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1: Ectal excipulum of a glabrous apothecium (Helvella leucopus var. populina). 2: Hyphal fascicle of the outer layer of a subpubescent stipe (H. griseoalba). 3: Hyphal fascicle of the ectal excipulum of a pubescent apothecium (H. costifera)-a: short, b: large. 4: Pleurorhynchous asci (H. pocillum). 5: Aporhynchous asci (H. crassitunicata); 6: Common paraphyses (H. subglabra). 7: Paraphyses with a thick-walled brown cap or collar at the apex (H. paraphysitorquata). 8: Thickwalled paraphyses (H. crassitunicata). 9: Ascospores (H. solitaria)-a: ornamented, b: smooth. Scale bars: 1-8 = 20 µm, 9 = 8 µm.
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Abstract — Sixteen type and some additional specimens representing fifteen species assigned to Helvella were studied with the aim of providing new data for their improved delimitation and recognition. Specimens were received on loan from DAOM, FH, H, K, MA, MICH, S and UPS. A synoptic key to the studied species is provided to contribute to a better...
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... layer are a "textura angularis" (almost isodiametric cells without intracellular spaces) ( Korf 1952, Dissing 1966, Eckblad 1968, Weber 1972). These cells are arranged in rows perpendicular to the surface. In glabrous ascomata, the ectal excipulum external layer is a palisade of clavate hyphal end-cells, ending more or less at the same level ( Fig. 1) (Weber 1972). Hairy ascomata have an ectal excipulum outer layer that Eckblad (1968) described as "consisting of more or less swollen chains of cells often converging into distinct clusters. " These clusters are called "hyphal fascicles" (Weber 1972) or "chains of cells and/or fascicled hyphal chains" (Abbott & Currah 1997). ...
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... used to refer to the surface of the apothecial ectal excipulum and stipe outer layer are: glabrous (Fig. 1), sub-pubescent (for fascicled hyphal chains < 50 μm long; Fig. 2), and pubescent (for fascicled hyphal chains > 50 μm long; Fig. 3). As the hairs' length can vary in the last case, their measurements are given in the descriptions. See also "Results & ...
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... studied sixteen type specimens and present only new information or data that differs from the protologue or previous publications. In addition to the characters established by previous authors (Dissing 1966b;Abbott & Currah 1997), we found that pigmentation of apothecial cells, stipe tissue anatomy, and the reddish (hemiamyloid) reaction in Melzer's reagent (Figs 10-11) are important characters that contribute to species recognition in Helvella. Because we found no variation in the basal mycelium of the stipe, which is universally composed of smooth-walled, hyaline, rr-, interwoven hyphae, we do not include it in the species descriptions. ...
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... the value of pigmentation in the structures, we think it is important to study it from two perspectives: a) location and b) intensity. Concerning location, pigment may occur in the cytoplasm, in the wall, or encrusted on the wall (Figs 19-21). We observed that while the first two are constant within species, encrusted pigments are variable. ...
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... example, in H. costifera there are specimens with encrusted paraphyses and others without such incrustations. On the matter of pigment intensity, we return to cotton blue used first by Dissing (1964) to note that there are species where the cytoplasmic pigment color is so intense that it is still visible even after adding cotton blue, while others have lighter pigment no longer visible when adding the colorant (Figs 12-17). We realized that this characteristic is constant among specimens of the same species. ...
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... ascospore ornamentation, we make the following points. 1) Ornamented ascospores are not restricted to H. subgen. Macropodes (here represented only by H. macropus, Fig. 18a) as Abbott & Currah (1997) proposed. We have also observed ornamented ascospores (generally only in mature ascospores outside the asci) in the type specimens of H. leucopus var. populina (Fig. 18b), H. paraphysitorquata (Fig. 18c), H. solitaria, H. subglabra, and H. ulvinenii. 2) We agree with Weber (1972) that it is the mature ...
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... ornamentation, we make the following points. 1) Ornamented ascospores are not restricted to H. subgen. Macropodes (here represented only by H. macropus, Fig. 18a) as Abbott & Currah (1997) proposed. We have also observed ornamented ascospores (generally only in mature ascospores outside the asci) in the type specimens of H. leucopus var. populina (Fig. 18b), H. paraphysitorquata (Fig. 18c), H. solitaria, H. subglabra, and H. ulvinenii. 2) We agree with Weber (1972) that it is the mature Helvella ascospores that are verrucose, contrary to the assumption of Dissing (1964) and Dissing & Nannfeldt (1966) that ornamented ascospores are the immature ones. 3) When citing ascospore sizes, it is ...
