Abstract
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A synopsis of the saddle fungi (Helvella: Ascomycota) in Europe – species delimitation, taxonomy and typification
Abstract
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 for Helvella species 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 of Helvella specimens from Europe. We combine the morphological and phylogenetic information from 55 Helvella species from Europe with a small sample of Helvella species from other regions of the world. Little intraspecific variation was detected within the species using these molecular markers; hsp and rpb2 markers 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 European Helvella species, designate neo- and epitypes for 30 species, and describe seven Helvella species new to science, i.e., H. alpicola, H. alpina, H. carnosa, H. danica, H. nannfeldtii, H. pubescens and H. scyphoides.
INTRODUCTION
Helvella is a widespread, speciose genus of apothecial ascomycetes (Pezizomycetes: Pezizales) whose members are found in terrestrial biomes of the Northern and Southern Hemispheres. The genus contains many of the larger and charismatic species of the order Pezizales and comprises a range of elaborate apothecia, from cupulate to saddle-shaped, and convex to campanulate, including species with folded and lobed caps seated on a simple, ribbed or furrowed stipe.
Although easily separated from other macrofungi by conspicuous polymorphic apothecia, it is surprisingly difficult to distinguish between Helvella species. Historically, the shape and colour and outer surface characters have been emphasized in species discrimination, whilst microanatomy of the sterile and fertile structures has added few characters of value in species recognition. Weber (1972), after extensive studies of Michigan collections of Helvella, concluded that: “in summary, most morphological and anatomical characters exhibit nearly continuous variation in the genus as a whole. Each species, however, represents only one area on the spectrum of variation for each character, and several species may share a given part of the spectrum for a character. Thus a group of characters must be used to delimit species”.
Linnaeus (1737, 1753) proposed the genus Elvela (= Helvella, orth. var.) for Elvela mitra (= Helvella crispa ss. Fries (1822)). Fries (1822) erected the family Elvellaceae, which was later referred to as Helvellaceae (Corda 1842). Generic delimitation of the morphologically diverse Helvella has varied. Fuckel (1870) segregated two new genera, i.e., Macroscyphus and Paxina, and Boudier (1907a) recognized altogether five genera, i.e., Leptopodia, Cyathipodia, Acetabula, Macropodia and Helvella s.str., for what was later merged into a broadly defined Helvella (Quélet 1886, Nannfeldt 1932, 1937, Dissing 1966a, b, Weber 1972, 1975, Häffner 1987, Abbott & Currah 1997). In its widest circumscription over the last century, Helvella encompassed the monotypic genera Underwoodia (Peck 1890, Eckblad 1968, Harmaja 1974), Wynnella (Boudier 1885, 1907, Harmaja 1974, Abbott & Currah 1997), Pindara (Velenovský 1934, Landvik et al. 1999) and the hypogeous Helvella astieri (Korf 1973a).
A number of Helvella taxa dating from late 18th (pre-Friesian) and early 19th century taxonomic works, have been variously interpreted over time. Most taxonomists, including Fries himself, applied a broad species concept allowing for a lot of morphological variation among the Helvella species that were accepted in their monumental works (Afzelius 1783, Fries 1822, Dissing 1966a, b, Weber 1972, 1975, Abbott & Currah 1997). These works also listed a number of heterotypic synonyms for many Helvella species, mostly assessed based on similarities of published descriptions rather than as a result of comparative studies of type specimens. Furthermore, the iconic European Helvella species names, e.g., H. lacunosa, H. corium, H. elastica and H. acetabulum have been applied to specimens belonging to a number of phylogenetically related or unrelated Helvella taxa in regional or local reports on the genus, and misidentifications of Helvella spp. in the literature and on the web are common. The more comprehensive 20th and 21st century taxonomic works on the genus Helvella in Europe include the works of Dissing (1966a, b, 2000), Harmaja (1977b, 1979), Häffner (1987) and Van Vooren (2010, 2014). Dissing (1966b) recognised 26 species of Helvella in Europe, a list which was later expanded to 28 (Dissing 1972) and to 38 in a later survey of the species confined to the restricted area of the Nordic countries (Dissing 2000). Häffner (1987) accepted altogether 41 species from Germany and the rest of the European continent.
The development of molecular systematics and the possibility of employing DNA barcode sequences as a more robust tool to identify specimens of closely related species have recently and to some extent been applied to taxonomic studies of Helvella. Landvik et al. (1999) used nrDNA sequences to characterise a small subset of Helvella species from Norway and emphasized the general limitation of single locus analyses and the unsuitability of ITS to infer phylogeny and discriminate species across the breadth of the genus. Nguyen et al. (2013) used ITS and LSU nrDNA sequences and morphology to discriminate among at least four distinct species of the H. lacunosa species complex in western North American, and Landeros et al. (2015) used partial LSU sequences to study infrageneric groups of Helvella from Europe and North America. Recently, Ariyawansa et al. (2015) used the ITS, Zhao et al. (2015) ITS and a concatenated multilocus dataset of ITS, LSU, tef, rpb2 and mcm7, and Zhao et al. (2017) a dataset of ITS, LSU and tef to discriminate and describe new species of the H. lacunosa, H. crispa and H. monachella morphospecies complexes in China. To this array of genes we add hsp as an informative additional gene of utility in species identification in Helvella.
We use sequence data and a simplified genealogical concordance phylogenetic species recognition (GCPSR) (Avise & Ball 1990, Taylor et al. 2000, Dettman et al. 2003) approach to resolve species-level lineages and phylogenetic species of Helvella in Europe. By using this molecular approach in combination with morphological data, we reassess species limits within Helvella in Europe. We hypothesize that well-supported terminal clades based on more than one locus and unique singletons linked to geographic separation in our phylograms represent biological species of Helvella. Another major objective of this study has been to undertake a nomenclatural revision and propose typifications of Helvella spp. in Europe to stabilize name use and bring the nomenclature in accordance with the Code (ICN 2012). This will hopefully form a basis for further studies to critically assess the biogeography and ecology of Helvella species worldwide.
MATERIALS AND METHODS
The present morphological and molecular synopsis of Helvella species in Europe is based on a selection of Helvella specimens from Europe, America, Asia and Oceania stored at the herbaria (fungaria) of Copenhagen (C), Harvard (FH) and Oslo (O). All sequenced specimens used in this study were given a number for easier recognition throughout the study (H001-H1030). The studied material also includes a large collection of helvellas made by the senior author in the 1980s and 1990s (Herb. T. Schumacher, now deposited in O). The fungarium collections were supplemented with 28 freshly collected specimens made by the authors in 2009. Colour codes of fresh apothecia are based on Kornerup & Wanscher (1961). We used water and Cotton Blue in lactic acid (CB) to observe microanatomical features of hymenium and receptacle (excipular tissues) of fresh and rehydrated materials. We present an artificial key to 55 species, provide full descriptions of seven new species and 12 emended species concepts, and refer to good discriminating descriptions of the rest of the treated species of Helvella in Europe. Macromorphological and microanatomical terminology used in descriptions and key follow Dissing (1966a, b), Dissing & Lange (1967), Eckblad (1968), Weber (1972, 1975) and Korf (1973b).
We use sequence data from four loci to obtain an initial estimate of the genetic diversity represented by 432 collections of dried and fresh specimens of Helvella, each annotated by a ‘H’ number. In addition to a broad sample of European Helvella collections, we included some collections from the American and Asian continents in order to evaluate and better understand the intra- and intercontinental variations and distributions of the pertinent species.
DNA extraction and PCR
DNA was extracted using the E.Z.N.A HP Fungal DNA kit (Omega Bio-Tek, Norcross, GA, USA) protocol for dried samples, except that tissue was not ground, but frozen and thawed two times in the extraction buffer, and subsequently vortexed thoroughly with one tungsten bead. Samples collected by the authors were stored directly in extraction buffer and thus, were not dried before DNA extraction, but extracted using the same protocol as above.
Efforts were made to PCR amplify parts (270–700 bp) of four genetic loci that had been found useful in resolution of lower level relationships in other fungal groups; i.e., the translation elongation factor 1-α (tef), the large subunit RNA polymerase II (rpb2), the heat shock protein 90 (hsp) and the nuclear LSU ribosomal DNA (including D1–D2 domains) (Johannesson et al. 2000, Taskin et al. 2010, O’Donnell et al. 2011, Bonito et al. 2013, Hansen & Olariaga 2015).
A combination of previously published and newly designed primers was used (Table 1). Universal rDNA primers and primer pairs published for rpb2 and hsp failed to produce amplicons from many of the fungarium specimens. Therefore, new primer pairs were designed for rpb2 and hsp, while Helvella specific rDNA primers from Landeros et al. (2015) were used to amplify the parts of the nuclear LSU ribosomal DNA. We initially used the hsp primer pair from Johannesson et al. (2000) and the rpb2-7cf (Liu et al. 1999) and rpb2-9f (Taskin et al. 2010) to generate a few sequences that were used to design new internal primers using Primer3 (Koressaar & Remm 2007, Untergrasser et al. 2012) within in Geneious (http://www.geneious.com; Kearse et al. 2012). The Helvella specific rpb2 forward primer is only 2 bp different from the rpb2-3r primer from Taskin et al. (2010).
Table 1
Locus1 | Forward primer sequence (5’-3’) | Reverse primer sequence (5’-3’) |
---|---|---|
LSU | H_LSUf12: AGCGGAGGAAAGAAACCAACA | H_LSUr22: TCCCAACAGCTATGCTCCTAC |
rpb2 | H_rpb2r2: TCCACAATCTGCATCCCGATTCGC | H_rpb2f: CCAGACATGGACAGAAGGTTGAGCT |
hsp90 | H_hspf : CRGGCATCCGGGTGACGTAAT | H_hspr: AGGGKGTTGTCGACTCCGAGG |
tef | EF595F3: CGTGACTTCATCAAGAACATG | EF1160R3: CCGATCTTGTAGACGTCCTG |
1 LSU: 28s large subunit ribosomal RNA, domains D1\D2; rpb2: RNA polymerase II; hsp: heat shock protein 90; tef: translation elongation factor 1-alpha.
2 From Landeros et al. (2015). They added a G at the 3’end of forward primer and TC at the 3’end of the reverse primer, which were not used in the current study.
3 From Kauserud & Schumacher (2001).
We generated amplicons for the four loci using the following PCR protocols: 4 min at 95 °C, followed by 40 (50 for LSU) cycles of 25 s (30 s for LSU and hsp) at 95 °C, 30 s at 53 °C (52 °C for LSU, 58 °C for hsp) and 60 s at 72 °C, followed by a 10 min extension at 72 °C and a indefinite hold at 10 °C. For all PCR reactions, we used the PuReTaq Ready-To-Go PCR Beads (GEhealthcare, Waukesha, WI) in 25 μL reactions (12 μL for hsp and rpb2). We purified 8 μL PCR products with 0.2 μL ExoSAP-IT or ExoSTAR (GEhealthcare) and 1.8 μL H2O. Sequencing was performed at the in-house ABI-laboratory, Department of Biosciences, UiO or GATC Biotech (Constance, Germany). All sequences were assembled and edited using Geneious R4-7. All sequences have been deposited in GenBank and accession numbers are given in Table 2.
Table 2
Species | Type/Sample ID/Fungaria1 | Locality | Collectio year | GenBank accession numbers2 | |||
---|---|---|---|---|---|---|---|
LSU | tef | hsp | rpb2 | ||||
H. acetabulum | epitype, H410, C-F Fungi Exs. Suec. 1354 | Sweden, Östergötland | 1945 | KY773154 | – | KY784506 | KY772752 |
H126, O-178001 | Norway, Møre og Romsdal | 2004 | KY772978 | – | KY784259 | KY772509 | |
H127, O-178005 | Norway, Møre og Romsdal | 2004 | – | KY772872 | KY784260 | KY772510 | |
H128, O-284420 | Norway, Vestfold | 2006 | KY772979 | KY772873 | KY784261 | KY772511 | |
H133, O-286533 | Norway, Vestfold | 2005 | KY772984 | KY772875 | KY784266 | KY772516 | |
H134, O-64925 | Norway, Østfold | 2001 | KY772985 | KY772876 | KY784267 | KY772517 | |
H225, O-253212 | Norway, Oppland | 1984 | KY773055 | KY772894 | KY784344 | KY772594 | |
H226, O-253213 | Norway, Oppland | 1984 | KY773056 | – | KY784345 | KY772595 | |
H261, C-F-21174 | Iceland, Nordur-Mulasysla | 1983 | KY773082 | – | KY784376 | KY772625 | |
H286, O-253214 | Norway, Oppland | 1983 | – | – | KY784399 | KY772648 | |
H287, O-253215 | Norway, Oppland | 1982 | KY773095 | – | KY784400 | KY772649 | |
H409, C-F Fungi Exs. Suec. 3261 | Sweden, Öland | 1967 | KY773153 | – | KY784505 | KY772751 | |
H427, C-F-45314 | Czech Republic, Vändra | 1960 | KY773159 | – | KY784521 | KY772764 | |
H485, O-253211 | Norway, Finnmark | 1961 | – | – | KY784564 | – | |
H. aestivalis | H007, O-253216 | Norway, Oppland | 2007 | KY772905 | – | KY784187 | – |
H024, O-253217 | Norway, Oppland | 2009 | KY772917 | – | KY784200 | KY772444 | |
H185, O-253218 | Norway, Oppland | 2001 | KY773028 | – | KY784312 | KY772563 | |
H288, C-F-45329 | Kyrgyz Republic, Thien Shan | 1967 | KY773096 | – | KY784401 | KY772650 | |
H355, O-129530 | Norway, Oppland | 1957 | – | – | KY784456 | – | |
H. alpestris | epitype, H014, O-253221 | Norway, Oppland | 2007 | KY772909 | KY772826 | KY784191 | KY772435 |
H013, O-253220 | Norway, Oppland | 2007 | KY772908 | KY772825 | KY784190 | KY772434 | |
H031, O-253222 | Norway, Oppland | 2009 | KY772922 | KY772840 | – | KY772451 | |
H036, O-253223 | Norway, Oppland | 2007 | KY772927 | KY772842 | KY784210 | – | |
H042, O-253224 | Norway, Oppland | 2009 | KY772932 | KY772847 | – | KY772457 | |
H246, O-253219 | Norway, Oppland | 1997 | KY773073 | – | KY784364 | – | |
H460, O ex DAOM574891 | Canada, Nunavut | 2014 | – | – | KY784542 | KY772789 | |
H483, O-253225 | Norway, Oppland | 1984 | – | – | KY784562 | KY772807 | |
H. alpicola | H231, O-253226 | Switzerland, Graubünden | 1984 | KY773061 | – | KY784349 | KY772598 |
holotype, H175, O-185924 | Norway, Nordland | 1988 | KY773020 | – | KY784304 | KY772554 | |
H. alpina | H336, O-253227 | Canada, British Columbia | 1994 | KY773116 | – | KY784439 | KY772690 |
H349, C-F-63820 | Greenland, Qeqertarssuaq | 1977 | – | – | KY784450 | KY772701 | |
holotype, H223, O-253228 | France, Savoie | 1992 | KY773054 | – | KY784343 | KY772593 | |
H. arctoalpina | H003, O-253232 | Norway, Oppland | 2001 | – | KY772818 | KY784183 | – |
H011, O-253233 | Norway, Oppland | 2007 | – | KY772823 | – | – | |
H012, O-253234 | Norway, Oppland | 2007 | KY772907 | KY772824 | KY784189 | – | |
H026, O-253235 | Norway, Oppland | 2009 | – | KY772835 | KY784202 | KY772446 | |
H030, O-253236 | Norway, Oppland | 2009 | – | KY772839 | KY784206 | KY772450 | |
H033, O-253237 | Norway, Oppland | 2009 | KY772924 | KY772841 | KY784207 | KY772453 | |
H040, O-253238 | Norway, Oppland | 2009 | KY772931 | KY772846 | – | KY772456 | |
H083, O-253239 | Norway, Oppland | 1984 | KY772955 | KY772865 | KY784235 | KY772482 | |
H084, O-253240 | Norway, Oppland | 1996 | KY772956 | KY772866 | KY784236 | KY772483 | |
H086, O-253241 | Norway, Oppland | 1966 | – | – | KY784238 | KY772485 | |
H087, O-253242 | Norway, Oppland | 1966 | – | – | KY784239 | KY772486 | |
H173, O-69063 | Norway, Nordland | 2008 | – | – | KY784302 | KY772552 | |
H182, O-253243 | Norway, Oppland | 2001 | – | KY772885 | KY784310 | – | |
H255, O-253403 | Norway, Oppland | 1997 | KY773077 | – | KY784371 | KY772620 | |
H337, C-F-56723 | Canada, Nunavut | 1984 | – | – | KY784440 | KY772691 | |
H338, C-F-56722 | Canada, Nunavut | 1984 | – | – | KY784441 | KY772692 | |
H342, C-F-34334 | Iceland, Nordur-Mulasysla | 1993 | KY773120 | – | KY784445 | KY772696 | |
H354, O-174714 | Norway, Oppland | 1957 | – | – | KY784455 | KY772706 | |
H356, O-253244 | Norway, Oppland | 1984 | KY773124 | – | KY784457 | KY772707 | |
H359, C-F-50652 | Greenland, Ella Isl. | 1957 | KY773127 | – | KY784460 | KY772710 | |
H360, O-253229 | France, Val d’Isere | 1992 | KY773128 | – | KY784461 | KY772711 | |
H456, O-253230 | Norway, Hordaland | 2014 | – | – | – | KY772785 | |
H472, O-253231 | Norway, Hordaland | 2014 | – | – | KY784553 | KY772800 | |
holotype, H293, O-72616 | Norway, Hordaland | 1959 | – | – | KY784406 | KY772655 | |
H. atra | epitype, H406, C-F Fungi Exs. Suec. 2066 | Sweden, Uppland | 1938 | – | – | KY784502 | KY772748 |
H016, O-253251 | Norway, Oppland | 2007 | KY772911 | KY772828 | KY784193 | KY772437 | |
H151, O-281641 | Norway, Nordland | 2006 | KY773000 | KY772881 | – | KY772534 | |
H233, O-253245 | Georgia, Caucasus | 1988 | KY773063 | – | KY784351 | KY772600 | |
H375, O-253246 | Georgia, Caucasus | 1988 | KY773138 | – | KY784474 | KY772723 | |
H462, O-253249 | Norway, Nordland | 2010 | – | – | KY784544 | KY772791 | |
H464, O-253250 | Norway, Nordland | 2005 | – | – | KY784545 | KY772792 | |
H465, O-253252 | Norway, Oppland | 1981 | – | – | KY784546 | KY772793 | |
H467, C-F-45782 | Denmark, Zealand | 1999 | – | – | KY784548 | KY772795 | |
H481, O-291384 | Norway, Sør-Trøndelag | 2009 | – | – | KY784560 | KY772805 | |
H484, O-253247 | Norway, Finnmark | 1961 | – | – | KY784563 | – | |
H487, O-253248 | Norway, Finnmark | 1961 | – | – | KY784566 | KY772809 | |
H. bicolor | epitype, H289, C-F-45334 | Sweden, Gästrikland | 1978 | KY773097 | – | KY784402 | KY772651 |
H056, O-253253 | Norway, Akershus | 2009 | KY772940 | – | KY784220 | KY772467 | |
H057, O-253254 | Norway, Akershus | 2009 | KY772941 | – | KY784221 | KY772468 | |
H129, O-65375 | Norway, Hedmark | 2002 | KY772980 | – | KY784262 | KY772512 | |
H147, O-105346 | Norway, Vestfold | 2000 | KY772996 | – | KY784280 | KY772530 | |
H. calycina | epitype, H022, O-253255 | Norway, Oppland | 2009 | KY772915 | KY772833 | KY784198 | KY772442 |
H382, C-F-21122 | Denmark, Klim Bjerg | 1993 | KY773143 | – | KY784480 | KY772727 | |
H384, C-F-41038 | Denmark, Klim Bjerg | 1999 | KY773144 | – | KY784482 | KY772729 | |
H387, C-F-71286 | Denmark NE Jutland | 1998 | KY773147 | – | KY784484 | KY772732 | |
H. capucina | epitype, H208, O-253256 | France, Savoie | 1992 | KY773041 | – | KY784331 | KY772582 |
H009, O-253258 | Norway, Oppland | 2007 | – | – | KY784188 | KY772432 | |
H034, O-253259 | Norway, Oppland | 2007 | KY772925 | – | KY784208 | KY772454 | |
H035, O-253260 | Norway, Oppland | 2007 | KY772926 | – | KY784209 | KY772455 | |
H046, O-253261 | Norway, Oppland | 2009 | KY772935 | KY772849 | KY784216 | KY772461 | |
H082, O-253262 | Norway, Oppland | 1983 | – | – | KY784234 | KY772481 | |
H089, O-253263 | Norway, Oppland | 1996 | KY772958 | – | KY784240 | KY772487 | |
H201, O-253264 | Norway, Oppland | 1989 | KY773036 | – | KY784325 | KY772576 | |
H202, O-253265 | Norway, Oppland | 1989 | KY773037 | – | KY784326 | KY772577 | |
H203, O-253266 | Switzerland, Graubünden | 1984 | – | – | KY784327 | KY772578 | |
H211, O-253257 | France, Savoie | 1992 | KY773043 | – | KY784333 | KY772584 | |
H397, C-F-92107 | Canada, NW Territory | 1971 | – | – | KY784494 | KY772741 | |
H459, O ex DAOM574890 | Canada, Nunavut | 2014 | – | – | KY784541 | KY772788 | |
H488, C-F-50768 | Greenland, Mestervig | 1982 | – | – | KY784567 | KY772810 | |
H. capucinoides | H113, FH | USA, Maine | 1997 | KY772969 | – | KY784251 | KY772501 |
H398, C-F-92116 | USA, Michigan | 1968 | – | – | KY784495 | KY772742 | |
H. carnosa | holotype, H146, O-68100 | Norway, Oslo | 2006 | KY772995 | – | KY784279 | KY772529 |
KH.10.277, S | Sweden, Gotland | 2010 | KY6600423 | – | – | KY6600443 | |
H. compressa | H112, FH | USA, California | 1998 | KY772968 | – | KY784250 | KY772500 |
H. confusa | H008, O-253269 | Norway, Oppland | 2007 | KY772906 | – | – | KY772431 |
H092, O-253272 | Switzerland, Graubünden | 1984 | KY772960 | – | KY784242 | KY772489 | |
H155, O-71947 | Norway, Aust Agder | 1998 | – | – | – | KY772538 | |
H192, O-253270 | Norway, Østfold | 1987 | – | – | – | KY772570 | |
H253, O-253271 | Norway, Østfold | 1981 | – | – | – | KY772618 | |
H279, C-F-92132 | Sweden, Uppland | 1965 | KY773091 | – | – | KY772642 | |
H313, C-F-58808 | Norway, Nordland | 1998 | KY773104 | – | – | KY772670 | |
H320, C-F-92125 | Norway, Nordland | 1979 | KY773107 | – | – | KY772676 | |
H331, C-F-56845 | Switzerland, Graubünden | 1979 | KY773113 | – | KY784435 | KY772686 | |
H386, C-F-55016 | Norway, Nordland | 1981 | KY773146 | – | – | KY772731 | |
H437, O-253268 | Norway, Buskerud | 2014 | KY773164 | – | KY784529 | KY772772 | |
H438, C | Denmark, Northern Jutland | 1986 | KY773165 | – | – | KY772773 | |
H439, O-129435 | Norway, Oppland | 1987 | KY773166 | – | KY784530 | KY772774 | |
H446, C-F-45513 | India, Kupwara | 1967 | – | – | KY784532 | KY772776 | |
H449, C-F-56513 | USA, Oregon | 1991 | – | – | – | KY772779 | |
isotype, H477, C-F-70807 | Denmark, Northern Jutland | 1965 | – | – | – | KY772802 | |
H. corbierei | epitype, H392, C-F-57133 | Switzerland Graubünden | 1982 | – | – | KY784489 | KY772737 |
H178, O-253273 | Norway, Oppland | 2001 | KY773023 | – | KY784307 | KY772557 | |
H191, O-253274 | Norway, Oppland | 2001 | – | – | KY784317 | KY772569 | |
H232, O-253275 | Switzerland, Graubünden | 1984 | KY773062 | – | KY784350 | KY772599 | |
H346, C-F-56503 | USA, Oregon | 1991 | KY773121 | – | KY784448 | KY772699 | |
H390, C-F-53800 | Norway, Nordland | 1972 | – | – | KY784487 | KY772735 | |
H391, C-F-57087 | Switzerland, Graubünden | 1982 | – | – | KY784488 | KY772736 | |
H. corium | epitype, H352, C-F-71638 | Denmark, Mid Zealand | 1984 | – | – | KY784453 | KY772704 |
H242, O-362201 | Svalbard, Kongsfjord | 1986 | KY773071 | – | KY784360 | KY772609 | |
H248, O-253277 | Norway, Hordaland | 1996 | KY773075 | – | KY784366 | KY772614 | |
H292, C-F-92111 (holotype of H. arctica) | Sweden, Norrbotten | 1928 | KY773099 | – | KY784405 | KY772654 | |
H294, C-F-16568 | Russia, N Ural Mts | 1990 | KY773100 | – | KY784407 | KY772656 | |
H324, C-F-55580 | Svalbard, Bunsow Land | 1985 | KY773111 | – | KY784429 | KY772680 | |
H328, C-F-34476 | Russia, Ural Mts | 1992 | – | – | KY784432 | KY772683 | |
H434, O-253280 | Norway, Troms | 1989 | KY773162 | – | KY784527 | KY772770 | |
H436, O-253281 | Svalbard, Kongsfjord | 1988 | KY773163 | – | KY784528 | KY772771 | |
H451, O-253278 | Norway, Hordaland | 2014 | KY773171 | – | KY784535 | KY772781 | |
H453, O-253279 | Norway, Hordaland | 2014 | KY773173 | – | KY784537 | KY772783 | |
H. costata | H100, FH | USA, California | 1998 | KY772962 | – | KY784244 | KY772491 |
H. costifera | epitype, H298, C-F Fungi Exs. Suec. 2061 | Sweden, Uppland | 1948 | – | – | KY784409 | – |
H131, O-68514 | Norway, Akershus | 2007 | KY772982 | KY772874 | KY784264 | KY772514 | |
H209, O-253282 | Norway, Oppland | 1985 | KY773042 | – | KY784332 | KY772583 | |
H227, O-253285 | Norway, Sør-Trøndelag | 1984 | KY773057 | – | KY784346 | KY772596 | |
H247, O-253283 | Norway, Oppland | 1998 | KY773074 | KY772900 | KY784365 | KY772613 | |
H347, O-253284 | Norway, Oppland | 1982 | KY773122 | – | KY784449 | KY772700 | |
H383, C-F-92113 | Denmark, Zealand | 1984 | – | – | KY784481 | KY772728 | |
H385, C-F-53538 | Norway, Finnmark | 1986 | KY773145 | – | KY784483 | KY772730 | |
H388, C-F-53529 | Norway, Finnmark | 1986 | – | – | KY784485 | KY772733 | |
H. crassitunicata | H222, O-253286 | Canada, British Columbia | 1994 | KY773053 | – | KY784342 | KY772592 |
isoparatype, H299, C-F-92106 | USA, Washington | 1952 | – | – | – | KY772658 | |
H. crispa | epitype, H408, C-F Fungi. Exs. Suec. 2062 | Sweden, Blekinge | 1946 | – | – | KY784504 | KY772750 |
H062, O-253287 | Sweden, Hallan | 2009 | KY772946 | – | KY784226 | KY772472 | |
H105, FH | England, Surrey | 1994 | KY772965 | – | – | KY772495 | |
H107, FH | Latvia, Riga | 1982 | – | – | – | KY772496 | |
H135, O-18774 | Norway, Akershus | 2004 | KY772986 | – | KY784268 | KY772518 | |
H235, O-360158 | Norway, Nordland | 2005 | KY773065 | – | KY784353 | KY772602 | |
H405, C-F Fungi exs. Suec. 3263 | Sweden, Öland | 1960 | – | – | KY784501 | KY772747 | |
H. danica | H058, O-253288 | Norway, Akershus | 2009 | KY772942 | – | KY784222 | KY772469 |
H111, FH | Russia, St. Petersburg | 1982 | – | – | – | KY772499 | |
H177, O-129543 | Norway, Finnmark | 1983 | KY773022 | – | KY784306 | KY772556 | |
H396, C-F-64512 | Denmark, Zealand | 1974 | – | – | KY784493 | – | |
holotype, H263, C-F-85205 | Denmark. Jutland | 2008 | KY773083 | – | KY784378 | KY772627 | |
H. dryadophila | H010, O-253306 | Norway, Oppland | 2007 | – | KY772822 | – | KY772433 |
H180, O-253307 | Norway, Oppland | 1997 | KY773024 | KY772883 | KY784309 | KY772559 | |
H183, O-253308 | Norway, Oppland | 1996 | KY773026 | KY772886 | KY784311 | KY772561 | |
H190, O-253309 | Norway, Oppland | 2005 | – | – | – | KY772568 | |
H234, O-362204 | Svalbard, Kongsfjord | 1986 | KY773064 | – | KY784352 | KY772601 | |
H321, C-F-55579 | Svalbard, Bunsow Land | 1985 | KY773108 | – | KY784426 | KY772677 | |
H322, C-F-55597 | Svalbard, Bunsow Land | 1985 | KY773109 | – | KY784427 | KY772678 | |
H323, C-F-55578 | Svalbard, Bunsow Land | 1985 | KY773110 | – | KY784428 | KY772679 | |
H335, C-F-56772 | Canada, Nunavut, Ellesmere Isl. | 1984 | KY773115 | – | KY784438 | KY772689 | |
H378, C-F-52701 | Greenland, Mestervig | 1882 | KY773140 | – | KY784477 | KY772725 | |
H379, C-F-50346 | Greenland, Ella Isl. | 1982 | KY773141 | – | KY784478 | – | |
holotype, H302, O-72617 | Norway, Oppland | 1957 | – | – | KY784412 | KY772661 | |
H. elastica | epitype, H372, C-F-29130 | Sweden, Uppland | 1946 | – | – | KY784471 | – |
H066, O-253311 | Sweden, Hallan | 2009 | KY772950 | KY772858 | KY784230 | KY772476 | |
H329, C-F-51672 | Russia, Sakha | – | – | – | KY784433 | KY772684 | |
H414, C-F Fungi Exs. Suec. 271 | Sweden, Uppland | 1935 | – | – | KY784509 | – | |
H443, O-253310 | Norway, Akershus | 2009 | KY773167 | – | KY784531 | KY772775 | |
H. ephippioides | H085, O-253267 | Japan, Honshu | 1983 | KY772957 | KY772867 | KY784237 | KY772484 |
H. fallax | epitype, H339, C-F-84621 | France, Savoie | 1992 | KY773117 | – | KY784442 | KY772693 |
H002, O-253350 | Norway, Oppland | 2001 | KY772901 | – | – | KY772429 | |
H018, O-253351 | Norway, Oppland | 2009 | KY772913 | KY772830 | KY784195 | KY772439 | |
H032, O-253352 | Norway, Oppland | 2009 | KY772923 | – | – | KY772452 | |
H162, O-280610 | Norway, Hedmark | 2006 | KY773009 | – | KY784292 | KY772543 | |
H163, O-242201 | Norway, Møre og Romsdal | 2001 | KY773010 | – | KY784293 | – | |
H168, O-63684 | Norway, Hedmark | 1998 | KY773015 | – | KY784298 | KY772548 | |
H169, O-66873 | Norway, Oppland | 2002 | KY773016 | – | KY784299 | KY772549 | |
H219, O-253348 | France, Savoie | 1992 | KY773050 | – | KY784339 | – | |
H221, O-253349 | France, Savoie | 1992 | KY773052 | – | KY784341 | – | |
H244, O-360225 | Norway, Troms | 2005 | KY773072 | – | KY784362 | KY772611 | |
H256, O-253353 | Norway, Oppland | 1996 | KY773078 | – | KY784372 | KY772621 | |
H395, C-F-45484 | Sweden, Lule Lappmark | 1970 | – | – | KY784492 | KY772740 | |
H402, O-253354 | Norway, Oppland | 1985 | KY773150 | – | KY784498 | KY772745 | |
H419, C-F-45301 | Norway, Nordland | 1979 | – | – | KY784514 | KY772757 | |
H428, C-F-57377 | Switzerland, Graubünden | 1984 | – | – | KY784522 | KY772765 | |
H430, C-F-92108 | Canada, NW Territories | 1974 | KY773160 | – | KY784524 | KY772767 | |
H. fibrosa | H139, O-63973 | Norway, Hedmark | 2000 | – | – | KY784272 | KY772522 |
H164, O-88570 | Norway, Akershus | 1988 | KY773011 | – | KY784294 | KY772544 | |
H166, O-185923 | Norway, Vestfold | 1985 | KY773013 | – | KY784296 | KY772546 | |
H167, O-185919 | Norway, Vestfold | 1985 | KY773014 | – | KY784297 | KY772547 | |
H240, O-291352 | Norway, Sør-Trøndelag | 2008 | KY773069 | KY772898 | KY784358 | KY772607 | |
H243, O-291377 | Norway, Sør-Trøndelag | 2009 | – | KY772899 | KY784361 | KY772610 | |
H250, O-253313 | Georgia, Kaukasus | 1988 | – | – | KY784367 | KY772615 | |
H270, C-F-54475 | Norway, Nordland | 1975 | – | – | KY784385 | KY772633 | |
H343, C-F-92103 | Iceland, Merkurhraun | 1984 | – | – | KY784446 | KY772697 | |
H344, C-F-92102 | Iceland, Borgarfjördur | 1984 | – | – | KY784447 | KY772698 | |
H357, C-F-53774 | Norway, Nordland | 1972 | KY773125 | – | KY784458 | KY772708 | |
H369, C-F-92109 | Finland, Oulun Pohjanmaa | 1981 | KY773136 | – | KY784469 | KY772719 | |
H426, C-F-38089 | Denmark, Falster | 1998 | KY773158 | – | KY784520 | KY772763 | |
isoepitype, H413, C-F Fungi Exs. Suec 3262 | Sweden, Halland | 1956 | – | – | KY784508 | – | |
H. fistulosa | H109, FH | USA, Massachusetts | 1999 | KY772966 | – | KY784248 | KY772497 |
H114, FH | USA, Vermont | 2004 | – | – | KY784252 | – | |
H179, O-253315 | Norway, Akershus | 2001 | – | – | KY784308 | KY772558 | |
H205, O-253314 | Japan, Honshu | 1983 | KY773039 | – | KY784329 | KY772580 | |
H476, C | Iceland, Austur-Skaftafellsysla | 1981 | – | – | KY784556 | – | |
neotype, H241, O-291887 | Norway, Hordaland | 2009 | KY773070 | – | KY784359 | KY772608 | |
H. fusca | H304, C-F-89381 | Netherlands, Zuid-Holland | 1983 | – | – | KY784414 | KY772663 |
H305, C-F-92122 | Hungary, Kiskunsag National Park | 1978 | KY773101 | – | KY784415 | KY772664 | |
H. griseoalba | H306, C-F-92112 | USA, Michigan | 1973 | – | – | KY784416 | KY772665 |
H. helvellula | epitype, H308, C-F-92128 | Spain, Canary Islands, Hierro | 1977 | – | – | KY784418 | KY772667 |
H278, C-F-45507 | France, Manche | 1977 | KY773090 | – | KY784393 | KY772641 | |
H309, C-F-45469 | France, Region des Landes | 1981 | KY773103 | – | KY784419 | KY772668 | |
H. hyperborea | H389, C-F-54473 | Norway, Nordland | 1975 | – | – | KY784486 | KY772734 |
H491, C-F-45306 | Finland, Kuusamo | 1978 | – | – | KY784569 | KY772812 | |
H. hypocrateriformis | epitype, H301, C-F-45379 | Sweden, Uppland | 1948 | – | – | KY784411 | KY772660 |
H275, C-F-57126 (isotype of H. pulchra) | Switzerland, Graubünden | 1982 | – | – | KY784390 | KY772638 | |
H300, C-F-92131 (isotype of H. cupuliformis) | Sweden, Uppland | 1936 | – | – | KY784410 | KY772659 | |
H. juniperi | H415, C-F-42193 | Portugal, Minho | 1996 | – | – | KY784510 | KY772754 |
H. lactea | H262, C-F Fungi. Exs. Suec. 1355 | Sweden, Uppland | 1939 | – | – | KY784377 | KY772626 |
H374, C-F-39379 | Denmark, Zealand | 1961 | – | – | KY784473 | KY772722 | |
H. lacunosa | epitype, H407, C-F Fungi Exs. Suec. 2065 | Sweden, Femsjö | 1948 | KY773152 | – | KY784503 | KY772749 |
H039, O-253319 | Norway, Oppland | 2007 | KY772930 | KY772845 | KY784213 | – | |
H044, O-253320 | Norway, Oppland | 2009 | – | – | – | KY772459 | |
H045, O-253321 | Norway, Oppland | 2009 | KY772934 | – | KY784215 | KY772460 | |
H059, O-253316 | Norway, Akershus | 2009 | KY772943 | – | KY784223 | KY772470 | |
H150, O-280703 | Norway, Hordaland | 2005 | KY772999 | – | KY784283 | KY772533 | |
H153, O-285214 | Norway, Sør-Trøndelag | 2007 | KY773002 | – | KY784285 | KY772536 | |
H170, O-59885 | Norway, Finnmark | 1995 | KY773017 | – | KY784300 | KY772550 | |
H189, O-253322 | Norway, Oppland | 2001 | KY773032 | – | KY784316 | KY772567 | |
H327, C-F-34477 | Russia, Krasnoyarsk Krai | 1993 | KY773112 | – | KY784431 | KY772682 | |
H399, C-F-55985 | Greenland, Thule | 1987 | KY773149 | – | KY784496 | KY772743 | |
H470, O-253318 | Norway, Hordaland | 2014 | – | – | KY784551 | KY772798 | |
H471, C-F-19329 | Svalbard, Longyearbyen | 1988 | – | – | KY784552 | KY772799 | |
H474, C-F-53539 | Norway, Finnmark | 1986 | – | – | KY784554 | KY772801 | |
H486, O-253317 | Norway, Finnmark | 1986 | – | – | KY784565 | KY772808 | |
H. leucomelaena | epitype, H404, C-F Fungi Exs. Suec. 952 | Sweden, Gotland | 1938 | – | – | KY784500 | – |
H115, FH | USA, Massachusetts | 2006 | KY772970 | KY772870 | KY784253 | JX9437513 | |
H267, C-F-39492 | Denmark, Jutland | 1967 | – | – | KY784382 | KY772630 | |
H447, C-F-92127 | Chile, Santiago | 1966 | KY773168 | – | KY784533 | KY772777 | |
H450, C-F-38142 | Denmark, Zealand | 1998 | KY773170 | – | KY784534 | KY772780 | |
H. leucophaea | H274, C-F-45487 | France, Porquerelles | 1976 | KY773088 | – | KY784389 | KY772637 |
H. levis | epitype, H290, C-F-55188 | Denmark, Mid Zealand | 2001 | KY773098 | – | KY784403 | KY772652 |
H264, C-F-39370 | Denmark, Mid Zealand | 1968 | – | – | KY784379 | KY772627 | |
H265, C-F-39366 | Denmark, Jutland | 1979 | – | – | KY784380 | KY772628 | |
H266, C-F-39367 | Denmark, Zealand | 1974 | – | – | KY784381 | KY772629 | |
H393, C-F-40830 | Denmark, East Jutland | 2002 | KY773148 | – | KY784490 | KY772738 | |
H458, C | Denmark, NE Zeland | 2013 | – | – | KY784540 | KY772787 | |
H469, C-F-52988 | Denmark, NE Zealand | 1997 | – | – | KY784550 | KY772797 | |
H480, C-F-45517 | Germany, Hamburg | 1973 | – | – | KY784559 | – | |
H. macropus | epitype, H412, C-F Fungi Exs. Suec. 3266 | Sweden, Uppland | 1960 | – | – | KY784507 | KY772753 |
H060, O-253323 | Norway, Buskerud | 2009 | KY772944 | KY772853 | KY784224 | – | |
H069, O-292075 | Norway, Akershus | 2009 | – | KY772861 | KY784231 | KY772478 | |
H073, O-253326 | Norway, Oslo | 2009 | KY772954 | KY772863 | KY784233 | KY772480 | |
H118, FH | USA, Massachusetts | 2006 | KY772972 | – | – | KY772503 | |
H119, FH | Canada, Alberta | 1996 | KY772973 | KY772871 | KY784255 | KY772504 | |
H120, FH | Canada, Quebec | 1997 | KY772974 | – | KY784256 | KY772505 | |
H124, FH | USA, Massachusetts | 1998 | KY772977 | – | – | KY772508 | |
H141, O-168819 | Norway, Telemark | 2000 | KY772990 | – | KY784274 | KY772524 | |
H142, O-166155 | Norway, Telemark | 2004 | KY772991 | – | KY784275 | KY772525 | |
H157, O-370348 | Norway, Akershus | 2006 | KY773004 | – | KY784287 | KY772539 | |
H158, O-65347 | Norway, Hedmark | 2002 | KY773005 | – | KY784288 | – | |
H159, O-220323 | Norway, Hordaland | 2001 | KY773006 | – | KY784289 | KY772540 | |
H160, O-283072 | Norway, Hordaland | 2006 | KY773007 | – | KY784290 | KY772541 | |
H161, O-223871 | Norway, Møre og Romsdal | 2003 | KY773008 | – | KY784291 | KY772542 | |
H165, O-220225 | Norway, Møre og Romsdal | 2001 | KY773012 | – | KY784295 | KY772545 | |
H228, O-253324 | Norway, Oppland | 1980 | KY773058 | – | KY784347 | – | |
H238, O-291425 | Norway, Rogaland | 2009 | KY773067 | KY772896 | KY784356 | KY772605 | |
H239, O-291391 | Norway, Rogaland | 2009 | KY773068 | KY772897 | KY784357 | KY772606 | |
H257, O-253325 | Norway, Oppland | 2009 | KY773079 | – | KY784373 | KY772622 | |
H260, O-253327 | Norway, Østfold | 2013 | KY773081 | – | KY784375 | KY772624 | |
H351, C-F-56087 | USA, Alaska | 1980 | – | – | KY784452 | KY772703 | |
