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

December 2020
Fungal Systematics and Evolution 6(1)

DOI:10.3114/fuse.2020.06.05

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CC BY-NC-ND 4.0

Authors:

Inger Skrede

University of Oslo

Cecilie Mathiesen

University of Oslo

Trond Schumacher

University of Oslo

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.

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.

…

(Continued).

…

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. retinervis 2404-16HP1 [H1000]. J. H. retinervis ARAN3008363 [H1925]. Scale bars = 1 cm. All photographs taken by the collectors (confer Table 1).

…

Photographs of fresh apothecia and illustrations A. Authentic illustration 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. queletiana 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).

…

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).

…

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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

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

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

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

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

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

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

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

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

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

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-

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

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

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

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

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

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

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

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

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.

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

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

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

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

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.

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

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).

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

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).

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

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).

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

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).

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

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).

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

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).

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

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).

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

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).

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

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 (‘monachella’ orth. 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

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

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

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

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).

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

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

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

/ 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.

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

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:

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

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.

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

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.

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During investigations into fungal diversity in the Tibetan Plateau, two Helvella collections were made. Morphological characteristics and phylogenetic analyses of combined ITS and LSU sequence data revealed one collection represents a distinct new species that is introduced as Helvella cystidiata. Helvella cystidiata is characterized by having loosely saddle-shaped pileus, greyish to grey brown hymenium, lacunose stipe and inflated ectal excipulum cells. Furthermore, a known species Helvella lacunosa is reported in Tibetan Plateau. Detailed descriptions, illustrations and a phylogenetic tree to show the placement of two taxa are provided.

The Helvella corium species aggregate in Nordic countries – phylogeny and species delimitation

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Dec 2019

Mycologists have always been curious about the elaborate morphotypes and shapes of species of the genus Helvella . The small, black, cupulate Helvella specimens have mostly been assigned to Helvella corium , a broadly defined morpho-species. Recent phylogenetic analyses, however, have revealed an aggregate of species hidden under this name. We performed a multispecies coalescent analysis to re-assess species limits and evolutionary relationships of the Helvella corium species aggregate in the Nordic countries. To achieve this, we used morphology and phylogenetic evidence from five loci – heat shock protein 90 ( hsp ), translation elongation factor 1-alpha ( tef ), RNA polymerase II ( rpb2 ), and the 5.8S and large subunit (LSU) of the nuclear ribosomal DNA. All specimens under the name Helvella corium in the larger university fungaria of Norway, Sweden and Denmark were examined and barcoded, using partial hsp and/or rpb2 as the preferential secondary barcodes in Helvella . Additional fresh specimens were collected in three years (2015–2018) to obtain in vivo morphological data to aid in species discrimination. The H. corium species aggregate consists of seven phylogenetically distinct species, nested in three divergent lineages, i.e. H. corium , H. alpina and H. pseudoalpina sp. nov. in the /alpina-corium lineage, H. alpestris , H. macrosperma and H. nannfeldtii in the /alpestris-nannfeldtii lineage, and H. alpicola as a weakly supported sister to the /alpestris-nannfeldtii lineage. Among the seven species, the ribosomal loci expressed substantial variation in evolutionary rates, suggesting care in the use of these regions alone in delimitation of Helvella species. Altogether, 469 out of 496 available fungarium specimens were successfully barcoded.

A four-locus phylogeny of rib-stiped cupulate species of Helvella (Helvellaceae, Pezizales) with discovery of three new species

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Oct 2019

Helvella species are ascomycetous macrofungi with saddle-shaped or cupulate apothecia. They are distributed worldwide and play an important ecological role as ectomycorrhizal symbionts. A recent multi-locus phylogenetic study of the genus suggested that the cupulate group of Helvella was in need of comprehensive revision. In this study, all the specimens of cupulate Helvella sensu lato with ribbed stipes deposited in HMAS were examined morphologically and molecularly. A four-locus phylogeny was reconstructed using partial sequences of the heat shock protein 90, nuclear rDNA internal transcribed spacer region 2, nuclear large subunit ribosomal DNA and translation elongation factor 1-α genes. Three clades were revealed in Helvella sensu stricto. Twenty species were included in the analysis, of which 13 are distributed in China. Three new species, H. acetabuloides, H. sichuanensis and H. tianshanensis, are described and illustrated in detail. A neotype was designated for H. taiyuanensis. Helvella calycina is a new record for China, while Dissingia leucomelaena should be excluded from Chinese mycota. Hsp90 and ITS2 are recommended as useful supplementary barcodes for species identifications of the genus.

