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The genera Helvella and Dissingia (Ascomycota: Pezizomycetes) in Europe – Notes on species from Spain

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Phylogenetic analyses of 115 newly collected Helvella specimens from Spain using three genetic markers [heat shock protein 90 ( hsp ), RNA polymerase II second largest subunit ( rpb2 ) and the nuclear large subunit ribosomal DNA (LSU)] confirm the assignment of the Spanish collections to one Dissingia and 30 Helvella species. The analyses were supplemented with an additional sample of 65 Spanish and extralimital Helvella specimens from the fungaria of Oslo (O), Trondheim (TRH), Copenhagen (C), Uppsala (UPS), Stockholm (S) and Venice (MCVE). Nine species are described as new, i.e. Helvella fuscolacunosa , H. hispanica , H. iberica , H. inexpectata , H. neopallescens , H. phlebophoroides , H. poculiformis , H. retinervis , and H. terricola . We present photographs of a selection of fresh specimens and provide descriptions of all species of this diverse South European Mediterranean element of the genera in Europe.
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© 2020 Westerdijk Fungal Biodiversity Instute 65
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
Fungal Systemacs and Evoluon
doi.org/10.3114/fuse.2020.06.05
VOLUME 6
DECEMBER 2020
PAGES 65–93
INTRODUCTION
Helvella is a species-rich genus of apothecial ascomycetous fungi
(Pezizomycetes: Pezizales: Helvellaceae) of terrestrial biomes of
the northern and southern hemispheres. Many authors have
contributed to the species-level taxonomy of the genus, the more
comprehensive are those of Boudier (1905–1910), Nannfeldt
(1932, 1937), Dissing (1966a, b, Dissing & Nannfeldt 1966),
Weber (1972, 1975), Harmaja (1977b, 1979), Häner (1987),
Abbo & Currah (1997), Vooren (2010, 2014), Landeros et al.
(2012, 2015), and Skrede et al. (2017). In Dissing’s monograph
of “The genus Helvella in Europe” (Dissing 1966a), he recognised
26 species in Europe, a list that was later expanded to 28 (Dissing
1972) and to 38 in a survey 30 yr later, but then conned to
species of the Nordic countries only (Dissing 2000). Meanwhile,
Häner (1987) accepted 41 species of Helvella from Germany
and the European connent.
In a preceding study by Skrede et al. (2017), we used a
mullocus genealogical approach to re-assess species limits,
phylogeny and species-level taxonomy within the genus. Based
on a selecon of fungarium collecons and newly collected
specimens from North and Central Europe, Skrede et al. (2017)
assigned molecular specics and morphological characteriscs
to altogether 55 phylogenec species of Helvella in Europe. In
that work, however, specimens from Southern Europe were
seriously under-represented. This paved the way for a review of
Helvella specimens from Southern Europe using sequence data.
The present work adds nine species to the checklist of Helvella-
species from the European connent.
In a recent account on the generic limits and phylogeny of the
family Helvellaceae, Hansen et al. (2019) erected the segregate
genus Dissingia to encompass the morphological and molecular
disnct members, previously recognised as the /leucomelaena
lineage of Helvella sensu lato (Landeros et al. 2015, Skrede et
al. 2017). This genus has been included in this revisionary study.
Calonge & Arroyo (1990) published an annotated species list
of the genus Helvella in Spain, following the species concepts
in Dissing (1966b). Their material, deposited at the fungarium
of the Botanic Garden of Madrid, comprised 22 species and
one variety assumed to include all species that occurred in the
country. For a survey of early studies on Spanish Helvella species,
the reader is referred to the introductory notes and bibliography
presented in the work by Calonge & Arroyo (1990).
MATERIALS AND METHODS
Table 1 summarises Spanish and extralimital samples of Helvella
specimens included in this study that are deposited in the
fungaria of Copenhagen (C), Oslo (O), Sociedad de Ciencias
Aranzadi (ARAN), Stockholm (S), Trondheim (TRH), Uppsala
(UPS), and Venice (MCVE). In addion, Dissingia leucomelaena
specimens from Spain were included as a suitable outgroup in
the phylogenec analyses. A selecon of in situ photographs
of typical specimens of all species of the Spanish specimens is
given.
The morphological analyses follow the outline in Skrede et
al. (2017). For the ascospore measurements, 10 ejected mature
The genera Helvella and Dissingia (Ascomycota: Pezizomycetes) in Europe – Notes on species
from Spain
I. Skrede1*, L. Ballester Gonzalvo2, C. Mathiesen1, T. Schumacher1
1Department of Biosciences, University of Oslo, P.O. Box 1066, 0316 Oslo Norway
2Calle Octavio de Toledo 25, 50007, Zaragoza, Spain
*Corresponding author: inger.skrede@ibv.uio.no
Abstract: Phylogenec analyses of 115 newly collected Helvella specimens from Spain using three genec markers [heat
shock protein 90 (hsp), RNA polymerase II second largest subunit (rpb2) and the nuclear large subunit ribosomal DNA
(LSU)] conrm the assignment of the Spanish collecons to one Dissingia and 30 Helvella species. The analyses were
supplemented with an addional sample of 65 Spanish and extralimital Helvella specimens from the fungaria of Oslo
(O), Trondheim (TRH), Copenhagen (C), Uppsala (UPS), Stockholm (S) and Venice (MCVE). Nine species are described
as new, i.e. Helvella fuscolacunosa, H. hispanica, H. iberica, H. inexpectata, H. neopallescens, H. phlebophoroides, H.
poculiformis, H. renervis, and H. terricola. We present photographs of a selecon of fresh specimens and provide
descripons of all species of this diverse South European Mediterranean element of the genera in Europe.
Key words:
Helvella
molecular phylogeny
new taxa
Pezizales
Citaon: Skrede I, Ballester Gonzalvo L, Mathiesen C, Schumacher T (2020). The genera Helvella and Dissingia (Ascomycota: Pezizomycetes) in Europe
– Notes on species from Spain. Fungal Systemacs and Evoluon 6: 65–93. doi: 10.3114/fuse.2020.06.05
Effectively published online: 3 April 2020
Corresponding editor: P.W. Crous
© 2020 Westerdijk Fungal Biodiversity Instute
Skrede et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
66
Table 1. Locality, idencaon, and type informaon for Helvella and Dissingia specimens included in the study. The ID column contains type
informaon and sequencing fungarium idencaon numbers. The columns LSU, hsp and rpb2 contain GenBank accession numbers. Accession
numbers in italics are new accessions from this study, accession numbers in bold are included in maximum likelihood analyses resulng in Fig. 1.
Species ID LSU hsp rpb2 Locality Coll. date Collector
Collector ID
D. leucomelaena H1930 (O-F-256543) MN598130 MN626787 Spain, Navarra,
Viana
12.03.2016 A. Ezquerro –
AEA0010
H1931 (O-F-256544) MN598131 MN626788 Spain, Rioja,
Logrono
11.03.2017 A. Ezquerro,
L. Ballester –
LB17031101
H2892 (TRH-12603) MN598182 Spain, Balearic
Islands, Mallorca,
Galilea
04.02.1989 S. Sivertsen
H2893 (TRH-12606) MN598183 Spain, Balearic
Islands, Mallorca,
Galilea
29.12.1997 S. Sivertsen
H2894 (TRH-12615) MN598184 Spain, Balearic
Islands, Mallorca,
Galilea
29.12.1987 S. Sivertsen
H2895 (TRH-12616) MN598185 Spain, Balearic
Islands, Mallorca,
Formentor
07.04.1982 L. Sivertsen
H2896 (TRH-12619) MN598186 Spain, Balearic
Islands, Mallorca,
Lluch
08.04.1982 S. Sivertsen
H2898 (TRH-12628) MN598187 Spain, Balearic
Islands, Mallorca,
Son Torrella
28.01.1979 S. Sivertsen
H2899 (TRH-12629) MN598188 Spain, Balearic
Islands, Mallorca,
Son Torrella
28.01.1979 S. Sivertsen
H2900 (TRH-12630) MN598189 Spain, Balearic
Islands, Mallorca,
Massanella
30.01.1979 S. Sivertsen
H. acetabulum H1927 (O-F-256511) MN598127 MN626784 Spain, Cadiz,
Castellar de la
Fontera
05.03.2017 M. Plaza –
AH0170305
H1934 (O-F-256512) MN598134 Spain, Zaragoza,
Paniza
30.04.2006 L. Ballester –
LB06043001
H1935 (O-F-256513) MN598135 Spain, Rioja,
Banos de Rio Tobia
02.04.2016 A. Ezquerro –
AEA0026
H1936 (O-F-256514) MN598136 Spain, Girona, S.
Mar d’Empuries
12.03.2017 M. Perez de
Gregorio –
PG120317
H1940 (O-F-256515) MN598140 Spain, Rioja,
Sorzano
24.06.2013 R. Marnez,
L. Ballester –
LB13062401
H1942 (O-F-256516) MN598141 Spain, Rioja,
Ventas Blancas
29.04.2012 G. Munoz –
GM2507
H1943 (O-F-256517) MN598142 Spain, Rioja,
Almarza de
Cameros
28.04.2016 G. Munoz –
GM2920
H2876 (TRH-12717) MN598166 Spain, Balearic
Islands, Mallorca,
Lluch
08.04.1982 L. Sivertsen
© 2020 Westerdijk Fungal Biodiversity Instute
Dissingia and Helvella species from Spain
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
67
Table 1. (Connued).
Species ID LSU hsp rpb2 Locality Coll. date Collector
Collector ID
H2877 (TRH-12620) MN598167 Spain, Balearic
Islands, Mallorca,
Coma Freda
08.04.1982 E. Dahl
H2878 (TRH-12622) MN598168 Spain, Balearic
Islands, Mallorca,
Coma Freda
08.04.1982 G. Gulden
H2879 (TRH-12626) MN598169 Spain, Balearic
Islands, Mallorca,
Coma Freda
26.04.1984 L., S. Sivertsen
Epitype, H410 (C-F- Fungi.
Exs. Suec. 1354)
KY773154 KY784506 KY772752 Sweden,
Ôstergötland
14.09.1945 J. A. Nannfeldt
H. atra H1019 (O-F-256518) MN644503 MN626770 Spain, Bizkaia,
Gorliz-Berreaga
17.11.2012 A. Melendez,
A. Ayala
PIPE0966
H233 (O-253245) KY773063 KY784351 KY772600 Georgia, Caucasus 13.07.1988 E. Johannesen
Epitype, H406 (C-F- Fungi.
Exs. Suec. 2066)
KY784502 KY772748 Sweden, Uppland 13.07.1938 S. Lundell
H. calycina Epitype, H022 (O-253255) KY772915 KY784198 KY772442 Norway, Oppland,
Dovre
07.08.2009 T. Carlsen, T.
Schumacher, I.
Skrede
H382 (C-F-21122) KY773143 KY784480 KY772727 Denmark, Klim
Bjerg
18.10.1993 Jye, Lässøe
H384 (C-F-41038) KY773144 KY784482 KY772729 Denmark, Klim
Bjerg
16.07.1999 L. Hansen, J.
Vesterholt
H845 (O-F-256519) MN598102 Spain, Rioja, Autol 08.05.2009 G. Munoz –
GM1504
H846 (O-F-256520) MN598103 Spain, Palencia,
Duenas
30.04.2016 A. Calzada –
ACD1844
H847 (O-F-256521) MN598104 Spain, Rioja,
Sorzano
24.06.2013 R. Marnez,
L. Ballester –
LB13062404
H848 (O-F-256522) MN598105 Spain, Rioja,
Ventas Blancas
29.04.2015 R. Marnez,
L. Ballester –
RM2338
H849 (O-F-256523) MN598106 Spain, Valladolid,
Aldeamayor de
San Marn
14.05.2016 F. Bellido –
FB160514
H. cosfera H131 (O-68514) KY772982 KY784264 KY772514 Norway, Akershus 27.07.2007 P.A. Bergersen
H247 (O-253283) KY773074 KY784365 KY772613 Norway, Oppland.
Vågå
29.08.1998 S. Kjølner
H. crispa H135 (O-18774) KY772986 KY784268 KY772518 Norway, Akershus 15.10.2004 P. Marstad
H2624 (O-F-256524) MN598151 MN626795 Spain, Balearic
Islands, Menorca,
Binimo, Ferreries
08.12.2007 J. L. Melis –
GM071208-6
H2626 (O-F-256525) MN598153 MN626754 Spain, Huesca,
Villanúa
12.10.2007 L. Ballester –
LB07101201
H2634 (O-F-256526) MN598161 Spain, Huesca,
Bergua
03.11.2017 J. Garcia,
A. Hereza –
LB17110301
H2880 (TRH-12605) MN598170 Spain, Balearic
Islands, Mallorca,
Galilea
29.12.1997 S. Sivertsen
© 2020 Westerdijk Fungal Biodiversity Instute
Skrede et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
68
Table 1. (Connued).
Species ID LSU hsp rpb2 Locality Coll. date Collector
Collector ID
H2881 (TRH-12612) MN598171 Spain, Balearic
Islands, Mallorca,
Formentor
01.01.1988 S. Sivertsen
H2882 (TRH-12625) MN598172 Spain, Balearic
Islands, Mallorca,
Reg. Escorca
24.10.1984 T. Schumacher
Epitype, H408 (C-F- Fungi.
Exs. Suec. 2062)
KY784504 KY772750 Sweden, Blekinge 5-14.10.
1946
S. Lundell, S.
Wickland
H. dalgeri H1924 (O-F-256527) MN598125 MN626782 Spain, Girona,
Serinya
04.02.2001 M. Perez de
Gregorio –
PG250217
H2984 (O-F-256528) MN598202 MN626814 Spain, Albacete,
Masegoso
02.05.2009 A. Rodriguez –
AR090502
H. danica H058 (O-253288) KY772942 KY784222 KY772469 Norway, Akershus 16.09.2009 T. Schumacher,
I. Skrede
Holotype, H263 (O-253288) KY773083 KY784378 KY772627 Denmark, Jutland 18.09.2008 J. Heilmann-
Clausen
H. elasca H066 (O-253311) KY772950 KY784230 KY772476 Sweden, Hallan 16.09.2009 G. Holden
H329 (C-F-51672) KY784433 KY772684 Russia, Sakha 21.08.1999 H. Knudsen
H443 (O-253310) KY773167 KY784531 KY772775 Norway, Akershus 00.00.2009 Ø. Stensrud
H. brosa H343 (C-F-92103) KY784446 KY772697 Iceland,
Merkurhraun
26.08.1984 S. Elborne
H357 (C-F-53774) KY773125 KY784458 KY772708 Norway, Nordland 11.09.1975 H. Dissing
H. fusca H2632 (O-F-256529) MN598159 MN626800 Spain, Rioja,
Ventas Blancas
09.04.2016 L. Ballester –
LB16040903
H304 (C-F-89381) KY784414 KY772663 Netherlands, Zuid-
Holland
14.05.1983 C. Bas
H305 (C-F-92122) KY773101 KY784414 KY772663 Hungary,
Kiskunsag Naonal
Park
11.05.1978 M. Babos, A.
Friesz
Epitype, H817 (S-F-124424) JX993065 MN598098 MN626765 Italy, Ex Herb.