H. macrosperma | H029, O-253328 | Norway, Oppland | 2007 | KY772921 | KY772838 | KY784205 | KY772449 |
H047, O-253329 | Norway, Oppland | 2009 | KY772936 | KY772850 | KY784217 | KY772462 | |
H050, O-253330 | Norway, Oppland | 2009 | KY772938 | KY772852 | – | KY772464 | |
H053, O-253331 | Norway, Oppland | 2009 | KY772939 | – | – | KY772466 | |
H. maculata | H303, C-F-45400 | USA, Alaska | 1973 | – | – | KY784413 | KY772662 |
H. monachella | epitype, H268, C-F-92121 | Hungary, Szelid | 1979 | – | – | KY784383 | KY772631 |
H269, C-F-92120 | Hungary, Szelid | 1965 | KY773084 | – | KY784384 | KY772632 | |
H. nannfeldtii | H017, O-253337 | Norway, Oppland | 2007 | KY772912 | KY772829 | KY784194 | KY772438 |
H028, O-253339 | Norway, Oppland | 2007 | KY772920 | KY772837 | KY784204 | KY772448 | |
H188, O-253340 | Norway, Oppland | 2005 | KY773031 | – | KY784315 | KY772566 | |
H212, O-253332 | France, Savoie | 1992 | KY773044 | KY772888 | – | KY772585 | |
H216, O-253333 | France, Savoie | 1992 | KY773048 | KY772891 | KY784337 | KY772589 | |
H254, O-253341 | Norway, Oppland | 1996 | KY773076 | – | KY784370 | KY772619 | |
H318, C-F-92123 | Norway, Nordland | 1974 | KY773106 | – | KY784424 | KY772674 | |
H362, O-253342 | Norway, Oppland | 1996 | KY773130 | – | KY784463 | KY772713 | |
H367, O-253343 | Norway, Oppland | 1985 | KY773134 | – | KY784467 | KY772717 | |
H452, O-253334 | Norway, Hordaland | 2014 | KY773172 | – | KY784536 | KY772782 | |
H454, O-253335 | Norway, Hordaland | 2014 | KY773174 | – | KY784538 | KY772784 | |
H457, O-253336 | Norway, Hordaland | 2014 | – | – | KY784539 | KY772786 | |
H479, O-253344 | Norway, Oppland | 1983 | – | – | KY784558 | KY772804 | |
holotype, H027, O-253338 | Norway, Oppland | 2009 | KY772919 | KY772836 | KY784203 | KY772447 | |
H. nigra | epitype, H063, O-253345 | Sweden, Hallan | 2009 | KY772947 | KY772855 | KY784227 | KY772473 |
H067, O-253346 | Sweden, Hallan | 2009 | KY772951 | KY772859 | – | – | |
H068, O-253347 | Sweden, Hallan | 2009 | KY772952 | KY772860 | – | KY772477 | |
H1029, O-253404 | Spain, La Rioha | 2014 | – | – | KY784573 | KY772815 | |
H1030, O-253405 | Spain, Basque Country | 2007 | – | – | KY784572 | KY772816 | |
H. oblongispora | H132, O-166316 | Norway, Oppland | 2004 | KY772983 | – | KY784265 | KY772515 |
H332, C-F-56844 | Switzerland, Graubünden | 1979 | – | – | – | KY772687 | |
H448, C-F-56914 | Switzerland, Graubünden | 1979 | KY773169 | – | – | KY772778 | |
H. pallescens | epitype, H138, O-66205 | Norway, Møre og Romsdal | 2003 | KY772988 | KY772878 | KY784271 | KY772521 |
H070, O-289039 | Norway, Møre og Romsdal | 2003 | KY772953 | KY772862 | KY784232 | KY772479 | |
H136, O-220306 | Norway, Hordaland | 2001 | KY772987 | KY772877 | KY784269 | KY772519 | |
H236, O-291458 | Norway, Rogaland | 2009 | – | KY772895 | KY784354 | KY772603 | |
H373, O-253355 | Norway, Østfold | 1992 | KY773137 | – | KY784472 | KY772721 | |
H. palustris | H001, O-253357 | Norway, Oppland | 2001 | – | KY772817 | KY784182 | KY772428 |
H019, O-253358 | Norway, Oppland | 2009 | KY772914 | KY772831 | KY784196 | KY772440 | |
H043, O-253359 | Norway, Oppland | 2009 | KY772933 | KY772848 | KY784214 | KY772458 | |
H181, O-253360 | Norway, Oppland | 1996 | KY773025 | KY772884 | – | KY772560 | |
H184, O-253361 | Norway, Oppland | 1996 | KY773027 | KY772887 | – | KY772562 | |
H195, O-253356 | Japan, Honshu | 1983 | KY773033 | – | KY784319 | – | |
H350, C-F-55330 | Finland, Kuusamo | 1978 | KY773123 | – | KY784451 | KY772702 | |
H468, O-253362 | Norway, Oppland | 1982 | – | – | KY784549 | KY772796 | |
H. panormitana | epitype, H064, O-253363 | Sweden, Hallan | 2009 | KY772948 | KY772856 | KY784228 | KY772474 |
H130, O-171969 | Norway, Møre og Romsdal | 2001 | KY772981 | – | KY784263 | KY772513 | |
H137, O-284515 | Norway, Hordaland | 2007 | – | – | KY784270 | KY772520 | |
H143, O-65394 | Norway, Hordaland | 2002 | KY772992 | – | KY784276 | KY772526 | |
H144, O-178718 | Norway, Møre og Romsdal | 2003 | KY772993 | – | KY784277 | KY772527 | |
H145, O-203499 | Norway, Møre og Romsdal | 2005 | KY772994 | – | KY784278 | KY772528 | |
H154, O-167560 | Norway, Hordaland | 2005 | KY773003 | – | KY784286 | KY772537 | |
H237, O-360894 | Norway, Møre og Romsdal | 2006 | KY773066 | – | KY784355 | KY772604 | |
H. paraphysitorquata | isotype, H271, C-F-45305 | Spain, Teruel | 1988 | KY773085 | – | KY784386 | KY772634 |
H. pezizoides | epitype, H061, O-253366 | Sweden, Hallan | 2009 | KY772945 | KY772854 | KY784225 | KY772471 |
H065, O-253367 | Sweden, Hallan | 2009 | KY772949 | KY772857 | KY784229 | KY772475 | |
H196, O-253365 | Norway, Vestfold | 1981 | – | – | KY784320 | – | |
H204, O-253364 | Japan, Honshu | 1983 | KY773038 | – | KY784328 | KY772579 | |
H230, C-F ex GZU109.86 | Nepal, Langtang | 1986 | KY773060 | – | – | – | |
H418, C-F-45301 | Denmark, Jutland | 1968 | – | – | KY784513 | – | |
H420, C-F-45505 | Norway, Vestfold | 1981 | – | – | KY784515 | KY772758 | |
H431, C-F-52986 | Denmark N Jutland | 1997 | KY773161 | – | KY784525 | KY772768 | |
H. philonotis | H005, O-253294 | Norway, Oppland | 2001 | KY772903 | KY772820 | KY784185 | – |
H006, O-253295 | Norway, Oppland | 2007 | KY772904 | KY772821 | KY784186 | – | |
H015, O-253296 | Norway, Oppland | 2007 | KY772910 | KY772827 | KY784192 | KY772436 | |
H023, O-253297 | Norway, Oppland | 2009 | KY772916 | KY772834 | KY784199 | KY772443 | |
H037, O-253298 | Norway, Oppland | 2007 | KY772928 | KY772843 | KY784211 | – | |
H038, O-253299 | Norway, Oppland | 2007 | KY772929 | KY772844 | KY784212 | – | |
H048, O-253300 | Norway, Oppland | 2009 | KY772937 | KY772851 | KY784218 | KY772463 | |
H051, O-253301 | Norway, Oppland | 2009 | – | – | KY784219 | KY772465 | |
H193, O-253305, holotype of H. dovrensis | Norway, Oppland | 1989 | – | – | KY784318 | KY772571 | |
H198, O-253302 | Norway, Oppland | 1989 | – | – | KY784322 | KY772573 | |
H199, O-253303 | Norway, Oppland | 1989 | KY773035 | – | KY784323 | KY772574 | |
H200, O-253304 | Norway, Oppland | 1989 | – | – | KY784324 | KY772575 | |
H214, O-253291 | France, Savoie | 1992 | KY773046 | KY772889 | KY784335 | KY772587 | |
H218, O-253292 | France, Savoie | 1992 | KY773049 | KY772892 | KY784338 | KY772590 | |
H220, O-253293 | France, Savoie | 1992 | KY773051 | KY772893 | KY784340 | KY772591 | |
H361, C-F-57376 | Switzerland, Graubünden | 1984 | KY773129 | – | KY784462 | KY772712 | |
H381, O-253289 | Austria, Tirol | 1990 | KY773142 | – | KY784479 | KY772726 | |
H475, O-253290 | Austria, Tirol | 1990 | – | – | KY784555 | – | |
holotype, H492, C-F-45481 | Iceland, Hveragerdi | 1959 | – | – | KY784570 | KY772813 | |
paratype, H272, C-F-45486 | Iceland, Grøndalur | 1959 | KY773086 | – | KY784387 | KY772635 | |
H. phlebophora | H197, O-253368 | Japan, Honshu | 1983 | KY773034 | – | KY784321 | KY772572 |
H273, C-F-45405 | Iceland, Austur Skaftafellsysla | 1981 | KY773087 | – | KY784388 | KY772636 | |
H. pubescens | H251, O-253369 | Norway, Østfold | 1986 | – | – | KY784368 | KY772616 |
H333, C-F-56499 | USA, Oregon | 1991 | – | – | KY784436 | – | |
H334, C-F-56502 | USA, Oregon | 1991 | KY773114 | – | KY784437 | KY772688 | |
holotype, H121, FH | Canada, Alberta | 1994 | KY772975 | – | KY784257 | KY772506 | |
H. pulla | epitype, H149, O-69282 | Norway, Møre og Romsdal | 2008 | KY772998 | – | KY784282 | KY772532 |
H259, O-253370 | Norway, Østfold | 2013 | KY773080 | – | KY784374 | KY772623 | |
H. queletiana | neotype, H403, C-F-45303 | Denmark, Zealand | 1995 | KY773151 | – | KY784499 | KY772746 |
H. rivularis | H025, O-253380 | Norway, Oppland | 2009 | KY772918 | – | KY784201 | KY772445 |
H245, O-253381 | Norway, Oppland | 1996 | – | – | KY784363 | KY772612 | |
H312, C-F-86705 | Norway, Nordland | 1975 | – | – | KY784420 | KY772669 | |
H319, C-F-92124 | Norway, Nordland | 1972 | – | – | KY784425 | KY772675 | |
H363, O-253382 | Norway, Oppland | 2001 | KY773131 | – | KY784464 | KY772714 | |
H365, O-253383 | Norway, Oppland | 1983 | KY773132 | – | KY784465 | KY772715 | |
H366, O-253384 | Norway, Oppland | 1983 | KY773133 | – | KY784466 | KY772716 | |
H368, O-253385 | Norway, Oppland | 1984 | KY773135 | – | KY784468 | KY772718 | |
H424, O-253386 | Norway, Oppland | 1982 | KY773157 | – | KY784519 | KY772762 | |
holotype, H276, C-F-59447 | Norway, Nordland | 1972 | KY773089 | – | KY784391 | KY772639 | |
H. scyphoides | holotype, H140, O-65348 | Norway, Hedmark | 2002 | KY772989 | KY772879 | KY784273 | KY772523 |
H. semiobruta | H307, C-F-45467 | Spain, Valdepenas | 1975 | KY773102 | – | KY784417 | KY772666 |
isotype, H277, C-F-45507 | France, Rhone | 1972 | – | – | KY784392 | KY772640 | |
H. silvicola | H281, C-F-60841 | Kyrgyz Republic, Tianshan Interior | 1967 | – | – | KY784395 | KY772644 |
H282, C-F-57385 | Switzerland, Graubünden | 1984 | KY773092 | – | KY784396 | KY772645 | |
H283, C-F-56847 | Switzerland, Graubünden | 1979 | KY773093 | – | KY784397 | KY772646 | |
H489, O-253387 | Switzerland, Graubünden | 1984 | – | – | KY784568 | KY772811 | |
H. solitaria | epitype, H370, C-F Fungi Exs. Suec. 3267 | Sweden, Uppsala | 1942 | – | – | KY784470 | KY772720 |
H004, O-253374 | Norway, Oppland | 2001 | KY772902 | KY772819 | KY784184 | KY772430 | |
H021, O-253375 | Norway, Oppland | 2009 | – | KY772832 | KY784197 | KY772441 | |
H080, O-253376 | Norway, Oppland | 1989 | – | KY772864 | – | – | |
H090, O-253371 | France, Savoie | 1992 | KY772959 | KY772868 | KY784241 | KY772488 | |
H122, FH | Canada, Alberta | 1996 | KY772976 | – | KY784258 | KY772507 | |
H171, O-129536 | Norway, Akershus | 1987 | KY773018 | – | KY784301 | KY772551 | |
H174, O-129533 | Norway, Sør-Trøndelag | 1987 | KY773019 | – | KY784303 | KY772553 | |
H176, O-58891 | Norway, Oslo | 1996 | KY773021 | – | KY784305 | KY772555 | |
H186, O-253377 | Norway, Oppland | 2005 | KY773029 | – | KY784313 | KY772564 | |
H187, O-253378 | Norway, Oppland | 2005 | KY773030 | – | KY784314 | KY772565 | |
H215, O-253379 | Norway, Sør-Trøndelag | 1985 | KY773047 | KY772890 | KY784336 | KY772588 | |
H252, O-253372 | Georgia, Kaukasus | 1988 | – | – | KY784369 | KY772617 | |
H314, C-F-54683 | Norway, Nordland | 1981 | KY773105 | – | KY784421 | KY772671 | |
H315, C-F-54680 | Norway, Nordland | 1981 | – | – | KY784422 | KY772672 | |
H316, C-F-54682 | Norway, Nordland | 1981 | – | – | KY784423 | KY772673 | |
H326, C-F-34410 | Russia, Khatanga | 1993 | – | – | KY784430 | KY772681 | |
H330, C-F-57390 | Switzerland, Graubünden | 1984 | – | – | KY784434 | KY772685 | |
H340, C-F-34316 | Iceland, Sudur-Mulasysla | 1993 | KY773118 | – | KY784443 | KY772694 | |
H341, C-F-34321 | Iceland, Nordur-Mulasysla | 1993 | KY773119 | – | KY784444 | KY772695 | |
H353, O-129541 | Norway, Oppland | 1957 | – | – | KY784454 | KY772705 | |
H358, O-253373 | Norway, Hedmark | 1980 | KY773126 | – | KY784459 | KY772709 | |
H. sp. ‘INDIA’ | H421, C-F-45704 | India, Udhampur | 1967 | – | – | KY784516 | KY772759 |
H. sp. ‘JAPAN 1’ | H206, O-253388 | Japan, Honshu | 1983 | KY773040 | – | KY784330 | KY772581 |
H. sp. ‘JAPAN 2’ | H093, O-253389 | Japan, Honshu | 1983 | KY772961 | – | KY784243 | KY772490 |
H. sp. ‘JAPAN 3’ | H213, O-253390 | Japan, Honshu | 1991 | KY773045 | – | KY784334 | KY772586 |
H. sp. ‘JAPAN 4’ | H461, O-253391 | Japan, Honshu | 1983 | – | – | KY784543 | KY772790 |
H478, O-253392 | Japan, Honshu | 1983 | – | – | KY784557 | KY772803 | |
H. sp. ‘JAPAN 5’ | H466, O-253393 | Japan, Honshu | 1983 | – | – | KY784547 | KY772794 |
H. sp. ‘JAPAN 6’ | H482, O-253394 | Japan, Honshu | 1983 | – | – | KY784561 | KY772806 |
H. sp. ‘PAPUA 1’ | H432, C-F-45531 | Papua New Guinea, Manki | 1972 | – | – | KY784526 | KY772769 |
H. sp. ‘PAPUA 2’ | H400, C-F-45434 | Papua New Guinea, Manki | 1971 | – | – | KY784497 | KY772744 |
H. sp. ‘USA 1’ | H416, C-F-58799 | USA, New York | 1989 | – | – | KY784511 | KY772755 |
H. sp. ‘USA 2’ | H103, FH | USA, California | 1988 | – | – | KY784246 | KY772493 |
H. sp. ‘USA 3’ | H376, C-F-92118 | USA, Idaho | 1972 | – | – | KY784475 | – |
H. sp. ‘USA 4’ | H104, FH | USA, Massachusetts | 1998 | KY772964 | KY772869 | KY784247 | KY772494 |
H. sp. ‘USA 5’ | H110, FH | USA, Iowa | 2008 | KY772967 | – | KY784249 | KY772498 |
H. sp. ‘USA 6’ | H291, C-F-92104 | USA, Michigan | 1979 | – | – | KY784404 | KY772653 |
H. sp. ‘USA 7’ | H377, C-F-92119 | USA, Michigan | 1940 | KY773139 | – | KY784476 | KY772724 |
H. sp. ‘USA 8’ | H429, C-F-92105 | USA, Michigan | 1956 | – | – | KY784523 | KY772766 |
H. stevensii | H297, C-F-45350, isotype of H. connivens | USA, Michigan | 1947 | – | – | KY784408 | KY772657 |
H394, C-F-92117 | USA, Michigan | 1969 | – | – | KY784491 | KY772739 | |
H. subglabra | topotype, H280, C-F-65405 | USA, Michigan | 1972 | – | – | KY784394 | KY772643 |
H. sublicia | epitype, H417, C-F-39823 | Denmark, Zealand | 1993 | KY773155 | – | KY784512 | KY772756 |
H148, O-70080 | Norway, Akershus | 1994 | KY772997 | KY772880 | KY784281 | KY772531 | |
H229, O-253312 | Norway, Akershus | 1988 | KY773059 | – | KY784348 | KY772597 | |
H422, C-F-39826 | Denmark, Zealand | 1993 | – | – | KY784517 | KY772760 | |
H423, C-F-53175 | Denmark, Zealand | 1999 | KY773156 | – | KY784518 | KY772761 | |
H. sulcata | epitype, H152, O-68095 | Norway, Oslo | 2006 | KY773001 | KY772882 | KY784284 | KY772535 |
H1027, O-253406 | Spain, La Rioja | 2013 | – | – | KY784571 | KY772814 | |
H. terrestris | H284, C | Norway, Nord Trøndelag | 1983 | KY773094 | – | KY784398 | KY772647 |
H. vespertina | H102, FH | USA, California | 2013 | KY772963 | – | KY784245 | KY772492 |
H116, FH | USA, Maine | 2000 | KY772971 | – | KY784254 | KY772502 |
1 C-F = Copenhagen Fungal Herbarium, Natural History Museum of Denmark, University of Copenhagen; O = Mycological Herbarium, Natural History Museum, University of Oslo; FH = Farlow Herbarium, Harvard University Herbaria and Libraries, Harvard University.
2 GenBank accession numbers are provided for the following markers: partial 28S large ribosomal sub unit (LSU), translation elongation factor 1-alpha (tef), heat shock protein 90 (hsp) and RNA polymerase II (rpb2). Sequences included in the three-gene alignment are indicated in bold. Sequences included in the two-gene alignment of hsp and rpb2 are indicated in italics. Sequences included in both alignments are in bold italics.
3 Sequences obtained from GenBank.
Phylogenetic inference
DNA sequences from each locus were aligned separately using MAFFT v. 7.017 (Katoh & Standley 2013) within Geneious R9.0. The alignments were then improved manually. Alignments of each partition were analysed individually by Maximum Likelihood (ML) and Bayesian Inference phylogenetic methods, as implemented in RAxML 7.2.8 (Stamatakis 2006) and MrBayes 3.2.2 (Ronquist & Huelsenbeck 2003) in Geneious. For the ML analyses, the GTRCAT approximation was used for each partition. Bootstrap analyses were performed with 1 000 pseudoreplicates. PartitionFinder v. 1.1.1 was used to find the best-fit models for Bayesian analyses (Lanfear et al. 2012). Bayesian inferences were performed using MrBayes, applying the GTR + GAMMA model for hsp and rpb2 and the GTR + GAMMA + I model for LSU. Two independent runs, each with three cold and one heated Markov Chain Monte Carlo chains, were started from a random staring tree. The MCMC chains lasted for four million generations, saving trees every 1 000th generation. The posterior probabilities (BPP) were calcucated after burn-in phase (400 trees), which was determined from the marginal likelihood scored of the initially sampled trees. The average split frequencies of the two runs were < 0.01, indicating the convergence of the MCMC chains.
In order to obtain a balanced dataset for the phylogenetic analysis we attempted to include 2–3 individuals for each evolutionary unit. Thus, from the initial alignments and phylogenetic trees of all specimens we pruned the dataset, keeping as much as possible genetic and geographic diversity.
The continental distribution of species based on our included sample was visualized in a Venn diagram (Fig. 1) using the online tool Venny (Oliveros 2007–2015).
To understand and compare the diversity of the various species groups we analysed the average pairwise within-species and within-groups-of-species-diversity for each locus using Mega 6.06 (Tamura et al. 2013). These analyses were based on the same specimens as included in the final phylogenetic analyses. In order to further compare the phylogenetic informativeness (PI) of the different loci we produced a phylogenetic informativeness profile for each locus using the online tool PhyDesign (López-Giráldez & Townsend 2011). For these analyses an ultrametric tree based on the ML tree of three loci (hsp, rpb2 and LSU) was constructed in the R package Ape (Paradis 2012, Paradis et al. 2004, R Core Team 2014). Separate analyses including more specimens from selected species groups were also analysed.
Nomenclature
Roughly 170 valid species names are available to Helvella spp. when excluding subspecific epithets (cf. Index Fungorum, http://www.indexfungorum.org/).
Taking into account the many new phylogenetic species that have been recognised among the earlier inferred morphospecies complexes of Helvella in this synopsis, we assume that many heterotypic synonyms listed in Helvella literature may prove to be ‘good’ species if their taxonomic status can be re-assessed based on adequate type studies. As a consequence, we have been reluctant to accept early heterotypic synonyms of species if they are not resting on morphological and molecular comparisons of type specimens. At the end, we have compiled a list of 108 binomials not treated in this synopsis but referred to or retained in Helvella or its segregate genera by previous authors and also suspected by us to represent good species or taxonomic synonyms of Helvella species (Appendix 1).
It is against this background we typify the European names in designating lectotypes and epitypes, or neotypes, for many early Helvella names where the original material is an illustration (iconotype), does not exist, is poorly preserved, or will be much too old to yield useful DNA sequence data for an identification (cf. Richard et al. 2015). For epitype designations we use the criterion that the epitype shall originate from the original continent (and preferably the same climatic zone and habitat) as judged from the protologue, and that DNA sequence data should have been obtained from at least one protein-coding locus.
RESULTS
Sequence amplification and assembly
Four loci, which included a 515 bp region of the protein-coding gene tef, a 269 bp region of the protein-coding gene hsp, a 347 bp region of the protein-coding gene rpb2 (including the 7–11 regions) and a 697 bp fragment at the 5’ end of LSU (including the D1 and D2 regions), were sampled to delimitate phylogenetic species of Helvella.
Amplicons were not produced for all taxa. Hsp and rpb2 were the most successful single-copy protein coding loci obtained from DNA extractions of old, dried fungarium material. The tef primers worked well on DNA extractions from freshly collected material but failed to regularly produce amplicons from DNA extracted from old fungarium material. For this reason tef was excluded from our ‘whole set’ analyses. Although, we would like to emphasize that an analysis of a small alignment (88 accessions of 26 taxa, 515 bp) of tef produced a tree that was concordant with the results of the other three loci (analyses not shown).
Based on initial analyses of hsp and rpb2 sequences of a collection of 432 Helvella specimens, we selected 183 specimens, which together represented the full range of genetic diversity sampled, as representative for all independently evolving units in our dataset.
For the final analyses we produced two alignments: one including all species for the concatenated dataset of hsp and rpb2 – consisting of 178 accessions and 616 bp –, and a second alignment including all species with an LSU sequence for the concatenated dataset of hsp, rpb2 and LSU consisting of 118 accessions and 1 314 bp. All alignments were submitted to Dryad (http://datadryad.org/).
Phylogenies derived from the separate hsp, tef, rpb2 and LSU datasets did not exhibit any incongruence. Incongruence was evaluated by presence of conflicting nodes with bootstrap support above 70 in the ML trees. Trees constructed from the combined four locus –, three locus – and two locus datasets produced a topology similar to those constructed from individual marker datasets. Nevertheless, the four loci discriminated clades at somewhat different level. Hsp and rpb2 consistently yielded high levels of species discrimination power in all clades and in a multilocus combination were the two most informative loci for inferring species-level relationships/lineages in Helvella. LSU provided more variable levels of resolution in the different clades. For instance, Helvella palustris consistently formed a distinct clade with H. philonotis in the hsp, rpb2 and tef gene trees, but showed no resolution in the LSU rDNA gene tree.
The phylogenetic Informativeness (PI) analyses (Appendix 2) and the diversity measures are presented in Table 3. This demonstrated that LSU had the highest PI overall and per site (Appendix 2). Moreover, LSU showed high PI that dropped quickly at a more recent time. Hsp and rpb2 evolved at a more similar rate than LSU, as reflected in the PI values and diversity measures, and were informative for a wider evolutionary time span.
Table 3
LSU1 | LSU1 | hsp1 | hsp1 | rpb21 | rpb21 | two-gene tree3 | two-gene tree3 | three-gene tree3 | three-gene tree3 | |
---|---|---|---|---|---|---|---|---|---|---|
Clades and lineages | all2 | per site2 | all2 | per site2 | all2 | per site2 | BP4 | BPP4 | BP4 | BPP4 |
entire alignment | 22.0 | 0.044 | 16.4 | 0.061 | 19.0 | 0.064 | ||||
without outgroup | 20.7 | 0.041 | 16.1 | 0.060 | 18.5 | 0.062 | ||||
acetabulum-solitaria lineage | 20.7 | 0.046 | 16.3 | 0.061 | 8.6 | 0.029 | 64 | 0.908 | 84 | 1 |
alpestris-nannfeldtii lineage | 22.5 | 0.049 | 2.7 | 0.010 | 1.05 | 0.003 | 96 | 1 | 100 | 1 |
alpina-corium lineage | 8.7 | 0.017 | 7.3 | 0.027 | 5.0 | 0.017 | 100 | 1 | 100 | 1 |
atra-pallescens lineage | 0 | 0 | 4.0 | 0.015 | 1.8 | 0.006 | 1 | 99 | 1 | |
bicolor-elastica lineage | 2.3 | 0.005 | 8.1 | 0.030 | 2.3 | 0.008 | 42 | – | 70 | 0.984 |
capucina-danica lineage | 7.3 | 0.015 | 4.6 | 0.017 | 6.8 | 0.022 | 87 | 0.712 | 95 | 1 |
corbierei-stevensii lineage | 7.0 | 0.014 | 4.5 | 0.017 | 4.4 | 0.014 | – | 0.938 | 97 | 1 |
costifera lineage | 2.0 | 0.004 | 1.2 | 0.005 | 1.5 | 0.006 | 91 | 0.990 | 98 | 1 |
crispa lineage | 5.0 | 0.010 | 5.6 | 0.021 | 6.0 | 0.020 | 98 | 1 | 100 | 1 |
elastica clade | 7.7 | 0.015 | 7.4 | 0.027 | 8.5 | 0.028 | 90 | 1 | 99 | 1 |
fallax-pezizoides lineage | 9.4 | 0.019 | 8.1 | 0.034 | 7.1 | 0.026 | 84 | 0.912 | 82 | 0.993 |
fibrosa-macropus lineage | 8.2 | 0.016 | 9.1 | 0.038 | 8.9 | 0.034 | 81 | 1 | 88 | 0.998 |
fusca-lactea lineage | – | – | 4.3 | 0.016 | 3.3 | 0.011 | 89 | 1 | 95 | 1 |
helvellula lineage | 3.3 | 0.007 | 2.6 | 0.010 | 2.0 | 0.007 | 100 | 1 | 100 | 1 |
hypocrateriformis lineage | – | – | 3.0 | 0.011 | 3.0 | 0.010 | 93 | 1 | – | – |
lacunosa-nigra lineage | 8.7 | 0.017 | 4.5 | 0.017 | 4.1 | 0.014 | 61 | 0.932 | 36 | – |
lacunosa clade | 10.8 | 0.021 | 9.4 | 0.035 | 8.9 | 0.026 | 99 | 1 | 100 | 1 |
leucomelaena lineage | 16.9 | 0.034 | 11.1 | 0.041 | 9.0 | 0.026 | – | – | 84 | 0.998 |
palustris-philonotis lineage | 0 | 0 | 2.1 | 0.008 | 2.0 | 0.007 | 100 | 1 | 98 | 1 |
rivularis-sublicia lineage | 8.5 | 0.017 | 7.6 | 0.028 | 9.7 | 0.033 | 99 | 1 | 99 | 0.918 |
Species | ||||||||||
H. acetabulum | 0 | 0 | 2.0 | 0.007 | 1.3 | 0.004 | 100 | 1 | 100 | 1 |
H. aestivalis | 18.0 | 0.036 | 2.0 | 0.009 | 0 | 0 | – | – | – | – |
H. alpestris | 5.0 | 0.010 | 1 | 0.004 | 0 | 0 | 76 | 0.999 | 100 | 1 |
H. alpicola | 0 | 0 | 2.0 | 0.008 | 0 | 0 | 99 | 1 | 100 | 1 |
H. alpina | 5.0 | 0.011 | 2.3 | 0.009 | 0.7 | 0.002 | 100 | 1 | 100 | 1 |
H. arctoalpina | 2.0 | 0.004 | 0.0 | 0 | 1.0 | 0.003 | – | – | 100 | 1 |
H. atra | 0 | 0 | 0.3 | 0.001 | 0.5 | 0.002 | 93 | 0.993 | 100 | 1 |
H. bicolor | 0 | 0 | 0.7 | 0.002 | 0.7 | 0.002 | 100 | 1 | 100 | 1 |
H. calycina | 1.0 | 0.002 | 1 | 0.004 | 0 | 0 | 95 | 0.999 | 98 | 1 |
H. capucina | 0 | 0 | 0 | 0 | 0.5 | 0.002 | 96 | 1 | 100 | 1 |
H. capucinoides | – | – | 0 | 0 | 0 | 0 | – | – | – | – |
H. carnosa | 1.0 | 0.002 | 5.0 | 0.036 | 0 | 0 | 96 | 1 | 99 | 1 |
H. confusa | 0 | 0 | 0.7 | 0.002 | 1.3 | 0.004 | – | – | 100 | 1 |
H. corbierei | – | – | 2.7 | 0.01 | 1.3 | 0.004 | 69 | 0.992 | 89 | 1 |
H. corium | 0 | 0 | 0.7 | 0.002 | 0.5 | 0.002 | 100 | 1 | 100 | 1 |
H. costifera | 2.7 | 0.005 | 0 | 0 | 3.3 | 0.011 | 65 | 0.824 | 93 | 0.996 |
H. crispa | 5.0 | 0.010 | 2.7 | 0.01 | 3.3 | 0.010 | 90 | 0.997 | 98 | 0.998 |
H. danica | 0 | 0 | 0 | 0 | 1.3 | 0.004 | 100 | 1 | 100 | 1 |
H. dryadophila | 1.0 | 0.002 | 0 | 0 | 0.7 | 0.002 | 100 | 1 | 100 | 1 |
H. elastica | 0 | 0 | 1.3 | 0.005 | 0 | 0 | 97 | 1 | 100 | 1 |
H. fallax | 0.7 | 0.001 | 0.7 | 0.002 | 0.7 | 0.002 | 99 | – | 89 | 0.940 |
H. fibrosa | 1.0 | 0.002 | 0 | 0 | 0 | 0 | 100 | 1 | 100 | 1 |
H. fistulosa | 3.3 | 0.007 | 2.0 | 0.007 | 0.7 | 0.002 | 94 | 1 | 99 | 1 |
H. fusca | – | – | 0 | 0 | 0.0 | 0 | 100 | 1 | – | – |
H. helvellula | 0 | 0 | 0 | 0 | 0 | 0.002 | 91 | 0.999 | 97 | 1 |
H. hyperborea | – | – | 1.0 | 0.004 | 0 | 0 | 100 | 1 | – | – |
H. hypocrateriformis | – | – | 0 | 0 | 0.7 | 0.002 | 99 | 0.971 | – | – |
H. lactea | – | – | 0 | 0 | 0 | 0 | 100 | 0.998 | – | – |
H. lacunosa | 0 | 0 | 1.3 | 0.005 | 0 | 0 | 92 | 0.624 | 100 | 1 |
H. leucomelaena | 11.0 | 0.020 | 0 | 0 | 0.7 | 0.002 | 99 | 1 | 95 | 1 |
H. levis | 1.0 | 0.002 | 0 | 0 | 0 | 0 | 98 | 0.889 | 100 | 1 |
H. macropus | 0 | 0 | 0 | 0 | 0 | 0 | 97 | 0.952 | 100 | 1 |
H. macrosperma | 0 | 0 | 0 | 0 | 0 | 0 | 96 | – | 100 | 0.999 |
H. monachella | – | – | 0 | 0 | 0.0 | 0.0 | 100 | 1 | – | – |
H. nannfeldtii | 7.0 | 0.015 | 1.3 | 0.005 | 0 | 0 | 84 | 1 | 100 | 1 |
H. nigra | – | – | 0.7 | 0.002 | 0.7 | 0.002 | 100 | 1 | – | – |
H. pallescens | 0 | 0 | 2.0 | 0.007 | 2.0 | 0.007 | 98 | 1 | 94 | 0.999 |
H. palustris | 0 | 0 | 2.0 | 0.007 | 0 | 0 | 99 | 1 | 96 | 0.997 |
H. panormitana | 1.3 | 0.003 | 0.7 | 0.002 | 1.3 | 0.004 | 86 | 1 | 99 | 1 |
H. pezizoides | 0 | 0 | 0.5 | 0.002 | 0.5 | 0.002 | 100 | 0.997 | 100 | 1 |
H. philonotis | 0 | 0 | 0.7 | 0.001 | 0 | 0 | 96 | 0.693 | 88 | 1 |
H. phlebophora | 0 | 0 | 1.0 | 0.004 | 1.0 | 0.003 | 100 | 1 | 71 | 0.978 |
H. pubescens | 5.0 | 0.010 | 0 | 0 | 1.0 | 0.003 | 100 | 1 | 100 | 1 |
H. pulla | 0.0 | 0 | 0 | 0 | 0 | 0 | 100 | 1 | 100 | 1 |
H. rivularis | 0.7 | 0.001 | 0 | 0 | 0 | 0 | 100 | 1 | 100 | 1 |
H. semiobruta | 5.0 | 0.010 | 2.0 | 0.007 | 2.0 | 0.007 | – | – | – | – |
H. silvicola | – | – | 2.0 | 0.007 | 0.7 | 0.002 | 100 | 1 | – | – |
H. solitaria | 0 | 0 | 0 | 0 | 0.7 | 0.002 | 100 | 1 | 100 | 1 |
H. stevensii | – | – | 0 | 0 | 0 | 0 | 99 | 1 | – | – |
H. sublicia | 8.0 | 0.016 | 0.7 | 0.002 | 0 | 0 | 100 | 1 | 100 | 1 |
H. sulcata | – | – | 1.0 | 0.004 | 0 | 0 | 100 | 1 | – | – |
H. vespertina | 0 | 0 | 0 | 0 | 0 | 0 | 100 | 1 | 100 | 1 |
H. sp. ‘JAPAN 4’ | – | – | 0 | 0 | 1.0 | 0.003 | 96 | 1 | – | – |
1 partial 28S large ribosomal sub unit (LSU), heat shock protein 90 (hsp) and RNA polymerase II (rpb2).
2 ‘all’ indicates total pairwise nucleotide diversity, while ‘per site’ indicates pairwise nucleotide diversity per site.
3 Two-gene tree denotes phylogenies based on a hsp and rpb2 alignment, three-gene tree denotes phylogenies based on a hsp, rpb2 and LSU alignment.
4 BP denotes Maximum Likelihood Bootstrap Pseudoreplicates and BPP denotes Bayesian Posterior Probability. ‘–’ denotes missing value.
Inference of species phylogenies
We selected the maximum likelihood (ML) tree based on the hsp and rpb2 concatenated alignment as representative for the species lineages in Helvella (Fig. 2), as this alignment included most of the taxa with more than one accession and the maximum likelihood and Bayesian methods produced concordant topologies (not shown). Support values from Bayesian posterior probability (BPP) analyses were added to the ML tree. In addition, ML bootstrap (BP) and BPP from the analyses of the three-locus alignment (including LSU) were added, since LSU gave additional support to some branches.
We recovered 55 Helvella species from Europe based on concordance of the single gene phylogenies and two- and/or three-gene phylogenies. In addition, 27 extralimital non-European species or single-specimen lineages were supported by genetic divergence from their sisters in two or three of the protein-coding gene trees. A lineage was considered to be genealogical concordant if it was found in more than one locus and had BP and BPP support values greater than 70 % and 0.95, respectively.
Initial analyses suggested that H. aestivalis (R. Heim & L. Rémy) Dissing & Raitv. (1974) does not belong in the genus Helvella. However, blast searches, using LSU sequences as the query, indicated that this species is part of the Helvellaceae, and therefore was used as an outgroup to Helvella in our analyses. The placement of H. aestivalis will be resolved in a separate study, thus the species is not treated here.
In our phylogeny specimens of H. silvicola formed a well-supported group as sister to the rest of the Helvella species. The phylogenetic tree exposed some well-supported major clades and a number of resolved and unresolved infrageneric lineages. To improve the ‘readability’ of the tree, we have outlined and named 5 nested clades (A–E) and 18 lineages to be compared and discussed. The first lineage, named the leucomelaena lineage (clade A), encompasses the cosmopolitan H. leucomelaena, the European H. oblongispora and H. confusa, and the North American H. crassitunicata. Between this lineage and the rest of the helvellas encompassed in clade B, we find the morphologically highly divergent H. terrestris, which constitutes a very long branch of its own that may be subject to long-branch attraction. However, analysing a small subset of samples did not alter the placement of this species (analyses not shown). Within clade B we find a lineage, although unsupported, named the acetabulum-solitaria lineage (Fig. 2), which encompasses a species assemblage of H. solitaria, H. dryadophila, H. acetabulum, H. arctoalpina and the North American H. costata. A crispa lineage of H. crispa, H. maculata and H. leucophaea constitutes a species assemblage of its own nested outside a well-supported large lacunosa clade (clade C). The lacunosa clade includes 14 European and 8 non-European species and recovered many sister group relationships with high support: i.e., H. lactea (two-gene support: BPP = 0.996/BP = 99) with H. fusca (1/100) constituting a fusca-lactea lineage; H. pallescens (0.975/90) with H. atra (0.996/93) constituting an atra-pallescens lineage; H. palustris (1/95) with H. philonotis (0.885/73) constituting a palustris-philonotis lineage, H. lacunosa with H. vespertina and H. nigra, constituting a lacunosa-nigra lineage, and a helvellula lineage of H. juniperi, H. helvellula and H. semiobruta. The next two lineages to diverge are the alpestris-nannfeldtii lineage and the alpina-corium lineage, which include five species previously considered to constitute a single morphospecies, i.e., H. corium s.lat. The last lineage in clade B is the costifera lineage that includes H. calycina and H. costifera. The rest of the supported lineages are found within clade D. This clade encompasses lineages and species with cylindrical (not ribbed) stipes, i.e., a hypocrateriformis lineage, a fallax-pezizoides lineage, an unsupported fibrosa-macropus lineage, a rivularis-sublicia lineage, and a well-supported elastica clade (clade E), in addition to several single species lineages from Oceania, North America and Asia. The elastica clade contains 10 European and 5 non-European species and a number of supported sister group relationships, i.e., a corbierei-stevensii lineage, a bicolor-elastica lineage, and a capucina-danica lineage.
Seven independent evolving lineages are represented by single collections in our phylogram, i.e., H. scyphoides, H. semiobruta, H. juniperi, H. paraphysitorquata, H. leucophaea, H. queletiana and H. terrestris, plus an additional 22 extralimital non-European singletons, which represent a combination of discrete morphological and molecular data. These single-specimen lineages are interpreted as putative species of Helvella.
Species diversity and distribution
Helvella is a highly diverse genus, however, very little within-species divergence of the three loci hsp, rpb2 and LSU were observed (Table 3). Nevertheless, some species show more intraspecific diversity than others, e.g., H. costifera and H. crispa have highest within-species diversity. The diversity analyses also suggested that LSU evolved in a more uneven rate than hsp and rpb2. For some species, e.g., in H. nannfeldtii and H. leucomelaena (Table 3), LSU has evolved very quickly, but in others much slower, e.g., in H. palustris and H. philonotis. The hsp and rpb2 mostly evolved at a similar rate. The inferred lineages also showed variable degree of genetic diversity. For instance, the acetabulum-solitaria lineage showed high diversity but the elastica lineage contained comparably low genetic diversity. This also matched the branch lengths in our two-gene tree.
The Venn diagram demonstrated that most species are endemic to one continent (Fig. 1). The Venn diagram of course reflects that this study has mainly focused on European species. Of the 55 species in Europe, 36 were found only in Europe. Of the 29 species collected in North America, 16 were endemic to North America, while 10 were shared between Europe and North America. One species was present in East Asia, Americas and Europe, while two species were present in West Asia, Americas and Europe. Of the intercontinental species, some shared identical sequences from the different continent, e.g., for collections of H. pezizoides from Denmark and Japan and H. solitaria from Canada and Norway.
TAXONOMY
Nomenclature and typification
We propose 29 lectotypes, 28 epitypes and two neotypes for European Helvella species and resurrect 12 old binomials to be applied to newly recovered species, i.e., two species described by Schaeffer (1774), two species described by Bergeret (1783), one species described by Persoon (1799), two species described by Holmskjold (1799), one species described by Albertini & Schweinitz (1805), one species described by Raddi (1807), one species described by Schweinitz (1822), one species described by Fries (1822), and one species described by Inzenga (1865). Seven species are described as new, i.e., H. alpicola, H. alpina, H. carnosa, H. danica, H. nannfeldtii, H. pubescens and H. scyphoides, and full descriptions are provided for 12 emended species where the current concepts encompassed several related or unrelated species. Citations to precise descriptions and illustrations of the accepted European Helvella species of our sample are given.
