The genus Otidea is one of the more conspicuous members of the Pyronemataceae, with high speciesdiversity in hemiboreal and boreal forests. The genus is morphologically coherent and in previous higher-level multigeneanalyses it formed a highly supported monophyletic group. Species delimitation within Otidea is controversialand much confusion has prevailed in the naming of taxa. To provide a phylogenetic hypothesis of Otidea, elucidatespecies diversity and limits we compiled a four-gene dataset including the nuclear LSU rDNA and three nuclearprotein-coding genes (RPB1, RPB2 and EF-1α) for 89 specimens (total 4 877 nucleotides). These were selected froma larger sample of material studied using morphology and 146 ITS (ITS1-5.8S-ITS2) and 168 LSU rDNA sequencesto represent the full genetic diversity. Using genealogical concordance phylogenetic species recognition (GCPSR),Bayesian and maximum likelihood analyses of the individual datasets resolved 25 species of Otidea. An additionaleight singletons are considered to be distinct species, because they were genetically divergent from their sisters.Sequences of multiple genes were included from 13 holotypes, one neotype and three epitypes. Otidea angusta,O. myosotis and O. papillata f. pallidefurfuracea are nested within O. nannfeldtii, O. leporina and O. tuomikoskii,respectively and are considered synonyms. Otidea cantharella var. minor is shown to be a distinct species. Fivenew species were discovered: O. oregonensis and O. pseudoleporina for North America; and O. borealis, O. brunneoparvaand O. subformicarum for Europe. The analyses of the individual four gene datasets yielded phylogeniesthat were highly concordant topologically, except for the RPB1 that showed supported conflict for some nodes inBayesian analysis. Excluding the RPB1 from the combined analyses produced an identical topology to the four-genephylogeny, but with higher support for several basal nodes and lower support for several shallow nodes. We argueto use the three-gene dataset to retrieve the maximum support for the higher-level relationships in Otidea, but stillutilise the signal from the RPB1 for the delimitation and relationships of closely related species. From the four generegions utilised, EF-1α and RPB1 have the strongest species recognition power, and with higher amplification successEF-1α may serve as the best secondary barcoding locus for Otidea (with ITS being a primary). The phylogenyfrom the three- and four-gene datasets is fully resolved and strongly supported in all branches but one. Two majorclades, as part of six inclusive clades A–F, are identified – and ten subclades within these: A) O. platyspora andO. alutacea subclades, and B) O. papillata, O. leporina, O. tuomikoskii, O. cantharella, O. formicarum, O. unicisa,O. bufonia-onotica and O. concinna subclades. Morphological features in Otidea appear to be fast evolving andprone to shifts, and are poor indicators of higher-level relationships. Nevertheless, a conspicuous spore ornament isa synapomorphy for the O. unicisa subclade (/Otideopsis); all other species in Otidea have smooth or verruculose (inSEM) spores. Exclusively pale to bright yellow apothecia and straight to curved, broadly clavate to distinctly capitateparaphyses are synapomorphies for a restricted O. concinna subclade (/Flavoscypha). The curved to hooked apicesof the paraphyses is suggested to be a symplesiomorphic trait for the genus. The reaction of resinous exudateson the outermost excipular cells that coalesce into amber drops in Melzer’s reagent is likely an ancestral state forclade B. We estimate that Otidea consists of 47 species worldwide, based on all available information (includingmorphology, ITS or LSU sequences, and literature descriptions). Three fifths of the species occur in Europe, with20 species recognised as endemic. At least 14 species occur in North America and 17 in Asia, with eight and tenspecies considered endemic to each continent, respectively. Our knowledge about Otidea in Asia is still fragmentaryand the diversity likely much higher.

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

The easily recognised genus Otidea is subjected to numerous problems in species identification. A numberof old names have undergone various interpretations, materials from different continents have not been compared andmisidentifications occur commonly. In this context, Otidea is monographed, based on our multiple gene phylogeniesassessing species boundaries and comparative morphological characters (see Hansen & Olariaga 2015). All namescombined in or synonymised with Otidea are dealt with. Thirty-three species are treated, with full descriptions andcolour illustrations provided for 25 of these. Five new species are described, viz. O. borealis, O. brunneoparva,O. oregonensis,O. pseudoleporina and O. subformicarum. Otidea cantharella var. minor and O. onotica var. brevisporaare elevated to species rank. Otideopsis kaushalii is combined in the genus Otidea. A key to the species of Otideais given. An LSU dataset containing 167 sequences (with 44 newly generated in this study) is analysed to placecollections and determine whether the named Otidea sequences in GenBank were identified correctly. Fourty-ninenew ITS sequences were generated in this study. The ITS region is too variable to align across Otidea, but had lowintraspecific variation and it aided in species identifications. Thirty type collections were studied, and ITS and LSUsequences are provided for 12 of these. A neotype is designated for O. cantharella and epitypes for O. concinna,O. leporina and O. onotica, along with several lectotypifications. The apothecial colour and shape, and spore charactersare important for species identification. We conclude that to distinguish closely related or morphologicallysimilar species, a combination of additional features are needed, i.e. the shape of the paraphyses, ectal excipulumstructure, types of ectal excipulum resinous exudates and their reactions in Melzer’s reagent and KOH, tomentumand basal mycelium colours and exudates. The KOH reaction of excipular resinous exudates and basal myceliumare introduced as novel taxonomic characters.