Rehm In Trento
05.1888 G. Bresadola
H835 (S-F-023173) MN598099 MN626753 Spain, Valladolid,
La Cisterniga,
Fuentes de Duero
21.04.1991 J. C. Santos
H. fuscolacunosa H1020 (O-F-256530) MN644493 Spain, Huesca,
Banastas
08.12.2012 J. Garcia, L.
Ballester –
LB12120804
H1024 (O-F-256531) MN644494 MN598086 MN626772 Spain, Rioja,
Villarroya
15.05.2016 G. Munoz –
GM2930
Holotype, H2883 (TRH-
12618)
MN644495 MN598173 Spain, Balearic
Islands, Mallorca,
Formentor
07.04.1982 L. Sivertsen
H2889 (TRH-12623) MN644496 MN598179 Spain, Balearic
Islands, Mallorca,
Lluch
24.10.1984 T. Schumacher
H. griseoalba H306 (C-F-92112) KY784416 KY772665 USA, Michigan 09.06.1973 C. Nimke
H. helvellula H1928 (O-F-256532) MN598128 MN626785 Spain, Badajoz,
Campillo de
Llerena
11.03.2017 M. Plaza –
AH0170311-1
H278 (C-F-45506) KY773090 KY784393 KY772641 France 20.04.1977 R. Schumacher
© 2020 Westerdijk Fungal Biodiversity Instute
Dissingia and Helvella species from Spain
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
69
Table 1. (Connued).
Species ID LSU hsp rpb2 Locality Coll. date Collector
Collector ID
Epitype, H308 (C-F-92128) KY784418 KY772667 Spain, Canary
Islands, Hierro,
toward Taibique
07.01.1977 R. Korf, Fogel,
Hennebert, L.
Kohn
H309 (C-F-45469) KY773103 KY773103 France, Region des
Landes
01.03.1981 F. Candoussau
H521 (O-F-256533) MN598090 Spain, Rioja,
Villarroya
02.05.2010 G. Munoz –
GM1832
H523 (O-F-256534) MN598085 MN626760 Spain, Rioja,
Avellaneda
14.04.2007 R. Marnez –
RM321
H525 (O-F-256535) MN598093 Spain, Rioja,
Entrena
21.02.2015 R. Marnez,
L. Ballester –
LB15022101
H836 (S-F-023260) MN598100 MN626766 Spain, Valladolid,
La Parrilla, Monte
Bayón
28.03.1991 J. C. Santos
H. hispanica H1023 (O-F-256536) MN644504 MN598112 MN626771 Spain, Rioja, Santa
Lucia
06.04.2015 A. Melendez –
PIPE0833
Holotype, H1929 (O-F-
256537)
MN598129 MN626786 Spain, Rioja,
Ventosa
11.03.2017 L. Ballester –
LB17031103
H. hypocrateriformis H275 (C-F-57126) KY784390 KY772638 Switzerland,
Graubünden
07.09.1982 H. Dissing
Epitype, H301 (C-F-85205) KY784411 KY772660 Sweden, Uppland 21.07.1948 R. Molander
H. iberica H1016 (O-F-256538) MN644597 MN598108 MN626767 Spain, Rioja,
Tudelilla
03.04.2016 G. Munoz –
GM2907
Holotype, H1018 (O-F-
256539)
MN644498 MN598109 MN626769 Spain, Rioja, El
Rasillo
13.04.2013 R. Marnez –
RM2373
H. inexpectata H1017 (O-F-256540) MN598192 MN626768 Spain, Rioja,
Sojuela
05.01.2013 R. Marnez –
RM1088
Holotype, H2621 (O-F-
256541)
MN598148 MN626792 Spain, Rioja,
Sojuela
05.03.2016 A. Ezquerro –
AE160305
H. juniperi Holotype, H2973 (ex MCVE
20997 (no 501))
MN598194 MN626807 Italy, Lu, Viareggio 16.03.1996 G. Baiano,
M. Filippa, D.
Garofoli
Isotype, H2974 (ex MCVE
20998 (no 501/a))
MN598195 MN626808 Italy, Lu, Viareggio 16.03.1996 G. Baiano,
M. Filippa, D.
Garofoli
H415 (C-F-42193) KY784510 KY772754 Portugal, Minho, S
of S. Bartolomeu
do Mar, N of
Esposende
15.11.2000 P. Boisen
Hansen
H. lactea H262 (C-F Fungi. Exs. Suec.
1355)
KY784377 KY772626 Sweden, Uppland 25.07.1939 N. Hylander
H374 (C-F-39379) KY784473 KY772722 Denmark, Zealand 27.08.1961 H. Dissing
H. lacunosa H153 (O-285214) KY773002 KY784285 KY772536 Norway, Sør-
Trøndelag
10.06.2007 R. Haugan –
S07415
Epitype, H407 (C-F Fungi
Exs. Suec. 2065)
KY773152 KY784503 KY772749 Sweden, Femsjö.
Skatåker
04.10.1948 S. Lundell, G.
Haglund
H. leucophaea H2628 (O-F-256545) MN598155 MN626797 Spain, Huesca,
Banastás
08.12.2012 J. Garcia, L.
Ballester –
LB12120805
H. macropus H073 (O-253326) KY772954 KY784233 KY772480 Norway, Oslo 03.09.2009 M. F. M.
Bjorbaekmo
© 2020 Westerdijk Fungal Biodiversity Instute
Skrede et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
70
Table 1. (Connued).
Species ID LSU hsp rpb2 Locality Coll. date Collector
Collector ID
H1933 (O-F-256546) MN598133 Spain, Huesca,
Cerler
12.09.2015 C. Rojo, L.
Ballester –
LB15091205
H1948 (O-F-256547) MN598147 Spain, Rioja,
Lumberas
04.10.2008 G. Munoz –
GM1324
H257 (O-253325) KY784373 KY772622 Norway, Oppland,
Dovre
08.08.2009 T. Carlsen, T.
Schumacher, I.
Skrede – TS09-
36
H351 (C-F-56087) KY784452 KY772703 USA, Alaska,
Anchorage
14.08.1980 H. Dissing
H. maculata H303 (C-F-45400) KY784413 KY772662 USA, Alaska,
Eklutna Lake
24.08.1973 V. L. Wells, P.
E. Kempton
H. monachella H2631 (O-F-256548) MN598158 MN626799 Spain, Rioja,
Ventas Blancas
29.04.2015 R. Marnez,
L. Ballester –
LB15042904
Epitype, H268 (C-F-92121) KY784373 KY772622 Hungary, Szelid 14.05.1979 M. Babos,
L. Albert, J.
Bukei, A. Friesz
H269 (C-F-92120) KY773084 KY784384 KY772632 Hungary, Szelid 24.05.1965 I. Ferencz,
E. Toth, E.
Vessey, S.
Vasali
H843 (S-F-022128) MN598101 Spain, Valladolid,
Tudela de Duero,
Fuente del Rey
18.04.1990 J. C. Santos
H. neopallescens H1021 (O-F-256549) MN598110 Spain, Rioja,
Ventosa
18.01.2014 L. Ballester –
LB14011801
Holotype, H1022 (O-F-
256550)
MN644500 MN598111 Spain, Rioja,
Logrono
07.02.2015 R. Marnez –
RM2397
H1025 (O-F-256551) MN598113 MN626773 Spain, Bizkaia,
Gorliz-Urezarantza
18.01.2014 A. Melendez –
PIPE0753
H1026 (O-F-256552) MN598114 Spain, Rioja,
Sorzano
28.03.2013 L. Ballester –
LB13062402
H1032 (O-F-256553) MN598117 Spain, Rioja,
Villarroya
27.04.2006 G. Munoz –
GM495
H2639 (O-F-256554) MN598165 MN626806 Spain, Bizkaia,
Derio
03.10.2017 A. Melendez –
PIPE1063
H2884 (TRH-12607) MN644501 MN598174 Spain, Balearic
Islands, Mallorca,
Galilea
29.12.1997 S. Sivertsen
H2885 (TRH-12609) MN644502 MN598175 Spain, Balearic
Islands, Mallorca,
Formentor
01.01.1988 S. Sivertsen
H2886 (TRH-12610) MN598176 Spain, Balearic
Islands, Mallorca,
Lluch
28.12.1987 S. Sivertsen
H2887 (TRH-12611) MN598177 Spain, Balearic
Islands, Mallorca,
Puert de Canonge
29.12.1987 S. Sivertsen
H2888 (TRH-12613) MN598178 Spain, Balearic
Islands, Mallorca,
Puert de Canonge
29.12.1987 S. Sivertsen
© 2020 Westerdijk Fungal Biodiversity Instute
Dissingia and Helvella species from Spain
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
71
Table 1. (Connued).
Species ID LSU hsp rpb2 Locality Coll. date Collector
Collector ID
H2890 (TRH-12624) MN598180 Spain, Balearic
Islands, Mallorca,
Lluch
24.10.1984 W. Holm
H2891 (TRH-12627) MN598181 Spain, Balearic
Islands, Mallorca,
Reg. Escorca
24.10.1984 T. Schumacher
H2983 (O-F-256555) MN598201 MN626813 Spain, Rioja,
Clavijo
01.12.2018 R. Marnez –
RM2482
H401 (C-F-49767) MN598089 France, Avignon 22.10.1974 H. Dissing
H. nigra Epitype, H063 (O-253345) KY772947 KY772947 KY784227 Sweden, Hallan,
Halmstad
16.09.2009 G. Holden
H1029 (O-F-256556) MN644491 KY784573 KY772815 Spain, Rioja,
Castroviejo
09.08.2014 A. Melendez,
R. Marnez,
C. Perez –
PIPE0829
H1030 (O-F-256557) KY784572 KY772816 Spain, Bizkaia,
Urkiola
19.10.2007 G. Munoz –
GM1085
H2622 (O-F-256558) MN644492 MN598149 MN626793 Spain, Navarra,
Suarbe
07.10.2017 C. Echavarri –
CE17100701
H. pallescens H136 (O-220306) KY772987 KY784269 KY772519 Norway,
Hordaland, Ulvik
28.08.2001 G. Flatabø
Epitype, H138 (O-66205) KY772988 KY784271 KY772521 Norway, Møre og
Romsdal, Norddal
29.08.2003 D. Holtan
H2630 (O-F-256559) MN598157 MN626798 Spain, Rioja,
Sojuela
24.06.2013 R. Marnez,
L. Ballester –
LB13062407
H2633 (O-F-256560) MN598160 MN626801 Spain, Zaragoza,
Tarazona
01.11.2017 J. Genova, L.
Ballester –
LB17110103
H2638 (O-F-256561) MN598164 MN626805 Spain, Rioja,
Montemediano de
Cameros
23.06.2007 R. Marnez –
RM378
H. palustris H019 (O-253358) KY772914 KY784196 KY772440 Norway, Oppland,
Dovre
07.08.2009 T. Carlsen, T.
Schumacher, I.
Skrede
H043 (O-253359) KY772933 KY784214 KY772458 Norway, Oppland,
Dovre
07.08.2009 T. Carlsen, T.
Schumacher, I.
Skrede
H350 (C-F-55330) KY773123 KY784451 KY772702 Finland, Kuusamo 15.08.1978 H. Dissing
H. panormitana Epitype, H064 (0-253363) KY772948 KY784228 KY772474 Sweden, Hallan,
Laholm
16.09.2009 C. C. Mellberg
H145 (O-203499) KY772994 KY784278 KY772528 Norway, Møre og
Romsdal, Stordal
18.09.2005 P. Larsen
H2627 (O-F-256563) MN598154 MN626796 Spain, Huesca,
Villanúa
12.10.2007 L. Ballester –
LB07101202
H. paraphysitorquata Isotype, H271 (C-F-45305) KY773085 KY784386 KY772634 Spain, Teruel 03.06.1988 Arroyo, Calon
H. pezizoides Epitype, H061 (O-253366) KY772945 KY784225 KY772471 Sweden, Hallan,
Halmstad
16.09.2009 S. Lund
H2636 (O-F-256564) – – MN626756 Spain, Rioja,
Castroviejo
09.08.2014 R. Marnez –
RM2274
H431 (C-F-52986) KY773161 KY784525 KY772768 Denmark, N
Jutland
06.10.1997 M.
Christensen, K.
Hansen
© 2020 Westerdijk Fungal Biodiversity Instute
Skrede et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
72
Table 1. (Connued).
Species ID LSU hsp rpb2 Locality Coll. date Collector
Collector ID
H. phlebophora H273 (C-F-45405) KY773087 KY784388 KY772636 Iceland, Austur
Skaafellsysla
20.08.1981 H. F. Gøtzsche
– 84/31
H. phlebophoroides Holotype, H1031 (O-F-
256565)
MN644506 MN598116 MN626775 Spain, Rioja,
Anguiano
02.07.2011 R. Marnez –
RM2193
H1709 (C-F-39531) MN598087 MN626777 Denmark, Jutland,
Hansed Skov N
Horsens
22.08.1960 H. Dissing
H1846 (C-F-17373) MN644507 MN598088 MN626779 Denmark, NEZ,
København
31.07.1985 J. Vesterholt
H. platypodia H1792 (C-F-39481) MN598120 Denmark, Zealand,
Allindelille, N
Ringsted
24.08.1961 B. & M. Lange,
H. Dissing
H1795 (C-F-39482) MN598121 Denmark, Zealand,
Lellinge, E Køge
30.08.1961 L. Hansen,
M. Lange, H.
Dissing
H1796 (C-F-64490) MN598122 Denmark, EJ,
Tudsedamsmosen
at Varna
13.10.1981 S. A. Elborne
Epitype, H1939 (O-F-
256566)
MN598139 MN626790 Spain, Rioja,
Sorzano
11.06.2013 L. Ballester –
LB13061103
H. poculiformis Holotype, H1944 (O-F-
256567)
MN598143 MN626791 Spain, Zaragoza 10.11.2009 G. Munoz –
GM1702
H. pulla Epitype, H149 (O-253370) KY772623 KY784282 KY772532 Norway, Møre og
Romsdal
26.09.2008 T. Læssøe
H259 (O-253370) KY773080 KY784374 KY772623 Norway, Øsold 18.09.2013 K. Varenius
H2623 (O-F-256568) MN598150 MN626794 Spain, Rioja,
Montemediano de
Cameros
21.06.2008 F. Marnez –
FM574
H2629 (O-F-256569) MN598156 MN626758 Spain, Rioja,
Sorzano
11.06.2013 L. Ballester –
LB13061101
H2635 (O-F-256570) MN598162 MN626803 Spain, Rioja,
Castroviejo
02.08.2014 R. Marnez –
RM1161
H2977 (O-F-256571) MN598196 MN626757 Spain, Rioja,
Hornos de
Moncalvillo
19.12.2018 R. Marnez,
L. Ballester –
LB18121901
H. queleana H1716 (C-F-39372) MN598119 MN626778 Denmark, Zealand,
Gribskov
05.06.1973 P. M. Petersen,
H. Dissing
H2981 (O-F-256572) MN598200 MN626812 Spain, Rioja,
Canales de la
Sierra
20.06.2009 R. Marnez –
RM0749
Neotype, H403 (C-F-45303) KY773151 KY784499 KY772746 Denmark, Zealand,
Boserup Skov
29.04.1995 B. T. Olsen
H. renervis H1000 (O-F-256573) MN598107 Spain, Zamora,
Villalazan
24.04.2016 H. Perez –
2404-16HP1
Holotype, H1925 (ex
ARAN3008363)
MN598126 MN626783 Spain, Gipuzkoa,
San Sebasan
21.05.2009 J. M. Lekuona
H1926 (ex ARAN5037657) MN598193 MN626755 Spain, Navarra,
Huitzi-Basakaitz
24.06.2009 J. M. Lekuona
H778 (UPS-F-006175) MN598096 MN626763 Sweden, Skåne,
Ivö
09.08.1987 B. Hägg
© 2020 Westerdijk Fungal Biodiversity Instute
Dissingia and Helvella species from Spain
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
73
Table 1. (Connued).