For some species, our taxon sampling and data acquisition (i.e., incomplete molecular datasets) are not yet ideal for a consummate description of the species. Two new species descriptions are based on morphology and DNA sequence information from only one specimen. Being fully aware that single specimen descriptions cannot reflect intraspecific variation, we still believe our data represent characters so discrete that it is considered highly unlikely they fall within the variation range of another species. We therefore find it justified keeping these entities on a formally recognized level of species.
We provide hsp sequences for 55 European and 27 extralimital species of Helvella, tef sequences for 25 European and two extralimital species, rpb2 sequences for 55 European and 25 extralimital species, and LSU nrDNA sequences for 55 European and 11 extralimital and putative Helvella species. This includes partial sequences from one to three genetic loci for 14 holo- or isotypes and 30 newly designated neo- and epitypes of species recorded in this study. Heterotypic synonymies inferred from identical partial protein-coding gene sequences of type specimens are provided for four taxa, i.e., H. arctica, H. cupuliformis, H. dovrensis and H. pulchra.
In a situation where sufficient material has not been collected across the whole distributional area we add under Notes molecular specifics for each locus to the species descriptions. This is intended to help in specimen identification and to guide future re-assessment of useful characters for species discrimination within Helvella spp. in Europe and worldwide.
In the present synopsis, we refer to good illustrations and descriptions or describe de novo the Helvella species from Europe known by us. We also present an artificial key to the species based on morphological characters.
Morphology
Ascocarps (apothecia) in Helvella are either cupulate or capitate, subsessile or usually distinctly stipitate; when cupulate, cup deeply cupulate to saucer-shaped to discoid (planar), sometimes cup laterally compressed; when capitate, cap regularly campanulate to bi- to trilobate to irregularly saddle-shaped with cap edge free or partly attached to stipe, apothecial margin (cup margin or cap edge) recurved (deflexed) or adnate (reflexed) towards the stipe and sometimes firmly fused with it; hymenium whitish to greyish to brownish to black when fresh, receptacle surface (apothecial underside) concolorous or discoloured, glabrous or pubescent; stipe when present terete or ribbed and furrowed, inside solid or hollow or chambered. In some cupulate species the ribbed stipe becomes much reduced so that almost sessile apothecia occur. Asci are cylindrical, 8-spored, operculate, with an aporhynchous or pleurorhynchous base. Ascospores ellipsoid to ellipso-fusoid, hyaline, generally unsculptured or minutely verrucose in fully mature ascospores, containing a large central guttule when mature, tetranucleate. Paraphyses filiform, septate, not much inflated, or inflated (enlarged) at tips to a clavate or subcapitate apex.
KEY TO EUROPEAN SPECIES OF HELVELLA
1. Apothecium with a ± prominent distinct stipe. . . . . . . . . . . . . . 2
1. Apothecium subsessile, stipe poorly differentiated or absent . . . . . . . . . . . . . . 13
2. Stipe ribbed and furrowed over entire length, internally solid or chambered, sometimes ribs present on receptacle surface. . . . . . . . . . . . . . 3
2. Stipe terete, even, internally solid or hollow, occasionally with grooves at base and/or apex, tissues well differentiated. . . . . . . . . . . . . . 30
3. Apothecium regularly cupulate to planar. . . . . . . . . . . . . . 4
3. Apothecium capitate, convex to saddle shaped or irregularly lobed. . . . . . . . . . . . . . 16
4. Stipe with ribs extending onto receptacle surface; ribs commonly branching and interconnected by cross-veins . . . . . . . . . . . . . . 5
4. Stipe with blunt ribs not or scarcely continuing onto the receptacle surface. . . . . . . . . . . . . . 11
5. Hymenium light brown (6D5) to brown (6D8), sometimes with a violet tinge (11E5), drying brown (6-7D8) to dark brown (7F4-8). Receptacle surface light brown above, paler yellowish (4C7) to cream (4A3) below, delicately pubescent; ribs on receptacle surface branching, sharp-angled, rarely interconnected by cross-veins, not reaching the margin. Fruiting in spring (April to June). . . . . . . . . . . . . . H. acetabulum
5. Hymenium grey (3-6D-E1-3) or light brown (5-6D6-8) to dark brown (6-9E-F5-8). Fruiting in summer and autumn (July to October). . . . . . . . . . . . . . 6
6. Stipe long and slender, inflated above, with prominent blunt ribs extending half way up the receptacle surface. Hymenium light brown (5-6D6-8), drying darker (5-6E-F7-8), upper part of receptacle concolorous, lower receptacle surface yellowish white (3A2) to pale yellow (3A3-4); stipe and ribs light to greyish orange (5A-B4-6) . . . . . . . . . . . . . . H. hyperborea
6. Stipe low, 1–5 cm high, with prominent ± branching ribs extending onto the receptacle surface. . . . . . . . . . . . . . 7
7. Apothecium light brown (5-6D6-8) to dark brown (6-9E-F5-8); ribs reaching lower to upper third of receptacle surface. In Dryas vegetation in arctic-alpine habitats. . . . . . . . . . . . . . 8
7. Apothecium grey (3-6D) or pale greyish brown (6E3), ribs prominent, reaching or almost reaching the cup edge. Not in Dryas vegetation. . . . . . . . . . . . . . 9
8. Hymenium brown (6D-E5-8) to dark brown (6-9E-F5-8), receptacle surface paler, drying blackish. Ribs blunt, double-edged, with little branching, reaching lower third of the receptacle surface. . . . . . . . . . . . . . H. dryadophila
8. Hymenium dark brown (6-9E-F5-8), drying brownish black (5-9H8); receptacle surface paler brown, sharp-angled, rarely interconnected by cross-veins, reaching upper half of receptacle surface in fully expanded specimens. . . . . . . . . . . . . .. . . . . . . . . . . . . . H. arctoalpina
9. Apothecium with prominent blunt to angular ribs and grooves on underside of apothecium. Hymenium brown when dried (6D7-8), receptacle covered by greyish white pruina (B1). . . . . . . . . . . . . . H. oblongispora
9. Hymenium grey (E1-2) to brownish (6E5-8), exterior of cup (receptacle) pubescent. . . . . . . . . . . . . . 10
10. Hymenium greyish (6-7E1-3), drying dark greyish brown (6E3), stipe distinct with 5–6 prominent yellowish white blunt-edged ribs (4A2-3) extending onto receptacle, ribs not interconnected. . . . . . . . . . . . . . H. costifera
10. Hymenium brownish (6-8E5-8), receptacle greyish brown (6E3), stipe distinct with 5–8 prominent blunt-edged yellowish grey (4B2-3) ribs extending onto receptacle, in old fruit bodies ribs interconnected by shallow concentric parallel connections. . . . . . . . . . . . . . H. calycina
11. Hymenium black (19-24H8) or greyish black (19-24H1). Apothecium at first regularly cupulate, then laterally compressed with a ± recurved margin; hymenium black (20-21H8), receptacle and stipe greyish black (H1), with ± ‘warts’ (tufts of hyphoid hairs); stipe short, often compressed, with two to four grooves and blunt-edged shallow ribs that continue shortly onto the receptacle, stipe pale greyish (B1) towards the base. . . . . . . . . . . . . . H. philonotis
11. Hymenium brown (5-9D-E5-8) or greyish brown (6-7E3-4). . . . . . . . . . . . . . 12
12. Apothecium 1.8–8 cm broad; stipe 1–8 cm long, 0.5–2 cm broad. Hymenium dark brown (6-9E-F5-8) to brownish black (5-9H8); stipe white to pale yellow (1-4A1-3), with 4–7 regular blunt ribs; when mature apothecia laterally compressed and sometimes recurved and irregular in shape, receptacle paler, greyish to greyish brown; frequently under Salix. (High elevation specimens have been distinguished as H. ulvinenii.). . . . . . . . . . . . . . H. solitaria
12. Apothecium 1–3.5 cm broad; stipe 0.5–4 cm long, 0.4–1.3 cm broad. Hymenium greyish brown (6-7E3-4) to dark brown (5-9E5-6), receptacle paler; stipe yellowish white to yellowish grey (2-4A-B2-3), with 2–4 regular, blunt ribs . . . . . . . . . . . . . . H. confusa
13. Apothecium subsessile or with a short, narrow stipe, rabbit-ear shaped, 2–4.5 cm broad, 3–6 cm high; hymenium brownish red (10D-E7-8); receptacle surface glabrous, brownish orange (6C6-8), towards the base pale yellow to white (4A1-3). . . . . . . . . . . . . . H. silvicola
13. Apothecium cupulate or planar, hymenium and exterior brown (6D-E5-8) to dark brown (6-9E-F5-8) or black 20-21H8; stipe short (subsessile) and indistinct, with grooves. . . . . . . . . . . . . . 14
14. Apothecium cupulate to planar with an irregular lobed apothecial margin; hymenium and outside black (H8) . . . . . . . . . . . . . .. . . . . . . . . . . . . . H. helvellula
14. Apothecium cupulate with a crenate margin; hymenium greyish brown to dark chocolate brown (6D-F3-8) . . . . . . . . . . . . . . 15
15. Receptacle copper brown (7-8C-D8); hymenium dark chocolate brown (6-8E-F8), without ribs on receptacle surface. In Dryas vegetation in arctic-alpine habitats. . . . . . . . . . . . . .. . . . . . . . . . . . . .‘H. aestivalis’
15. Receptacle light brown to brown (6D-E3-5), turning pale brown (5-6C-D4-6) to whitish when dried, with conspicuous blunt ribs and grooves on lower half. Under conifers, usually Pinus. . . . . . . . . . . . . .H. leucomelaena
16. Hymenium white to cream (3-5A1-2) or golden to brownish yellow (5B-C7-8); receptacle surface villose; edge of cap free or in part fused with stipe. . . . . . . . . . . . . . 17
16. Hymenium grey, brown or black; receptacle surface smooth or at most finely sub-pubescent; edge of cap free or in part fused with stipe. . . . . . . . . . . . . . 21
17. Hymenium white to cream (3-5A1-2), receptacle surface pubescent. . . . . . . . . . . . . . 18
17. Hymenium golden to brownish yellow (5B-C7-8), receptacle surface finely pubescent. . . . . . . . . . . . . . H. leucophaea
18. Apothecium saddle-shaped to irregularly lobed, edge of cap free. . . . . . . . . . . . . . 19
18. Apothecium saddle-shaped to irregularly lobed, edge of cap adnate and fused with stem at 3–4 points. . . . . . . . . . . . . . 20
19. Apothecium irregularly lobed, receptacle surface pubescent, light greyish to brownish orange (5-6B-C4-5); stipe robust, with sharp, partly anastomosing longitudinal ribs. Asci 270–320 μm long. . . . . . . . . . . . . . H. crispa
19. Apothecium saddle-shaped, margin reflexed towards the stipe, hymenium white to pale yellow (3-4A1-3), drying brownish orange (5-6C5-7); stipe short and broad, with longitudinal distantly interconnected ribs. Asci 240–260 μm long. . . . . . . . . . . . . . H. lactea
20. Apothecium saddle-shaped to campanulate; hymenium and stipe white to pale grey (3-5A-B1-2), drying brownish yellow (5B-C7-8); stipe long and slender, with four to five longitudinal distant ribs. Asci 280–310 um long. . . . . . . . . . . . . .. . . . . . . . . . . . . . H. pallescens
20. Apothecium irregularly lobed to saddle-shaped; receptacle surface smooth, lacking raised veins, stipe lacunose. . . . . . . . . . . . . .. . . . . . . . . . . . . . H. lacunosa (albino)
21. Hymenium brown (6D-E5-8) to dark brown (6-9E-F5-8) when fresh, drying darker; stipe white. . . . . . . . . . . . . . 22
21. Hymenium grey (B-F1-2) or black (H8) when fresh. . . . . . . . . . . . . . 23
22. Apothecium irregularly lobed, cap edge partly attached to stipe; stipe robust (broad), lacunose, with prominent, longitudinal ribs, extending onto receptacle surface and here forming elevated, branching veins . . . . . . . . . . . . . . H. fusca
22. Apothecium hemispherical in shape; stipe with sharp, elevated ribs extending onto and ending blunt on the receptacle surface, occasionally with a few veins from termination points; fruiting in spring. . . . . . . . . . . . . . H. queletiana
23. Apothecium saddle-shaped or irregularly lobed, edge of cap partly fused with stipe; stipe with 4–8 prominent, longitudinal, parallel ribs. . . . . . . . . . . . . . 24
23. Apothecium convex, flat hemispherical or saddle-shaped; edge of cap free. . . . . . . . . . . . . . 25
24. Apothecium dark grey to black (F-H1-8), bi- to trilobate, cap edge remaining adnate towards stipe in mature specimens; hymenium black, often undulate-rugose, receptacle concolorous or dark greyish black (F-G1); stipe with 3–6 parallel ribs and grooves. . . . . . . . . . . . . . H. palustris
24. Apothecium convex to flat hemispherical; stipe greyish white (B1), contrasting the shining black hymenium (H8); with conspicuous sharp-edged longitudinal ribs extending onto receptacle surface and here building a network of radiating, raised veins. . . . . . . . . . . . . . H. phlebophora
25. Apothecium short-stipitate to medium-sized, cap saddle-shaped with lobes reflexed towards the stipe, up to 2 cm across; hymenium black (H8); stipe to 1.5 cm tall, with 3–4 inconspicuous, rounded ribs and grooves, subglabrous, grey to greyish black (B-F1-2), whitish below. Growing with Cistus and Pinus in the Mediterranean region. . . . . . . . . . . . . .. . . . . . . . . . . . . . H. semiobruta
25. Apothecium long-stalked; medium-sized to large; cap margin partly fused with stem at 3–6 points. . . . . . . . . . . . . . 26
26. Apothecium hemispherical to slightly saddle-shaped, sometimes cap edge in part deflexed; hymenium black (H8), stipe grey above (5-7C-E1-2), whitish below, with 3–6 prominent longitudinal, blunt-edged, parallel ribs, partly attached to cap margin at 2–3 points. Growing with Juniperus in the Mediterranean region . . . . . . . . . . . . . . H. juniperi
26. Apothecium saddle-shaped to irregularly lobed; hymenium dark grey to black, stipe concolorous or slightly paler than hymenium, with 4–8 longitudinal, ± anastomosing parallel ribs partly attached to apothecial margin at 3–6 points, ± lacunose. . . . . . . . . . . . . . 27
27. Apothecium from 1–4 cm broad, from 3–6 cm high; hymenium black (H8); stipe slightly paler than hymenium, with 4–6 partly anastomosing longitudinal ribs, slightly lacunose. . . . . . . . . . . . . . H. sulcata
27. Apothecium large, from 1.5–6 cm broad, 3–15 cm high; hymenium black (H8) or brownish black (5-7F-G6-8) to dark violet brown (10-12F7-8); stipe robust (broad), lacunose, with prominent, anastomosing parallel ribs. . . . . . . . . . . . . . 28
28. Apothecium large, saddle-shaped; hymenium dark grey (F1) to greyish black (G1) when fresh, drying darker; stipe concolorous or slightly paler, lacunose, with 4–6 partly double-edged ribs, attached at 4–6 points to the adnate apothecial margin. . . . . . . . . . . . . . H. nigra
28. Hymenium black or dark violet brown; stipe concolorous or grey to whitish. . . . . . . . . . . . . . 29
29. Apothecium 1–5 cm broad, 3–12 cm high, saddle-shaped or irregularly lobed; hymenium and stipe blackish with a dark violet brown tinge (10-12F-H7-8), stipe with 4–7 sharp-edged, partly double-edged ribs, attached at 3–4 points to the reflexed apothecial margin. . . . . . . . . . . . . . H. atra
29. Apothecium large, 3–6 cm broad, 4–15 cm high, saddle-shaped or irregularly lobed, hymenium black (H8), often wrinkled-folded; stipe pale grey to white, lacunose, with 5–8 sharp-edged, commonly double-edged ribs, attached at 4–6 points to the apothecial margin; parts of margin free and often deflexed. . . . . . . . . . . . . . H. lacunosa
30. Apothecium cupulate to planar. . . . . . . . . . . . . . 31
30. Apothecium convex, bi- to trilobate saddle-shaped, or irregularly lobed. . . . . . . . . . . . . . 43
31. Apothecium stipitate-cupulate to discoid, cup 0.4–0.8 cm diam, stipe 0.3–0.8 cm high, 0.1–0.2 cm broad; hymenium purplish grey (12-14D3) to dark greyish black (F-G1), outside concolorous or paler, pubescent; stipe greyish to whitish. Ascospores fusoid, 50–60 × 10–15 μm. . . . . . . . . . . . . .. . . . . . . . . . . . . . H. terrestris
31. Apothecium larger. . . . . . . . . . . . . . 32
32. Hymenium and receptacle black (H8). . . . . . . . . . . . . . 33
32. Hymenium grey (D-F1-2), greyish brown (5-11D-F1-2) or yellowish brown (5C-D4-8); receptacle surface and stipe pubescent or villose. . . . . . . . . . . . . . 37
33. Apothecium 1–5 cm across, at first regularly cupulate, later slightly compressed and folded with irregular lobed margin; receptacle surface villose, sometimes with whitish scales (‘warts’) of fascicled hyphoid hairs towards apothecial margin; stipe black, 0.5–5 cm high, 0.2–0.7 cm broad, pubescent, occasionally with 2–3 irregular ribs extending onto the base of the receptacle. . . . . . . . . . . . . . H. corium
33. Apothecium regularly cupulate to discoid, 0.5–2 cm across; stipe black or whitish below. . . . . . . . . . . . . . 34
34. Apothecium short-stipitate, regularly cupulate to slightly irregular and compressed when post-mature, receptacle surface and stipe subtomentose; stipe greyish black above, white below, 0.5–2.0 cm high, 0.2–0.3 cm broad, deeply buried in soil, sometimes with a few inconspicuous furrows. . . . . . . . . . . . . . H. nannfeldtii
34. Stipe entirely black . . . . . . . . . . . . . . 35
35. Stipe slender, 1–3.5 cm high, 0.2–0.5 cm broad, terete when fresh, with some inconspicuous furrows when dried, subpubescent. . . . . . . . . . . . . . H. alpina
35. Stipe relatively short, 0.4–1.5 cm high, 0.2–0.4 cm broad, terete, cylindrical, densely tomentose. . . . . . . . . . . . . . 36
36. Hairs dark brownish black, partly fascicled and partly scattered, distributed on receptacle surface, up to 200 μm long. Asci 270–310 × 12–16 μm; ascospores 18.5–22.5 × 11.0–13.8 μm; paraphyses clavately enlarged at tips. . . . . . . . . . . . . .. . . . . . . . . . . . . . H. alpestris
36. Hairs dark brownish black, mostly aggregated in conspicuous, pyramidal fascicles, hairs up to 300 μm long at the apothecial margin. Asci 290–330 × 16–20 μm; ascospores 18.5–22.5 × 11.5–13.8 μm; paraphyses irregularly gnarled and slightly bent at tips. . . . . . . . . . . . . . H. macrosperma
37. Hymenium grey (D-F1-2) or dark greyish brown (5-11D-E2-3), receptacle surface and stipe villose; ascospores verruculose after discharge. . . . . . . . . . . . . . 38
37. Hymenium yellowish brown (5-6C-D4-8) or grey (D-E1-2) to dark greyish brown (6D-E2-3) or black when fresh, receptacle and stipe pubescent . . . . . . . . . . . . . . 39
38. Apothecium 1–3 cm across, hymenium grey (5-7C-E1-2); stipe long and slender, 3–7 cm high, solid. Ascospores ellipso-fusoid (acuminate), to 27 μm long. . . . . . . . . . . . . . H. macropus
38. Apothecium 0.5–2.0 cm across, hymenium dark greyish brown (5-6E-F1-6); stipe 1.5–4 cm long, pale grey to brownish. Ascospores ellipsoid with rounded ends, to 23 μm long. . . . . . . . . . . . . . H. fibrosa
39. Apothecium 1.0–3.0 cm across; hymenium and receptacle surface yellowish brown (5-6C-D4-8) drying dark brown (6F5-8); stipe cream to whitish, usually shorter than cup diam. . . . . . . . . . . . . . H. hypocrateriformis
39. Apothecium regularly cup-shaped to discoid, cup 0.5–3.5 cm across, hymenium grey, brown or greyish black. . . . . . . . . . . . . . 40
40. Hymenium greyish black (F-G1); receptacle dark greyish (E-F1), receptacle and stipe densely tomentose; stipe greyish to greyish white below, 0.2–0.3 cm broad, 0.5–1.8 cm high, with 2–3 shallow grooves at base. . . . . . . . . . . . . . H. alpicola
40. Hymenium grey or brown . . . . . . . . . . . . . . 41
41. Cup 0.5–2.0 cm across; hymenium grey (D-E1-2) to dark greyish brown (6D-E2-3), receptacle surface conspicuously villose, concolorous; stipe 1.5–3.5 cm long, greyish above, yellowish white below. . . . . . . . . . . . . . H. rivularis
41. Cup 1.5–3.5 cm across; hymenium pale greyish yellow (3-4B-C4-8) to pale brown (5-6D4-8) to brown, turning dark brown when dried. . . . . . . . . . . . . . 42
42. Apothecium cupulate to discoid; cap 2–3.5 cm across; hymenium greyish yellow (3-4B-C4-8) when fresh, drying dark yellowish brown (5F5); receptacle pale grey (B1-2), delicately pubescent; stipe light grey to whitish, pubescent, 1.5–3 cm high. . . . . . . . . . . . . . H. pubescens
42. Apothecium cupulate; hymenium dark brown (5-6F5-8) when dried, receptacle and stipe greyish (E1) to whitish, tomentose. . . . . . . . . . . . . . H. scyphoides
43. Cap remaining reflexed (adnate) towards the stipe. . . . . . . . . . . . . . 44
43. Cap saddle-shaped or irregularly lobed with a ± deflexed margin. . . . . . . . . . . . . . 50
44. Cap grey or black; receptacle and stipe concolorous or paler. . . . . . . . . . . . . . 45
44. Cap and stipe with contrasting colours; hymenium grey, brown or brownish black; stipe white to yellowish. . . . . . . . . . . . . . 46
45. Apothecium dark grey to black (F1-H1), slender, up to 6 cm high, cap 1–3 cm across, in mature specimens cap bi- to trilobate and reflexed towards stipe (connivent saddle-shaped); receptacle surface greyish (E-F1-2) when dried, subtomentose at margin, becoming almost glabrous towards stipe attachment; stipe solid, greyish, paler towards base, pubescent below. . . . . . . . . . . . . . H. fallax
45. Apothecium pale grey to medium grey (C-E1), saddle-shaped, bilobate, apothecial margin reflexed throughout development, receptacle and stipe concolorous, pubescent, attached to deeply decayed wood. . . . . . . . . . . . . . H. pulla
46. Cap regularly bi- to trilobate to campanulate; hymenium greyish brown (5-8D-E3) to light brown (5-6D4-8) to dark brown (5-6E5-8) to brownish black (6-9F-G6-8); flesh (receptacle tissues) gelatinous. . . . . . . . . . . . . . 47
46. Cap campanulate or bilobate; hymenium brownish yellow (5C6-8) to light brown (5-6D4-8) to brown (6-7D5-8), drying darker; flesh not gelatinous. . . . . . . . . . . . . . 49
47. Apothecium long and slender, 3–15 cm in length, cap bi- to trilobate to campanulate; hymenium greyish brown to light brown to brown, stipe whitish to cream. . . . . . . . . . . . . . 48
47. Apothecium short-stipitate, cap campanulate to bi- to trilobate, hymenium dark brown to brownish black (6E-G6-8), stipe up to 2.5 cm high, solid, glabrous, broadly attached to cap underside, pure white when fresh, drying yellowish . . . . . . . . . . . . . . H. capucina
48. Hymenium light brown to brown (5-7D4-8); sometimes decoloured; receptacle glabrous. . . . . . . . . . . . . . H. elastica
48. Hymenium greyish brown (5-8D-E3), receptacle subpubescent. . . . . . . . . . . . . . H. panormitana
49. Cap campanulate; hymenium brownish yellow (5C6-8), apothecia commonly clustered. . . . . . . . . . . . . . H. danica
49. Cap regularly bilobate; hymenium brown (5-6E4-8), drying brownish black (6F-G5-8); receptacle surface delicately pubescent; stipe yellowish when dried. . . . . . . . . . . . . . H. carnosa
50. Receptacle surface pubescent to villose, apothecial margin partly inrolled (deflexed) over hymenium . . . . . . . . . . . . . . 51
50. Receptacle surface glabrous. . . . . . . . . . . . . . 54
51. Apothecium white to cream when fresh, drying light brown. . . . . . . . . . . . . . H. corbierei
51. Apothecium grey, brown or black. . . . . . . . . . . . . . 52
52. Apothecium grey; cap irregularly saddle-shaped, at first deflexed, in fully expanded specimens often campanulate or bilobate with reflexed margin; hymenium and receptacle greyish brown (6D-E2-3); stipe paler grey (B-C1-2) towards base. . . . . . . . . . . . . . H. sublicia
52. Apothecium dark brown or black. . . . . . . . . . . . . . 53
53. Apothecium irregularly saddle-shaped and lobed, hymenium brown to dark brown (6E-F6-8), drying blackish, stipe consisting of 2–3 terete strands, free or fused at top half, whitish above, brownish below, delicately pubescent. . . . . . . . . . . . . .. . . . . . . . . . . . . . H. paraphysitorquata
53. Apothecium saddle-shaped, occasionally with a deflexed margin, in mature specimens cap occasionally bilobate and reflexed towards stipe (connivent saddle-shaped), 0.5–1.5 cm across; receptacle surface pubescent to villose; stipe 0.3–0.5 cm thick, up to 5 cm high, at first solid, then partly hollow (fistulose), pubescent above, glabrous below. . . . . . . . . . . . . .. . . . . . . . . . . . . . H. pezizoides
54. Cap regularly saddle-shaped to campanulate to irregularly lobed; light brown (5-6D6-8) to brown (5-9D-E5-8) to dark brown (6-9E-F5-8). . . . . . . . . . . . . . 55
54. Cap bi- to trilobate, hymenium brown (6D-E7-8) apothecial margin free and reflexed towards stipe (adnate), stipe terete, solid or hollow. . . . . . . . . . . . . . H. fistulosa
55. Hymenium greyish yellow (3B-C6-7) to brownish yellow (5C6-8), drying pale brown (5-6D4-6); cap regularly saddle-shaped, lobes deflexed; stipe at first solid, then hollow. Ascospores broadly ellipsoid, 17–21 × 13.5–17 μm. . . . . . . . . . . . . .. . . . . . . . . . . . . . H. levis
55. Hymenium dark brown (6-7E-F7-8) or blackish (6-7G-H7-8). . . . . . . . . . . . . . 56
56. Hymenium dark brown (6-7E7-8), drying blackish; cap regularly saddle-shaped or with irregularly deflexed lobes; stipe solid or hollow, 2–6 cm long, 0.3–0.6 cm broad. Ascospores ellipsoid, 19–22 × 11–13.5 μm. . . . . . . . . . . . . . H. bicolor
56. Hymenium dark brownish black to black (6-7G-H7-8), cap complex saddle-shaped with irregularly deflexed lobes; stipe robust, hollow, 2–6 cm long, 0.5–2 cm broad. Ascospores ellipsoid, 21–24 × 13–15 μm; preferable habitat sand dunes. . . . . . . . . . . . . . H. monachella
Helvella acetabulum (L.) Quél., Hymenomyc., Fasc. Suppl. (Alencon): 102. 1874 — Fig. 3c
Basionym. Peziza acetabulum L., Sp. Pl. 2: 1181. 1753; Fr., Syst. Mycol. 2: 44. 1822.
Synonyms. Octospora acetabulum (L.) Timm, Fl. Megapolit.: 260. 1788.
Aleuria acetabulum (L.) Gillet, Champ. France Discomyc.: 36. 1879.
Paxina acetabulum (L.) Kuntze, Revis. Gen. Pl. 2: 864. 1891.
Acetabula acetabulum (L.) Underw. & Earle, Alabama Agric. Exp. Sta. Agric. Coll. Bull. 80: 200. 1897.
Phleboscyphus acetabulum (L.) Clem., Crypt. Form. Coloradens.: 298. 1907.
Fungioides acetabuliforme Vaill., Bot. Par.: 57. 1727, nom. inval.
Macroscyphus acetabuliforme Gray, Nat. Arr. Brit. Pl. 1: 672. 1821.
Lectotype. Vaill., Bot. Par. t. 13, f. 1. 1727, selected by Dissing (1966b). Epitype designated here: Sweden, Östergötland, Gryt parish, Säterön, 14 June 1945, J.A. Nannfeldt 7885 (C-Fungi Exs. Suec. 1354, ‘Helvella acetabulum (L.: Fr.) Quél.’). MycoBank MBT375375
Peziza sulcata Pers., Syn. Meth. Fung. 2: 643. 1801; Fr., Syst. Mycol. 2: 44. 1822.
Acetabula sulcata (Pers.) Fuckel, Jahrb. Nassauischen Vereins Naturk. 23–24: 330. 1870 (‘1869–1870’).
Paxina sulcata (Pers.) Kuntze, Revis. Gen. Pl. 2: 864. 1891.
Lectotype designated here: Pers., Syn. Meth. Fung. 2, t. 5, f. 1. 1801. MycoBank MBT375686.
Misapplied name
– Acetabula sulcata sensu Boud., Icon. Mycol., livr. 16: no 357, pl. 246. 1907 (preliminary text with ‘circulaires’) (= ? Helvella solitaria P. Karst.).
Illustrations — Dissing (1964: f. 2), Harmaja (1977b: f. 1–2).
Description of the species — Harmaja (1977b: 48).
Specimens examined/sequenced. Czech Republic, Vändra, 4 Aug. 1960, A. Raitviir [H427] (C-F-45314). – Iceland, Nordur-Mulasysla, Hengifossargil, 6 Aug. 1983, J. Vesterholt [H261] (C-F-21174; ‘H. hyperborea Harmaja’ det. Harmaja). – Norway, Møre og Romsdal, Fræna, Skotten, road side on calcareous ground, 15 m asl, 3 June 2004, J.B. Jordal [H126] (O-178001); Møre og Romsdal, Fræna, Talstadhesten, road side on calcareous ground, c. 310 m asl, 11 June 2004, J.B. Jordal [H127] (O-178005); Vestfold, Sande, Mørkhassel, along road under Quercus, 10 m asl, 28 May 2006, O. Karlsen [H128] (O-284420); Vestfold, Nøtterøy, Torød, on mossy grass garden, 20 May 2005, A. Aronsen [H133] (O-286533); Østfold, Halden, Kjeøya, moldy deciduous forest on calcareous soil, 8 June 2001, I.-J. Seem & J. Iversby [H134] (O-64925); Oppland, Dovre, Grimsdalen, at Buåi, on the ground in subalpine birch forest, 29 July 1984, T. Schumacher & K. Østmoe TS 99.84 [H226] (O-253213), TS 100.84 [H225] (O-253212); ibid., 7 July 1982, TS 142.82 [H287] (O-253215); Dovre, Grimsdalen, Kattuglehøi, southern slope, 19 July 1983, T. Schumacher & K. Østmoe TS 33.83 [H286] (O-253214); Finnmark, F.-E. Eckblad 19.61 [H485] (O-253211). – Sweden, Östergötland, Gryt parish, Säteröen, close to Quercus, 14 June 1945, J.A. Nannfeldt 7885 [H410] (C-Fungi Exs. Suec. 1354 epitype); Öland. Böda parish, Byerum, by the road to Bränslegårdarna, 31 May 1967, R. Schöldström [H409] (C-Fungi Exs. Suec. 3261).
Notes — Harmaja (1976, 1977b, 1979) was particularly insightful when he concluded there were more recognisable species in the H. acetabulum morphospecies complex than previously thought. We inferred a highly-supported lineage of H. acetabulum and the morphologically and molecularly related H. arctoalpina and H. costata, and H. dryadophila (Table 3). Helvella costata, an apparent distinct American taxon of the H. acetabulum morphospecies complex, also fruits in spring and is here adopted for a genetically divergent specimen from California, USA (see Extralimital species below). The five collections of H. acetabulum from the Czech Republic, Sweden and Iceland, plus some 11 additional collections from Norway, share three unique single nucleotide polymorphisms (SNPs) within hsp, two in rpb2 and one within the LSU (see alignment: Dryad). The tef sequence was identical among five collections from different corners of Norway (see alignment: Dryad).
Helvella alpestris Boud., Bull. Soc. Bot. France 41: CCXL. 1894 — Fig. 4c, f
Synonyms. Cyathipodia corium var. alpestris (Boud.) Boud., Hist. Classif. Discomyc. Europe: 39. 1907.
Leptopodia alpestris (Boud.) Grelet, Bull. Soc. Bot. Centre-Ouest, Nouv. sér. 3: 86. 1934.
Holotype. Switzerland, Zermatt, Grand St. Bernard, Aug. 1894 (PC), not examined. Epitype designated here: Norway, Oppland, Dovre, Grimsdalen, Jegerhøi, 1 Aug. 2007, T. Schumacher TS 45.07 (O-253221). MycoBank MBT375376.
Misapplied names
– Helvella corium forma alpestris (Boud.) J. Favre, Ergebn. Wiss. Untersuch. Schweiz. Nationalparkes 5 (33): 27. 1955 (= ? H. philonotis Dissing).
– Leptopodia murina Boud. var. alpestris (Boud.) R. Heim & L. Rémy, Bull. Soc. Mycol. France 48: 58. 1932 (= ? H. alpina Skrede, T. Carlsen & T. Schumach.).
Illustrations — Boudier (1895: pl. 2, f. 2), Dissing (1966a: f. 11j).
Apothecia shallow stipitate-cupulate to -discoid, cup 0.5–1.8 cm broad, stipe terete, solid, c. 0.2–0.4 cm broad, 0.4–1.5 cm high, hymenium and outside black, receptacle and stipe densely tomentose. Medullary excipulum of textura intricata, hyphae 2–5 μm broad. Outer excipulum of brown-walled globose to angular cells, 10–20 μm diam, gradually arranged perpendicularly to the surface and forming a layer of closely packed cells extending into dark brown-walled multicellular, scattered hyphoid hairs or form bundles from the excipulum exterior, hairs generally 40–150 μm, at the margin to 200 μm long, cells 10–15 μm broad, 10–40 μm long. Asci pleurorhynchous, 270–310 × 12–16 μm. Ascospores ellipsoid, 18.5–22.5 × 11.0–13.8 μm. Paraphyses filiform, 2–2.5 μm broad, septate, brownish almost from base, colour increasing in intensity towards the gnarled, clavate tips, 4.5–6.0 μm broad.
Specimens examined/sequenced. Canada, Nunavut, Kitikmeot Region, above Bloody Falls, on moist, seasonally wet sand in shallow gully in sand hills, 19 July 2014, J.M. Saarela, P.C. Sokoloff, R.D. Bull 4182 [H460] (O ex DAOM-574891). – Norway, Oppland, Dovre, Grimsdalen, Jegerhøi, 1 Aug. 2007, T. Schumacher TS 45.07 [H014] (O-253221 epitype); Dovre, Grimsdalen, Tverrådalen, 6 Aug. 2009, T. Carlsen, I. Skrede & T. Schumacher TS 11.09 [H042] (O-253224); ibid., 15 Aug. 1997, T. Schumacher [H246] (O-253219); ibid., 31 July 2007, T. Schumacher TS 33.07 [H036] (O-253223); ibid., 1. Aug. 2007, T. Schumacher TS 20.07 [H013] (O-253220); ibid., 7 Aug. 2009, T. Carlsen, I. Skrede & T. Schumacher TS 06.09 [H031] (O-253222); ibid., 7 Aug. 2009, T. Carlsen, I. Skrede & T. Schumacher TS 11.09 [H042] (O-253224); Grimsdalen, Tverrgjelbekken, on silt and moss underneath Salix arbuscula, 26 July 1984, T. Schumacher & K. Østmoe TS 70.84 [H483] (O-253225).
Notes — The holotype of H. alpestris from Zermatt in Switzerland, located in PC and depicted by Dissing (1966a), is old and presumably unsuitable for DNA extraction. It is here supported with an epitype specimen from an alpine locality in Norway, adding sequences of hsp, tef, rpb2 and LSU as molecular characteristics to the species. Based on morphology and molecular data, H. alpestris is sister species to H. macrosperma from which it diverges in one substitution in hsp, four substitutions in tef, one substitution in rpb2 and seven substitutions in LSU.
Helvella alpicola Skrede, T. Carlsen & T. Schumach., sp. nov. — MycoBank MB820315; Hsp barcode GenBank KY784304; Rpb2 barcode GenBank KY772554; Fig. 5c, e
Etymology. From Latin ‘thriving in mountainous areas’.
Holotype. Norway, Nordland, Saltdal, Junkerdalsura, 27 Aug. 1988, A.E. Torkelsen 180.88 (O-185924).
Apothecia stipitate-cupulate, cup 0.5–1.5 cm broad, hymenium greyish black (G1), receptacle dark greyish (E-F1), receptacle surface and stipe densely tomentose, stipe greyish to whitish below, 0.2–0.3 cm broad, 0.5–1.8 cm high, with 2–3 shallow grooves at base. Medullary excipulum of loose textura intricata, hyphae 2–5 μm broad, hyaline. Outer excipulum of textura angularis, cells 10–27 μm diam, the strong pubescence of the outer receptacle surface due to long fascicled tufts of multiseptate, subhyaline, hyphoid hairs, 60–250 μm long; individual cells cylindrical to drum-shaped, 20–40 × 10–25 μm, with conspicuous brown pigments at septa. Asci pleurorhynchous, 290–330 × 15–18 μm. Ascospores ellipsoid, 20–22.6 × 11.8–13.5 μm. Paraphyses c. 2.5 μm below, septate, brownish, gradually enlarged to 5–6 μm at the clavate tips.
Specimens examined/sequenced. Norway, Nordland, Saltdal, Junkerdalsura, 27 Aug. 1988, A.E. Torkelsen 180.88 [H175] (O-185924 holotype). – Switzerland, Graubünden, Inn at Resgia, on soil at the river, 26 Aug. 1984, H. Dissing [H231] (O-253226).
Notes — Helvella alpicola possesses a tomentose receptacle covered by short-celled hyphoid hairs (Fig. 5c, e). It resembles H. rivularis in shape and colour. The species is recorded from two distant localities, e.g., in Northern Norway (Arctic) and the Swiss Alps. The sequenced specimens are invariable in rpb2, but differ in two hsp sites. Helvella alpicola forms a divergent lineage in our phylograms.
Helvella alpina Skrede, T. Carlsen & T. Schumach., sp. nov. — MycoBank MB820322; Hsp barcode GenBank KY784343; Rpb2 barcode GenBank KY772593; Fig. 4e
Etymology. From Latin ‘occurring in mountainous regions’.
= ? Helvella murina Boud. var. alpestris R. Heim & L. Rémy, Bull. Soc. Mycol. France 48: 58. 1932.
Holotype. France, Savoie, Plan des Evettes, 26 Aug. 1992, T. Schumacher F15.92 (O-253226).
Apothecia regularly cupulate-stipitate, black all over, cup 0.8–2.0 cm across, receptacle surface subpubescent, commonly with white crystalline deposits at the margin, stipe slender, 0.2–0.3 cm thick, 1.0–3.5 cm long, solid to hollow, occasionally with a few longitudinal grooves. Medullary excipulum of textura intricata, hyphae 2–5 μm broad, hyaline. Outer excipulum of textura angularis, cells 10–25 μm diam, intermixed with subhyaline to brown-walled hyphae, turned perpendicularly to receptacle surface and forming scattered 3–8-celled, brown-walled, branching hyphoid hairs, 30–100 μm long, individual cells ovoid to subglobose, up to 20 μm broad. Asci aporhynchous, 220–290 × 14–18 μm. Ascospores ellipsoid, 16.6–19.2 × 11.5–13.2 μm. Paraphyses 2.0–2.8 μm broad, septate, brownish along the whole length, gradually enlarged to 4.0–6.5 μm at the subcapitate tips.
Specimens examined/sequenced. Canada, British Columbia, Whistler National Park, on the ground under Salix in subalpine spruce forest, 13 Aug. 1994, T. Schumacher [H336] (O-253227). – France, Savoie, Plan des Evettes, 26 Aug. 1992, T. Schumacher F15.92 [H223] (O-253226 holotype). – Greenland, Qeqertarssuaq, Godhavn, the plain E of the Arctic station, in moss carpet of Paludella squarrosa with Salix arctophila, 11 Aug. 1977, P.M. Petersen [H349] (C-F-63820).
Notes — This new species may have been described from the French Alps as a variety (var. alpestris) of H. murina (Heim & Rémy 1932). There are, however, no authentic specimens left to support this assumption (Dissing, pers. comm). Macromorphologically, H. alpina resembles H. corium and H. alpestris, but has a more slender stipe two to three times longer than cap height and width. The receptacle appears smooth to the naked eye but is covered with minute, scattered distributed hyphoid hairs using a 10× hand lens. Helvella alpina and H. corium form a highly divergent lineage in our phylogeny characterised by black, stipitate-cupulate apothecia and asci with an aporhynchous ascus base. Asci and ascospores of the two species are in the same range, but apothecial shape, hairs and habitat easily distinguish them. The three collections from arctic and alpine localities in Greenland, France and Canada share identical rpb2 sequences, but vary in three hsp sites.
Helvella arctoalpina Harmaja, Karstenia 17: 58. 1977 — Fig. 3a, f
Holotype. Norway, Hordaland, Eidfjord, Dyranut, 1 Aug. 1959, F.-E. Eckblad (O-72616).
Synonyms. Acetabula barlae Boud., Hist. Classif. Discomyc. Europe: 40. 1907.
Paxina barlae (Boud.) Seaver, North Amer. Cup-fungi (Operculates): 205. 1928.
Helvella dalgeri Donadini, Bull. Soc. Linn. Provence 36: 136. 1985 ‘1984’.
Lectotype. France, Nice, Feb. 1883, D. Barla, selected by Dissing 1966a.
Illustrations — Boudier (1905–1910: pl. 245, as Acetabula barlae), Harmaja (1977b: f. 3–4), Schumacher & Mohn Jenssen (1992: 13).
Descriptions of the species — Harmaja (1977b: 48), Schumacher & Mohn Jenssen (1992: 13).