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... points. 1) Ornamented ascospores are not restricted to H. subgen. Macropodes (here represented only by H. macropus, Fig. 18a) as Abbott & Currah (1997) proposed. We have also observed ornamented ascospores (generally only in mature ascospores outside the asci) in the type specimens of H. leucopus var. populina (Fig. 18b), H. paraphysitorquata (Fig. 18c), H. solitaria, H. subglabra, and H. ulvinenii. 2) We agree with Weber (1972) that it is the mature Helvella ascospores that are verrucose, contrary to the assumption of Dissing (1964) and Dissing & Nannfeldt (1966) that ornamented ascospores are the immature ones. 3) When citing ascospore sizes, it is important to indicate what kind of ...
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... pubescent hyphal fascicles in the stipe while H. griseoalba does not (glabrous) or has shorter (subpubescent) hyphal fascicles; 2) H. costifera can have light brown paraphyses while H. griseoalba has only hyaline ones, and 3) in H. griseoalba the terminal cells of the apothecial hyphal fascicles have brown pigments that are visible in cotton blue (Fig. 13) while in H. costifera the light brown pigments in the cell walls are not visible in cotton blue (Fig. 12). = Helvella costifera Nannf., in Lundell & Nannfeldt, Fungi Exsicc. Suec., Fasc. 41-42: 37, 1953 Asci 240-315 × 15-18 μm. Paraphyses thin-walled, hyaline, pale brown to brown in mass, pigment deposits in cytoplasm and cell walls, ...
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... hyphal fascicles; 2) H. costifera can have light brown paraphyses while H. griseoalba has only hyaline ones, and 3) in H. griseoalba the terminal cells of the apothecial hyphal fascicles have brown pigments that are visible in cotton blue (Fig. 13) while in H. costifera the light brown pigments in the cell walls are not visible in cotton blue (Fig. 12). = Helvella costifera Nannf., in Lundell & Nannfeldt, Fungi Exsicc. Suec., Fasc. 41-42: 37, 1953 Asci 240-315 × 15-18 μm. Paraphyses thin-walled, hyaline, pale brown to brown in mass, pigment deposits in cytoplasm and cell walls, some with brown encrusted pigment on walls. Medullary excipulum hyaline to light brown. Stipe inner and ...
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Citations
... however, morphological characteristics have inherent limitations as the main means of identifying and classifying Helvella in history. In Species Fungorum, more than 300 names attributable to the genus of Helvella are recorded, but with the development of molecular biology and DNa analysis, many species were synonymized (o 'Donnell et al. 1997, Landeros et al. 2012, hansen et al. 2019, Løken et al. 2020. as a universal fungal barcode, ItS is not suitable for addressing the phylogenetics and species differentiation of Helvella due to its limited molecular data in genBank and a low amplification success rate (Skrede et al. 2017. ...
... häffner (1987) and abbott & Currah (1997) considered it was a synonym of H. atra J. könig. however, Landeros et al. (2012) examined the holotype specimen (Helvella subglabra, C-F-65405) and identified morphological differences distinguishing it from H. atra. Later, Skrede et al. (2017) confirmed H. subglabra as a distinct species through phylogenetic analysis. ...
During the investigation of Helvella in Yunnan, China, four saddle-like specimens were collected and studied. Through a comprehensive analysis of both morphological traits and phylogenetic data from LSU and hsp90 sequences, the four collections were identified and are described as two new species, i.e., Helvella liquii and H. pseudoatra. Helvella liquii stands out with its distinctive features, including the fusion of the pileus margin with the stipe, a slightly rugose coffee blackish surface on the receptacle, and relatively broad apex of the paraphyses. Helvella pseudoatra is characterized by a pubescent receptacle surface that exhibits shaded or dark grey to greyish brown. Additionally, the upwardly white to brown stipe with a basally white to greyish color further distinguishes this species. To provide a comprehensive understanding of these newly discovered taxa, detailed descriptions and illustrations of both Helvella liquii and H. pseudoatra species are accurately documented.