Species ID LSU hsp rpb2 Locality Coll. date Collector
Collector ID
H779 (UPS-F-122183) MN598097 MN626764 Sweden, Närke,
Hallsberg
31.07.1981 S. Ryman
H. semiobruta H1922 (O-F-256574) MN598123 MN626780 Spain, Rioja,
Logrono, Pinar del
Gitano
11.03.2017 A. Ezquerro,
L. Ballester –
LB17031102
Isotype, H277 (C-F-45507) MN644499 KY784392 KY772640 France, Rhone,
D’Aubagne
15.12.1972 J. C. Donadini
H2901 (TRH-12614) MN598190 Spain, Balearic
Islands, Mallorca,
Galilea
27.12.1987 S. Sivertsen
H522 (O-F-256575) MN598091 MN626759 Spain , Rioja,
Ventosa
28.02.2015 R. Marnez,
L. Ballester –
RM2302
H524 (O-F-256576) MN598092 Spain, Rioja,
Lardero
30.01.2016 R. Marnez –
AC5335
H526 (O-F-256577) MN598094 MN626761 Spain, Rioja,
Lardero
21.03.2015 R. Marnez,
L. Ballester –
LB15032101
H529 (O-F-256578) MN598095 MN626762 Spain, Alava,
Laguardia
12.03.2016 A. Ezquerro –
AE16031201
H1366 (C-F-45419) MN598118 MN626776 Sweden, Gotland,
at Skansudd S of
Fröjel
28.05.1968 P. M. Petersen
H. solitaria H1923 (O-F-256579) MN598124 MN626781 Spain, Huesca,
Villanua
23.05.2016 J. Garcia, L.
Ballester –
LB16052302
H1937 (O-F-256580) MN598137 Spain, Rioja,
Ventas Blancas
29.04.2015 R. Marnez,
L. Ballester –
LB15042901
H1938 (O-F-256581) MN598138 Spain, Rioja,
Sorzano
11.06.2013 L. Ballester –
LB13061102
H1945 (O-F-256582) MN598144 Spain, Rioja,
Ventas Blancas
29.04.2007 G. Munoz –
GM1011
H1946 (O-F-256583) MN598145 Spain, Rioja,
Tudelilla
29.04.2012 G. Munoz –
GM2506
H2902 (TRH-12608) MN598191 Spain, Balearic
Islands, Mallorca,
Escorca, Pollenza
02.04.1985 L & S.
Sivertsen
H370 (C-F-45467) KY784470 KY772720 Spain, Valdepenas 06.07.1975 Lundell,
Morand
H. sublicia H1932 (O-F-256584) MN598132 MN626789 Spain, Rioja,
Castroviejo
09.07.2016 R. Marnez –
RM2390
H1947 (O-F-256585) MN598146 Spain, Asturias,
Covandonga
22.07.2011 G. Munoz –
GM2192
H2625 (O-F-256586) MN598152 Spain, Huesca,
Villanova
01.08.2006 R.Tena & L.
Ballester –
LB06080101
H2637 (O-F-256587) MN598163 MN626804 Spain, Rioja, Santa
Coloma
18.09.2014 R. Marnez –
RM2286
Epitype, H417 (C-F-39823) KY773155 KY784512 KY772756 Denmark, Zealand,
Sorø
02.10.1993 K. Hansen
H. sulcata H1027 (O-F-256588) MN644505 MN598115 MN626774 Spain, Rioja,
Clavijo
22.11.2013 R. Marnez –
RM2238
© 2020 Westerdijk Fungal Biodiversity Instute
Skrede et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
74
Table 1. (Connued).
Species ID LSU hsp rpb2 Locality Coll. date Collector
Collector ID
Epitype, H152 (O-69282) KY773001 KY784284 KY772535 Norway, Oslo 13.10.2006 O. Smith, A.-S.
Karlsson
H2979 (O-F-256589) MN598198 MN626810 Spain, Huesca,
Barbaruens
28.09.2018 R. Marnez,
L. Ballester –
LB18092809
H2980 (O-F-256590) MN598199 MN626811 Spain, Huesca,
Barbaruens
28.09.2018 F. Pancorbo,
L. Ballester –
LB18092802
H. terricola Holotype, H2978 (O-F-
256562)
MN598197 MN626809 Spain, Huesca,
Cerler
28.09.2018 F. Cervera, L.
Ballester –
LB18092810
spores from one individual were measured and the minimum –
mean – maximum values are reported.
DNA was extracted from all specimens using the E.Z.N.A.®HP
Fungal DNA Kit (Omega Biotek D3195), following the slightly
modied procedure from Skrede et al. (2017).
The three genec markers heat shock protein 90 (hsp), RNA
polymerase II second largest subunit (rpb2) and nuclear ribosomal
large subunit DNA (LSU) were amplied using PuReTaq Ready-To-
Go PCR Beads (GEhealthcare, Waukesha, WI), and puried with
ExoSAP-IT (GEhealthcare, Waukesha, WI). For primer sequences,
detailed PCR condions, and sequencing techniques, see the
procedure in Skrede et al. (2017). In addion to the sequences
produced in the present study, a representave selecon of
sequences from Helvella species from other geographic regions
were downloaded from GenBank and included for reference.
Informaon on specimens studied and all new and retrieved
sequences with accession numbers in GenBank is given in Table
1. The sequences were automacally aligned using the MUSCLE
v. 3.8.425 (Edgar 2004) plugin in Geneious Prime 2019.0.4
(Biomaers, Auckland). Alignments of each marker were
analysed individually by the Maximum Likelihood (ML) method
implemented in RAxML v. 8.2.11 (Stamatakis 2006), using the
GTRCAT approximaon. The best ML tree from the analysis of a
concatenated, paroned alignment of three genec markers is
presented in Fig. 1. In this concatenated dataset, we allowed for
the inclusion of some missing data by perming individuals with
only LSU, rpb2 and hsp, hsp and LSU or rpb2 and LSU. Bootstrap
analyses using 1 000 pseudoreplicaons were included in all
ML analyses. The concatenated alignment is available in Dryad
(hp://datadryad.org).
The ML analyses were the basis for species delimitaon.
A simplied genealogical concordance phylogenec species
recognion (Avise & Ball 1990, Taylor et al. 2000, Deman et al.
2003) was used, where terminal clades with bootstrap support
> 70 % from two or more genec markers were considered to be
species. Some intraspecic variaon was allowed.
RESULTS
Phylogeny
A total of 118 hsp, 69 rpb2 and 17 LSU sequences were
produced and submied to GenBank. The nal concatenated
alignment was 1 163 bp (279, 346, 538 bp for hsp, rpb2 and
LSU, respecvely). The ML phylogenec tree has high support
on most nodes toward the ps of the branches and is highly
discriminang among species (Fig. 1). Although the basal nodes
are mainly not supported, most of the main lineages and clades
in Skrede et al. (2017) were retrieved.
The resulng ML analyses demonstrated the presence of
27 Spanish species. Spanish representaves of four addional
species, i.e. H. calycina, H. crispa, H. macropus and H. pezizoides
were not included in the concatenated dataset, since only hsp
sequences were obtained (Table 1). Thus, 31 Spanish species
are included in the present survey. The sequenced specimens
from our samples are assigned to major lineages and clades
across the genus Helvella. Nine independent lineages are
dened as new species, i.e. H. phlebophoroides, H. hispanica,
H. neopallescens, H. terricola, H. fuscolacunosa, H. iberica, H.
inexpectata, H. renervis and H. poculiformis, of which seven are
nested in the /lacunosa clade (indicated as A in Fig. 1). The other
two new species are found elsewhere in the phylogeny, one as a
sister species to the /lacunosa clade (i.e. H. renervis) and one
as sister to H. sublicia (i.e. H. poculiformis).
Taxonomy
Dissingia leucomelaena (Pers.) K. Hansen & X.H. Wang,
Persoonia 42: 198. 2019. Fig. 2A.
Basionym: Peziza leucomelas Pers., Mycol. Eur. 1: 219. 1822.
Synonym: Helvella leucomelaena (Pers.) Nannf., in Lundell &
Nannfeldt, Fungi exs. Suec. Fasc. 19-20: 21, no. 952. 1941.
See Skrede et al. (2017) for addional synonyms.
Illustraon: Dissing (1966b: g. 8, as H. leucomelaena).
Apothecium spitate-cupulate, cup 2–8 cm broad, 1–5 cm
deep, with crenate margin and expanded into irregular lobes,
hymenium greyish brown to dark brownish black, oen with a
violaceus nge, receptacle surface pubescent, upper part whish
to dark greyish brown, below gradually paler to yellowish white.
Spe short, 0.5–2 cm high, 0.6–1 cm broad, with blunt irregular
ribs and grooves, with internal chambers. Medullary excipulum
thick, of dense textura intricata, hyphae 3–5 µm broad. Outer
excipulum of textura angularis, cells 10–24 µm, interspersed
with some narrow hyphae of prismac cells, outermost cells
in rows turning out perpendicular to receptacle surface giving
rise to tus of fascicled hyphae consisng of club-shaped cells.
Asci aporhynchous, 320–380 × 12–14 µm. Ascospores ellipsoid,
19.4–21.6–24.0 × 10.6–12.1–13.0 µm. Paraphyses brownish, 4
– 5 µm broad below, clavately enlarged to 7–11 µm at the ps.
© 2020 Westerdijk Fungal Biodiversity Instute
Dissingia and Helvella species from Spain
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
75
Specimens examined/sequenced: See Table 1.
Notes: This short-spitate deeply cupulate species was recently
re-combined in the new segregate genus Dissingia (Hansen et
al. 2019), a genus molecularly and morphologically segregated
from Helvella s. s. based on phylogenec evidence (paraphyly)
and asci that arise from simple septa. The oen semi-immersed
growth habit, with the whish spe buried in the soil,
characterises the species.
Helvella acetabulum (L.) Quel., Hymenomycetes, Fasc. Suppl.
(Alençon): 102. 1874. Fig. 2B, C.
Basionym: Peziza acetabulum L., Sp. Pl. 2: 1181. 1753; Fr., Syst.
Mycol. 2: 44. 1822
Synonyms: See Skrede et al. (2017).
Illustraon: Harmaja (1977b: gs 1–2).
Apothecium spitate-cupulate, 2.5–10 cm broad, by 3–8 cm
high, hymenium pale brown to medium brown, drying dark
brown, receptacle concolourous or pale yellowish brown, sub-
pubscent. Spe 1.5–7.5 cm high, 1.5–5 cm broad, broad above
and narrowing towards the base, hollow inside, outside whish
to yellowish, with 5 to 7 prominent, mostly double-edged sharp
ribs branching upwards to the aachment point about half
way to the apothecial margin. Medullary excipulum of textura
intricata, hyphae 4–8 µm broad, light brown-walled. Outer
excipulum of textura angularis to textura prismaca, individual
cells 10–25 µm diam, outermost cells prismac, hyaline, 5–15
× 4–8 µm, arranged in rows that turn out perpendicular to
receptacle surface. Asci pleurorhynchous, 310–380 × 13–16 µm.
Ascospores ellipsoid, uniguulate, 15.5–17.4–19.2 × 10.5–11.4
13.2 µm. Paraphyses 3–5 µm below, brownish along their whole
length, slightly enlarged at the clavate ps.
Specimens examined/sequenced: See Table 1.
Note: The Spanish collecons are morphologically and
molecularly idencal to specimens recorded from Northern and
Central Europe (cf. Skrede et al. 2017).
Helvella atra König in Oeder, Flora Danica 3 (9): 7. 1770. - nom.
sanct. Fig. 3C.
Synonym: Leptopodia atra (König in Oeder) Boud., Icon. Mycol.,
liste prélim.: 2. 1904.
See Skrede et al. (2017) for addional synonyms.
Misapplied name: Helvella atra Holmsk., Beata ruris oa Fungis
Danicis impensa 2: 47. 1799.
Illustraons: Oeder (1770: pl. 534, g. 1), Boudier (1906: no. 121,
pl. 228, as H. lacunosa, Filippa & Baiano (2011: gs 9–11, as H.
lacunosa).
Apothecium spitate-capitate, saddle-shaped or irregularly
lobed, 1–4 cm broad, 3–12 cm high, cap margin adnate and
aached to spe at 3 to 4 points, hymenium blackish with a dark
violet brown nge, even or wrinkled, receptacle surface greyish
brown to black, smooth, with inconspicuous ribs on outer
surface. Spe 0.5–2.5 cm broad, naked, greyish brown to black,
paler towards base, hollow and chambered inside, outside with
4 to 7 sharp and partly double-edged prominent ribs aached
to the reexed apothecial margin at 3 to 4 points, between ribs
with deep grooves and lacunae (‘holes’). Medullary excipulum
of textura intricata, hyphae 3–5 mm broad. Outer excipulum of
textura angularis, cells 8–15 µm in diam, hyaline, outermost cells
club-shaped, 10–30 × 5–15 µm. Asci pleurorhynchous, 260–10 ×
14–18 µm. Ascospores ellipsoid, 15.5–17.1–18.5 × 10–11.3–12.5
µm. Paraphyses subhyaline to light brown, 3–4 µm broad below,
increasingly brown towards the 6–8 µm broad clavate ps.
Specimens examined/sequenced: See Table 1.
Notes: An account of the nomenclatural history of this medium-
size greyish black species of the H. lacunosa morpho-species
complex was given by Filippa & Baiano (2011) and Skrede et
al. (2017). Helvella atra is apparently less common in Southern
compared to Northern Europe.
Helvella calycina Skrede et al., Persoonia 39: 221. 2017. Fig.
2D–F.
Basionym: Boletus calyciformis Baara, Fungorum Arimin.: 25.
1759; Fr., Syst. Mycol. 2: 45. 1822.
Synonyms: See Skrede et al. (2017).
Illustraon: Dissing (1966b: g. 16, as H. cosfera).
Apothecium short-spitate, at rst cupulate, then expanded
and irregularly folded, 2–7.5 cm broad, 1.5–4.5 cm high;
hymenium brownish, receptacle surface greyish brown,
pubescent to furfuraceous; spe with 5 to 8 prominent, blunt-
edged, yellowish-white ribs dichotomously branched when
extending onto receptacle surface, not reaching the margin in
fully developed specimens, receptacle with concentric, parallel
connecons between ribs, making the surface somewhat
wrinkled. Medullary excipulum of hyaline textura intricata,
hyphae 3–5 µm broad. Outer excipulum of textura prismaca
to textura angularis, outermost cells brown-walled. Asci
pleurorhynchous, cylindrical, 240–280 × 11–14 µm. Ascospores
ellipsoid, uniguulate, 14.5–16.2–17.6 × 9.5–10.8–12.0 µm.
Paraphyses 2.5–3.5 µm broad below, straight, septate, gradually
enlarged to 4–6 µm at the ps.
Specimens examined/sequenced: See Table 1.
Notes: The species was re-named and re-described in Skrede et al.
(2017), based on a taxon originally described from Italy as Boletus
calyciformis by Baara (1759). Helvella calycina seems common
throughout Europe, and was recently also reported from China
(Wang et al. 2019). The North European and Spanish specimens
are molecularly idencal and diverge genecally from its sister
species H. cosfera in one consistent hsp and two rpb2 SNPs.
Helvella crispa (Scop.) Fr., Syst. Mycol. 2: 14. 1822. Fig. 4B.
Basionym: Phallus crispus Scop., Fl. Carniol., ed. 2, 2: 475. 1772;
Fr., Syst. Mycol. 2: 14. 1822.
Synonym: Helvella pallida Schae., Fung. Bavar. Palat. Nasc. 4:
112. 1774.
See Skrede et al. (2017) for addional synonyms.
Illustraons: Boudier (1909b: no. 490, pl. 225), Dissing (1966b:
g. 21).