Specimens examined/sequenced. Canada, Ellesmere Island, Alexandra Fjord, 23 July 1984, H. Dissing [H338] (C-F-56722); ibid., [H337] (C-F-56723). – France, Savoie, Val d’Isere, 2550 m asl, in Dryas association, 31 Aug. 1992, T. Schumacher [H360] (O-253229). – Greenland, Ella Island, St. Elvdal, most calcareous sand in Dryas association, 6 Aug. 1982, H. Dissing & S. Sivertsen [H359] (C-F-50652). – Iceland, Nordur-Mulasysla, Hengifossargill, in dry heath vegetation with Dryas octopetala, 6 Aug. 1993, H. Knudsen [H342] (C-F-34334). – Norway, Hordaland, Eidfjord, Dyranut, in Dryas carpets, 1 Aug. 1959, F.-E. Eckblad [H293] (O-72616 holotype); Ulvik, Finse, Bio 101 peak, in Dryas association, 24 July 2014, T. Schumacher [H472] (O-253231); ibid., 29 July 2014, T. Schumacher [H456] (O-253230); Oppland, Lom, Høyrokampen, in Dryas association, 1400–1440 m asl, 29 Aug. 1957, F.-E. Eckblad [H354] (O-174714); Oppland, Dovre, Grimsdalen, Tverråi, along the slopes in Dryas vegetation, 27 July 1984, T. Schumacher & K. Østmoe D 83.84 [H083] (O-253239); ibid., 16 Aug. 1996, S. Landvik & T. Schumacher TS 199.96 [H084] (O-253240); ibid.,16 Aug. 2001, T. Schumacher [H003] (O-253232); ibid., 2. Aug. 2007, T. Schumacher TS 13.07 [H011] (O-253233); ibid., T. Schumacher TS 27.07 [H012] (O-253234); ibid., 7 Aug. 2009, T. Carlsen, I. Skrede & T. Schumacher TS 2.09 [H030] (O-253236); ibid., TS 7.09 [H033] (O-253237); ibid., TS 8.09 [H040] (O-253238); Grimsdalen, Tverrgjelet, in moist Dryas vegetation, 26 July 1984, T. Schumacher & K. Østmoe D 75.84 [H356] (O-253244); ibid., 20 Aug. 1997, T. Schumacher 35.97 [H255] (O-253403); ibid.,15 Aug. 2001, T. Schumacher [H182] (O-253243); ibid., 8 Aug. 2009, T. Carlsen, I. Skrede & T. Schumacher TS 24.09 [H026] (O-253235); Grimsdalshytta, 3 Aug. 1966, A. Danielsen & A.-E. Torkelsen [H086] (O-253241); ibid., [H087] (O-253242); Nordland, Ballangen, Håfjellet, on calcareous soil along path, 26 July 2008, D. Holtan & P.G. Larsen PL 41.2008 [H173] (O-69063).
Notes — The sequenced specimens from Canada and Greenland have one common rpb2 nucleotide character that separates them from the European specimens. European specimens are genetically uniform. Dissing (1966a) examined a specimen of Acetabula barlae in PC, which he considered to be a suitable lectotype specimen, and referred it to his concept of H. acetabulum. He noted that “the very characteristic dark greenish olive colours, seen in fresh specimens of H. acetabulum, are not mentioned in Boudier’s otherwise detailed description” (Dissing 1966a). In our opinion, Boudier’s plate of A. barlae (Boudier 1905–1910: pl. 245) depicts a typical specimen of H. arctoalpina, as does Favres record of A. barlae from the Swiss Alps (Favre 1955: pl. 4, f. 1). Helvella arctoalpina and H. acetabulum form a well-supported sublineage of the acetabulum-solitaria lineage in our phylogeny. Helvella arctoalpina differs from H. acetabulum by one substitution in rpb2, six in hsp, three in tef and 10 in LSU.
Helvella atra Oeder, Fl. Dan. 3 (9): 7. 1770 — Fig. 6f
Basionym. Helvella atra Oeder, Fl. Dan. 3, fasc. 9: 7. 1770; Fr., Syst. Mycol. 2: 19. 1822.
Synonym. Leptopodia atra (Oeder) Boud., Icon. Mycol., liste prélim.: 2. 1904.
Lectotype. Oeder, Fl. Dan. 3 (9): t. 534, f. 1. 1770 (‘Elvela’), selected by Filippa & Baiano (2011). Epitype designated here: Sweden, Uppland, Uppsala, about 300 m NW of Grindstugan, 13 July 1938, S. Lundell (C-Fungi Exs. Suec. 2066 ‘H. lacunosa Afzel. ex Fr. – dwarfed form’). MycoBank MBT375395.
Illustrations — Oeder (1770: pl. 534, f. 1), Filippa & Baiano (2011: f. 9–10, f. 11 as H. lacunosa).
Apothecia stipitate-capitate, saddle-shaped or irregularly lobed, 1–4 cm broad, 3–12 cm high, cap margin adnate and attached to stipe at 3–4 points; hymenium and stipe blackish with a dark violet brown tinge (10-12F-H7-8), even or wrinkled; receptacle surface greyish brown to black, smooth, with inconspicuous ribs on outer surface; stipe 0.5–2.5 cm broad, naked, with 4–7 sharp-edged, partly double-edged ribs, attached at 3–4 points to the reflexed apothecial margin greyish brown to black, paler towards base, ribs prominent, sharp, partly anastomosing and double-edged, with ‘holes’ (lacunae) between ribs, stipe chambered inside. Medullary excipulum of textura intricata, hyphae 3–5 μm broad. Outer excipulum of textura globulosa-angularis, cells 8–15 μm diam, hyaline, heavily staining in CB, outermost cells club-shaped, 10–30 × 5–15 μm. Asci pleurorhynchous, 260–10 × 14–18 μm. Ascospores ellipsoid, 15.5–18.5 × 10–12.5 μm. Paraphyses straight, septate, subhyaline to light brown below, increasingly brown above, 3–4 μm broad, clavate to 6.5 μm at tips.
Specimens examined/sequenced. Denmark, Zealand, Møn, Ulvshale, 13 June 1999, T. Læssøe [H467] (C-45782). – Georgia, Kaukasus, Kasbeghi, Gvethi valley, sandy soil along path, 200 m asl, 13 July 1988, E. Johannesen [H233] (O-253245); ibid., [H375] (O-253246). – Norway, Oppland, Dovre, Grimsdalen, at Buåi, on sandy soil along the road, 12 Aug. 1981, T. Schumacher & K. Østmoe 165.81 [H465] (O-253252); Grimsdalen, Jegerhøi towards Verkenseter, 1 Aug. 2007, T. Schumacher TS 43.07 [H016] (O-253251); Sør-Trøndelag, Oppdal, Loslia, in calcareous pasture land above tree limit, 995 m asl, 20 Aug. 2009, J.B. Jordal [H481] (O-291384); Nordland, Bodø, Sjunkhatten National Park, Steigtindvann, at the trail, 23 June 2010, L. Ryvarden [H462] (O-294033); Nordland, Steigen, Liland, Bru, 21 July 2006, N. Hagen 74.06 [H151] (O-281641); ibid., Andøy, Slettneset, in sand dunes on calcareous soil, 13 Sept. 2005, P.H. Larsen 234 [H464] (O-360146); Finnmark, F.-E. Eckblad 64.61 [H487] (O-253248); Finnmark, F.-E. Eckblad 99.61 [H484] (O-253247). – Sweden, Uppland, Uppsala, about 300 m NW of Grindstugan, 13 July 1938, S. Lundell [H406] (C-Fungi Exs. Suec. 2066 epitype).
Notes — Helvella atra was originally described and depicted in Flora Danica (Oeder 1770), but redescribed two years later by Zoega (1772) based on a specimen from Iceland collected by J. König. The illustration and descriptions leave little doubt that the fungus belongs to the H. lacunosa morphospecies complex, as also concluded by Dissing (1964, 1966b). Fries (1822) adopted Oeder’s name for a broadly defined species which included Königs fungus (Oeder 1770, Zoega 1772 – erroneously referred to as published by König) as well as H. atra in the sense of Afzelius (1783) and Holmskjold (1799), and H. nigricans (Persoon 1796, 1801). In our opinion Afzelius’ and Holmskjold’s ‘H. atra’ and Persoon’s H. nigricans represent a distinct species for which we have adopted the binomial H. fallax (see this species below). Dissing selected Holmskjold’s plate (1799: pl. 25) as the lectotype of H. atra, but should better have used Oeder’s plate as an interpretive legitimate type, since H. atra Oeder represents the older sanctioned homonym (cf. ICN Articles 53.1, 53.2, 15. Note 1 (ICN 2012)). We therefore re-instate Oeder’s plate as a lectotype and supports it with an epitype. Although a specimen from Iceland would have been the ideal epitype, we have selected a specimen from Sweden, widely distributed as an exsiccate specimen, to represent this species. The sequenced specimens from subalpine Norway have one rpb2 site polymorphism that separates them from the rest of the sequenced specimens. Two LSU sequences in GenBank [KC122770, KC122771] treated as H. lacunosa in Landeros et al. (2015) cluster with H. atra (cf. Appendix 3).
Helvella bicolor Raddi, Mem. Mat. Fis. Soc. Ital. Sci. 13, 2: 354. 1807 ‘1806’ — Fig. 7e, i
Lectotype designated here: Raddi, Mem. Mat. Fis. Soc. Ital. Sci. 13, 2: 362, pl. 13, f. 10. 1807. MycoBank MBT375687. Epitype designated here: Sweden, Gästrikland, Göskegruvan i Hofors, calcareous soil under Picea, 9 Sept. 1978, E. Sundström (C-F-45334). MycoBank MBT375464.
Synonyms. Helvella albella Quél., Compt. Rend. Assoc. Franc. Avancem. Sci. 24: 621. 1896 ‘1895’ (‘Elvela’)
Leptopodia albella (Quél.) Boud., Icon. Mycol., liste prélim.: 2. 1904.
Lectotype designated here: Quél., Compt. Rend. Assoc. Franc. Avancem. Sci. 24: t. 6, f. 16. 1896 (‘Elvela’). MycoBank MBT375688.
Illustrations — Raddi (1807: pl. 13, f.10), Dissing (1966b: f. 37, as H. albella).
Description of the species — Dissing (1966b: 136, as H. albella).
Specimens examined/sequenced. Norway, Vestfold, Lardal, Nerli, coniferous forest with Betula and Populus, 29 July 2000, P. Marstad [H147] (O-105346); Hedmark, Åmot, Kvannbekken National Reserve, calcareous river cleft, 30 Aug. 2002, T.S. Nilsen [H129] (O-65375); Akershus, Eidsvoll, Eidsvollbygningen, 16 Sept. 2009, I. Skrede TS 40.09 [H057] (O-253254); Akershus, Hurdal, 15 Sept. 2009, I. Skrede TS 41.09 [H056] (O-253253). – Sweden, Gästrikland, Göskegruvan i Hofors, on calcareous soil under Picea, 9 Sept. 1978, E. Sundström [H289] (C-F-45334 epitype).
Notes — The original description and illustration of H. bicolor Raddi (1807), which fully cover the present concept of H. albella of Quélet’s (1896), seems to have escaped recognition by earlier mycologists, including Fries (1822, 1832) and Dissing (1966b). We have chosen the plate and figure of Raddi’s (1807) as a lectotype and supported it with an epitype from Sweden, adding partial sequences from hsp, rpb2 and LSU to the species characteristics. According to Dissing (1966b), there is no authentic material of H. albella left in PC. The five collections studied by us have one polymorphic rpb2 and one polymorphic hsp site among them. A number of synapomorphies in the hsp and rpb2 loci separates the species from the sisters of the bicolor-elastica lineage of the elastica clade (clade E).
Helvella calycina Skrede, T. Carlsen & T. Schumach., nom. nov. — MycoBank MB820325
Etymology. From Latin ‘trumpet-like’.
Basionym. Boletus calyciformis Battarra, Fungorum Arimin.: 25. 1759; Fr., Syst. Mycol. 2: 45. 1822.
Synonyms. Peziza calyciformis (Battarra) Fr., Syst. Mycol. 2: 45. 1822.
Acetabula calyciformis (Battarra) Sacc., Syll. Fung. 8: 61. 1889.
Paxina calyciformis (Battarra) Kuntze, Revis. Gen. Pl. 2: 864. 1891.
– non Helvella calyciformis Batsch, Elench. Fung., cont. Prim. (Halle): 27. 1786 (‘Elvela’) = Peziza calycina α (var.) Fr., Syst. Mycol. 2: 91. 1822.
Lectotype designated here: Battarra, Fungorum Arimin.: t. III, f. C, 1759. MycoBank MBT375689. Epitype designated here: Norway, Oppland, Dovre, Grimsdalen, Storberget, 8 Aug. 2009, T. Carlsen, I. Skrede & T. Schumacher TS 16.09 (O-253255). MycoBank MBT375389.
Illustration — Dissing (1966b: f. 16, as H. costifera).
Apothecia at first short-stipitate-cupulate, when expanding more irregularly folded, 2.0–7.5 cm broad, 1.5–4.5 cm high. Hymenium brownish (6-8E5-8), receptacle surface greyish brown 6E3), slightly pubescent, stipe with 5–8 prominent blunt-edged yellowish grey ribs (4B2-3) extending onto receptacle but not reaching the margin, ribs apically dichotomously branched and interconnected by shallow concentric parallel connections in old fruit bodies making receptacle surface ‘wrinkled’. Medullary excipulum of hyaline textura intricata, hyphae c. 3–5 μm broad. Outer excipulum of textura prismatica to textura angularis, outermost cells with brown walls. Asci pleurorhynchous, 240–280 × 11–14 μm. Ascospores ellipsoid, 14.5–17.6 × 9.5–12.0 μm. Paraphyses 2.5–3.5 μm broad, straight, septate, gradually enlarged to 4–5 μm at the tips.
Specimens examined/sequenced. Denmark, Klim Bjerg, Klim W of Fjerritslev, 18 Oct. 1993, J. & T. Læssøe & A. Møller [H382] (C-F-21122); ibid., 16 July 1999, P.B. Hansen & J. Vesterholt JV 153.99 [H384] (C-F-41038); Northeast Jutland, Hjørring, under Picea in Klitplantage (Rubjerg Knude Plantage), 4 Aug. 1998, D. Boertmann DB 98007 [H387] (C-F-71286). – Norway, Oppland, Dovre, Grimsdalen, Storberget, 8 Aug. 2009, T. Carlsen, I. Skrede & T. Schumacher TS 16.09 [H022] (O-253255 epitype).
Notes — Helvella calycina is here typified based on the basionym Boletus calyciformis Battarra (1759). Fries adopted Peziza calyciformis for this cupulate, prominently ribbed autumnal species of Battarra’s and compared it with the vernal Peziza acetabulum (Fries 1822). Later, he described Peziza costata (Fries 1851), which was recombined in Helvella under the new legitimate name of H. costifera by Nannfeldt (Lundell & Nannfeldt 1953). Neither Fries nor Nannfeldt considered Boletus calyciformis as a possible earlier name for Peziza costata (= Helvella costifera). We now know there are at least two prominently ribbed cupulate Helvella species that occur in Europe sympatrically in autumn, i.e., H. costifera and H. calycina. Three specimens of Helvella from Mexico referred to as Helvella aff. costifera (JX993057, JX993058, JX993059; cf. Appendix 3) are, judged from the published LSU sequence, concluded to represent H. calycina. Helvella calycina differs from its sister species H. costifera in two consistent rpb2 and one hsp substitution plus some inconstant polymorphies, including one hsp and five rpb2 sites.
Helvella capucina Quél., Bull. Soc. Bot. France 24: 327. 1878 ‘1877’ — Fig. 7g
Synonym. Leptopodia capucina (Quél.) Boud., Hist. Classif. Discomyc. Europe: 37. 1907.
Lectotype. Quél., Bull. Soc. Bot. France 24: t. 6, f. 3. 1878, selected by Van Vooren (2010). Epitype designated here: France, Savoie, Val d’Isere, Gorges du Mal, 2300 m asl, on river bed, 31 Aug. 1992, T. Schumacher (O-253256). MycoBank MBT375396.
Illustrations — Dissing (1964: f. 3), Schumacher & Mohn Jenssen (1992: 15).
Descriptions of the species — Dissing (1964: 113), Schumacher & Mohn Jenssen (1992: 15).
Specimens examined/sequenced. Canada, Northwest Territories, Keewatin district, Rankin Inlet, NW of airstrip, on sandy ground with moss, 21 Aug. 1971, E. & M. Ohenoja 43.1971 [H397] (C-F-92107); Nunavut, Kitikmeot Region, above Bloody Falls, at Coppermine River, 19 July 2014, J.M. Saarela, P.C. Sokoloff & R.D. Bull 4183 [H459] (O- ex DAOM 574890). – France, Savoie, Val d’Isere, Gorges du Mal, 2300 m asl, on river bed, 31 Aug. 1992, T. Schumacher F 74.92 [H208] (O-253256 epitype); Savoie, Val d’Isere, Plan des Evettes, 2500 m asl, on river terrace, 26 Aug. 1992, R. Fellner F 1.82 [H211] (O-253257). – Greenland, Mestervig, around Blyminen, on naked soil in river bed, 14 Aug. 1982, H. Dissing & S. Sivertsen [H488] (C-F-50768). – Norway, Oppland, Dovre, Grimsdalen, Buåi, at the river, 17 Sept. 1983, T. Schumacher & K. Østmoe D164.83 [H082] (O-253262); Grimsdalen, Tverrgjelet, on silt along Tverrgjelbekken, 4 Aug. 1989, T. Schumacher G 39.89 [H201] (O-253264); ibid., G.40.89 [H202] (O-253265); ibid., 16 Aug. 1996, S. Landvik & T. Schumacher SL 202.96 [H089] (O-253263); Grimsdalen, Veslegrimsa, on river bed, 8 Aug. 2009, T. Carlsen, I. Skrede & T. Schumacher TS 20.09 [H046] (O-253261); Grimsdalen, Tverrådalen, on moist silt on river slope, 1 Aug. 2007, T. Schumacher TS 14.07 [H035] (O-253260); Grimsdalen, Grimsa at Storberget, on river terrace, 3 Aug. 2007, T. Schumacher TS 38.07 [H034] (O-353259); Grimsdalen, Jegerhøi SE toward Verkenseter, on soil along path, 3 Aug. 2007, T. Schumacher TS 50.07 [H009] (O-253258). – Switzerland, Graubünden, Las Palüds, 1750 m asl, on soil, 29 Aug. 1984, H. Dissing [H203] (O-253266).
Notes — The species is genetically uniform; the Greenland collection has one hsp substitution not found in the rest of the specimens, which are identical. The species appears to be restricted to the arctic-alpine biome. Helvella pygmaea, originally described from Northern Norway (Sommerfelt 1826), and the name sanctioned by Fries (1828), presumably represents an older name for this species, however, there is neither an original illustration nor material of H. pygmaea left in Sommerfelt’s herbarium in O to support this assumption.
Helvella carnosa Skrede, T. Carlsen & T. Schumach., sp. nov. — MycoBank MB820323; hsp barcode GenBank KY784279; rpb2 barcode GenBank KY772529; Fig. 7b
Etymology. From Latin ‘fleshy in consistency’.
Holotype. Norway, Oslo, Malmøykalven, 13 Oct. 2006, O. Smith (O-68100).
Apothecia stipitate-capitate, regularly bilobate, cap 0.5–1.5 cm broad, apothecial margin inflexed (adnate); hymenium brown (5-6E4-8), drying brownish black (6F-G5-8); receptacle surface delicately pubescent; stipe hollow, 3–4 cm high, by 0.3–0.5 cm broad, yellowish and slightly compressed when dried. Medullary excipulum of dense textura intricata, hyphae 3–5 μm broad. Outer excipulum of subhyaline to light brown textura angularis, cells 20–35 × 10–30 μm, outermost cells club-shaped turning out perpendicularly to the surface forming clusters of loose, short-celled, hyaline hyphoid hairs heavily stained in CB. Asci pleurorhynchous, 270–300 × 13–15 μm. Ascospores ellipsoid, 18.0–19.8 × 11.8–12.6 μm. Paraphyses straight, c. 3 μm broad, light brown along the whole length, at tips clavately enlarged to 5–8 μm.
Specimen examined/sequenced. Norway, Oslo, Malmøykalven, 13 Oct. 2006, O. Smith [H146] (O-68100 holotype).
Notes — This new species of the elastica clade is described based on a single specimen from Norway. The molecular specifics are supported with hsp, rpb2 and LSU GenBank sequences of a second specimen from Sweden (cf. Fig. 2). Helvella carnosa resembles H. bicolor in colour, but has a subpubescent exterior and smaller asci and ascospores. In summary, five rpb2, eight hsp and eight LSU substitutions separate it from H. elastica; 11 rpb2, six hsp and eight LSU substitutions from H. danica; 11 rpb2 and six hsp substitutions from H. fistulosa; six rpb2, 16 hsp and 11 LSU substitutions from H. bicolor; and 11 rpb2, seven hsp, and 10 LSU substitutions from H. panormitana.
Helvella confusa Harmaja, Karstenia 17: 43. 1977 — Fig. 3g
Holotype. Denmark, Northern Jutland, Klitmøller, 15 May 1965, K. Toft & H. Dissing (‘H. solitaria’) (H). Isotype (C-F-70807 ‘H. solitaria’).
Synonym. ? Helvella pedunculata Harmaja, Karstenia 18: 57. 1978.
Holotype. Finland, Pohjois-Pohjanmaa, Kiiminki, Keskikylä, 2 July 1975, E. & M. Ohenoja (OULU), not examined.
Misapplied name
– Helvella solitaria sensu Dissing, Dansk Bot. Ark. 25: 41. 1966.
Illustration — Dissing (1966b: f. 10, as H. solitaria).
Descriptions of the species — Dissing (1966b: 42, as H. solitaria), Harmaja (1977a: 43, 1979: 36).
Specimens examined/sequenced. Denmark, Northern Jutland, Klitmøller, on calcareous soil under Picea, 15 May 1965, K. Toft & H. Dissing [H477] (C-F-70807 isotype); ibid., in coniferous tree plantage, 12 June 1981, T. Læssøe 0110 [H438a] (C); Northern Jutland, Nystrup Plantage, Klitmøller, 31 May 1986, D. Boertmann [H438] (C-F-71647). – India, Kupwara, Lolab Valley, on soil in coniferous forest, 2 May 1967, K.S. Thind [H446] (C-F-45513). – Norway, Østfold, Fredrikstad, Kråkerøy, Fuglevik, roadside with spruce, elm and birch, 20 June 1987, R. Kristiansen [H192] (O-253270); Onsøy, Engalsvik, in spruce forest on gravelly sand, 20 June 1981, R. Kristiansen [H253] (O-253271); Buskerud, Gol, in moss along rivulet, 9 June 2014, K. Sæbø [H437] (O-253268); Oppland, Lunner, Søndre Oppdalen, on calcareous ground, 12 July 1987, T.E. Brandrud [H439] (O-129435); Aust-Agder, Grimstad, Landvik, in rich deciduous forest at the road, 24 Apr. 1998, I.-L. Fonneland [H155] (O-71947); Dovre, Grimsdalen, Tollevshaugen, in pine forest reserve, 4 Aug. 2007, T. Schumacher & T. Vrålstad TS 077.07 [H008] (O-253269); Nordland, Rana, Ørtfjellmoen bridge, spring vegetation in calciphilic spruce forest, 20 Aug. 1979, S. Sivertsen [H320] (C-92125); ibid., 24 Aug. 1981, H. Dissing [H386] (C-F-55016); Nordland, Rana, Dunderland, in river bed, 31 Aug. 1998, H. Dissing & S. Sivertsen AAS 08.98 [H313] (C-F-58808). – Sweden, Uppland, Älvkarleby parish, 3 km NE of Gårdskär, on gravelly roadside among bark debris in coniferous forest, 30 June 1965, O. Eriksson & N. Lundquist [H279] (C-F-92132 Fungi Exs. Suec. 3268 ‘H. solitaria P. Karst.’). – Switzerland, Graubünden, Mota Jüda, Val Plavna, 1600 m asl, along path, 30 Aug. 1979, H. Dissing [H331] (C-F-56845); Las Palüds, 1750 m asl, on soil, 29 Aug. 1984, H. Dissing [H092] (O). – USA, Oregon, Sahale Falls, Hood River Meadows, road to Cloud Cap, road side on soil, 9 July 1991, H. Dissing [H449] (C-F-56513).
Notes — We have adopted the binomial H. confusa for this widespread species. Harmaja (1977a) argued that Dissing had misunderstood Karsten’s concept of H. solitaria and introduced the name H. confusa for this species. A Danish specimen that was depicted in Dissing (1966b: f. 10), was selected as the holotype (Harmaja 1977a), a specimen re-examined here by us. Later, Harmaja (1978) erected an additional taxon, H. pedunculata, to include parts of the specimens previously cited as H. solitaria sensu Dissing. Dissing (1966b) noted the close relationship between H. confusa (as H. solitaria) and H. leucomelaena, but kept them separate based on colour, anatomy and ascospore characteristics. The distinct sulcate stipe in H. confusa was maintained as a feature distinguishing it from H. leucomelaena (Dissing 1966b).
We obtained partial sequences from hsp, rpb2 and LSU as molecular specifics to the species circumscription. Two unique rpb2 substitutions separate the non-European from the European specimens, which on their hand are identical. The Indian and North American specimens are separated by one rpb2 substitution. A specimen from Spain [JX 993070] published as H. leucomelaena by Landeros et al. (2015), shares the LSU nrDNA sequence of H. confusa (cf. Appendix 3). Four rpb2, three hsp and five LSU substitutions distinguish H. confusa from the sister species H. leucomelaena.
Helvella corbierei (Malençon) Van Vooren & Frund, Bull. Mycol. Bot. Dauphiné-Savoie 198: 9. 2010
Basionym. Leptopodia corbierei Malençon, Bull. Soc. Mycol. France 43: 95. 1927.
Lectotype. Malençon, Bull. Soc. Mycol. France 43: t. 6, f. 7, selected by Van Vooren & Frund (2010). Epitype designated here: Switzerland, Graubünden, Ramosch, along the river Inn at Resgia, 7 Sept. 1982, H. Dissing (C-F-57133). MycoBank MBT375397.
Misapplied name
– Helvella stevensii sensu Dissing, Dansk Bot. Ark. 25: 125. 1966.
Illustrations — Malençon (1927: pl. 6, f. 7), Dissing (1966b: f. 34, as H. stevensii), Van Vooren & Frund (2010: 7).
Description of the species — Van Vooren & Frund (2010: 6).
Specimens examined/sequenced. Norway, Oppland, Dovre, Grimsdalen, along the road at Storberget, 16 Aug. 2001, T. Schumacher [H178] (O-253273); ibid., 17 Aug. 2001, T. Schumacher [H191] (O-253274); Nordland, Rana, Krogstrand, 40 km E Mo i Rana, in small stream, 7 Sept. 1972, H. Dissing [H390] (C-F-53800). – Switzerland, Graubünden, Ramosch, along the river Inn at Resgia, 7 Sept. 1982, H. Dissing [H392] (C-F-57133 epitype); ibid., 26 Aug. 1984, H. Dissing [H232] (O-253275); Schmelboden, along road and river Landwasser to Brombänz, 2300 m asl, 4 Sept. 1982, L. Petrini & H. Dissing [H391] (C-F-57087). – USA, Oregon, National Forest Lane Co., Mac Donald Forest, Benton Co., on moss grown trunk along stream, 6 July 1991, H. Dissing Oregon 40.91 [H346] (C-F-56503).
Notes — Van Vooren & Frund (2010) adopted the binomial H. corbierei for this species, a disposition approved by us. Helvella corbierei represents the European H. stevensii sensu Dissing (Dissing 1966b, 2000). To support the lectotype (iconotype) selected by Van Vooren & Frund (2010), we have designated an epitype specimen from Switzerland in order to stabilize the use of the name. The sequenced specimens share identical rpb2 sequences and vary in one hsp site. The specimen from Western USA (Oregon) deviates from the European collections by two rpb2 and three hsp substitutions. Two specimens from North America referred to as H. compressa by Landeros et al. (2015) belong here (cf. Appendix 3).
Helvella corium (O. Weberb.) Massee, Brit. Fungus-Fl. 4: 463. 1895 — Fig. 4a
Basionym. Peziza corium O. Weberb., Pilze Nord-Deutschl. 1: 7. 1873.
Synonyms. Aleuria corium (O. Weberb.) Gillet, Champ. France Discomyc. 2: 39. 1880.
Scypharia corium (O. Weberb.) Quél., Enchir. Fung.: 83. 1886.
Lachnea corium (O. Weberb.) W. Phillips, Man. Brit. Discomyc. 204. 1887.
Macropodia corium (O. Weberb.) Sacc., Syll. Fung. 8: 159. 1889.
Fuckelina corium (O. Weberb.) Kuntze, Revis. Gen. Pl. 2: 852. 1891.
Sarcoscypha corium (O. Weberb.) J. Schröt., Krypt.-Fl. Schlesien (Breslau) 3: 59. 1893.
Cyathipodia corium (O. Weberb.) Boud., Icon. Mycol., liste prélim.: 2. 1904.
Leptopodia corium (O. Weberb.) Boud., Hist. Classif. Discomyc. Europe: 239. 1907.
Cowlesia corium (O. Weberb.) Nieuwl., Amer. Midl. Naturalist 4: 380. 1916.
Paxina corium (O. Weberb.) Seaver, North Amer. Cup-fungi (Operculates): 208. 1928.
Holotype. Poland, Georgenberg Landeck, May 1870, Weberbauer (WRSL), not examined. Epitype designated here: Denmark, Mid Zealand, Kirke Hvalsø, Brødlesgård, 2 July 1984, U. Søchting (C-F-71638). MycoBank MBT375398.
Helvella arctica Nannf., Svensk Bot. Tidskr. 31: 60. 1937.
Cyathipodia arctica (Nannf.) M.M. Moser in Gams, Kl. Krypt.-Fl. (Stuttgart) 2a: 89. 1963.
Paxina arctica (Nannf.) E.K. Cash, J. Wash. Acad. Sci. 44(2): 45. 1954.
Holotype. Sweden, Torne Lappmark, Jukkasjärvi, E of Abiskojok, Aug. to Sept. 1928, J.A. Nannfeldt 1523 (UPS). Isotype (C-F-92111- Fung. Exs. Suec. 369 as ‘H. arctica Nannf.’).
Illustrations — Nannfeldt (1937: pl. 1–2), Dissing (1966b: f. 19), Schumacher & Mohn Jenssen (1992: 17).
Description of the species — Nannfeldt (1937: 56, 60 as H. arctica).
Specimens examined/sequenced. Denmark, Mid Zealand, Kirke Hvalsø, Brødlesgård, 2 July 1984, U. Søchting [H352] (C-F-71638 epitype). – Norway, Hordaland, Ulvik, Finse, at railway station, 29 July 2014, T. Schumacher [H453] (O-253279); Finse, Blåisen, on 1960 moraine, 11 Aug. 1996, K. Høiland [H248] (O-253277); ibid., 27 July 2014, T. Schumacher [H451] (O-253278); Troms, Tromsø, Ringheim, on sandy soil, 23 July 1989, A. Holst Jensen & S. Såstad [H434] (O-253280). – Russia, Tjumenskaja oblast, northern Urals, Labytnangi, with Salix on rich clay, 14 Aug. 1990, H. Knudsen [H294] (C-F-16568); Ural Mts, Krasnij Khamei, 40 km W of Labytnangi, 20 July 1992, H. Kotiranta [H328] (C-F-34476). – Svalbard, Bunsow Land N Longyearbyen, around Gipselva, among Dryas and Salix polaris, 10 Aug. 1985, H. Dissing [H324] (C-F-55580); Kongsfjord, Ny-Ålesund, Hollenderhaugen at Thiisbukta, on silt near the shore, 3 Aug. 1986, G. Gulden & K.M. Jenssen GG 142.86 [H242] (O-362201); Kongsfjord, Ny-Ålesund, at the Bay river, 5 Aug. 1988, T. Schumacher [H436] (O-253281). – Sweden, Torne Lappmark, Jukkasjärvi, E of Abiskojok, Aug. to Sept. 1928, J.A. Nannfeldt 1523 [H292] (C-F-92111-Fung. Exs. Suec. 369 isotype of H. arctica Nannf.).
Notes — Traditionally, H. corium has been interpreted as a highly variable species colonizing a range of habitats and biomes on the European continent. We have examined a selection of specimens curated as H. corium in the fungaria of (O) and (C) and have discovered five phylogenetic species, viz. H. corium s.str., H. alpina, H. nannfeldtii, H. alpestris and H. macrosperma. The H. corium morphospecies complex forms two well-supported lineages, i.e., an alpina-corium lineage and an alpestris-nannfeldtii lineage. Helvella corium and H. alpina differ in 10 hsp and 10 rpb2 substitutions. The alpina-corium lineage has morphological features that separate it from the alpestris-nannfeldtii lineage, i.e., smaller asci (< 300 μm in length), an aporhynchous ascus base, and ascospores less than 20 μm long, compared to asci exceeding 300 μm in length, a pleurorhyncous ascus base, and ascospores up to 23 μm in length in the alpestris-nannfeldtii lineage. Macromorphologically, the most notable features that separate H. corium and H. alpina are apothecium size and stipe length; in H. corium the breadth of the apothecium and the stipe length are approximately 1 : 1 and the tomentum of hyphoid hairs forms macroscopical ‘warts’, while in H. alpina the apothecia are slender with a stipe two to three times longer than apothecium breadth and with a receptacle surface of scattered distributed short hairs. Helvella corium occurs from sea level to high altitude elevations, while the black ‘look-alike’ H. alpina has a more restricted distribution at higher altitudes (subalpine and alpine zone) of Northern and Central Europe and Canada. The holotype of H. corium in WRSL is old and presumably unsuitable for molecular identification and is therefore supported by an epitype from Denmark. The sequenced specimens of H. corium from Norway, Denmark, Svalbard and Russia show variation in one rpb2 and one hsp character.
Helvella costifera Nannf., in Lundell & Nannfeldt, Fungi Exsicc. Suecici Fasc. 41–42: 37, no. 2061. 1953 — Fig. 3h
Basionym. Peziza costata Fr., Acta Reg. Soc. Sci. Ups., Ser. 5, 1: 120. 1851.
Synonym. Acetabula costifera (Nannf.) Benedix, Westfälische Pilzbriefe 5: 113. 1965.
Holotype. Sweden, Uppland, Uppsala, Botanical Garden, E. Fries sub nomen Peziza costata Fr. (UPS), not examined. Epitype designated here: Sweden, Uppland, Uppsala, Carolinaparken, 9 July 1948, J.A. Nannfeldt no. 9956 (C-Fungi Exs. Suec. 2061). MycoBank MBT375399.
Misapplied name
– Acetabula ancilis sensu Boud., Icon. Mycol., livr. 7: no 181, pl. 244. 1906 (preliminary text with ‘circulaires’).
Illustration — Landeros et al. (2012: f. 23).
Apothecia regularly stipitate-cupulate, cup 1.5–4 cm broad, 1–4 cm high, hymenium greyish (6-7E1-3), drying dark greyish brown (6E3), receptacle surface subpubescent, concolorous with hymenium when fresh, when dried pale greyish brown; stipe distinct, hollow, with 5–6 prominent yellowish white blunt-edged ribs (4A2-3) extending onto receptacle, ribs dichotomously branched apically, not interconnected, remaining whitish in dried specimens. Outer excipulum of textura prismatica to textura angularis, outermost cells with light brown walls. Asci pleurorhynchous, 230–270 × 12–15 μm. Ascospores broadly ellipsoid, 14.2–16.6 × 10.6–13.6 μm. Paraphyses 2.0–3.2 μm broad, straight, septate, gradually enlarged to 4.5 μm at tips.
Specimens examined/sequenced. Denmark, Zealand, Copenhagen, Fredriksberg Park, 7 July 1984, J. Vesterholt JV 615.84 [H383] (C-F-92113). – Norway, Akershus, Bærum, Øverland Arboretum, 27 July 2007, P.A. Bergersen [H131] (O-68514); Oppland, Dovre, Grimsdalen, Kvannbekken at Gravhøi, 12 Sept. 1982, T. Schumacher & K. Østmoe 507.82 [H347] (O-253284); Grimsdalen, Tverråi, 940 m asl, Aug. 1985, T. Schumacher [H209] (O-253282); Vågå, Brunskardsknappene, 1180–1200 m asl, on soil, 29 Aug. 1998, S. Kjølner [H247] (O-253283); Sør-Trøndelag, Oppdal, Vinstradalen at Ryphuskollen, on rich calcareous soil, 11 Aug. 1984, T. Schumacher & K. Østmoe TS 15.84 [H227] (O-253285); Finnmark, Cirgojokka, Vassbotndalen, Bjørnelva to Scaccavaggi, in Dryas vegetation, 25 Aug. 1986, H. Dissing [H388] (C-F-53529); ibid., 27 Aug. 1986, H. Dissing [H385] (C-F-53538). – Sweden, Uppland, Uppsala, Carolinaparken, 9 July 1948, J.A. Nannfeldt 9956 [H298] (C-Fungi Exs. Suec. 2061 epitype).
Notes — We have identified two widespread species of the H. costifera morphospecies complex, i.e., H. costifera and H. calycina nom. nov. (= Boletus calyciformis Battarra), separated by one hsp and two rpb2 substitutions. Dissing (1966b) included both species in his concept of H. costifera stating that “collections with a darker, brownish colour have been seen from Norway”. Some of these specimens also have a whitish margin, due to colourless hyphae in the outer excipulum. These latter specimens apply to H. calycina (see this species above). Landeros et al. (2012) examined an isolectotype specimen of H. costifera and the holotype specimen of H. hyperborea, and placed them into synonymy based on morphological features. We have found good reasons to keep these two species apart (see H. hyperborea).
Helvella crispa (Scop.) Fr., Syst. Mycol. 2: 14. 1822 — Fig. 8c
Basionym. Phallus crispus Scop., Fl. Carniol., ed. 2, 2: 475. 1772; Fr., Syst. Mycol. 2: 14. 1822.
Synonym. Costapeda crispa (Scop.) Falck, Sluzowce Monogr., Suppl. 3: 401. 1923.
Lectotype. P. Micheli, Nov. Pl. Gen., t. 86, f. 7, 1729, selected by Dissing (1966b). Epitype designated here: Sweden, Blekinge parish, Karlskrona, Wämö, 5–14 Oct. 1946, S. Lundell & S. Wikland (C-Fungi exs. Suec. 2062). MycoBank MBT375400.
Helvella pallida Schaeff., Fung. Bavar. Palat. Nasc. 4: 112. 1774.
Helvella crispa a. (forma) pileo albo, pallescente Fr., Syst. Mycol. 2: 14. 1822 (Helvella ‘albida’ Schaeff. = lapsus calami for H. pallida Schaeff.).
Lectotype designated here: Schaeff., Fung. Bavar. Palat. Nasc. 3: t. 282. 1772 (‘Elvela vicesima nona’). MycoBank MBT375690.
Helvella alba Bergeret, Phytonom. Univ. 1: t. 145. 1783 (= H. crispa a. (forma) fide Fries (1822)).
Helvella nivea Schrad., J. Bot. 2: 66. 1799 (= H. crispa a. (forma) fide Fries (1822)).
Illustrations — Boudier (1905–1910: pl. 225), Dissing (1966b: f. 21).
Description of the species — Dissing (1966b: 85).
Specimens examined/sequenced. England, Surrey, Oxshott Heath, roadside, 16 Oct. 1994, E.W. Brown [H105] (FH). – Latvia, Riga, on soil, 6 Sept. 1982, D. Pfister [H107] (FH). – Norway, Akershus, Asker, Leangbukta, under pine and beech, 15 Oct. 2004, P. Marstad 326.04 [H135] (O-187744); Nordland, Andøy, Oksbåsen, sand dunes at sea shore, 13 Sept. 2005, B.H. Larsen 235 [H235] (O-360158). – Sweden, Blekinge parish, Karlskrona, Wämö, 5–14 Oct. 1946, S. Lundell & S. Wikland [H408] (C-Fungi Exs. Suec. 2062 epitype); Öland, Böda parish, Byerum, along wayside in forest, 24 Aug. 1960, R. Schöldström [H405] (C-Fungi exs. Suec. 3263); Halland, Spendshult, rich soil, 17 Sept. 2009, L. Walter & A. Molia [H062] (O-253287).
Notes — We have selected an epitype specimen from Southern Sweden that deviates from the Latvian and North Norwegian specimens in two rpb2 characters.
Helvella danica Skrede, T. Carlsen & T. Schumach., sp. nov. — MycoBank MB820330; hsp barcode GenBank KY784378; rpb2 barcode GenBank KY772627; Fig. 7f
Etymology. From Latin ‘described from Denmark’.
Holotype. Denmark, Jutland E, Hobro Østerskov, 18 Sept. 2008, J. Heilmann-Clausen (C-F-85205).
Misapplied names
–Helvella esculenta sensu Hornem., Fl. Dan., fasc. 26: pl.1559. 1816.
–Helvella pulla sensu Rostr. in Lange, Nomenclator ‘Florae Danicae’: 54. 1887.
Illustration — Hornemann (1816: pl.1559, as H. esculenta).
Apothecium stipitate-capitate, small to medium-sized, cap campanulate or slightly irregularly lobate, 1.2–3.0 cm broad, of similar height, hymenium brownish yellow (5C6-8), receptacle surface white to yellowish, smooth; stipe short, 0.5–0.8 cm broad, 1.5–3.5 cm high, whitish to yellowish, hollow, often with grooves towards the base. Medullary excipulum of loosely interwoven textura intricata, hyphae 3–5 μm broad. Outer excipulum of isodiametric textura angularis to textura globulosa, innermost cells angular, subhyaline, turning out in rows of clavate to subglobose cells perpendicularly to receptacle surface, cells heavily stained in CB. Asci pleurorhynchous, 300–340 × 14–16 μm. Ascospores ellipsoid, 19.5–22.4 × 12.2–13 μm. Paraphyses subhyaline, 3–5 μm broad, septate, broadly cylindrical in upper part, generally broadest below tips (spadiceiform), to 10–14 μm broad.
