KH.11.112 (S); δappland, Jokkmokk,
by Kassavare mountain, Köpenhamn, under Picea abies, 1 Sept. 2011,
K. Hansen & I. Olariaga, KH.11.110 (S); δappland, Jokkmokk, ‘Nornaskogen’
by Ållojaur, mossy Picea forest on rich ground, 29 Aug. 2011, K. Hansen &
I. Olariaga, KH.11.16 (S); δappland, Kuouka, 15 km SE εessaure, herb-rich
Picea forest on rich ground, among mosses, 3 Sept. 2011, K. Hansen &
I. Olariaga, KH.11.68 (S); ibid., KH.11.69 (S); δappland, 5 km SE Vuollerim,
part of Andersviksravinerna, Rävabacken Nature Reserve, herb-rich Picea
forest, 21 Aug. 2000, M. Karström, εK200061 (S); ibid., under Picea on
mossy ground, together with Cudonia confusa, 2 Sept. 2011, K. Wiking,
KH.11.59 (S); Småland, Ryssby, Gärdsholmen, Björnö, coniferous forest,
6 Sept. 1930, H.G. Bruun (UPS F-146485); ibid., on deep moss in coniferous forest (UPS F-146486); Uppland, Uppsala, Sävja, Norra δunsen Nature
Reserve, Picea forest, on soil, thick layer of litter and mosses, 28 Aug. 2008,
J. Santos, JS.08.47 (S). – SwitzErland, Wallis Canton, δiddes, Palazuit, in
side of brook under Picea abies, 17 Aug. 2008, M. Carbone (εCVE 24217).
Notes — Otidea cantharella can be recognised by broadly
ear-shaped apothecia, with yellow to orange tones, often with
a well-developed stipe, and large spores. No other species of
Otidea have the spore size of O. cantharella, in combination
with those macroscopic characters.
195
Otidea cantharella is associated with Picea abies, and often
produces apothecia on anthills or thick needle layers. Furthermore, we have often found apothecia of O. cantharella and
Cudonia confusa together (Fig. 15a), and once with Spathularia
rufa, both species belonging to the Rhytismatales. An apparent
closer association in the same fairy ring has also been observed
once. Interestingly, Bresadola noted the association with C. confusa in one of his collections (UPS F-629361) studied by us. The
presence of C. confusa in the same collecting spot may give
a first plausible field identification of O. cantharella. εoreover,
it suggests a possible biotrophic association between O. cantharella and C. confusa.
Nomenclatural notes — This species has been referred to
as O. caligata (Nyl.) Sacc. (Nannfeldt 1966, Dissing 2000).
Nevertheless, Harmaja (2009a) came to the conclusion that
the name O. cantharella must refer to the large-spored species
treated here, with which some authors have later disagreed
(Carbone 2010b, Van Vooren 2011b). Harmaja stressed the
following characters from the protologue supporting the usage
of the name as presented here: ear-shaped, stipitate apothecia,
the yellow colour of Cantharellus cibarius and the occurrence
in Picea forests in southern Sweden. We have studied material
from Småland (UPS F-146485, UPS F-146486), from where
Fries described O. cantharella. It is therefore very likely that
Fries had in mind the species described here when he coined
the name O. cantharella, although he had only seen dried
material, as indicated by the abbreviation ‘v. s.’ (vidi siccam,
seen dried). Since no original material is known to exist, we
propose a neotype that will attach the name O. cantharella to
the Otidea species described here. This stabilises the interpretation proposed by Harmaja (2009a), also used by εornand &
Courtecuisse (2005), and long before adopted by Bresadola
(1900: 102), and should serve to settle the interpretation of
O. cantharella. Our ITS-δSU sequences of four O. cantharella
collections from Sweden and France are identical.
12. Otidea propinquata (P. Karst.) Harmaja, Karstenia 15: 32.
1976 — Fig. 16
Basionym. Peziza propinquata P. Karst., Not. Sallsk. Fauna Fl. Fenn.
Forh. 10: 110. 1869.
Lectotype. Finland, Tavastland, εessuby, 7 Oct. 1860, P.A. Karsten
(H6010807) !, selected by Nannfeldt (1966).
= Otidea abietina f. nigra Rick, Oesterr. Bot. Z. 48: 62. 1898.
≡ Otidea abietina var. nigra (Rick) Sacc., Syll. Fung. 14: 746. 1899.
Lectotype. auStria, Vorarlberg, an der Gamp, im Nadelwald, 1700 m,
Sept. 1897, Rick (S-F9962) !, indicated by Nannfeldt (1966).
= Otidea indivisa Velen., εonograph. Discom. Bohemiae 1: 355. 1934.
Lectotype. CzECh rEpubliC, Karlštejn N, Oct. 1922, J. Fechtner (PRε
149147) !, selected by Nannfeldt (1966).
Misapplied names
– Pseudotis abietina sensu Boudier, Icon. εycol. livr. 7: nº. 131, pl. 333.
1906 (preliminary text with ‘circulaires’).
– Otidea cochleata sensu Breitenbach & Kränzlin, Fung. Switzerland 1: 84.
1984.
Apothecia gregarious or caespitose, 7–20 mm high, 15–35
mm wide, obconical to broadly cup-shaped, entire, very rarely
split, regular or sometimes undulate in the margin, stipitate.
Hymenium ochre brown (5C7, 6D5, 6D6) to dark reddish brown
(6F6, 6F7, 7F7, 7F8), when dried ochre brown (6B7, 6C7).
Receptacle surface concolorous, orange brown (6F6, 6F7),
slightly hygrophanous, in drying yellowish brown (5C6, 5C7),
when dried orange brown (6D6, 6D7, 6E7), furfuraceous to
warty, sometimes wrinkled at the base. Warts hemispherical,
gregarious, concolorous, sometimes darker than the background, brown. Stipe 7–16 × 2–7 mm. Basal tomentum and
mycelium abundant, white to very pale brown (5A3), very pale
brown when dried. Spores ellipsoid and often narrowing toward
196
Persoonia – Volume 35, 2015
Fig. 16 Otidea propinquata. a. Apothecia; b. spores in water †; c. paraphyses in KOH †; d. wart of the ectal excipulum in water †; e. ectal excipulum in KOH †;
f. basal mycelium in water† (a: KH.11.21; b–f: KH.09.99). — Scale bars = 10 µm; † = dried material.
the poles, sometimes very slightly inequilateral, with two large
and several smaller guttules, smooth, hyaline, (18–)19–21 ×
10–12.5 µm (δm = 19.3–20 µm, Wm = 10.9–11.6 µm, Qm = 1.6–
1.7; n = 4). Paraphyses hooked, of the same width or often
enlarged at apices, 3–5 µm wide, often with 1–3 notches or
forked at apices, when fresh containing small, refractive, light
yellow guttules; when dried hyaline. Asci 231– 275 × 12–16
µm. Apothecial section 1100–1500 µm thick. Subhymenium
c. 100–120 µm thick, of dense textura intricata, visible as a
darker brown zone. Medullary excipulum of textura intricata,
600–800 µm thick, hyphae 4–9 µm wide, sometimes slightly
swollen, thin- to thick-walled, very pale brown, sometimes with
brown resinous exudates at septa. Ectal excipulum of textura
angularis 100–130 µm, cells rather thick-walled, pale brown,
17–50 × 10–40 µm. Surface with conical to broadly conical
warts, 50–75 µm high, formed by short, fasciculate hyphoid
hairs, of 2 – 3 subglobose to elongated cells, constricted at
septa or not, 8–11 µm wide, sometimes with a pale brown thick
gelatinous sheath. Resinous exudates abundant, yellow-brown
to brown, turning brownish red in KOH, dissolving in εδZ. Basal
mycelium of 3–6 µm wide, hyaline to pale brown hyphae, unchanged in KOH, smooth or normally with regularly arranged,
spheroid to rod-shaped, resinous exudates, dissolving in εδZ
and partially in KOH.
Specimens examined. CzECh rEpubliC, Praha, Karlštejn, 9 Oct. 1922,
F. Fechtner (UPS F-629369, syntype of O. indivisa); Sept. 1924 (UPS F-629367,
syntype of O. indivisa). – dEnmark, NE Sjælland, Asserbo Plantage, under
Pinus and Picea among needles, 7 Oct. 1975, H. Knudsen (C-F-87203).
I. Olariaga et al.: A monograph of Otidea
– FranCE, Isère, δans-en-Vercors, on the ground, on Picea leaf litter, 17
Sept. 2008, J. Cavet, NV 2008.09.15 (dupl. S). – italy, Trentino-Alto Adige,
Sopramonte, ad acus abiegnos in sylvis coniferis, Aug. 1898, G. Bresadola
(UPS F-629357). – SwEdEn, Jämtland, Sällsjö surroundings, in young stand
of Picea abies, on rich ground, 28 Aug. 2009, K. Hansen & I. Olariaga,
KH.09.99 (S); Jämtland, Östersund, Andersön Nature Reserve, under Pinus
and Picea, on rich ground, among mosses, 28 Aug. 2009, K. Hansen &
I. Olariaga, KH.09.94 (S); ibid., KH.09.103 (S); Jämtland, Östersund, Ändsjön
Nature Reserve, in rich Picea forest, 26 Aug. 2009, K. Hansen & I. Olariaga,
KH.09.81 (S); ibid., 31 Aug. 2009, KH.09.123 (S); δappland, Jokkmokk,
‘Nornaskogen’ by Ållojaur, mossy Picea forest on rich ground, 28 Aug.
2011, K. Hansen & I. Olariaga, KH.11.21 (S); δappland, Jokkmokk, Ultevis
Fjällurskog Nature Reserve, Sitoätno, Picea mossy forest on rich ground,
among leaf litter, 31 Aug. 2011, K. Hansen & I. Olariaga, KH.11.30 (S); ibid.,
KH.11.35 (S); δappland, S of Kvikkjobb-Kabla FUR Nature Reserve, by
Kassavare mountain, Köpenhamn, young Picea stand by a road, 1 Sept.
2011, K. Hansen & I. Olariaga, KH.11.53 (S); δappland, εessaure, herbrich Picea forest, 29 Aug. 2008, M. Karström, εK0834 (S); δappland, 17 km
WSW of Vuollerim, Slubbojaureskogen, herb-rich Picea forest, 8 Aug. 2002,
M. Karström, εK0222 (S); Närke, Knista, δekhyttan, under Picea, in thick litter
layer, in lime rich forest, 12 Sept. 2008, J. Santos, JS.08.67 (S); Uppland,
Uppsala, Sätra Nature Reserve, on litter on lime rich soil, under Picea, 23
Sept. 2008, J. Santos & K. Hansen, JS.08.98 (S). – USA, Washington, δake
Crescent, under fir, 28 Oct. 1935, A.H. Smith (UPS F-629351).
Notes — Otidea propinquata is easily recognised by the
stipitate, entire, brown apothecia, large spores, and notched
or forked paraphyses. Otidea daliensis shares brown, entire
apothecia with O. propinquata, but differs in having sessile
apothecia, with purplish brown tones, and basal mycelium that
lacks abundant resinous exudates.
There has been uncertainty surrounding the correct name of
this taxon. A number of early authors used the epithet abietina
to refer to O. propinquata (Boudier 1906, Bresadola 1933). In
agreement with Harmaja (2009a) and Carbone (2010c), we
consider Peziza abietina a nomen confusum. Nannfeldt (1966)
referred to O. propinquata as O. indivisa and argued that the
type of O. propinquata was conspecific with O. cantharella (as
caligata). Harmaja (1976) examined the lectotype of O. propinquata and found it to be clearly distinct from O. cantharella and
O. indivisa, a later synonym. This is confirmed by our study of
the type of O. propinquata. Otidea propinquata occurs in Picea
forests on calcareous ground. It is widespread in Northern Fennoscandia and present in central Europe.
Otidea formicarum clade
Apothecia ear- to cup-shaped, split, ochre to reddish brown.
Spores small, 9.5–12 µm long. Basal mycelium with abundant
yellow resinous exudates. Associated with conifers.
Species — Otidea formicarum, O. nannfeldtii, O. subformicarum, O. aff. subformicarum, Otidea sp. ‘b’.
13. Otidea formicarum Harmaja, Karstenia 15: 31. 1976 — Fig.
5f, 17
Holotype. Finland, Etelä-Karjala, εiehikkäla, Savanjärvi, on anthill in
spruce forest, 26 Sept. 1970, L. Fagerström (H6003549) !
Apothecia gregarious to caespitose, 8–22 mm high, 5–20 mm
wide, broadly ear-shaped, upper margin rounded, then expanding and sometimes in the end becoming cup-shaped and flattened, split, stipitate or sessile. Hymenium yellowish brown
(5C7, 5C8) to reddish brown (6C7, 6C8, 6D8), when dried ochre
(4A5, 4A6) to brownish ochre (5B7). Receptacle surface yellowish brown (5C6) to reddish brown (6C6), hygrophanous,
in drying ochre (5B7), when dried ochre (4A5, 4A6), finely
warty, smooth at the base. Warts broadly conical to rounded,
gregarious, concolorous, sometimes slightly darker than the
background, brown. Stipe 3 – 6 × 2–4 mm. Basal tomentum
and mycelium abundant, light yellow-ochre (5A2). Spores ellip-
197
soid, seldom very slightly inequilateral, with two large guttules,
smooth, hyaline, 9.5–11(–11.5) × (5.5–)6–7 µm (δm = 10–10.7
µm, Wm = 6–6.9 µm, Qm = 1.6–1.7; n = 7). Paraphyses curved
to hooked, of the same width or slightly enlarged at apices, 3–4
µm wide, with up to two slightly swollen areas, occasionally
notched, sometimes when fresh containing small, refractive,
light yellow or green guttules; when dried small, refractive,
hyaline to light yellow granules. Asci 160–232 × 10.5–13 µm.
Apothecial section 700–800 µm thick. Subhymenium c. 80–100
µm thick, of dense textura intricata, visible as a yellowish brown
zone, cells cylindrical to swollen, densely arranged. Medullary
excipulum 400–500 µm thick, differentiated into two parts: a)
textura angularis underneath the subhymenium, 120–200 µm
thick, cells 6–15 µm broad; b) textura intricata, hyphae sometimes slightly swollen, thin-walled to thick-walled, 3–11 µm wide,
hyaline to very pale yellow, sometimes with pale yellow resinous exudates at septa. Ectal excipulum of textura angularis,
70–120 µm, cells thin-walled, light yellow, 17–40 × 11–25 µm.
Surface with broadly conical warts, 45–60 µm high, formed by
short, fasciculate, hyphoid hairs, of 2–3 elongated cells, 7–12
µm wide, not or slightly constricted at septa, sometimes with
a gelatinous sheath. Resinous exudates abundant, brownish
yellow, dissolving into amber drops in εδZ. Basal mycelium of
3–5 µm wide, hyaline to very light yellow hyphae, unchanged
in KOH, with regularly arranged, very small, spheroid resinous
exudates, dissolving in εδZ, partially and more slowly in KOH.
Specimens examined. Finland, Etelä-Häme, δammi, Evo, Vahtervehmas,
Kotinen virgin forest, 6 Sept. 1988, H. Harmaja (H6003551); Etelä-Häme,
δoppi, Topeno, Piimästennummi, on a huge anthill, under Picea, 16 Sept.
2011, S. Huhtinen 11/65 (TUR); Perä-Pohjanmaa, Rovaniemi, Välijoki, calciferous Picea forest, on anthill, 25 Aug. 2011, T. Kekki, TK223 (TUR); Uusimaa,
Elimäki, Villikkala, δääksynmäki, mesic heath spruce forest, on anthill, 22 Oct.
2005, U. Nummela-Salo & P. Salo (H6003550); Varsinais-Suomi, Koski Tl.,
Hongisto, on old anthill under Picea abies, with Betula sp., Salix caprea and
Pinus sylvestris, 8 Aug. 1998, M.-L. & P. Heinonen (TUR 124728); VarsinaisSuomi, δieto, Suopohja, SE of Päivärinne, along the road by the house EskoUkura, on old anthill under Picea abies, 2 Oct. 2009, K. Ruottinen (TUR-A
183242). – FranCE, Haute Savoie, Thorens-Glières, plateau des Glières,
under Picea, on the ground, 22 Sept. 2006, L. Francini, NV 2006.09.11 (S).
– norway, Nordland, Grane, Holmvassdalen Nature Reserve, on old anthill
under Picea, 29 Oct. 2009, J. Lorås (S-F244372). – SwEdEn, Dalarna, Stora
Tuna, between Falubäcken and Övre εorbygge fäbod, abundant in the lower
part of an active anthill, 22 Sept. 1963, R. Morander (UPS F-146725); Närke,
Askersund, Orkarebäckens Nature Reserve, calcareous forest, on an active anthill under Picea, 11 Sept. 2008, J. Santos, JS.08.63 (S); ibid., under
Picea in litter, JS.08.62 (S); Uppland, Bondkyrka, Gottsundabergen, 1 Oct.
1927, J.A. Nannfeldt (UPS F-146706); Uppland, Bondkyrka, Nåsten, S of
δäbyvad station, among coniferous needles, 16 Sept. 1932, J.A. Nannfeldt
(UPS F-146709); Uppland, Älvkarleby, Billuddens Nature Reserve, under
Pinus and Picea on sandy ground, at the base of a Pinus, 15 Sept. 2011,
J.C. Zamora & I. Olariaga, KH.11.104 (S); Ångermanland, Ullanger, Håll, in
the eastern slope of εt εoberget, c. 0.5 km SW of p. 35,29, on the ground,
14 Sept. 1974, R. Moberg (UPS F-146735).
Notes — Otidea formicarum is characterised by relatively
small, reddish brown, broadly ear-shaped apothecia, small
spores, and by its habitat, often occurring on anthills. For a
comparison with the sister species O. subformicarum see under
that species below. Otidea nannfeldtii resembles O. formicarum,
but differs in often having more narrowly ear-shaped and paler
coloured apothecia, sometimes with pink tones in the hymenium. Dried apothecia of O. nannfeldtii have, on the contrary,
darker colours than O. formicarum. Otidea pseudoleporina and
O. formicarum share apothecial shape, but O. pseudoleporina
can be distinguished by the ochre-orange colour, and comparatively narrower spores (Qm = 1.7–1.9).
The original description of O. formicarum was based on several
collections made on anthills in Finland (Harmaja 1976), and
more recent material has been cited from anthills (Harmaja
2009a, b), mainly associated with Picea. εost of the material
198
examined by us was also from anthills, but at least one of our
Swedish finds (KH.11.104) was from Pinus needle litter, which
shows the habitat of O. formicarum partly overlaps with the
habitat of O. subformicarum.
14. Otidea nannfeldtii Harmaja, Karstenia 15: 31. 1976 — Fig.
5a, d, 18
Holotype. Finland, Ahvenanmaa, δemland, Nåtö, spruce forest near Övergård, 17 Sept. 1972, C.-A. Haeggström (H6002902) !
= Otidea angusta Harmaja, Karstenia 48: 35. 2009.
Holotype. Finland, Varsinais-Suomi, δohja, Jalassaari, Ahtiala, E of Heimo
house, mixed somewhat moist rich woods with Picea, Betula, Corylus etc.
on somewhat calcareous soils, 23 Aug. 1965, H. Harmaja (H6010804) !
Persoonia – Volume 35, 2015
Misapplied names
– Otidea papillata sensu Van Vooren et al., Bull. εycol. Bot. Dauphiné-Savoie
188: 52. 2008.
Apothecia gregarious to caespitose, 8–35 mm high, 5–15 mm
wide, initially long, narrowly ear-shaped, sometimes expanding
and becoming broadly ear-shaped, split, stipitate or sessile.
Hymenium ochre (4A5), orangish ochre (5B4, 5C4) or pale
brown (6D6, 6D7), sometimes with pink tones or entirely pinkish (6A4), when dried orange-ochre (5A5) to reddish brown
(6C6). Receptacle surface brown orange (5B4, 5C4) to pale
brown (6D6–7), slightly hygrophanous, in drying ochre (4A5),
when dried reddish brown (7D7, 7E7), finely warty, smooth at
the base. Warts conical, gregarious, concolorous, sometimes
distinctly darker than the background, brown. Stipe 4 – 8 ×
Fig. 17 Otidea formicarum (KH.11.104)*. a. Apothecia; b. spores; c. paraphyses; d. ectal excipulum; e. resinous exudates of the ectal excipulum; f. basal
mycelium. — Scale bars = 10 µm; * = all fresh material.
I. Olariaga et al.: A monograph of Otidea
2–3 mm. Basal tomentum and mycelium abundant, white to
light yellow (5A2) or ochre (5A3). Spores ellipsoid, sometimes
slightly inequilateral, with two large guttules, smooth, hyaline,
(9 –)9.5 –10.5(–11.5) × 5.5 – 6.5(–7) µm (δm = 9.8 –10.9 µm,
Wm = 5.5–6.6 µm, Qm = 1.6–1.9; n = 10). Paraphyses curved
to hooked, of the same width or slightly broader at apices,
2.5–5 µm wide, without notches, rarely with up to two slightly
swollen areas or forked at apices, when fresh containing small,
refractive, light yellow guttules; when dried hyaline to light yellow. Asci 137–190 × 8–10 µm. Apothecial section 650–900 µm
thick. Subhymenium c. 80–100 µm thick, visible as a yellowish
brown zone, of cylindrical to swollen cells, densely arranged.
Medullary excipulum of textura intricata, 400–600 µm thick,
differentiated into two parts: a) textura angularis underneath
the subhymenium, 100–150 µm thick, cells 13–32 × 12–21
199
µm; b) textura intricata, 400–450 µm thick, hyphae 4–9 µm
wide, thin-walled to slightly thick-walled, hyaline to very pale
yellow, sometimes with pale yellow resinous exudates at septa.
Ectal excipulum of textura angularis, sometimes of a textura
prismatica, 80–120 µm, cells thin-walled, hyaline to light brown,
16–40 × 11–22 µm. Outer part of conical to broadly conical
warts, 45–85 µm high, formed by short, fasciculate, hyphoid
hairs, of 2–3(–4) subglobose to elongated cells, constricted
at septa, 5–9 µm wide, sometimes with a gelatinous sheath.
Resinous exudates abundant, yellow brown, dissolving into
amber drops in εδZ, reddish brown in KOH. Basal mycelium
of 3–5.5 µm wide, septate, hyaline to very light yellow hyphae,
unchanged in KOH, with very small, regularly arranged, spheroid, yellow resinous exudates, dissolving in εδZ, partially and
more slowly in KOH.
Fig. 18 Otidea nannfeldtii. a. Apothecia; b. spores*; c. paraphyses*; d. ectal excipulum*; e. hyphoid hairs with gelatinous sheath*; f. amber drops on the
outermost ectal excipulum cells in εelzer’s reagent † (a–e: KH.10.302; f: S-F249387). — Scale bars = 10 µm; * = fresh material; † = dried material.
200
Specimens examined. Finland, Varsinais-Suomi, δohja, Jalassaari, Ahtiala, Alho, very close to the Ahtiala Nature Reserve, below a spruce tree in
mixed forest on fairly calcareous soil, 29 Sept. 1978, H. Harmaja (S-F249387,
ex H6017194 as O. lohjaënsis nom. prov.). – FranCE, Hautes Alpes, ProvenceAlpes-Côte-d’Azur, δa Bâtie-εontsaléon, δes Chariots du Buech, au sol dans
la litière d’aiguilles de pins (P. sylvestris), 26 Oct. 2008, N. Van Vooren, NV
2008.10.01 (dupl. S). – italy, Abruzzo, Pietracamela (TE), Prati di Tivo, on
soil in mixed forest, mainly with Larix but also Picea and Pinus, 2 Oct. 2009,
B. De Ruvo (S-F257096); Calabria, Celico (CS), Contrada Colamauci, under
Pinus and Picea abies, 16 Sept. 2009, C. Lavorato, Cδ 091116-17 (dupl. S);
Calabria, εorano Calabro (CS), Campotenese, under Pinus sylvestris, 7 Dec.