Apothecium spitate-capitate, 2–7 cm broad, irregularly bi- to
tri-lobate, saddle-shaped, with inrolled, free margin, hymenium
© 2020 Westerdijk Fungal Biodiversity Instute
Skrede et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
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76
D. leucomelaena (H1930) Spain
D. leucomelaena (H1931) Spain
H. solitaria (H370) Sweden
H. solitaria (H1923) Spain
H. maculata (H303) USA
H. leucophaea (H2628) Spain
H. leucophaea (H274) France
H. crispa (H408) Sweden
H. crispa (H2634) Spain
H. crispa (H2624) Spain
H. crispa (H135) Norway
H. acetabulum (H1927) Spain
H. acetabulum (H410) Sweden
H. griseoalba (H306) USA
H. dalgeri (H1924) Spain
H. dalgeri (H2984) Spain
H. costifera (H131) Norway
H. costifera (H247) Norway
H. calycina (H382) Denmark
H. calycina (H384) Denmark
H. calycina (H022) Norway
H. queletiana (H1716) Denmark
H. queletiana (H2981) Spain
H. queletiana (H403) Denmark
H. phlebophoroides (H1031) Spain
H. phlebophoroides (H1846) Denmark
H. phlebophoroides (H1709) Denmark
H. sulcata (H152) Norway
H. sulcata (H1027) Spain
H. sulcata (H2980) Spain
H. sulcata (H2979) Spain
H. atra (H1019) Spain
H. atra (H233) Georgia
H. atra (H406) Sweden
H. hispanica (H1929) Spain
H. hispanica (H1023) Spain
H. neopallescens (H1025) Spain
H. neopallescens (H2639) Spain
H. neopallescens (H2983) Spain
H. neopallescens (H2885) Spain
H. neopallescens (H1022) Spain
H. neopallescens (H2884) Spain
H. pallescens (H138) Norway
H. pallescens ( H136) Norway
H. pallescens (H2638) Spain
H. pallescens (H1726) Denmark
H. pallescens (H2630) Spain
H. pallescens (H2633) Spain
H. terricola (H2978) Spain
H. palustris (H019) Norway
H. palustris (H043) Norway
H. palustris (H350) Finland
H. fuscolacunosa (H1020) Spain
H. fuscolacunosa (H1024) Spain
H. fuscolacunosa (H2889) Spain
H. fuscolacunosa (H2883) Spain
H. fusca (H2632) Spain
H. fusca (H304) Netherlands
H. fusca (H305) Hungary
H. fusca (H817) Italy
H. lactea (H374) Denmark
H. lactea (H262) Sweden
H. iberica (H1018) Spain
H. iberica (H1016) Spain
H. juniperi (H415) Portugal
H. juniperi (H2974) Italy
H. juniperi (H2973) Italy
H. semiobruta (H1366) Sweden
H. semiobruta (H277) France
H. semiobruta (H1922) Spain
H. semiobruta (H522) Spain
H. semiobruta (H526) Spain
H. semiobruta (H529) Spain
H. helvellula (H523) France
H. helvellula (H308) Spain
H. helvellula (H1928) France
H. helvellula (H309) France
H. helvellula (H836) Spain
H. helvellula (H278) France
H. inexpectata (H1017) Spain
H. inexpectata (H2621) Spain
H. nigra (H063) Sweden
H. nigra (H2622) Spain
H. nigra (H1029) Spain
H. nigra (H1030) Spain
H. lacunosa (H153) Norway
H. lacunosa (H407) Sweden
H. plebophora (H273) Iceland
H. retinervis (H779) Sweden
H. retinervis (H1925) Spain
H. retinervis (H778) Sweden
0.02
H. hispanica
H. neopallescens
H. fuscolacunosa
H. iberica
H. inexpectata
H. retinervis
H. phlebophoroides
Holo-/Isotype
Epi-/Neotype
A
H. terricola
Fig. 1. Maximum likelihood tree of Spanish and other members of Helvella using parts of the RNA polymerase II second largest subunit (rpb2), heat
shock protein 90 (hsp) and nuclear ribosomal large subunit DNA (LSU). Branches supported by ML bootstrap analyses (>70) are thickened. Dissingia
leucomelaena is used as outgroup. Sequences from holo-/isotype (red) and epi-/neotype (green) are colour-coded. Green boxes indicate species
described in the present study. Sequences from Spain are in bold.
© 2020 Westerdijk Fungal Biodiversity Instute
Dissingia and Helvella species from Spain
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
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77
H. scyphoides (H2544) Norway
H. scyphoides (H140) Norway
H. platypodia (H1939) Spain
H. hypocrateriformis (H301) Sweden
H. hypocrateriformis (H275) Switzerland
H. macropus (H073) Norway
H. macropus (H257) Norway
H. macropus (H351) USA
H. fibrosa (H343) Iceland
H. fibrosa (H357) Norway
H. paraphysitorquata (H271) Spain
H. poculiformis (H1944) Spain
H. sublicia (H417) Denmark
H. sublicia (H2637) Spain
H. sublicia (H1932) Spain
H. elastica (H066) Sweden
H. elastica (H443) Norway
H. elastica (H329) Russia
H. danica (H263) Denmark
H. danica (H058) Norway
H. monachella (H268) Hungary
H. monachella (H269) Hungary
H. monachella (H2631) Spain
H. panormitana (H064) Norway
H. panormitana (H2627) Spain
H. panormitana (H145) Norway
H. pezizoides (H061) Sweden
H. pezizoides (H431) Denmark
H. fallax (H018) Norway
H. fallax (H339) France
H. pulla (H2635) Spain
H. pulla (H259) Norway
H. pulla (H149) Norway
H. pulla (H2623) Spain
H. poculiformis
Fig. 1. (Connued).
white to light greyish white, drying yellowish to brownish,
receptacle surface even, pubescent, pale to medium brown.
Spe prominent, 3–10 cm high, 1–3 cm broad, widest below,
pure white when fresh, with densely set irregular longitudinal
anastomosing ribs and dep grooves, hollow with longitudinal
chambers inside. Medullary excipulum of textura intricata,
hyphae 5–7 µm broad. Outer excipulum of textura angularis,
cells irregular in shape and size, 12–25 × 5–25 µm, outermost
cells prismac to club-shaped, arranged in fascicled rows forming
tus of hyphae, walls pale brown. Asci pleurorhynchous, 260–
310 × 12–16 µm. Ascospores ellipsoid, uniguulate, 17.8–19.0–
21.4 × 10.8–11.9–12.6 µm. Paraphyses 3–4 µm below, slightly
enlarged to 6–9 µm at the clavate ps.
Specimens examined/sequenced: See Table 1.
Notes: European specimens of H. crispa, including the Spanish
specimens, are morphologically uniform, but molecularly
variable, however, sll constung a single disnct species in
our phylogeny (cf. Fig. 1).
Helvella dalgeri Donadini, Bull. Soc. linn. Provence 35: 136. 1985
[1984]. Fig. 2G, H.
Basionym: Acetabula barlae Boud., Hist. Class. Discom. Eur.: 40.
1907.
non Helvella barlae Boud. & Pat., J. Bot., Paris 2: 445. 1888.
Illustraon: Boudier (1908b): no. 349, pl. 245 (as Acetabula
barlae).
Apothecium regularly short-spitate cupulate, cup 3–5 cm
broad, spe 0.5–1.5 cm high, 0.7–1.5 cm broad, hymenium black,
receptacle dark greyish black, with conspicuous, delicate white
ribs extending from the spe onto the receptacle; ribs on spe
and receptacle high and prominent, sharp-edged, branching,
aachment points of ribs in periphery at the mid part of the
receptacle surface. Medullary excipulum of compact textura
intricata, hyphae mulseptate, 4–7 µm broad. Outer excipulum
of textura angularis, cells 20–35 × 10–20 µm, outermost cells
cylindrical to drum-shaped forming dense rows of cells turning
out perpendicular to receptacle surface. Asci cylindrical,
pleurorhynchous, 280–340 × 14–18 µm, with a long, exuous,
spore-free lower part. Ascospores ellipsoid, 18.2–19.2–20.5 ×
9.4–10.1–11.4 µm, with one large internal guule. Paraphyses
3–4 µm broad below, enlarged to a clavate apical cell segment
20–30 µm long, by 5–7 µm broad.
Specimens examined/sequenced: See Table 1.
Notes: This relavely large, greyish black, short-spitate species
was described as Acetabula barlae by Boudier (1908b). Since the
earlier name Helvella barlae Boud. & Pat. (1888) exists, Donadini
(1985) proposed the new name H. dalgeri for this South European
species. The species was misinterpreted by Skrede et al. (2017)
who, based on Boudier’s plate and descripon (Boudier 1908b),
placed Acetabula barlae in synonymy with Helvella arctoalpina
Harmaja. The prominent white ribs on cup and spe, which
contrast with the dark-coloured receptacle surface, characterise
the gross morphology of the species. In our phylogeny (Fig. 1)
Helvella dalgeri is nested with H. griseoalba in a highly supported
clade sister to the /cosfera lineage in our phylogeny (cf. Fig. 1).
A hypogeous Helvella specimen from Masegoso (Albacete) was
shown molecularly idencal to H. dalgeri (cf. Fig. 2G and Table 1).
Helvella fusca Gillet, Champ. Fr., Discom. 1: 9. 1879. Fig. 5B.
Synonyms: See Skrede et al. (2017).
© 2020 Westerdijk Fungal Biodiversity Instute
Skrede et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
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78
Illustraons: Gillet (1879: pl. 4), Dissing (1966b: g. 23), Landeros
& Korf (2012: gs 1–4).
Apothecium spitate-capitate, 1.8–3.5 cm broad, 0.7–4.5 cm
high, irregularly bi- to tri- to tetralobate, saddle-shaped, with
adnate, partly fused margin, hymenium light to medium to dark
brown when fresh, drying brownish black, receptacle greyish
white to pale brown, sub-pubescent, with low, anastomosing
densely packed ribs on the receptacle surface. Spe white to
yellowish brown, 3–7 cm high, 1–2.5 cm broad, with longitudinal
Fig. 2. Photographs of apothecia in their natural habitat. A. Dissingia leucomelaena AEA0010 [H1930]. B. Helvella acetabulum LB13062401 [H1940].
C. H. acetabulum GM2507 [H1942]. D. H. calycina GM1504 [H848]. E. H. calycina RM2338 [H845]. F. H. calycina FB160514 [H849]. G. H. dalgeri
AR090502 [H2984]. H. H. dalgeri PG250217 [H1924]. I. H. renervis 2404-16HP1 [H1000]. J. H. renervis ARAN3008363 [H1925]. Scale bars = 1 cm.
All photographs taken by the collectors (confer Table 1).
© 2020 Westerdijk Fungal Biodiversity Instute
Dissingia and Helvella species from Spain
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
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79
anastomosing ribs and deep, partly lacunose furrows. Medullary
excipulum of loosely interwoven textura intricata, hyphae 3–8 µm
broad, hyaline. Outer excipulum of textura angularis to textura
prismaca, brownish, outermost cells in rows perpendicular to
receptacle surface, cells cylindrical to club-shaped, partly in few-
celled fascicles (tus). Asci pleurorhynchous, 260–320 × 14–16
µm. Ascospores uniguulate, 17.4–18.7–20.8 × 11.0–12.1–13.2
µm. Paraphyses 3–4 µm broad below, with yellowish brown
content, enlarged to 5–8 µm at the clavate ps.
Specimens examined/sequenced: See Table 1.
Fig. 3. Photographs of fresh apothecia and illustraons A. Authenc illustraon of Helvella sulcata deposited in the fungarium of Uppsala (UPS),
photograph taken by Åsa Kruys. B. H. sulcata RM2238 [H1027]. C. H. atra PIPE0966 [H1019]. D. H. queleana RM0749 [H2981]. E. H. phlebophoroides
RM2193 [H1031]. Scale bars = 1 cm. Photographs B, C, D and E are taken by the collectors (confer Table 1).
© 2020 Westerdijk Fungal Biodiversity Instute
Skrede et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
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80
Notes: Helvella fusca is a spring and early summer occurring
species characterised by a spitate saddle-shaped apothecium
with brown hymenium and a white, lacunose spe with
delicate anastomosing ribs extending onto the receptacle
surface. Helvella fusca occurs in associaon with Populus and
is likely to form ectomycorrhiza with this host. See Notes of H.
fuscolacuonsa for informaon on how to disnguish H. fusca
from the morphologically similar species H. fuscolacuosa.
Helvella fuscolacunosa Skrede & T. Schumach. sp. nov. Myco-
Bank MB834620. — Fig. 5A.
Etymology: Pertaining to the dark brown colour nged
with greyish and blackish (adj. fuscous) of the hymenium in
combinaon with the ribbed, lacunose spe.
Typus: Spain, Balearic Islands, Mallorca, Formentor, 7 Apr. 1982,
S. Sivertsen (holotype TRH-12618).
Apothecium spitate-capitate, 2.5–4.2 cm broad; irregularly
saddle-shaped, trilobate, deexed, that may mask its general
shape; margin involute and wavy. Hymenium grey-brown to
almost black, surface with ± bulges; receptacle pale grey, sub-
pubescent, occasionally with inconspicuous veins. Spe whish,
hollow, with internal chambers, up to 5 cm high and 2 cm broad,
sub-pubescent, with 5 to 8 partly anastomosing, sharp ribs and
deep grooves and a few lacunae between, ribs connuing onto
the receptacle surface. Medullary excipulum of densely packed
textura intricata, hyphae 3–7 µm broad, light brown-walled.
Outer excipulum of textura angularis with some interwoven
hyphae between, cells 7–30 × 5–20 µm, outermost cells
cylindrical to club-shaped in rows terminang in a subcapitate
up to 45 µm long end cell. Asci cylindrical, pleurorhynchous,
270–300 × 13–17 µm, ascospores oen transversely arranged
in young asci. Ascospores broadly ellipsoid, uniguulate, 15.6–
16.4–18.8 × 11.0–11.8–12.5 µm. Paraphyses 3–4 µm broad
below, straight, enlarged to 6–8 µm at the clavate ps.
Specimens examined/sequenced: See Table 1.
Notes: Helvella fuscolacunosa shares stature and hymenium
colour with H. fusca and H. neopallescens. However, while
the spe is interspersed with some lacunae (“holes”) in
H. fuscolacunosa and H. fusca, these are lacking in H.
neopallescens. Microanatomically, the ascospores in H.
fuscolacunosa are smaller and broader than in H. fusca and
H. neopallescens. Molecularly, H. fuscolacunosa deviates
consistently in four hsp, three rpb2 and 18 LSU substuons
from H. fusca, and in nine hsp, seven rpb2 and 25 LSU
substuons from H. neopallescens.
Fig. 4. Photographs of apothecia in their natural habitat. A. Helvella leucophaea LB121208 [H2628]. B. H. crispa LB07101201 [H2626]. C. H. nigra
GM1085 [H1030]. D. H. juniperi MCVE 20997 [H2973]. Scale bars = 1 cm. All photographs taken by the collectors (confer Table 1).
© 2020 Westerdijk Fungal Biodiversity Instute
Dissingia and Helvella species from Spain
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
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81
Helvella helvellula (Durieu) Dissing, Rev. Mycol. 31: 204. 1966.
Fig. 6E, F.
Basionym: Peziza helvellula Durieu, Expl. Sci. Algerie 1: t. 27, f.
11. 1849.
Synonyms: See Skrede et al. (2017).
Illustraons: Durieu (1849): t. 27, f. 11, as Peziza helvellula,
Boudier (1908a): no. 346, pl. 247, as Acetabula clypeata), Caro
et al. (2007: pp. 33, 35, 41).
Apothecium subsessile to short-spitate-cupulate, 1.2–2.5
cm broad; cupulate to irregular cupuliform to discoid, margin
undulate and gently involute, hymenium black, receptacle
surface furfuraceous, blackish, slightly sooty (fuliginous) towards
the spe. Spe short, solid, cylindrical to aened, 0.7–1.2 cm
high, by 0.3–0.7 cm broad, somewhat thickened at the base,
with 1 to 2 longitudinal grooves, at the top concolourous with
the receptacle, paler towards the base which is almost always
totally buried. Medullary excipulum of textura intricata, hyphae
3–5 µm broad. Outer excipulum of textura angularis, cells of
uneven size, 10–25 µm broad, interspersed with some inated,
larger cells 20–45 × 10–15 µm. terminang into cylindrical to
club-shaped cells towards the surface. Asci pleurorhynchous,
320–390 × 16–20 µm. Ascospores ellipsoid, obtuse, with one
large and several small internal guules, 21.8–24.0–27.3 ×
12.8–13.8–15.4 µm. Paraphyses 3–4 µm broad below, slightly
enlarged to 5–8 µm at the irregularly clavate ps.