Specimens examined/sequenced. Denmark, Jutland E, Hobro Østerskov, 18 Sept. 2008, J. Heilmann-Clausen [H263] (C-F-85205 holotype); Zealand, Gentofte, Insulin Mosen, rich soil together with Parascutellinia carneosangjuinea and H. elastica, 18 Sept. 1974, D. Boertmann [H396] (C-F-64512). – Norway, Akershus, Eidsvoll, at Eidsvollbygningen, in garden, 16 Sept. 2009, I. Skrede & T. Schumacher [H058] (O-253288); Finnmark, Alta, Altaelva at Lille Raipas, on sandy soil in upper inundation zone, 7 Aug. 1983, H. Edvardsen & K. Høiland [H177] (O-129543). – Russia, St. Petersburg, near town, on soil, 28–29 Aug. 1982, D. Pfister [H111] (FH).
Notes — This short-stipitate, campanulate, medium-sized species with brown hymenium and white stipe of the elastica clade (clade E) appears to have been overlooked by early taxonomists, except Hornemann (1816) who depicted the species in Flora Danica, but misapplied the binomial H. esculenta for this species. Helvella danica is distinguishable macromorphologically by the short stipe and often clustered growth habit of the medium brown (hymenium) and white (stipe) apothecia, by large asci and ascospores, and by a distinctive shape (spadiceiform) of the paraphyses under the microscope. The species resembles H. bicolor and H. capucina when fresh; the two latter species have a darker brown hymenium. Our sample of H. danica constitutes a well-supported sublineage of the elastica clade (clade E), separated from a sublineage of H. fistulosa, H. panormitana, H. monachella and H. capucina of the capucina-danica lineage by four hsp and eight rpb2 substitutions.
Helvella dryadophila Harmaja, Karstenia 17: 58. 1977 — Fig. 3d
Holotype. Norway, Oppland, Lom, Høyrokampen, 29 Aug. 1957, F.-E. Eckblad (O-72617)
Illustration — Harmaja (1977b: f. 5).
Description of the species — Harmaja (1977b: 48).
Specimens examined/sequenced. Canada, Ellesmere Island, Alexandra Fjord, near Fox hill, in moss among Dryas and Salix arctica, 1 Aug. 1984, H. Dissing [H335] (C-F-56772). – Greenland, Ella Island, E of Lauge Kock’s house, near Lange sea, on soil with Dryas, 4 Aug. 1982, H. Dissing & S. Sivertsen [H379] (C-F-50346); Mestervig, behind Nyhavn, in sand at the riverside, 17 Aug. 1982, H. Dissing & S. Sivertsen [H378] (C-F-52701). – Norway, Oppland, Lom, Høyrokampen, 29 Aug. 1957, F.-E. Eckblad [H302] (O-72617 holotype); Dovre, Grimsdalen, Tverrgjelet, in Dryas association on moist slopes, 16 Aug. 1996, S. Landvik & T. Schumacher SL 201.96 [H183] (O-253308); ibid., 20 Aug. 1997, S. Landvik & T. Schumacher TS 35.97 [H180] (O-253307); ibid., 21 Aug. 2005, T. Schumacher [H190] (O-253309); ibid., 2 Aug. 2007, T. Schumacher [H010] (O-253306). – Svalbard, Bunsow Land, in Dryas association, 10 Aug. 1985, H. Dissing [H323] (C-F-55578); ibid., [H321] (C-55579); ibid., [H322] (C-F-55597); Kongsfjord, Gluudneset, ‘Gåsebu’, 2 Aug. 1986, G. Gulden & K. Mohn Jenssen GG 111.86 [H234] (O-363304).
Notes — The sequenced specimens, including collections from Svalbard, Greenland, Norway and Canada, vary in one rpb2 site. Abbott & Currah (1997) compared the type specimens of H. dryadophila, H. arctoalpina and H. verruculosa, and placed them in synonymy under the latter, older name. Landeros et al. (2012) studied the types of H. dryadophila and H. verruculosa and concluded they were conspecific. However, they did not study the type of H. arctoalpina. Since the older name H. verruculosa may apply to either of the two, i.e., H. arctoalpina or H. dryadophila, it is unclear to us to which name Saccardos’ species applies. We have therefore avoided using the binomial H. verruculosa in the present synopsis (see Appendix 1).
Helvella elastica Bull., Herb. France 6: t. 242. 1785 — Fig. 7d
Basionym. Helvella elastica Bull., Herb. France 6: t. 242. 1785; Fr., Syst. Mycol. 2: 21. 1822.
Synonyms. Leptopodia elastica (Bull.) Boud., Icon. Mycol., liste prélim.: 2.1904 (preliminary text with ‘circulaires’).
Tubipeda elastica (Bull.) Falck, Mykol. Untersuch. Ber. 1: 401. 1923.
Lectotype. Bull., Herb. France 6: t. 242, f. A–B, D–G. 1786, selected by Abbott & Currah (1997). Epitype designated here: Sweden, Uppland, Älvkarleby parish, Billudden, N Källvik, on bare sandy soil, 29 Sept. 1946, G. Sandberg & R. Santesson (C-F-29130 Fungi Exs. Suec. 2064 ‘H. elastica Bull. ex Fr.’). MycoBank MBT375401.
Illustration — Bulliard (1786: t. 242, f. A–B, D–G).
Apothecia stipitate-capitate, cap 1.0–3.5 cm broad, 1.5–3.0 cm high, at first irregularly lobed to bilobate with an adnate apothecial margin adhered to stipe, then sometimes irregularly folded with margin free from stipe; hymenium light brown to brown (5-7D4-8), sometimes decoloured, receptacle surface subpubescent, whitish to yellowish, concolorous with stipe; stipe terete, 0.5–1.2 cm broad, 3–15 cm high, solid to hollow, depressed and flat when dry. Subhymenium and medullary excipulum of highly gelatinous hyphae, embedded in an amorphic gelatinous matrix, hyphae c. 3 μm broad. Outer excipulum of angular cells up to 40 × 30 μm diam, outermost cells giving rise to 2–4-celled club-shaped extensions making up the subpubescent surface. Asci pleurorhynchous, 330–360 × 15–17 μm. Ascospores oblong ellipsoid, 19–22 × 10.6–12.2 μm. Paraphyses c. 3 μm broad below, septate, light brown in colour, at tips irregular clavate to 6–10 μm broad.
Specimens examined/sequenced. Norway, Nannestad, Åsgreina, Damvegen, in mossy garden, 30 Aug. 2009, Ø. Stensrud TS 35.09 [H443] (O-253310). – Russia, Sakha, Khangalasskiy Ulus, Sasabyt River 2 km NW Bestyakh, under Larix on calcareous soil, H. Knudsen [H329] (C-F-51672). – Sweden, Uppland, Älvkarleby parish, Billudden, N Källvik, on bare sandy soil, 29 Sept. 1946, G. Sandberg & R. Santesson [H372] (C-F-29130 Fungi Exs. Suec. 2064 ‘H. elastica Bull. ex Fr.’ epitype); Uppland, Vassunda parish, Örsand, close to lake Mälaren, 27 Oct. 1935, E. Krantz [H414] (C- Fungi Exs. Suec. 271 ‘H. elastica Bull. ex Fr.’); Halland, Halmstad, Almberget, 16 Sept. 2009, G. Holden [H066] (O-253311).
Notes — Helvella elastica, as circumscribed here, represents a much narrower species concept than commonly outlined in literature. As the name indicates, the species is elastic due to the gelatinous receptacle and stipe tissues. The gelatinous matrix of the excipulum also characterises the morphologically similar but genetically divergent H. panormitana and H. capucina. Dissing (1966b) included H. panormitana, H. fistulosa, as well as the American H. capucinoides (1912), H. gracilis, and H. adherens (Peck 1902) in his broad concept of H. elastica. Zhuang (2004) described a new species of the elastica group, i.e., H. jimsarica from China, distinguished from H. elastica on the basis of larger ascocarps and occurrence on rotten wood. Helvella elastica deviates from the putative North American H. capucinoides in two rpb2 characters (see this taxon under non-European taxa). A published LSU sequence from Estonia [AJ 972411] referred to as H. albella, is suggestive of H. elastica (Landeros et al. 2015) (cf. Appendix 3).
Helvella fallax Quél., Bull. Soc. Bot. France 23: 331. 1876 — Fig. 9a
Lectotype designated here: Quél., Bull. Soc. Bot. France 23: t. 3, f. 15. 1876. MycoBank MBT375691. Epitype designated here: France, Savoie, Bonneval-sur-Arc, Rau du Chalanson, 2300–2500 m asl, 2 Sept. 1992, E. Horak (C-F-84621). MycoBank MBT375402.
Synonym. Helvella nigricans Pers., Observ. Mycol. 1: 72. 1796, nom. illeg. (homonym).
– non Helvella nigricans Schaeff., Fung. Bavar. Palat. Nasc. 4: 102. 1774 (‘Elvela’) (= H. lacunosa Afzel.).
Misapplied names
– Helvella atra sensu Fr., Syst. Mycol. 2: 19. 1822.
– Helvella atra sensu Holmsk., Beata Ruris Otia Fung. Dan. 2: 47. 1799.
– Helvella atra sensu Dissing, Dansk Bot. Ark. 25: 121. 1966.
– Leptopodia atra sensu Boud., Icon. Mycol., liste prélim.: 2. 1904.
Illustrations — Holmskjold (1799: pl. 25, as H. atra), Boudier (1905–1910: pl. 238, as L. atra), Filippa & Baiano (2011: f. 13–15, as H. atra).
Descriptions of the species — Holmskjold (1799: 47 as H. atra), Dissing (1964: 112, 1966b: 121 as H. atra).
Specimens examined/sequenced. Canada, Northwest Territories, District of Keewatin, Baker Lake NE, sandy river shore with Salix herbacea, 16 Aug. 1974, E. & M. Ohenoja [H430] (C-F-92108 ex Herb. Univ. Ouluensis). – France, Savoie, Bonneval-Sur-Arc, Rau du Chalanson 2300–2500 m asl, 2 Sept. 1992, E. Horak [H339] (C-F-84621 epitype); Savoie, Val d’Isere, Gorges du Mal, on river terrace, 2250 m asl, 31 Aug. 1992, T. Schumacher F 76.92 [H219] (O); Savoie, Les Vallions, Nat. Park Vanoise, on river terrace, 2600 m asl, 28 Aug. 1992, T. Schumacher F 28.92 [H221] (O). – Norway, Hedmark, Løten, Rokoberget, 17 Oct. 2006, R. Haugan, S06005 [H162] (O-280610); Hedmark, Kongsvinger, Bogeråsen, in spruce forest, 6 Aug. 1998, I. Tangen & K.H. Amundsen [H168] (O-63684); Oppland, Dovre, Grimsdalen, Tverrådalen, 13 Aug. 1985, T. Schumacher [H402] (O-253354); ibid., 12. Aug. 1996, A. Holst Jensen [H256] (O-253353); ibid., 7 Aug. 2009, T. Carlsen, T. Schumacher & I. Skrede TS 10.09 [H018] (O-253351); Grimsdalen, Veslegrimsa, on river terrace, 8 Aug. 2009, T. Carlsen, T. Schumacher & I. Skrede TS 19.09 [H032] (O-253352); Grimsdalen, 16 Aug. 2001, T. Schumacher [H002] (O-253350); Oppland, Lom, N Bøvertjønnen, 950 m asl, 20 July 2002, R.Y. Berg [H169] (O-66873); Møre og Romsdal, Sunndal, Sunndalsfjorden N Hisdalen, under Corylus, 11 Sept. 2001, J.B. Jordal [H163] (O-242201); Nordland, Rana, Hammarnes, 15 km N Mo i Rana, rich mouldy soil, 18 Aug. 1979, H. Dissing [H419] (C-F-54652); Troms, Gratangen, Finnesletta, calcareous pasture land, 10 Sept. 2005, B.H. Larsen [H244] (O-360225). – Sweden, Lule Lappmark, Gällivarre Parish, 6 km SE Ritsemjokk train station, moist fen at rivulet with Sphagnum and dwarf willows, 22 Aug. 1970, G. Lohammar [H395] (C-F-45484). – Switzerland, Graubünden, Albula Pass, near Crap Alv, 2040 m asl, 30 Aug. 1984, T. Schumacher [H428] (C-F-57377).
Notes — We have adopted the binomial H. fallax for this species, a species described and depicted from France by Quélet (1876). Helvella nigricans, which was referred to as a synonym of H. atra by Fries (1822, 1832), represents an older illegitimate name (homonym) for the species. Persoon (1796) compared his fungus with H. atra sensu Afzel., leaving no doubt about which fungus he had in hand. Fries’ broad concept of H. atra, including H. atra J. König, H. atra sensu Holmsk. (1799) and Afzelius (1783) and H. nigricans Pers. as well, was not accepted by Dissing (1966b), who adopted Holmskjolds concept of H. atra as an available name for the present species. Since Fries (1822, 1832) clearly assigned and credited König to the sanctioned epithet of ‘atra’, we have reserved H. atra for König’s fungus and adopted Quélet’s name (i.e., H. fallax) for the illegitimate H. nigricans. Helvella fallax is here lectotypified by Quélet’s original figure, supported with an epitype specimen from France that provides partial sequences from four genes, e.g., hsp, tef, rpb2 and LSU as molecular specifics to the species. The sequenced specimens of H. fallax from France have identical LSU and hsp sequences, but deviate from the rest of the collections from Northern Europe in one rpb2 and three LSU substitutions. A collection from Estonia [AJ 972413] recorded by Landeros et al. (2015, as H. atra) is also nested here (cf. Appendix 3).
Helvella fibrosa (Wallr.) Korf, Mycotaxon 103: 311. 2008 — Fig. 9b
Basionym. Peziza fibrosa Wallr., Fl. Crypt. Germ. 2: 498. 1833 (typus: Octospora villosa Hedw.).
Synonyms. Macropodia fibrosa (Wallr.) Sacc., Syll. Fung. 8: 160. 1889.
Octospora villosa Hedw., Descr. Micro-anal. Musc.: 54. 1789.
Peziza macropus var. villosa (Hedw.) Pers., Syn. Meth. Fung. 2: 646. 1801.
Peziza macropus β (var.) hirta (Hedw.: Fr.) Fr., Syst. Mycol. 2: 57. 1822.
Fuckelina villosa (Hedw.) Kuntze, Revis. Gen. Pl. 2: 852. 1891.
Cyathipodia villosa (Hedw.) Boud., Icon. Mycol., liste prélim.: 2. 1904.
Leptopodia villosa (Hedw.) Arnould, Bull. Soc. Mycol. France 9: 111. 1893.
Helvella villosa (Hedw.) Dissing & Nannf., Svensk Bot. Tidskr. 60: 330. 1966, nom. illeg. (homonym)
Helvella dissingii Korf, Mycotaxon 31: 381. 1988.
Lectotype. Hedw., Descr. Micro-anal. Musc.: t. 19, f. B, no. 3. 1789, selected by Dissing & Nannfeldt (1966). Epitype. Sweden, Halland, Onsala parish, Presse, Björkhamra, 8 July 1956, F. Karlvall 6944 - Fungi Exs. Suec. 3262 ‘H. chinensis’ (Velen.) Nannf. & L. Holm (UPS), selected by Korf (2008).
Peziza sublicia Holmsk., Beata Ruris Otia Fung. Dan. 2: 26. 1799.
Lectotype designated here: Holmsk., Beata Ruris Otia Fung. Dan. 2: pl. 10. 1799. MycoBank MBT375692.
Illustration — Dissing & Nannfeldt (1966: pl. 2, as H. villosa).
Descriptions of the species — Dissing & Nannfeldt (1966: f. 3, pl. 2, as H. villosa), Dissing (1966b: 67, as H. villosa), Weber (1972: 171, as H. villosa).
Specimens examined/sequenced. Denmark, Falster, Gedesby, in garden under Corylus, 7 July 1998, P. Corfixen [H426] (C-F-38089). – Finland, Oulun Pohjanmaa, Haukipudas, Niemeläntörmä, Höyhtyä farm, 26 July 1981, T. Ulvinen [H369] (C-F-92109). – Georgia, Kaukasus, Kasbhegi, Gvethi valley, sandy soil with herbs, along trail, 2000 m asl, 13 July 1988, E. Johannesen [H250] (O-253313). – Iceland, Rang., Merkurhraun, forest near Merkihvoll, among moss on gravel in Betula forest, 26 Aug. 1984, S.A. Elborne [H343] (C-F-92103); Borg., Borgarfjördur, Hafnarskogur, 18 Aug. 1984, H.F. Gøtzsche [H344] (C-F-92102). – Norway, Akershus, Bærum, Kalvøya, on calcareous soil under Corylus, 28 Aug. 1988, T. Schumacher [H164] (O-88570); Vestfold, Sande, Bekkestranda, 12 Sept. 1985, P. Marstad 135.85 [H166] (O-185923); Vestfold, Tønsberg, Sem, Slagentangen, 11 Sept. 1985, P. Marstad [H167] (O-185919); Hedmark, Stange, Roa nordre, pasture land, 22 Sept. 2000, S. Sivertsen, P.G. Larsen, T.S. Nilsen, A. Molia & T. Pousi [H139] (O-63973); Sør-Trøndelag, Oppdal, Vinstradalen, Nordistuseter at the road, rich pasture land, 28 Aug. 2008, J.B. Jordal [H240] (O-291352); Oppdal, Loslia, rich pasture land, 1000 m asl, 20 Aug. 2009, J.B. Jordal [H243] (O-291377); Nordland, Rana, Rausandsaksla, 15 km E Mo i Rana, calcareous soil, 11 Sept. 1975, T. Schumacher [H270] (C-F-54475); Rana, Store Røvatn, 20 km NE of Mo i Rana, on calcareous soil under Salix, 6 Sept. 1972, H. Dissing [H357] (C-F-53774). – Sweden, Halland, Onsala parish, Presse, Björkhamra, 8 July 1956, F. Karlvall 6944 ‘H. chinensis’ (Velen.) Nannf. & L. Holm’ isoepitype [H413] (C-Fungi Exs. Suec. 3262).
Notes — Helvella fibrosa apparently has a worldwide distribution. It has been confused with H. macropus on many occasions (cf. Dissing & Nannfeldt 1966). Korf (2008) gave an extensive treatment of name use and nomenclature of this characteristic species. There exist several 18th century descriptions and coloured iconographs of H. fibrosa, although being presented with other binomials, i.e., in Hedwig (1789: 54, pl. 19, f. B, no. 3, as Octopsora villosa), Persoon (1796: 26, pl. 1, f. 2. as Peziza macropus var. β. lacunosa) and Holmskjold (1799: 26, pl. 10, as Peziza sublicia). The hymenium is darker (greyish brown) and the receptacle surface less villose in H. fibrosa compared to H. macropus. Moreover, the ascospores are broadly ellipsoidal and not acuminate as in H. macropus. The material studied by us is invariable in hsp and rpb2 sequences, except for the Georgian specimen that deviates in one rpb2 bp character. Häffner (1987), Abbott & Currah (1997) and Landeros et al. (2015) placed H. pallidula in synonymy with H. chinensis (= H. fibrosa), a disposition which needs to be re-assessed based on molecular data. Helvella ‘Japan 1’ and Helvella ‘Japan 2’ are morphologically similar to H. fibrosa, but deviates in a number of rpb2 and hsp characters.
Helvella fistulosa Alb. & Schwein., Consp. Fungorum Lusat.: 299. 1805 — Fig. 7h
Basionym. Helvella fistulosa Alb. & Schwein., Consp. Fungorum Lusat.: 299. 1805; Fr., Syst. Mycol. 2: 21. 1822.
Synonyms. Leptopodia fistulosa (Alb. & Schwein.) Boud., Hist. Classif. Discomyc. Europe: 37. 1907.
Helvella elastica forma fistulosa (Alb. & Schwein.) Rehm, in Winter, Rabenh. Krypt.-Fl., ed. 2 (Leipzig) 1.3 (lief. 53): 1184. 1895.
Neotype designated here: Norway, Hordaland, Ulvik, Ossete, Nipane, on lime-phyllite gravel, 5 Sept. 2009, G. Flatabø [H241] (O-291887). MycoBank MBT375403.
Apothecia medium-sized, 0.5–1 cm broad, 1–2 cm high, bi- to trilobate to irregularly lobed, margin reflexed and free; hymenium brown (6D-E7-8), outside subpubescent, yellowish, concolorous with stipe; stipe terete, solid or hollow, inflated and occasionally reddish brown when dried, 0.3–0.6 cm broad, by 2–7 cm high. Medullary excipulum of loosely interwoven textura intricata, hyphae 3–4 μm broad. Outer excipulum a mixture of broad-celled textura intricata, hyphae 6–10 μm broad and textura angularis, angular cells 20–35 μm diam, towards surface cells more club-shaped, to 40–50 μm long, forming 3–4-celled, ramified extensions on the excipulum exterior. Asci pleurorhynchous, 300–320 × 14–17 μm. Ascospores ellipsoid, 19.0–21.5 × 12–13.2 μm. Paraphyses straight, c. 3 μm broad below, subhyaline, septate, clavate to subcapitate to 6–8 μm at the tips.
Specimens examined/sequenced. Iceland, Austur-Skaftafellsysla, Skaftafell, 20 Aug. 1981, H.F. Gøtzsche HFG 81.32 [H476] (C-F-45406). – Japan, Honshu, Tochigi, Okunikko, Chuzenjikohan, Asegata, on soil in forest, 25 Aug. 1983, T. Schumacher [H205] (O-253314). – Norway, Akershus, Bærum, pathway in forest, 16 Aug. 2001, T. Schumacher [H179] (O-253315); Hordaland, Ulvik, Ossete, Nipane, on lime-phyllite gravel, 5 Sept. 2009, G. Flatabø [H241] (O-291887 neotype). – USA, Massachusetts, Worchester county, Lancaster, Devens Reserve Forces Training Area, on soil and debris in hemlock-Northern hardwood forest, 9 July 1999, B. Neill, G. Valiant, W. Metclaf, M. Jacobs, D. Pfister, K. Griffith D-435 [H109] (FH); Vermont, Windsor county, Norwich, on soil in hemlock forest with some pines and occasional hardwoods, 29 Apr. 2004, K. Griffith [H114] (FH).
Notes — Helvella fistulosa is morphologically almost indistinguishable from H. danica, however, molecular data clearly discriminate between the two: five hsp, six rpb2 and eight LSU substitutions separate the species. We have selected a collection from Norway as a neotype for this intercontinental species, here supplied with partial sequences of hsp, rpb2 and LSU. The North American, European and Japanese specimens vary by one rpb2 bp substitution.
Helvella fusca Gillet, Champ. France Discomyc. 1: 9. 1879
Synonyms. Helvella fusca var. bresadolae Boud., Icon. Mycol., liste prélim.: 2. 1904.
Helvella fusca var. gyromitroides Chenant., in Pelé & Chenantais, Bull. Soc. Sci. Nat. Ouest France, sér. 4, 1: 75. 1921.
– non Helvella elastica (Bull.: Fr.) var. fusca Bull., Herb. France 2: 299. 1791.
Helvella elastica (Bull.: Fr.) c. fusca Bull.: Fr., Syst. Mycol. 2: 21. 1822.
– non Helvella sulcata α. (var.) fusca Afzel., Kongl. Vetensk. Acad. Nya Handl. 4: 305, t. 10, f. 1. 1783 (= Helvella sulcata a. (forma) fusca Afzel.: Fr., Syst. Mycol. 2: 16. 1822).
Lectotype. Gillet, Champignons de France, Discom., t. 4. 1879, selected by Landeros & Korf (2012). Epitype. Italy, Al Deserto, 21 May 1898, Bresadola (S-F-124428), selected by Landeros & Korf (2012), not examined.
Illustrations — Gillet (1879: pl. 4), Dissing (1966b: f. 23), Landeros & Korf (2012: f. 1–4).
Descriptions of the species — Dissing (1966b: 92), Landeros & Korf (2012: 434).
Specimens examined/sequenced. Hungary, Kiskunsag National Park, Bugac, in populeto (Populus canescens, P. alba), 11 May 1978, M. Babos & A. Friesz [H305] (C-F-92122). – Netherlands, Zuid-Holland, Oostvoorne, in sandy dunes, 14 May 1983, C. Bas [H304] (C-F-89381).
Notes — Helvella fusca is a spring and early summer species characterized by a stipitate saddle-shaped apothecium, brownish hymenium, and a white, lacunose stipe with ribs extending and ramifying onto the receptacle surface. It grows in association with Populus and is presumed to be an ectomycorrhizal partner with this host. Dissing (1966b) referred H. fusca to his section Crispae based on the saddle-shaped apothecium and the pubescent outside, a disposition that get no support from our phylogeny. The description of H. fusca in Abbott & Currah (1997) probably applies to H. maculata, as also concluded by Landeros & Korf (2012). Landeros et al. (2015) recently supplied a LSU nrDNA sequence from the epitype specimen of H. fusca, which deviates in two bp substitutions from the above cited specimens from Hungary (cf. Appendix 3).
Helvella helvellula (Durieu) Dissing, Rev. Mycol. 31: 204. 1966
Basionym. Peziza helvellula Durieu, Expl. Sci. Algerie 1: t. 27, f. 11. 1849.
Synonyms. Geopyxis helvellula (Durieu) Sacc., Syll. Fung. 8: 65. 1889.
Acetabula helvellula (Durieu) Maire, Bull. Soc. Hist. Nat. Afrique N. 8: 176. 1917.
Lectotype designated here: Durieu, Expl. Sci. Algerie 1: t. 27, f. 11. 1849. MycoBank MBT375693. Epitype designated here: Spain, Canary Islands, Hierro, 4.5 km toward Taibique, SW San Andres, on mossy soil, 7 Jan. 1977, R.P. Korf, R. Fogel, G.L. Hennebert & L.M. Kohn (C-F-92128 Mycoflora of Macaronesia 1412). MycoBank MBT375404.
Misapplied names
– Acetabula clypeata sensu Boud.. Bull. Soc. Mycol. France 3: 146. 1887.
– Paxina clypeata sensu Linder in Polunin, Bull. Natl. Mus. Canada, Biol. ser. 26, 97: 257. 1947.
Illustration — Boudier (1905–1910: pl. 247, as Acetabula clypeata).
Description of the species — Dissing (1966b: 45).
Specimens examined/sequenced. France, Cote Atlantique, Region des Landes, Mimizan, with Pinus maritima, March 1981, F. Candoussau [H309] (C-F-45469); Dpt. Manche, Carteret au nord de phare, 15–25 m asl, 20 Apr. 1977, R. Schumacker 770420/01 [H278] (C-F-45506). – Spain, Canary islands, Hierro, 4.5 km toward Taibique, SW San Andres, on mossy soil, 7 Jan. 1977, R.P. Korf, R. Fogel, G.L. Hennebert & L.M. Kohn [H308] (C-F-92128 ‘Mycoflora of Macaronesia 1412’ epitype).
Notes — This cupulate, black, subsessile Mediterranean species was described by Durieu (1849) based on a specimen from Morocco collected by Montagne. There are no specimens of this species in Montagne’s Herbarium in Paris (PC) (Dissing 1966a). The species is here typified with the original plate of Durieu’s, supported by a widely distributed epitype specimen from the Canary Islands, Spain. According to Dissing (1966a), materials in Boudier’s herbarium, described and depicted as Acetabula clypeata (Boudier 1905–1910: pl. 247) represent H. helvellula. In our phylogeny H. helvellula clusters with H. juniperi and H. semiobruta in a well-supported divergent lineage of the lacunosa clade (clade C).
Helvella hyperborea Harmaja, Karstenia 18: 57. 1978 — Fig. 3e
Holotype. Finland, Kuusamo, Juuma, Jäkälävuoma, 27 Aug. 1970, H. Harmaja (H), not examined.
Illustrations — Harmaja (1979: f. 1–2), Landeros et al. (2012: f. 27).
Description of the species — Harmaja (1979: 35).
Specimens examined/sequenced. Finland, Kuusamo, Liikasenvaara, slope of hill Korvasvaara, 23 Aug. 1978, H. Harmaja [H491] (C-F-45306). – Norway, Nordland, Rana, Rausandaksla, 15 km E Mo i Rana, on calcareous soil, 11 Sept. 1975, T. Schumacher [H389] (C-F-54473).
Notes — Helvella hyperborea belongs in the morphospecies complexes of H. acetabulum and H. costifera/H. calycina, but is clearly distinct judged from the hsp and rpb2 barcode sequences; altogether 12, 11 and 19 rpb2 substitutions distinguish it from the morphologically similar H. costifera, H. calycina and H. acetabulum, respectively. The specimens from Nordland in Northern Norway and Kuusamo in Finland share identical hsp and rpb2 sequences. Landeros et al. (2012) studied the morphology of the type specimen of H. hyperborea and placed it in synonymy with H. costifera. A few additional collections from the Nordic countries, determined and referred to as H. hyperborea by Harmaja (annotations on herbarium specimens), have been sequenced by us and are referred to H. acetabulum, H. calycina or H. costifera. The above cited collection from Finland was photographed in fresh condition and depicted as the archetypic H. hyperborea in Harmaja (1979: f. 1). An attempt to sequence the holotype specimen of H. hyperborea awaits future studies.
Helvella hypocrateriformis Schaeff., Fung. Bavar. Palat.
Nasc. 4: 102. 1774 (‘Elvela’) — Fig. 5a
Synonyms. Peziza hypocrateriformis (Schaeff.) Wallr., Fl. Crypt. Germ. 2: 498. 1833, nom. illeg. (homonym).
– non Peziza hypocrateriformis Bergeret, Phytonom. Univ. 1: t. 169. 1783 (= Helvella macropus (Pers.) P. Karst.).
Macropodia hypocrateriformis (Schaeff.) Sacc., Syll. Fung. 8: 159. 1889.
Cyathipodia hypocrateriformis (Schaeff.) Boud., Hist. Classif. Discomyc. Europe: 39. 1907.
Lectotype designated here: Schaeff., Fung. Bavar. Palat. Nasc. 2: t. 152. 1763 (‘Elvela quinta’). MycoBank MBT375694. Epitype designated here: Sweden, Uppland, Uppsala, Stadsskogen, 21 July 1948, R. Molander (C-F-45378). MycoBank MBT375405.
Helvella cupuliformis Dissing & Nannf., Svensk Bot. Tidskr. 60: 326. 1966.
Cyathipodia cupuliformis (Dissing & Nannf.) J. Breitenb. & F. Kränzl., Champ. Suisse 1. Ascomycetes: 58. 1981.
Holotype. Sweden, Uppland, Uppsala, Kåbo, 30 July 1936, K.G. Ridelius - Fungi Exs. Suec. 3264 (UPS). Isotype. ibid. (C-F-92131).
= Helvella pulchra Dissing, Biblioth. Mycol. 150: 17. 1993.
Holotype. Switzerland, Graubünden, Ramosch, Resgia, along the river Inn, 7 Sept. 1982, H. Dissing Sch 82-151 (ZT), not examined. Isotype. (C-F-57126).
Illustrations — Schaeffer (1763: pl. 152, as ‘Elvela quinta’), Dissing (1966b: f. 17, as H. cupuliformis), Dissing & Nannfeldt (1966: pl. 1, as H. cupuliformis).
Descriptions of the species — Dissing & Nannfeldt (1966: 326, as H. cupuliformis), Dissing (1966b: 71 as H. cupuliformis; 1993: 17, as H. pulchra).
Specimens examined/sequenced. Sweden, Uppland, Uppsala, Stadsskogen, near Kåbo, 21 July 1948, R. Molander [H301] (C-F-45379 epitype); ibid., 30 July 1936, K.G. Ridelius [H300] (C-F-92131, Fungi Exs. Suec. 3264 isotype of H. cupuliformis). – Switzerland, Graubünden, Ramosch, along the river Inn, at the bridge near Resgia, under Picea, 1100 m asl, 7 Sept. 1982, H. Dissing [H275] (C-F-57126 isotype of H. pulchra).
Notes — Helvella hypocrateriformis was referred to as a Peziza similar to P. macropus (= H. macropus) and P. macropus var. hirta (= H. fibrosa) by Fries (1832). Based on Schaeffer’s plate (1763: pl. 152) and Fries’ recognition of Schaeffer’s fungus as a species close to H. fibrosa, we conclude that H. hypocrateriformis represents an older, available name for H. cupuliformis. We have resurrected Schaeffer’s name for this fungus, selecting Schaeffer’s plate as lectotype supported with a sequenced epitype specimen from Sweden. The synonymies of H. pulchra and H. cupuliformis with H. hypocrateriformis are established here based on hsp and rpb2 sequence comparisons; the isotypes of H. pulchra and H. cupuliformis share identical hsp and rpb2 sequences, while the selected epitype of H. hypocrateriformis, collected in the same area as the type specimen of H. cupuliformis, deviates from the latter in one rpb2 bp character. Landeros et al. (2015) supplied an LSU sequence from paratype material of H. cupuliformis in UPS, which deviates from the isoparatype of H. cupuliformis in C sequenced by us. This latter specimen is selected here as an epitype of H. hypocrateriformis, thus merging the concepts of H. hypocrateriformis and H. cupuliformis. Zhuang (2004) erected a variety ‘crassa’ for Chinese specimens of H. cupuliformis with large ascocarps and somewhat broader ascospores than the type variety. This variety probably deserves specific rank.
Helvella juniperi M. Filippa & Baiano, Riv. Micol. 42: 100. 1999
Holotype. Italy, Viareggio, (LU), 16 Mar. 1996, Baiano, Filippa & Garofoli (MCVE-501), not examined.
Illustration — Filippa & Baiano (1999: 100).
Description of the species — Filippa & Baiano (1999: 100).
Specimen examined/sequenced. Portugal, Minho, S of S. Bartolomeu do Mar N of Esposende, in sandy soil under Cistus salvifolius in the dunes, 15 Nov. 2000, P. Boisen Hansen [H415] (C-F-42193).
Notes — This morphologically distinct species of the helvellula lineage of the lacunosa clade (clade C) deviates from the subsessile H. helvellula in two rpb2 and five hsp substitutions, and from H. semiobruta in three rpb2 and five hsp substitutions.
Helvella lactea Boud., Icon. Mycol., liste prélim.: 2. 1904 — Fig. 8a
Lectotype. France, Meaux, June 1902, D. Dumée (PC), selected by Dissing (1966a), not examined.
Illustrations — Dissing (1966a: f. 9b, f. 3, 7e–f; 1966b: f. 29).
Descriptions of the species — Dissing (1966a: 206; 1966b: 108).
Specimens examined/sequenced. Denmark, Zealand, Dyrehaven, N of Copenhagen, rich soil in deciduous forest, 27 Aug. 1961, H. Dissing [H374] (C-F-39379). – Sweden, Uppland, Uppsala, Botanical garden, in sparse loamy lawn, under oaks and hazels, 25 July – 2 Aug. 1939, N. Hylander [H262] (C-Fungi Exs. Suec. 1355).
Notes — Dissing selected a lectotype in Boudier’s herbarium in PC which morphologically matched material from Sweden and Denmark studied by him. The description and illustrations of H. lactea in Dissing (1966b) is representative of the narrowed concept of H. lactea, as reassessed here. Two specimens listed by Dissing (1966b) from Sweden and Denmark have been sequenced by us. We conclude there are two similar species that have been commonly affiliated to the broad concept of H. lactea, i.e., H. lactea and H. pallescens. It may well be that Boudier had the two species in hand when recording H. lactea and illustrating parts of the type material in Icones (Boudier 1905–1910). This conclusion rests on the observation that the slender white specimen depicted on plate 226 of the ‘Icones’ is more indicative of H. pallescens than H. lactea s.str. We also studied a North American white counterpart to H. lactea, referred to as Helvella ‘USA 5’ in the present synopsis (see non-European species), which differs by seven rpb2 and five hsp substitutions.
Helvella lacunosa Afzel., Kongl. Vetensk. Acad. Nya Handl. 4: 303. 1783 — Fig. 6a–b
Basionym. Helvella lacunosa Afzel., Kongl. Vetensk. Acad. Nya Handl. 4: 303. 1783; Fr., Syst. Mycol. 2: 15. 1822.
Synonyms. Helvella lacunosa Fr., Observ. Mycol. 2: 301. 1818, nom. illeg. (homonym).
Costapeda lacunosa (Afzel.) Falck, Sluzowce Monogr., Suppl. (Paryz) 3: 401. 1923.
Helvella lacunosa Afzel. a. (forma) major Fr., Syst. Mycol. 2: 15. 1822 (= H. mitra Schaeff., where epithet ‘mitra’ is an unintentional 1822 error for epithet ‘nigricans’).
Helvella nigricans Schaeff., Fung. Bavar. Palat. Nasc. 4: 102. 1774 (‘Elvela’)
– non Helvella nigricans Pers., Observ. Mycol. 1: 72. 1796 (= H. fallax Quél.).
Lectotype designated here: Schaeff., Fung. Bavar. Palat. Nasc. (Ratisbonae) 2: t. 154. 1763 (‘Elvela septima’). MycoBank MBT375733. Epitype designated here: Sweden, Femsjö parish, Femsjö, N of Skatåker, 4 Oct. 1948, S. Lundell & G. Haglund (C-Fungi Exs. Suec. 2065). MycoBank MBT375406.
Illustrations — Schaeffer (1763: pl. 154, as H. nigricans), Boudier (1905–1910: pl. 228).
Apothecia stipitate-capitate, saddle-shaped, bi- to trilobate, cap 3–6 cm broad, 2–4.5 cm high, apothecial margin reflexed or slightly deflexed when fully mature, attached to stipe at 4–8 points; hymenium black (G-H8), even or wrinkled; receptacle dark grey to greyish black (F-G8), drying blackish, smooth; stipe whitish to greyish, 4–15 cm long, 1.5–3(–6) cm broad, naked, chambered inside and outside with 5–8 sharp-edged, commonly double-edged and partly anastomosing ribs and deep ‘holes’ (lacunae) inbetween. Medullary excipulum of textura intricata, hyphae 3–5 mm broad. Outer excipulum of intermixed broad-celled textura intricata and textura globulosa-angularis, outermost cells club-shaped with brownish walls, 10–30 × 5–15 μm, with short 2–4-celled hyaline outgrowths. Asci pleurorhynchous, 250–290 × 14–17 μm. Ascospores narrowly ellipsoid, 17.0–19 × 9–11.5 μm. Paraphyses straight, subhyaline to light brown below, increasingly brown above, 3–4 μm broad, septate, clavate to 6–8 μm at the tips.
Specimens examined/sequenced. Greenland, Thule, Airbase, N river at Dundas road, clayey silt along coast, 4 Aug. 1987, H. Dissing [H399] (C-F-55985). – Norway, Akershus, Ullensaker, Aurtjern, albino morphotype, 12 Sept. 2009, Ø. Stensrud [H059] (O-253316); Hordaland, Ulvik, Finse, Bio 101-peak, 28 July 2014, T. Schumacher [H470] (O-253318); Kvam, Treskårana, Gjerde – Oma, mixed deciduous forest, 24 Aug. 2005, W. Holm WH 7 [H150] (O-280703); Oppland, Dovre, Grimsdalen, along road at Storberget, 20 Aug. 2001, T. Schumacher [H189] (O-253322); ibid., 8 Aug. 2009, T. Carlsen, I. Skrede & T. Schumacher [H044] (O-253320), [H045] (O-253321); Grimsdalen, Tollevshaugen, 3 Aug. 2007, T. Schumacher [H039] (O-253319); Sør-Trøndelag, Ørland, Storfosna, Haugan, in garden, 6 Oct. 2007, R. Haugan, S07415 [H153] (O-285214); Finnmark, Nordkapp, Magerøya, Kobbholen, among Dryas, 12 Aug. 1995, L. Ryvarden 38048 [H170] (O-59885); Vassbotndalen, from Bjørnelva to Skacca vaggi, in subalpine birch forest, 400 m asl, 27 Aug. 1986, S. Sivertsen 44.86 [H474] (C-53539); Finnmark, F.-E. Ecckblad 189b.61 [H486] (O-253317). – Russia, River Kotuy Khan at junction with river Kotuy, under Larix gmelinii, 19 Aug. 1993, S. Å. Hanssen [H327] (C-F-34477). – Svalbard, Longyearbyen, among mosses, 19 Aug. 1988, H. Knudsen [H471] (C-F-19329). – Sweden, Femsjö parish, Femsjö, N of Skatåker, 4 Oct. 1948, S. Lundell & G. Haglund [H407] (C- Fungi Exs. Suec. 2065 epitype).
Notes — The H. lacunosa morphospecies complex comprises at least four phylogenetic species with ± restricted distributions in Europe, i.e., H. atra, H. nigra, H. sulcata and H. lacunosa s.str. Pale (albino) forms may be mistaken for H. crispa but can be distinguished readily by the smooth outer surface and partial fusing of cap margin with the stipe in H. lacunosa. Morphologically, H. lacunosa resembles H. atra and H. nigra; however, H. lacunosa possesses a receptacle surface and stipe remaining light-coloured and strongly contrasting the hymenium colour when dried. Fries (1822) adopted Afzelius’ name for his broadly defined H. lacunosa and referred to Schaeffer’s description and plate of H. nigricans (erroneously as ‘mitra’ Schaeff.) as synonymous to H. lacunosa forma major (a.) in his sanctioning work (Fries 1822, 1832). Helvella lacunosa is here typified with Schaeffer’s plate, supported with an epitype specimen from Sweden. The rpb2 barcode sequence of H. lacunosa is identical among the sequenced specimens.
Helvella leucomelaena (Pers.) Nannf., in Lundell & Nannfeldt, Fungi Exsicc. Suecici Fasc. 19–20: 21, no. 952. 1941
Basionym. Peziza leucomelas Pers., Mycol. Eur. 1: 219. 1822.
Synonyms. Aleuria leucomelaena (Pers.) Gillet, Champ. France Discomyc.: 37. 1879 (‘leucomelas’).
Acetabula leucomelaena (Pers.) Sacc., Syll. Fung. 8: 61. 1889 (‘leucomelas’).
Paxina leucomelaena (Pers.) Kuntze, Revis. Gen. Pl. 2: 864. 1891 (‘leucomelas’).
Peziza sulcata Pers. b. (forma) Fr., Syst. Mycol. 2: 44. 1822.
Lectotype. Netherlands, Herb. Persoon L 8945 – 6, (L), selected by Abbott & Currah (1997). Epitype designated here: Sweden, Gotland, Klintehamn parish, at the railway station, 7 May 1938, S. Lundell & E. Åberg (C-Fungi Exs. Suec. 952 ‘Helvella leucomelas’ (Pers.) Nannf.). MycoBank MBT375407.
Acetabula calyx Sacc., Mycoth. Veneta: no. 168. 1873 (fide Landeros et al. 2015)
Paxina calyx (Sacc.) Kuntze, Revis. Gen. Pl. 2: 864. 1891.