2009, C. Lavorato, Cδ 091207-01 (dupl. S). – SwEdEn, Gotland, Ollajvs Nature
Reserve, close to δjugarn, under Picea and Pinus on calcareous ground,
31 Sept. 2010, K. Hansen, K. Gillen & I. Olariaga, KH.10.302 (S); ibid., under
Picea and Pinus on rich calcareous soil, 27 Sept. 2011, H. Tuovila & S. Huhtinen, KH.11.115 (S); δappland, Jokkmokk, ‘Nornaskogen’ by Ållojaur, under
Picea among mosses, K. Olofsson, 1 Sept. 2011, KH.11.45 (S); δappland,
εessaure, 3 miles NW Vuollerim, herb rich Picea forest, 1 Sept. 2005,
M. Karström, εK0536 (S); δappland, Västra Tjetnekbäcken, 9 km E of
Jokkmokk, under Picea abies, K. Olofsson, 1 Sept. 2011, KH.11.112 (S);
Uppland, Stockholm, Enebyberg, Rinkebyskogen, on soil under Picea and
deciduous trees in side of footpath, 29 Sept. 2008, J. Santos, JS.08.103
(S). – USA, Oregon, Douglas Co., Thielsen Creek, under conifers, 13 Oct.
2010, R. Helliwell, rh101310 (OSC).
Otidea cf. nannfeldtii — dEnmark, δøvenholm Skov, δangsø, 25 km W of
Grenaa, 30 Sept. 1968, H. Folkmar (C-F-48295), (C-F-48296), (C-F-48297).
Notes — Otidea nannfeldtii is characterised by ochre to light
brown, narrowly ear-shaped apothecia, small spores and resinous exudates on the ectal excipulum turning reddish brown in
KOH. It resembles other species with ear-shaped apothecia and
small spores, such as O. formicarum, O. papillata, O. pseudoleporina and O. tuomikoskii. The most similar species is O. formicarum (see under O. formicarum). Otidea tuomikoskii is separated from O. nannfeldtii by higher and more densely placed
warts on the apothecial outer surface, along with the yellow
reaction of the excipulum in KOH. Otidea pseudoleporina has
a brighter ochre-orange hymenium, and the resinous exudates
of the outer excipulum only partly convert into reddish grey
heterogeneous drops in KOH.
Otidea nannfeldtii, as treated here, shows some phylogenetic
structure (Fig. 1; but see Hansen & Olariaga 2015). Recently
O. angusta was described as distinct from O. nannfeldtii,
based on a few subtle characters, i.e. taller and slightly thicker
fleshed apothecia, with very faintly brownish basal mycelium,
paraphyses with shorter and thinner apical cells, and maybe
smaller spores (Harmaja 2009a). At that time only a single
collection of each species was known. Based on studies of
additional collections, we did not find correlations between
any morphological characters and the subgroups supported
by our multiple molecular phylogenies. Therefore, we propose
O. angusta be placed in synonymy with O. nannfeldtii. Supporting this, our study of the holotype of O. angusta revealed larger
spores (9.5–11 × 5–5.8 µm) than cited in the protologue (8–9.8
× 4.5–5.2 µm), and broader paraphyses (2–3 µm) with longer
terminal cells (up to 67 µm). We have also observed both tall,
narrow and rounded apothecia of O. nannfeldtii in the same
spots, and we consider the apothecial shape to vary during
development. Otidea lohjaënsis, proposed by Harmaja (2009a)
as a provisional name, is also suggested to be conspecific
with O. nannfeldtii. A collection identified by Harmaja as
O. lohjaënsis (S-F249387, ex-H6017194) showed excipular
resinous exudates dissolving into amber drops in εδZ (Fig.
18f), contrary to the key character given for O. lohjaënsis (i.e.
not responding to εδZ). The variation in the ITS region within
O. nannfeldtii as recognised here is high, but displays no length
variation (alignment 790 bp long). The ITS sequences of the
Finnish holotypes of O. angusta and O. nannfeldtii show 26 bp
differences. At the same time, however, the ITS sequence of
the Swedish KH.10.302 shows 33 and 32 bp differences from
the O. nannfeldtii and O. angusta holotypes, respectively. The
Persoonia – Volume 35, 2015
North American rh101310 shows 14 and 20 bp differences from
the two holotypes, O. nannfeldtii and O. angusta, respectively.
The collections showing no δSU sequence variation (Fig. 1),
show also no, or only 1–2 bp differences in the ITS region. No
variation was found between the holotype of O. angusta and
the Finnish S-F249387 (identified as the provisional O. lohjaënsis by Harmaja).
Otidea nannfeldtii was previously known only from South West
Finland (Harmaja 1976, 2009a). Here we report it from other
areas in Europe, and for the first time from North America.
15. Otidea subformicarum Olariaga, Van Vooren, ε. Carbone
& K. Hansen, sp. nov. — εycoBank εB809252; ITS barcode GenBank: Kε010054; Fig. 19, 21
Etymology. Referring to its similarity to O. formicarum.
Holotype. Spain, Huesca, Bielsa, Ermita de Nª Señora de Pineta, 1270 m,
42.638039, 0.180532 (decimal format), under Pinus sylvestris and Abies alba
on calcareous ground, 13 Oct. 2012, J.C. Campos & J. Herranz (S-F242696).
Apothecia gregarious to caespitose, 13–32 mm high, 10–20 mm
wide, broadly ear-shaped, soon becoming deeply cup-shaped,
split, margin sometimes lobate, sessile or stipitate. Hymenium
orange brown (6C6, 6C8) to dark reddish brown (6D7, 6D8),
slightly hygrophanous, when dried orange brown (7A8). Receptacle surface orange brown (6B7, 6C7) to reddish brown
(6D8), hygrophanous, in drying orange ochre (6A6, 6A7), when
dried ochre brown (5B6), brown (6C8) or reddish brown (7D8),
finely furfuraceous, smooth at the base. Warts present near
the margin, flat and rounded, concolorous, brown. Stipe 3–4
× 2–3 mm. Basal tomentum and mycelium pale yellow (4A3).
Spores ellipsoid, slightly inequilateral, with two large guttules,
rarely with one to several smaller granules, smooth, hyaline,
10.5–12 × 6–6.5 µm (δm = 11.1–11.7 µm, Wm = 6.1–6.7 µm,
Qm = 1.7–1.9; n = 4). Paraphyses broadly hooked, sometimes
curved, sometimes enlarged at the apices to 2.5–3 µm wide,
without notches, sometimes with a brown matter covering the
apices, when fresh containing small, pale yellow guttules; when
dried heterogeneous, pale yellow. Asci 184 – 237 × 11–11.5
µm. Apothecial section 800 –1200 µm thick. Subhymenium
c. 100–150 µm thick, of dense textura intricata, with tendency to
textura angularis towards the medullary excipulum, visible as a
darker orange brown zone, cells 2.5–6(–15) µm wide, with scattered brown resinous exudates. Medullary excipulum 300–700
µm thick, of textura intricata, hyphae thin-walled to slightly
thick-walled, 3–12 µm wide, very pale yellow to pale brownish
yellow, sometimes with brown resinous exudates at septa. Ectal
excipulum of textura angularis, (85–)100 –120 µm, cells thinwalled, very pale brown, (9–)17– 22(–35) × (6–)11.5–23 µm.
Surface with narrowly to broadly conical warts, 45–65 µm high,
formed by short, fasciculate, hyphoid hairs, of 2–3 cylindrical to
ovoid cells, 7–10 µm wide, not or slightly constricted at septa.
Non-warted areas with single hyphoid hairs, of 2–3 cylindrical
to ovoid cells, (5–)7–12 µm wide, not or slightly constricted at
septa, sometimes with a gelatinous sheath. Resinous exudates
abundant, pale yellowish brown, dissolving into amber drops
in εδZ, turning slightly darker and partly dissolving in KOH.
Basal mycelium of 2–5.5 µm wide, hyaline to very light yellow
hyphae, unchanged in KOH, with refractive, pale yellow drops
on the surface, dissolving in εδZ.
Specimens examined. italy, Cosenza, Calabria, Colamauci, Celico, under
Pseudotsuga menziesii, C. Lavorato, 28 Sept. 2005, Cδ 050928-30 (dupl.
S-F256978). – Spain, Canary Islands, La Palma, under Pinus canariensis,
26 Nov. 2008, J. Fernández Vicente, P. Iglesias, F. Hidalgo, J.R. Undagoitia,
S. Lequerica & R. Martínez (S-F256979); δa Rioja, Clavijo, under Pinus
sylvestris, on the ground, C.M. Pérez del Amo & R. Gil, 3 Jan. 2009, private
herb. CεP 1179, Rε 1095 (dupl. S-F256980); εadrid, Bustarviejo-Canencia,
Puerto de Canencia, 3 Oct. 1979, E. Álvarez (AH44526).
I. Olariaga et al.: A monograph of Otidea
201
Fig. 19 Otidea subformicarum apothecia. a. S-F242696, holotype; b. S-F256980; c. S-F256979; d. Cδ 050928-30. — Photos: b. C. Pérez del Amo; c. J.
Fernández Vicente; d. C. δavorato.
Other specimens examined. Otidea aff. subformicarum — mExiCo, Salazar, Parque Nacional εiguel Hidalgo, Abies forest, 23 Sept. 2007, M. Hernández (FH301035); Veracruz, Cofre de Perote, Camino de los Conejos,
Los Lescados, montane forest of Pinus teocote, P. montezumae, Arbutus
jalapensis, 18 Sept. 2007, M.E. Smith (FH301036).
Notes — Otidea subformicarum is closely related to O. formicarum based on both morphological and molecular characters.
Both have broadly ear- to cup-shaped apothecia and small
spores, and are associated with conifers. Diagnostic features
of O. subformicarum are the orange-brown to reddish brown
apothecial colours, and especially the long and narrow spores
compared to related species. Otidea formicarum has shorter
and comparatively more rounded spores (Fig. 20). All O. subformicarum apothecia were collected under Pinus, Pseudotsuga
O. formicarum
O. nannfeldtii
O. pseudoleporina
O. subformicarum
O. aff. subformicarum
O. sp. ‘b’
6.9
Wm (µm)
6.7
6.5
6.3
6.1
5.9
9.7
10.2
10.7
11.2
11.7
Lm (µm)
Fig. 20 εean spore length and width in collections of species in the O. formicarum clade and O. pseudoleporina, based on 20 spores from each
collection.
menziesii or Abies. The ecology of O. formicarum partly overlaps (see further under that species). Otidea subformicarum
appears to have a southern European distribution, whereas
O. formicarum is known only from Fennoscandia and the Alps.
Otidea subformicarum forms a strongly supported clade in
the ITS-δSU phylogeny (Fig. 2). The four ITS sequences of
O. subformicarum, from Spain and Italy, are nearly identical
(only S-F256979 differs in 1 bp), except for a small variation
in the minisatellites. The ITS nucleotide diversity is 0.71 % in
the four minisatellites unique to O. subformicarum. The five
ITS sequences of O. formicarum, from Finland, Norway and
Sweden, are likewise almost identical (only JS.08.63 differs
in 2 bp). The ITS sequences of O. subformicarum and O. formicarum show many nucleotide differences (40 / 747 bp). Thus
the interspecific ITS nucleotide diversity is much higher (5.35 %)
than the intraspecific diversity (in O. subformicarum 0.07 %
without the minisatellites and in O. formicarum 0.10 %).
Two collections from εexico, FH301035 and FH301036 (here
referred to as O. aff. subformicarum), belong to the O. formicarum clade based on our molecular phylogenetic analyses
(Fig. 1, 2). Although analyses of the combined ITS-δSU dataset
did not resolve the relationships among these three lineages
(Fig. 2) and the δSU phylogeny place FH301035 as a strongly
supported sister taxon to two collections of O. subformicarum,
our three- and four-gene phylogenies (Hansen & Olariaga 2015)
suggest the two European species share a most recent common ancestor, and the εexican collections diverged earlier. The
spore size and shape of the εexican collections conform to
those of O. subformicarum (Fig. 20), but the apothecial colours
differ in these collections. Fresh material has not been available
to us, but the photo of fresh apothecia of FH301035 shows eggyellow apothecia, clearly distinct from O. subformicarum. Photos
of apothecia of FH301036 have a dark purplish brown colour,
Persoonia – Volume 35, 2015
202
which has not been observed in O. subformicarum. Based on
these colour differences and molecular characters, we suggest
that those collections belong to an additional undescribed species, possibly endemic to North America. Further collections are
needed to get insights into the variation and delimitation of this
taxon for its formal description.
Outside the O. formicarum clade, a few species have similar
spores to O. subformicarum. These differ by a combination
of other features: Otidea tuomikoskii by narrowly ear-shaped
apothecia, a receptacle surface with high warts (55–177 µm
high), an ectal excipulum that turns yellow in KOH and typically
orange ochre basal tomentum; and O. nannfeldtii by apothecia
often having yellow tones, sometimes pink stains, and above
all by the shorter spores (δm = 10–10.7 µm).
Otidea unicisa clade
Apothecia with ochre yellow tones. Basal tomentum with ochre
tones. Spores ellipsoid, with warts ± ridges or spinose. Resinous
exudates on the outermost ectal excipulum cells dissolve and
exude bright yellow pigment in KOH.
Species — Otidea kaushalii, O. unicisa, O. yunnanensis.
16. Otidea kaushalii (J. εoravec) K. Hansen & Olariaga, comb.
nov. — εycoBank εB810994; Fig. 22
Basionym. Sowerbyella kaushalii J. εoravec, εycol. Helv. 2: 94. 1986.
≡ Aleurina kaushalii (J. εoravec) W.Y. Zhuang & Korf, εycotaxon 29:
312. 1987.
≡ Otideopsis kaushalii (J. εoravec) J. εoravec, εycol. Helv. 3: 138.
1988.
Fig. 21 Otidea subformicarum (S-F242696, holotype). a. Spores*; b. paraphyses and asci*; c. ectal excipulum in water showing reddish brown resinous exudates†; d. ectal excipulum in εelzer’s reagent showing amber drops †; e. basal mycelium in water †; f. basal mycelium in εelzer’s reagent†. — Scale bars = 10
µm; * = fresh material; † = dried material.
I. Olariaga et al.: A monograph of Otidea
Holotype. india, West Bengal, Darjeeling, Batasi, on soil and decayed
wood in angiosperm forest, alt. 7600 f, 6 Sept. 1979, R. Kaushal, PAN 18169.
Isotypes herb. J. εoravec (CUP 61814, C-F-60847 !).
Apothecia 16–65 mm high, 12–62 mm wide, broadly ear-shaped,
split, stipitate. Hymenium dirty grey to very faintly incarnate,
greyish yellow, orange-ochre (5A6) when dried. Receptacle surface dark orange brown (6E7, 6E8) when dried, densely warty.
Warts conical, acute or blunt, densely gregarious, darker than
the background, dark reddish brown. Stipe 3 – 32 mm long,
3–10 mm wide. Basal tomentum and mycelium ochre (5A4).
Spores ellipsoid to slightly subfusoid, inequilateral, with one or
two large and/or a few small guttules, with thin, often curved,
spines, up to 1(–1.5) µm high, denser at the poles, sometimes
203
united in short ridges, hyaline, 14–17 × 7–9 µm (δm = 14.9–
15.7 µm, Wm = 7.7–7.9 µm, Qm = 1.9–2; n = 2). Paraphyses
curved to hooked, sometimes enlarged at apices, 2.5–5 µm
wide, when dried containing small, yellowish refractive granules. Apothecial section 700–800 µm thick. Asci 188–213 ×
11–13 µm. Medullary excipulum of textura intricata, hyphae
thick-walled, hyaline to very pale yellow. Ectal excipulum of
textura globulosa-angularis, 70–90 µm thick, of 3–4 cell layers, cells thin-walled, pale yellowish brown, 12–46 × 13–33
µm. Surface with conical warts, densely placed, 40–140 µm
high, 50–137 µm wide, formed by globose to elongated cells,
8–11.5 µm broad. Resinous exudates abundant, reddish brown,
amorphous, and/or many yellowish brown, crystal-like, oblate
Fig. 22 Otidea kaushalii. a. Apothecia; b. spores in water †; c. spores in Cotton Blue†; d. paraphyses in KOH †; e. ectal excipulum and warts with abundant
crystal-like, resinous exudates in water†; f. close-up of crystal-like, oblate spheroid exudates on outermost ectal excipulum cells in water † (a–d, insertion on f:
T. δæssøe 6236; e, f: PAN 18169, isotype). — Scale bars = 10 µm; † = dried material. — Photos: a. T. δæssøe.
204
spheroid, 6–8 × 7.5–11 µm, striate bodies, with a constricted
centre, not dissolving in εδZ, dissolving and exuding bright
yellow pigment in KOH. Basal mycelium of 4.5–6 µm wide,
thin-walled to slightly thick-walled, very pale yellow hyphae,
unchanged in KOH, with regularly arranged, spheroid, yellow
resinous exudates, dissolving in εδZ, unchanged in KOH.
Specimens examined. malaySia, Sabah Kinabalu, on rotten wood, 2 εar.
1999, T. Læssøe 6236 (C, dupl. BORH).
Notes — Otidea kaushalii is closely related to O. yunnanensis based on our analyses of the δSU (Fig. 1) and morphology.
Already εoravec (1988) stated this, when he combined O. kaushalii in Otideopsis. The distinctly spiny spore ornamentation in
these two species appears identical; see SEε photographs of
the spores of the holotype of O. kaushalii in εoravec (1986)
Persoonia – Volume 35, 2015
and of O. yunnanensis in δiu & Zhuang (2006). Both species
have a receptacle surface with densely placed, high, dark brown
warts and a lighter coloured hymenium, i.e. greyish yellow,
cream to yellow. Otidea kaushalii is distinguished by the smaller
spores, and possibly by the ectal excipulum of large, yellowish
brown globose to angular cells (Fig. 22e). We report here for
the first time a so far unique type of crystal-like, oblate spheroid, striate exudates on the outermost ectal excipulum cells
of O. kaushalii (Fig. 22e, f) that might be confined to O. kaushalii within Otidea. These are abundant in both the isotype and
the εalaysian material. A third collection of O. kaushalii has
been reported by Zhuang & Korf (1987) from Xizang, China. The
εalaysian and Chinese collections have smaller apothecia with
shorter or almost no stipe (apothecia up to 19 × 18 mm; stipe
3 × 3 mm in the εalaysian material, Fig. 22a) as compared to
Fig. 23 Otidea unicisa. a. Apothecium; b. apothecia†; c. spores in water †; d. spores in Cotton Blue †; e. paraphyses in water †; f. ectal excipulum in water†
(a: JK12082101; b: H7003343; c–f: KH.06.06). — Scale bars = 10 µm; † = dried material. — Photos: a. J. Karakehian.
I. Olariaga et al.: A monograph of Otidea
the holotype (apothecia up to 65 × 62 mm; stipe 32 × 10 mm).
The apothecia of O. yunnanensis have also been described
with varying stipe length (6 –25 × 4–6 mm).
17. Otidea unicisa (Peck) Harmaja, Karstenia 26: 44. 1986. —
Fig. 23
Basionym. Peziza unicisa Peck, Rep. (Annual) New York State εus.
Nat. Hist. 26: 81. 1874.
≡ Sowerbyella unicisa (Peck) J. εoravec, Czech εycol. 47: 266. 1994.
Holotype. USA, New York, δewis County, Croghan, Felt House, ground
in woods, Sept. (NYSf 3283).
Misapplied names
– Otidea grandis sensu Kanouse, εycologia 41: 672. 1949; δiu & Zhuang,
Fung. Diversity 23: 188. 2006.
Apothecia gregarious to caespitose, 12–25 mm high, 12–35 mm
wide, initially broadly ear-shaped, then becoming cup-shaped,
split, stipitate or sessile. Hymenium ochraceous yellow, sometimes with pink tinges, yellowish ochre (5A5, 5B5), orange-ochre
(5A6) when dried. Receptacle surface ochraceous yellow, when
dried dark orange brown (6E7, 6F7), ochre-brown (5A4) towards
the base, warty, occasionally with shallow ribs at the base.
Warts rounded, gregarious, concolorous or slightly darker than
the background, reddish brown. Stipe 4 – 8 × 2–4 mm. Basal
tomentum and mycelium abundant, yellowish ochre (5A4).
Spores ellipsoid to slightly fusoid, inequilateral, with two large
guttules, often with a few smaller guttules, with small, low warts,
often irregular ridges, denser at the poles, hyaline, (13–)14–
15.5(–16.5) × 6.5–8.5 µm (δm = 14.6–15.2 µm, Wm = 7.1–8 µm,
Qm = 1.8–2.1; n = 3). Paraphyses curved to hooked, sometimes
enlarged at apices, 2.5–4 µm wide, sometimes with 1–2 low
notches, when dried containing refractive, pale yellow granules.
Asci 181–197 × 8–11 µm. Apothecial section 750–900 µm thick.
Subhymenium c. 70–120 µm thick, of cylindrical cells, densely
arranged. Medullary excipulum of textura intricata, 500–700 µm
thick, hyphae thin-walled, 3.5–9 µm wide, hyaline to very pale
yellow, without resinous exudates at septa. Ectal excipulum of
textura angularis, 70–95 µm thick, cells thin-walled, hyaline to
light yellow, 11–30 × 6.5–18 µm. Surface with conical warts,
35–85 µm high, formed by fasciculate, short, hyphoid hairs,
of globose to elongated cells, constricted at septa, 6–11 µm
wide. Resinous exudates abundant, dark yellowish to reddish
brown, dissolving in part and converting into reddish particles
in εδZ, turning brighter yellow in KOH. Basal mycelium of
4.5–6 µm wide, thin-walled to slightly thick-walled, hyaline to
very pale yellow hyphae, unchanged in KOH, with regularly
arranged, spheroid, yellow resinous exudates, dissolving in
εδZ, unchanged in KOH.
Specimens examined. USA, εassachusetts, Carlisle, Great Farm, on
woody debris, 8 July 2006, L. Millman, KH.06.06 (FH); εassachusetts, Carlisle, Towle Conservation δand, 15 July 2006, L. Millman (FH301030); εassachusetts, Purgatory Charm, 9 July 2003, Z. Wang, ZW Geo 65-Clark (S);
εichigan, Cut R., εackinac Co., in frondose woods, 10 Aug. 1949, A.H. Smith
33020 (UPS F-629380); εichigan, N of Hessel, εackinac Co., in frondose
woods, 15 Aug. 1949, H. Imshaug 3458 (UPS F-629377); εichigan, δuce
Co., Tahquamenon Falls State Park, 21 Aug. 1951, A.H. Smith 39070 (UPS
F-629382); εichigan, W of Detour on ε134, Chippewa Co., in beech-maple
woods, 14 Aug. 1949, H. Imshaug 3348 (UPS F-629379); New Hampshire,
Shelburne (UPS F-629427); North Carolina, εacon Co., near summit of
Standing Indian εountain, on duff and buried wood of Betula, 1 Aug. 1969,
H.H. Burdsall 2605 (dupl. H7003343); West Virginia, on light loam and leaf
mold, 15 July 1896, L.W. Nuttall, Ellis 868 (UPS F-630023); West Virginia,
εonongahela National Forest, Dolly Sods Wilderness, Wildlife Trail (TR 560),
on soil among leaf litter under Fagus grandifolia, Acer pennsylvanicum, and
Betula sp., 21 Aug. 2012, J. Karakehian, JK12082101 (FH, dupl. S).
Notes — Otidea unicisa is easily recognised by the spore ornamentation of low, delicate warts and short ridges. The dried
specimens are typically bicoloured, with dark brown outside
and orange-ochre hymenium.
205
The spore ornamentation of O. unicisa was for a long time overlooked (Harmaja 1986). The ornamentation is clearly visible at
1 000×, especially at the poles where it is more prominent. We
agree with Harmaja (2009a) that what Kanouse (1949) described under the name O. grandis is O. unicisa. The bicoloured
apothecia with brown outside and orange hymenium, along
with the ornamented spores, and spore sizes in the range of
O. unicisa, support this view. δiu & Zhuang (2006) also gave a
collection with the typical spore ornament of O. unicisa under
the name O. grandis (for SEε of the ornament see their f. 8).
Both studies mention the Boudier plate 328 (n°. 134, 1905; as
O. grandis), which shows bicoloured apothecia. Otidea unicisa
has not been recorded outside Eastern North America, and
in our opinion the Boudier plate 328 shows typical O. bufonia
apothecia. Corroborating this, two Boudier specimens labelled
O. grandis (UPS F-629342 and likely the collection used for
plate 328: PC0093644), studied by us, are in fact O. bufonia.
For further comments on O. grandis see Excluded, dubious
and imperfectly known taxa.