Specimens examined/sequenced: See Table 1.
Notes: Helvella helvellula is a subsessile to short-spitate
black, cupulate species, described by Durieu (1849) based
on a specimen in Herb. C Montagne (PC). Dissing (1966a) did
not nd any original specimen of this Mediterranean species
in Montagne’s Herbarium in Paris (PC). Caro et al. (2007)
later succeeded in nding an original specimen from Morocco
collected by M. Roussel in 1836 and deposited in Montagne’s
herbarium in PC (see photograph on p. 41 in Caro et al.). This
specimen, which apparently is part of the original descripon
of Durieu’s species is probably too old to provide DNA for
sequencing, thus the epitype selected by Skrede et al. (2017)
from the Canary Islands is sll in place to provide necessary
informaon for a molecular idencaon of the species.
The Spanish collecons are from acid soil, at altudes
from 500 to 1 100 m. a.s.l., always associated with Cistus, i.e.
Cistus albidus, C. salviifolius and C. laurifolius. Short-spitate
specimens (see Fig. 6F) may be confused with H. semiobruta,
which occurs in the same habitat, but, the apothecia in H.
semiobruta are spitate-capitate, with a bi- to trilobate convex
cap and an elongate hollow spe.
Helvella hispanica Skrede & T. Schumach. sp. nov. MycoBank
MB834621. Fig. 7C, D.
Etymology: Referring to its place of origin (in Spanish language).
Typus: Spain, Ventosa, 11 Mar. 2017, L. Ballester LB17031103
(holotype O-F-256537).
Apothecium spitate-capitate, cap 2.5–4.5 cm broad, irregular
saddle-shaped, adnate, margin generally free but aached to
the spe at 2 to 4 points, hymenium dark greyish black to black,
even or wrinkled folded, receptacle paler, greyish, smooth,
with branching, blunt ribs. Spe cylindrical, hollow, 2–7 cm
high, 2–4.5 cm broad, whish to grey, with 3 to 7 longitudinal,
prominent, partly double-edged blunt ribs and deep grooves,
occasional with some elongate lacunae. Medullary excipulum of
dense textura intricata, hyphae 3–6 µm broad. Outer excipulum
of textura angularis to textura prismaca, brownish, angular
cells 10–20 µm interspersed with prismac cells in rows, 8–14
µm broad, turning out perpendicular to receptacle surface,
outermost cells club-shaped. Asci pleurorhynchous, 260–310
× 12–16 µm. Ascospores elongate-ellipsoid, 16.8–17.8–19.6 ×
8.2–9.5–11.2 µm, with one large internal oil drop. Paraphyses
straight, 3–5 µm below, indisnctly enlarged at the clavate ps.
Specimens examined/sequenced: See Table 1.
Notes: This new species is nested in the /atra-pallescens lineage,
as delineated by Skrede et al. (2017). It resembles H. lacunosa in
the ribbed, whish spe, the greyish-black to black hymenium,
and the greyish-black outer cap surface. The apothecial margin
Fig. 5. Photographs of apothecia in their natural habitat. A. Helvella fuscolacunosa LB12120804 [H1020]. B. H. fusca LB160409 [H2632]. Scale bars =
1 cm. Both photographs taken by the collectors (confer Table 1).
© 2020 Westerdijk Fungal Biodiversity Instute
Skrede et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
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82
is generally free from the spe. It genecally diverges from
its sister species H. atra in two hsp, one rpb2 and ve LSU
substuons (cf. Fig. 1).
Helvella iberica Skrede & T. Schumach. sp. nov. MycoBank
MB834622. Fig. 6A, B.
Etymology: Referring to its occurrence on the Iberian Peninsula.
Typus: Spain, El Rasillo, 13 Apr. 2013, R. Marnez RM2373
(holotype O-F-256539).
Apothecium spitate-capitate, cap 1.5–3.5 cm broad, irregularly
saddle-shaped with 2 to 4 lobes, adnate; margin free, hymenium
dark greyish black to black, even or wrinkled folded, receptacle
paler, light grey to greyish black, smooth, with branching blunt
ribs connuing from spe. Spe cylindrical, 2–4 cm high, 0.7–1.5
cm broad, whish to greyish, with deep, longitudinal, undulang
furrows and blunt ribs, with some ± lacunae between. Medullary
excipulum of loose textura intricata, hyphae 4–6 µm broad.
Outer excipulum of textura angularis, individual cells 12–24 ×
10–20 µm, light brown-walled, outermost cells club-shaped
to sub-capitate. Asci pleurorhynchous, 260–310 × 15–21 µm.
Ascospores elongate ellipsoid, 18.2–20.4–21.8 × 10.8–12.0–13.8
µm, with one large internal oil drop. Paraphyses straight, 4–6
µm below, slightly bent at ps, enlarged to 7–9 µm at the clavate
ps.
Specimens examined/sequenced: See Table 1.
Notes: Helvella iberica is nested in the /helvellula lineage,
as delineated by Skrede et al. (2017) (cf. Fig. 1). It resembles
H. juniperi in colour and stature (Filippa & Baiano 1999).
However, the free apothecial margin and ve hsp and one rpb2
substuons separate it from the laer. See also Fig. 4D and
‘Notes’ under H. inexpectata and H. semiobruta below.
Helvella inexpectata Skrede & T. Schumach. sp. nov. MycoBank
MB834623. Fig. 6G, H.
Etymology: Referring to the unexpected occurrence of yet
another species of the /helvellula lineage on the Iberian
Peninsula.
Typus: Spain, Sojuela (La Rioja), 5 Mar. 2016, A. Ezquerro
AE160305 (holotype O-F-256541).
Apothecium spitate-capitate, cap black to greyish black, 1.5–
4 cm broad, in young specimens bi- to trilobate, later more
irregular in shape, margin gently involute, deexed and adhered
to the spe, hymenium greyish black to black, receptacle surface
greyish, smooth to furfuraceous, covered by ny veins. Spe
cylindrical, 1.2–4 cm high, by 0.5–4.5 cm broad, hollow with a
few internal chambers, concolourous with receptacle or paler
above, whish below, with six to eight blunt, longitudinal ribs
and deep grooves. Medullary excipulum of dense textura
intricata, hyphae 2.8–5.5 µm broad, pale brown-walled. Outer
excipulum of densely packed rows of angular to club-shaped
cells, 10–30 × 7–15 µm, strongly constricted at septa, turning out
perpendicular to receptacle surface. Asci pleurorhynchous, 280–
320 × 14–18 µm. Ascospores oblong ellipsoidal, 18.4–19.6–21.4
× 10.6–11.7–12.8 µm, with one large and several small internal
guules. Paraphyses straight, 2.5–3.5 µm below, enlarged to
5–6.5 µm at the clavate ps.
Specimens examined/sequenced: See Table 1.
Notes: Helvella inexpectata has a convex cap and a short greyish
white spe with longitudinal blunt ribs and deep grooves
between. It is nested with H. helvellula, H. iberica, H. juniperi
and H. semiobruta in a highly divergent /helvellula lineage of the
/lacunosa clade (Fig. 1, cf. also Skrede et al. 2017). The species
resembles H. juniperi (Fig. 4D) and H. iberica (Fig. 6A, B) in shape
and colour, but has smaller and more oblong, narrow ascospores
than these ‘look alike’ species. The Spanish specimens of H.
iberica and the Italian holo- and isotype specimens of H. juniperi,
newly sequenced by us, dier in three and two hsp, and one and
two rpb2 substuons, respecvely.
Helvella leucophaea (Ba.) Pers., Obs. mycol. 2: 19. 1799. Fig.
4A.
Basionym: Boletus leucophaea Baara, Fungorum Arimin.: 25.
1759.
Synonym: Helvella pithyophila Boud., J. Bot. 1: 218. 1887.
Illustraons: Trania (1809: pl. 28, as H. leucophaea Pers.),
Boudier (1887: pl. 3; 1904b: no. 17, pl. 227, as H. pithyophila).
Apothecium spitate-capitate, apothecia irregular saddle-
shaped, 3–7 cm broad, hymenium pale olive brown, uneven
bulged (wrinkled), margin crenate, reexed, receptacle
sub-pubescent, ochraceous brown, with low, branching,
interconnected ribs. Spe cylindrical, 6–10 cm high, by 1.5–3.5
cm broad, grey to dark greyish brown, sub-pubescent, with deep,
longitudinal, irregular furrows and sharp-edged ribs, inside
of spe with uneven-sized longitudinal chambers. Medullary
excipulum of interwoven septate, brown, branching hyphae, 3–5
µm broad. Outer excipulum of textura angularis, cells mostly 10–
20 × 6–15 µm, outermost layer of cylindrical to club-shaped 3-
to 5-celled protruding hyphae, 10–15 µm broad. Asci cylindrical,
pleurorhynchous, 290–350 × 14–16 µm. Ascospores ellipsoid,
with one large and numerous small internal oil drops, 16.2–
18.8–20.4 × 10.2–10.8–12.0 µm. Paraphyses subhyaline, 3–4 µm
broad below, enlarged to 6–10 µm at the irregular clavate ps,
terminal cell thick-walled with incrusted, adhesive pigments.
Specimen examined/sequenced: See Table 1.
Notes: Helvella leucophaea resembles H. crispa in size and
stature. However, the hymenium and spe in fresh ascomata of
the former are greyish to greyish brown, drying caramel brown.
The two individuals from France and Spain are molecularly
idencal. Four consistent rpb2 and two hsp substuons
separate H. leucophaea and its sister species H. crispa of the /
crispa lineage (cf. Skrede et al. 2017).
Helvella macropus (Pers.) P. Karst., Not. Sällsk. F. Fl. Fenn. Förh.
11: 224. 1870. Fig. 8F.
Basionym: Peziza macropus Pers., Ann. Bot. (Usteri) 15: 26.
1795; Fr., Syst. Mycol. 2: 57. 1822.
Synonyms: See Skrede et al. (2017).
Illustraons: Dissing & Nannfeldt (1966: gs 4, 5a–e, pl. III),
Weber (1972: gs 23–24).
© 2020 Westerdijk Fungal Biodiversity Instute
Dissingia and Helvella species from Spain
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
83
Fig. 6. Photographs of apothecia in their natural habitat. A. Helvella iberica GM2907 [H1016]. B. H. iberica RM2373 [H1018]. C. H. semiobruta
RM2302 [H522]. D. H. semiobruta AC5335 [H524]. E. H. helvellula RM321 [H523]. F. H. helvellula LB15002101 [H525]. G. H. inexpectata RM1088
[H1017]. H. H. inexpectata AE160305 [H2621]. Scale bars = 1 cm. All photographs taken by the collectors (confer Table 1).
© 2020 Westerdijk Fungal Biodiversity Instute
Skrede et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
84
Apothecium spitate-cupulate, cup ± regular saucer-shaped,
1.5–3 cm broad, hymenium yellowish to greyish brown,
receptacle concolourous, villose. Spe terete, 2.5–12 cm high,
0.3–0.5 cm broad, pubescent. Medullary excipulum of loosely
interwoven textura intricata, hyphae generally 4–6 µm broad,
brownish, some hyphae with inated cells 10–20 µm broad,
constricted at septa. Ectal excipulum of textura intricata,
innermost cells 10–30 µm broad, turning into fascicled rows
of cylindrical cells 20–35 × 10–15 µm in outermost excipulum,
some hyphae extending into 50–200 µm long fascicled hyphoid
hairs. Asci pleurorhynchous, 240–300 × 13–16 µm. Ascospores
ellipsoid-fusoid, with one large and two to three medium
large internal guules, 19.5–23.4–25.8 × 9–10.5–12.0 µm.
Paraphyses 3–4 µm below, gradually enlarged to 5–9 µm at the
clavate ps.
Specimens examined/sequenced: See Table 1.
Notes: Our molecular survey of Helvella species in Europe
and the Nordic countries conrms the existence of ve or
six phylogenec species in the H. macropus morphospecies
complex (including H. brosa), of which H. macropus is the most
common species in Europe.
Fig. 7. Photographs of apothecia in their natural habitat. A. Helvella neopallescens LB14011801 [H1021]. B. H. neopallescens LB13062402 [H1026].
C. H. hispanica LB16022002 [H528]. D. H. hispanica PIPE-0833 [H1023]. E. H. terricola LB18092810 [H2978]. F. H. pallesens RM378 [H2638]. Scale bars
= 1 cm. All photographs taken by the collectors (confer Table 1).
© 2020 Westerdijk Fungal Biodiversity Instute
Dissingia and Helvella species from Spain
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
85
Helvella monachella (Scop.) Fr., Syst. Mycol. 2: 18. 1822. Fig. 8B.
Basionym: Phallus monacella Scop., Fl. Carniol., ed. 2, 2: 478.
1772; Fr., Syst. Mycol. 2: 18. 1822 (‘monachellaorth. mut.), non
Helvella monacella Schae., Fung. Bavar. Palat. Nasc. 4: 106.
1774 (‘Elvela’), nom. inval. (= Helvella sulcata Afzel.).
Synonyms: Helvella albipes Fuckel, Jahrb. Nassau. Vereins
Naturk. 23–24: 334. 1870.
Helvella leucopus Pers., Mycol. Eur. 1: 213. 1822.
Illustraons: Fuckel (1870: pl.5, g. 2), Dissing (1966b: g. 38, as
H. leucopus), Moravec (1980: pl. 4, as H. leucopus), Van Vooren
(2010: 43, photo 12).
Apothecium spitate-capitate, cap 1.5–4 cm broad, saddle-
shaped or irregularly lobate, deexed, margin occasionally
reexed, free, hymenium dark brown to brownish black,
receptacle smooth, whish to pale greyish. Spe terete, hollow,
whish, 3–7 cm high, 1–2 cm broad, oen with irregular groves.
Medullary excipulum of textura intricata, hyphae 3–6 µm broad,
some hyphae heavily pigmented, blackish. Outer excipulum of
textura angularis, cells cylindrical to club-shaped, 15–40 × 6–20
µm, arranged in rows turning out perpendicular to receptacle
surface. Asci pleurorhynchous, 260–310 × 15–18 µm. Ascospores
ellipsoid, 20.6–22.0–23.8 × 12.7–14.1–15.2 µm, with one large
internal oil drop. Paraphyses straight, 3–5 µm below, enlarged to
6–8 µm broad at the clavate ps.
Specimens examined/sequenced: See Table 1.
Note: The sequenced specimens from Spain have hsp and rpb2
sequences idencal to the rest of the sequenced specimens
from Europe.
Helvella neopallescens Skrede & T. Schumach. sp. nov. Myco-
Bank MB834625. Fig. 7A, B.
Etymology: Referring to it being genecally similar to H.
pallescens.
Typus: Spain, Rioja, Logrono, La Grajera, 7 Feb. 2015, R. Marnez
RM2397 (holotype O-F-256550).
Apothecium spitate-capitate, cap 2.5–4.5 cm broad, irregularly
saddle-shaped, margin reexed but aached to spe at 3 to
4 points, hymenium dark brownish to greyish black, even,
receptacle paler, greyish, smooth, with ± anastomosing, delicate
veins on the surface. Spe whish to greyish to dark grey, highly
irregular in shape and length, ± cylindrical, 2–6 cm high, 0.8–
2.5 cm broad, with 5–7 longitudinal blunt-edged ribs, hollow,
with large chambers inside. Medullary excipulum of densely
interwoven textura intricata, hyphae 3–5 µm broad. Outer
excipulum of irregular textura angularis, cells relavely broad,
15–35 × 12–25 µm, outermost cells club-shaped, short-celled.
