Abstract

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.

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).

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Fig. 1

Venn diagram of the localities where specimens have been collected, depicting the level of continental endemism and inter- and intracontinental distributions of Helvella in our sample set. Blue indicates species collected in Europe, yellow indicates species collected in North and South America, green indicates species collected in East Asia (Japan), red indicates species collected in West Asia (including India).

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.

RESULTS

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.

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Fig. 2

Maximum Likelihood (ML) tree of European (and some extralimital) members of Helvella using parts of the RNA polymerase II (rpb2) and the heat shock protein 90 (hsp) loci. Branches supported by both ML bootstrap analyses (> 70) and posterior probability (> 0.95) from Bayesian inference are in bold. Branches that are supported only if part of the nuclear ribosomal large subunit (LSU) is added are bold and red. Sequences from holo-/isotype (blue) and epi-/neotype specimens (green) are colour-coded. Sequences from holo-/isotypes of synonymous taxa are indicated with blue asterix: *¹ = holotype of Helvella dovrensis; *² = isotype of Helvella arctica; *³ = isotype of Helvella pulchra; *⁴ = isotype of Helvella cupuliformis; *⁵ = isotype of Helvella connivens.

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.

TAXONOMY

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

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Fig. 3

Photos of fresh (a–b) and dried (c–h) apothecia. a. H. arctoalpina (O-253395); b. H. solitaria (O-253396); c. H. acetabulum (C-Fungi Exs. Suec. 3261); d. H. dryadophila (O-72617, holotype); e. H. hyperborea (C-F-45306); f. H. confusa (C-F-70807, isotype); g. H. costifera (O-253397 ex TROM-F11436); h. H. arctoalpina (O-72616, holotype). — Scale bars = 1 cm. — Photos: a–b: T. Schumacher; c–h: I. Skrede.

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

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Fig. 4

Photos of fresh (f) and (a–e) dried apothecia. a. H. corium (C-F-92111, holotype of H. arctica); b. H. nannfeldtii (O-253343); c. H. alpestris (O ex DAOM574891); d. H. macrosperma (O-253330); e. H. alpina (O-253228, holotype); f. H. alpestris (O-253400). — Scale bars = 1 cm. — Photos: a–e: I. Skrede; f: T. Schumacher.

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

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Fig. 5

Photos of fresh (e) and dried (a–d) apothecia. a. H. hypocrateriformis (C-F-92131, isotype of H. cupuliformis); b. H. scyphoides (O-65348, holotype); c. H. alpicola (O-185924, holotype); d. H. rivularis (C-F-59447, holotype); e. H. alpicola (O-253402). — Scale bars = 1 cm. — Photos: a–d: I. Skrede; e: T. Schumacher.

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

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Fig. 6

Photos of fresh (b–c) and dried (a, d–f) apothecia. a. H. lacunosa (C-Fungi Exs. Suec. 2065, epitype); b. H. lacunosa (O-253398); c. H. palustris (O-253399); d. H. philonotis (C-F-45481, holotype); e. H. nigra (O-253345, epitype); f. H. atra (C-Fungi Exs. Suec. 2066, epitype). — Scale bars = 1 cm. — Photos: a, d–f: I. Skrede; b–c: T. Schumacher.

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

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Fig. 7

Photos of dried apothecia. a. H. panormitana (O-253363, epitype); b. H. carnosa (O-68100, holotype); c. H. levis (C-F-55188, epitype); d. H. elastica (C-F-29130, epitype); e. H. bicolor (O-105346); f. H. danica (O-253288); g. H. capucina (O-253256); h. H. fistulosa (O-253315); i. H. bicolor (C-45334, epitype). — Scale bars = 1 cm. — Photos: I. Skrede.

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.