18. Otidea yunnanensis (B. δiu & J.Z. Cao) W.Y. Zhuang &
C.Y. δiu, Fung. Diversity 23: 188. 2006
Basionym. Otideopsis yunnanensis B. δiu & J.Z. Cao, Shanxi Univ. J.,
Nat. Sci. Ed. 4: 70. 1987.
Holotype. China, Yunnan, Dulong River, δang Tuan, on ground in forest,
30 Aug. 1982, D.C. Zhang (HKAS 12150).
Otidea yunnanensis was described when the monotypic genus
Otideopsis was erected. It was separated from Otidea because
of the ornamented spores and paraphyses with fused apical
portion (δiu & Cao 1987). Upon re-examination of the type
material, δiu & Zhuang (2006) discovered that the paraphyses
had free apices. They also concluded that the spores were with
two large and several small guttules, and not multi-guttulate as
originally described by δiu & Cao (1987). Our δSU phylogeny
confirms that O. yunnanensis is deeply nested within Otidea
(Fig. 1). The species is easily recognised within Otidea by the
large spores with fine, curved spines. Different spore sizes have
however, been reported from the holotype: 18–20 × 8–10 µm
(including ornament, δiu & Cao 1987) and 15–21 × 9.7–10.5
µm (excluding ornament, εoravec 1988), and including one
additional collection: 16.5 – 20 × 7.6 –10 µm (δiu & Zhuang
2006). The very wide spore measurements given by εoravec
overlap somewhat with the spore length of the closely related
O. kaushalii (for further comparisons see under O. kaushalii).
Otidea bufonia-onotica clade
Apothecia dark brown or ochre yellow. Basal tomentum with
brown or yellow tones, especially when dried. Spores fusoid or
ellipsoid, smooth. Resinous exudates on the ectal excipulum
converting into reddish particles or melting into amber drops.
Species — Otidea brevispora, O. bufonia, O. fusconigra ad
interim, O. mirabilis, O. onotica, O. purpurea, O. smithii.
19. Otidea brevispora (W.Y. Zhuang) Olariaga & K. Hansen,
comb. & stat. nov. — εycoBank εB808974
Basionym. Otidea onotica var. brevispora W.Y. Zhuang, εycotaxon 94:
368. 2006 ‘2005’.
Holotype. China, Yunnan, Baoshan, 24 July 2003, Z.L. Yang (HKAS
43003) !
Otidea onotica var. brevispora was distinguished from O. onotica based on shorter spores (Zhuang 2006). Our study of the
O. brevispora holotype confirmed the smaller spores (9.5–10.5
× 5.5–6 µm; δm = 9.9 µm, Wm = 5.6 µm, Qm = 1.75). Otherwise
O. brevispora shares all macro- and microscopic features with
206
O. onotica, including the yellow reaction of the basal mycelium
in KOH. Our εδ analyses, with an δSU GenBank sequence of
the Chinese holotype, show O. onotica var. brevispora is a sister
taxon to a clade of O. onotica specimens (Fig. 1). Based on this
and the smaller spores we consider it to be a distinct species.
20. Otidea bufonia (Pers.) Boud., Hist. Classif. Discomyc.
Europe: 52. 1907. — Fig. 4b, c, 5b, 24
Basionym. Peziza bufonia Pers., εycol. Eur. 1: 225. 1822: Fr., Syst.
εycol. 2: 54. 1822.
≡ Geopyxis bufonia (Pers.) Sacc., Syll. Fung. 8: 73. 1889.
Lectotype designated here: FranCE, in sylvula Vincennes, Aug. 1816 (δ
0116690 / 911.81.97, Persoon herbarium) !; εycoBank εBT178089.
= Peziza umbrina Pers., Observ. εycol. 2: 77. 1799.
≡ Scodellina umbrina (Pers.) Gray, Nat. Arr. Brit. Pl. 1: 668. 1821.
Persoonia – Volume 35, 2015
≡ Peziza cochleata var. umbrina (Pers.) Fr., Syst. εycol. 2: 50. 1822:
Fr. loc. cit (‘a umbrina’).
≡ Otidea umbrina (Pers.) Bres., Fungi Trident. Ser. 2, fasc. 11–13: 68.
1898.
Lectotype designated here: Sowerby, Col. Fig. Engl. Fung. 1: t. 5. 1797
(as Peziza cochleata); εycoBank εBT200088.
= Peziza pseudobadia Cooke, εycographia part 4: 176. 1877.
≡ Aleuria pseudobadia (Cooke) Gillet, Champ. France Discomycetes:
38. 1879 (‘pseudo-badia’).
≡ Geopyxis pseudobadia (Cooke) Sacc., Syll. Fung. 8: 69. 1889 (‘pseudobadia’).
Holotype. FranCE, εérignac, sur un mur, 1814 (K(ε) 195314, ex herb.
Cooke).
= Otidea pedunculata Velen., εonogr. Discomyc. Bohemiae 1: 354.
1934.
Lectotype designated here: CzECh rEpubliC, Hrusice, Aug. 1924, J. Velenovský (PRε 147622) !; εycoBank εBT200086.
Fig. 24 Otidea bufonia*. a, b. Apothecia; c. spores; d. paraphyses; e. ectal excipulum; f. basal mycelium (a: KH.09.172; b: JS.08.55; c–f: KH.09.171) — Scale
bars = 10 µm; * = all fresh material. — Photos: b. J. Santos.
I. Olariaga et al.: A monograph of Otidea
Misapplied names
– Otidea grandis sensu Boudier, Icon. εycol. livr. 6: n°. 134, pl. 328. 1905
(preliminary text with ‘circulaires’).
– Peziza cochleata sensu Sowerby, Col. Fig. Engl. Fung.1: t. 5. 1797.
Apothecia gregarious, rarely caespitose, 15–45 mm high, 17–
32 mm wide, initially ear-shaped, then soon expanding and becoming deeply cup-shaped, split, stipitate or sessile. Hymenium
initially orange brown (6C6) sometimes olivaceous brown (4D6),
then dark orange brown (6F6, 7E8), when dried greyish brown
(5E3, 5E4), slightly purple. Receptacle surface dark brown
(6E4–6E7), sometimes pale rusty brown (6D8, 6E8) or purplish
brown (6E3) or with olivaceous tones, slightly hygrophanous,
in drying slightly paler, when dried dark orange brown (6E5,
6F4), warty, seldom slightly wrinkled at the base. Warts conical to flattened, gregarious, dark brown, sometimes distinctly
darker than the background. Stipe 5–14 × 7–10 mm, often hollow and felty inside. Smell weak; taste mild. Basal tomentum
and mycelium abundant, brownish white to light brown (6B3).
Spores narrowly fusoid, rarely ovoid, inequilateral, with two
large guttules, very rarely with a third small guttule, smooth,
hyaline to pale yellowish, (12–)13–16.5(–18) × 6–7.5(–8) µm
(δm = 12.4–16.1 µm, Wm = 6.3–7.3 µm, Qm = 1.9–2.5; n = 8).
Paraphyses hooked, a few curved, of the same width or slightly
enlarged at apices, 3.5–5(–7) µm wide, without notches or
rarely with a notch on the underside, when fresh containing
small, refractive, light brownish yellow guttules; when dried
light brownish yellow. Asci 143 –172 × 10 –12 µm. Apothecial section 1000 –1300 µm thick. Subhymenium c. 70 –100
µm thick, visible as a darker zone, composed of cylindrical
to swollen cells, densely arranged, with scattered brownish
resinous exudates at septa. Medullary excipulum of textura
intricata, 500–700 µm thick, hyphae thick-walled, 3.5–9 µm
wide, hyaline to light brown, some covered with strikingly dark
brown, striate resinous exudates. Big crystal-like aggregates
sometimes present among the hyphae, dissolving in KOH,
visible in εδZ. Ectal excipulum of textura angularis, 80–100
µm thick, cells thin-walled, brownish, 16–31 × 11–25(–30) µm.
Surface with broadly conical warts, 40–80 µm high, formed by
short, fasiculate hyphoid hairs, of 3–4 subglobose to elongated
cells, constricted at septa, 6–11 µm wide, sometimes with a
gelatinous sheath. Resinous exudates abundant, dark brown,
partly dissolving and converting into small reddish particles in
εδZ, partly dissolving in KOH. Basal mycelium of 3.5–4.5 µm
wide, hyaline to brown hyphae, with oily, light brown drops on
the surface, sometimes crystalloid and rod-shaped.
Specimens examined. CzECh rEpubliC, Prague-East district, infra Klokočná, in carpineto, 5 Oct. 1931, J. Velenovský (PRε 150074, syntype of
O. pedunculata). – dEnmark, Fyn, Juelsberg Skov, N of Nyborg, along roadside in Fagus forest, 28 Sept. 1986, D. Boertman (C-F-47790); E Jylland,
δøvenholm Skovene, Eldrup Skov, in grass along road, 23 Sept. 1979, E. Andersen (C-F-86688); E Jylland, Vejle Nørreskov, at roadside on rich soil
under Populus tremula, Alnus, Corylus, Betula, Fraxinus, Fagus, etc., 13
Sept. 1993, J. Vesterholt (C-F-20453); NE Jylland, Rubjerg Knude, under
Abies, 18 Sept. 1989, C. Lange & J. Vesterholt (C-F-25955); NW Jylland,
Nystrup Plantage near Thisted, 1 Sept. 1972, K. Toft (C-F-48018); W Jylland, Bordrup Plantage, in Picea and Pinus plantation along forest road on
sandy ground, 17 Sept. 2011, M. Sasa (C-F-94240); δolland, Favrsted Skov,
among mosses, on calcareous soil, by forest-road under Fagus, 4 Oct. 2007,
K. Hansen & B. Kullman, KH.07.37 (S); Sjælland, Feldskoven near Sorø,
14 Aug. 1974, M. Lange (C-F-47995); N Sjælland, Gribskov, under Fagus
in the forest, 19 Sept. 1971, H. Dissing (C-F-48019); ibid., 8 Oct. 1973, P.M.
Pedersen (C-F-48015); Sjælland, Horserød Hegn, 6 km W of Helsingør, along
roadside, 17 Sept. 1964, H. Dissing (C-F-48000). – Finland, Varsinais-suomi,
δohja, Virkkala, Pähkinäniemi, herb-rich deciduous forest on calcareous, mull
soil, with Corylus avellana, Betula pendula, Populus tremula, Picea abies
and Pinus sylvestris, 28 Aug. 1997, J. Vauras 12503F (TUR-A, dupl. S). –
FranCE, Fontainebleau, on the ground under Quercus and Abies, Oct. 1876,
E. Boudier (PC0093644, as O. grandis); Forêt de Châtellerault (δ 0111782,
Herb. Persoon, as Peziza umbrina); δandes, Contis, under Pinus pinaster
on sandy soil, 6 Dec. 2009, J.L. Teres, KH.09.249 (S); δoire-Atlantique,
207
Bourgneuf-en-Retz, 5 Nov. 2009, G. Moyne, NV 2009.11.01 (S); Picardie,
Compiègne, Oct. 1892, E. Boudier (UPS F-629342, as O. grandis); Saône-etδoire, Robin, on the ground under Corylus and Quercus, J.-P. Dechaume, NV
2008.09.12 (dupl. S); Vendée, Jard-sur-εer, in dune forest with dominance
of Pinus pinaster and Quercus ilex, G. Ouvrard, NV 2006.11.07 (S); Vosges,
Rambervillers, forêt de Saint-Hélène, under Alnus in a peatbog, 5 Oct. 2006,
M. Hurtu, NV 2006.10.12 (S). – italy, Caorle (VE), Brussa, Vallevecchia,
under Pinus pinea close to the sea, 4 Nov. 2001, E. Campo (S-F257089);
Ceva (CN), 9 km from the city on the road to Viola, on the side of a track
under Quercus pubescens, Castanea sativa and Corylus avellanea, 16 Sept.
2005, M. Carbone (S-F257088); Deiva εarina (SP), under Quercus ilex,
5 Nov. 2008, M. Carbone (S-F257090); Piemonte, Vignole Borbera (Aδ), Fraz.
Variano superiore, in soil under Quercus pubescens and Castanea sativa, 19
Oct. 2010, M. Carbone (S-F257087). – Spain, Barcelona, entre Sant Feliu de
Codines y Castellterçol, forest with Quercus ilex and Pinus sylvestris, on calcareous ground, 14 Oct. 1976, C. Montoliu (AH44527); Huesca, Javierregay,
under Quercus rotundifolia and Q. humilis humus, 5 Dec. 2009, F. Prieto &
A. González s.n. (S); Navarre, Arbizu, in mixed forest, 14 Nov. 2009, J. Martín
(ARAN-Fungi A5048005); Navarre, Etxauri mendatea, under Quercus ilex,
2 Dec. 2006, J.M. Lekuona (ARAN-Fungi A5053019); Navarre, δete, under
Quercus faginea and Q. rotundifolia on calcareous ground, 19 Dec. 2009,
J.L. Teres & P.M. Pasaban, KH.09.248 (S); Valladolid, Tudela de Duero,
Puente Hierro, under Quercus ilex, J. Santos (S-F22110). – SwEdEn, Gotland,
near Visby, Värnhem, under Fagus and Quercus robur on rich ground, with
Hepatica nobilis, 22 Sept. 2009, E. Bohus-Jensen, K. Hansen & I. Olariaga,
KH.09.189 (S); Skåne, Degeberga, εörkavad, broadleaf forest, 27 Sept.
2001, S.-Å. Hanson, SÅH 2001-253 (C); Skåne, Helsinborg, Gyhult, broadleaf
forest dominated by Fagus and Quercus, on the ground, 13 Oct. 1994, S.-Å.
Hanson, SÅH 16457 (C); Uppland, Stockholm, Enebyberg, Rinkebyskogen,
on rich soil in deciduous forest, under Corylus, but also Tilia, Quercus and
Betula, 1 Sept. 2008, J. Santos, JS.08.55 (S); ibid., 17 Sept. 2008, JS.08.79
(S); Uppland, Uppsala, Hågadalen-Nåsten Nature Reserve, Predikstolen,
under Quercus robur, Picea abies, Corylus and Salix, on rich bare ground,
17 Sept. 2009, K. Hansen & I. Olariaga, KH.09.171 (S); ibid., KH.09.172 (S);
Uppland, Vaksala, near Törnby, Ekbacken, grassy ground with bare patches
of soil, close to a Quercus robur tree, 13 Sept. 2010, K. Gillen & I. Olariaga,
KH.10.199 (S); Öland, Algustrum, Hönstorp, 500 m SO of the village, grazed
mixed deciduous forest, 19 Sept. 1993, T. Knutsson, TK93-209 (S); Öland,
δånglöt, Åstad, c. 75 m from Björkerumsvägen, c. 550 S of the T-crossing
in Åstad, under Corylus, 6 Sept. 1998, T. Knutsson, TK98-208 (S). – USA,
εichigan, Cheboybogan Co., Colonial Pt, Burt δ., in beech-maple woods,
9 Aug. 1951, A.H. Smith 37560 (UPS F-629510); εichigan, Emmet Co.,
Pellston, in beech-maple woods, 11 Aug. 1951, A.H. Smith 37654 (UPS
F-629511); εinnesota, Itasca State Park, on soil in mesic deciduous woods of
red oak, bur oak, birch, hazelnut, Ostrya, 28 July 2010, R.A. Healy, RH1218
(εIN 933332); εinnesota, δake Alexander SNA, on soil in mixed woods with
canopy or red oak, Ostrya, white birch, 16 Aug. 2011, R.A. Healy, RH1393
(εIN 9333323). – without loCality (δ 0111780, Herb. Persoon, as Peziza
umbrina); (δ 0111781, Herb. Persoon, as Peziza umbrina).
Notes — Otidea bufonia is macroscopically characterised
by the cup-shaped apothecia, with a dark brown outside and a
brown basal tomentum, especially in dried specimens. εicroscopically, the narrowly fusoid spores and the presence of hyphae with striate resinous exudates in the medullary excipulum
are important diagnostic characters. Otidea mirabilis is very
similar to O. bufonia (see under O. mirabilis).
The hymenium colour shows a high variability in O. bufonia (Fig.
24a, b); we have observed orange brown, olivaceous or dark
brown tones within a single collection. Also the colour of the
receptacle surface varies and can be with rusty to purplish
brown or olivaceous tones. If purplish tones are present it may
be confused with O. mirabilis. The pigmented, striate, crystallike exudates on the hyphae of the medullary excipulum are
considered a unique feature for O. bufonia (Korf & Zhuang
1991, Harmaja 2009a, Parslow & Spooner 2013), and have
been used to separate O. bufonia from O. mirabilis (Carbone
et al. 2010, Van Vooren 2010). Our results strengthen this view.
We have observed striate exudates in all the material examined
(Fig. 4b, c). The size and abundance of the striate exudates
are variable, and often restricted to the outermost part of the
medullary excipulum (Fig. 4b). In O. mirabilis we have only
observed biflabellate, crystal-like exudates (Fig. 4d).
208
An exceptional O. bufonia specimen with deviant spore characters was discovered (NV 2009.11.01), in which the spores
were ovoid and considerably shorter than in the rest of the
material. This collection is nested within the O. bufonia clade,
along with typical collections.
Nomenclatural notes — A specimen from the Persoon herbarium (911.81.97, δ0116690) labelled as O. bufonia by Persoon can be considered type material (Harmaja 2009a, b), being
collected prior to the publication of O. bufonia. We designate it
here as the lectotype to stabilise the name. It includes a single
dark brown apothecium, with uniseriate, smooth, subfusoid
spores, 14 –16 × 5.8 –7 µm, curved paraphyses, numerous
dark brown, striate exudates on the hyphae of the medullary
Persoonia – Volume 35, 2015
excipulum, and outermost ectal excipular cells with dark brown
resinous exudates.
Otidea umbrina has been recognised as a separate species
(εedardi 1995, Dissing 2000), based mainly on different hymenium colour. The original description of O. umbrina and the
Sowerby plate to which Persoon (1799) referred, agree with
our concept of O. bufonia. Confirming Harmaja (2009a), three
of the four collections of O. umbrina studied by us and kept in
Persoon´s herbarium in δ belong to O. bufonia. Thus we regard
O. umbrina as a synonym of O. bufonia in accordance with other
authors (Dennis 1978). The name O. bufonia is sanctioned at
specific rank and thus has priority over O. umbrina, which is
sanctioned only at variety rank. The name O. grandis has been
used for O. bufonia collections with bicoloured hymenium, i.e.
Fig. 25 Otidea mirabilis*. a, b. Apothecia; c. spores; d. paraphyses; e. ectal excipulum; f. ectal excipulum wart in εelzer’s reagent, showing reddish reaction
of resinous exudates (a: KH.09.188; b: KH.10.294; c–f: KH.10.308). — Scale bars = 10 µm; * = all fresh material.
I. Olariaga et al.: A monograph of Otidea
orange-brown or olivaceous hymenium in contrast to a dark
brown outside (Boudier 1905, Van Vooren 2010). For the interpretation of O. grandis see Excluded, dubious and imperfectly
known taxa. We synonymise O. pedunculata with O. bufonia
based on the study of two syntypes, and lectotypify it with
PRε 147622. The lectotype shows dark brown, split apothecia, fusoid spores (13.5–15.5 × 5.5–6.5 µm, from 8 spores)
and importantly striate exudates on some of the hyphae of the
medullary excipulum. Harmaja (2009b) studied the holotype of
P. pseudobadia and found it is a later synonym of O. bufonia.
Only one collection of P. pseudobadia could be located in Kew
(K(ε) 195314); it is from εérignac as given in the protologue,
but from a wall, whereas Cooke wrote ‘on the ground’. In any
case, one of the apothecia corresponds well to the illustration
in the protologue, and we consider this the holotype (and the
material studied by Harmaja, although no annotation by him
could be found; pers. comm. B. Aguirre-Hudson).
21. Otidea mirabilis Bolognini & Jamoni in Jamoni, Funghi e
Ambiente 85 –86: 56. 2001. — Fig. 4d, 25
Holotype. italy, Piemonte, Alagna valsesia (VC), Val d’Otro, on soil under
Picea and Larix, alt. c. 1500 m, 3 Sept. 1999, D. Bolognini, GεFN 1951.
Isotype (S-F256929) !
Misapplied names
– Otidea leporina sensu Zhuang pro parte, εycotaxon 96: 367. 2005.
Apothecia gregarious or caespitose, 18–62 mm high, 9–52 mm
wide, initially ear-shaped, short or elongated, soon expanding
and becoming deeply cup-shaped, split, stipitate or sessile.
Hymenium initially reddish ochre (5B7, 6B7) or sometimes olivaceous brown (5F5, 5F6), then dark reddish ochraceous brown
(5D8) to dark purple brown (6F4–6F7), when dried ochraceous
brown (5D6) to purplish brown (6E5, 6E6). Receptacle surface
dark purple brown with bluish lilaceous shades (7F2, 7F3),
fading away with time to ochraceous dark-brown (5D7), then
dark purple brown (6F4–6F7), sometimes with greyish ochre
(4A3, 4B3) patches in unexposed parts, hygrophanous, in drying light purple brown (5D2, 6D2) to ochre brown (5B4), when
dried purple brown (6D4, 6F3, 6F4), warty, often wrinkled at the
base. Warts broadly conical to hemispherical, obtuse, gregarious, dark brown, sometimes distinctly darker than the background. Stipe 7–17 × 3–7 mm. Smell weak, vaguely resin-like;
taste mild, slightly bitter in the end. Basal tomentum and mycelium abundant, whitish to light brown (5A3). Spores narrowly
fusoid, inequilateral, with two large guttules, very rarely with
additional 1–3 small guttules, smooth, hyaline to pale yellowish, (13–)13.5–16(–17) × 6–7(–7.5) µm (δm = 14.1–15.4 µm,
Wm = 6.3–6.9 µm, Qm = 2.1–2.3; n = 9). Paraphyses curved to
hooked, of the same width to slightly enlarged at apices, 2.5–5
µm wide, without or occasionally with a few low notches towards
the apex, when fresh containing small, refractive, light brownish yellow guttules; when dried refractive, hyaline guttules.
Asci 178–204 × 9–11 µm. Apothecial section 1000 –1300 µm
thick. Subhymenium c. 100–150 µm thick, visible as a darker
zone, composed of irregularly, densely arranged, globose cells,
with scattered brownish resinous exudates at septa. Medullary excipulum of textura intricata, 600–900 µm thick, hyphae
4–11(–15) µm wide, sometimes slightly swollen, thin to thickwalled, hyaline to light brown, seldom with brown resinous
exudates at septa or biflabellate exudates, sometimes forming
cross-like aggregates. Ectal excipulum of textura angularis,
80 –100 µm thick, cells thin-walled to slightly thick-walled,
brownish, 8–30 × 8–18 µm. Surface with broadly conical warts,
35–55 µm high, formed by short, fasciculate, hyphoid hairs,
of 2 – 4 subglobose to elongated cells, constricted at septa,
6.5–10(–17) µm wide, sometimes with a gelatinous sheath.
Resinous exudates abundant, dark brown, partially dissolving
209
and turning reddish in εδZ. Basal mycelium of 3–6 µm wide,
light yellow to light brown hyphae, seldom with oily brownish
drops on the surface, with abundant small dark brown resinous
exudates, rounded to irregular.
Specimens examined. dEnmark, Bornholm, Rø Plantage, on needle layer
under Picea, 30 Sept. 2001, C. Lange, KH.01.09 (C). – Finland, Koillismaa,
Kuusamo, Oulanka National Park, Ampumavaara, south of the main road
to δiikanen, 14 Aug. 2010, M. Carbone (S-F257083). – FranCE, Isère, δansen-Vercors, 1300 m, on the ground, on litter, under Larix, 20 Sept. 2008,
J. Cavet, NV 2008.09.14 (dupl. S). – india, Uttarakhand, Kalika, Nainital, on
loose humus soil, 22 Sept. 1973, S. Chandes (UPS F-630072). – italy, Friuli
Venezia Giulia, Tarvisio (UD), Val Saisera-Valbruna, on calcareous soil under
Picea abies, Larix decidua and Fagus sylvatica, 27 Sept. 2010, G. Dose
(S-F257092). – norway, Rana, St. Alteren, 7 km W of εo i Rana, 4 Sept.