Asci cylindrical, pleurorhynchous, 260–310 × 12.5–15 µm.
Ascospores obtuse ellipsoidal, 16.1–16.8–18.2 × 9.2–9.6–11.4
µm, with one large internal oil drop. Paraphyses 3–5 µm below,
enlarged to 7–8 µm broad at the clavate to spadiceiform ps.
Specimens examined/sequenced: See Table 1.
Notes: Helvella neopallescens is nested in the /atra-pallescens
lineage, as delimited by Skrede et al. (2017). Morphologically,
it resembles H. hispanica, H. fuscolacunosa, and H. lacunosa by
the hollow, ribbed, and partly lacunose, whish spe and the
contrasng brown to blackish cap, but is genecally disnct
(cf. Fig. 1). It deviates from its morphologically dissimilar
sister species H. pallescens in two hsp, one rpb2 and four LSU
substuons.
Helvella nigra Berg., Phytonom. univ. 1: t. 147. 1783. Fig. 4C.
non Helvella nigra Peck, Bull. Torrey Bot. Club 26: 70. 1877, nom.
illeg. (homonym).
See Skrede et al. (2017) for synonyms and typicaon.
Illustraon: Bergeret (1783: pl. 147).
Apothecium spitate-capitate, 2–6 cm across, cap saddle-shaped,
bi- to trilobate, from 6 to 15 cm in height, apothecial margin
adnate, rmly aached to the spe at 4 to 6 points; hymenium
wrinkled, dark grey to greyish black when fresh, drying darker,
receptacle concolourous, smooth. Spe concolourous or slightly
paler, lacunose, receptacle surface concolourous or slightly
paler, smooth, with disnct, branching ribs. Spe 2.0–4.5 cm
broad, naked, ribbed, ash grey to almost black, longitudinal ribs
high and prominent, sharp, partly anastomosing and double-
edged, with deep ‘holes’ (lacunae) between ribs, spe hollow
and chambered inside. Medullary excipulum of textura intricata,
hyphae 3–5 µm broad. Outer excipulum thin, of intermixed
broad-celled textura intricata and textura globulosa-angularis,
the layer light brown, outermost cells club-shaped, hyaline,
10–30 × 5–15 µm, with loose outgrowth of few cells colouring
intensely in coon blue. Asci pleurorhynchous, 290–330 × 14–
16 µm. Ascospores ellipsoid, uniguulate, 16.5–17.8–19.0 ×
10.4–11.1–12 µm. Paraphyses straight, slightly bent at ps, light
brown in the whole length, 3–4 µm broad below, enlarged to
6–8 µm at the ps.
Specimens examined/sequenced: See Table 1.
Notes: Helvella nigra is characterised by greyish black ashen
colours, a tall, slender stature, a ribbed, lacunose spe, and a
reexed and lobed cap with prominent ribs on the receptacle
surface. The apically bent paraphyses (like a “golf-club”) are
characterisc under the lens. Eight hsp, three rpb2 and 25 LSU
characters disnguish H. nigra and H. lacunosa.
Helvella pallescens Schae., Fungi Bavar. Palat. Nasc. 4: 114.
1774 (‘Elvela’). Fig. 7F.
Synonym: Helvella sulcata β (var.) pallescens (Schae.) Fr., Syst.
Mycol. 2: 16. 1822.
Illustraon: Schaeer (1774: pl. 322).
Apothecium bilobate to campanulate to irregularly saddle-
shaped, cap 0.6–2 cm broad, by 2–3 cm high, underside
glabrous, edge of cap adnate (reexed) and fused with the spe
at 3 to 4 points, hymenium white to pale greyish drying brownish
yellow; receptacle surface and spe white to pale yellowish to
brown, drying brown. Spe long and slender, 0.5–1.5 cm broad,
4–6 cm high, hollow, with four to ve longitudinal ribs and a few
lacunae between. Medullary excipulum of an interwoven textura
intricata, hyphae of variable thickness, 4–10 µm broad. Outer
excipulum of textura globulosa to textura angularis, cells 20–40
µm diam, the outermost cells more elongated and prismac
© 2020 Westerdijk Fungal Biodiversity Instute
Skrede et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
86
giving rise to a cover of densely packed club-shaped cells on
receptacle surface. Asci pleurorhynchous, 280–310 × 14–15
µm, young asci with numerous cyanophilic ascospores with
large pustules adhering to the spore wall. Ascospores broadly
ellipsoid, 15.0–17.0–18.2 × 10.2–11.8–13.2 µm. Paraphyses
hyaline, 2–3 µm broad, septate, gradually enlarged into irregular
clavate ps, 6–10 µm broad.
Specimens examined/sequenced: See Table 1.
Notes: Helvella pallescens is a slender, whish to greyish species
that has been confused with H. lactea, with which it shares
similar colours and microanatomy. Eight rpb2 and nine hsp
substuons separate H. pallescens and H. lactea. The species
is nested in the /lacunosa clade (Fig. 1).
Helvella panormitana Inzenga, Funghi siciliani 1: 41. 1865. Fig.
8A.
Illustraons: Inzenga (1865: pl. 4. f.11–4).
Apothecium spitate-capitate, cap campanulate to bilobate,
1–2.5 cm broad, 1.5–3 cm high, long-spitate, margin free,
reexed towards spe, hymenium pale greyish brown,
receptacle smooth, white or pale yellow, concolourous with
spe, drying brownish orange. Spe 0.4–1 cm broad, by 3–15
cm high, solid to hollow, occasionally with grooves towards the
base. Subhymenium and medullary excipulum of hyphae, 3–4
µm broad, embedded in an amorphous gelanous matrix. Outer
excipulum a mixture of textura intricata and textura angularis,
hyphae up to 8 µm broad, angular cells generally 25–40 µm in
diam., partly inated to 60–70 µm diam, outermost cells giving
rise to densely packed rows of prismac to club-shaped cells
10–20 µm broad, 10–30 µm long, outermost cell club-shaped
and heavily staining in CB. Asci pleurorhynchous, 290–310 ×
14–16 µm. Ascospores ellipsoid, 19–22.2–24 × 11.5–12.7–13.5
µm. Paraphyses ca. 3 µm broad below, septate, light brown,
gradually enlarged to 6–8 µm at the clavate ps.
Specimens examined/sequenced: See Table 1.
Notes: Helvella panormitana was originally described from
Sicily, Italy (Inzenga 1865). The species deviates from the
morphologically similar H. elasca in 5 hsp, 11 rpb2 and 10 LSU
substuons. The species is widely distributed in Europe (cf.
Skrede et al. 2017).
Helvella pezizoides Afzel., Kongl. Vetensk. Acad. Nya Handl. 4:
308. 1783. - nom. sanct. Fig. 9A, B, D, F.
Synonym: Leptopodia pezizoides (Afzel.) Boud., Icon. Mycol.,
liste prelim.: 2. 1904.
Illustraons: Afzelius (1783: pl. 10, f. 2), Dissing (1966b: 117).
Apothecium involute, spitate, 0.5–1.5 cm broad, with edge
permanently deexed, hymenium black, receptacle surface
pubescent, ash grey. Spe brownish black, terete, pubescent,
1–2 cm high, 0.4–0.7 cm broad. Medullary excipulum of loose
textura intricata, hyphae hyaline, 4–7 µm broad, interspersed
with some disnct brownish black hyphae. Outer excipulum
of textura angularis, cells thick-walled, brownish, turning out
in dense rows perpendicular to receptacle surface, outermost
cells irregularly club-shaped, constricted at septa, forming two-
three-celled brownish dense individual hyphae to 50 µm long
by 15 µm broad. Asci pleurorhynchous, 260–300 × 12–15 µm.
Ascospores ellipsoid, 14.5–15.3–16.5 × 8.5–9.0–9.6 µm, with
one large guule in rehydrated material. Paraphyses straight,
3.5–4.5 µm broad below, gradually enlarged to 5.5–7 µm at the
clavate ps.
Specimens examined/sequenced: See Table 1.
Notes: Morphologically, H. pezizoides resembles H. fallax and H.
pulla in colour, but the pubescent receptacle and spe separate
it from the two laer species. In an ongoing study of Helvella in
the Nordic countries, we discovered an authenc collecon of
H. pezizoides in the fungarium of Stockholm (S), and a colour
plate of H. pezizoides in the fungarium of Uppsala (UPS) both
examined and prepared by Adam Afzelius himself. The original
specimen in S is three apothecia of Afzelius’ original material
mounted and glued to a sheet and led under the name of
Peziza (spitata), a name which was also referred to by Afzelius
in his original descripon of H. pezizoides (cf. Afzelius 1783, p.
308). The colour plate in UPS is a drawing of this fungarium
sheet (now in S), which was also reproduced in an inverted
version as a gure in the original work by Afzelius (1783, tab X,
g. 2). This gure was selected as a lectotype (iconotype) of H.
pezizoides by Dissing (1966b). The original collecon of Afzelius
was tentavely referred to H. villosa [= H. brosa (Wallr.) Korf]
by Dissing (1966b: 69).
Helvella phlebophoroides Skrede & T. Schumach. sp. nov. Myco-
Bank MB834601. Fig. 3E.
Etymology: Pertaining to the prominent ribbed (phlebus =
veined) outer surface of the receptacle similar to that observed
in H. phlebophora.
Typus: Spain, Rioja, Anguiano, 2 Jul. 2011, R. Marnez RM2193
(holotype O-F-256565).
Apothecium spitate-capitate, saddle-shaped to plane, cap 1.5–
3.5 cm broad, aached to spe at two to three points, when fully
developed with a deexed and free margin, hymenium greyish
black, surface even or wrinkled, receptacle smooth, pale grey to
whish. Spe grey to greyish white, 3–6 cm high, 1–1.5 cm broad,
with 4 to 7 partly double-edged, blunt longitudinal ribs and deep
furrows with a few lacunae between, ribs extending onto the
receptacle forming delicate, radial ribs and veins (cf. Fig. 3E).
Medullary excipulum of dense textura intricata, hyphae 4–6.5
µm broad. Outer excipulum of a mixture of textura angularis/
intricata, the outermost cells bound in loosely interconnected
rows of cells, brown-walled, 10–20 × 5–12 µm, turning out
perpendicular to receptacle surface, outermost cell club-shaped.
Asci cylindrical with a lengthy narrow sterile poron below,
pleurorhynchous, 280–320 × 12–15 µm. Ascospores ellipsoid,
with one large oil drop when fully mature, 15.2–16.6–17.8 ×
9.0–9.6–10.4 µm. Paraphyses straight, 4–5 µm broad, enlarged
to 6–8 µm at the ps.
Specimens examined/sequenced: See Table 1.
Notes: Helvella phlebophoroides is sister species to H. sulcata
in our phylogeny (cf. Fig. 1). Four hsp and three rpb2 characters
© 2020 Westerdijk Fungal Biodiversity Instute
Dissingia and Helvella species from Spain
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
87
separate the two species. The species occurs in Denmark as
well (Table 1). Helvella phlebophoroides may be confused
with H. phlebophora and H. queleana, due to the prominent
radiang ribs of the outer receptacle surface. However, H.
Fig. 8. Photographs of apothecia in their natural habitat. A. Helvella panormitana LB07101202 [H2627]. B. H. monachella LB150429 [H2631]. C. H.
poculiformis GM1702 [H1944]. D. H. sublicia RM22 [H2637]. E. H. sublicia RM2286 [H1932]. F. H. macropus LB15091205 [H1933]. G. H. platypodia
LB13061103 [H1939]. H. H. solitaria GM1011 [H1945]. I. H. solitaria GM2506 [H1946]. Scale bars = 1 cm. All photographs taken by the collectors
(confer Table 1).
© 2020 Westerdijk Fungal Biodiversity Instute
Skrede et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
88
phlebophoroides, like H. queleana has fewer and more
distantly set, irregularly radiang ribs on the receptacle surface
compared to H. phlebophora. These three species all form
disnct phylogenec groups. Helvella phlebophoroides and
H. queleana is separated by nine hsp, 10 rpb2 and 19 LSU
substuons, and H. phlebophoroides and H. phlebophora by 10
hsp, 14 rpb2 and 20 LSU substuons.
Helvella platypodia (Boud.) Donadini, Bull. Soc. linn. Provence
35: 136. 1985. [1984]. Fig. 8G.
Basionym: Cyathipodia platypodia Boud., Hist. Class. Discom.
Eu r.: 39. 1907.
Typus: Lectotype designated here (MycoBank MBT390951):
Boudier, Icones Mycol. Livr. 22: no. 467, pl. 241, as C. platypodia
1909a. Epitype designated here (MycoBank MBT390950): Spain,
Rioja, Sorzano, under Corylus avellana, 11 Jun. 2013, L. Ballester
LB13061103 (epitype O-F-256566).
Illustraon: Boudier (1909a: no. 467, pl. 241, as C. platypodia).
Apothecium cupulate-spitate, small to medium-sized, 1.5–3.5
cm broad, by 1–2.5 cm high, regular cupulate to saucer-shaped
and recurved when fully mature, hymenium and receptacle
greyish to olive grey, concolourous, receptacle and spe
pubescent, drying rusty brown, hymenium drying dark brown.
Spe whish to yellowish, compressed (aened - ‘platypodia’),
solid, to 3.5 cm high, 0.5–1.5 cm broad, with 2 to 3 longitudinal
grooves. Medullary excipulum of textura intricata, hyphae
3–6 µm broad, septate. Outer excipulum of textura angularis,
cells elongate club-shaped in rows, 20–30 × 10–15 µm, cells
in periphery constricted at septa, outermost cells protruding
into scaered loose tus of fascicled, mulseptate hyphae
(hyphoid hairs), 40–150 µm long, by 10–25 µm broad. Asci
pleurorhynchous, 270–310 × 13–15 µm. Ascospores ellipsoid,
16.5–18.1–19.0 × 11.0–11.9–12.5 µm, with one large internal oil
drop and some smaller guules at the poles. Paraphyses 2–3.5
µm broad, not much enlarged to 5–6 µm at the ps.
Specimens examined/sequenced: See Table 1.
Notes: We have resurrected the name H. platypodia for this
species, lectotypied by Boudier’s plate (iconotype), and
supported with an epitype from Spain, which contributes paral
hsp and rpb2 sequences to the species. Helvella platypodia
belongs in the /hypocrateriformis lineage, as delineated by
Skrede et al. (2017).
Helvella platypodia is a small to medium-sized grey to olive
grey species with a contrasng white, compressed, furrowed
spe. The ascoma turn rusty brown when dried, matching the
colours and descripon (in dry condion) in Boudier (1909a),
and then resembling miniature ascomata of Helvella solitaria
(see this species below). The Spanish collecon is on acid
soil in associaon with Corylus avellana and Populus nigra.
The species occurs in Denmark as well (cf. Table 1). Based on
gross morphology in dried condion, H. platypodia has been
intermixed with the genecally unrelated H. solitaria (= H.
quelei) of the /acetabulum - solitaria lineage with less greyish
but more brown-coloured hymenium and more elongate, larger
ascospores.
Helvella poculiformis Skrede & T. Schumach. sp. nov. MycoBank
MB834627. Fig. 8C.
Etymology: Pertaining to the cup-shaped apothecia in this
species.
Typus: Spain, Zaragoza, 10 Nov. 2009, G. Munoz GM1702
(holotype O-F-256567).