... As a frequently encountered and important fungal group, the systematics and species diversity of Helvella have been widely studied by mycologists from different parts of the world, especially from Europe and North America (Dissing 1966a, b, Korf 1972, Weber 1972, 1975, Harmaja 1979, Häffner 1987, Abbott & Currah 1997, Landeros et al. 2012, Nguyen et al. 2013, Skrede et al. 2017, 2020, Hansen et al. 2019, Løken et al. 2019). The ascoma shape and colour, pubescence on the apothecium receptacle surface, ascospore size and shape are highlighted in the identification of Helvella species. ...
... Sequences of all DNA regions generated in this study were deposited in GenBank. Other sequences of Helvella and related species obtained from GenBank are based on published literature (Nguyen et al. 2013, Landeros et al. 2012, Hyde et al. 2016, Wang et al. 2016, 2023a, Zhao et al. 2016a, b, Skrede et al. 2017, 2020, Hansen et al. 2019, Løken et al. 2019, or selected by using the BLASTn search function of the NCBI database to find similar matches with taxa in Helvella. For ITS sequence data, the genus search tool in GenBank database was used for retrieving Helvella-related ITS sequences. ...
Helvella is a widespread, frequently encountered fungal group appearing in forests, but the species diversity and molecular phylogeny of Helvella in China remains incompletely understood. In this work, we performed comprehensive phylogenetic analyses using multilocus sequence data. Six datasets were employed, including a five-locus concatenated dataset (ITS, nrLSU, tef1-α, rpb2, hsp), a two-locus concatenated dataset (ITS, nrLSU), and four single-locus datasets (ITS) that were divided based on the four different phylogenetic clades of Helvella recognized in this study. A total of 1 946 sequences were used, of which 713 were newly generated, including 170 sequences of ITS, 174 sequences of nrLSU, 131 sequences of tef1-α, 107 sequences of rpb2 and 131 sequences of hsp. The phylogeny based on the five-locus concatenated dataset revealed that Helvella s. str. is monophyletic and four phylogenetic clades are clearly recognized, i.e., Acetabulum clade, Crispa clade, Elastica clade, and Lacunosa clade. A total of 24 lineages or subclades were recognized, 11 of which were new, the remaining 13 corresponding with previous studies. Chinese Helvella species are distributed in 22 lineages across four clades. Phylogenetic analyses based on the two-locus concatenated dataset and four single-locus datasets confirmed the presence of at least 93 phylogenetic species in China. Among them, 58 are identified as known species, including a species with a newly designated lectotype and epitype, 18 are newly described in this paper, and the remaining 17 taxa are putatively new to science but remain unnamed due to the paucity or absence of ascomatal materials. In addition, the Helvella species previously recorded in China are discussed. A list of 76 confirmed species, including newly proposed species, is provided. The occurrence of H. crispa and H. elastica are not confirmed although
both are commonly recorded in China.
... Same as previous studies, this study also found that the morphology of saddle fungi is not enough for species definition alone (Dissing 1966, Landeros et al. 2012, Skrede et al. 2017, Løken et al. 2020. Recently, studies on the complex of Helvella in China have found that some Chinese species are endemic and different from those in Europe and America . ...
The brown cupulate saddle fungi with villus surface in China, have long been named as Helvella macropus, a well-known European and North American species. In this study, Helvella fruiting bodies from Austria, China (the Tibetan Plateau), and Germany were examined. Phylogenetic analyses of combined LSU, TEF and HSP90 sequence data showed that the Chinese H. macropus-like samples represent two phylogenetic species, one of which is H. macropus s.s., and the other is H. guttata, which is introduced as new species here. Helvella guttata and H. macropus are described and illustrated in detail and compared with their similar species. In addition, a key to the morphologically recognizable taxa of the H. macropus complex is provided.
... The genus includes a series of elaborate ascomata, from cupulate to saddle-shaped and/or from lobed to folded apothecia, which are located on terete, ribbed, or furrowed stipes (Skrede et al., 2017). Helvella ascomata usually produce a wide range of colors such as white, creamy white, gray, and brown to black (Landeros et al., 2012;Skrede et al., 2017). Some Helvella species form ectomycorrhizal symbioses with plants in several families, such as Fagaceae, Pinaceae, and Salicaceae (Nguyen et al., 2013;Hwang et al., 2015). ...