1972, H. Dissing (C-F-87187). – SwEdEn, Gotland, Ala, Näsmyr, under Picea
and Pinus sylvestris on calcareous ground, 30 Sept. 2010, E. Bohus-Jensen,
K. Hansen, K. Gillen & I. Olariaga, KH.10.294 (S); Gotland, δjugarn, forest
close to Kaupungs Fridhem, under Picea abies on calcareous ground, under
a cliff, 27 Sept. 2010, K. Hansen, K. Gillen & I. Olariaga, KH.10.279 (S);
Gotland, δojsta hed, Russpark, grazed forest with Pinus sylvestris, on calcareous ground, 2 Oct. 2010, K. Hansen, K. Gillen & I. Olariaga, KH.10.308 (S);
Gotland, near Visby, Rävhagen, under Pinus sylvestris, with small Quercus
robur, Helianthemum sp., Melampyrum pratense, Gallium verum, on sandy
and acidified soil, 22 Sept. 2009, E. Bohus-Jensen, K. Hansen & I. Olariaga,
KH.09.188 (S); Gotland, Tofta, Smågårde naturskog Nature Reserve, Tofta
strand, under Pinus sylvestris on calcareous ground, 28 Sept. 2010, K. Hansen,
K. Gillen & I. Olariaga, KH.10.285 (S); ibid., KH.10.288 (S).
Otidea cf. mirabilis — SwEdEn, Öland, Gräsgård, Solberga, Stora Alvaret,
1300 m S of the village, 17 Oct. 1993, T. Knutsson, TK93-276 (S).
Notes — Otidea mirabilis is characterised by dark brown
apothecia, purple to lilaceous-bluish shades in the receptacle
surface of especially young apothecia, fusoid spores, and when
present, biflabellate, crystal-like exudates in the medullary excipulum. Otidea mirabilis and O. bufonia are strongly supported
as sister species in our multigene phylogeny (Hansen & Olariaga 2015). Thorough morphological comparisons show that
O. bufonia differs from O. mirabilis in lacking lilaceous-bluish
shades, and having brown striate exudates on some hyphae of
the medullary excipulum. Otidea smithii is distinguished from
O. mirabilis by the shorter spores with lower Qm (1.9–2.0).
Otidea mirabilis was described with emphasis on the lilaceous
tones of the outside, in contrast to the paler ochre to olivaceous
hymenium (Jamoni 2001). Part of our material shows this colour pattern, but collections with dark brown hymenium have
also been observed. Some authors have noted the absence
of pigmented resinous exudates in the medullary excipulum of
O. mirabilis (Carbone et al. 2010, Van Vooren 2010). We have,
however, observed brown crystal-like exudates in KH.10.308
and the holotype. They are flabellate and forming cross-like aggregates (Fig. 4d), clearly different from those present in O. bufonia (Fig. 4b, c).
Coniferous forests on calcareous ground are the typical habitat
of O. mirabilis. Two Chinese collections assigned to O. leporina
by Zhuang (2006) and sequenced by δiu & Zhuang (2006) are
resolved in the O. mirabilis clade in our δSU tree (Fig. 1). Based
on morphology we report it also from India (UPS F-630072).
22. Otidea onotica (Pers.) Fuckel, Jahrb. Nassauischen Vereins Naturk. 23–24: 330. 1870. — Fig. 26
Basionym. Peziza onotica Pers., Syn. εeth. Fung.: 637. 1801: Fr., Syst.
εycol. 2: 48. 1822.
≡ Scodellina onotica (Pers.) Gray, Nat. Arr. Brit. Pl. 1: 668. 1821.
≡ Peziza leporina var. onotica (Pers.) P. Karst., Bidrag Kannedom Finlands Natur Folk 19: 41. 1871 (‘P. leporina * P. onotica’).
Lectotype designated here: Sowerby, Col. Fig. Engl. Fung. 1: t. 79. 1797
(as Peziza leporina). Epitype designated here: SwEdEn, Gotland, Ollajvs
Nature Reserve, close to δjugarn, calcareous ground in mossy Picea forest,
27 Sept. 2010, K. Hansen, K. Gillen & I. Olariaga, KH.10.284 (S); εycoBank
εBT178083.
= Peziza rosea Schumach., Enum. Pl. 2: 416. 1803.
210
Misapplied names
– Otidea concinna sensu Boudier, Icon. εycol. livr. 26: n°. 552, pl. 325. 1909
(preliminary text with ‘circulaires’).
– Peziza leporina sensu Sowerby, Col. Fig. Engl. Fung.1: t. 79. 1797.
Apothecia gregarious to caespitose, 25–100 mm high, 14–80
mm wide, initially long and narrowly ear-shaped, then soon
expanding, and becoming deeply cup-shaped, split, stipitate
or sessile. Hymenium initially light yellow (4A3 – 4A5), then
ochraceous yellow (4A6, 4A7, 5A6, 5A7, 5B6, 5B7), in some
parts light orange (5A3, 5A4), often with pink tones or entirely
pinkish (6A4), sometimes with red dots, when dried orangeochre (5A5, 5A6) to reddish ochre (6B7, 6B8). Receptacle
surface ochraceous yellow (4A6, 4A7), slightly hygrophanous,
in drying slightly paler, when bruised sometimes brownish ochre
(5B7) in the margin, when dried brownish ochre with orange
Persoonia – Volume 35, 2015
tinge (5B7, 5B8), slightly warty, sometimes wrinkled at the base
when old. Warts broadly conical, gregarious, concolorous or
sometimes distinctly darker than the background, brown. Stipe
13–27 × 4–14 mm. Smell weak; taste mild. Basal tomentum
and mycelium abundant, whitish to light yellow (4A2, 5A2).
Spores ellipsoid to broadly ellipsoid, inequilateral, with two large
guttules, smooth, hyaline, (11–)12–13.5(–14) × (5.5–)6–7 µm
(δm = 12.1–13.3 µm, Wm = 6.2–6.8 µm, Qm = 1.8–2; n = 10).
Paraphyses curved to hooked, of the same width or slightly
broader at apices, 2.5–4.5 µm wide, without notches or rarely
with a slightly swollen area on the underside, when fresh containing small, refractive, light yellow guttules; when dried hyaline
to pale yellow guttules. Asci 138–233 × 9.5–12 µm. Apothecial
section 850 –1400 µm thick. Subhymenium c. 100 –120 µm
thick, of dense textura intricata, visible as a darker zone, cells
Fig. 26 Otidea onotica*. a, b. Apothecia; c. spores; d. paraphyses; e. ectal excipulum; f. basal mycelium (a: KH.10.284, epitype; b: KH.09.132; c – f:
KH.09.165). — Scale bars = 10 µm; * = all fresh material.
I. Olariaga et al.: A monograph of Otidea
cylindrical to swollen, densely arranged. Medullary excipulum
of textura intricata, 400–600 µm thick, hyphae 5–7(–12) µm
wide, thick-walled, hyaline. Ectal excipulum of textura angularis,
80–110 µm thick, cells 13–55 × 11–28 µm, thin-walled, light
yellow. Surface with conical warts, 85–105 µm high, formed by
short, fasciculate hyphoid hairs, of 2–3 subglobose to elongated
cells, constricted at septa, 11–14 µm wide. Resinous exudates
abundant, yellow, dissolving into amber drops in εδZ. Basal
mycelium of 3.5–6 µm wide, hyaline to very light yellow hyphae,
turning yellow in KOH, with very small, regularly arranged,
spheroid, resinous exudates on the surface, dissolving in εδZ,
partially and more slowly in KOH.
Specimens examined. CzECh rEpubliC, South Bohemia, Netolice, Sept.
1922, Hampl (PRε 148341). – dEnmark, Amager, Kongelunden, soil along
forest path, 21 July 1998, K. Hansen, T. Læssøe & C. Lange, KH.98.107 (C);
εøn, Fanefjord Skov, under conifers on calcareous influenced soil, 30 Sept.
2008, H. Knudsen (C-F-89691); Sjælland, Geelskov, 12 km N of Copenhagen,
under Fagus, 18 Sept. 1963, L. Hansen & A. Kjøller (C-F-47985). – FranCE,
Landes, Contis phare, under Pinus pinaster on sandy soil, 9 Sept. 2009,
J.L. Teres (ARAN-Fungi A8200131C). – italy, Piemonte, Vinadio (CN), San
Bernolfo, Picea abies and Abies alba forest, with presence of Fagus sylvatica,
29 Sept. 2008, M. Carbone (εCVE 23277). – norway, Nord-Trøndelag,
Gjøråsvika, δeksvik, under Picea on rich ground, among mosses, 3 Sept.
2009, K. Hansen & I. Olariaga, KH.09.132 (S); under Corylus and Picea on
rich ground, KH.09.136 (S). – Spain, Gipuzkoa, Aia, Amezketalardiko bidegurutzea, under Abies alba, 3 Oct. 2009, J.L. Teres (ARAN-Fungi A3033701A);
εadrid, Sierra de Guadarrama, deciduous forest, among Quercus pyrenaica
leaves, 4 Oct. 1981, G. Moreno (AH2528); Navarre, Arbizu, mixed forest, 14
Nov. 2009, J. Martín (ARAN-Fungi A5048003); Navarre, Orokieta, δoiandi, in
Picea abies plantation, 10 Oct. 2009, Aranzadi ZE (ARAN-Fungi A50411742). – SwEdEn, Skåne, Vittskövle, Segesholm, Herremöllan, broadleaf forest,
27 Sept. 2001, S.-Å. Hanson, SÅH 2001-266 (C); Uppland, Stockholm,
Norra Järvafältet, Hansta Nature Reserve, on rich ground under Picea abies
with Corylus avellana, 15 Sept. 2009, K. Hansen & I. Olariaga, KH.09.164
(S); ibid., KH.09.165 (S), KH.09.166 (S); Uppland, Uppsala, forest-covered
hill NNE of Naturicum, coniferous forest with Quercus and Corylus, 4 Oct.
2011, R. Sundin, KH.11.108 (S); Uppland, Uppsala, Norra δunsen Nature
Reserve, δunsentorpet, on rich organic soil, under dead standing Picea, 28
Aug. 2008, J. Santos, JS.08.48 (S); Uppland, Uppsala, Vänge, Fiby urskog
Nature Reserve, old Picea forest, in the middle of a path, 17 Sept. 2009,
K. Hansen & I. Olariaga, KH.09.175 (S). – USA, εinnesota, Cedar Creek
ESR, on sandy soil in oak savannah, canopy of pin oak, hazelnut, 18 July
2011, R.A. Healy, RH1121 (εIN 933307); εinnesota, δake Alexander SNA,
on soil, mixed woods, canopy of red oak, paper birch, Ostrya, 29 July 2010,
R.A. Healy, RH1222 (εIN 933311); εinnesota, Wild River State Park εany,
on moist, sandy soil in mesic deciduous woods with oak, 23 July 2010,
R.A. Healy, RH1199 (εIN 933309); Oregon, Benton Co., Philomath, Wood
Creek Road, 18 Nov. 1996, E.T. Peterson (OSC 56759); Washington, δower
Tahoma Creek, εt Rainier National Park, 27 Aug. 1948, A.H. Smith 30764
(UPS F-629424); Washington, Nisqually R., εt Rainier National Park, 30
Aug. 1948, H.A. Imshaug 2116 (UPS F-629438).
Notes — Otidea onotica is one of the most common and wellknown species of the genus. It is characterised by rather large,
ochraceous yellow apothecia, often with a pinkish tinge, dots
or stains in the hymenium, and spores of unusual size within
Otidea. The presence of pinkish tones varies considerably.
Apothecia ranging from entirely pink to completely devoid
of pinkish tones have been observed in the same locality, in
KH.09.132 and KH.09.136, respectively. These two collections
have identical ITS sequences (GenBank accessions Kε010103
and JN942772). A consistent character, here reported for the
first time, is the yellow reaction of the basal mycelium in KOH.
Otidea unicisa resembles O. onotica macroscopically, but is
distinguished by ornamented spores.
Nomenclatural notes — Carbone (2009) revised the nomenclature of O. onotica. He considered the plate by Sowerby (1797,
as Peziza leporina) the only element seen by Persoon, and
thus, the holotype. However, Persoon did indirectly refer to
two elements by giving the habitat in the protologue as beech
forests (faginetis) and citing Sowerby’s plate, which shows an
oak leaf. Thus, the plate is not the holotype; i.e. the one element
on which the author based the name (Art. 9.1 ICN; εcNeill et
211
al. 2012). The use of the term holotype could be corrected to
lectotype according to Art. 9.9, but this article is not applicable
since Carbone’s holotype indication does not fulfil Art. 7.10 of
the ICN (it does not include the phrase ‘designated here’ or
equivalent) (εcNeill et al. 2012). Carbone (2009) selected in
addition an epitype from Persoon´s herbarium. But as an epitype must refer to the type it interprets and there was no validly
selected type in 2009, the epitypification by Carbone is also
not valid. We therefore typify O. onotica here, by designating
as lectotype the Sowerby plate, supported by a newly collected
epitype with multiple gene sequences and colour photographs.
The selection of an epitype from Sweden is justified by the
name being sanctioned (Fries saw and studied living material
as indicated by ‘v. v.’), and our ITS sequences of O. onotica
(4, 5, 6, 7, 8, 9) from Denmark, Italy, Norway and Sweden
(including the epitype) being identical. The δSU sequences of
O. onotica (4, 5) from Denmark and Sweden differ only 2 bp
from the other collections.
23. Otidea purpurea (ε. Zang) Korf & W.Y. Zhuang, εycotaxon
22: 507. 1985
Basionym. Acetabula purpurea ε. Zang, Acta Bot. Yunnan. 1: 101. 1979.
Holotype. China, Tibet, Zayu, on the ground in forest with Pinus yunnanensis, 1 Sept. 1976, M. Zang 670 (HKAS 5670) !
Notes — Otidea purpurea is probably closely related to
O. mirabilis. It is also characterised by dark brown apothecia
with lilaceous tones, but it clearly differs in the smaller spores
(8.8–10 × 4.5–5.2 µm). Two additional taxa, O. subpurpurea
and O. bicolor, should be compared with O. purpurea. Otidea
subpurpurea likewise has violaceous tones on the outside,
but has larger spores and asci (Zhuang & Yang 2008). Otidea
bicolor has dark brown apothecia and small spores (Zhuang
2010), and is probably also closely related or conspecific with
O. purpurea, based on the spore size. We treat here O. purpurea in the O. bufonia clade, because of the dark brown apothecia
and brown basal tomentum.
24. Otidea smithii Kanouse, Pap. εichigan Acad. Sci., Part 1.
24: 28. 1939 ‘1938’ — Fig. 27
Holotype. USA, California, Crescent City, 18 Nov. 1937, A.H. Smith 8843
(εICH 14408) !
Apothecia gregarious or caespitose, 32–70 mm high, 13–40
mm wide, initially narrowly ear-shaped, short or elongated,
finally expanding, at late stages becoming deeply cup-shaped,
split, normally shortly stipitate. Hymenium purple-brown (6D5)
to dark purple-brown (7E5), then ochraceous brown (5D8),
when dried very pale purplish brown (5A2). Receptacle surface
dark purple-brown (7E5), sometimes with lilaceous shades or
lighter ochre-brown (6C5) patches in unexposed parts, slightly
hygrophanous, in drying dark ochraceous brown (6D7), when
dried reddish brown (6E5, 6E6), warty, sometimes wrinkled
at the base. Warts conical, obtuse or acute, gregarious, dark
brown, sometimes slightly darker than the background. Stipe
12–26 × 5–12 mm. Basal tomentum and mycelium abundant,
whitish to brown (5B2, 5C2). Spores ellipsoid, sometimes fusoid, inequilateral, with two large guttules, smooth, hyaline,
12–14(–14.5) × 6–7.5 µm (δm = 12.5–13.6 µm, Wm = 6.4–7.1
µm, Qm = 1.9–2.0; n = 5). Paraphyses curved to hooked, of the
same width to slightly enlarged at apices, 3–5 µm wide, sometimes with 1–3 notches or with a low notch near the apex, when
dried containing refractive, pale yellow guttules. Asci 175–251 ×
9–11 µm. Apothecial section 800–1300 µm thick. Subhymenium
c. 140–180 µm thick, of dense textura intricata, visible as a
darker zone, hyphae intermixed with a few subglobose cells,
with scattered brownish resinous exudates at septa. Medullary excipulum of textura intricata, 700–950 µm thick, hyphae
212
Persoonia – Volume 35, 2015
Fig. 27 Otidea smithii (ECV3345). a. Apothecia; b. spores in water †; c. paraphyses in water †; d. ectal excipulum in water †. — Scale bars = 10 µm; † = dried
material. — Photos: a. E. Vellinga.
Fig. 28 Otidea borealis (S-F242694, holotype). a, b. Apothecia; c. close-up of apothecia showing a warty external receptacle and dentate margin; d. Young
narrowly cup-shaped apothecium. — Photos: ε. Carbone.
213
I. Olariaga et al.: A monograph of Otidea
4–14(–21) µm wide, sometimes slightly swollen, thin-walled to
slightly thick-walled, light brown, seldom with light brown resinous exudates at septa. Ectal excipulum of textura angularis,
70–100 µm thick, cells thin-walled to slightly thick-walled, pale
brown, 13–30 × 8–15 µm. Surface with conical warts, 34–71
(–102) µm high, formed by short, fasciculate, hyphoid hairs,
of 2 – 5 subglobose to elongated cells, constricted at septa,
6–14 µm wide, sometimes with a gelatinous sheath. Resinous
exudates rather abundant on the outside, dark brown, partly
dissolving and turning reddish in εδZ. Basal mycelium of 3–5
µm wide, light to darker brown hyphae, sometimes with pale
brown drops on the surface, with abundant small, rod-shaped
to irregular, brown resinous exudates.
Specimens examined. Canada, British Columbia, Goldstream Park, Victoria, on decayed wood under mature Douglas fir, 21 Sept. 1968, J. Ginns
1212 (UPS F-629486). – USA, California, Alameda Co., Berkeley, δe Conte
Avenue, in lawn under Betula and Cedrus, 25 Oct. 2005, E. Vellinga,
ECV3345 (S); California, Del Norte Co., California State Park, δake Earl
Wildlife Area, access by Sand Hill Road, with Abies grandis, Picea sitchensis, Polystichum munitum, Rubus spectabilis, Vaccinum ovatum, 15 Dec.
1997, M. Madsen & R. Davis (OSC 56823); Oregon, Benton Co., Philomath,
Woods Creek Road, 15 Nov. 1997, E.T. Peterson (OSC 56811); Washington,
Pierce Co., εt Rainier National Park, δower Tahoma Creek, 30 Oct. 1996,
E.T. Peterson (OSC 56753); ibid., 18 Oct. 1997 (OSC 56799); Washington,
δewis Co., Gifford-Pinchot National Forest, Camp Creek Falls Trail, on litter
with Acer circinatum, A. macrophyllum, Tsuga heterophylla, Berberis nervosa,
Polystichum munitum, Linnaea borealis, Rubus sp., Goodyera oblongifolia,
6 Nov. 1997, E. Hathaway & E. Millian (OSC 56830).
Notes — Otidea smithii is closely related to O. bufonia and
O. mirabilis, as shown by morphological and molecular characters. These three species share dark brown apothecia, brown
basal tomentum and dark brown resinous exudates in the ectal
excipulum. Otidea smithii is distinguished by shorter spores with
a lower Qm value, and typically narrower, ear-shaped apothecia.
The unique, biflabellate or striate exudates in the medullary
excipulum, as seen in O. bufonia and O. mirabilis (Fig. 4c, d),
respectively, are not present in O. smithii. Only pigmented resinous exudates at the hyphal septa in the medullary excipulum
are rarely found in O. smithii (as in Fig. 4a). Otidea smithii is so
far only known from Western North America (Kanouse 1949,
Peterson 1998). Our ITS sequence of O. smithii (JN942771)
is identical to the ITS sequence of the holotype (AF072065).
Otidea concinna clade
Apothecia cup-shaped, split or sometimes entire, often with bright
yellow tones. Spores small, 10–12.5 µm long, smooth. Paraphyses typically straight and claviform at apices. Basal mycelium with scattered reddish or yellow resinous exudates.
Species — Otidea borealis, O. caeruleopruinosa, O. concinna,
O. flavidobrunneola, O. lactea, O. minor, O. oregonensis, O. phlebophora, O. rainierensis, O. sinensis, O. tianshuiensis, O. sp. ‘a’.
brown (5D7) in the margin, hygrophanous, when dried reddish
brown (6D5, 6D6), warty, seldom slightly wrinkled at the base.
εargin sometimes finely dentate and darker. Warts broadly
conical, gregarious, light to dark brown. Stipe well developed,
3–12 × 2–6 mm. Basal tomentum and mycelium white to very
pale yellow (3A2), when dried very pale yellow (3A2). Spores
broadly ellipsoid, symmetrical, with two large guttules, smooth,
hyaline, (10–)10.5–11.5 × 6–7 µm (δm = 10.5 µm, Wm = 6.5
µm, Qm = 1.7; n = 1). Paraphyses straight to curved, very few
hooked, 2–2.5 µm wide, at apices up to 3–4.5(–5) µm wide,
without notches, terminal segment 33–47 µm long, narrowly
claviform to claviform, when dried containing slightly refractive,
hyaline granules. Asci 167–198 × 9–10.5 µm. Apothecial section 1 000 µm thick. Subhymenium c. 100 µm thick, of dense
textura intricata, cells 2–4 µm thick. Medullary excipulum 600
µm thick, of textura intricata, hyphae thin-walled to slightly
thick-walled, 4–7 µm wide, hyaline, without resinous exudates.
Ectal excipulum of textura prismatica, 80 µm thick, cells thinwalled, hyaline, unchanged to very pale yellow in KOH, 10–21
× 8.5–11 µm. Surface with broadly conical warts, 63–90 µm
high, formed by fasciculate, short, hyphoid hairs, of 3–6 subglobose to ovoid cells, constricted at septa, 6–12 µm wide.
Non-warted parts with single, 2–5-celled hyphoid hairs, with
cylindrical to claviform upper cell, 45–73 × 8–10 µm. Resinous
exudates golden brown, partly dissolving in εδZ (amber drops
not observed), bright yellow and partly dissolving in KOH. Basal
mycelium of 3–4.5 µm wide, very pale yellow hyphae, bright
yellow in KOH, with spheroid to rod-shaped, reddish yellow
resinous exudates, dissolving in εδZ, partially dissolving and
turning bright yellow in KOH.
Other specimens examined. Otidea sp. ‘a’ — SwEdEn, Lappland, Arvidsjaurs, δillån Allmänningreservat, 8 km SV Järvträsk, herb-rich Picea forest
on calcareous ground, 12 Sept. 2009, M. Karström, εK0942 (S); ibid., 10
Sept. 2010, εK1081 (S).
Notes — Within the O. concinna clade, O. borealis is distinguished by broadly ear-shaped apothecia, with a pale yellowish
ochre hymenium and a darker ochre yellow outer surface with
small brown warts. εicroscopically, the spores are proportionally broader than in closely related species, and the mycelium
at the base of the apothecia turns bright yellow in KOH (tested
in dried specimen only; Fig. 30f).
At least four species of the O. concinna clade have bright
yellow outer apothecial colours, similar to O. borealis. Otidea
concinna appears morphologically most similar, but besides the
citrine-yellow outer receptacle, it differs in the narrower spores
(Fig. 29) with higher Qm value. Otidea phlebophora and O. minor
are distinguished by the often entire apothecia with ribs or
anastomosing veins at the base and distinctly narrower spores.
Otidea oregonensis differs from O. borealis in typically having
O. borealis
O. concinna
7
25. Otidea borealis ε. Carbone, Olariaga, K. Hansen & Van
Vooren, sp. nov. — εycoBank εB809250; ITS barcode
GenBank Kε010023; Fig. 28, 30
Holotype. Finland, Koillismaa, Kuusamo, Juuma, Jäkälävuoma, western
part of Jäkälävuoma gorges, many apothecia in moist soil among mosses
at brookside, under Picea abies, 16 Aug. 2010, M. Carbone (S-F242694).
Isotype (TUR-A 198578).
O. kauffmanii type
6.6
O. minor
6.4
Wm (µm)
Etymology. Referring to a supposedly boreal distribution.
6.8
O. oregonensis
6.2
O. phlebophora
6
O. rainierensis type
5.8
O. sp. ‘a’
5.6
5.4
Apothecia gregarious to caespitose, 12–22 mm high, 8–15 mm
wide, initially narrowly ear-shaped, upper margin subacute to
rounded, then broadly ear-shaped, split, stipitate. Hymenium
pale yellowish ochre (4A2, 5A2), when dried greyish ochre
(4A2, 5A2). Receptacle surface ochraceous yellow (4A5,
5A5), yellowish brown (5B5, 5B6) in some parts, sometimes
5.2
5
10.1
10.6
11.1
11.6
12.1
12.6
Lm (µm)
Fig. 29 εean spore length and width in collections of species in the core of
the O. concinna clade, based on 20 spores from each collection.