Apothecium spitate-cupulate, 1–2.5 cm broad, by 1.5–3.5
cm high, regular cupulate to saucer-shaped, hymenium and
receptacle dark grey to greyish, concolourous, receptacle drying
rusty brown, hymenium dark brown. Spe terete, cylindrical,
whish to greyish white, 0.3–0.5 cm broad. Medullary excipulum
of textura intricata, hyphae brownish, 3–4 µm broad. Outer
excipulum of textura angularis, cells elongate club-shaped of
even size, 15–20 × 8–15 µm, outermost cells extending into
scaered mul-celled hyphae (hyphoid hairs), 30–80 µm long,
7–15 µm broad. Asci pleurorhynchous, 240–280 × 11.5–15 µm,
ascospores ± obliquely arranged in asci. Ascospores ellipsoid,
slightly narrowing towards the poles, 18.5–19.2–20.5 × 9.2–
10.0–11.0 µm, with one large and several minor internal guules
at the poles. Paraphyses 2.5–3.5 µm broad, enlarged to 6–9 µm
at the slightly spadiceiform, clavate ps.
Specimen examined/sequenced: See Table 1.
Notes: Helvella poculiformis resembles H. sublicia in shape
and colour and is nested in the /rivularis – sublicia lineage of
Skrede et al. (2017). The ascoma is regularly cupulate to saucer-
shaped, with a dark greyish black cup and a contrasng whish
to greyish, terete spe. Helvella poculiformis diers from H.
sublicia in three hsp and four rpb2 substuons.
Helvella pulla Holmsk., Beata ruris oa Fung. Dan. 2: 49. 1799.
- nom. sanct. Fig. 9C, E.
Synonym: Helvella fuliginosa Schae., Fung. Bavar. Palat. Nasc.
4: 113. 1774.
Illustraons: Holmskjold (1799: pl. 26), Schaeer (1774: pl. 320,
as Helvella fuliginosa).
Apothecium spitate-capitate, cap 0.5–1.5 cm broad, 0.5–1.5 cm
high saddle-shaped, bilobate and involute (reexed) throughout
development, hymenium pale grey to medium grey, receptacle
concolourous, subpubescent. Spe grey, terete or slightly
compressed, sub-pubescent, greyish, 3–6 cm long, 0.3–0.5 cm
broad, rmly aached to deeply decayed wood. Medullary
excipulum of loosely interwoven textura intricata, hyphae ca.
2.5–4.5 µm broad, subhyaline, intermixed with some broader,
brown-walled hyphae. Outer excipulum of textura prismaca,
hyphae light brown-walled, 5–15 µm broad, arranged in rows
turning out perpendicular to receptacle surface, outermost cells
club-shaped, 15–30 × 10–20 µm. Asci pleurorhynchous, 275–310
× 13–15 µm. Ascospores ellipsoid, 17.6–18.3–19.0 × 10.6–11.4
12.0 µm. Paraphyses subhyaline, 3–4 µm broad below, gradually
enlarged to 8–12 µm at the ps.
Specimens examined/sequenced: See Table 1.
Notes: This grey to blackish species of the /fallax - pezizoides
lineage may resemble H. poculiformis and H. sublicia (of the
© 2020 Westerdijk Fungal Biodiversity Instute
Dissingia and Helvella species from Spain
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
89
/ rivularis - sublicia lineage) in colour, however, both ascoma
ontogeny (permanently saddle shaped and bilobate in H. pulla
versus cupulate to applanate to compressed in H. poculiformis
and H. sublicia) and microanatomy (ascospore mean values 18.3
× 11.4 µm in H. pulla versus 19.2 × 10.0 µm in H. poculiformis
and 20.1 × 12.5 µm in H. sublicia) as well as molecular data easily
discriminate among the three.
Fig. 9. Photographs of apothecia, an illustraon and a fungarium specimen. A. Helvella pezizoides RM2274 [H2636]. B. Authenc plate of H. pezizoides
deposited in the fungarium of Uppsala (UPS). C. H. pulla LB13061101 [H2629]. D. Original specimen of H. pezizoides, glued to cardboard in the
fungarium of Stockholm (S). E. H. pulla RM1161 [H2635]. F. The envelope of the original material of H. pezizoides, annotated Peziza spitata. Scale
bars = 1 cm. Photographs A, C and E are taken by the collectors (confer Table 1) and photographs B, D and F by Åsa Kruys.
© 2020 Westerdijk Fungal Biodiversity Instute
Skrede et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
90
Helvella queleana Sacc. & Trav., in Sacc., Syll. Fung. 19: 850.
1910. Fig. 3D.
Synonyms: See Skrede et al. (2017).
Illustraon: ? Weber (1972: g. 34, as H. phlebophora?).
Apothecium spitate-capitate, cap hemispheric, convex to
aened, 2–3.5 cm broad, 0.5–1.5 cm high, margin free to
deexed toward spe, hymenium grey to light brownish grey,
receptacle light grey to cream, drying darker, subtomentose,
with conspicuous, branched, blunt ribs. Spe 2.5–5 cm long, 0.8–
1.5 cm broad, pubescent, paler or concolourous with receptacle,
inside chambered, outside with 3 to 5 double-edged, sharp ribs
connuing onto the receptacle surface. Medullary excipulum of
textura intricata, hyphae septate, branching, hyaline, 4–10 µm
broad. Outer excipulum of angular to club-shaped cells arranged
in a palisade towards the surface, individual cells 15–35 × 5–20
µm, hyaline to brown-walled, outermost cells in rows forming
hyaline hyphoid short-celled hairs. Asci pleurorhynchous, 270–
320 × 12–17 µm. Ascospores ellipsoid, 17.0–17.8–19.2 × 11.2–
12.3–13.5 µm. Paraphyses 3–4 µm broad below, subhyaline to
pale brown-walled, at ps enlarged to 5–7 µm broad.
Specimens examined/sequenced: See Table 1.
Notes: Helvella queleana is nested in an unsupported single-
species lineage of the /lacunosa clade (clade B), as delineated
by Skrede et al. (2017), a posion retained in the present
phylogeny (cf. Fig. 1). The photograph and descripon of
Helvella phlebophora in Weber (1972, g. 34) is suggesve of
H. queleana.
Helvella renervis Skrede & T. Schumach. sp. nov. MycoBank
MB834628. Fig. 2I, J.
Etymology: Referring to the veined (nerved) receptacle surface
forming an incomplete network (reculum).
Typus: Spain, Gipuzkoa, San Sebasan, 21 May 2009, J.M.
Lekuona (holotype ARAN3008363).
Apothecium shortly spitate-cupulate, cup 1.5–3.2 cm broad,
hymenium greyish brown, smooth, oen wrinkled, receptacle
subpubescent, paler, greyish to whish, with prominent
branching ribs and interconnected veins forming a ± reculate
paern, ribs reaching the margin or not, arising from a short
spe. Spe indisnct, ribbed, 0.4–1 cm high. Medullary
excipulum of ghtly interwoven textura intricata, hyphae 3–6
µm broad. Outer excipulum of angular to prismac cells, 10–20
× 5–20 µm broad, protruding in a loosely interwoven fasciculate,
3- to 6-celled tus. Asci cylindrical, pleurorhynchous, 240–280
× 12–16 µm. Ascospores broadly ellipsoid to oval, uniguulate,
14.0–14.6–15.8 × 11.3–12.0–13.3 µm, with one large internal oil
drop. Paraphyses straight, 2.5–3.5 µm broad below, enlarged to
5–7.5 µm at the clavate to subcapitate ps.
Specimens examined/sequenced: See Table 1.
Notes: The species belongs in the H. cosfera morphospecies
complex, but constutes a lineage of its own in our phylogeny (cf.
Fig. 1). Helvella renervis is disnguished from H. cosfera and
H. calycina, by two hsp and 12 rpb2, and four hsp and 12 rpb2
substuons, respecvely. Helvella renervis has also turned
up among recently sequenced specimens of the H. cosfera
morpho-species complex from the fungarium in Stockholm (S),
Sweden (see Table 1).
Helvella semiobruta Donadini & Berthet, Bull. Soc. mycol. Fr. 91:
555. 1976. (1975). Fig. 6C, D.
Illustraon: Donadini & Berthet (1975: gs 1–8).
Apothecium spitate-capitate, cap convex, 1.5–3 cm broad, in
young specimens folded and convex, later bilobate or trilobate and
irregular in shape, margin undulate, gently involute or straight, in
mature specimens deexed and welded to the aachment points
of the spe, hymenium smooth, even or gibbose, folded, black,
receptacle surface smooth to furfuraceous, at rst greyish white,
then greyish black in old specimens, covered by branched ribs
and veins. Spe 1.2–4 cm high, 0.5–2 cm broad, hollow, with
1 to 3 internal chambers, outside with three to ve blunt and
broad ribs and grooves, concolourous with receptacle or paler
above, whish below, in premature specimens spe compressed
with less protruding ribs. Medullary excipulum of dense textura
intricata, hyphae 2.8–6.6 µm broad, with pale brown walls. Outer
excipulum of brownish rows of angular to prismac cells, 10–30
× 7–15 µm turning out perpendicular to receptacle surface. Asci
pleurorhynchous, 300–370 × 14–18 µm. Ascospores oblong
ellipsoidal, 19.8–22.1–24.5 × 12.2–13.8–14.8 µm, with one large
and several small internal guules. Paraphyses straight, 3–4 µm
below, enlarged to 6–9 µm at the clavate ps.
Specimens examined/sequenced: See Table 1.
Notes: This Mediterranean species has a convex cap and a short
greyish white spe with a few longitudinal, blunt ridges and
grooves. It is nested with H. helvellula, H. iberica, H. inexpectata
and H. juniperi in a highly divergent lineage of the /lacunosa
clade (Fig. 1, see also Skrede et al. 2017). A specimen from
Gotland, Sweden, has been idened among specimens of the
H. lacunosa morphospecies complex deposited in the fungarium
of Stockholm (S). Filippa et al. (2013) gave an account of the
morphology and ecology of the species in the Mediterranean
region.
The Spanish specimens of H. semiobruta are all associated
with Cistus albidus and collected from January to March between
500 to 700 m a.s.l. Helvella semiobruta may be confused with
H. juniperi, with which it shares habitat and period of ascoma
formaon. In H. juniperi the spitate-capitate ascoma is more
irregularly saddle-shaped and has more numerous, prominent,
longitudinal ribs and deep furrows on a cavernous, hollow spe.
Genecally, three hsp and three rpb2 substuons separate
H. semiobruta and H. juniperi. Helvella semiobruta also shares
habitat and fruing me with H. helvellula, which is cupulate
and never capitate as in H. semiobruta.
Helvella solitaria P. Karst., Bidr. Känn. Finl. Nat. Folk 19: 37.
1871. Fig. 8H, I.
Synonyms: Helvella quelei Bres., Rev. Mycol. 4: 211. 1882.
Helvella ulvinenii Harmaja, Karstenia 19: 42. 1979.
See Skrede et al. (2017) for addional synonyms.
Illustraons: Dissing (1964: g. 9; 1966b: g. 18, as H. quelei),
Dissing & Nannfeldt (1966: pl. 2: 2, as H. quelei), Weber (1972:
© 2020 Westerdijk Fungal Biodiversity Instute
Dissingia and Helvella species from Spain
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
91
gs 27–28, as H. quelei), Harmaja (1977a: g. 1, 1979: g. 8,
as H. ulvinenii), Schumacher & Mohn Jenssen (1992: 21, as H.
quelei).
Apothecium spitate-cupulate, cup to saucer-shaped, laterally
compressed, 2.5–5 cm broad, hymenium dark greyish brown to
brownish black, even, receptacle concolourous or paler, delicately
pubescent. Spe 1.5–6 cm high, by 0.8–2 cm broad, whish or
pale greyish, with 3 to 6 blunt ribs and grooves connuing onto
the base of the cup. Medullary excipulum of densely interwoven
textura intricata, hyphae 3–7 µm broad. Outer excipulum of
textura angularis, cells slightly elongate, 25–50 × 20–40 µm,
outermost cells giving rise to 3 to 6-celled hyphae in fascicled
tus, outermost cells cylindrical to club-shaped, 10–26 × 8–14
µm. Asci pleurorhynchous, 270–330 × 13–15 µm. Ascospores
ellipsoid, uniguulate, 18.2–20.1–21.8 × 11.4–12.5–13.2 µm.
Paraphyses 3–4 µm broad below, brown-walled, with brownish
granular content, enlarged to 5–8 µm at the clavate ps.
Specimens examined/sequenced: See Table 1.
Notes: The specimens from Spain show no intraspecic hsp
and rpb2 variaon, as also observed among the Scandinavian
populaons (Skrede et al. 2017).
Helvella sublicia Holmsk., Beata ruris oa Fung. Dan. 2: 51.
1799. Fig. 8D, E.
Synonym: Helvella ephippium Le v., Ann. Sci. Nat. Bot., sér. 2, 16:
240. 1841.
See Skrede et al. (2017) for addional synonyms.
Illustraons: Holmskjold (1799: pl. 27), Léveillé (1841: pl. 15 g.
7, as H. ephippium), Dissing & Lange (1967: gs 1a, 7 d, as H.
ephippium).
Apothecium spitate-cupulate, at rst cupulate and laterally
compressed with inrolled margin, then expanding to saddle-
shaped, adnate or reexed, occasionally applanate, 0.5–2 cm
broad, hymenium greyish to dark grey, occasionally whish,
even, receptacle sub-pubescent to villose, concolourous with
hymenium. Spe solid, terete, or compressed with one or
two inconspicuous grooves below, concolourous with cap or
paler to whish. Medullary excipulum of textura intricata,
hyphae 3–5 µm broad, walls brownish, intermixed with some
blackish hyphae. Outer excipulum of textura angularis, cells
20–44 × 8–25 µm, outermost cells cylindrical to club-shaped in
rows perpendicular to receptacle surface, 20–40 × 10–15 µm,
outermost cells protruding to tus of fascicled hyphae forming
the villi of receptacle. Asci cylindrical, pleurorhynchous, 260–
300 × 14–17 µm. Ascospores ellipsoid, uniguulate, 18.8–20.2–
21.5 × 10.7–11.6–12.4 µm. Paraphyses 2–3 µm below, slightly
enlarged to 4–7 µm at the clavate ps.
Specimens examined/sequenced: See Table 1.
Notes: Helvella sublicia diers from other saddle-shaped
Helvella species in its greyish colours and pubescent receptacle
surface. The hsp and rpb2 sequences of the Spanish and Nordic
specimens are idencal.
Helvella sulcata Afzel., K. svenska Vetensk.-Acad. Handl. 4: 305.
1783. - nom. sanct. Fig. 3A, B.
Synonyms: See Skrede et al. (2017) for addional synonyms.
Illustraon: Afzelius (1783: tab. 1, g. 1), Schaeer (1763: pl.
162, as ‘Elvela decima qvarta’).
Apothecium spitate-capitate, cap 1–4 cm broad, 3–6 cm
high, saddle-shaped, bi- to trilobate, adnate, margin aached
to spe at 3 to 4 points, hymenium black, even or slightly
wrinkled, receptacle surface greyish brown, smooth, without
ribs on receptacle surface. Spe 2–5 cm high, 0.5–3 cm broad,
(apothecial height : spe height = 1 : 1), naked, prominently
ribbed, ribs sharp and deep, partly double-edged and
anastomosing, with deep ‘holes’ (lacunae) between, 4–6 along
the circumference, almost no spe esh. Medullary excipulum
of loose textura intricata, hyphae ca. 4–5 µm broad. Outer
excipulum of angular to prismac cells arranged in rows turning
out perpendicular to receptacle surface, individual cells 8 – 15
µm, outermost cells more loosely arranged, club-shaped, 10–30
× 5–15 µm. Asci cylindrical, pleurorhynchous, 290–320 × 13–16
µm. Ascospores broadly ellipsoid, 14.8–16.2–16.8 × 10.5–12.1–
13.2 µm. Paraphyses 3–4 µm broad, light brown below, medium
brown above, enlarged to 5–6.5 µm at the clavate ps.
Specimens examined/sequenced: See Table 1.