Pindara revisited – evolution and generic limits in Helvellaceae

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Jun 2019

The Helvellaceae encompasses taxa that produce some of the most elaborate apothecial forms, as well as hypogeous ascomata, in the class Pezizomycetes (Ascomycota). While the circumscription of the Helvellaceae is clarified, evolutionary relationships and generic limits within the family are debatable. A robust phylogeny of the Helvellaceae, using an increased number of molecular characters from the LSU rDNA, RPB2 and EF-1α gene regions (4 299 bp) and a wide representative sampling, is presented here. Helvella s.lat. was shown to be polyphyletic, because Helvella aestivalis formed a distant monophyletic group with hypogeous species of Balsamia and Barssia. All other species of Helvella formed a large group with the enigmatic Pindara (/Helvella) terrestris nested within it. The ear-shaped Wynnella constitutes an independent lineage and is recognised with the earlier name Midotis. The clade of the hypogeous Balsamia and Barssia, and H. aestivalis is coherent in the three-gene phylogeny, and considering the lack of phenotypic characters to distinguish Barssia from Balsamia we combine species of Barssia, along with H. aestivalis, in Balsamia. The closed/tuberiform, sparassoid H. astieri is shown to be a synonym of H. lactea; it is merely an incidental folded form of the saddle-shaped H. lactea. Pindara is a sister group to a restricted Helvella, i.e., excluding the /leucomelaena lineage, on a notably long branch. We recognise Pindara as a separate genus and erect a new genus Dissingia for the /leucomelaena lineage, viz. H. confusa, H. crassitunicata, H. leucomelaena and H. oblongispora. Dissingia is supported by asci that arise from simple septa; all other species of Helvellaceae have asci that arise from croziers, with one exception being the /alpina-corium lineage of Helvella s.str. This suggests ascus development from croziers is the ancestral state for the Helvellaceae and that ascus development from simple septa has evolved at least twice in the family. Our phylogeny does not determine the evolutionary relationships within Helvella s.str., but it is most parsimonious to infer that the ancestor of the helvelloids produced subsessile or shortly stipitate, cup-shaped apothecia. This shape has been maintained in some lineages of Helvella s.str. The type species of Underwoodia, Underwoodia columnaris, is a sister lineage to the rest of the Helvellaceae.

A synopsis of the saddle fungi (Helvella: Ascomycota) in Europe - Species delimitation, taxonomy and typification

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Dec 2017

Helvella is a widespread, speciose genus of large apothecial ascomycetes (Pezizomycete:Pezizales) that are found in terrestrial biomes of the Northern and Southern Hemispheres. This study represents a beginning on assessing species limits and applying correct names forHelvellaspecies based on type material and specimens in the university herbaria (fungaria) of Copenhagen (C), Harvard (FH) and Oslo (O). We use morphology and phylogenetic evidence from four loci - heat shock protein 90 (hsp), translation elongation factor alpha (tef), RNA polymerase II (rpb2) and the nuclear large subunit ribosomal DNA (LSU) - to assess species boundaries in an expanded sample ofHelvellaspecimens from Europe. We combine the morphological and phylogenetic information from 55Helvellaspecies from Europe with a small sample ofHelvellaspecies from other regions of the world. Little intraspecific variation was detected within the species using these molecular markers;hspandrpb2markers provided useful barcodes for species delimitation in this genus, while LSU provided more variable resolution among the pertinent species. We discuss typification issues and identify molecular characteristics for 55 EuropeanHelvellaspecies, designate neo- and epitypes for 30 species, and describe sevenHelvellaspecies new to science, i.e.,H. alpicola,H. alpina,H. carnosa,H. danica,H. nannfeldtii,H. pubescensandH. scyphoides.

Notes on the Genus Helvella in North America

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