214
Persoonia – Volume 35, 2015
Fig. 30 Otidea borealis (S-F242694, holotype)†. a. Spores in water; b. paraphyses and asci in KOH; c. ectal excipulum in water, with resinous exudates on the
outside; d. ectal excipulum turning bright yellow in KOH, resinous exudates partly dissolving; e. basal mycelium in water; f. basal mycelium in KOH. — Scale
bars = 10 µm; † = all dried material.
obconical apothecia, sometimes with strongly rugulose base,
citrine-yellow colour in the outer receptacle when young, narrower spores (Fig. 29), and the mycelium at the base of the
apothecia does not turn yellow in KOH. The North American
O. rainierensis (= O. kauffmanii) appears to have some yellowish
apothecial pigment (see further under O. rainierensis). Nevertheless, the spores of O. rainierensis are longer than the spores
of O. borealis (Fig. 29). Furthermore, our ITS-δSU phylogenies
(Fig. 3) suggest that these species are distinct from O. borealis.
Two Swedish collections, εK0942 and εK1081, constitute another undescribed species (Otidea sp. ‘a’) nested within the
O. concinna group. It resembles O. borealis in spore shape
(only slightly longer in Otidea sp. ‘a’: 11–12(–13) × 6–7 µm;
Lm = 11.2–11.6 µm, Wm = 6.6–6.8 µm, Qm = 1.6–1.7; n = 2) and
yellow reaction of the basal mycelium in KOH. The two Otidea
sp. ‘a’ collections differ from O. borealis in the brownish ochre
receptacle surface, without yellow tones, and ochre to pinkish
ochre hymenium. Also our analyses of the ITS-δSU regions
support O. sp. ‘a’ and O. borealis as distinct species (Fig. 3).
Since only dried material has been available to us, we postpone
a formal description of O. sp. ‘a’ until fresh and more abundant
material, with good colour photographs, becomes available.
I. Olariaga et al.: A monograph of Otidea
26. Otidea caeruleopruinosa Harmaja, Karstenia 48: 37. 2009
— Fig. 31
Holotype. Finland, Varsinais-Suomi, δohja, Jalassaari, 20 Sept. 1978,
H. Harmaja (H6010805) !
Apothecia gregarious to caespitose, 31– 60 mm high, 17–50
mm wide, initially ear-shaped, upper margin rounded, then cupshaped, split, margin sometimes lobulate, stipitate or sessile.
Hymenium yellowish brown (5C6), when dried yellowish brown
with a faint olivaceous tint (4D6) or orange-ochre (6B6, 6C6).
Receptacle surface upper half concolorous with hymenium or
slightly duller (5C5, 5C6), lower half more whitish (pubescent
pruinose), slightly hygrophanous, when dried concolorous with
hymenium towards the margin, below cream, or pale reddish
brown (6C6), furfuraceous to very finely warty, seldom slightly
ribbed at the base. Warts broadly conical, gregarious, paler
or bluish. Stipe not well developed, rooting, up to 11 × 5 mm.
Basal tomentum and mycelium abundant, when fresh ochraceous white, when dried light ochre (5A2). Spores narrowly
ellipsoid to ellipsoid, often inequilateral, with two large guttules,
smooth, hyaline, (10.5–)11–12.5 × 5.5–6.5 µm (δm = 11.2–11.8
µm, Wm = 5.9–6.1 µm, Qm = 1.9; n = 3). Paraphyses straight
or curved, or broadly hooked, of the same width or broader at
apices, 2–4(–5) µm wide, without notches, terminal segment
26–78 µm long, when fresh containing few, small, weakly refractive, pale guttules; when dried refractive, hyaline granules. Asci
144–200 × 9–11 µm. Apothecial section 700–950 µm thick.
Subhymenium c. 70–80 µm thick, of 2.5–5 µm wide cylindrical
cells, with scattered swollen, 12–18 µm wide cells, densely arranged. Medullary excipulum of textura intricata, 550–700 µm
thick, hyphae thin to thick-walled, 3–9 µm wide, intercalated with
215
swollen cells, 12–18 µm diam, hyaline, without resinous exudates. Ectal excipulum of textura prismatica-angularis, 100–120
µm, cells thin-walled, pale brown, sometimes yellow in KOH,
10–32 × 10–22 µm. Surface with conical to broadly conical
warts, 60–80 µm high, formed by short, fasciculate hyphoid
hairs, of 2 – 4 ovoid to cylindrical cells, 6 –10 µm wide, not
constricted at septa. Non-warted parts with single 2–7-celled
hyphoid hairs, of globose to cylindrical cells, 20–33 × 9–17
µm. Resinous exudates abundant, yellow to reddish brown,
dissolving in εδZ, bright yellow and partly dissolving in KOH
and the outer excipulum turning bright yellow. Basal mycelium
of 3.5–6(–8) µm wide hyphae, with spheroid to rod-shaped,
reddish yellow resinous exudates, dissolving in εδZ, more
slowly and partially dissolving in KOH, turning yellow.
Specimens examined. Spain, Girona, Ripollés, Setcases, under Corylus
avellana, Betula verrucosa and Buxus sempervirens, on calcareous soil, 26
Aug. 2010, M. Tabarés & S. Santamaría, εT 10082601 (dupl. S). – SwEdEn,
Södermanland, Nynäshamn, Herrhamra, on soil under Fagus, in narrow forest area along the road, 18 Sept. 2013, I.-L. Walter, KH.13.48 (S); Uppland,
Älvkarleby, V from the train station, 24 Sept. 1950, G. Fåhraeus & G. Stenlid
(UPS F-146664).
Notes — The main diagnostic characters of O. caeruleopruinosa are cup-shaped, split apothecia, whitish to pale cream hymenium, grey outside and rather small spores. Harmaja (2009a)
described O. caeruleopruinosa with strong emphasis on the
bluish shades of the warts on the apothecial outside, a character that has not been observed in our Swedish, or the Iberian
material (Van Vooren et al. 2011). A third gathering from the
Iberian site does not show bluish tones either (ε. Tabarés, pers.
comm.).
Fig. 31 Otidea caeruleopruinosa. a. Apothecia; b. apothecium showing basal tomentum†; c. spores in water †; d. paraphyses in water †; e. ectal excipulum in
water † (a: εT 11080205; b–e: H6010805, holotype). — Scale bars = 10 µm; † = dried material. — Photos: a. ε. Tabarés; b. J. Kearey.
216
Otidea caeruleopruinosa belongs to the O. concinna clade, and
it shares several characters with the other members of the
clade, such as cup-shaped apothecia, small spores, and an
ectal excipulum of textura prismatica to textura angularis. It is
morphologically most similar to O. flavidobrunneola (see Notes
under that species).
Persoonia – Volume 35, 2015
holm, Naturhistoriska riksmuseet, in front of the Botany building, under a
big Quercus robur tree, by a row of Populus, 20 Sept. 2009, K. Hansen &
I. Olariaga, KH.09.183 (S); εycoBank εBT178084.
= Helvella pyxidata Schaeff., Fung. Bavar. Palat. Nasc. 4: 111. 1774
(‘Elvela’).
≡ Peziza marsupium var. pyxidata (Schaeff.) Pers., Syn. εeth. Fung. 2:
640. 1801 (‘ß pyxidata’).
≡ Scodellina pyxidata (Schaeff.) Gray, Nat. Arr. Brit. Pl. 1: 669. 1821.
27. Otidea concinna (Pers.) Sacc., Syll. Fung. 8: 96. 1889 —
Fig. 32
Misapplied names
– Flavoscypha cantharella sensu Harmaja, Karstenia 14: 107. 1974.
Basionym. Peziza concinna Pers., εycol. Eur. 1: 221. 1822: Fr., Syst.
εycol. 2: 49. 1822.
≡ Helvella scutellata Schaeff., Fung. Bavar. Palat. Nasc. 4: 101. 1774
(‘Elvela’).
Lectotype designated here: Schaeffer, Fung. Bavar. Palat. Nasc. 2: t. 150,
f. 1. 1763 (‘Elvela tertia’). Epitype designated here: SwEdEn, Uppland, Stock-
Apothecia gregarious to caespitose, 17–71 mm high, 12–66 mm
wide, initially broadly ear-shaped, then soon expanding and becoming deeply cup-shaped, split, often broader above, very
rarely entire when young, stipitate or broadly sessile. Hymenium
pale yellowish ochre (3A3, 4A5) to pale greyish ochre (5A2),
Fig. 32 Otidea concinna*. a, b. Apothecia; c. spores; d. paraphyses; e. ectal excipulum; f. basal mycelium (a, c, e, f: KH.09.183, epitype; b: KH.09.217;
d: KH.09.176). — Scale bars = 10 µm; * = all fresh material.
I. Olariaga et al.: A monograph of Otidea
sometimes with rose stains or spots, when dried light yellowish
grey (2A2) to very pale yellowish ochre (3A2, 3A3). Receptacle
surface bright citrine yellow (3A5–3A7), sometimes with a greenish hue (2A6), slightly hygrophanous, in drying slightly paler,
fading to brownish ochre (4B5) in age, when dried yellow (3A7),
furfuraceous, sometimes warty in young apothecia, sometimes
shallowly wrinkled at the base. Warts flat to broadly conical,
leaving a reticulum among them, concolorous. Stipe not well
developed, rooting, 7–8 × 4–5 mm. Smell sweet, fruity, weak;
taste mild. Basal tomentum and mycelium abundant, white to
very pale cream (4A2), sometimes with very pale purplish tones,
when dried white to very pale yellow (4A2). Spores narrowly ellipsoid to ellipsoid, inequilateral, with two large guttules, smooth,
hyaline, (10–)10.5–12 × (5–)5.5–6.5 µm (δm = 10.8–11.2 µm,
Wm = 5.6–6 µm, Qm = 1.8–2; n = 8). Paraphyses straight to
bent, seldom broadly hooked, at apices claviform to abruptly
capitate, 2.5–6(–8.5) µm wide, without notches, terminal segment 19–66 µm long, when fresh content hyaline homogenous
or of small, refractive, hyaline to pale yellow granules; when
dried refractive, hyaline granules. Asci 176 –196 × 10 –11.5
µm. Apothecial section 1200 –2100 µm thick. Subhymenium
c. 60–90 µm thick, composed of cylindrical cells, 2–4 µm wide,
with scattered swollen cells, 9–13 µm wide, densely arranged.
Medullary excipulum 600 –1000(–1500) µm thick, of textura
intricata, hyphae cylindrical to slightly swollen, 3–11(–15) µm
wide, thin to thick-walled, hyaline, without resinous exudates.
Ectal excipulum of textura prismatica-angularis, 80–100 µm
thick, cells thin-walled, hyaline to very pale yellow, sometimes
light reddish in KOH, 12 – 35 × 6 – 22 µm. Surface with low
broadly conical warts, 60–70 µm high, formed by fasciculate,
parallel, short, hyphoid hairs, of 4–6 ovoid cells, constricted
at septa, 5–9.5 µm wide. Non-warted parts with single, 2–6celled, hyphoid hairs, uppermost cell narrowly claviform to
subcapitate, 20–45 × 4.5–7 µm, sometimes with a gelatinous
sheath. Resinous exudates abundant, yellow or reddish, dissolving into amber drops in εδZ, turning bright yellow in KOH.
Basal mycelium of cylindrical to slightly swollen, hyaline to very
pale yellow hyphae, 3–4.5(–8.5) µm wide, yellow in KOH, with
scattered, spheroid to rod-shaped, yellow or reddish resinous
exudates, dissolving in εδZ, partially in KOH.
Specimens examined. dEnmark, NE Sjælland, Jægersborg Dyrehave,
8 Sept. 1979, H. Knudsen (C-F-87186); NW Jylland, Nystrup Plantage, ‘Kridtstien’, on calcareous soil with Abies, 14 Sept. 1985, T. Læssøe (C-F-81617).
– EStonia, Saaremaa, Abruka, ad terram, 16 Sept. 1966, K. Kalamees (UPS
F-629562). – Finland, Varsinais-Suomi, Turku, Ispoinen Katariinanlaakso,
W part of the reserve, in rich, essentially deciduous woods with Corylus, Quercus, Tilia, etc. on bare clayey mull soil, 22 Aug. 1977, H. Harmaja (S-F249360,
ex-H6015773). – FranCE, Saône-et-δoire, δa Grande-Verrière, Senavelle, on
the ground under Pseudotsuga menziesii, 12 Sept. 2008, J.-P. Dechaume, NV
2008.09.13 (dupl. S). – Spain, Huesca, Javierregay, humus of Quercus rotundifolia and Q. humilis, 5 Dec. 2009, F. Prieto & A. González, KH.09.250 (S);
Huesca, Sallent de Gállego, δanuza, 14 Oct. 2006, I. Olariaga (BIO-Fungi
13002); Huesca, Yebra de Basa, close to Sta Orosia chapel, in Fagus sylvatica forest, 11 Oct. 2009, P. Siljeström (ARAN-Fungi A8700091). – SwEdEn,
Gotland, Ala, Stenstugu, near a big Quercus robur tree in meadow, on rich
ground, among grass, 26 Sept. 2009, E. Bohus-Jensen, K. Hansen & I. Olariaga, KH.09.217 (S); Gotland, δojsta, δojsta slot, under Corylus on a chalkrich slope, 19 Sept. 2000, T. Knutsson, TK2000-078 (S); Närke, Havsta, Bruntorpskärret, under conifers, 10 Aug. 2008, B. Wasstorp, JS.08.59 (S); Skåne,
Degeberga, Segesholm, on a steep slope on bare ground, under Fagus and
Ulmus, 23 Sept. 2010, K. Hansen, K. Gillen, I. Olariaga, KH.10.256 (S);
Skåne, Fjälkstad, Balsberget, on rich ground among leaf litter under Fagus,
20 Sept. 2010, K. Hansen, K. Gillen & I. Olariaga, KH.10.212 (S); Skåne,
Kristianstad, N. δingenäset, Storskogen, on ground in broadleaf forest,
18 Sept. 1996, S.-Å. Hanson s.n. (C); Uppland, Uppsala, Carolinaparken,
under Corylus avellana, 7 Sept. 2010, S. Ryman, K. Hansen, K. Gillen &
I. Olariaga, KH.10.180 (S); Uppland, Uppsala, Hågadalen-Nåsten Nature
Reserve, Predikstolen, under Quercus robur, Ulmus glabra, Sorbus aucuparia, with Hepatica nobilis, Geum urbanum, Convallaria majalis, on rich
ground, 19 Sept. 2009, K. Hansen & I. Olariaga, KH.09.176 (S); ibid., 6 Oct.
2010, KH.10.182 (S).
217
Notes — Otidea concinna is characterised by cup-shaped,
split apothecia, citrine yellow receptacle surface, along with paraphyses predominantly straight and claviform at apices. Otidea
minor and O. phlebophora share with O. concinna a yellow
receptacle surface, but they have anastomosing ribs at the
apothecial base and narrower spores (Fig. 29). Otidea oregonensis is distinguished from O. concinna by the often obconical
apothecia, sometimes with blunt ribs at the base.
The bright yellow receptacle surface is one of the main diagnostic characters of O. concinna, but we have observed the yellow
colour can disappear with age, and the outer surface becomes
brownish ochre (Fig. 32b) thus making it difficult to recognise
the species. Otidea rainierensis is a North American taxon that
shows an apothecial shape similar to O. concinna. Nevertheless, bright citrine yellow tones are absent in O. rainierensis,
and our molecular data support it as distinct from O. concinna
(Fig. 1, 3). For further comments see under O. rainierensis.
Nomenclatural notes — No authentic material has been
located in Persoon’s herbarium in δ. We therefore select one
element in Schaeffer’s plate (t. 150, f. 1) as the lectotype, because it most closely resemble Persoon’s description, showing
two or more large, convolute, caespitose, sessile, cup-shaped
apothecia with a citrine (although light) outer surface. Persoon (1822) collected the species in ‘sylvula Vincennes prope
Parisios’ and Schaeffer’s plate is from Bavaria, Germany. Fries
sanctioned the name, but did not see material (‘v. ic.’, seen from
icones). Nevertheless, we select a newly collected epitype from
Sweden (Fig. 32a, c, e, f), associated with Quercus (the habitat
described by Persoon) and backed by sequences of multiple
gene regions (ITS, δSU, PRB1, RPB2 and EF1) of O. concinna
collections from southern Europe (Spain) and from the epitype
being identical (Hansen & Olariaga 2015), acknowledging the
species is widely distributed in Europe.
28. Otidea flavidobrunneola Harmaja, Karstenia 48: 38. 2009
— Fig. 33
Holotype. Finland, Varsinais-Suomi, δohja, Jalassaari, Ahtiala, Alho,
Ahtiala Nature Reserve, rich, predominantly deciduous (Quercus, Corylus
etc.) woods on calcareous soils, 20 Sept. 1978, H. Harmaja (H6010806) !
Apothecia gregarious to caespitose, 15–70 mm high, 8–50 mm
wide, initially narrowly to broadly ear-shaped, then cup-shaped,
split, margin sometimes lobulate, shortly stipitate or sessile.
Hymenium cream or pale yellowish (4A2, 4A3), when dried
orange-ochre (5B6). Receptacle surface pale brownish ochre
(5A3, 5A4), slightly hygrophanous, when dried dark reddish
brown (5E5), furfuraceous to very finely warty, seldom shallowly
wrinkled at the base. Warts flattened, gregarious, concolorous. Stipe not well developed. Basal tomentum and mycelium
abundant, white to pale cream (4A2) when fresh, ochre (5A4)
to orange-ochre (6A4) when dried. Spores narrowly ellipsoid
to ellipsoid, inequilateral, with two large guttules, very rarely
with a third small guttule, smooth, hyaline, 9.5–11 × (4.5–)5–6
µm (δm = 10–10.6 µm, Wm = 5.1–5.5 µm, Qm = 1.9–2; n = 5).
Paraphyses curved, a few straight or broadly hooked, of the
same width or slightly broader at apices, 3–4 µm wide, without
notches or slightly sinuous underside, uppermost cell 40–76 µm
long, when fresh containing yellow guttules; when dried hyaline
guttules or granules. Asci 127–170 × 9 –10 µm. Apothecial
section 900–1900 µm thick. Subhymenium c. 100 µm thick, of
dense textura intricata, visible as a yellowish zone, cells cylindrical to slightly swollen. Medullary excipulum 600–1300 µm
thick, of textura intricata, hyphae 5–13 µm wide, thick-walled,
sometimes with a thinner outer wall, hyaline, without resinous
exudates. Ectal excipulum of textura prismatica, sometimes of
218
Persoonia – Volume 35, 2015
Fig. 33 Otidea flavidobrunneola. a. Apothecia; b. apothecium showing basal tomentum†; c. spores in water †; d. paraphyses in water †; e. ectal excipulum in
water †; f. ectal excipulum in KOH†; g. basal mycelium in water † (a, c: KH.09.153; b, d–g: H6010806, holotype). — Scale bars = 10 µm; † = dried material. —
Photos: b. J. Kearey.
a textura angularis, 100–120 µm, cells thin-walled, yellowish,
distinctly citrine yellow in KOH, 18–38 × 11–13 µm. Surface with
broadly conical warts, 40–70 µm high, formed by fasciculate,
short hyphoid hairs, of 3–4 ovoid to elongated cells, constricted
at septa, 5–9 µm wide, sometimes with a gelatinous sheath.
Non-warted parts with 2–3-celled hyphoid hairs, with claviform
uppermost cell, more rarely cylindrical, 33 – 55 × 8 –13 µm.
Resinous exudates abundant, yellowish brown, partially dissolving and turning slightly reddish in εδZ, partly dissolving,
bright yellow and appearing gelatinous in KOH. Basal mycelium of 4–6 µm wide, hyaline to very pale yellow hyphae, with
scattered swollen septa, with rounded to rod-shaped, yellow
resinous exudates, slowly dissolving in KOH, quickly in εδZ.
Specimens examined. Finland, Uusimaa, Nurmijärvi, parish centre, under
Quercus, 16 Sept. 1987, P. Askola 2220 (H); Uusimaa, Nurmijärvi, parish
centre, clayey soil under Quercus, 11 Aug. 1988, P. Askola 2334 (H); Uusimaa, Nurmijärvi, parish centre, the margin of the park by the vicarage, under
Quercus, 18 Aug. 1988, P. Askola 2360 (H). – norway, Nord-Trøndelag, δeksvik, Gjøråsvika, on rich ground, on slope under Corylus and Picea, 3 Sept.
2009, K. Hansen & I. Olariaga, KH.09.153 (S). – SwEdEn, Uppland, Bondkyrka,
Vårdsätra naturpark, 17 Aug. 1927, H. Svensson (UPS F-146554); Uppland,
Uppsala, in front of the prison, on bare ground under deciduous trees, 14 Sept.
1938, R. Gustafsson (UPS F-146718).
Notes — Otidea flavidobrunneola is macroscopically characterised by cup-shaped, split apothecia, especially with age very
pale yellowish brown outside and cream-white to pale yellow
hymenium. εicroscopically, the small spores and the yellow
I. Olariaga et al.: A monograph of Otidea
ectal excipulum turning brighter yellow in KOH are diagnostic.
The apothecia and basal tomentum become characteristically
darker upon drying: the outside turns dark brown, the hymenium and the basal tomentum orange ochre (Fig. 33b). Otidea
caeruleopruinosa and O. flavidobrunneola are morphologically
very similar, but differ in the spore sizes (δm = 11.2–11.8 µm
vs 10.1–10.6 µm) and colours of the basal tomentum in dried
specimens. According to Harmaja (2009a), O. flavidobrunneola
is reminiscent of O. bufonia, but the latter has a darker brown
basal tomentum and larger, narrowly fusoid spores. We report
here the first finds outside Finland.
29. Otidea lactea J.Z. Cao & δ. Fan in Cao et al., εycologia
82: 735. 1990
Holotype. China, Heilongjiang province, Yichun City, on ground (rotten
wood?) under broadleaf trees, 6 Sept. 1987, J.Z. Cao (HεAS 61359, exεHSU 1803).
Notes — We were not able to get the material of O. lactea
on loan, but it appears to be a distinct species based on the
entire, cup-shaped, cream white apothecia and paraphyses
with broadly clavate to subglobose apices (Cao et al. 1990).
Also, in our δSU phylogeny (including a GenBank sequence of
the holotype, DQ443447 from δiu & Zhuang (2006)), O. lactea
forms a distinct sister lineage (εδ 71 %, PP 99 %) to a clade of
seven collections of O. minor from Europe (εδ 95 %, PP 98 %,
Fig. 1). Otidea minor differs from O. lactea in the (most often)
split apothecia, with a yellow outer surface and often more
narrow, straight to bent, subclaviform to claviform paraphyses
apices. Otidea lactea has the characteristic broad apices of
the paraphyses of the O. concinna clade, which however, in
O. lactea becomes hooked with age (Cao et al. 1990). The holotype was originally deposited in εHSU (Cao et al. 1990), but
later transferred to HεAS (confirmed by Hong-εei δu, HεAS).
30. Otidea minor (Boud.) Olariaga & K. Hansen, comb. & stat.
nov. — εycoBank εB808975; Fig. 34
Basionym. Otidea cantharella var. minor Boud., Icon. εycol. livr. 23: n°.
411. 1909 (preliminary text with ‘circulaires’).
≡ Flavoscypha cantharella var. minor (Boud.) Häffner, Rheinland Pfäl.
Pilzj. 4: 36. 1994.
Lectotype designated here: Boud., Icon. εycol. livr. 23: n°. 411, pl. 326.
1909; εycoBank εBT178086.
Misapplied names
– Otidea cantharella sensu δundell & Nannfeldt, Fungi Exs. Suec. 1–2: 93.
1934.
– Flavoscypha cantharella sensu Dennis, Brit. Ascomyc.: pl. 8 D. 1978.