Illustration — Dissing (1966b: f. 8).
Description of the species — Dissing (1966b: 36).
Specimens examined/sequenced. Chile, Santiago, El Tabo, in pine wood, 11 July 1966, W. Lazo [H447] (C-F-92127). – Denmark, Jutland, Nystrup Plantage 15 km NW Thisted, calcareous soil, 25 Nov. 1967, K. Toft [H267] (C-F-39492); Zealand, Hellerup Church Yard, in gravel under Pinus, 9 May 1998, T. Læssøe 4940 [H450] (C-F-38142). – Sweden, Gotland, Klintehamn parish, at the railway station, along sandy road in coniferous wood, almost covered by sand, 7 May 1938, S. Lundell & E. Åberg [H404] (C-Fungi Exs. Suec. 952, ‘Helvella leucomelas’ (Pers.) Nannf. epitype). – USA, Massachusetts, Worchester Co., at Bolton Lime Quarry, 7 May 2006, K. Hansen & G. Lewis-Gentry KH.06.01 [H115] (FH).
Notes — We have selected an epitype specimen to support the lectotype in Weber (1972) and Landeros et al. (2012) observed that H. leucomelaena has aporhynchous asci, an observation we concur with. Landeros et al. (2015) compared LSU sequences from the type specimen of A. calyx with recent collections of H. leucomelaena and concluded they were the same. A specimen from Australia [KC012682] in Landeros et al. (2015) also matches our concept of H. leucomelaena (cf. Appendix 3).
Helvella leucophaea (Battarra) Pers., Observ. Mycol. 2: 19. 1799
Basionym. Boletus leucophaea Battarra, Fungorum Arimin.: 25. 1759.
Synonyms. Helvella crispa c. (forma) lutescens, sicca subfulva Fr., Syst. Mycol. 2: 14. 1822.
Lectotype designated here: Battarra, Fungorum Arimin.: t. III, f. B, 1759. MycoBank MBT375735.
Helvella pityophila Boud., J. Bot. 1: 218. 1887 (‘pithyophila’).
Helvella crispa var. pityophila (Boud.) Donadini, Bull. Soc. Linn. Provence 28: 75. 1975.
Lectotype: France, Fontainebleau, Oct. 1885, Herb. Boudier (PC), not examined.
Illustrations — Trattinia (1809: pl. 28, as H. leucophaea Pers.), Boudier (1887: pl. 3; 1905–1910: pl. 227, as H. pithyophila).
Description of the species — Boudier (1905–1910: 119, as H. pithyophila).
Specimen examined/sequenced. France, Porquerelles, terrain sabloneux, 10 Dec. 1976, J.C. Donadini [H274] (C-F-45487).
Notes — This species resembles H. crispa in size and stature; however, the hymenium in fresh ascocarps is light brown, drying caramel brown and the receptacle and stipe are whitish. The paraphyses are thread-like and not significantly inflated at the clavate tips compared to the abruptly and subcapitately inflated paraphyses in H. crispa. Four rpb2 and two hsp substitutions separate H. leucophaea and the sister species H. crispa. The LSU nrDNA sequences are insufficient to discriminate among the two. The selection of an epitype specimen is postponed until an extended study of the H. crispa morphospecies complex could be undertaken.
Helvella levis Bergeret, Phytonom. Univ. 1: t. 149. 1783 — Fig. 7c
Lectotype designated here: Bergeret, Phytonom. Univ. 1: t. 149. 1783. MycoBank MBT375700. Epitype designated here: Denmark, M Zealand, Hejede Overdrev by Tjørnehus, 14 Sept. 2001, B.W. Pedersen (C-F-55188). MycoBank MBT375468.
Synonyms. Helvella latispora Boud., Bull. Soc. Mycol. France 14: 16. 1898.
Leptopodia latispora (Boud.) Boud., Icon. Mycol., liste prélim.: 2. 1904.
Lectotype. France, Helvella no. 480, ‘H. latispora’ Boud. (PC-Herb. Boudier), selected by Dissing (1966b).
Illustrations — Bergeret (1783: pl. 149), Boudier (1898: pl. 3, f. 2; 1905–1910: pl. 233, as Leptopodia latispora), Van Vooren (2010: 43, photo 11, as H. latispora).
Description of the species — Dissing (1966b: 135, as H. latispora).
Specimens examined/sequenced. Denmark, M Zealand, Hejede Overdrev by Tjørnehus, 14 Sept. 2001, B.W. Pedersen [H290] (C-F-55188 epitype); Lolland, Kristiansaede Forest, 6 km W Maribo, 5 Oct. 1968, P.M. Petersen [H264] (C-F-39370); Zealand, Copenhagen, Kongelunden, 25 Oct. 1974, H. Dissing [H266] (C-F-39367); Jutland, Pamhule Forest, on rich mouldy soil, 2 Sept. 1979, I. Weng [H265] (C-F-39366); East Jutland, W of Kongsted, Elbodalen, NW Fredericia, in mixed deciduous forest, 4 Oct. 2002, J. Vesterholt JV 660.02 [H393] (C-F-40830); NE Zealand, Gurre Sø, on soil under Urtica, 8 Oct. 1997, K. Hansen & T. Læssøe, KH 109.97 [H469] (C-F-52988); Konabbe Forest, 24 Sept. 2013, T. Kehlet [H458] (C). – Germany, Hamburg E, Sachsenwald, Friedrichsruh, under beech and oak, 12 Sept. 1973, I. Friedrichsen & Lorch [H480] (C-F-45517).
Notes —This species has commonly been treated under the younger, synonymous name H. latispora Boud. Helvella levis was placed in synonymy with H. elastica by Fries (1822). In our opinion, the description and figure of Bergeret’s (1783) is not representative for H. elastica, but for the deflexed, saddle-shaped fungus commonly referred to as H. latispora in literature. Dissing’s descriptions (1966a, b) of H. latispora fully cover Bergeret’s fungus, which is lectotypified here with Bergeret’s original plate (iconotype), supported with hsp, rpb2 and LSU sequences from a newly selected epitype specimen from Denmark.
Helvella macropus (Pers.) P. Karst., Not. Sallsk. Fauna Fl. Fenn. Forh. 11: 224. 1870 — Fig. 9c
Basionym. Peziza macropus Pers., Ann. Bot. (Usteri) 15: 26. 1795; Fr., Syst. Mycol. 2: 57. 1822.
Synonyms. Peziza macropus Pers., Ann. Bot. (Usteri) 15: 26. 1795, var. macropus.
Peziza macropus var. hirta Pers., Ann. Bot. (Usteri) 15: 26. 1795.
Helvella macropus (Pers.) P. Karst., Not. Sallsk. Fauna Fl. Fenn. Forh. 11: 224. 1870, var. macropus.
Macroscyphus macropus (Pers.) Gray, Nat. Arr. Brit. Pl. 1: 672. 1821.
Macropodia macropus (Pers.) Fuckel, Jahrb. Nassauischen Vereins Naturk. 23–24: 331. 1870 (‘1869–1870’).
Aleuria macropus (Pers.) Gillet, Champ. France Discomyc.: 35. 1879.
Lachnea macropus (Pers.) W. Phillips, Man. Brit. Disomyc.: 207. 1887.
Cowlesia macropus (Pers.) Nieuwl., Amer. Midl. Naturalist 3: 300. 1916.
Cyathipodia macropus (Pers.) Dennis, British Cup Fungi: 7. 1960.
Lectotype designated here: Pers., Observ. Mycol. 1: pl.1, f. 3. 1796. MycoBank MBT375701. Epitype designated here: Sweden, Uppland, Skogstibble parish, Rosenberg, 4 Sept. 1960, R. Santesson 13997 (C-Fungi Exs. Suec. 3266). MycoBank MBT375408.
Helvella hispida Schaeff., Fung. Bavar. Palat. Nasc. 4: 108. 1774.
Paxina hispida (Schaeff.) Seaver, North Amer. Cup-fungi (Operculates): 205. 1928.
Macropodia macropus var. hispida (Schaeff.) Killerm., Kryptog. Forsch. 2: 36. 1929.
Lectotype designated here: Schaeff., Fung. Bavar. Palat. Nasc. 2: pl. 167. 1763 (‘Elvela decima nona’). MycoBank MBT375702.
Peziza hypocrateriformis Bergeret, Phytonom. Univ. 1: t. 169. 1783.
Lectotype designated here: Bergeret, Phytonom. Univ. 1: t. 169. 1783. MycoBank MBT375736.
Illustrations — Dissing & Nannfeldt (1966: f. 4, 5a–e, pl. III), Weber (1972: f. 23–24).
Descriptions of the species — Dissing & Nannfeldt (1966: 333), Dissing (1966b: 62), Weber (1972: 164, as H. macropus var. macropus).
Specimens examined/sequenced. Canada, Alberta, Devonian Botanic Garden, Devon, 24 Aug. 1996, S.P. Abbott [H119] (FH); Quebec, 10 km NW Montreal at Biological Research Station, 2 Aug. 1997 [H120] (FH). – Norway, Østfold, Moss, Jeløya, Kjeldstadvik, humid deciduous forest, 18 Sept. 2013, T. Læssøe [H260] (O-253327); Oslo, Skullerud, on soil in deciduous forest, 3 Sept. 2009, M. Bjorbækmoe [H073] (O-253326); Akershus, Røyken, Nærsnes, 10 Sept. 2009, K. Bjørum [H060] (O-253323); Akershus, Oppegård, Bestemorstranda, rich spruce forest, 19 Sept. 2009, C. Christiansen [H069] (O-292075); Akershus, Ski, N Busterud gård, in rich spruce forest, 16 Sept. 2006, A. Pedersen [H157] (O-370348); Oppland, Dovre, Grimsdalen, Buåi, in birch forest at the river, 23 July 1980, T. Schumacher [H228] (O-253324); Oppland, Dovre, Grimsdalen, Storberget, in birch forest, 8 Aug. 2009, T. Carlsen, I. Skrede & T. Schumacher TS 36.09 [H257] (O-253325); Hedmark, Åmot, Kvannbekken National Reserve, mixed rich coniferous forest, 30 Aug. 2002, P. Marstad [H158] (O-65347); Telemark, Drangedal, Henneseid, Trillingtjernåsen W, in mixed pine-oak forest, 15 July 2004, T.E. Brandrud TEB 95.04 [H142] (O-166155); Kragerø, Storkollen, valley between Storkollen and Lindvikskollen, in rich mixed forest, 5 Aug. 2000, T.E. Brandrud TEB 70.00 [H141] (O-168819); Rogaland, Vindafjord, Kvaløy, rich deciduous forest, 11 Sept. 2009, E. Dalen, L. Dalen, J.B. Jordal [H238] (O-291425); Sokndal, Skaråsen, rich soil under Corylus, 9 Sept. 2009, J.I. Johnsen & J.B. Jordal [H239] (O-291391); Hordaland, Ulvik, Veånerbekken, bare soil in Fraxinus dominated forest, 28 Aug. 2001, G. Flatabø [H159] (O-220323); Eidfjord, Hæreid, pasture land with Betula, 6 Oct. 2006, Å. Oterhals [H160] (O-283072); Møre og Romsdal, Stordal, Stiggjeldet, deciduous forest, 5 Sept. 2003, D. Holtan & P. Larsen 78.2003 [H161] (O-223871); Sunndal, Sunndalsfjorden N Løvika, under Corylus, 11 Sept. 2001, J.B. Jordal [H165] (O-220225). – Sweden, Uppland, Skogstibble parish, Rosenberg, in forest, 4 Sept. 1960, R. Santesson 13997 [H412] (C-Fungi Exs. Suec. 3266 epitype). – USA, Alaska, Anchorage, between Anchorage and Denali National Park, among leaves of deciduous trees, 14 Aug. 1980, H. Dissing [H351] (C-F-56087); Massachusetts, Carlisle, Great Farm, 8 July 2006, L. Millman [H118] (FH); Massachusetts, Concord, Eastbrook Woods, 25 June 1998, D. Pfister [H124] (FH).
Notes — Persoon’s and Fries’ original concept of Peziza macropus included the European H. macrospora as well as H. fibrosa (= Octospora villosa). These two species are readily separable based on hymenium colour and microanatomical characters (see also H. fibrosa and artificial key). Helvella macropus is typified with Persoon’s original plate in the 1796 issue of his work, which includes a coloured plate, and supported with a widely distributed epitype (exsiccate) specimen capturing specific barcode sequences to the species discrimination. Our molecular survey confirms the existence of at least three species of the H. macropus morphospecies complex, i.e., H. macropus s.str. with a mainly European and North American distribution, Peziza subclavipes (= H. brevis) in USA and Mexico (cf. Landeros et al. 2015: f. 3) (not included in our dataset) and H. ephippioides S. Imai reported from Japan (Imai 1932) (see extralimital Helvella species below).
Helvella macrosperma (J. Favre) R. Fellner & Landa, Česka Mykol. 45 (1–2): 35. 1991 — Fig. 4d
Basionym. Helvella arctica Nannf. var. macrosperma J. Favre, Ergebn. Wiss. Untersuch. Schweiz. Nationalparkes 5, 33: 199. 1955.
Synonym. Helvella corium var. macrosperma (J. Favre) Bizio, Franchi & M. Marchetti, Riv. Micol. 41: 232. 1998.
Holotype. Switzerland, Val Nüglia 2550 m asl, leg. J. Favre, not examined.
Illustrations — Favre (1955: pl. 1, f. 2).
Apothecia short-stipitate, regularly cupulate, cap 0.5–1.8 cm diam, 0.5–1.5 cm high; stipe terete, solid, 0.4–1.5 cm high, 0.2–0.4 mm broad; hymenium and receptacle black (H1), receptaculum and stipe densely tomentose, hyphoid hairs mostly aggregated in conspicuous, pyramidal fascicles on the receptacle surface and stipe. Medullary excipulum of loosely interwoven textura intricata, hyphae 3–5 μm broad, short-celled; outer excipulum of brown-walled globose to angular cells, 10–20 μm diam, gradually arranged perpendicularly to the surface and forming a layer of closely packed cells that extend into dark brown-walled multicellular hyphoid hairs on the excipulum exterior, hairs generally 40–150 μm, at the margin to 300 μm in length, in some parts forming dense bundles, individual cells 10–15 μm broad, 10–40 μm long, constricted at septa with conspicuous deposits of incrusted pigments on the interior of the cell walls. Asci pleurorhynchous, 290–330 × 16–20 μm. Ascospores broadly ellipsoid, 18.5–22.5 × 11.5–13.8 μm. Paraphyses filiform, 2–2.5 μm broad, septate, with brownish pigments in their whole length, colour increasing in intensity towards the gnarled, clavate tips, 4.5–6.0 μm broad.
Specimens examined/sequenced. Norway, Dovre, Grimsdalen, Veslegrimsa, river bed, 2 Aug. 2007, T. Schumacher TS 54.07 [H029] (O-253328); Dovre, Tverrlibekken, 7 Aug. 2009, T. Carlsen, I. Skrede & T. Schumacher TS 21.09 [H047] (O-253329); ibid., 7 Aug. 2009, T. Carlsen, I. Skrede & T. Schumacher TS 28.09 [H050] (O-253330); ibid., 7. Aug. 2009, T. Carlsen, I. Skrede & T. Schumacher TS 31.09 [H053] (O-253331).
Notes — The species was originally described from the Swiss Alps (Favre 1955, as H. arctica Nannf. var. macrosperma). It is reported recently also from the French Alps (Van Vooren 2015). Helvella macrosperma is closely related to H. alpestris, from which it is distinguished by the prominent tufts of dark-pigmented hyphoid hairs on the receptacle surface and stipe. The microanatomy of ectal excipulum, ascospores and parahyses also discriminate among the two. One hsp, one rpb2, four tef, and seven LSU substitutions separate H. macrosperma and H. alpestris.
Helvella monachella (Scop.) Fr., Syst. Mycol. 2: 18. 1822
Basionym. Phallus monacella Scop., Fl. Carniol., ed. 2, 2: 476. 1772; Fr., Syst. Mycol. 2: 18. 1822 (‘monachella’ orth. mut.).
– non Helvella monacella Schaeff., Fung. Bavar. Palat. Nasc. 4: 106. 1774 (‘Elvela’), nom. inval. (= Helvella sulcata Afzel.).
Lectotype designated here: Battarra, Fungorum Arimin.: t. 2, f. H, sub ‘Boletus albus pileolo complicatus nigro’, 1759. MycoBank MBT375704. Epitype designated here: Hungary, Szelid, Bacs-Kiskun, in populeto-saliceto, 14 May 1979, M. Babos, L. Albert, J. Bükei & A. Friesz (C-F-92121 ‘Helvella leucopus Pers.’ ex Herb. Musei Hist. Nat. Hung. Budapest, dupl.). MycoBank MBT375409.
Synonyms. Helvella leucopus Pers., Mycol. Eur. 1: 213. 1822.
Lectotype. France. Herb. Persoon no. 910.261-997 (L) selected by Dissing (1966a), not examined.
Helvella albipes Fuckel, Jahrb. Nassauischen Vereins Naturk. 23–24: 334. 1870. ‘1869–1870’.
Lectotype. Austria, Fung. rhen.1240 ‘H. albipes’ (BM), selected by Dissing (1966b), not examined.
Illustrations — Fuckel (1870: pl. 5, f. 2), Dissing (1966b: f. 38 (as H. leucopus), Moravec (1980: pl. 4, as H. leucopus), Van Vooren (2010: 43, photo 12).
Descriptions of the species — Fuckel (1870: 334, as H. albipes), Dissing (1966b: 138 as H. leucopus).
Specimens examined/sequenced. Hungary, Szelid, Bacs-Kiskun, in populeto-saliceto, 14 May 1979, M. Babos, L. Albert, J. Bükei & A. Friesz [H268] (C-F-92121 ‘Helvella leucopus Pers.’ epitype); Örkeny, ad terram in populeto, 24 May 1965, I. Ferencz, E. Toth, E. Vessey & S. Vasadi [H269] (C-F-92120 ex Herb. Musei Hist. Nat. Hung. Budapest).
Notes — The rpb2 and hsp sequences are identical among the sequenced specimens. Helvella monachella Scop. was sanctioned by Fries (1822), who listed H. leucopus as a presumable taxonomic synonym (Fries 1832). Fries (1822) misinterpreted Scopoli’s species when he also included H. spadicea as a forma b of H. monachella (Fries 1822), this latter fungus presumably representing a Gyromitra species, as also concluded by Dissing (1966b). We have selected Battarra’s original illustration of ‘Boletus albus pileolo complicatus nigro’ (Battarra 1759) as lectotype (‘iconotype’) for the original fungus of Scopoli’s (1772), since it was referred to in Fries’ original description of the species. Helvella albipes Fuckel (1870) is a more recent heterotypic synonym. The lectotype of H. monachella is supported with an epitype from Hungary, supplying rpb2, hsp and LSU sequences for the purpose of a precise application of the name (ICN article 9.8).
Helvella nannfeldtii Skrede, T. Carlsen & T. Schumach. sp. nov. — MycoBank MB820326; hsp barcode GenBank KY784203; rpb2 barcode GenBank KY772447; Fig. 4b
Etymology. Named in honour of the Swedish botanist and mycologist Jon Axel Nannfeldt.
Holotype. Norway, Oppland, Dovre, Grimsdalen, Veslegrimsa, 8 Aug. 2009, T. Carlsen, T. Schumacher & I. Skrede TS 22.09 (O-253338).
Apothecia short-stipitate-cupulate, black, 1–3 cm broad, 1–3 cm high; stipe terete, usually deeply buried in the soil, 0.2–0.3 cm broad, 0.5–2.0 cm long, greyish black above, whitish below, with or without grooves at base, receptacle surface pubescent, towards margin with long slender tufts of short-segmented, subhyaline to light brown-walled hyphoid hairs to 200 μm long staining in cotton blue, cells irregular in shape and generally much constricted at the septa. Medullary excipulum of loosely interwoven textura intricata, hyphae 3–5 μm broad. Outermost excipulum of textura angularis, cells dark brown, thick-walled, 15–30 μm diam, intermixed with broad short-segmented brownish hyphae forming a textura intricata. Asci pleurorhyncous, 230–270 × 15–19 μm. Ascospores ellipsoid, 18–21 × 11–13 μm. Paraphyses straight, dark brown along upper two thirds, 2–3 μm broad, septate, gradually enlarged to 5–6 μm at the subcapitate tips.
Specimens examined/sequenced. France, Savoie, Val d’Isere, Gorges du Mal, 2400 m asl, 31 Aug. 1992, T. Schumacher F 83.92 [H212] (O-253332); Savoie, Bon Valle, Sur Arc, 2 Sept. 1992, T. Schumacher F 94.92 [H216] (O-253333). – Norway, Oppland, Dovre, Grimsdalen, Veslegrimsa, 8 Aug. 2009, T. Carlsen, T. Schumacher & I. Skrede TS 22.09 [H027] (O-253338 holotype); ibid., 2 Aug. 2007, T. Schumacher TS 66.07 [H028] (O-253339); Grimsdalen, Tverrgjelet, on river bed, deeply immersed in sand, 6 Aug. 1996, A.H. Jensen & T. Schumacher [H254] (O-253341); ibid., 2 Aug. 2007, T. Schumacher TS 60.07 [H017] (O-253337); Grimsdalen, Tverråi, 13 Aug. 1985, T. Schumacher [H367] (O-253343); Grimsdalen, Grimsa, on river terrace at the river, 25 Aug. 1996, T. Schumacher [H362] (O-253342); ibid., 17 Aug. 2005, T. Schumacher [H188] (O-253340); Grimsdalen, Buåi, at the river, 17 Sept. 1983, T. Schumacher & K. Østmoe D 168.83 [H479] (O-253344); Hordaland, Ulvik, Finse, Blåisen, 27 July 2014, T. Schumacher [H452] (O-253334), [H454] (O-253335), 31 July 2014, K. Høiland [H457] (O-253336); Nordland, Rana, Virvassdalen, Beveråa, river bed, 30 July 1974, S. Sivertsen [H318] (C-92123 ex TRH).
Notes — Helvella nannfeldtii shows superficial resemblance to H. corium and H. macrosperma; however, the short cylindrical stipe, which is whitish below, helps discriminate the two latter species in the field. Helvella nannfeldtii forms a well-supported lineage with H. alpestris and H. macrosperma in our phylogeny. Six hsp, two rpb2, four tef and 43 LSU substitutions, and two hsp, one rpb2, three tef and 41 LSU substitutions separate H. alpestris and H. macrosperma, respectively.
Helvella nigra Bergeret, Phytonom. Univ. 1: t. 147. 1783 — Fig. 6e
– non Helvella nigra Peck, Bull. Torrey Bot. Club 26: 70. 1899, nom. illeg. (homonym).
Lectotype designated here: Bergeret, Phytonom. Univ. 1: t. 147. 1783. MycoBank MBT375705. Epitype designated here: Sweden, Halland, Halmstad, Almeberget, 16 Sept. 2009, G. Holden (O-253345). MycoBank MBT375466.
Misapplied name
– Helvella lacunosa sensu Fr., Syst. Mycol. 2: 15. 1822.
Illustration — Bergeret (1783: pl. 147).
Apothecia stipitate-capitate, 2–6 cm across, cap saddle-shaped, bi- to trilobate, from 6–15 cm in height, apothecial margin adnate, firmly attached to the stipe at 4–6 points; hymenium wrinkled, dark grey (F1) to greyish black (G1) when fresh, drying darker; stipe concolorous or slightly paler, lacunose, receptacle surface concolorous or slightly paler, smooth, with distinct, branching ribs; stipe 2.0–4 cm broad, naked, ribbed, ash grey to almost black, longitudinal ribs high and prominent, sharp, partly anastomosing and double-edged, with deep ‘holes’ (lacunae) inbetween ribs, stipe 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 few-celled outgrowths taking much colour in CB. Asci pleurorhynchous, 290–330 × 14–16 μm. Ascospores ellipsoid, 16.5–19 × 10.5–12 μm. Paraphyses straight, slightly bent at tips, light brown in the whole length, 3–4 μm broad below, enlarged to 6–8 μm at the tips.
Specimens examined/sequenced. Spain, La Rioja, Castroviejo, 9 Aug. 2014, A. Melendez, R. Martinez, C. Perez PIPE 0829 [H1029] (O-253404); Basque Country, Urkiola, 19 Oct. 2007, G. Munoz GM 1085 [H1030] (O-253405). – Sweden, Halland, Halmstad, Almeberget, beech forest, 16 Sept. 2009, G. Holden [H063] (O-253345 epitype); Halmstad, Holkåsen, 17 Sept. 2009, G. Stenström [H068] (O-253347); Halmstad, Biskopstorp, 17 Sept. 2009, K.H. Larsson [H067] (O-253346).
Notes — We have resurrected the binomial H. nigra for this fungus, referring to a species placed in synonymy with H. lacunosa by Fries (1822). Helvella nigra can be distinguished by the large sized, reflexed and lobed apothecium with prominent ribs on the outer surface firmly adhered onto the stipe, and microscopically by apically bent paraphyses (like a ‘golf-club’). The Swedish specimens studied by us occur on soil and deeply decayed wood. The selected lectotype of Bergeret’s is supported here with an epitype from lowland Sweden supplying hsp, rpb2, tef and LSU sequences. Eight hsp, three rpb2 and 25 LSU nrDNA characters distinguish H. nigra from H. lacunosa. An LSU sequence in GenBank (AJ 544211) from Germany labelled H. lacunosa, matches the specimens of H. nigra (cf. Appendix 3).
Helvella oblongispora Harmaja, Karstenia 18: 57. 1978
Holotype. Germany, Bavaria, Munich, Isar river, 23 July 1969, A. Einhellinger (C).
Illustration — Harmaja (1979: f. 7).
Description of the species — Harmaja (1979: 36).
Specimens examined/sequenced. Germany, Bavaria, Munich, Isar, 23 July 1969, A. Einhellinger (C holotype). – Norway, Oppland, Lunner, Muttagruvene naturminne, on calcareous gravel, 27 Aug. 2004, T.E. Brandrud & E. Bendiksen TEB 278.04 [H132] (O-166316). – Switzerland, Graubünden, Mota Jüda, Val Plavna, on path with small stones, 30 Aug. 1979, H. Dissing [H332] (C-F-56844); Graubünden, Oftenpass, near bridge under Raitschana, at P-place N National Park, 5 Sept. 1979, H. Dissing [H448] (C-F-56914).
Notes — Helvella oblongispora is morphologically similar to H. costifera and H. calycina; however, the aporhynchous ascus base and species specific DNA sequences confirm its affiliation to the leucomelaena lineage together with H. leucomelaena, H. confusa and H. crassitunicata. We have examined the holotype specimen under the lens, but were unsuccessful in extracting amplifiable DNA from this specimen.
Helvella pallescens Schaeff., Fung. Bavar. Palat. Nasc. 4: 114. 1774 (‘Elvela’) — Fig. 8b
Synonym. Helvella sulcata β (var.) pallescens (Schaeff.) Fr., Syst. Mycol. 2: 16. 1822.
Lectotype designated here: Schaeff., Fung. Bavar. Palat. Nasc. 4: t. 322. 1774 (‘Elvela tricésima quinta’). MycoBank MBT375713. Epitype designated here: Norway, Møre og Romsdal, Norddal, Stiggjelet, 29 Aug. 2003, D. Holtan (O-66205). MycoBank MBT375410.
Illustrations — Schaeffer (1774: pl. 322), ? Boudier (1905–1910: pl. 226, as H. lactea).
Apothecia bilobate to campanulate to irregularly saddle-shaped, cap 0.6–2 cm broad, by 2–3 cm high, underside glabrous, edge of cap adnate (reflexed) and fused with the stipe at 3–4 points; hymenium and stipe white to pale greyish, hymenium white to pale grey (3-5A-B1-2), drying brownish yellow (5B-C7-8); receptacle surface and stipe pale yellowish to brown, drying brown; stipe long and slender, 0.5–1.5 cm broad, 4–6 cm high, hollow, with 4–5 longitudinal ribs and a few lacunae inbetween. Medullary excipulum of an interwoven textura intricata, hyphae of variable thickness, 4–10 μm broad. Outer excipulum of textura globulosa-angularis, cells 20–40 μm diam, outermost cells more elongated and prismatic giving rise to a cover of densely packed club-shaped cells on receptacular surface. Asci pleurorhynchous, 280–310 × 14–15 μm. Young asci with numerous cyanophilic ascospores with large pustules adhered to the spore wall. Ascospores broadly ellipsoid, 15.0–18.2 × 10.2–13.2 μm. Paraphyses hyaline, 2–3 μm broad, septate, gradually enlarged into irregular clavate tips, 6–10 μm broad.
Specimens examined/sequenced. Norway, Møre og Romsdal, Norddal, Stiggjeldet, rich moldy soil under Corylus avellana, 29 Aug. 2003, D. Holtan [H138] (O-66205 epitype); ibid., [H070] (O-289039); Østfold, Hvaler, Asmaløy, Fonten, on soil in rich deciduous forest, 29 Aug. 1992, Roy Kristiansen RK 49784 [H373] (O-253355); Rogaland, Hjelmeland, Hetlandsbygda W Kvitefjell, on soil under Corylus, 27 Sept. 2009, O. Førland & J.B. Jordal [H236] (O-291458); Hordaland, Ulvik, Veånesbekken, on soil in deciduous forest under Betula, 28 Aug. 2001, G. Flatabø [H136] (O-220306).
Notes — This small and slender, whitish to greyish species has apparently been confused with H. lactea, which shares a similar colour and micromorphological characters; however, H. pallescens has a more slender and narrow stipe and sharper ribs that extend onto the proximal parts of the receptacle keeping the margin firmly attached to the stipe. Microanatomical features that distinguish it from H. lactea are longer asci (280–310 μm in H. pallescens vs 240–260 μm in H. lactea) and differently shaped paraphyses. The species is nested with H. atra and H. sulcata as the closest species (sisters) in the lacunosa clade (clade C). Eight rpb2 and nine hsp substitutions separate H. pallescens and H. lactea. We have selected Schaeffer’s plate 322 as lectotype (Schaeffer 1774) and supported it with sequences of rpb2, tef, and LSU from an epitype specimen from Norway.
Helvella palustris Peck, Ann. Rep. N. Y. State Mus. Nat. Hist. 33: 31. 1880 — Fig. 6c
Holotype. USA, New York, Manlius Center, Aug., C.H. Peck (NYS), not examined.
Illustration — Weber (1972: f. 40).
Description of the species — Weber (1972: 189).
Specimens examined/sequenced. Finland, Kuusamo, Juuma, The Vuomas, in gravel pit, 25 Aug. 1978, H. Dissing [H350] (C-55330). – Japan, Honshu, Gumma Prefecture, Tone-gum, Katashima-mura, Jujo Seishi Co. Forest, on soil, 26 Aug. 1983, T. Schumacher [H195] (O-253356). – Norway, Oppland, Dovre, Grimsdalen, Buådalen, in spring horizon in the subalpine birch forest, 11 Sept. 1982, T. Schumacher & K. Østmoe 479.82 [H468] (O-253362); Grimsdalen, at Grimsdalshytta, 16 Aug. 2001, T. Schumacher [H001] (O-253357); Grimsdalen, Tverrådalen, moist slope along river, 25 Aug. 1996, T. Schumacher [H184] (O-253361); ibid., 7 Aug. 2009, T. Carlsen, I. Skrede & T. Schumacher TS 12.09 [H019] (O-253358); ibid., TS 13.09 [H043] (O-253359); Grimsdalen, at margin of minerotrophic fen, 23 Aug. 1996, S. Landvik & T. Schumacher TS 204.96 [H181] (O-253360).
Notes — This characteristic species of the lacunosa clade is a medium-sized, bi- to trilobate black Helvella species in which the cap is reflexed towards a slender and longitudinally ribbed stipe. The typical habitat is along rivulets and in deep moss carpets of rich, minerotrophic fens in subalpine areas. Weber (1972) compared the type specimens of H. palustris and H. philonotis and concluded they were conspecific. Based on morphology and DNA barcode sequences we have found good reasons to distinguish these two species (see H. philonotis). Landeros et al. (2015) collected a specimen from the USA under the name of H. aff. sulcata [JX993082] that seems to belong here (cf. Appendix 3).
Helvella panormitana Inzenga, Funghi Siciliani 1: 41. 1865 — Fig. 7a
Synonym. Leptopodia panormitana (Inzenga) Boud., Hist. Classif. Discomyc. Europe: 37. 1907.
Lectotype designated here: Inzenga, Funghi Siciliani 1: pl. 4, f.11-4. 1865. MycoBank MBT375706. Epitype designated here: Sweden, Halland, Laholm, Hallandsåsen N, on soil in beech forest, 16 Sept. 2009, C.C. Mellberg (O-253363). MycoBank MBT375469.
Illustrations — Inzenga (1865: pl. 4, f.11-4).
Apothecia stipitate-capitate, cap campanulate to bilobate, 1–2.5 cm broad, 1.5–3 cm high, long-stipitate, margin free, reflexed towards stipe, hymenium pale greyish brown (5-8D-E3, receptacle smooth, white or pale yellow (3A4), concolorous with stipe, drying brownish orange (5C4-6); stipe 0.4–1 cm broad, by 3–15 cm high, solid to hollow, occasionally with grooves towards the base. Subhymenium and medullary excipulum of highly gelatinous hyphae, 3–4 μm broad, embedded in an amorphous gelatinous matrix. Outer excipulum a mixture of textura intricata and textura angularis, hyphae to 8 μm broad, angular cells generally 25–40 μm diam, partly inflated to 60–70 μm diam, outermost cells giving rise to densely packed rows of prismatic 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, 19–24 × 11.5–13.5 μm. Paraphyses c. 3 μm broad below, septate, light brown in colour, gradually enlarging towards the 6–8 μm broad clavate tips.
Specimens examined/sequenced. Norway, Hordaland, Bømlø, Grønåsvågen, under Corylus, 1 Sept. 2002, A. Knutsen [H143] (O-65394); Bømlø, Hildeneset, pasture land in deciduous forest, 17 Sept. 2007, A. Knutsen [H137] (O-284515); Granvin, Kvanndal E, boreonemoral rich broad-leaved deciduous forest, 25 Sept. 2005, E. Bendiksen EB 290.05 [H154] (O-167560); Møre og Romsdal, Ålesund, Magerholm, under Corylus, 8 Oct. 2006, P.G. Larsen [H237] (O-360894); Møre og Romsdal, Stordal, Seljeneset, rich deciduous forest with Corylus, Betula and Populus, 18 Sept. 2005, P.G. Larsen [H145] (O-203499); Møre og Romsdal, Sykkylven, Klungregylet, rich soil in hazel wood, 18 Sept. 2001, D. Holtan [H130] (O-171669); Møre og Romsdal, Midsund, Otrøya, Trelvika, in SW exposed Corylus forest, 10 Sept. 2003, J.B. Jordal [H144] (O-178718). – Sweden, Halland, Laholm, Hallandsåsen N, on soil in beech forest, 16 Sept. 2009, C.C. Mellberg [H064] (O-253363 epitype).
Notes — We have adopted the name H. panormitana for this species of the elastica clade (clade E). The species was originally described from Sicilia, Italy, by Inzenga (1865), but was later referred to as a synonym of H. elastica (Inzenga 1869). Two rpb2, one hsp and seven LSU substitutions separate the Swedish and Norwegian specimens. Helvella panormitana deviates from the morphologically similar H. elastica in five hsp, 11 rpb2 and 10 LSU substitutions.
Helvella paraphysitorquata I. Arroyo & Calonge, in Calonge & Arroyo, Mycotaxon 39: 210. 1990
Holotype. Spain, Teruel, Albarracin, 3 June 1988, I. Arroyo & F.D. Calonge (MA-Fungi 24512). Isotype (C-F-45305).
Illustrations — Calonge & Arroyo (1990: f. 2), Landeros et al. (2012: f. 30).
Descriptions of the species — Calonge & Arroyo (1990: 210), Landeros et al. (2012: 54).
Specimen examined/sequenced. Spain, Teruel, Albarracin, 3 June 1988, I. Arroyo & F.D. Calonge [H271] (C-F-45305 isotype).
Notes — A molecularly and morphologically unique species, separated from its sister species H. pubescens by 11 rpb2 and 11 hsp and from H. rivularis by eight rpb2 and 10 hsp substitutions, respectively. Landeros et al. (2015) published an LSU nrDNA sequence of a specimen referred to as H. ephippium [JN 048874] that shows sequence similarity to H. paraphysitorquata (cf. Appendix 3).
Helvella pezizoides Afzel., Kongl. Vetensk. Acad. Nya Handl. 4: 308. 1783
Basionym. Helvella pezizoides Afzel., Kongl. Vetensk. Acad. Nya Handl. 4: 308. 1783; Fr., Syst. Mycol. 2: 20. 1822.
Synonym. Leptopodia pezizoides (Afzel.) Boud., Icon. Mycol., liste prélim.: 2. 1904.
Lectotype. Afzel., Kongl. Vetensk. Acad. Nya Handl. 4: t. 10, f. 2, 1783, designated by Dissing (1966b). Epitype designated here: Sweden, Halland, Halmstad, Almeberget, moist beech forest, 16 Sept. 2009, S. Lund (O-253366). MycoBank MBT375411.
Illustrations — Afzelius (1783: pl.10, f. 2), Dissing (1966b: f. 32b).
Description of the species — Dissing (1966b: 117).
Specimens examined/sequenced. Denmark, Jutland, Oksbøl Plantage, 13 km W Varde, 27 Oct. 1968, P.M. Petersen [H418] (C-F-45301); North Jutland, Fosdalen, Vor Frues kilde, soil on slope, 6 Oct. 1997, M. Christensen & K. Hansen KH-97-96 [H431] (C-F-52986). – Japan, Honshu, Tochigi Prefecture, Okunikko, Kotoku Bokujo, 26 Aug. 1983, T. Schumacher J119.83 [H204] (O-253364). – Nepal, Langtang Area, N Kyangjin, moraines of Langtang Lirung Glacier, 4000 m asl, 9 Sept. 1986, J. Poelt [H230] (C ex Herb. GZU 109.86). – Norway, Vestfold, Horten, Karljohansvern, 21 July 1981, P. Marstad & S. Aase [H196] (O-253365); Vestfold, Tønsberg, along road near lawn with Quercus, 21 July 1981, P. Marstad & R. Kristiansen [H420] (C-F-45505). – Sweden, Halland, Halmstad, Almeberget, moist beech forest, 16 Sept. 2009, S. Lund [H061] (O-253366 epitype); ibid., [H065] (O-253367).
Notes — The excellent description by Afzelius (1783), including a lot of details about apothecial ontogeny, leaves no doubt about the identity of this small to medium-sized, brownish black species of the fallax-pezizoides lineage. Helvella pezizoides resembles H. pulla in microanatomical features; however, the hymenium is dark brownish black to black in H. pezizoides and grey to greyish brown in H. pulla. We have selected a specimen from the collecting ground of Afzelius’ in Sweden as an epitype, highlighting the original concept of H. pezizoides.
Helvella philonotis Dissing, Bot. Tidsskr. 60: 117. 1964 — Fig. 6d
Holotype. Iceland, Hveragerdi, Mt Tindar, in moss in a seeping spring, 31 July 1959, M. Lange 130 (C-F-45481).
Synonyms. Helvella dovrensis T. Schumach., Mycotaxon 43: 34. 1992.
Holotype. Norway, Oppland, Dovre, Grimsdalen, Tverrliseter, 5 Aug. 1989, T. Schumacher 54.89 (O-253305)
? H. queletii Bres. var. alpina R. Heim & L. Rémy, Bull. Soc. Mycol. France 48: 53. 1932.
Misapplied names
– Helvella alpestris sensu Häffner, Beih. Z. Mykol. 7: 60. 1987.
– Helvella corium forma alpestris sensu J. Favre, Ergebn. Wiss. Untersuch. Schweiz. Nationalparkes 5, 33: 27. 1955.
Illustrations — Dissing (1964: f. 8), Schumacher & Mohn Jenssen (1992: 19, as H. dovrensis).
Descriptions of the species — Dissing (1964: 117; 1966b: 112), Schumacher (1992: 34, as H. dovrensis), Schumacher & Mohn Jenssen (1992: 19, as H. dovrensis).
Specimens examined/sequenced. Austria, Tirol, Obergurgl, Rotmoostal, 2300–2460 m asl, 19 Aug. 1990, T. Schumacher [H475] (O-253290); ibid., 23 Aug. 1990, T. Schumacher [H381] (O-253289). – France, Savoie, Les Vallions, Vanoise National Park, 2600 m asl, on calcareous soil, 28 Aug. 1992, T. Schumacher F 28.92 [H220] (O-253293); Savoie, Val d’Isere, Gorges du Mal, on river terraces, 31 Aug. 1992, T. Schumacher F 75.92 [H214] (O253291); ibid., F 82.92 [H218] (O-253292). – Iceland, Hveragerdi, Mt Tindar, in moss in a seeping spring, 31 July 1959, M. Lange 130 [H492] (C-F-45481 holotype); Grøndalur, near Kloarfjell, on mosses in a seeping spring, 30 July 1959, M. Lange 90 [H272] (C-F-45486 paratype). – Norway, Oppland, Dovre, Grimsdalen, Tverrliseter, on sandy soil, 5 Aug. 1989, T. Schumacher G 54.89 [H193] (O-253305 holotype of H. dovrensis); Grimsdalen, Veslegrimsa, on river terrace, 2 Aug. 2007, T. Schumacher TS 57.07 [H038] (O-253299); ibid., 8 Aug. 2009, T. Carlsen, I. Skrede, T. Schumacher TS 32.09 [H023] (O-253297); Grimsdalen, Tverrådalen, on slopes towards the river Tverråi, 7 Aug. 1989, T. Schumacher G 71.89 [H200] (O-253304); ibid., 31 July 2007, T. Schumacher TS 7.07 [H037] (O-253298); Grimsdalen at Storberget, 8 Aug. 2009, T. Carlsen, I. Skrede, T. Schumacher TS 26.09 [H048] (O-253300); Grimsdalen, Buåi, in the valley at the Buåi estuary, on calcareous soil, 5 Aug. 1989, T. Schumacher G 49.89 [H198] (O); Grimsdalen, Tverrgjelet, 2 Aug. 2007, T. Schumacher TS 64.07 [H015] (O-253296); 8 Aug. 2009, T. Carlsen, I. Skrede, T. Schumacher TS 29.09 [H051] (O-253301); Grimsdalen, Grimsdalshytta, 5 Aug. 1989, T. Schumacher G 15.89 [H199] (O-253303); ibid.,16 Aug. 2001, T. Schumacher [H005] (O-253294); ibid., 31 July 2007, T. Schumacher TS 06.07 [H006] (O-253295). – Switzerland, Graubünden, Albula Pass, N of the road near Crap Alv, 30 Aug. 1984, T. Schumacher Sch 42.84 [H361] (C-F-57376).