Notes: Five hsp, nine rpb2, and 15 LSU substuons disnguish
H. sulcata and H. lacunosa.
Skrede et al. (2017) lectotypied H. sulcata with an authenc
gure (iconotype) from the original work by Afzelius (1783, tab
1, g. 1). A depicon of an authenc colour plate of H. sulcata
prepared by Adam Afzelius himself, located in the fungarium of
Uppsala (UPS), is shown in Fig. 3A. This colour plate constutes
the prototype for the gure of H. sulcata in Afzelius’ work
(Afzelius 1783, plate X, g. 1).
Helvella terricola Skrede & T. Schumach. sp. nov. MycoBank
MB834629. Fig. 7E.
Etymology:dwelling on soil” as opposed to its “look-alike”
sister species H. palustris.
Typus: Spain, Huesca, Cerler, 28 Sep. 2018, F. Cervera & L.
Ballester LB18092810 (holotype O-F-256562).
Apothecium spitate-capitate, cap 1.5–4 cm broad, 1–2 cm high,
saddle-shaped, bi- to trilobate, deexed but margin not aached
to the spe, hymenium greyish black, surface rugose with age,
receptacle concolourous with hymenium or paler, glabrous or
subtomentose, with ribs radiang from spe apex. Spe light grey
to dark grey 3–7.5 cm long, 0.5–1.5 cm thick, slightly tapering
above, solid, with 5 to 8 single- or double-edged longitudinal ribs.
Medullary excipulum of textura intricata, hyphae hyaline, 4–8
µm broad. Outer excipulum of textura globulosa-angularis, cells
20–40 µm diam., hyaline or light brown-walled, the outermost
cells more elongated and prismac giving rise to a palisade of
densely packed club-shaped cells turning out perpendicular to
receptacle surface. Asci pleurorhynchous, 260 – 320 x 12 – 15
µm. Ascospores ellipsoid, 15.0–17.8–19.5 × 10.8–11.5–12.6
µm. Paraphyses light brown-walled, 2–3 µm broad, septate,
gradually enlarged into an irregular clavate p, 5–10 µm broad.
Specimen examined/sequenced: See Table 1.
© 2020 Westerdijk Fungal Biodiversity Instute
Skrede et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
92
Notes: Helvella terricola resembles H. palustris in shape and
colour, but diers genecally in three hsp and three rpb2
substuons. The two species dier substanally in ecology;
H. palustris grows in minerotrophic fens and marshes while H.
terricola grows on naked, moist soil.
DISCUSSION
This follow-up study of Dissingia and Helvella of species from
Spain focuses on Mediterranean South European species and
adds to the current knowledge of the species composion and
diversity of Helvella in Europe. This study complements the
previous synopsis by Skrede et al. (2017) which was primarily
focused on North European species. The Spanish samples
amount to 31 species of Dissingia (1) and Helvella (30) and in
total we add 11 species to the list of European helvellas, of
which nine are described as new. Altogether, 67 species are
now re-evaluated and molecularly characterized from Europe
(Skrede et al. 2017, Løken et al. 2019). Two of the newly
described species are also found in other European countries,
i.e. H. renervis in Sweden and H. phlebophoroides in Denmark
(cf. Table 1). It is worth menoning that H. semiobruta, with a
mainly South European distribuon, has been detected from
the island of Gotland, Sweden, which has a geology and climate
comparable to that of the Mediterranean countries (cf. Table 1).
The diversity illustrated in our survey is astonishing based on the
limited number of collecons (115) available. This underlines
the extent of the unexplored diversity of Helvella and is highly
suggesve that many more species of Helvella await discovery
and descripon, in parcular in regions where few collecons
have been made and are available in fungaria.
The Spanish samples also highlight the extensive radiaon
and diversicaon of e.g. the /helvellula lineage. This highly
divergent lineage of the /lacunosa clade detected by Skrede et
al. (2017), included three species, i.e. H. helvellula, H. semiobruta
and H. juniperi. In the present outline, aer incorporaon of two
addional species, i.e. H. iberica and H. inexpectata, this lineage
comprises ve species (cf. Fig. 1). It is noteworthy that H. juniperi,
which is known with certainty from Italy and Portugal (c f. Table 1),
did not turn up among the Spanish samples. Instead, the two new
species H. iberica and H. inexpectata were detected, apparently
sharing the same type of habitat as H. juniperi. It is expected that
H. juniperi is also present in Spain. Moreover, the morphological
diversicaon of species of this lineage is remarkable, considering
the common ancestry of the group. Both spitate and sessile, as
well as cupulate and capitate species characterised have derived
among these genecally very closely related species.
ACKNOWLEDGEMENTS
We thank all the collectors for this study, both for collecng the
material, but also for taking excellent photographs of fresh ascomata
to be published in the synopsis. We thank curator Raaella Trabucco
at the MCVE Herbarium for the arrangement of loan of holo- and
paratype material of H. juniperi, and assistant Giovanni Robich at
MCVE for sending an in situ photograph of the holotype specimen of
H. juniperi to be published. The help from curator Åsa Kruys at the
UPS Fungarium who has taken photographs of one original collecon
of Helvella pezizoides and authenc illustraons (plates) of Afzelius’
Helvella pezizoides and H. sulcata deposited in the UPS Fungarium to
be presented here is gratefully acknowledged. We thank Prof. Donald
H. Pster and one anonymous reviewer for construcve comments and
valuable correcons to an earlier version of this manuscript. Funding for
this research was provided by a grant from the Norwegian Biodiversity
Informaon Centre to T.S. and I.S. (grant no. 45/15).
Conict of interest: The authors declare that there is no conict
of interest.
REFERENCES
Abbo SP, Currah RS (1997). The Helvellaceae: systemac revision
and occurrence in northern and northwestern North America.
Mycotaxon 62: 1–125.
Afzelius A (1783). Svamp-slägtet Helvella. Kungliga Vetenskaps-
akademiens Nya Handlingar 4: 299–313, 1 pl.
Avise JC, Ball Jr RM (1990). Principles of genealogical concordance
in species concepts and biological taxonomy. Oxford Surveys in
Evoluonary Biology 7: 45–67.
Baara AJA (1759). Fungorum Agri Ariminensis Historia. 2nd ed.,
Favenae: Typus Marnianis, Austria.
Bergeret J (1783). Phytonomatotechnie universelle. Vol. 1. Paris, France.
Boudier [JL] E (1887). Noce sur les Discomycètes gurés dans les
dissins inédits de Dunal conservés à la Faculté de Montpellier.
Bullen de la Société mycologique de France 3: 88–96. pl. 8.
Boudier [JL] E (1904a). Icones Mycologicae - liste préliminaire des noms
des 600 espèces de Champignons qui doivent être gurés dans les
six series de Boudier, Icones Mycologicae, 4 pp., Paul Klincksieck,
Paris, France.
Boudier [JL] E (1904b). Icones Mycologicae, livr. 1. Klincksieck, Paris,
France.
Boudier [JL] E (1906). Icones Mycologicae, livr. 8. Klincksieck, Paris,
France.
Boudier [JL] E (1908a). Icones Mycologicae, livr. 18, Klincksieck, Paris,
France.
Boudier [JL] E (1908b). Icones Mycologicae, livr. 20, Klincksieck, Paris,
France.
Boudier [JL] E (1909a). Icones Mycologicae, livr. 22, Klincksieck, Paris,
France.
Boudier [JL] E (1909b). Icones Mycologicae, livr. 24, Klincksieck, Paris,
France.
Calonge FD, Arroyo I (1990). Notes on the genus Helvella in Spain.
Mycotaxon 39: 203–217.
Caro V, Baiano G, Filippa M (2007). Una species picamente
mediterranea presente anche in Italia: Helvella helvellula.
Mycologia Montenegrina 10: 31–48.
Deman JR, Jacobson DJ, Turner E, et al. (2003). Reproducve isolaon
and phylogenec divergence in Neurospora: comparing methods of
species recognion in a model eukaryote. Evoluon 57: 2721–2741.
Dissing H (1964). Studies in arcc and subarcc Discomycetes. I. The
Genus Helvella. Botanisk Tidsskri 60: 108–128.
Dissing H (1966a). A revision of collecons of the genus Helvella L.
ex St-Amans emend. Nannf. in the Boudier Herbarium. Revue de
Mycologie 31: 189–224, pl. 1–4 (gs. 8–11).
Dissing H (1966b). The genus Helvella in Europe with special emphasis
on the species found in Norden. Dansk Botanisk Arkiv 25: 1–172.
Dissing H (1972). Specic and generic delimitaon in the Helvellaceae.
Persoonia 6: 425–432.
Dissing H (2000). Pezizales Bessey. In: Hansen, L, Knudsen, H (eds.).
Nordic Macromycetes Vol. 1. Ascomycetes: 55–127. Nordswamp,
Copenhagen, Denmark.
© 2020 Westerdijk Fungal Biodiversity Instute
Dissingia and Helvella species from Spain
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
93
Dissing H, Lange M (1967). Notes on the genus Helvella in North
America. Mycologia 59: 349–360.
Dissing H, Nannfeldt JA (1966). Helvella cupuliformis sp. nov., H. villosa
(Hedw. ex O. Kuntze) comb. nov., H. macropus (Pers. ex Fr.) Karst.,
and their allies. Svensk Botanisk Tidskri 60: 325–337.
Donadini, JC (1985). Étude des Discomycètes (I). Addif et commentaires.
Bullen de la Société Linnéenne de Provence 35: 131–138. (1984)
Donadini JC, Berthet P (1975). Une Helvelle nouvelle: Helvella
semiobruta nov. sp. Bullen de la Société mycologique de France
91: 553–555.
Durieu MC (1849). Flore d’Algérie. Exploraon scienque de l’Algerie
1: 441–600.
Edgar, RC (2004) MUSCLE: mulple sequence alignment with high
accuracy and high throughput. Nucleic Acids Research 32: 1792–
1797.
Filippa M, Baiano G (1999). Helvella juniperi sp. nov. Nuova specie
raccolta sul litorale rrenico. Rivista di Micologia 42: 99–118.
Filippa M, Baiano G (2011). Helvella atra König: Fries nomenclatura e
picazione. Ascomycete.org. 2 (4): 57–64.
Filippa M, Baiano G, Baglivo A, et al. (2013). Helvella semiobruta:
rivulatazione di una specie mediterránea. Rivista di Micologia 2013
(3): 195–210.
Fuckel L (1870). Symbolae mycologicae. Beiträge zur Kenntnis der
rheinischen Pilze. Jahrbücher des Nassauischen Vereins für
Naturkunde 23–24: 1–459. (1869–1870)
Gillet GC (1879). Champignons de France. Les Discomycètes 1. Alencon,
France.
Häner J (1987). Die gaung Helvella, morphologie und taxonomie.
Beihee zur Zeitschri für Mykologie 7: 1–165.
Hansen K, Schumacher T, Skrede I, et al. (2019). Pindara revisited –
evoluon and generic limits in Helvellaceae. Persoonia 42: 186–204.
Harmaja H (1977a). A note on Helvella solitaria (syn. H. quelei) and H.
confusa n. sp. Karstenia 17: 40–44.
Harmaja H (1977b). A revision of the Helvella acetabulum group
(Pezizales) in Fennoscandia. Karstenia 17: 45–58.
Harmaja H (1979). Studies on cupulate species of Helvella. Karstenia
19: 33–45.
Holmskjold T (1799). Beata ruris oa Fungis Danicis impensa 2.
København, Denmark.
Inzenga G (1865). Funghi Siciliani. Centuria prima. Palermo, Italy.
Landeros F, Korf, RP (2012). Nomenclatural notes 13. An incorrect
neotype designaon and provision for a lectotype and an epitype
for Helvella fusca. Mycotaxon 119: 431–438.
Landeros F, Iturriaga T, Guzmán-Dávalos L (2012). Type studies in
Helvella (Pezizales) 1. Mycotaxon 119: 35–63.
Landeros F, Iturriaga T, Rodriguez A, et al. (2015). Advances in the
phylogeny of Helvella (Fungi: Ascomycota), inferred from nuclear
ribosomal LSU sequences and morphological data. Revista
Mexicana de Boversidad 86: 856–871.
Léveillé JH (1841). Descripon de quelques espèces nouvelles des
champignons. Annales des Sciences Naturelles Botanique, 2e Série
16: 235–242, pl. 14–15.
Løken SB, Skrede I, Schumacher T (2020). The Helvella corium species
aggregate in Nordic countries – phylogeny and species delimitaon.
Fungal Systemacs and Evoluon 5: 169–186.
Moravec J (1980). Helvella leucopus Pers. in Czechoslovakia.
(Discomycetes, Helvellaceae). Ceska Mykologie 34: 214–216, pl. IV.
Nannfeldt JA (1932). Bleka Stenmurklan, Gyromitra gigas (Krombh.)
Cke. Friesia 1: 34–45.
Nannfeldt JA (1937). Contribuons to the mycoora of Sweden. 4. On
some species of Helvella, together with a discussion of the natural
anies within Helvellaceae and Pezizaceae trib. Acetabuleae.
Svensk Botanisk Tidskri 31: 47–66, 2 plates.
Oeder GC (1770). Abbildungen der Panzen, welche in den Königreichen
Dännemark und Norwegen, in den Herzogthümern Schlesswig und
Holstein, und in den Grafschaen Oldenburg und Delmenhorst
wild wachsen, zu Erläuterung des unter dem Titel Flora Danica,
auf königl. Befehl veranstalteten Werkes von diesen Panzen /
herausgegeben von dem Verfasser des bemeldten Werkes Georg
Chrisan Oeder, Doctor der Arzenenkunst [sic] und Königlichen
professor der Botanik. Flora Danica 3(9). Copenhagen, Denmark.
Schaeer JC (1763). Fungorum qui in Bavaria et Palanatu circa
Rasbonam nascuntur icones navis coloribus expressae. Tomus 2.
Henrici Godofredi Zunkelii, Rasbonae, Germany.
Schaeer JC (1774). Fungorum qui in Bavaria et Palanatu circa
Rasbonam nascuntur icones navis coloribus expressae. Tomus 4.
Henrici Godofredi Zunkelii, Rasbonae, Germany.
Schumacher T, Mohn Jenssen K (1992). Discomycetes from the Dovre
Mountains, Central South Norway. Arcc and alpine fungi, Vol. 4.
Soppkonsulenten A/S, Oslo, Norway.
Skrede I, Carlsen T, Schumacher T (2017). A synopsis of the saddle fungi
(Helvella: Ascomycota) in Europe – species delimitaon, taxonomy
and typicaon. Persoonia 39: 201–253.
Stamatakis A (2006). RAxML-VI-HPC: maximum likelihood-based
phylogenec analyses with thousands of taxa and mixed models.
Bioinformacs 22: 2688–2690.
Taylor JW, Jacobson DJ, Kroken S, et al. (2000). Phylogenec species
recognion and species concepts in fungi. Fungal Genecs and
Biology 31: 21–32.
Trania L (1809). Die essbaren Schwämme des Oesterreichischen
Kaiserstaates. Wien & Trieste, Austria.
Van Vooren N (2010). Notes sur le genre Helvella L. (Ascomycota,
Pezizales). 1. Le sous-genre Elascae. Bullen mycologique et
botanique Dauphiné-Savoie 199: 27–60.
Van Vooren N (2014). Notes sur le genre Helvella L. (Ascomycota,
Pezizales). 2. Les sous-genres Cupuliformes et Macropodes. Bullen
mycologique et botanique Dauphiné-Savoie 212: 29–47.
Wang X-C, Liu T-Z, Chen S-L, et al. (2019). A four-locus phylogeny of
rib-sped cupulate species of Helvella (Helvellaceae, Pezizales) with
discovery of three new species. MycoKeys 60: 45-67.
Weber NS (1972). The genus Helvella in Michigan. The Michigan
Botanist 11: 147–201.
Weber NS (1975). Notes on western species of Helvella. I. Beihee Nova
Hedwigia 51: 25–38, pl. 7-8.
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