Apothecia gregarious to caespitose, 8 – 33 mm high, 4 – 41
mm wide, broadly ear-shaped or shallowly cup-shaped, often
elongated on one side, normally split, often stipitate. Hymenium
light ochre (3A2, 3A3) to yellowish grey (4A3), sometimes with
weak rose stains or spots, when dried yellowish ochre (4A4,
4A5) to ochre (5A4). Receptacle surface citrine yellow (2A6,
2A7), slightly hygrophanous, in drying slightly paler, fading to
brownish ochre (4B5, 4B6) in age, when dried reddish ochre
(6B6–6D6) to reddish brown (5B4, 5B5), slightly furfuraceous
to slightly warty, some apothecia with low ridges coming from
the stipe, restricted to the base or reaching 1/2–1/3 of the apothecium, sometimes interconnected by low veins. Warts minute,
conical, concolorous to brownish. Stipe often well developed,
cylindrical, 2–12 × 2–5 mm. Basal tomentum and mycelium
abundant, white, seldom with purplish tones, very pale yellow
(4A2) when dried. Spores narrowly ellipsoid, often inequilateral,
with two large guttules, smooth, hyaline, (10–)10.5–12.5(–13)
× (4.5–)5–5.5(–6.2) µm (δm = 10.7–12.3 µm, Wm = 5–5.6 µm,
Qm = 2.1– 2.3; n = 10). Paraphyses straight to bent, seldom
curved, subclaviform to claviform at apices, 3–8 µm wide, with-
219
out notches, uppermost cell 20–71 µm long, when fresh with
homogeneous content or seldom containing slightly refractive,
light yellow granules at apices, often with a yellow refractive
body (Fig. 34d); when dried small, slightly refractive, hyaline
granules. Asci 153 –169 × 9 –10.5 µm. Apothecial section
750–1000 µm thick. Subhymenium c. 100 µm thick, of dense
textura intricata, hyphae 3–6 µm wide. Medullary excipulum
of textura intricata, 500–700 µm thick, hyphae 4–9(–13) µm
wide, sometimes slightly swollen, thin to thick-walled, hyaline
to very pale yellow, without resinous exudates. Ectal excipulum
of textura prismatica-angularis, 80–150 µm, cells thin-walled,
very pale yellow, unchanged in KOH, 10–33 × 6–15 µm. Surface with low flattened warts, 35–53 µm high, formed by short,
fasciculate, hyphoid hairs, of 2–3 ovoid cells, constricted at
septa, 6–9 µm wide. Non-warted parts with 2–4-celled hyphoid
hairs, with subclaviform to claviform uppermost cell, 43–58 ×
7.5–13 µm. Resinous exudates abundant, yellow or reddish
to brownish yellow, dissolving into amber drops in εδZ. Basal
mycelium of 3–7 µm wide, very pale yellow, slightly swollen
hyphae, not changing in KOH, with very scattered amorphous
or rod-shaped, yellow resinous exudates, dissolving in εδZ.
Specimens examined. dEnmark, Jylland, Nystrup Plantage, Kridtstien, on
calcareous soil under Abies, 1 Nov. 2007, T. Læssøe, Tδ-13332 (C); N Jylland,
Rold Skov, Buderupholm Bjergskov, by fence of Cypripedium, calcareous soil
along roadside, deciduous forest, 14 Sept. 1998, K. Hansen, KH.98.84 (C);
Jylland, Vorsø, under Salix caprea, 25 Sept. 1981, T. Læssøe, Tδ-0684 (C);
ibid., on base of Salix caprea, 8 Sept. 1982, T. Læssøe, Tδ-0754 (C); Sjælland, Geelskov, 10 km N of Copenhagen, 3 Aug. 1950, M. Lange (C-F-47967).
– Finland, Helsinki, Toukola, Koreankatu, Acer, Betula, Populus tremula, Salix
caprea, Sambucus, Sorbus, Aegopodium podagraria, Urtica dioica, 8 Sept.
1992, R. Saarenoksa 24592 (H); Varsinais-Suomi, δohja, Pähkinäniemi, very
rich, somewhat dry grass-herb forest with calcareous ground, 1 Aug. 1997,
U. Nummela-Salo & P. Salo 4051 (H); Varsinais-Suomi, δohja, Virkkala,
NE-slope of Pähkinäniemi, very rich, somewhat dry grass-herb forest with
calcareous soil, nearby Corylus avellana, Populus tremula, Betula pendula,
12 Sept. 2006, U. Nummela-Salo & P. Salo 10724 (H). – italy, Calabria, Acri
(CS), Croce di Greca, 14 Sept. 1995, C. Lavorato, Cδ 950914-01 (dupl. S).
– SwEdEn, Gotland, Fårö, Avanäset, under Pinus sylvestris, on humus,
sandy soil, 27 Sept. 2011, J.C. Zamora, KH.11.103 (S); Gotland, δojsta hed,
Russpark, grazed forest with Pinus sylvestris, on calcareous ground, 2 Oct.
2010, K. Hansen, K. Gillen & I. Olariaga, KH.10.311 (S); Småland, Stenbrohult, Stockanäs SSV of Stenbrohults kyrka, under Pyrus, Salix and Prunus
domestica, 9 Aug. 2011, G. Aronsson (UPS F-548414); Södermanland,
Södertalje, εörkö, Oaxen, in rich soil in deciduous forest, Corylus avellana,
Salix caprea, with Lactarius citriolens, 10 Sept. 1994, P. Höljer (H7003652);
Uppland, Uppsala, the park in front of the prison, on bare soil, among needles
etc. under Abies, 16 Aug. 1932, S. Lundell (S-F108335, Fungi Exs. Suec.
93); Öland, Högby, Horns kungsgårds Nature Reserve, under Corylus,
6 Aug. 2000, T. Knutsson, TK2000-057 (S).
Notes — Otidea minor is recognised macroscopically by
apothecia with a yellow outside, and shallow ribs and veins
at the base of at least some apothecia. Otidea concinna and
O. oregonensis can be distinguished by the broader spores
with a lower Qm (Fig. 29). In fresh material of O. minor, some
paraphyses had a strikingly, yellowish refractive body in the
upper part, a so far unique feature within Otidea. To assess its
taxonomic value, this feature has to be checked in additional
fresh material and in closely related species.
Otidea minor has been confused with O. phlebophora due to the
presence of ribs or veins at the base of the apothecia (δundell &
Nannfeldt 1934, Dennis 1978). Harmaja (2009a) proposed the
provisional name O. subconcinna for O. minor as circumscribed
here. After examining two collections annotated by him, we
consider the Finnish material to be conspecific with our Swedish finds, based on both morphological and molecular data.
Although Harmaja (2009a) did not directly compare O. minor
(as O. subconcinna) and O. phlebophora, he stated in the key
that O. phlebophora has ‘other tinges of yellow’ and mostly
straight paraphyses. We could not confirm these differences
in the material we examined. Instead, O. minor can be distin-
220
guished from O. phlebophora by the presence of at least some
apothecia without ribs, and ribs when present shallower and
less anastomosing, as well as normally split apothecia. In spite
of their morphological similarity, O. minor and O. phlebophora
are not sister species in our four-gene phylogeny, but both
are deeply nested within the O. concinna clade (Hansen &
Olariaga 2015; see also Fig. 3). Boudier´s plate under Otidea
cantharella var. minor shows shallow ribs at the base of some
apothecia, agreeing with our material, and as considered by
Harmaja (2009a). No original material could be located in PC,
and we therefore designate Boudier’s plate as the lectotype.
Persoonia – Volume 35, 2015
31. Otidea oregonensis K. Hansen & Olariaga, sp. nov. —
εycoBank εB808973; ITS barcode GenBank Kε010048;
Fig. 35
Etymology. Named after Oregon, the area where most of the specimens
of this species have been collected.
Holotype. USA, Oregon, Douglas Co., Umpqua National Forest, Diamond
δake Ranger District, under Pseudotsuga menziesii and Abies concolor, 7 Nov.
2010, J. Moore, εoorefun 58 (OSC). Isotype (S).
Apothecia gregarious to caespitose, 12–48 mm high, 23–80
mm wide, shallowly to deeply cup-shaped, sometimes elongated on one side or obconical, split, seldom entire, sometimes
stipitate. Hymenium greyish white (2A2) to pale ochre (4A2–
4A4), sometimes with rose stains or spots, when dried pale
ochre (4A2 – 4A4). Receptacle surface bright citrine yellow
Fig. 34 Otidea minor (KH.10.311)*. a. Apothecia; b. apothecium showing veined outer receptacle; c. spores; d. paraphyses showing a crystallized body; e. ectal
excipulum; f. ectal excipulum in εelzer’s reagent, resinous exudates mostly washed away, showing free hypoid hairs. — Scale bars = 10 µm; * = fresh material.
I. Olariaga et al.: A monograph of Otidea
(2A6–2A8), slightly hygrophanous, fading to brownish ochre
(4B5, 4B6) in age, when dried brownish ochre (4A4, 4B4, 5B4),
furfuraceous, sometimes some apothecia wrinkled-veined at
the base, seldom with short ribs reaching up 1/3 to the margin, partly covered by white tomentum from the base. Warts
sometimes present toward the base, minute, flat, concolorous.
Stipe sometimes well developed, cylindrical, 3–18 × 3–11 mm.
Smell mild. Basal tomentum and mycelium abundant, white,
when dried very pale yellow (4A2). Spores narrowly ellipsoid
to ellipsoid, sometimes inequilateral, with two large guttules,
rarely with a few small granules, smooth, hyaline, 10 –11.5
× 5.5 – 6(– 6.5) µm (δm = 10.4 –11.3 µm, Wm = 5.6 – 5.9 µm,
Qm = 1.8 – 2; n = 7). Paraphyses bent to curved, sometimes
straight, subclaviform to capitate at apices, 3–6(–8) µm wide,
221
without notches, uppermost cell 28–70 µm long, when dried
containing small, weakly refractive, hyaline granules. Asci
171–203 × 9–10.5 µm. Apothecial section 1000–1400(–2000)
µm thick. Subhymenium 100–140 µm thick, of dense textura
intricata, hyphae 2–4.5 µm wide, with scattered swollen, up to
12 µm wide cells. Medullary excipulum 650–850(–1400) µm
thick, of textura intricata, hyphae 5–9(–17) µm wide, sometimes swollen, thin- to thick-walled, hyaline, without resinous
exudates. Ectal excipulum of textura prismatica to textura
angularis, 70–100 µm thick, cells thin-walled, very pale yellow,
brighter yellow in KOH, 16–37 × 8–14 µm. Surface with low
warts, 35–70 µm high, formed by short, fasciculate hyphoid
hairs, of 2–4 ovoid cells, constricted at septa, 5–9 µm wide.
Non-warted parts with 2–4-celled, hyphoid hairs, with claviform
Fig. 35 Otidea oregonensis. a. Apothecia; b. apothecia showing faded colour in the outer receptacle; c. spores in water †; d. paraphyses in water †; e. ectal
excipulum in water †; f. ectal excipulum in KOH† (a, c–f: εoorefun 58, holotype; b: rh139). — Scale bars = 10 µm; † = dried material. — Photos: a. J. εoore;
b. R. Helliwell.
222
uppermost cell, 33–55 × 7–12 µm. Resinous exudates often
abundant, yellow, dissolving into amber drops in εδZ, brighter
yellow in KOH. Basal mycelium of 3–5(–8) µm wide, sometimes
slightly swollen, very pale yellow hyphae, not changing in KOH,
with very scattered, spheroid to rod-shaped, yellow resinous
exudates, dissolving in εδZ.
Specimens examined. USA, Oregon, Jackson Co., Rogue River National
Forest, under Pseudotsuga menziesii, Abies concolor, Pinus ponderosa,
2 Dec. 1999, B. Schroeter (OSC 72950); ibid., 27 Oct. 1990, D. McKay,
NSW6354 (OSC 132740, dupl. S); Oregon, Josephine Co., Bureau of δand
εanagement, εedford District, Grants Pass Resource Area, δittle Pickett
Creek, under Pseudotsuga menziesii, Calocedrus decurrens, Lithocarpus
densiflorus, Quercus chrysolepis, 19 Dec. 2000, R. Meyer (OSC 108041);
Oregon, εarion Co., near Breitenbush Hot Springs Community, 27 Oct.
1996, J. Trappe (OSC 56745); Oregon, Umpqua NF, Diamond δake RD,
under Pseudotsuga menziesii, 2 Nov. 2010, J. Moore, εoorefun 31 (S); ibid.,
old growth forest with Abies concolor, Pseudotsuga menziesii, 2 Nov. 2010,
R. Helliwell, rh139 (OSC); Washington, δewis Co., Gifford Pinchot National
Forest, Camp Creeks Falls Trail, 6 Nov. 1997, M. Castellano (OSC 56829).
Notes — Otidea oregonensis is characterised by a citrine
yellow receptacle surface, often a wrinkled to veined, or shallowly ribbed apothecial base, and relatively broad spores. The
apothecia are typically obconical cup-shaped, with a narrow
base. Otidea oregonensis was treated as O. rainierensis Kanouse by Peterson (1998), primarily based on the presence of
swollen apices of the paraphyses. Also, one of the paratypes
of O. rainierensis (EGS2179), sequenced by Peterson (1998),
is nested within the O. oregonensis lineage in our ITS-δSU
phylogeny (Fig. 3). Nevertheless, our sequences of the holotype
of O. rainieriensis, from four different gene regions, show it
belongs to another lineage, well separated from O. oregonensis
(Hansen & Olariaga 2015). Kanouse´s concept of O. rainier-
Persoonia – Volume 35, 2015
ensis was therefore mixed. The ITS-δSU phylogeny, including
a larger sampling of the O. concinna clade, likewise resolves
O. oregonensis as a distinct species.
Otidea phlebophora and O. minor, so far only known from
Europe, resemble O. oregonensis in the cup-shaped apothecia with yellow outside and swollen paraphyses. Otidea
phlebophora differs macroscopically from O. oregonensis in
predominantly entire apothecia, with always a strongly veinedribbed base reaching up 1/2 to the margin, while only small
veins are present in some apothecia of O. oregonensis. Based
on the material examined here, the slightly narrower spores in
O. phlebophora (Qm = 2–2.1 vs Qm = 1.8–2 in O. oregonensis)
are a constant and reliable character to separate these species
(Fig. 29). Otidea minor tends to have smaller, shallower and
more broadly cup-shaped apothecia than O. oregonensis. The
spores of O. minor have a higher Qm value (2.1–2.3) that does
not overlap with the Qm of O. oregonensis.
32. Otidea phlebophora (Berk. & Broome) Sacc., Syll. Fung.
8: 97. 1889 — Fig. 36
Basionym. Peziza phlebophora Berk. & Broome, Ann. εag. Nat. Hist.,
ser. III, 18: 122. 1866.
≡ Flavoscypha phlebophora (Berk. & Broome) Harmaja, Karstenia 14:
107. 1974.
Lectotype designated here: England, North Somerset, Brislington, 16
Sept. 1853 (K(ε) 144045, ex Herb. C.E. Broome) !; εycoBank εBT178087.
Apothecia gregarious, 8–12 mm high, 3–40 mm wide, shallowly to deeply cup-shaped, sometimes elongated on one side,
entire, seldom split, often stipitate. Hymenium pale ochre (4A2,
4A3), sometimes with rose stains or spots, when dried yellowish
Fig. 36 Otidea phlebophora (JV06-385). a. Apothecia; b. spores in water †; c. paraphyses in water †; d. ectal excipulum in water †. — Scale bars = 10 µm;
= dried material. — Photos: a. J. Vesterholt.
†
223
I. Olariaga et al.: A monograph of Otidea
ochre (4B4, 5B6, 5C6) to reddish brown (6C5, 6C6). Receptacle surface bright citrine yellow (3A7, 2A7), sometimes with a
greenish hue (2A6), slightly hygrophanous, fading to brownish
ochre (4B5) in age, when dried yellowish brown (5B6, 5C6) to
reddish brown (6C6, 6D6), furfuraceous, with high ridges in the
basal part of all apothecia reaching up 1/2–1/3 to the margin,
interconnected by veins, partly covered by white basal tomentum. Warts minute, flat to rounded, appressed, concolorous.
Stipe often well developed, cylindrical, 1–10 × 1.5–3 mm. Basal
tomentum and mycelium abundant, white, very pale ochre (5A2)
when dried. Spores narrowly ellipsoid, sometimes inequilateral,
with two large guttules, smooth, hyaline, (9.5–)10–11.5(–12)
× (4.5 –)5 – 5.5(– 6.2) µm (δm = 10.2 –11 µm, Wm = 4.9 – 5.1
µm, Qm = 2–2.1; n = 5). Paraphyses straight to bent, cylindrical to claviform at apices, 2–3.5 µm wide, without notches,
uppermost cell 23–46 µm long, when dried containing small,
rather refractive, hyaline guttules. Asci 130–153 × 8–9 µm.
Apothecial section 650–1000 µm thick. Subhymenium 80–100
µm thick, of dense textura intricata, hyphae 2 – 3 µm wide.
Medullary excipulum of textura intricata, 450–650 µm thick,
hyphae 4–7(–14) µm wide, sometimes swollen, thick-walled,
hyaline to very pale yellow, without resinous exudates. Ectal
excipulum of textura prismatica-angularis, 80–100 µm thick,
cells thin-walled, hyaline to very pale yellow, unchanged in KOH,
15–21 × 6–11 µm. Surface with low flattened warts, 40–60 µm
high, formed by short, fasciculate hyphoid hairs, of 3–4 ovoid
cells, constricted at septa, 8–12 µm wide. Non-warted parts
with 2–3-celled hyphoid hairs, with claviform uppermost cell,
30–60 × 7–13 µm. Resinous exudates abundant, yellow, dissolving into amber drops in εδZ. Basal mycelium of 3.5–7 µm
wide, slightly swollen, very pale yellow hyphae, not changing
in KOH, with very scattered, spheroid to rod-shaped, yellow
resinous exudates, dissolving in εδZ.
Specimens examined. dEnmark, NE Jylland, Rubjerg Knude Plantage,
under Abies in large fairy ring, 28 Aug. 2006, J. Vesterholt & L. Vesterholt,
JV06-385 (C); NE Sjælland, Ravnsholts Hegn, under Picea abies, 30 Aug.
1999, B.W. Pedersen (C-F-71506). – Finland, Varsinais-Suomi, δohja,
Jalassaari, Alho, by the Ahtiala manor, in rich forest with Quercus robur,
Corylus avellana, Prunus padus, cultivated Larix sibirica and Betula, 30 Aug.
1967, H. Harmaja (H6010675). – SwEdEn, Dalarna, Husby, Husby värdhus,
on lawn under Quercus, Tilia, Larix and Acer, 31 Aug. 1958, R. Morander
(UPS F-144691); Skåne, Helsinborg, Jordbodalen by Harlyckan, on sandy
ground in deciduous forest under Quercus, 13 July 1995, S.-Å Hanson, SÅH
30601 (C); Uppland, Djurö, Runmarö, Södersunda, in the city, on the ground
under Syringa, 18 Sept. 1949, G. Haglund & R. Rydberg (S-F108338, UPS
F-144689); Uppland, Uppsala, Sunnersta, Almlund, calcareous, humus-rich
clay, 23 Aug. 1986, J. Nitare (UPS F-119845). – UK, Northampshire, King´s
Cliffe, on soil, 1853, M.J. Berkeley (K(ε) 144046, syntype of P. phlebophora);
North Somerset, Brislington, The Beeches, Sept. 1853, C.E. Broome (K(ε)
194582, syntype of P. phlebophora).
Other specimen examined. Otidea integra — italy, Sopramonte, 1892,
G. Bresadola (S-F108342). δocality not specified, in silvis mixtis, 1892, G. Bresadola (PC 124965).
Notes — Otidea phlebophora is primarily characterised by
high anastomosing ribs and veins towards the apothecial base.
Diagnostic characters are in addition, the predominantly entire
apothecia, with a citrine yellow outside. For a comparison with
O. concinna, O. minor and O. oregonensis see Notes under
those species. Harmaja (1986) elevated O. integra to species
rank based on smaller apothecia and broader, curved paraphyses. The likely original specimen of O. integra collected in 1892
and kept in Bresadola’s herbarium (S-F108342), shows curved
paraphyses up to 5 µm broad, the same as Harmaja (1986,
2009a) gave for O. phlebophora. The paraphyses in the O. phlebophora material examined by us were straight to bent, and
otherwise similar to the original material of O. integra. In spite of
the morphological similarity, the ITS2 sequence of O. integra (281
bp obtained) is different from the ITS sequences of the material
assigned to O. phlebophora by us, and we preliminary accept
O. integra as a separate species. The position of O. integra is
without support in our ITS-δSU phylogeny (Fig. 3).
Nomenclatural notes — Harmaja (1974: 107) indicated a lectotype of O. phlebophora at K, but he gave no collection number
and the typification was not achieved. We have studied three
of four syntypes at Kew and select here the richest collection
containing ten apothecia as the lectotype. It conforms to the
current interpretation of the name, with several entire apothecia
(i.e. without a split), a ribbed-veined base seen on two of the
apothecia (the others with the base glued to the cardboard),
spores in the range 9.5 –11.7 × 4.7– 5.9 µm (δm = 10.6 µm,
Wm = 5.1 µm), paraphyses straight and enlarged at apices, and
abundant yellow resinous exudates in the ectal excipulum that
dissolve into amber drops in εδZ.
33. Otidea rainierensis Kanouse, εycologia 41: 674. 1949
Holotype. USA, Washington, Pierce County, Lower Tahoma Creek, εT
Rainier National Park, 23 Aug. 1948, A.H. Smith 30553 (εICH 14410) !
= Otidea kauffmanii Kanouse, εycologia 41, 6: 673. 1949.
Holotype. USA, εichigan, δakeland, 18 July 1915, C.H. Kauffman (εICH
14409) !
Kanouse (1949) only had dried material of O. rainierensis and
described the outside of the apothecia as ‘ochraceous buff ’,
‘cinnamon buff ’ to ‘wood brown’ and the hymenium as ‘avellaneous’, ‘vinaceous buff’ to ‘drab grey’. For O. kauffmanii she
had notes on fresh material and she separated it from O. rainierensis based on the presence of yellow tones in the apothecia
(outside ‘chamois’ to ‘ochraceous’, hymenium ‘cream buff ’). We
believe O. rainierensis does have yellow apothecial tones, as
we observed small yellow resinous exudates (in water) in the
type collection of O. rainierensis, and as observed in closely
allied species the yellow colour can fade and almost disappear
with age (Fig. 32b). Different spore sizes were also used to
distinguish the two species. The spores of the holotype of O.
kauffmanii are however, larger than noted in the protologue,
11.5–12.5 × 5.5–6.5 µm, δm = 11.9 µm, Wm = 6 µm, Qm = 2
(spores 8–10(–12) × 5–6(–7) µm in the protologue), and thus
overlapping with the spores of the holotype of O. rainierensis,
11–12 × 6.5–7 µm, δm = 11.7, Wm = 6.7 µm, Qm = 1.7. We suggest O. kauffmanii and O. rainierensis constitute a single species, based on molecular and morphological study of the type
material (Fig. 3). Our ITS sequences of the holotypes differ by
4 bp. See also comments on O. microspora under Excluded,
dubious and imperfectly known taxa. Otidea rainierensis is
characterised by a smooth apothecial base, long and relatively
broad spores, compared to the rest of the species in the O. concinna clade, and by paraphyses with abruptly enlarged, broadly
clavate to globose apices.
EXCLUDED, DUBIOUS AND IMPERFECTLY KNOwN
TAXA
Cochlearia (Cooke) δambotte, εém. Soc. Roy. Sci. δiege, ser.
2. 14: 323. 1888
Nom. illegit. Art. 53.1, non Cochlearia δ., Sp. Pl. 2: 647. 1753 (Cruciferae).
Basionym. Peziza subg. Cochlearia Cooke, εycographia part 6: 252
(‘Index Systematicus’). 1879.
Notes — Eckblad (1968) selected Peziza cochleata as the
type species for the genus Cochlearia considering it a synonym
of Otidea. However, Rifai (1968) had already selected Peziza
aurantia as the type species, herewith making Cochlearia a
synonym of Aleuria. He felt too much doubt had surrounded
the identity of P. cochleata and it better not be selected as the
type species.
224
Otidea abietina (Pers.) Fuckel, Jahrb. Nassauischen Vereins
Naturk. 23–24: 330. 1870 ‘1869 –1870’
Basionym. Peziza abietina Pers., Neues εag. Bot. 1: 113. 1794: Fr., Syst.
εycol. 2: 47. 1822.
≡ Discina abietina (Pers.) Rehm, Rabenh. Krypt.-Fl., ed. 2, 3: 977. 1896.