Notes — Helvella philonotis may have been described from the French Alps as a variety (var. alpina) of H. queletii (= H. solitaria) by Heim & Rémy (1932). There are no authentic specimens left to support this assumption (Dissing 1964, Heim in litt.). The synonymy of H. dovrensis with H. philonotis is established here based on hsp, rpb2 and LSU sequence similarity of holotype specimens of the two taxa. Helvella philonotis differs from H. palustris by three rpb2, one hsp and no LSU substitutions. Although genetically uniform, the two species are easily separated based on macromorphology and ecology, i.e., in H. philonotis the apothecia are short-stipitate and cupulate to laterally compressed to irregular saddle-shaped with a recurved margin and with a greyish black stipe with irregular grooves and blunt ribs scarcely continuing onto the receptacle surface. The Icelandic material represents a long-stalked form of the species; in H. palustris the apothecia are bi- to trilobately saddle-shaped with an adnate reflexed margin and a stipe with distinct, longitudinal, parallel ribs. Helvella philonotis seems restricted to the arctic-alpine biome of Europe while H. palustris seems confined to rich minerotrophic fens (mires) in boreal and boreo-nemoral forest biomes. The Norwegian specimens of H. alpestris in Landeros et al. (2012, 2015) represent H. philonotis. Recent records of H. dovrensis from Quercus root tips at high altitudes in Southwest China may represent another species (Hwang et al. 2015).
Helvella phlebophora Pat. & Doass. in Pat., Tab. Analyt. Fung. Ser. 1, 5: 208. 1886
Synonyms. Globopilea phlebophora (Pat. & Doass.) Beauseign., Contr. Etude Fl. Mycol. Landes: 205. 1926.
Lectotype designated here: Patóuillard, Tab. Analyt. Fung. Ser.1, 5: f. 468. 1886. MycoBank MBT375707. Epitype designated here: Iceland, Austur-Skaftafellsysla, Skaftafell, 20 Aug. 1981, H.F. Gøtzsche HFG 84-31 (C-F-45405 epitype). MycoBank MBT375412.
Illustrations — Dissing (1966a: f. 8b, f; 1966b: f. 30), Weber (1972: f. 34).
Descriptions of the species — Dissing (1966b: 110), Weber (1972: 182).
Specimens examined/sequenced. Iceland, Austur-Skaftafellsysla, Skaftafell, 20 Aug. 1981, H.F. Gøtzsche HFG 84-31 [H273] (C-F-45405). – Japan, Honshu, Tochigi, Asegata, Chuzenjikohan, Okunikko, 22 Aug. 1983, T. Schumacher J 28.83 [H197] (O-253368).
Notes — No authentic material of this characteristic species was located by Dissing (1966b). One substitution in rpb2 distinguishes the two collections from Japan and Iceland.
Helvella pubescens Skrede, T. Carlsen & T. Schumach., sp. nov. — MycoBank MB820328; hsp barcode GenBank KY784257; rpb2 barcode GenBank KY772506
Etymology. From Latin ‘downy with short soft hairs’; pertaining to hairy stipe and receptacle.
Holotype. Canada, Alberta, Wagner Natural Area near Spruce Grove, on the ground under Picea, 6 July 1994, S.P. Abbott (FH).
Apothecia stipitate-cupulate to disc-shaped, cap 2–3.5 cm broad, 0.5–1 cm high, slightly compressed; hymenium greyish yellow (3-4B-C4-8) when fresh, drying dark yellowish brown (5F5); receptacle pale grey (B1-2), delicately pubescent; stipe slender, terete, towards base with inconspicuous grooves, pubescent, 1.5–3 cm long, 0.3–0.6 cm broad, light grey to whitish, drying greyish, densely covered by pyramidal ‘warts’, i.e., bundles of hyphoid hairs. Medullary excipulum of textura intricata, individual hyphae hyaline, 3–4 μm broad. Outer excipulum of textura angularis; outermost layer with cells densely packed in rows consisting of drumshaped to clubshaped cells, 15–40 μm diam, outermost cells giving rise to short-segmented hyaline, hyphoid hairs to 200 μm long, partly arranged in fascicled tufts, hairs with drum-shaped cells constricted at septae and heavily staining in CB. Asci pleurorhynchous, 270–290 × 13–15 μm. Ascospores ellipsoid, 17.0–20.8 × 10–11.8 μm. Paraphyses subhyaline, septate, 3–4 μm broad below, enlarged to 6–8 μm at the tips.
Specimens examined/sequenced. Canada, Alberta, Wagner Natural Area near Spruce Grove, on the ground under Picea, 6 July 1994, S.P. Abbott [H121] (FH holotype). – Norway, Østfold, Skjeberg, Hafslundsparken, among mosses on rich soil under Corylus, 16 Aug. 1986, R. Kristiansen RK 45.86 [H251] (O-253369). – USA, Oregon, Benton Co., National Forest Lane CO., McKenzie District, on soil in moist ditch along road, 5 July 1991, H. Dissing Oregon 36.91 [H333] (C-56499); Oregon, Benton Co., National Forest Lane CO., MacDonald Forest, on moss grown trunk, along stream, 6 July 1991, H. Dissing Oregon 39.91 [H334] (C-56502).
Notes — Specimens of H. pubescens form a lineage of its own in our phylogeny. The American specimen is separated from the European (Norwegian) and Canadian specimens by two rpb2 and two hsp substitutions. A specimen from Mexico (JX 993064) referred to as H. aff. ephippium in Landeros et al. (2015) belongs here (cf. Appendix 3). The material from Alberta, Canada, and from Østfold, Norway, which was referred to as H. rivularis by Abbott & Currah (1997) also belongs here. Helvella pubescens may be confused with H. scyphoides, but the hymenium and stipe have more contrasting colours compared to the latter. Moreover, they belong in different lineages (cf. Fig. 2).
Helvella pulla Holmsk., Beata Ruris Otia Fung. Dan. 2: 49. 1799 — Fig. 9d
Basionym. Helvella pulla Holmsk., Beata Ruris Otia Fung. Dan. 2: 49. 1799; Fr., Syst. Mycol. 2: 20. 1822.
Synonyms. Leptopodia pulla (Holmsk.) Boud., Icon. Mycol., liste prélim.: 2. 1904.
Lectotype designated here: Holmsk., Beata Ruris Otia Fung. Dan. 2: t. 26. 1799. MycoBank MBT375708. Epitype designated here: Norway, Møre og Romsdal, Nesset, Eikesdalen, beneath Rangåfjellet, 26 Sept. 2008, T. Læssøe (O-69282). MycoBank MBT375413.
Helvella fuliginosa Schaeff., Fung. Bavar. Palat. Nasc. 4: 113. 1774. (‘Elvela’)
Helvella elastica Bull. b. (forma); Fr., Syst. Mycol. 2: 21. 1822.
Lectotype designated here: Schaeff., Fung. Bavar. Palat. Nasc. 4, t. 320. 1774. (‘Elvela tricesima tertia’). MycoBank MBT375709.
Illustrations — Schaeffer (1774: pl. 320), Holmskjold (1799: pl. 26).
Apothecia stipitate-capitate, cap 0.5–1.5 cm broad, 0.5–1.5 cm high, saddle-shaped, bilobate and involute (reflexed) throughout development; hymenium pale grey to medium grey (C-E1), receptacle concolorous, subpubescent; stipe grey, terete or slightly compressed, subpubescent, greyish, 3–6 cm long, 0.3–0.5 cm broad, firmly attached to deeply decayed wood. Medullary excipulum of loosely interwoven textura intricata, hyphae c. 2–4 μm broad, subhyaline, intermixed with some broader, brown-walled hyphae. Outer excipulum of a dense textura intricata to prismatica, hyphae 5–15 μm broad, partly brown-walled, short-segmented, turning out perpendicular to receptacle surface singly or in loose bundles, outermost cells inflated and club-shaped, 15–30 × 10–20 μm, heavily staining in CB. Asci pleurorhynchous, 275–310 × 13–15 μm. Ascospores ellipsoidal, 17.6–19.0 × 10.6–12.0 μm. Paraphyses subhyaline, relatively broad below, 3–4 μm, gradually enlarged along upper half to 8–12 μm at the tips.
Specimens examined/sequenced. Norway, Møre og Romsdal, Nesset, Eikesdalen, beneath Rangåfjellet, 26 Sept. 2008, T. Læssøe [H149] (O-69282 epitype); Østfold, Moss, Jeløya, Kjelstadvik, deciduous forest, under tree trunk, 18 Sept. 2013, K. Varenius [H259] (O-253370).
Notes — We have found good reasons to resurrect Holmskjold’s name for this characteristic greyish species of the fallax-pezizoides lineage. The species was thoroughly described and illustrated by Holmskjold (1799) and the name sanctioned by Fries (1822). No authentic specimen of Holmskjold’s remains at the Botanical Museum in Copenhagen. We also refer H. fuliginosa to this species, typified with Schaeffer’s original plate. Helvella pulla resembles H. sublicia in colours; however, ascocarp ontogeny and microanatomical details discriminate among the two.
Helvella queletiana Sacc. & Traverso in Sacc., Syll. Fung. 19: 850. 1910
Synonyms. Helvella venosa Quél., Compt. Rend. Assoc. Franc. Avancem. Sci. 9: 672. 1881, nom. illegit. (homonym).
= Helvella phlebophora Sacc., Syll. Fung. 8: 20. 1889, nom. illegit. (homonym).
Neotype designated here: Denmark, Zealand, Boserup, 29 Apr. 1995, B.T. Olsen (C-F-45303). MycoBank MBT375414.
Illustration — Dissing (1966a: f. 8d–e).
Description of the species — Dissing (1966b: 95).
Specimen examined/sequenced. Denmark, Zealand, Boserup, under Fagus, 29 April 1995, B.T. Olsen [H403] (C-F-45303 neotype).
Notes — Helvella queletiana is a spring fungus similar to H. phlebophora in colours and folding of the apothecium; however, there are morphological details as well as eight rpb2 substitutions that distinguish the two species.
Helvella rivularis Dissing & Sivertsen, Bot. Tidsskr. 75: 101. 1980 — Fig. 5d
Holotype. Norway, Nordland, Rana, Krokstrand, Tørrbekken, 7 Sept. 1972, H. Dissing & S. Sivertsen MO 95.72 (C-F-59447).
Illustration — Dissing & Sivertsen (1980: f. 2).
Description of the species — Dissing & Sivertsen (1980: 101).
Specimens examined/sequenced. Norway, Nordland, Rana, Krokstrand, Tørrbekken, 7 Sept. 1972, H. Dissing & S. Sivertsen MO 95.72 [H276] (C-F-59447 holotype); ibid., 7 Sept. 1975, H. Dissing & S. Sivertsen [H312] (C-F-86705); Oppland, Dovre, Grimsdalen, at Tverråi, on calcareous soil on river terraces and slopes along the river, 6 Aug. 1984, T. Schumacher & K. Østmoe D 128.84 [H368] (O-253385); ibid., 25 Aug. 1996, T. Schumacher [H245] (O-253381); Grimsdalen, Veslegrimsa, on gravel on river bed, 15 Aug. 2001, T. Schumacher [H363] (O-253382); Grimsdalen, Tverrgjelbekken, on river terrace, 8 Aug. 2009, T. Carlsen, T. Schumacher & I. Skrede TS 25.09 [H025] (O-253380); Grimsdalen, Buåi, in rivulet running into Buøi, 17 Sept. 1983, T. Schumacher & K. Østmoe D 167.83 [H365] (O-253383); Buådalen, on rich, moldy soil in subalpine birch forest under Aconitum septentrionale, 8 Aug. 1982, T. Schumacher & K. Østmoe 149.82 [H424] (O-253386); Grimsdalen, Grimsa at Storberget, dry fen vegetation under Salix in subalpine birch forest, 18 Sept. 1983, T. Schumacher & K. Østmoe D 226.83 [H366] (O-253384); Nordland, Rana, Dunderland, Messingåa, 8 Sept. 1972, S. Sivertsen & H. Dissing [H319] (C-F-92124).
Notes — Helvella rivularis has a bicentric subalpine to alpine distribution in Norway. Records of the species from other areas and countries need to be re-assessed based on a DNA barcoding approach. Some specimens referred to as H. rivularis in Abbott & Currah (1997), including specimens from Canada and the lowlands of Norway, are shown here to belong elsewhere.
Helvella scyphoides Skrede, T. Carlsen & T. Schumach., sp. nov. — MycoBank MB820329; hsp barcode GenBank KY784273; rpb2 barcode GenBank KY772523; Fig. 5b
Etymology. From Latin ‘cup-shaped’ apothecium.
Holotype. Norway, Hedmark, Åmot, Kvannbekken, 30 Aug. 2002, T. Pousi (O-65348).
Apothecia regularly stipitate-cupulate, cup 1.5–3 cm broad, 1–2 cm high; hymenium greyish brown (5-6D3), drying dark brown (5-6F5-8), receptacle surface paler, drying greyish (E1), densely pubescent; stipe slender, terete, towards base with inconspicuous grooves, whitish to yellowish, 0.4–0.5 cm broad, 2–3.5 cm high, densely covered by pyramidal ‘warts’ (bundles of hyphoid hairs). Medullary excipulum of textura intricata, individual hyphae hyaline, 3–4 μm broad. Outer excipulum of textura angularis to textura prismatica, interspaced with broad, thick-walled hyaline hyphae; outermost layer with cells densely packed in rows, cells 15–30 μm diam, outermost cells giving rise to short-segmented hyaline, hyphoid hairs, 100–150 μm long, consisting of drum-shaped to club-shaped cells, partly arranged in fascicled tufts, heavily staining in CB. Asci pleurorhynchous, 270–295 × 13–15 μm. Ascospores ellipsoid, 19.8–21.6 × 11.7–12.8 μm. Paraphyses subhyaline to light brown, 3–4 μm broad below, septate, enlarged to 6–8 μm at the clavate tips.
Specimen examined/sequenced. Norway, Hedmark, Åmot, Kvannbekken, 30 Aug. 2002, T. Pousi [H140] (O-65348 holotype).
Notes — This new species, recorded on the basis of a single specimen from Norway, forms a distinct lineage in our phylogeny. Helvella scyphoides resembles the sister species H. hypocrateriformis in shape and colour, but the stipe is longer and stipe and receptacle more densely pubescent. Five rpb2 and six hsp substitutions distinguish H. scyphoides and H. hypocrateriformis. A specimen from Estonia, recorded as H. aff. cupuliformis in Landeros et al. (2015), is concluded to belong here based on the published LSU nrDNA sequence (cf. Appendix 3).
Helvella semiobruta Donadini & Berthet, Bull. Soc. Mycol. France 91: 555. 1975
Holotype. France, Rhone, D’Aubagne, 15 Dec. 1972, J.C. Donadini (LY P.B. 949). Isotype. (C-F-45507).
Illustrations — Donadini & Berthet (1975: f. 1–8).
Description of the species — Donadini & Berthet (1975: 553).
Specimens examined/sequenced. France, Rhone, D’Aubagne, 15 Dec. 1972, J.C. Donadini [H277] (C-F-45507 isotype). – Spain, Valdepenas, Cindad Real, 25 Mar. 1975, F. Calonge [H307] (C-F-45467).
Notes — This Mediterranean species has a convex cap and a short greyish stipe with a few longitudinal blunt ridges and grooves. It is nested with H. helvellula and H. juniperi in a highly divergent lineage of the lacunosa clade (clade C).
Helvella silvicola (Beck) Harmaja, Karstenia 14: 103. 1974
Basionym. Otidea silvicola Beck, in Sacc., Syll. Fung. 8: 97. 1889.
Synonyms. Wynnella silvicola (Beck) Nannf., Ann. Bot. Fenn. 3: 309. 1966.
Peziza atrofusca Beck, Fl. Hernstein: 131. 1884, nom. illeg. (homonym).
Otidea atrofusca Beck ex Rehm, in Winter, Rabenh. Krypt.-Fl., ed. 2, 1.3 (lief 43): 1027. 1894, nom. inval. (Art. 36.1(c)).
Wynnea atrofusca R. Heim, Bull. Soc. Mycol. France 41: 442. 1926 (‘1925’), nom. illeg. (Art. 52.1).
Wynnella atrofusca Svrček, Česka Mykol. 17: 45. 1963, nom. illeg. (Art. 52.1).
Holotype. Austria, Niederösterreich, Schwarzau, in silvis abietinis, June 1883, Beck (PRC-1372 ex Herb. Beck (‘Peziza atrofusca n. sp.’), not examined.
Otidea neglecta Massee, Grevillea 22: 66. 1894.
Misapplied name
– Wynnella auricula (Schaeff.) Boud. sensu Boud., Icon. Mycol., liste prélim.: 2. 1904.
Illustrations — Boudier (1905–1910: pl. 250, as Wynnella auricula), Svrček (1963: pl. 48), Van Vooren (2013: photos).
Descriptions of the species — Boudier (1905–1910, vol. 4: 134, as Wynnella auricula), Parslow & Spooner (2009: 102), Van Vooren (2013: 172).
Specimens examined/sequenced. Kirghisia, Tianshan Interior, Montes Naryntau, 22 July 1967, A. Raitviir [H281] (C-F-60841). – Switzerland, Graubünden, Las Paluda, on the ground under Pinus, 1750 m asl, 29 Aug. 1984, H. Dissing [H489] (O); Graubünden, Mota Jüda, Val Plavna, along path, 1600 m asl, 30 Aug. 1979, E. Horak & H. Dissing [H283] (C-F-56847); Graubünden, along the river E of Suras, Mats da Nossa Donna, 31 Aug. 1984, H. Dissing [H282] (C-F-57385).
Notes — The species has previously been accommodated in the genus Wynnella, which, except for the ear-shaped apothecium, is morphologically similar to Helvella. Dissing (1972) concluded that the ear-shape and the horny consistency of the dried apothecium were the only differences to separate Wynnella from Helvella. Previous data also suggested a grouping of Wynnella with Helvella (O’Donnell et al. 1997, Landvik et al. 1999). Wynnella occupies an isolated position outside the rest of the helvellas in our phylogenetic trees, which may support its generic status within the Helvellaceae.
Helvella solitaria P. Karst., Bidrag Kannedom Finlands Natur Folk 19: 37. 1871 — Fig. 3b
Synonyms. Peziza solitaria P. Karst., Not. Sallsk. Fauna Fl. Fenn. Forh. 10: 111. 1869, nom. illeg. (homonym).
Lachnea solitaria (P. Karst.) Bizz. & Sacc., Mycoth. Veneta no. 323. 1876.
Holotype. Finland. Mustiala, in horto ad marg. Rivula, 21 Sept. 1866, P. Karsten PAK 3288 (H), not examined. Epitype designated here: Sweden, Uppland, Uppsala, Stadsskogen, W of Skogshall, 13 July 1942, R. Gustafsson & S. Lundell (C- Fungi Exs. Suec. 3267 ‘Helvella queletii Bres.’). MycoBank MBT375415.
Helvella queletii Bres., Rev. Mycol. 4: 211. 1882.
Acetabula queletii (Bres.) Benedix, Kulturpfl. 10: 365. 1962.
Holotype. Italy, Bosee di lance sopra Terzolaj, May 1882, Bresadola (S), not examined.
Helvella ulvinenii Harmaja, Karstenia 19: 42. 1979, fide Landeros et al. (2015).
Illustrations — Dissing (1964: f. 9, 1966b: f. 18), Dissing & Nannfeldt (1966: pl. 2, 2), Weber (1972: f. 27–28), Harmaja (1977a: f. 1, 1979: f. 8, as H. ulvinenii), Schumacher & Mohn Jenssen (1992: 21).
Descriptions of the species — Dissing (1964: 118, 1966b: 73), Weber (1972: 169), Harmaja (1979: 42), Schumacher & Mohn Jenssen (1992: 21).
Specimens examined/sequenced. Canada, Alberta, Gregoire Lake, Provincial Park near Fort McMurray, on the ground at roadside under Populus tremuloides, 26 July 1996, S.P. Abbott [H122] (FH). – France, Savoie, Gorge du Mal, 2400 m asl, on sandy soil, 31 Aug. 1992, R. Fellner F 64.92 [H090] (O-253371). – Georgia, Kaukasus, Kasberghi, Gvethi valley, on sandy soil rich in herbs along trail, 2000 m asl, 13 July 1988, E. Johannesen [H252] (O-253372). – Iceland, Sudur-Mulasysla, Hallormsstadur, Mörkin nursery and arboretum, on soil, 4 Aug. 1993, H. Knudsen [H340] (C-F-34316); Nordur-Mulasysla, Holl in Hjaltastadarthingha, on sandy soil, 5 Aug. 1993, O. Aas [H341] (C-F-34321). – Norway, Akershus, Nesodden, Flaskebekk, Bellevue, on grass in garden, 26 June 1987, A.E. Torkelsen 80.87 [H171] (O-129536); Oslo, Hovedøya, under detritus, 9 June 1996, L. Winter [H176] (O-58891); Oppland, Dovre, Grimsdalen, Tverråi, on silt among mosses underneath Salix shrubs, 7 Aug. 1989, T. Schumacher G66.89 [H080] (O-253376); ibid., 16 Aug. 2001, T. Schumacher [H004] (O-253374); ibid., 21 Aug. 2005, T. Schumacher [H186] (O-253377); ibid., 21 Aug. 2005, T. Schumacher [H187] (O-253378); Oppland, Dovre, Grimsdalen, Storberget, 8 Aug. 2009, T. Carlsen, I. Skrede & T. Schumacher TS 15.09 [H021] (O-253375); Oppland, Lom, Høyrokampen, 1400 m asl, in Dryas association, 29 Aug. 1957, F.-E. Eckblad [H353] (O-129541); Hedmark, Folldal, Råtåsjøhøi, 6 Aug. 1980, on soil, E. Johannesen [H358] (O-253373); Sør-Trøndelag, Oppdal, Vårstigen to Mt Knutshø, north-western slopes in birch forest on rich, calcareous soil, 14 Aug. 1985, T. Schumacher [H215] (O-253379); ibid., N. Knutshø, 14 Aug. 1987, A.-E. Torkelsen 148.87 [H174] (O-129533); Nordland, Rana, Sdr. Bjøllåvatn 20 km W of Lønsdal, on soil in Dryas association, 17 Aug. 1981, H. Dissing Rana 81.012 [H315] (C-F-54680); ibid., Rana 81.014 [H316] (C-F-54682); ibid., Rana 81.015 [H314] (C-F-54683). – Russia, Lukunskij forest reserve, NE Khatanga, on silty soil with mosses on lake embankment, 23 Aug. 1993, H. Knudsen [H326] (C-F-34410). – Sweden, Uppland, Uppsala, Stadsskogen, W of Skogshall, on clayey soil in coniferous plantation, 13 July 1942, R. Gustafsson & S. Lundell; ibid., 6 July 1948, S. Lundell & R. Morander [H370] (C-Fungi Exs. Suec. 3267 epitype). – Switzerland, Graubünden, S – Charl, Val Sesvanna, 2300–2400 m asl, under Pinus, 31 Aug. 1984, S. Redhead [H330] (C-F-57390).
Notes — Dissing (1966b) adopted Bresadola’s name (H. queletii) for this fungus. Harmaja (1977a) and Landeros et al. (2012), after a re-study of the holotype specimen of Peziza solitaria, concluded that the name H. solitaria was the older and correct name for this fungus. Since the holotype specimen is old and presumably cannot be critically identified by a barcode sequence, we have selected an epitype specimen from Sweden to support the typification. The morphology and LSU nrDNA sequence of the holotype specimen of H. ulvinenii [JX 993085], which was examined by Landeros et al. (2012, 2015), matches H. solitaria in all details and represents the high altitude form of H. solitaria (cf. Appendix 2). This alpine morphotype has also been observed in our sample from alpine areas in Europe. A specimen from the USA designated H. acetabulum [KC122805], and two specimens from Spain and one from Germany recorded by Landeros et al. (2015), also belong here (cf. Appendix 2). The world-wide distributed collections of H. solitaria studied by us vary in only one rpb2 character. Helvella solitaria is nested in an unsupported acetabulum-solitaria lineage in our phylogeny. It differs from its sister species H. dryadophila in 16 rpb2, 12 hsp, 14 tef and 35 LSU substitutions.
Helvella sublicia Holmsk., Beata Ruris Otia Fung. Dan. 2: 51. 1799 — Fig. 9e
– non Peziza sublicia Holmsk., Beata Ruris Otia Fung. Dan. 2: 26, pl. 10. 1799 (= H. fibrosa (Wallr.) Korf).
Lectotype designated here: Holmsk., Beata Ruris Otia Fung. Dan. 2: pl. 27. 1799. MycoBank MBT375710. Epitype designated here: Denmark, Zealand, Sorø Sønderskov, 2 Oct. 1993, K. Hansen (C-F-39823). MycoBank MBT375416.
Synonyms. Helvella ephippium Lév., Ann. Sci. Nat., Bot., sér. 2, 16: 240. 1841.
Leptopodia ephippium (Lév.) Boud., Hist. Classif. Discomyc. Europe: 37. 1907.
Lectotype. Lév., Ann. Sci. Nat., Bot., sér 2, 16, t. 15, f. 7. 1841.
?Peziza helvelloides Fr., Summa Veg. Scand. 2: 348. 1849, nom. illeg. (homonym).
Illustrations — Holmskjold (1799: pl. 27), Léveillé (1841: pl. 15, f. 7, as H. ephippium), Dissing & Lange (1967: f. 1a, 7d, as H. ephippium).
Descriptions of the species — Holmskjold (1799: 51), Dissing (1966b: 114, as H. ephippium), Dissing & Lange (1967: 355).
Specimens examined/sequenced. Denmark, Zealand, Sorø Sønderskov, 2 Oct. 1993, K. Hansen [H417] (C-F-39823 epitype); Hareskoven, 29 Sept. 1993, K. Hansen & S.K. Sandal [H422] (C-F-39826); Møn, Klinteskoven at Nælderenden, soil in roadside, 30 Oct. 1999, K. Hansen KH-99-25 [H423] (C-F-53175). – Norway, Akershus, Asker, Rabben, moldy soil in deciduous forest, 2 Oct. 1994, G. Gulden 88.94 [H148] (O-70080); Bærum, Kalvøya, on calcareous soil, 28 Aug. 1988, T. Schumacher [H229] (O-253312).
Notes — Helvella sublicia has commonly been treated under the younger synonymous name of H. ephippium. Holmskjold’s fungus was apparently overlooked in Fries’ sanctioning works (Fries 1822, 1828, 1832), but was probably recognized later by him under the new name Peziza helvelloides (Fries 1849). Helvella sublicia differs from the other European saddle-shaped Helvella species in its greyish colours and pubescent receptacle surface. Dissing & Nannfeldt (1966) compared H. sublicia (as H. ephippium) with H. fibrosa (as H. villosa) and concluded they were probably closely related, a conclusion which gains no support by our molecular phylogeny (cf. Fig. 2). We observed that the hsp and rpb2 sequences were identical among the Norwegian and Danish specimens.
Helvella sulcata Afzel., Kongl. Vetensk. Acad. Nya Handl. 4: 305. 1783
Basionym. Helvella sulcata Afzel., Kongl. Vetensk. Acad. Nya Handl. 4: 305. 1783; Fr., Syst. Mycol. 2: 15. 1822.
Lectotype designated here: Afzel., Kongl. Vetensk. Acad. Nya Handl. 4: t. 10, f. 1, 1783. MycoBank MBT375711. Epitype designated here: Norway, Oslo, Malmøykalven, in grass field on calcareous soil, 13 Oct. 2006, O. Smith & A.-S. Karlsson (O-68095). MycoBank MBT375417.
Synonyms. Helvella monacella Schaeff., Fung. Bavar. Palat. Nasc. 4: 106. 1774. (‘Elvela’), nom. inval.
Helvella lacunosa Afzel. b. (forma) minor, stipites nigricante Fr., Syst. Mycol. 2: 15. 1822.
Lectotype designated here: Schaeff., Fung. Bavar. Palat. Nasc. 2: t. 162. 1763, ‘Elvela decima qvarta’ (= Helvella monacella Schaeff.). MycoBank MBT375712.
Illustration — Schaeffer (1763: pl. 162, as ‘Elvela decima qvarta’).
Apothecia stipitate-capitate, cap 1–4 cm broad, 3–6 cm high, saddle-shaped, bi- to trilobate, adnate, margin attached to stipe at 3–4 points; hymenium black, even or slightly wrinkled; receptacle surface greyish brown (5DE4-7), smooth, without ribs on receptacle surface; stipe 2–5 cm high, 0.5–3 cm broad, (apothecial height : stipe height = 1 : 1), naked, prominently ribbed, ribs sharp and deep, partly double-edged and anastomosing, with deep ‘holes’ (lacunae) inbetween, 4–6 along the circumference, almost no stipe flesh. Medullary excipulum of loose textura intricata, hyphae c. 4–5 μm broad. Outer excipulum of more broad-celled textura intricata, the hyphae turning out parallel and arranged in dense, prismatic, hyaline cell rows perpendicular to apothecial surface, cells 8–15 μm broad, outermost cells club-shaped, 10–30 × 5–15 μm, loosely arranged, dense colouring in CB. Asci pleurorhynchous, 290–320 × 13–16 μm. Ascospores broadly ellipsoid, 14.8–16.8 × 10.5–13.2 μm. Paraphyses straight, light brown below, increasingly brown-coloured above, 3–4 μm broad, clavately enlarged to 6.5 μm at the tips.
Specimens examined/sequenced. Norway, Oslo, Malmøykalven, in grass field on calcareous soil, 13 Oct. 2006, O. Smith & A.-S. Karlsson [H152] (O-68095 epitype). – Spain, La Rioja, Clavijo, 22 Nov. 2013, R. Martinez RM 2238 [H1027] (O-253406).
Notes — The species is apparently rare in Northern Europe, but may have been confused with H. lacunosa (cf. Dissing 1964, 1966b). Afzelius (1783) was uncertain whether H. sulcata deserved specific rank when he described H. sulcata and recorded seven additional species of Helvella from Sweden. He also regretted not having had access to Schaeffer’s monumental work during his investigations (Afzelius 1783). Schaeffer’s illustration (1763) and description (1774) of Elvela monacella appear to match H. sulcata in all details. As H. monachella was sanctioned by Fries for another fungus (Fries 1822), the name is unavailable for Afzelius’ fungus. We have found good reasons to retain H. sulcata as a specific taxon, clearly distinguishable from H. lacunosa and other representatives of the H. lacunosa morphospecies complex in morphological and molecular details. We have selected Afzelius’ original illustration as a lectotype (iconotype) and supported it with a newly collected epitype from Norway. Nine rpb2, five hsp and 15 LSU substitutions distinguish H. sulcata and H. lacunosa. A specimen from the USA named H. lacunosa in Nguyen et al. (2013) has identical LSU sequence [KC-122796] with our specimen of H. sulcata from Norway (cf. Appendix 3).
Helvella terrestris (Velen.) Landvik, Mycologia 91: 283. 1999
Basionym. Pindara terrestris Velen., Monogr. Discom. Bohemiae: 341. 1934.
Lectotype selected by Van Vooren (2014): Czech Republic, Mnichovice, at the river in summer, (PRM-147368), not examined.
Illustration — Van Vooren (2014: 43).
Description of the species — Landvik et al. (1999: 283).
Specimen examined/sequenced. Norway, Nord-Trøndelag, Verdal, Ramsås, along rivulet with Trichophaea sp., 28 Aug. 1983, S. Sivertsen 83-138 [H284] (C).
Notes — This morphologically and molecularly highly diverged species was discussed in detail in Landvik et al. (1999) (cf. Fig. 2).
EXTRALIMITAL SPECIES
In addition to the 55 European Helvella species treated above, we discovered 27 extralimital, non-European species lineages among the sequenced specimens that are shortly commented upon.
Helvella capucinoides Peck, Bull. New York State Mus. Nat. Hist. 157: 27. 1912
Holotype. USA, Massachusetts, North River, Warren Co., 27 Sept. 1911, Peck (NYS), not examined.
Specimens examined/sequenced. USA, Michigan, Washtenaw Co., Stinchfield Woods, 13 Oct. 1968, N.J. Smith 2147 [H398] (C-F-92116); Maine, Pratt Island, W. Southport, on mossy garden soil, 21 Sept. 1997, D. Pfister [H113] (FH).
Notes — Helvella capucinoides may represent an available name for this American taxon. The species is apparently also known from Mexico (Landeros et al. 2015; as H. albella (JX 993063)). The two North American collections share identical rpb2 and hsp sequences. Peck described three American species of the H. elastica-group, i.e., H. gracilis (1872), H. adhaerens (1902) and H. capucinoides (1912). These taxa were reconsidered by Dissing & Lange (1967), who merged them into their broad concept of H. elastica. Authentic specimens of H. capucinoides were illustrated in Dissing (1966b: 129, f. 35). Two rpb2, three hsp and three LSU bp characters distinguish H. capucinoides and H. panormitana.
Helvella compressa (Snyder) N.S. Weber, Beih. Nova Hedwigia 51: 35. 1975
Basionym. Paxina compressa Snyder, Mycologia 28: 486. 1936.
Holotype. USA, Washington, Eaton, 5 May 1934, L.C. Snyder (WTU). Isotype (NY), not examined.
Illustration — Weber (1975: pl. 8).
Description of the species — Weber (1975: 35).
Specimen examined/sequenced. USA, California, Fort Oro, leaf duff of deciduous wood (oak), 14 Feb. 1998, K. Richmond [H112] (FH).
Notes — The specimen in FH supplied partial sequences from hsp, rpb2 and LSU. A morphologically similar but molecularly divergent species was recently reported from the American continent under this name (Landeros et al. 2015); however, the published LSU sequences [JX 993052, AY544655] are suggestive of H. corbierei (cf. Appendix 3).
Helvella costata Schwein., Syn. Fung. Carolinae Sup. Sec. Obs.: 88. 1822
Specimen examined/sequenced. USA, California, Berkley, off RTE24, Fish Ranch Road, 18 Feb. 1998, B. Neill [H100] (FH).
Notes — The cited specimen located in FH is tentatively referred to H. costata. It may eventually serve as a neotype for this American taxon. It is distinguishable from the European H. acetabulum by three rpb2, seven hsp and seven LSU characters. Two specimens from USA [KC 122798, KC 122799], recorded as H. leucomelaena in Landeros et al. (2015), belong here (cf. Appendix 3). Helvella unicolor sensu Abbot & Currah (1997) may represent this species as well.
Helvella crassitunicata N.S. Weber, Beih. Nova Hedwigia 51: 30. 1975
Holotype. USA, Washington, Mount Rainier, Narada Falls, 10 Aug. 1948, A.H. Smith 30052 (MICH-11561), not examined.
Illustration — Landeros et al. (2012: f. 24).
Descriptions of the species — Weber (1975: 30), Landeros et al. (2012: 47).
Specimens examined/sequenced. Canada, British Columbia, Whistler Nat. Park, on soil in subalpine spruce forest, 13 Aug. 1994, T. Schumacher [H222] (O-253286). – USA, Washington, Pierce Co., Mount Rainier Nat. Park, Gobler’s Knob, 12 Oct. 1952, A.H. Smith 40875 [H299] (C-F-92106 paratype).
Notes — Landeros et al. (2012) examined the morphology of the holotype specimen in MICH. We have supplied a partial rpb2 sequence from a paratype specimen (Smith 40875) of H. crassitunicata located in C.
Helvella ephippioides S. Imai, Bot. Mag. Tokyo 46: 172. 1932
Isotype. Japan, Hokkaido, Nopporo, Ishikari, ad terram in silvis, 7 Aug. 1927, S. Imai (UPS), not examined.
Specimen examined/sequenced. Japan, Honshu, Tochigi Pref., Okunikko, S Yonoko lake, 23 Aug. 1983, T. Schumacher J 71.83 [H085] (O-253267).
Notes — Among the Helvella collections from Japan studied by us, we discovered this morphologically and molecularly well-characterised species, which should be compared with authentic specimens of H. ephippioides (Imai 1932). Helvella ephippioides shares macromorphological and ascospore characteristics (acuminate spores) with H. macropus and was placed in synonymy with H. macropus by Dissing (1966b). We have supplied hsp, tef, rpb2 and LSU sequences from the above cited specimen. Helvella subfusispora in Liu et al. (1985) should also be compared with H. ephippioides.
Helvella griseoalba N.S. Weber, Michigan Bot. 11: 162. 1972
Holotype. USA, Michigan, Cheboygan Co., Univ. of Michigan Biological Station, Douglas Lake, 10 June 1968, N.J. Smith (MICH), not examined.
Illustrations — Weber (1972: f. 22), Landeros et al. (2012: f. 26).
Descriptions of the species — Weber (1972: 162), Landeros et al. (2012: 49).
Specimen examined/sequenced. USA, Michigan, Washtenaw Co., Crooked Lake, 9 June 1973, under Quercus, C. Nimke 347 [H306] (C-F-92112).
Notes — Landeros et al. (2012) compared the type specimens of H. costifera and H. griseoalba and found several morphological features that separated the two. We have studied material of H. griseoalba from Washtenow Co., Michigan (9 June 1973 C. Nimke 347), determined by Nancy Smith Weber and deposited in C. This specimen deviates from H. costifera in five hsp and nine rpb2 characters. Landeros et al. (2015) supplied an LSU sequence from the holotype of H. griseoalba deposited in MICH. A few European collections assigned to H. griseoalba in the fungarium of C have been re-examined by us and are referred to either H. costifera, H. calycina or H. confusa.
Helvella maculata N.S. Weber, Beih. Nova Hedwigia 51: 27. 1975
Holotype. USA. Michigan, Idaho Bonner Co., Hoodoo Mountain, 5 Oct. 1968, H.V. Smith & N.J. Smith 2124 (MICH), not examined.
Illustrations — Weber (1975: pl. 7, f. 1–2), Landeros et al. (2012: f. 29).
Description of the species — Weber (1975: 27).
Specimen examined/sequenced. USA, Alaska, at Eklutna Lake, 24 Aug. 1973, V.L. Wells & P.E. Kempton 1577 [H303] (C-F-45400).
Notes — The cited specimen forms a single specimen lineage in our phylogeny (cfr. Fig. 2).
Helvella stevensii Peck, Bull. Torrey Bot. Club. 31: 182. 1904
Synonyms. Leptopodia stevensii (Peck) Le Gal, Rev. Mycol. 2: 9. 1937.
Holotype. USA, Michigan, Detroit, June 1903, R.H. Stevens (NYS), not examined.
Helvella connivens Dissing & M. Lange, Mycologia 59: 351. 1967.
Holotype. USA, Michigan, Cross Village, along roadside, 21 July 1947, Brooks, Lange & A.H. Smith (MICH). Isotype. ibid., (C-F-45350).
Illustrations — Weber (1972: f. 43–46).
Descriptions of the species — Weber (1972: 193, 1975: 35).
Specimens examined/sequenced. USA, Michigan, Cross Village, along roadside, 21 July 1947, Brooks, Lange & A.H. Smith [H297] (C-F-45350 isotype of H. connivens); Michigan, Cross Village area, Wycamp Lake Road, Cheboygan Co., 8 July 1969, N.J. Smith 2209 [H394] (C-F-92117).
Notes — Dissing & Lange (1967) and Dissing (2000) included North American as well as European specimens in their broad concept of H. stevensii. The European specimens of H. stevensii sensu Dissing & Lange (1967) are here referred to H. corbierei. Weber (1972) and Landeros et al. (2015) compared authentic material of H. stevensii and H. connivens and concluded they were the same, a disposition concurred by us.
Helvella subglabra N.S. Weber, Michigan Bot. 11: 179. 1972
Holotype. USA, Michigan, Wahtenaw Co., Stinchfield Woods, near Dexter, 13 Oct. 1968, N.J. Smith 2145 (MICH), not examined.
Illustrations — Weber (1972: f. 32–33), Landeros et al. (2012: f. 35).
Descriptions of the species — Weber (1972: 179), Landeros et al. (2012: 58).
Specimen examined/sequenced. USA, Michigan, Washtenaw Co., Pickerel Lake, under white pine, 4 Oct. 1972, N.J. Smith [H280] (C-F-65405 topotype).
Notes — Helvella subglabra is nested in the fallax-pezizoides lineage. Abbott & Currah (1997) and Landeros et al. (2012) examined the holotype specimen of H. subglabra and placed it in synonymy with H. pezizoides. This is contradicted by our results. Although similar to H. pezizoides in shape, H. subglabra is greyish brown in colour and distinguished molecularly from H. pezizoides by three rpb2 and two hsp substitutions.
Helvella vespertina N.H. Nguyen & Vellinga, Mycologia 105: 1281. 2013
Holotype. USA, California, Sonoma, Salt Point State Park, 15 Jan. 2012, T.W. Osmundson (UC-1999204), not examined.
Illustration — Nguyen et al. (2013: f. 1D–J).
Description of the species — Nguyen et al. (2013: 1281).
Specimens examined/sequenced. USA, California, San Francisco, Land’s End, 4 Nov. 1981, R. Trial [H102] (FH); Maine, Washington Co., at Humbolt Research Institute, on station ground, 1 July 2000, S. LaGreca [H116] (FH).
Notes — This species of the H. lacunosa morphospecies complex was recently described from North America. The species has not been recorded from Europe.
Helvella sp. ‘INDIA’
Specimen examined/sequenced. India, Udhampur, Sanasar, on humid soil in Pinus excelsa forest, 10 Aug. 1967, K.S. Thind [H421] (C-F-45704).
Notes — This distinctive Asian specimen belongs in the elastica clade (clade E) (cf. Fig. 2).
Helvella sp. ‘JAPAN 1’
Specimen examined/sequenced. Japan, Honshu, Tochigi Pref., Okunikko, Chuzenjikohan, Asegata, forest floor, 22 Aug. 1983, T. Schumacher J 35.83 [H206] (O-253388).
Notes — Morphologically, the Japanese specimen resembles H. fibrosa, which is the sister species to ‘Japan 1’ and ‘Japan 2’ in our molecular phylogeny.