... The gross morphology of H. atroides is very similar to that of typical H. atra J. König s.s. However, the hymenial surface and stipe of H. atroides are usually dark gray to brownish black when young, while in the latter species, it is generally completely black (Landeros et al., 2012). ...
... Skrede et al. (2017) indicated that the color of fresh apothecia could change dramatically during drying. In microanatomical features, the characteristics of ectal excipulum, especially the shape, color, and distribution of the outer hyphoid hairs, are of special diagnostic value in morphologically similar species (Landeros et al., 2012). In addition, the ascus base (aporhynchous or pleurorhynchous), as well as the pigmentation and the shape of paraphyses upperpart also helped in species discrimination of some Helvella (Landeros et al., 2012;Skrede et al., 2017). ...
Most species of Helvella have been described from temperate regions in Asia, Europe, and North America, but little is known about the genus from tropical regions. In this report, phylogenetic analyses of 11 newly collected saddle-like fungi from northern Thailand using three genetic markers [the nuclear large subunit ribosomal DNA (LSU), the heat shock protein 90 (HSP90), and the translation elongation factor 1-alpha (TEF)] confirm their assignment in Helvella. Two species were described as new, i.e., Helvella atroides and H. orentitomentosa, and two species, i.e., H. fistulosa and H. rugosa, were reported for the first time in Thailand. Details of macro- and microscopic characters and illustrations were provided for each species. To date, seven species of Helvella have been recorded in Thailand, and a key for identifying the Thai Helvella species was provided here.
... The genus includes a series of elaborate ascomata, from cupulate to saddle-shaped and/or from lobed to folded apothecia, which are located on terete, ribbed, or furrowed stipes (Skrede et al., 2017). Helvella ascomata usually produce a wide range of colors such as white, creamy white, gray, and brown to black (Landeros et al., 2012;Skrede et al., 2017). Some Helvella species form ectomycorrhizal symbioses with plants in several families, such as Fagaceae, Pinaceae, and Salicaceae (Nguyen et al., 2013;Hwang et al., 2015). ...
... The gross morphology of H. atroides is very similar to that of typical H. atra J. König s.s. However, the hymenial surface and stipe of H. atroides are usually dark gray to brownish black when young, while in the latter species, it is generally completely black (Landeros et al., 2012). ...
... Skrede et al. (2017) indicated that the color of fresh apothecia could change dramatically during drying. In microanatomical features, the characteristics of ectal excipulum, especially the shape, color, and distribution of the outer hyphoid hairs, are of special diagnostic value in morphologically similar species (Landeros et al., 2012). In addition, the ascus base (aporhynchous or pleurorhynchous), as well as the pigmentation and the shape of paraphyses upperpart also helped in species discrimination of some Helvella (Landeros et al., 2012;Skrede et al., 2017). ...
... In previous years, the diagnosis of fungi was based on the phenotypic and microscopic form such as the shape of the ascus and the shape of the ripples in apothecium (Landeros, et al. 2015). But recently when the molecular identification was established, Helvella has been extensively studied by mycologists from North America , Europe, Australia and Asia ( Løken et al. 2020). ...
... Dried material was rehydrated and analyzed under compound microscope as described by Landeros et al. (2012). Briefly, hand sections were mounted in water or Melzer's reagent and measurements of spores, asci, paraphysis, width of subiculum, were registered. ...
Helvella lacunosa is a species complex, with Helvella lacunosa s.s. not currently distributed in America. The objective of this study was to resolve the taxonomy of specimens from central Mexico identified as Helvella lacunosa s.l. associated with Abies religiosa forests. The nuclear ITS and LSU regions were PCR-amplified and sequenced from dry herbaria specimens. Phylogenetic analyses were based on Parsimony, Maximum Likelihood and Bayesian Inference approaches. Sequences of Helvella from A. religiosa forests grouped into a well-supported lineage within the North American clade together with Helvella dryophila (associated with Quercus in western USA) and Helvella vespertina (associated with conifer forests in western USA). Therefore, we describe and illustrate Helvella jocatoi as a new species based on phylogeny and morphological traits. In central Mexico, this taxon is an edible mushroom known as "gachupín", has high cultural importance and is sold in large quantities. The description of this new species restricted to A. religiosa forests has implications for its conservation since its habitat is endangered.