≡ Pseudotis abietina (Pers.) Boud., Icon. εycol. list prél. 600 sp.: 3 (unnumbered page). 1904.
Notes — Nannfeldt (1966) stated that the original material
belongs to Peziza badia, but Harmaja (2009a) and Carbone
(2010c) reported two collections in Persoon’s herbarium under
O. abietina representing O. propinquata and a third collection
O. bufonia. If typified with elements belonging to either O. propinquata or O. bufonia, the name O. abietina would take priority
over any of those. Otidea abietina is the type species of the
genus Pseudotis (Boud.) Boud. The identity of Pseudotis will
thus remain open until O. abietina is clarified or typified. Another
way to typify O. abietina would be to select an element belonging to Peziza badia. However, this choice should be studied
more thoroughly, since it would make Pseudotis available as
a genus name for the Peziza depressa-Ruhlandiella lineage, if
Peziza is split into smaller genera in the future (Hansen et al.
2005). For the time being we regard O. abietina as a nomen
ambiguum, as several others (Harmaja 2009a, Carbone 2010c,
Parslow & Spooner 2013).
Otidea alba Velen., εonograph. Discom. Bohemiae 1: 354.
1934
Holotype. CzECh rEpubliC, Karlštejn, Sept. 1924, Fechtner (PRε 149788) !
Notes — The holotype specimen has the typical oblong
spores of the O. alutacea complex, (13.5–)14.5–16.5(–17.5)
× 6.5–7.5 µm (δm = 14.9 µm, Wm = 6.9, Qm = 2.1). Judging from
the spore size, O. alba might represent either O. alutacea s.str.
or the O. alutacea clade 3b. It should be considered in future
studies of the complex.
Otidea aurantia (Pers.) εassee, Brit. Fungus-Fl. 4: 448. 1895
Basionym. Peziza aurantia Pers., Observ. εycol. 2: 76. 1800: Fr., Syst.
εycol. 2: 49. 1822.
Notes — This is the type species of Aleuria, A. aurantia
(Pers.: Fr.) Fuckel.
Otidea aurantia var. atromarginata (W. Phillips & Plowr.)
εassee, Brit. Fungus-Fl. 4: 449. 1895
Basionym. Peziza aurantia var. atromarginata W. Phillips & Plowr., Gard.
Chron. 17: 191. 1882.
Notes — The blood red disc with short, obtuse, dark brown,
3–4 septate hairs, giving the margin a dark appearance suggest
this may be a species of Melastiza. The spores are tuberculate
with thread-like appendages.
Otidea aurantia var. stipitata (W. Phillips) εassee, Brit. FungusFl. 4: 448. 1895
Basionym. Peziza aurantia var. stipitata W. Phillips, εan. Brit. Discom.:
57. 1887.
Notes — The small, bright scarlet apothecia with a ‘stem
equalling the height of the cup, 4 mm’ and the ornamented
spores suggest this is a species of Aleuria or Sowerbyella. It was
described as a variety of Aleuria aurantia (as Peziza aurantia
Oed.). Ramsbottom (1914) cited O. aurantia var. stipitata as a
synonym of Sowerbyella rhenana (as Aleuria rhenana Fuckel),
but the type material is presumably lost (Spooner & Yao 1995)
and no modern interpretation can be provided.
Persoonia – Volume 35, 2015
Otidea auriculariiformis Henn., Hedwigia 36: 232. 1897
Holotype. brazil, A. Glaziou no. 20181 (S-F9965, ex Herb. Sydow) !
Notes — This species belongs to the genus Phillipsia, Sarcoscyphaceae. The large spores, (30.5 –)31.5 – 36.5(– 37) ×
12.5–14.0 µm (δm = 33.2 µm, Wm = 13.1 µm, from 13 spores),
are ellipsoid, inequilateral in profile view, smooth or with faint
cyanophobic, parallel, longitudinal ridges. Asci seem thickwalled, with an internal eccentric thickened apical pad. The
medullary excipulum is of interwoven hyphae, running mostly
parallel with the outer surface and the ectal excipulum is a
narrow band of textura prismatica, with the long axes of the
cells parallel to the exterior. The tropical distribution and the
substrate, suggested in the diagnosis to be wood, are typical
for Phillipsia.
Otidea cinerascens Velen., Novit. εycol.: 152 .1947
Holotype. CzECh rEpubliC, εoravia, Žarošice, Aug. 1940, V. Vacek (PRε
151779).
Notes — The grey-ochraceous apothecia and spores with
parallel sides suggest O. cinerascens belongs to the O. alutacea complex. Type not studied by us, but annotated in 2009 by
B. Spooner as O. alutacea (a photograph of the collection and
annotation provided by Jan Holec, PRε).
Otidea cochleata (δ.) Fuckel, Jahrb. Nassauischen Vereins
Naturk. 23–24: 329. 1870. ‘1869 –1870’
Basionym. Peziza cochleata δ., Sp. Pl. 4: 183. 1753: Fr., Syst. εycol.,
Index: 129. 1832.
≡ Cochlearia cochleata (δ.) δambotte, Fl. εycol. Belgique 1: 323. 1880.
Notes — The interpretation of the original description is
difficult, but O. cochleata has been treated as a taxon in the O.
alutacea group (e.g. δundell & Nannfeldt 1938, Dissing 2000,
εornand & Courtecuisse 2005, Zhuang 2006), probably following Bulliard (1791: plate 154 as Peziza cochleata). Part of the
Bulliard plate (f. b) has now been selected as the lectotype for
O. alutacea (Carbone 2010a). The name O. cochleata should
be considered in future revisions of the O. alutacea complex.
For a review of the nomenclatural history of O. cochleata see
Carbone (2010a) and Parslow & Spooner (2013).
Otidea darjeelensis (Berk.) Sacc., Syll. Fung. 10: 4. 1892
Basionym. Peziza darjeelensis Berk., Hooker’s J. Bot. Kew Gard. εisc.
3: 202. 1851.
Notes — Rifai (1968) stated that the type specimen of P. darjeelensis at Kew has iodine positive asci and echinulate spores,
and does not belong to Otidea. Two collections of P. darjeelensis
are present in Kew that may represent original material: india,
Sikkim, J.D. Hooker (K(ε) 177412, ex Herb. Berkeley); and
india, Sikkim (K(ε) 177413, ex Herb. Cooke). No annotation
label by Rifai was found (B. Aguirre-Hudson, pers. comm. ), but
he most likely studied the collection from Berkeley’s herbarium,
because he listed J.D. Hooker as the collector. The collection
K(ε) 177413 might be a part of K(ε) 177412, because Cooke
(1876, f. 215) illustrated P. darjeelensis from specimens in
Berkeley’s herbarium. Both collections should be studied and
a lectotype selected.
Otidea dochmia (Berk. & ε.A. Curtis) Sacc., Syll. Fung. 8:
95. 1889
Basionym. Peziza dochmia Berk. & ε.A. Curtis, J. δinn. Soc. Bot. 10:
364. 1869.
≡ Phillipsia dochmia (Berk. & ε.A. Curtis) Seaver, N. Amer. Cup-fungi,
Operc.: 184. 1928.
I. Olariaga et al.: A monograph of Otidea
≡ Aurophora dochmia (Berk. & ε.A. Curtis) Rifai, Verh. Kon. Ned. Akad.
Wetensch., Afd. Natuurk., sect. 2, 57: 52. 1968.
Notes — This is the type species of the genus Aurophora.
Rifai (1968) distinguished Aurophora from Phillipsia by its fanshaped apothecia and the presence of a gelatinous matrix in
the medullary excipulum.
Otidea domingensis (Berk.) Sacc., Syll. Fung. 8: 97. 1889
Basionym. Peziza domingensis Berk., Ann. εag. Nat. Hist., ser. II, 9: 201.
1852.
Notes — This is the type species of the genus Phillipsia,
P. domingensis (Berk.) Berk. (see Hansen et al. 1999).
Otidea doratophora (Ellis & Everh.) Sacc., Syll. Fung. 8: 96.
1888
225
Notes — The original sense of O. grandis corresponds most
likely to a species of the Peziza depressa-Ruhlandiella lineage
(Hansen et al. 2005) due to the lack of a split. No original material of O. grandis seems to be kept in Persoon´s herbarium (δ).
The name O. grandis has been used for O. bufonia (Boudier
1905; specimen PC0093644 is O. bufonia) or O. unicisa (Kanouse 1949, δiu & Zhuang 2006). We regard it here as a nomen
dubium and confusum in agreement with Harmaja (2009a).
Otidea grandis var. scheremetjeffii Henn., Hedwigia 42, 3:
(116). 1903
Notes — Hennings (1903) described this taxon based on
specimens kept in formalin and the colours provided in the
protologue are probably imprecise. The spore size can fit
O. bufonia or O. onotica, but there is no type material extant in
B or S, and a precise interpretation cannot be proposed here.
Basionym. Peziza doratophora Ellis & Everh., J. εycol. 1: 90. 1885.
Notes — The small spores and asci, along with the pointed
paraphyses, suggest this taxon does not belong to Otidea. Cash
(1953) proposed O. doratophora is a synonym of Ionomidotis
irregularis (Schwein.) E.J. Durand (as Midotis irregularis).
Otidea euplecta (Cooke) Sacc., Syll. Fung. 8: 97. 1889
Basionym. Peziza euplecta Cooke, εycographia part 3: 125. 1876.
Holotype. USA, Alabama, Peters 4560 (K(ε) 161851, ex Herb. Berkeley
as Peziza phlebophora var.) !
Notes — The type is in a poor condition: one apothecium immature; the other infected. But the species likely belongs to
Sarcoscypha, Sarcoscyphaceae. The asci are inamyloid, thickwalled and with an eccentrically placed, thickened operculum,
spores are ellipsoid, slightly inequilateral, 19–21 × 10.5–11 µm,
smooth, and paraphyses straight, filiform, branching above. The
excipulum is composed of interwoven hyphae that give rise on
the outside to shallow pustules.
Otidea felina (Pers.) Bres., Fungi Trident. ser. 2, fasc. 14:
103. 1900
Basionym. Peziza felina Pers., εycol. Eur. 1: 223. 1822.
Holotype. FranCE, prope Pariseos, sylvula εeudon (δ0116774, Herb.
Persoon).
Notes — Van Vooren & Carbone (2012) revised the holotype
and demonstrated that it belongs to the O. alutacea group.
Parslow & Spooner (2013) considered O. felina a synonym of
O. alutacea. Further studies on the O. alutacea group should
consider the name O. felina, which might be epitypified for an
unequivocal interpretation.
Otidea fibrillosa εassee, Brit. Fungus-Fl. 4: 449. 1895
≡ Pseudaleuria fibrillosa (εassee) J. εoravec (‘Pseudoaleuria’), Acta
εus. εorav. Sci. Biol. 88: 51. 2003.
Otidea harperiana Rehm, Ann. εycol. 2: 34. 1904
Holotype. USA, Ohio, Blue εountains, on ground, 6 June 1903, Harper
333 (S-F9961, ex Herb. Rehm, ‘Herb. R.A. and A.ε. Harper 333’) !
Notes — This species is closely related or conspecific with
Peziza phyllogena Cooke. The asci are strongly amyloid in
εδZ with a general bluing over the apex and the spores are
ornamented with irregular, low, separate warts that are higher
and more densely placed at the poles (forming ‘pole caps’).
Our spore measurements from the holotype, 18–19.5 × 8–9
µm (δm = 18.6 µm, Wm = 8.5 µm, from 15 spores) are larger
than those given in the protologue, 15 –17 × 5–7 µm.
Otidea hirneoloides (Berk.) Sacc., Syll. Fung. 8: 96. 1889
Basionym. Peziza hirneoloides Berk. in Berkeley & Curtis, J. δinn. Soc.
Bot. 10: 365. 1869.
≡ Phillipsia hirneoloides (Berk.) Berk., J. δinn. Soc. Bot. 18: 388. 1881.
Notes — The wood-inhabiting, ear-shaped apothecia, and
especially, the cymbiform spores suggest this name is to be
referred to Phillipsia. Hansen et al. (1999) suggested P. hirneoloides belongs to the Phillipsia domingensis complex.
Otidea lechria (Berk. & Broome) Sacc., Syll. Fung. 8: 97. 1889
Basionym. Peziza lechria Berk. & Broome, J. δinn. Soc. Bot. 14: 103.
1875.
Holotype. Sri lanka, on rotten wood, Nov. 1867, G.H.K. Thwaites
(K(ε)161847, ex Herb. Berkeley).
Notes — B. Spooner annotated the holotype in 2008, and
noted it has amyloid asci and belongs to Peziza.
Otidea lilacina R. Heim & δ. Remy, Bull. Soc. εycol. France
48: 65. 1932
Notes — According to εoravec (2005) this is a species of
Pseudaleuria, P. fibrillosa.
Notes — The ornamented, multi-guttulate spores and straight
paraphyses suggest that this taxon does not belong to Otidea.
No original material could be traced in PC (B. Buyck, pers.
comm.).
Otidea grandis (Pers.) Boud., Bull. Soc. εycol. France 9:
10. 1893
Otidea lobata Rodway, Pap. & Proc. Roy. Soc. Tasmania: 116.
1925 ‘1924’
Basionym. Peziza grandis Pers., Ann. Bot. Usteri 15: 27. 1795.
≡ Peziza abietina var. grandis (Pers.) Pers., εycol. Eur. 1: 233. 1822.
≡ Aleuria grandis (Pers.) Gillet, Champ. France Discomyc.: 42. 1879.
≡ Scodellina grandis (Pers.) Seaver, N. Amer. Cup-fung., Operc.: 186.
1928.
Notes — Rifai (1968) stated that the type specimen appears
to represent the inoperculate genus Discinella.
226
Otidea luculenta (Cooke) εassee, Brit. Fungus-Fl., 4: 450.
1895 (‘leuculenta’)
Basionym. Peziza luculenta Cooke, εycographia part 3: 121. 1876.
Notes — The entire, orange apothecia and straight paraphyses with orange granules suggest this name does not belong
to Otidea. Nannfeldt (1966) noted that O. luculenta has ‘other
affinities’ than Otidea, but did not provide a generic placement.
Otidea luteonitens (Berk. & Broome) εassee, Brit. FungusFl. 4: 449. 1895
Basionym. Peziza luteonitens Berk. & Broome, Ann. εag. Nat. Hist., ser.
II, 7: 180. 1851.
Notes — This name is currently placed in Aleuria as Aleuria
luteonitens (Berk. & Broome) Gillet.
Otidea micropus (Pers.) Sacc., Syll. Fung. 8: 98. 1889
Basionym. Peziza micropus Pers., Icon. Desc. Fung. 2: 30. 1800: Fr.,
Syst. εycol. 2: 54. 1822.
Notes — This name is a synonym of Peziza varia (Hedw.:
Fr.) Fr. sensu Hansen et al. (2002).
Otidea microspora (Kanouse) Harmaja, Karstenia 15: 32. 1976
Basionym. Otidea alutacea var. microspora Kanouse, εycologia 41: 668.
1949.
Notes — Kanouse (1949) described this taxon as a variety
of O. alutacea. She indicated two different collections as the
type; A.H. Smith 9351 after the diagnosis and A.H. Smith 17699
in the material examined. We have studied A.H. Smith 9351
(εICH 14406, dupl. UPS F-629985 !) and it has far larger spores
than stated in the protologue (13–15.5 × 7–8 µm vs 9–10 ×
5.5–6.5 µm). The oblong spores, an ectal excipulum of textura
angularis, and the absence of yellow pigment in KOH indicate
it belongs to the O. alutacea complex. As for A.H. Smith 17699
(UPS F-629996 !), the spores match the original description.
The apothecial shape is similar to that of O. rainierensis. A GenBank ITS sequence of a paratype of O. microspora (A.H. Smith
30502) differs only 1 bp from the ITS sequence of the holotype
of O. rainierensis (Fig. 3), but we prefer not to select a lectotype
until all the original material has been examined. We therefore
treat the name as doubtful for the time being.
Otidea neglecta εassee, Grevillea 22: 66. 1894
Persoonia – Volume 35, 2015
species of Otidea. The Bucholtz herbarium was bought by the
FH, but no authentic material of O. olivacea could be located
there.
Otidea onotica var. ochracea (Fr.) Sacc., Syll. Fung. 8: 95.
1889
Basionym. Peziza onotica var. ochracea Fr., Syst. εycol. 2: 48. 1822: Fr.
loc. cit. (‘ß ochracea’).
≡ Peziza ochracea (Fr.) P. Karst., Not. Sallsk. Fauna Fl. Fenn. Forh. 10:
110. 1869.
≡ Otidea ochracea (Fr.) Seaver, Bull. δab. Nat. Hist. Iowa State Univ. 5:
45. 1904.
Notes — Fries (1822) stated that this variety is close to
O. onotica. It was elevated to species rank by Karsten (1869), who
later considered it a synonym of O. onotica (Karsten 1871). As
no original material exists and the original description is meagre,
we are not able to provide a good interpretation of this taxon.
Otidea pleurota (W. Phillips) Sacc., Syll. Fung. 8: 97. 1889
Basionym. Peziza pleurota W. Phillips in Cooke, εycographia part 5:
208. 1878.
≡ Iotidea pleurota (W. Phillips) Clem., Gen. Fungi: 175. 1909.
Holotype. England, 1877, W. Phillips (K(ε)29973).
Notes — Eckblad (1968) observed strongly amyloid asci in
the type and concluded it belongs to Peziza. B. Spooner annotated the type as ‘Peziza badiofusca ?’.
Otidea radiculata (Sowerby) Bres., Fungi Trident. ser. 2, fasc.
11–13: 72. 1898
Basionym. Peziza radiculata Sowerby, Col. Fig. Engl. Fung. 1: 46 (unnumbered page), t. 114. 1797: Fr., Syst. εycol. 2: 81. 1822.
Notes — This name is placed in Sowerbyella, as S. radiculata (Sowerby) Nannf. Yao & Spooner (2006) examined the
type at K and confirmed its placement.
Otidea reisneri Velen., České Houby 4 –5: 872. 1922
Notes — Svrček (1976) studied the type material and concluded it is a synonym of Sowerbyella radiculata (Sowerby: Fr.)
Nannf.
Otidea schulzeri Quél. in Schulzer, Hedwigia 24, 4: 150. 1885
Notes — This name was erected as a new name for O. auricula in the sense of Rehm (1883), Saccardo (1889) and Bresadola (1884, as Peziza). The species that these three authors
treated under the epithet auricula is Wynnella silvicola (Beck)
Nannf. in its current sense.
Notes — The apothecia of O. schulzeri were described as
elongated on one side, split, pale yellow-grey and pseudostipitate. However, the very thick flesh (3–4 mm), the straight
paraphyses and the spore size (20–28 µm) suggest this taxon
does not represent Otidea. Also Nannfeldt (1966) stated that
O. schulzeri could hardly belong to Otidea.
Otidea obtecta (Schwein.) Sacc., Syll. Fung. 8: 98. 1889
Otidea silvicola Beck in Sacc., Syll. Fung. 8: 97. 1889
Basionym. Peziza obtecta Schwein., Trans. Amer. Philos. Soc. ser. 2, 4:
170. 1832 ‘1834’.
Notes — This is Wynnella silvicola. The name was created
for Peziza atrofusca Beck, a later homonym for P. atrofusca
Berk. & ε.A. Curtis.
Notes — The original description gives stipitate, 1–1.5 cm
wide, cinnamon-coloured apothecia with a split, growing among
leaves. Seaver (1928) stated that the identity of O. obtecta is
uncertain. No original material could be located in PH and the
identification of this species cannot be inferred.
Otidea sparassis Quél., Rev. εycol. (Toulouse) 54: 65. 1892
Notes — The uniguttulate spores do not suggest an Otidea.
This taxon might refer to a sparassoid Helvella species.
Otidea olivacea Bucholtz, Bull. Soc. Imp. Naturalists εoscou
2: 325. 1897
Otidea subonotica Henn., Hedwigia 36: 232. 1897
Notes — The curved, uniguttulate spores and the occurrence
on a rotten trunk suggest that this taxon does not refer to a
Notes — The original description does not provide any discordant feature for Otidea, except it was reported from Brazil.
I. Olariaga et al.: A monograph of Otidea
No type material of O. subonotica exists in Hennings herbarium
in B (Carbone 2009) or in S, and its correct placement cannot
be inferred with certainty here.
Otidea succosa (Berk.) Thüm., εycoth. Univ. 15: no. 1411.
1879
Basionym. Peziza succosa Berk., Ann. εag. Nat. Hist., ser. I, 6: 358.
1841.
Notes — This is currently considered to be a species of
Peziza s.l. (Hansen et al. 2005).
Otidea tasmanica Rodway, Pap. & Proc. Roy. Soc. Tasmania:
116. 1925 ‘1924’
Notes — Rifai (1968) studied the type specimen of O. tasmanica in Kew and synonymised it with Peziza praetervisa
Bres. (sensu Dennis, as Rifai depicted ornamented spores
from the type specimen). Thus, O. tasmanica might be close
to, or a synonym of, Peziza subviolacea Svrček.
Otidea violacea A.δ. Sm. & Ramsb., Trans. Brit. εycol. Soc.
5: 237. 1916
Holotype. England, Warwickshire, W.B. Grove 1915 (K(ε) 30407, ex Herb.
W.B. Grove).
Notes — Parslow & Spooner (2013) examined the holotype
and concluded it is a species of Peziza (cf. azureoides Donadini).
Acknowledgements This monograph became more comprehensive thanks
to the great efforts of numerous mycologists who provided us with interesting
material. All our colleagues who contributed collections, discussed issues
or helped us in other ways are warmly thanked. Especially we thank Nancy
S. Weber, Rosanne Healy, εats Karström and Tapio Kekki for putting at our
disposal valuable material accompanied by colour photographs. We also
thank the curators and staff in the herbaria mentioned in the taxonomic
part, H.J.ε. Sipman (B), ε. Herrera and I. Salcedo (BIO), H. Knudsen (C),
P.G. Jamoni and D. Bolognini (GεFN), D.H. Pfister and G. δewis-Gentry
(FH), P. Salo and S. Stenroos (H), E.D. δiu (HKAS), Hong-εei δu (HεAS),
B. Aguirre-Hudson, B. Dentinger and B. Spooner (K), R. Rabeler and
P. Rogers (εICH), D. εcδaughlin (εIN), A.B. εujic and J. Spatafora (OSC),
B. Buyck (PC), A. Freire-Fierro (PH), J. Holec (PRε), S. Huhtinen (TUR,
TUR-A) and R. Berndt (ZH), for searching material and arranging loans. We
appreciate Kerstin Gillen´s assistance during fieldwork in 2010. Juan Santos
and Xiang-Hua Wang collected part of the material during their participation in the Swedish Pezizomycetes project. We are indebted to Xiang-Hua
Wang for sequencing a few important Otidea collections, for discussions and
help in editing the manuscript, and Seppo Huhtinen for instructive discussions on resinous exudates. We thank D.H. Hawksworth and δ.A. Parra for
nomenclatural advice. Donald H. Pfister and Trond Schumacher reviewed
the manuscript and we are grateful for their valuable comments and corrections. Funding for this research was provided by a grant from the Swedish
Taxonomy Initiative to K.H. (grant no. 143/07 1.4).
REFERENCES
Arpin N. 1969. δes caroténoïdes des Discomycètes: essai chimotaxinomique.
Bulletin εensuel de la Société δinnéenne de δyon 38 (suppl.): 1–169.
Batsch AJGK. 1783. Elenchus Fungorum. Halle, Germany.
Bonorden HF. 1851. Handbuch der allgemeinen mykologie als anleitung zum
studium derselben, nebst speciellen beiträgen zur vervolkommung dieses
zweiges der naturkunde. Schweizerbart, Stuttgart, Germany.
Boudier JδÉ. 1885. Nouvelle classification naturelle des Discomycètes charnus connus généralement sous le nom de Pezizes. Bulletin de la Société
εycologique de France 1: 91–120.
Boudier JδÉ. 1905. Icones εycologicae, livr. 6. Klincksieck, Paris, France.
Boudier JδÉ. 1906. Icones εycologicae, livr. 7. Klincksieck, Paris, France.
Boudier JδÉ. 1907. Histoire et classification des Discomycètes d’Europe.
Klincksieck, Paris, France.
227
Boudier JδÉ. 1908. Icones εycologicae, livr. 21. Klincksieck, Paris, France.