Helvella sp. ‘JAPAN 2‘
Specimen examined/sequenced. Japan, Honshu, Tochigi Pref., Okunikko, Chuzenjikohan, 22 Aug. 1983, T. Schumacher J 33.83 [H093] (O-253389).
Notes — The specimen is morphologically similar to H. fibrosa from which it is distinguished by 16 hsp and 12 rpb2 substitutions.
Helvella sp. ‘JAPAN 3‘
Specimen examined/sequenced. Japan, Fokushima Prefecture, Tadami, Mt Asakusa, on rotten trunk of Fagus crenulata, 30 Oct. 1991, L. Ryvarden 30206 [H213] (O-253390).
Notes — The specimen is sister to H. pallescens in our phylogeny.
Helvella sp. ‘JAPAN 4’
Specimens examined/sequenced. Japan, Honshu, Tochigi Pref., Okunikko, Chuzenjikohan, Asegata, forest floor, 25 Aug. 1983, T. Schumacher J 105.83 [H461] (O-253391); ibid., T. Schumacher J 104.83 [H478] (O-253392).
Notes — This species belongs in the H. lacunosa morphospecies complex. It possesses a greyish black hymenium and greyish stipe when fresh. It is distinguished from its sister H. atra by four hsp and two rpb2 substitutions.
Helvella sp. ‘JAPAN 5’
Specimen examined/sequenced. Japan, Honshu, Tochigi Pref., Okunikko, Karikami, along inlet creek S lake, 23 Aug. 1983, T. Schumacher J 70.83 [H466] (O-253393).
Notes — This specimen belongs in the H. lacunosa morphospecies complex. It possesses a greyish hymenium and stipe when fresh.
Helvella sp. ‘JAPAN 6’
Specimen examined/sequenced. Japan, Honshu, Tochigi Pref., Okunikko, Chuzenjikohan, Asegata, forest floor, 25 Aug. 1983, T. Schumacher J 103.83 [H482] (O-253394).
Notes — This specimen is morphologically similar to H. sublicia, but clusters in another lineage, i.e., the fallax-pezizoides lineage.
Helvella sp. ‘PAPUA 1’
Specimen examined/sequenced. Papua New Guinea, Manki, W Bulolo, Morobo District, on soil in Castanopsis forest, 24 Apr. 1972, E. Horak [H432] (C-F-45531).
Notes — This specimen was recorded as Helvella sp. (aff. H. pezizoides?) by Dissing (1979). It differs from H. pezizoides in two rpb2 and four hsp substitutions.
Helvella sp. ‘PAPUA 2‘
Specimen examined/sequenced. Papua New Guinea, Manki, W Bulolo, Morobo District, among litter under Lithocarpus and Castanopsis, 13 Oct. 1971, E. Horak [H400] (C-F-45434).
Notes — This specimen was referred to H. lacunosa by Dissing (1979) from which it diverges in seven rpb2 and six hsp characters. The stipe is whitish.
Helvella sp. ‘USA 1’
Specimen examined/sequenced. USA, New York, Ithaca, Six mile creek, among leaves and soil in deciduous wood, 24 Aug. 1989, H. Dissing, Cornell 28.89 [H416] (C-F-58799).
Notes — This specimen is nested in the fallax-pezizoides lineage. It shows morphological resemblance to H. fallax, but can be distinguished by a conspicuously long and narrow stipe. Five rpb2 bp and eleven hsp substitutions discriminate against the latter.
Helvella sp. ‘USA 2’
Specimen examined/sequenced. USA, California, Yolo Co., Stebbins Reserve, Cold Canyon, in duff under Laurus, Jan.–Feb. 1988, D. Pfister [H103] (FH).
Notes — This specimen of the H. lacunosa morphospecies complex is molecularly nested in the lacunosa clade (clade C).
Helvella sp. ‘USA 3’
Specimen examined/sequenced. USA, Idaho, Bonner Co., across the river from Priest River, 29 June 1972, N.S. Weber 3629 [H376] (C-F-92118).
Notes — This specimen belongs in the elastica clade (clade E). It resembles H. monachella in morphology from which it differs in 15 hsp characters. We were unsuccessful in obtaining an rpb2 sequence from this specimen.
Helvella sp. ‘USA 4’
Specimen examined/sequenced. USA, Massachusetts, Concord, Eastbrook, on calcareous soil, 25 June 1998, D. Pfister [H104] (FH).
Notes — Molecularly, the specimen represents an American relative of H. pallescens.
Helvella sp. ‘USA 5’
Specimen examined/sequenced. USA, Iowa, White Rock Conservancy, on soil in oak – hikory woods, 22 July 2008, R. Healy [H110] (FH).
Notes — This specimen belongs in the corbierei-stevensii lineage of the elastica clade (clade E). It differs from Helvella ‘USA 6’ in four rpb2 and three hsp substitutions and from H. levis in three rpb2 and one hsp substitutions.
Helvella sp. ‘USA 6’
Specimen examined/sequenced. USA, Michigan, Oakland Co., Highland State Recreation Area, Goose Pond, on dirt along path, 27 Aug. 1979, N.S. Weber 4784 [H291] (C-F-92104).
Notes — This specimen belongs in the corbierei-stevensii lineage of the elastica clade (clade E). Morphologically, the specimen resembles H. levis from which it is distinguished by four hsp and no rpb2 substitutions. It differs from H. stevensii in seven hsp and seven rpb2 substitutions and from H. corbierei in three hsp and four rpb2 substitutions.
Helvella sp. ‘USA 7’
Specimen examined/sequenced. USA, Michigan, Washtenaw Co., Ann Arbor, Saginaw Forest, under conifers, 11 July 1940, A.H. Smith 15183 [H377] (C-F-92119).
Notes — The above cited specimen from USA, Michigan, has been referred to as H. sulcata by Nancy Weber (annotation on herbarium specimen). It differs from European collections of H. fusca in three rpb2 characters and five hsp characters, from Helvella ‘USA 4’ in five rpb2 bp and five hsp characters, from Helvella ‘PAPUA 2’ in four rpb2 and 4 hsp characters, and from H. lactea in three rpb2 and four hsp characters.
Helvella sp. ‘USA 8’
Specimen examined/sequenced. USA, Michigan, Luce Co., Tahquamenon Falls State Park, Upper Falls, in humus in dense mixed woods, 14 Aug. 1956, H.D. Thiers 4093 [H429] (C-F-92105).
Notes — This specimen is nested in the fallax-pezizoides lineage. It differs from its sister taxon H. pezizoides in seven rpb2 and six hsp characters.
DISCUSSION
We discovered 83 genetically distinct species lineages based on sequence data from 432 Helvella specimens from Europe and some scattered localities on other continents. When we combined our molecular data with morphology and biogeography, our results suggest that there are at least 55 Helvella species in Europe. Our data are also suggestive of a fair number of undescribed species occurring on the American and Asian continents (Fig. 1). Formal recognition of American and Asian species is deferred, pending additional material from these regions.
Phylogenetic informative loci in Helvella
More than one molecular marker is needed in order to discriminate species when using a GCPSR approach. How many loci that are needed is widely discussed (Rokas et al. 2003, Aguileta et al. 2008, Balasundaram et al. 2015). Recent studies have shown that which molecular markers you choose to use is as important as how many (Aguileta et al. 2008, Balasundaram et al. 2015). The choice of partial sequences from a few genes with high phylogenetic informativeness can allow the construction of robust phylogenies and minimize the amount of sequences needed (Townsend 2007, Aguileta et al. 2008, Schmitt et al. 2009, Balasundaram et al. 2015). Development of new markers that are informative at different phylogenetic levels may thus be useful for inferring phylogenetic relationships and resolving species lineages.
In our study we analysed four loci, one of these – hsp – was not previously used for species delimitation in the Pezizales. However, hsp has proven useful at the intraspecific level in the pyrenomycete Daldinia locata (Johannesson et al. 2000) and in recognising cryptic species in the basidiomycete genera Heterobasidion and Serpula (Johannesson & Stenlid 2003, Kauserud et al. 2006, Balasundaram et al. 2015). The hsp marker was successfully amplified from most strains in our dataset, possessed high phylogenetic informativeness and was useful in delimiting species in Helvella. The partial hsp sequence amplified was only 270 bp, which may explain the success in obtaining sequences from old specimens.
The three other molecular markers (tef, rpb2 and LSU) of our dataset are widely used in fungal phylogenies, inclusive of the Pezizales. Recently, tef was suggested as a secondary barcode for fungi because of its ability to resolve closely related species (Hansen & Olariaga 2015, Stielow et al. 2015). However, while we experienced that the tef primers worked well on DNA extracted from fresh specimens, it usually failed to amplify DNA from even recently dried Helvella specimens. For this reason, tef was excluded from our final phylogenetic analyses, since we wanted to construct a phylogeny that contained a broad sample of taxonomically important (type) specimens of Helvella. Rpb2 has also been a successful inclusion in multilocus datasets to delimitate species in genera of the Pezizales, e.g., in Peziza (Hansen et al. 2005), Morchella (Taskin et al. 2010, 2012, O’Donnell et al. 2011, Du et al. 2012a), Tuber (Bonito et al. 2013) and Otidea (Hansen & Olariaga 2015). This marker also worked well across the breadth of the genus Helvella.
In fungi the LSU marker, or parts of LSU (D1/D2 region) combined with the ITS from the nuclear ribosomal DNA, has long been used to discriminate species on its own (Kurtzman & Robnett 1998, Fell et al. 2000, Schoch et al. 2012). It has recently also been employed as the sole locus to infer phylogenies and discriminate species of helvellas from the American continents (Nguyen et al. 2013, Landeros et al. 2015). The LSU marker proved useful in resolving some species relationships in our study as well and had the highest phylogenetic informativeness of the markers in our dataset (Appendix 2). However, the level of LSU divergence across the inferred lineages was not homogenous compared to the rpb2 and hsp data (Table 3). In the alpestris-nannfeldtii lineage the LSU diversity was very high, and the branch of H. nannfeldtii was exceptionally long compared to most lineages in our LSU tree. In this respect it differed from the other loci where the diversity in this lineage was comparable to the other lineages. Thus, by adding information from the diversity analyses to our understanding of the different loci we concluded that rpb2 and hsp outperformed the LSU nrDNA locus in species discrimination power.
The unsuitability of ITS in isolation to infer phylogeny and delimit species in Helvella was already demonstrated by Landvik et al. (1999). They reported that Helvella differed remarkably from other molecular studies of pezizomycete genera at that time in that the ITS nrDNA region was too divergent to align across the breadth of the genus and that even the usually highly conserved ssu and 5.8S nrDNA gene varied among the species. Nevertheless, later studies have shown that ITS may be a useful nrDNA region to include in a multilocus dataset in smaller genera and lineages and subgroups of highly divergent pezizomycete genera as well, e.g., in Morchella (Taskin et al. 2010, 2012, Du et al. 2012a, b), Otidea (Olariaga et al. 2015), and Helvella (Zhao et al. 2015, 2017).
Our results add to a number of studies within the Ascomycota that have shown that sequence data from single copy protein-coding genes are more informative than the nuclear ribosomal DNA in resolving species (Hansen et al. 2005, Hofstetter et al. 2007, Taskin et al. 2012, Hansen & Olariaga 2015). We therefore advocate for using the combined information from single copy genes such as hsp, tef and rpb2, and LSU in combination, for phylogenetic and systematic studies in Helvella.
Inference of species and lineages in Helvella
Historically, species recognition in Helvella, as in most groups of Pezizomycetes, has relied on a morphological species concept mostly drawing on macroscopic characters of the apothecium, including the hymenium, excipulum and stipe. The shape, colours and outer surface characters of the cap and stipe have been especially important characters for the discrimination of species. The research by Dissing (1964, 1966a, b) and Dissing & Nannfeldt (1966) emphasized the importance of paraphysis pigmentation, reflected in the colour of the hymenium and concluded that in most species hymenium colour is a constant feature, e.g., in H. crispa and H. lactea the colour is always whitish, in H. sublicia and H. pulla greyish, and in H. corium and H. helvellula blackish. According to Dissing (1966b), a larger variation in colour of the hymenium was allowed in some species ranging from pale greyish brown to brown to nearly black, e.g., in H. solitaria (as H. queletii), H. lacunosa and H. elastica. From field observations on fresh Helvella specimens and in working on dried fungarium samples, we observed that colours of fresh apothecia can change drastically even when specimens were gently dried. In general, we have adopted a more restricted set of colour range of hymenium and receptacle surface for some species guided by observations of fresh specimens of the pertinent species.
There are few distinctive microanatomical characters useful in species delimitation in Helvella. As already pointed out in the introduction, most species and species groups are described by a suite of graded characters shared by restricted or more extended groups of species. Dissing (1972) concluded that the (only) distinctive characters on species level were type of fruitbody, colour, and glabrous or pubescent outer surface of the receptacle and stipe. Except for H. macropus, the spores were of minor diagnostic value. We found the microanatomy of the ectal excipulum and specifically the shape, colour, and distribution of the outermost hyphoid hairs, whether fascicled or scatteredly distributed – or both – on the receptacle surface, to be of particular diagnostic value in morphologically similar groups, e.g., in the fallax-pezizoides, alpestris-nannfeldtii and the alpina-corium lineages. The pigmentation and shape of the apical portion of the paraphyses also helped in species identification in some species groups, e.g., in the crispa and the alpestris-nannfeldtii lineages. As already pointed out by Häffner (1987) and Landeros et al. (2015), a major two-division of Helvella species is observed in the shape of the ascus base, whether aporhynchous (no crozier) or pleurorhynchous (forked, with crozier). The leucomelaena lineage has species with an aporhynchous ascus base, which is also gained or retained in the alpina-corium lineage, that separates against the rest of the helvellas with a pleurorhynchous ascus base. For a detailed discussion on micromorphological features valued and used in species discrimination of helvellas, we refer to Dissing (1966b), Eckblad (1968), Weber (1972, 1975) and Landeros et al. (2012, 2015).
Our phylogenetic analyses revealed several Helvella morphospecies complexes that harboured pseudocryptic variation, i.e., species that can be recognized morphologically only after molecular systematic data unveiled their existence (Knowlton 1993, 2000). For instance, the morphologically similar species pairs of H. corium/H. macrosperma, H. lactea/H. pallescens and H. elastica/H. panormitana, which were initially indiscernable due to the lack of discriminating morphological characters, could be resolved as genealogically exclusive using sequence data. Our results indicate that these species pairs are genetically divergent and located in distantly related Helvella lineages.
Furthermore, our study showed that in Helvella, DNA sequence information contributed valuable diagnostic characters and could even serve as the backbone of taxonomic descriptions for many species. Nevertheless, it is assumed that a re-evaluation of macromorphological characters such as ascocarp ontogeny and shape and colour of the apothecia based on fresh specimens of the pertinent species may reduce the number of morphologically indistinguishable species and thus support the molecularly discernible phylogenetic Helvella species.
In the same way as morphology is not a good predictor of phylogeny, we observed that our sequence data was sometimes insufficient to fully resolve morphologically and ecologically dissimilar species pairs, e.g., H. juniperi and H. semiobruta of the helvellula lineage, and H. philonotis and H. palustris of the palustris-philonotis lineage. Thus, additional phylogenetically informative markers are needed to improve the resolution within these lineages.
The origin and evolution of the large and complex ascomata of Helvella have inspired several evolutionary hypotheses and proposals for infrageneric subdivisions of Helvella (Le Gal 1947, Dissing 1966b, Weber 1972, Abbott & Currah 1997, Landeros et al. 2015). These infrageneric classifications mostly rest on differences in ascocarp ontogeny and outer surface characters of receptacle and stipe in different species groups and have been reviewed recently by Landeros et al. (2015). Our phylogenies do not provide a robust framework for evaluating these morphology-based infrageneric classification schemes proposed by earlier workers, as we gave priority to include a few ribosomal and protein coding loci of short length (269–700 bp) to be successful in amplifing old as well as newly collected fungarium specimens, providing high levels of informativeness at the tips of the phylograms, at the expense of longer segments of genes necessary to develop a robust hypothesis of evolutionary relationships within the genus. To infer infrageneric groups that reflect evolutionary relationships it will be necessary to identify combinations of genes that produce a fully resolved backbone and describe a complete set of highly supported lineages within this large genus of the Pezizomycetes.
A few patterns worth mentioning, however, is the substipitate and externally ribbed and internally chambered or hollow inflated stipitate species groups nested in the early diverging branches of our trees, opposed to the later diverging branches, e.g., the alpestris-nannfeldtii, the alpina-corium lineages and clades C, D and E constituting species groups with cylindrical (terete), hollow or solid stipes. The hairiness of the outer receptacle surface and stipe that were given heavy weight in the infrageneric classifications proposed by Dissing (1966b), Weber (1972) and Abbott & Currah (1997), i.e., transitions from hairless to pubescent to villose apothecia, or reverse, seem to have occurred in many lineages, implying that these character states are not useful in delimiting monophyletic groups in Helvella.
DNA barcodes in Helvella
The molecular-based methods now provide an accessory approach for species identification. Once species are well established under a robust phylogenetic framework, it becomes possible to identify specimens to species by DNA barcoding of only one or a few loci. Our study repeatedly reminded us that a 100 % conclusive identification had to rely on barcode sequences in addition to morphological examination of the specimens under study.
ITS is accepted as the universal barcode for fungi by the Consortium for the Barcode of Life (Schoch et al. 2012). Using a universal barcode is advantageous, as was shown by Du et al. (2012b) in the comparably large genus Morchella. They determined the species identity of GenBank sequences of Morchella using the ITS barcode for species already delimited by a multilocus phylogeny and concluded that ITS sequences were useful in identifying 77 % of the known phylospecies and that at least 66 % of the named Morchella sequences in GenBank are misidentified. We assume that comparable results will be found for Helvella, thus it would be advantageous to provide ITS sequences for all Helvella species as well. ITS sequences for selected Helvella species have been obtained in some studies (Landvik et al. 1999, Nguyen et al. 2013, Ariyawansa et al. 2015, Landeros et al. 2015, Zhao et al. 2015, 2017). However, we only managed to amplify ITS for a few species in our initial trials. The difficulties in amplifying the ITS for some Helvella species when using universal primers were first documented by Weideman (1998), who found it profitable to produce selective ITS primers to amplify targeted Helvella groups from ectomycorrhizal root tips.
We observed the inter- and intraspecific barcode gaps of rpb2 and hsp in combination allowed for a conclusive identification of all European Helvella species of our sample. Nevertheless, it would be profitable also to obtain species-specific ITS sequences for barcoding purposes. Bearing in mind the difficulties in obtaining ITS amplicons for many Helvella species, which may possibly be caused by primer mismatch using the currently available ITS primers, it would be useful in the near future to develop Helvella specific ITS primers for barcoding purposes. Such primers would probably also improve the possibility of detecting Helvella ITS sequences from soil amplicon studies using ITS as the target sequence.
Consequently, at present we advocate using partial rpb2 and hsp gene sequences, which have a high PCR and sequencing success rate, as the primary DNA barcodes to molecularly identify old and fresh specimens of Helvella.
Biogeography and ecology
In their overviews of European and North American Helvella species, Dissing (1964, 1966a, b, Dissing & Lange 1967), Weber (1972, 1975) and Abbott & Currah (1997) included samples from different regions and continents that encompass many morphologically similar but distinct phylogenetic species of Helvella. This implies that biogeographic and ecological hypotheses of Helvella spp. are problematic because most reports in the literature include specimens with misapplied binomials. The level of disjunct intercontinental distribution of species in our dataset is limited; from the many species included from Europe and North America, most are endemic to one continent (Fig. 1). However, some species were found on more than one continent, e.g., H. alpestris, H. capucina and H. solitaria were all found in Europe and North America, H. phlebopora in Greenland and East Asia, and H. fistulosa in Europe, North America and East Asia. Our sampling efforts have been on Europe, thus, there are too few collections from other continents to confidently draw conclusions regarding the distribution of species. Nevertheless, because of the large proportion of endemism seen from our sample we suspect that more species are to be found on other continents, as also reflected by the many new published species from, e.g., China over the last few decades (Liu et al. 1985, Liu & Cao 1988, Cao & Liu 1990, Cao et al. 1990, Zhuang 2004, Zhao et al. 2015, 2017, Wang et al. 2016). High levels of endemisms have also been demonstrated in other comparably large Pezizalean genera. In Morchella only two out of 18 species from Eurasia were present in both Europe and Asia (O’Donnell et al. 2011), while in Otidea only 6 of 29 species in Europe were also present in North America (Hansen & Olariaga 2015).
Our results show that many closely related species are sympatric throughout their range, while others are partially sympatric or allopatric. For instance, the morphologically similar parasympatrically occurring H. corium, H. alpina, H. nannfeldtii and H. alpestris occupy overlapping habitats of the alpine biome of northern Europe. Further sampling on other continents and a re-examination of herbarium collections, including DNA sequences for species identification, will be necessary to re-assess the biogeography and ecology of Helvella worldwide.
Molecular ecological studies have documented the presence of Helvella spp. as part of the mycorrhizal fungal community of a broad spectrum of trees and shrubs (Weidemann 1998, Tedersoo et al. 2006, Hwang et al. 2015). In an LSU nrDNA phylogenetic study of potential pezizalean ectomycorrhizae, Tedersoo et al. (2006) obtained four unknown ectomycorrhizal sequences nested within Helvella. Two newly described species in North America, i.e., H. vespertina and H. dryophila were reported as ectomycorrhizal with gymnosperms and Quercus, respectively (Nguyen et al. 2013). More recently, ITS sequences representing three clades within Helvella were obtained from ectomycorrhizal root tips in addition to a local sample; however, members of two of the Helvella clades were not recovered from uncultured root tips (Hwang et al. 2015). Moreover, Hobbie et al. (2001) investigated isotopes of five Helvella samples that exhibited diverse levels of enrichment and depletion of γ 13C and γ 15N, and concluded that it was unclear whether Helvella has a mycorrhizal life style. We observed that some Helvella species are always found in close association with the same plants, e.g., H. dryadophila and H. arctoalpina with Dryas and H. fusca with Populus. Nevertheless, further investigation of these relationships, by sampling and sequencing ectomycorrhizal root tips, is needed to confirm ectomycorrhizal relationships among Helvella species.
Acknowledgments
We acknowledge all the collectors of Helvella who deposited specimens in fungarias in Oslo, Copenhagen and Farlow Herbaria for their dedicated efforts in collecting valuable material. We thank the following curators and staff for supplying material: Donald H. Pfister (FH), Katriina Bendiksen (O), Karl-Henrik Larsson (O) and Henning Knudsen (C). We thank Cecilie Mathiesen, Birgitte Lisbeth Thorbek, Synnøve Botnen and Anders B. Aas for assistance in the molecular laboratory. We thank Karen Hansen and Xianghua Wang for allowing the use of KH.10.277 H. carnosa in our analyses. Karen Hansen, Donald H. Pfister and Kerry L. O’Donnell reviewed the manuscript and we are most grateful for their valuable comments and corrections. We acknowledge ‘Nansenfondet og de dermed forbundne fond’ and the University of Oslo for funding.
INDEX TO SPECIES AND VARIETIES
Accepted names treated in this work are in bold-italic. The number refers to the pagenumber.
acetabulum 217
aestivalis 214
alba 225
albella 221
albida 225
albipes 232
alpestris 217
alpicola 219
alpina 219
arctica 232
arctica var. macrosperma 232
arctoalpina 220
atra 220
bicolor 221
calyciformis 221
calycina 221
capucina 222
capucinoides 239
carnosa 223
compressa 239
confusa 223
connivens 240
corbierei 224
corium 224
costata 239
costifera 224
crassitunicata 240
crispa 225
cupuliformis 229
dalgeri 220
danica 225
dissingii 228
dovrensis 235
dryadophila 226
elastica 226
ephippioides 240
ephippium 238
esculenta 225
fallax 226
fibrosa 228
fistulosa 228
fuliginosa 236
fusca 228
griseoalba 240
helvellula 229
hispida 232
hyperborea 229
hypocrateriformis 229
juniperi 230
lactea 230
lacunosa 230
latispora 231
leucomelaena 231
leucomelas 231
leucophaea 231
leucopus 232
levis 231
macropus 231
macrosperma 232
maculata 240
monachella 232
murina var. alpestris 218
nannfeldtii 233
nigra 233
nigricans 227
nivea 225
oblongispora 233
pallescens 234
pallida 225
palustris 234
panormitana 234
paraphysitorquata 235
pedunculata 223
pezizoides 235
philonotis 235
phlebophora 236
pithyophila 231
pubescens 236
pulchra 229
pulla 236
queletiana 236
queletii 235
rivularis 237
scyphoides 237
semiobruta 237
silvicola 237
solitaria 238
stevensii 240
subglabra 240
sublicia 238
sulcata 238
terrestris 239
ulvinenii 238
venosa 236
vespertina 240
villosa 228
Appendix 1
Extralimital Helvella taxa not treated in the present synopsis. Altogether 470 Helvella names, representing taxa originally described as Helvella or later referred to or excluded from Helvella for taxonomic or nomenclatural reasons, are summarized in Index Fungorum (http://www.indexfungorum.org). In addition to the currently accepted names for the various Helvella species, which are to be found in Species Fungorum (http://www.speciesfungorum.org), the Index includes all accepted or suspected synonymous, homonymous and unpriorable names of the accepted species of Helvella and its former segregate genera. We have compiled a list of names suspected by us to represent synonymous or applicable names to extant Helvella species, but not treated in our synopsis or not yet fully accounted for in the Index Fungorum. Many of these will have to be typified and eventually re-assessed in a future monograph of the genus. * = accepted species in Species Fungorum; # = uncertain species affinity in Index Fungorum.
*Helvella adhaerens Peck, Ann. Rep. N. Y. State Mus. Nat. Hist. 54: 956. 1902.
Dissing 1966b: 131 = H. elastica.
*Helvella aestivalis (R. Heim & L. Rémy) Dissing & Raitv., Eesti NSV Tead. Akad. Toim. Biol. Ser. 23: 105. 1974.
*Helvella affinis Velen., Ceske Houby 4–5: 890. 1922.
Dissing 1966b: 64 = H. macropus.
*Helvella agaricoides Kreh, Pilz- Kräuterfreund: 205. 1922.
Peziza amphora Quél., Bull. Soc. Bot. France 23: 331. 1877 ‘1876’.
Index Fungorum = H. solitaria; Dissing 1966b: 37 = H. leucomelaena.
*Helvella astieri Korf & Donadini, Rep. Tottori Mycol. Inst. 10: 397. 1973.
*Helvella aterrima Velen., Monogr. Discom. Bohemiae: 388. 1934.
Dissing 1966b: 102 = H. lacunosa.
*Helvella bachu Q. Zhao, Zhu L. Yang & K.D. Hyde, in Zhao Q, Sulayman M, Zhu X-t, et al., Mycologia 108: 830. 2017 ‘2016’.
#Helvella badia Saut., Flora, Regensburg 24 (20): 319. 1841.
Sauter 1841: 319 = H. affinis lacunosa.
Helvella barlae Boud. & Pat., J. Bot., Paris 2: 445. 1888.
Index Fungorum = H. crispa; Dissing 1966b: 88 = H. crispa.
*Helvella beatonii (Rifai) Harmaja, Karstenia 14: 103. 1974.
*Helvella branzeziana Svrček & J. Moravec, Česka Mykol. 22: 87. 1968.
#Helvella brevipes DC apud de Candolle & Lamarck, Fl. France, ed. 3 (Paris) 6: 28. 1815.
Dissing 1966b: 140 = H. leucopus.
Helvella brevipes Gillet, Champ. France Discomyc. 1: 11. 1879, nom. illeg. (Art. 53.1) (homonym).
*Helvella brevis (Peck) Harmaja, Karstenia 14: 104. 1974.
Abbott & Currah 1997: 75 = H. macropus.
*Helvella brevissima Peck, Bull. Torrey Bot. Club 30: 100. 1903.
*Helvella bulbosa Font Quer, Cavanillesia 3: 173. 1931.
Helvella bulbosa (Hedw.) Kreisel, Boletus, Schr.Reihe 1: 29. 1984, nom. illeg. (Art. 53.1) (homonym)
Index fungorum = H. macropus.
*Helvella chinensis (Velen.) Nannf. & L. Holm, in Lundell, Nannfeldt & Holm, Publ. Herb. Univ. Uppsala 18: 5. 1985.
#Helvella cinerea Vill., Hist. Pl. Dauphiné 3, 2: 1045. 1789.
Fries 1822: 20, 1832: 95 = H. pulla.
*Helvella cinerella Velen., Monogr. Discom. Bohemiae: 386. 1934.
Dissing 1966b: 115 = H. ephippium.
#Helvella cinereocandida Hazsl., Ertek. Természettud. Köréb. Magyar Tud. Akad. Ertec 11 (19): 14. 1882 ‘1881’.
*Paxina clypeata (Pers.) Linder, in Polunin, Bull. Natl. Mus. Canada, Biol. Ser. 26, 97: 257. 1947.
#Helvella conformis Pers., Mycol. Eur. 1: 216. 1822.
Fries 1822: 25, 1832: 95 = Verpa patula Fr. (?).
*Helvella constricta Boud., Hist. Classif. Discomyc. Europe: 36. 1907.
Dissing 1966b: 102 = H. lacunosa.
Helvella cookeana (Boud.) Sacc. & Traverso, in Sacc., Syll. Fung. 19: 844. 1910.
Index Fungorum = H. pezizoides; Dissing 1966b: 119 = H. pezizoides.
*Helvella cornuta Velen., Monogr. Discom. Bohemiae: 385. 1934.
Phallus costatus Batsch, Elench. Fung. (Halle): 129. 1783.
Index Fungorum = H. crispa.
#Helvella craterella (Hedw.) Quél., Enchir. Fung.: 274. 1886 (‘Helvela’)
Dissing 1966b: 64 = H. macropus.
*Helvella dissingii Korf, Mycotaxon 31: 381. 1988.
Korf 2008: 307 = H. fibrosa.
*Helvella dryophila Vellinga & N.H. Nguyen, in Nguyen NH, Landeros F, Garibay-Orijel, et al., Mycologia 105: 1278. 2013.
*Macropodia dubaleni (Boud.) Sacc. & Trotter, Syll. Fung. 22: 628. 1913.
Dissing 1966b: 131 = H. elastica.
*Paxina dupainii (Boud.) Seaver, North Amer. Cup-fungi (Operculates) (New York): 207. 1928.
Dissing 1966b: 75 = H. queletii.
*Helvella dura Velen., Monogr. Discom. Bohemiae: 386. 1934.
Dissing 1966b: 131 = H. elastica.
*Helvella engleriana Henn., Bot. Jahrb. Syst. 18 (4 (Beibl. 44)): 40. 1894.
#Helvella exarata Gillet, Champ. France Discomyc. 1: 9. 1879.
#Helvella fargesii Pat., J. Bot. 7: 344.1893.
Dissing 1966b: 131 = H. elastica.
*Helvella faulknerae Copel., Ann. Mycol. 2: 509. 1904.
Dissing 1966b: 75 = H. queletii.
*Helvella favrei Quél., Compt. Rend. Assoc. Franc. Avancem. Sci. 30: 496. 1902.
*Macropodia fechtneri Velen., Monogr. Discom. Bohemiae: 342. 1934.
*Helvella flavida Velen., Nov. Mycol. Noviss.: 157. 1947.
Dissing 1966b: 88 = H. crispa.
*Helvella foetida Velen., Monogr. Discom. Bohemiae: 387. 1934.
*Helvella fuegiana (Speg.) Eckblad, Nytt Mag. Bot. 15: 92. 1968.
#Helvella fuliginea Saut., Flora, Regensburg 24 (20): 319. 1841.
Sauter 1841: 319 = H. affinis lacunosa; Dissing 1966b: 102 = H. lacunosa.
*Helvella galeriformis B. Liu & J.Z. Cao, Acta Mycol. Sin. 7: 199. 1988.
*Helvella glutinosa B. Liu & J.Z. Cao, Acta Mycol. Sin. 7: 198. 1988.
#Helvella gracilis Peck, Ann. Rep. N.Y. State Mus. Nat. Hist. 24: 94. 1872 ‘1871’.
Dissing 1966b: 130 = H. elastica.
#Helvella grisea Clem., Bot. Surv. Nebraska 4: 8. 1896.
Dissing 1966b: 102 = H. lacunosa.
Helvella guepinioides Berk. & Cooke, in Cooke, Mycogr.: 198. 1879.
Index Fungorum = H. elastica; Dissing 1966b: 131 = H. elastica.
*Helvella hegani Copel., Ann. Mycol. 2: 510. 1904.
Dissing 1966b: 141 = H. leucopus.
*Helvella involuta Q. Zhao, Zhu L. Yang & K.D. Hyde, in Zhao QI, Tolgor B, Zhao Y, et al., Phytotaxa 239, 2: 135. 2015.
*Helvella javanica Penz. & Sacc., Malpigia 15: 1. 1901.
Dissing 1979: 156 = H. lacunosa.
*Helvella jiaohensis J.Z. Cao, L. Fan & B. Liu, Acta Mycol. Sin. 9: 184. 1990.
*Helvella jilinesis J.Z. Cao, L. Fan & B. Liu, Acta Mycol. Sin. 9: 185. 1990.
*Helvella jimsarica W.Y. Zhuang, Mycotaxon 90: 39. 2004.
Helvella klotzschiana Corda, in Sturm, Deutschl. Fl. 3: 121. 1831.
Index Fungorum = H. elastica; Dissing 1966b: 130 = H. elastica.
*Cyathipodia lerchenfeldii (Schulzer) Boud., Hist. Classif. Discomyc. Europe: 39. 1907.
*Cyathipodia longipes Boud., Hist. Classif. Discomyc. Europe: 39. 1907.
*Helvella ludovicae J. Kickx f., Fl. Crypt. Flandres 1: 502. 1867.
*Paxina macropus (Clem.) Seaver, North Amer. Cup-fungi (Operculates) (New York): 203. 1928.
*Helvella maroccana Har. & Pat., Bull. Soc. Mycol. France 20: 64. 1904.
*Helvella mesatlantica Malençon, Bull. Soc. Mycol. France 95: 121. 1979.
*Helvella minor (Velen.) Rauschert, Haussknechtia 4: 52. 1988.
Dissing 1966b: 68 = H. villosa.
Helvella mitra L., Sp. Pl. 2: 1180 (‘Elvela’), nom. ambiguum (fide Fries 1822: 14).
Helvella murina (Boud.) Sacc. & Traverso, in Sacc., Syll. Fung. 19: 849. 1910.
Index fungorum = H. ephippium; Dissing 1966b: 115 = H. ephippium.
*Acetabula murina Zeller, Mycologia 19: 139. 1927.
*Paxina nemoralis (Speg.) Raithelh., Metrodiana Sonderh. 2: 19. 1983.
Gamundi 1960: 278 = H. leucomelaena.
#Acetabula ochroleuca Velen., Ceske Houby 4–5: 862. 1922.
Dissing 1966b: 52 = H. acetabulum.
*Phleboscyphus olivaceus Clem., Bull. Torrey Bot. Club 30: 93. 1903.
*Helvella orienticrispa Q. Zhao, Zhu L. Yang & K.D. Hyde, in Zhao QI, Tolgor B, Zhao Y, et al., Phytotaxa 239: 136. 2015.
*Helvella pallidula N.S. Weber, Michigan Bot. 11: 171. 1972.
Abbott & Currah 1997: 42 = H. fibrosa (as H. chinensis).
*Helvella papuensis Dissing, Beih. Sydowia 8: 156. 1979.
Notes — According to Dissing (1979) the species belongs in the H. crispa group. We were unsuccessful in obtaining any barcode sequences from the isotype specimen (C-F-45480) in C.
Peziza percevali Berk. & Cooke, in Cooke, Mycogr., Vol. 1. Discom. (London) (3): 111. 1876.
Index fungorum = H. leucomelaena; Dissing 1966b: 39 = H. leucomelaena.
*Helvella pileata Clem., Bull. Torrey Bot. Club 30: 94. 1903.
*Helvella platycephala Benedix, Kulturpfl. 19: 172. 1972.
*Helvella platypodia (Boud.) Donadini, Bull. Soc. Linn. Provence 36: 136. 1985.
Dissing 1966b: 75 = H. queletii.
*Helvella pseudolacunosa Q. Zhao & K.D. Hyde, in Ariyawansa HA, Hyde KH, Jayasiri SC, et al., Fung. Diversity 75: 142. 2015.
*Helvella pseudoreflexa Q. Zhao, Zhu L. Yang & K.D. Hyde, in Zhao QI, Tolgor B, Zhao Y, et al., Phytotaxa 239, 2: 137. 2015.
#Helvella pygmaea Sommerf., Suppl. Fl. Lapp.: 288. 1826.
*Helvella quadrisulca Velen., Nov. Mycol. Noviss.: 157. 1947.
Dissing 1966b: 109 = H. lactea.
*Helvella quinqueloba Pers., Mycol. Eur. 1: 216. 1822.
Persoon 1822: 216 = Boletus albus, pileolo complicato nigro Battarra (1759: 24; pl. 2: f. H); Fries 1832: 97 = H. monachella.
*Phleboscyphus radicatus Clem., Bull. Torrey Bot. Club 30: 94. 1903.
#Peziza rehmiana Sacc., Michelia 1: 544. 1879.
*Helvella robusta S.P. Abbott, in Abbott & Currah, Mycotaxon 33: 242. 1988.
*Helvella rossica Velen., Nov. Mycol.: 204. 1939.
Dissing 1966b: 131 = H. elastica.
*Helvella rugosa Q. Zhao & K.D. Hyde, in Ariyawansa HA, Hyde KH, Jayasiri SC, et al., Fung. Diversity 75: 142. 2015.
Helvella schaefferi Boud., Hist. Classif. Discomyc. Europe: 36. 1907.
Dissing 1966b: 102 = H. lacunosa.
*Helvella scrobiculata Velen., Ceske Houby 4–5: 898. 1922.
Dissing 1966b: 75 = H. queletii.
*Acetabula simplex Rolland, Bull. Soc. Mycol. France 17: 117. 1901.
Dissing 1966b: 39 = H. leucomelaena.
*Helvella sinensis B. Liu & J.Z. Cao, in Liu, Du & Cao, Acta Mycol. Sin. 4: 214. 1985.
*Helvella solida Velen., Monogr. Discom. Bohemiae: 387. 1934.
Dissing 1966b: 131 = H. elastica.
Peziza stipitata H.J. Huds., Fl. Angl., ed. 2, 2: 639. 1778.
Index Fungorum = H. macropus.
*Paxina subclavipes (W. Phillips & Ellis) Seaver, North Amer. Cup-fungi (Operculates): 206. 1928.
Dissing 1966b: 64 = H. macropus.
Helvella subcostata Cooke, Mycographia 1: 90. 1879.
Index Fungorum = H. lacunosa; Dissing 1966b: 102 = H. lacunosa.
*Helvella subfusispora B. Liu & J.Z. Cao, in Liu B, Du F, Cao J-Z, Acta Mycol. Sin. 4: 211. 1985.
*Helvella sublactea Q. Zhao, Mei Wang & Y.C. Zhao, in Wang M, Zhao Y-C, Zhao QI, et al., Phytotaxa 253, 2: 136. 2016.
*Helvella subspadicea Q. Zhao, Zhu L. Yang & K.D. Hyde, in Zhao Q, Sulayman M, Zhu X-t, et al., Mycologia 108: 832. 2017 ‘2016’.
#Helvella tabacina Mont., Fl. Chil. 1: 396. 1854.
*Helvella taiyuanensis B. Liu, Du & J.Z. Cao, Acta Mycol. Sin. 4: 211. 1985.
*Helvella unicolor (Boud.) Dissing, Rev. Mycol. 31: 219. 1966.
Notes — In the description of H. unicolor, Dissing (1966a, b) reported that the species is vernal, which is in accordance with the neotype specimen selected by Dissing from France, and collected in April. Dissing’s description and drawings of the species are, however, based on a specimen in F. Petraks Flora Bohemiae et Moraviae exsiccate no. 2326 (as ‘Acetabula sulcata’ (Pers.) Fuckel) from Czechia, collected in September 1930. This specimen has been re-examined by us, but we were unsuccessful in obtaining any barcode sequences from this specimen. The immature specimens of Petrak’s are morphologically similar to H. costifera. According to Dissing (1966b), H. unicolor is distinguished from H. acetabulum and H. costifera by excipulum anatomy and exceptionally broad spores, 17–(19.2)–21 × 13–(14.6)–16 μm. The hymenium, as well as the upper portion of the receptacle surface and ribs are cacao-brown to leather-brown (tan), gradually becoming ochreous towards the base (Dissing 1966b). These latter observations could not be reconfirmed by us and the possibility of a mix up of two morphologically similar taxa under the name H. unicolor seems likely. The identity of H. unicolor, as well as the collections referred to as H. unicolor by Dissing (1966a, b) and by Abbott & Currah (1997) from North America, need to be reassessed based on a molecular approach.
*Macropodia urceolata Clem., Bull. Torrey Bot. Club 30: 91. 1903.
Dissing 1966b: 64 = H. macropus.
*Helvella vacini Velen., Nov. Mycol. Noviss.: 156. 1947.
Dissing 1966b: 68 = H. villosa.
#Helvella venosa Fr., Syst. Mycol. 2: 22. 1822.
#Helvella verpoides Fr., Summa Veg. Scand. 2, sectio Post.: 346. 1849.
*Helvella verruculosa (Berk. & M.A. Curtis) Harmaja, Karstenia 18: 57. 1978.
Notes — Abbott & Currah (1997) compared the type specimens of H. verruculosa, H. arctoalpina and H. dryadophila and concluded they were conspecific. Later, Landeros et al. (2012) compared the holotype specimens of H. verruculosa and H. dryadophila and placed them in synonymy for the following reasons: “We did not observe chambers in the stipe and neither the ribs on the sterile surface of the apothecium. In fact, for the last reason, H. verruculosa can be confused with H. solitaria and H. ulvinenii, which do not have ribs on the sterile surface”. The status of H. verruculosa needs to be re-assessed based on molecular data.
Acetabula vulgaris Fuckel, Jahrb. Nassauischen Vereins Naturk. 23–24: 330. 1870 ‘1869–1870’.
Dissing 1966b: 52 = H. acetabulum.
*Helvella xinjiangensis J.Z. Cao, L. Fan & B. Liu, Acta Mycol. Sin. 9: 186. 1990.
*Helvella zhongtiaoensis J.Z. Cao & B. Liu, Mycologia 82: 642.
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