... Helvella is a species-rich genus of apothecial ascomycetous fungi (Pezizomycetes: Pezizales: Helvellaceae) of terrestrial biomes of the northern and southern hemispheres. Many authors have contributed to the species-level taxonomy of the genus, the more comprehensive are those of Boudier (1905Boudier ( -1910, Nannfeldt (1932Nannfeldt ( , 1937, Dissing (1966a, b, Dissing & Nannfeldt 1966, Weber (1972Weber ( , 1975, Harmaja (1977bHarmaja ( , 1979, Häffner (1987), Abbott & Currah (1997), Vooren (2010Vooren ( , 2014, Landeros et al. (2012Landeros et al. ( , 2015, and Skrede et al. (2017). In Dissing's monograph of "The genus Helvella in Europe" (Dissing 1966a), he recognised 26 species in Europe, a list that was later expanded to 28 (Dissing 1972) and to 38 in a survey 30 yr later, but then confined to species of the Nordic countries only (Dissing 2000). ...
Phylogenetic analyses of 115 newly collected Helvella specimens from Spain using three genetic markers [heat shock protein 90 ( hsp ), RNA polymerase II second largest subunit ( rpb2 ) and the nuclear large subunit ribosomal DNA (LSU)] confirm the assignment of the Spanish collections to one Dissingia and 30 Helvella species. The analyses were supplemented with an additional sample of 65 Spanish and extralimital Helvella specimens from the fungaria of Oslo (O), Trondheim (TRH), Copenhagen (C), Uppsala (UPS), Stockholm (S) and Venice (MCVE). Nine species are described as new, i.e. Helvella fuscolacunosa , H. hispanica , H. iberica , H. inexpectata , H. neopallescens , H. phlebophoroides , H. poculiformis , H. retinervis , and H. terricola . We present photographs of a selection of fresh specimens and provide descriptions of all species of this diverse South European Mediterranean element of the genera in Europe.
... Nguyen et al. (2013) explored Helvella phylogeny using large-scale sequence analysis of LSU and the nuclear rDNA internal transcribed spacer region (ITS) and reported two new species from North America based on molecular and morphological evidence. On the basis of examinations of the type specimens and LSU sequence analysis, Landeros et al. (2012Landeros et al. ( , 2015 concluded that the sections Elasticae, Helvella, Lacunosae and Leucomelaenae were monophyletic. Skrede et al. (2017) studied molecular characteristics of 55 European species, described seven new species based on the sequence divergences of LSU, RPB2, TEF1 and heat shock protein 90 gene (Hsp90), and designated neotypes and epitypes for 30 of them. ...
Helvella species are ascomycetous macrofungi with saddle-shaped or cupulate apothecia. They are distributed worldwide and play an important ecological role as ectomycorrhizal symbionts. A recent multi-locus phylogenetic study of the genus suggested that the cupulate group of Helvella was in need of comprehensive revision. In this study, all the specimens of cupulate Helvella sensu lato with ribbed stipes deposited in HMAS were examined morphologically and molecularly. A four-locus phylogeny was reconstructed using partial sequences of the heat shock protein 90, nuclear rDNA internal transcribed spacer region 2, nuclear large subunit ribosomal DNA and translation elongation factor 1-α genes. Three clades were revealed in Helvella sensu stricto. Twenty species were included in the analysis, of which 13 are distributed in China. Three new species, H. acetabuloides, H. sichuanensis and H. tianshanensis, are described and illustrated in detail. A neotype was designated for H. taiyuanensis. Helvella calycina is a new record for China, while Dissingia leucomelaena should be excluded from Chinese mycota. Hsp90 and ITS2 are recommended as useful supplementary barcodes for species identifications of the genus.
... Within the last six years species and subgroups of Helvella s.lat. have received renewed attention ( Landeros et al. 2012, Nguyen et al. 2013, 2016b, Skrede et al. 2017). These studies have focused on species delimitation including a limited number of molecular characters and/ or morphological data. ...
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.