Boudier JδÉ. 1909a. Icones εycologicae, livr. 23. Klincksieck, Paris, France.
Boudier JδÉ. 1909b. Icones εycologicae, livr. 24. Klincksieck, Paris, France.
Boudier JδÉ. 1910. Icones εycologicae, livr. 29. Klincksieck, Paris, France.
Bresadola G. 1884. Fungi Tridentini novi, vel nondum delineati, descripti, et
iconibus illustrati. Ser. 1, fasc. 4–5: 43–70, pl. 46–75. εonauni, Trento, Italy.
Bresadola G. 1898. Fungi Tridentini novi, vel nondum delineati, descripti, et
iconibus illustrati. Ser. 2, fasc. 11–13: 47–81, pl. 151–195. Zippel, Trento,
Italy
Bresadola G. 1900. Fungi Tridentini novi, vel nondum delineati, descripti, et
iconibus illustrati. Ser. 2, fasc. 14: 83–118, pl. 196–217. Zippel, Trento, Italy.
Bresadola G. 1933. Iconographia mycologica. Vol. XXV. Trento, Italy.
Bulliard JBF. 1791. Herbier de la France. Tome 11. Paris, France.
Cao JZ, Fan δ, δiu B. 1990. Some species of Otidea from China. εycologia
82: 734–741.
Carbone ε. 2009. Il genere Otidea I. Sull’identità di Peziza onotica. Rivista
di εicologia 52: 11–28.
Carbone ε. 2010a. Il genere Otidea III. Identità e tipifizione di Peziza
alutacea. Bollettino dell’Associazione εicologica ed Ecologica Romana
80–81: 22–38.
Carbone ε. 2010b. Il genere Otidea IV. Prima parte. Otidea caligata, l’attuale
nome di Otidea abietina sensu Breitenbach & Kränzlin, con discussione
sull’abbandono dell’epiteto abietina. Schweizerische Zeitschrift für Pilzkunde 88: 14–17.
Carbone ε. 2010c. Il genere Otidea IV. Seconda parte. Otidea caligata,
l’attuale nome di Otidea abietina sensu Breitenbach & Kränzlin, con discussione sull’abbandono dell’epiteto abietina. Schweizerische Zeitschrift
für Pilzkunde 88: 64–66.
Carbone ε, Campo E, Vauras J. 2010. Records on Otidea mirabilis and
O. tuomikoskii from Finland. Karstenia 50: 25–34.
Carbone ε, Van Vooren N. 2010 ‘2009’. Il genere Otidea – II. Otidea fuckelii,
una nuova specie pubblicata per chiarire le differenti interpretazioni di
O. leporina. Rivista di εicologia 52: 313–330.
Cash EK. 1953. A record of the fungi named by J.B. Ellis. Part. II. The division
of mycology and disease survey, special publication 2: 167–345.
Clements FE, Shear Cδ. 1931. The genera of fungi. Wilson, New York, USA.
Cooke εC. 1876. εycographia seu Icones Fungorum. Part. 3. Williams &
Norgate, δondon, Great Britain.
Cooke εC. 1878. εycographia seu Icones Fungorum. Part. 5. Williams &
Norgate, δondon, Great Britain.
Dennis RWG. 1978. British Ascomycetes. 2nd edn. Cramer, Vaduz, Germany.
Dissing H. 2000. Pezizales Bessey. In: Hansen δ, Knudsen H (eds), Nordic
εacromycetes. Vol. 1. Ascomycetes: 55–127. Nordsvamp, Copenhagen,
Denmark.
Eckblad F-E. 1968. The genera of operculate Discomycetes. A reevaluation
of their taxonomy, phylogeny and nomenclature. Norwegian Journal of
Botany 15: 1–191.
Farr εδ, δeeusink JA, Stafleu FA. 1979. Index Nominum Genericorum
(Plantarum). Regnum Vegetabile vols. 100 –102. Bohn, Scheltema &
Holkema, Utrecht.
Franchi P, δami δ, εarchetti ε. 1999. Helvella leporina, nome corretto per
Helvella silvicola. Rivista di εicologia 42: 63–72.
Fries Eε. 1822. Systema mycologicum. II. Officina Berlingiana, Sweden.
Fuckel δ. 1870 ‘1869–1870’. Symbolae mycologicae. Jahrbücher des Nassauischen Vereins für Naturkunde 23–24: 1–459.
Gal ε le. 1947. Recherches sur les ornamentations sporales des Discomycètes operculés. Bibliotheca εycologica 28: 73 – 297. Reprint 1970.
Cramer, New York, USA.
Gonnermann W, Rabenhorst δ. 1869. εycologia Europaea. Abbildungen
sämmtlicher Schwämme Europas. 3. Dresden, Germany.
Gray SF. 1821. A natural arrangement of British plants. Baldwin, Cradock &
Joy, δondon, United Kingdom.
Greuter W, Brummitt RK, Farr E, et al. (eds). 1993. Names in current use
for extant plant genera. Regnum Vegetabile vol. 129. Koeltz Scientific
Books, Koenigstein.
Häffner J, Winterhoff W. 1989. Rezente Ascomycetenfunde VI. Otidea apophysata (Cooke & Phill.) Sacc. ein extrem seltener Öhrling. Beiträge zur
Kenntnis der Pilze εitteleuropas 5: 175–184.
Hansen K, δæssøe T, Pfister DH. 2002. Phylogenetic diversity in the core
group of Peziza inferred from ITS sequences and morphology. εycological
Research 106: 879–902.
Hansen K, δoBuglio KF, Pfister DH. 2005. Evolutionary relationships of the
cup-fungus genus Peziza and Pezizaceae inferred from multiple nuclear
genes: RPB2, ß-tubulin, and δSU rDNA. εolecular Phylogenetics and Evolution 36: 1–23.
Hansen K, Olariaga I. 2015. Species limits and relationships within Otidea
inferred from multiple gene phylogenies. Persoonia 35: 148–165.
228
Hansen K, Perry BA, Dranginis AW, et al. 2013. A phylogeny of the highly
diverse cup-fungus family Pyronemataceae (Pezizomycetes, Ascomycota)
clarifies relationships and evolution of selected life history traits. εolecular
Phylogenetics and Evolution 67: 311–335.
Hansen K, Pfister DH. 2006. Systematics of the Pezizomycetes - the operculate discomycetes. εycologia 98: 1029–1040.
Hansen K, Pfister DH, Hibbett DS. 1999. Phylogenetic relationships among
species of Phillipsia inferred from molecular and morphological data.
εycologia 91: 299–314.
Harmaja H. 1974. Flavoscypha, a new genus of the Pezizales for Otidea
cantharella and O. phlebophora. Karstenia 14: 105–108.
Harmaja H. 1976. New species and combinations in the genera Gyromitra,
Helvella and Otidea. Karstenia 15: 29–32.
Harmaja H. 1986. Studies on Pezizales. Karstenia 26: 41–48.
Harmaja H. 2009a. Studies in Otidea (Pezizales). Karstenia 48: 33–48.
Harmaja H. 2009b. A note on Otidea (Pezizales, Fungi). Phytotaxa 2: 49–50.
Hennings P. 1903. Beiträge zur Pilzflora Südamerikas. II. Hedwigia 36:
190–246.
Huelsenbeck JP, δarget B, Alfaro εE. 2004. Bayesian phylogenetic model
selection using reversible jump εarkov chain εonte Carlo. εolecular Biology and Evolution 21: 1123–1133.
Huhtinen S. 1990 ‘1989’. A monograph of Hyaloscypha and allied genera.
Karstenia 29: 45–252.
Jamoni PG. 2001. Validazione di nuovi taxa. Funghi e Ambiente 85–86: 56.
Jamoni PG. 2004. I funghi dell’ambiente alpino – XVIII. Funghi e Ambiente
94–95: 5–19.
Kanouse B. 1949. Studies in the genus Otidea. εycologia 41: 660–677.
Karsten PA. 1869. εonographia Pezizarum fennicarum. Notiser ur Sallskapets pro Fauna et Flora Fennica Forhandlingar 10: 99–206.
Karsten PA. 1871. εycologia Fennica. Pars prima. Discomycetes. Bidrag till
Kannedom of Finlands Natur och Folk 19: 1–263.
Kasparek F. 2000. Über einige bemerkenswerte Schlauchpilze. Der Tintling
19: 7–15.
Kimbrough JW. 1966. Studies in the Pseudoascoboleae. Canadian Journal
of Botany 44: 685–704.
Kirk Pε, Stalpers JA, Braun U, et al. 2013. A without-prejudice list of generic
names of fungi for protection under the International Code of Nomenclature
for algae, fungi, and plants. IεA Fungus 4: 381–443.
Korf RP. 1963. Discomycete flora of Asia, precursor II: A revision of the genera Acervus and Ascosparassis and their new positions in the Pezizales.
δloydia 26: 21–26.
Korf RP. 1972. Synoptic key to the genera of the Pezizales. εycologia 64:
937–994.
Korf RP. 1973a. Sparassoid ascocarps in the Pezizales and Tuberales.
Reports of the Tottori εycological Institute 10: 389–403.
Korf RP. 1973b. Discomycetes and Tuberales. In: Ainsworth GC, Sparrow
FK, Sussman S (eds), The fungi: An advanced treatise. Vol. 4A: 249–319.
Academic Press, New York, USA.
Korf RP, Zhuang WY. 1991. A preliminary discomycete flora of εacaronesia:
part 15, Terfeziaceae, and Otideaceae, Otideoideae. εycotaxon 40: 413–
433.
Kornerup A, Wanscher JH. 1961. Farver i farver. Politikens forlag, København, 1st edn.
Kumar S, Skjæveland Å, Orr RJS, et al. 2009. AIR: A batch-oriented web
program package for construction of supermatrices ready for phylogenomic
analyses. BεC Bioinformatics 10: 357.
δiu B, Cao JZ. 1987. Otideopsis yunnanensis gen. et sp. nov. of Pezizales
from China and its position in Pezizales system. Journal of Shanxi University, Natural Science edition 4: 70–73.
δiu CY, Zhuang WY. 2006. Relationships among some members of the genus Otidea (Pezizales, Pyronemataceae). Fungal Diversity 23: 181–192.
δundell S, Nannfeldt JA. 1934. Fungi Exsiccati Suecici, praesertim upsalienses Fasc. 1–2. Almqvist & Wiksells Boktryckeri, Sweden.
δundell S, Nannfeldt JA. 1938. Fungi Exsiccati Suecici, praesertim upsalienses. Fasc. 11–12. Almqvist & Wiksells Boktryckeri, Sweden.
δundell S, Nannfeldt JA, Holm δ. 1985. Fungi Exsiccati Suecici, prasertim
upsalienses. Fasc. 66 (No 3251–3300). Publications from the Herbarium,
University of Uppsala, Sweden 18: 1–18.
εaas Geesteranus RA. 1967. De fungi van Nederland 2a. Pezizales – deel
1, Discinaceae, Helvellaceae, εorchellaceae, Pezizaceae, Rhizinaceae.
Wetenschappelijke mededelingen van de Koninklijke Nederlandse Natuurhistorische Vereniging 69: 1–72.
εcNeill J, Barrie FR, Buck WR, et al. (eds). 2012. International Code of
Nomenclature for algae, fungi, and plants (εelbourne Code) adopted by
the Eighteenth International Botanical Congress εelbourne, Australia, July
2011. [Regnum Vegetabile No. 154.] Koeltz Scientific Books, Königstein,
Germany.
Persoonia – Volume 35, 2015
εedardi G. 1995. Considerazioni sul genere Otidea Fuck., 1870. Bolletino
del Circolo εicologico ‘G. Carini’ 29–30: 23–32.
εiller εA, Pfeiffer W, Schwartz T. 2010. Creating the CIPRES Science
Gateway for inference of large phylogenetic trees. In: Proceedings of the
Gateway Computing Environments Workshop (GCE), 14 Nov.: 1–8. New
Orleans, δA, USA.
εoravec J. 1986. A new species and two new combinations in the genus
Sowerbyella. εycologia Helvetica 2: 93–102.
εoravec J. 1988. Sowerbyella angustispora spec. nov. and Otideopsis kaushalii comb. nov. (Discomycetes, Pezizales, Pyronemataceae). εycologia
Helvetica 3: 135–142.
εoravec J. 2005. A world monograph of the genus Cheilymenia (Discomycetes, Pezizales, Pyronemataceae). δibri Botanici 21: 3–256.
εornand J, Courtecuisse R. 2005. δe genre Otidea et espèces affines en
France. Bulletin mensuel de la Société linnéenne de δyon 74, numero
spécial: 65–84.
Nannfeldt JA. 1937. Contributions to the mycoflora of Sweden. 4. On some
species of Helvella, together with a discussion of the natural affinities within
the Helvellaceae and Pezizaceae trib. Acetabuleae. Svensk Botanisk
Tidskrift 31: 47–66.
Nannfeldt JA. 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 Fuckel emend. Boud. and Sowerbyella Nannf. n.
gen. Svensk Botanisk Tidskrift 32: 108–120.
Nannfeldt JA. 1966. On Otidea caligata, O. indivisa and O. platyspora (Discomycetes, Operculatae). Annales Botanici Fennici 3: 309–318.
Nei ε. 1987. εolecular evolutionary genetics. Columbia University Press, USA.
Parslow ε, Spooner B. 2013. The British species of Otidea: overview and
the large spored species. εycosystema 32: 347–365.
Pérez-Butrón Jδ, Fernández-Vicente J. 2008. Otidea apophysata, en la
Península Ibérica. Errotari 5: 36–43.
Perry BA, Hansen K, Pfister DH. 2007. A phylogenetic overview of the
family Pyronemataceae (Ascomycota, Pezizales). εycological Research
111: 549–571.
Persoon CH. 1799. Observationes εycologicae. Pars secunda. δeipzig &
δuzern.
Persoon CH. 1822. εycologia europaea. I. Erlangae, Germany.
Peterson ET. 1998. Systematics of the genus Otidea in the Pacific Northwest.
Unpublished thesis. Oregon State University.
Pfister DH. 1979. A monograph of the genus Wynnea (Pezizales, Sarcoscyphacea). εycologia 71: 144–159.
Pfister DH, Halling RE. 1989. Ascosparassis heinricheri from Venezuela: an
extended distribution. εycotaxon 35: 283–285.
Quélet δ. 1886. Enchiridion fungorum in Europa media et praesertim in Gallia
vigentium. Douin, Paris, France.
Rahm E. 1958. Otidea pusilla nov. spec. Zwerg-Öhrling. Schweizerische Zeitschrift für Pilzkunde 36: 33–35.
Rambaut A. 2002. Se-Al. Sequence Alignment Editor. Version 2.0 alpha 11.
University of Oxford, Oxford. From http://tree.bio.ed.ac.uk/software/seal/.
Ramsbottom J. 1914. A list of the British species of Discomycetes arranged
according to Boudier’s system, with a key to the genera. Transactions of
the British εycological Society 4: 343–381.
Rehm H. 1883. Ascomyceten XIV. Hedwigia 22: 1–18 (reprint).
Rifai εA. 1968. The Australasian Pezizales in the herbarium of the Royal
Botanic Gardens Kew. Verhandelingen der Koninklijke Nederlandse Akademie van Wetenschappen, Afd. Natuurkunde, Tweede Reeks 57: 1–295.
Ronquist F, Teslenko ε, εark P van der, et al. 2012. εrBayes 3.2: Efficient
Bayesian phylogenetic inference and model choice across a large model
space. Systematic Biology 61: 539–542.
Saccardo PA. 1884. Conspectus generum discomycetum hucusque cognitorum. Botanisches Centralblatt 18: 213–256.
Saccardo PA. 1889. Sylloge Fungorum omnium hucusque cognitorum 8.
Patavii.
Schaeffer JC. 1763. Fungorum qui in Bavaria et Palatinatu circa Ratisbonam
nascuntur icons natives coloribus expressae. Tomus II. Regensburg, Germany.
Schaeffer JC. 1774. Fungorum qui in Bavaria et Palatinatu circa Ratisbonam
nascuntur icons natives coloribus expressae. Tomus IV. Regensburg, Germany.
Seaver FJ. 1904. The discomycetes of eastern Iowa. Bulletin from the laboratories of natural history of the State University of Iowa 5: 230–297.
Seaver FJ. 1927. A tentative scheme for the treatment of the genera of the
Pezizaceae. εycologia 19: 86–89.
Seaver FJ. 1928. The North American cup-fungi (operculates). Reprint 1961.
Hafner Publishing Company, New York, USA.
Smith εE, Healy RA. 2009. Otidea subterranea sp. nov.: Otidea goes below
ground. εycological Research 113: 858–866.
229
I. Olariaga et al.: A monograph of Otidea
Sowerby J. 1797. Coloured figures of English fungi or mushrooms. Vol. I.
δondon, England.
Spooner Bε, Yao Y-J. 1995. Notes on British taxa referred to Aleuria. εycological Research 99: 1515–1518.
Stamatakis A. 2006. Raxml-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics
22: 2688–2690.
Svrček ε. 1976. A taxonomic revision of Velenoský’s types of operculate
discomycetes (Pezizales) preserved in national museum, Prague. Sborník
národního muzea v Praze. Acta εusei Nationalis Pragae 32 B: 115–194.
Taylor JW, Jacobson DJ, Kroken S, et al. 2000. Phylogenetic species recognition and species concepts in fungi. Fungal Genetics and Biology 31: 21–32.
Thiers B (ed). 2014. Index Herbariorum. A global directory of public herbaria
and associated staff. New York Botanical Garden´s Virtual Herbarium. http://
sweetgum.nybg.org/ih/.
Van Vooren N. 2010. Note sur Otidea mirabilis (Pezizales). Ascomycete.
org 2: 33–35.
Van Vooren N. 2011a. δe genre Otidea V. Otidea apophysata et ses interprétations. Bulletin mycologique et botanique Dauphiné-Savoie: 165–174.
Van Vooren N. 2011b. Premiers signalements d’Otidea caligata (Nyl.) Sacc.
(Ascomycota, Pezizales) en France. Ascomycete.org 3: 61–64.
Van Vooren N, Armada F. 2011. Redécouverte d’Otidea platyspora Nannf.
(Ascomycota, Pezizales) en France. Bulletin mycologique et botanique
Dauphiné-Savoie 203: 57–62.
Van Vooren N, Carbone ε. 2012. The genus Otidea. VI. Otidea felina and its
interpretations. Ascomycete.org 4: 29–34.
Van Vooren N, Hairaud ε, Jindřich O. 2008. Otidea tuomikoskii, Otidea papillata et Otidea papillata f. pallidefurfuracea f. nov. trois taxons remarquables
appartenant au genre Otidea (Pezizales, Pyronemataceae). Bulletin mycologique et botanique Dauphiné-Savoie 188: 47–57.
Van Vooren N, Olariaga I, Tabarés ε. 2011. First record of Otidea caeruleopruinosa Harmaja (Ascomycota, Pezizales) in the Iberian Peninsula.
Ascomycete.org 3: 43–46.
Yao Y-J, Spooner Bε. 2006. Species of Sowerbyella in the British Isles, with
validation of Pseudoombrophila sect. Nannfeldtiella (Pezizales). Fungal
Diversity 22: 267–279.
Zhuang WY. 2006 ‘2005’. Notes on Otidea from Xinjiang, China. εycotaxon
94: 365–370.
Zhuang WY. 2010. Taxonomic assessment of some pyronemataceous fungi
from China. εycotaxon 112: 31–46.
Zhuang WY, Korf RP. 1987. Some new species and new records of discomycetes in China. II. εycotaxon 29: 309–314.
Zhuang WY, Korf RP. 1989. Some new species and new records of discomycetes in China. III. εycotaxon 35: 297–312.
Zhuang WY, Yang Zδ. 2008 ‘2007’. Some pezizalean fungi from alpine areas
of southwestern China. εycologia εontenegrina 10: 235–249.
INDEX to species, varieties and forms
Accepted names treated in the monograph are in bolditalic.
abietina 166, 167, 193, 195, 197, 224
abietina var. grandis 225
abietina f. / var. nigra 195
alba 185, 224
alutacea 175–177, 179, 181, 182, 224, 225
alutacea var. microspora 226
angusta 176, 198, 200
apophysata 167, 168, 178, 179, 180–182
atrofusca 226
aurantia 223, 224
aurantia var. atromarginata 224
aurantia var. stipitata 224
auricula 167, 185, 187, 226
auriculariiformis 224
azureoides 227
badiofusca 226
bicolor 178, 211
borealis 168, 171, 174, 176–178, 212, 213
boudieri 180
brevispora 174, 179, 205
brunneoparva 168, 179, 187, 191
bufonia 175, 176, 179, 185, 192, 205, 206, 209,
211, 213, 219, 224, 225
caeruleopruinosa 168, 174, 177, 179, 213,
215, 219
caligata 189, 193, 195, 197
cantharella 167, 178, 189, 191, 192, 193, 197,
216, 219
cantharella var. minor 188, 189, 219, 220
cinerascens 185, 224
cochleata 166, 167, 180, 183–185, 195, 206,
207, 223, 224
cochleata var. alutacea 182
cochleata var. umbrina 206, 175
concinna 167, 168, 174, 175, 177, 178, 189,
210, 213, 216, 219, 223
crassa 174, 185, 187
daliensis 167, 168, 178–180, 181, 192, 197
darjeelensis 224, 225
dochmia 224
domingensis 225
doratophora 225
euplecta 225
felina 179, 180, 185, 187, 225
fibrillosa 225
flavidobrunneola 177, 179, 213, 216, 217
formicarum 168, 171, 174, 176, 177, 179, 189,
197, 200, 201
fuckelii 185, 187
fusconigra 192, 205
grandis 205, 207, 208, 209, 225
grandis var. scheremetjeffii 225
harperiana 225
hirneoloides 225
indivisa 195–197
integra 168, 174, 178, 223
kauffmanii 171, 175, 213, 214, 223
kaushalii 174, 176, 177, 178, 202, 205
kunmingensis 178, 184, 185
lactea 174, 178, 179, 213, 219
lechria 225
leporina 166, 167, 171, 174, 175, 177, 179, 185,
188, 189, 192, 209–211
leporina f. major 185
leporina f. / var. minor 185, 195
leporina var. onotica 209
leporina var. rubescens 185
lilacina 225
lobata 225
lohjaënsis 200
luculenta 226
luteonitens 226
marsupium var. pyxidata 216
micropus 226
microspora 226
minor 174, 177, 178, 189, 213, 217, 219, 222
mirabilis 171, 174–176, 179, 192, 205, 207,
209, 211, 213
myosotis 185, 187
nannfeldtii 168, 175–177, 179, 185, 189, 191,
197, 198, 201, 202
neglecta 226
obtecta 226
ochracea 226
olivacea 192, 226
olivaceobrunnea 178, 192
onotica 166, 167, 174, 177–179, 205, 209,
225, 226
onotica var. brevispora 174, 205, 206
onotica var. ochracea 226
oregonensis 168, 174, 177, 178, 213, 217,
219, 220, 223
papillata 175, 179, 185, 189, 191, 198, 200
papillata f. pallidefurfuracea 189
pedunculata 206, 207, 209
phlebophora 167, 168, 174, 178, 213, 217, 219,
220, 222, 225
phyllogena 225
platyspora 178, 179, 180, 182
pleurota 226
praetervisa 227
propinquata 167, 177, 178, 191–193, 195, 224
pseudobadia 206, 209
pseudoleporina 168, 174, 176, 177, 179, 185,
188, 197, 200, 201
purpurea 179, 205, 211
pusilla 192
pyxidata 216
radiculata 226
rainierensis 174, 175, 179, 213, 214, 217, 222,
223, 226
reisneri 226
rhenana 224
rosea 209
schulzeri 226
scutellata 216
shimizuensis 167
silvicola 167, 185, 187, 226
sinensis 174, 178, 213
smithii 168, 171, 176, 179, 192, 205, 209, 211
sparassis 226
subconcinna 219
subformicarum 168, 171, 174, 179, 197, 198,
200
subonotica 226
subpurpurea 178, 211
subterranea 167, 178, 179, 181
subviolacea 227
succosa 227
tasmanica 227
tianshuiensis 178, 213
tuomikoskii 171, 177, 179, 185, 189, 200, 202
umbrina 184, 187, 206–208
unicisa 167, 168, 174, 177, 178, 202, 205,
211, 225
varia 226
violacea 227
yunnanensis 167, 174, 178, 179, 202, 204, 205