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Notes sur le genre Helvella L. (Ascomycota, Pezizales). 2. Les sous-genres Cupuliformes et Macropodes

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Abstract and Figures

The author proposes a taxonomical synthesis of the genus Helvella, subgenera Cupuliformes and Macropodes, containing the taxa with a cylindrical stipe, neither lacunose nor costate, and with a cupuliform cap. All the taxa described in this note are illustrated with colour photographs. Ancient illustrations are also used to complete the iconography
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Notes sur le genre Helvella L. (Ascomycota,
Pezizales)
2. Les sous-genres Cupuliformes et Macropodes
Nicolas VAN VOOREN
1
Résumé
Lauteur propose une synthèse taxinomique du genre Helvella, sous-genres Cupuliformes et Macropodes, comprenant
les taxons à pied cylindracé, non lacuneux ou côtelé, et à chapeau cupuliforme. Tous les taxons retenus dans cette
note sont illustrés par des photographies en couleur. D’anciennes illustrations viennent également compléter lico-
nographie.
Abstract
The author proposes a taxonomical synthesis of the genus Helvella, subgenera Cupuliformes and Macropodes, contai-
ning the taxa with a cylindrical stipe, neither lacunose nor costate, and with a cupuliform cap. All the taxa described
in this note are illustrated with colour photographs. Ancient illustrations are also used to complete the iconography.
Mots-clés
Macropodia, Cyathipodia, taxinomie.
Introduction


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





  
󰮏




Systématique
 



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


1
36, rue de la Garde, F-69005 Lyon – nicolas@vanvooren.info
Bull. mycol. bot. Dauphiné-Savoie, 212, p. 29-47 (2014)
DESCRIPTION
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 
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
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



















󰮏

  



30
Fig. 1






31
Matériel et méthode



󰮏




Descriptions


Helvella corium














































 













 
  








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

󰮐







 



 

  





Helvella cupuliformis












   
  ć 

 
34






























 








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  
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          
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 
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 


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󰮏
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
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
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35
 
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󰮏



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

 
Helvella fibrosa


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
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
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

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
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



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



󰮏

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





 




Helvella macrosperma

č





















37



 













  
 
      











Helvella rivularis







 

 
38






















󰮏


          







 










  







Helvella macropus








































 







39

   







   












 











 




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








     
  


 










󰮏







 





















 












 

43
Helvella macropus


alba

















Helvella terrestris













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
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... macrosperma (Favre 1955) from the Swiss Alps. The latter variety was later given specific rank by Fellner & Landa (1991), as has been followed by subsequent authors (Van Vooren 2014, 2015, Skrede et al. 2017. Contrary, in his monumental work on Helvella, Dissing (1966b) included H. alpestris, H. arctica and H. arctica var. ...
... The ascus development, whether aporhynchous or pleurorhynchous (Chadefaud 1943, Berthet 1964, is considered valuable for defining lineages and sections of Helvella (Weber 1972, Landeros et al. 2015, and even genera of the family Helvellaceae (Hansen et al. 2019). Aporhynchous asci seem important to delineate the /alpinacorium lineage (Häffner 1987), while pleurorhynchous asci characterise the /alpestris-nannfeldtii lineage (Van Vooren 2014, 2015, Skrede et al. 2017. Indeed, aporhynchous asci represent a synapomorphy for the /alpina-corium lineage (Fig. 1). ...
Article
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Mycologists have always been curious about the elaborate morphotypes and shapes of species of the genus Helvella . The small, black, cupulate Helvella specimens have mostly been assigned to Helvella corium , a broadly defined morpho-species. Recent phylogenetic analyses, however, have revealed an aggregate of species hidden under this name. We performed a multispecies coalescent analysis to re-assess species limits and evolutionary relationships of the Helvella corium species aggregate in the Nordic countries. To achieve this, we used morphology and phylogenetic evidence from five loci – heat shock protein 90 ( hsp ), translation elongation factor 1-alpha ( tef ), RNA polymerase II ( rpb2 ), and the 5.8S and large subunit (LSU) of the nuclear ribosomal DNA. All specimens under the name Helvella corium in the larger university fungaria of Norway, Sweden and Denmark were examined and barcoded, using partial hsp and/or rpb2 as the preferential secondary barcodes in Helvella . Additional fresh specimens were collected in three years (2015–2018) to obtain in vivo morphological data to aid in species discrimination. The H. corium species aggregate consists of seven phylogenetically distinct species, nested in three divergent lineages, i.e. H. corium , H. alpina and H. pseudoalpina sp. nov. in the /alpina-corium lineage, H. alpestris , H. macrosperma and H. nannfeldtii in the /alpestris-nannfeldtii lineage, and H. alpicola as a weakly supported sister to the /alpestris-nannfeldtii lineage. Among the seven species, the ribosomal loci expressed substantial variation in evolutionary rates, suggesting care in the use of these regions alone in delimitation of Helvella species. Altogether, 469 out of 496 available fungarium specimens were successfully barcoded.
... Helvella semiobruta has a single record from Gotland, Sweden, a distribution assumed to be governed by Editor-in-Chief Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst E-mail: p.crous@westerdijkinstitute.nl local climatic and ecological features on this island of the Baltic Sea. This species, which is common in the Mediterranean region (Filippa et al. 2013 (Dissing & Sivertsen 1980, Peric 2011, Van Vooren 2014. Eleven species are exclusively found in the arctic-alpine biome of Norway and Sweden, and partially in Greenland, Iceland and Svalbard, i.e., H. alpestris, H. alpina, H. arctoalpina, H. bresadolae, H. capucina, H. dryadophila, H. macrosperma, H. nordlandica, H. oroarctica, H. philonotis, H. pseudoalpina. ...
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Helvella is a species-rich genus, forming a large variation of astounding ascocarps in many different habitats. During the last decade, molecular markers and morphological characters have been combined to delimit and identify cryptic species in this genus. We report on a list of 54 species of Helvella s.s. in the Nordic region and describe five new species, i.e. H. bresadolae, H. convexa, H. japonica, H. nordlandica and H. oroarctica. The morphological and molecular characteristics of the new species and the emended / hypocrateriformis, / fibrosa-macropus, and / fallax�pezizoides lineages of Helvella s.s. are shortly commented upon. Further we include a discussion of the distribution of species in the Nordic region based on a large set of studied collections. The ecological versatility and variable geographic patterns of these species indicate that cryptic species may have contrasting ecology in their local habitats.
... Helvella L. is proposed by Linnaeus (1753). It's the most recognizable mushroom, which has been described and illustrated repeatedly by mycologists worldwide (Dissing 1966a, 1966b, Weber 1972, Harmaja 1979, Abbott & Currah 1997, Zhuang 2004, Nguyen et al. 2013, Vooren 2014, Zhao et al. 2015, Hansen et al. 2019, Løken et al. 2020. Some Helvella species are edible and popular as food all over the world (e.g., H. acetabulum L. Quél, H. bachu Q. Zhao, Zhu L. Yang & K.D. Hyde, H. crispa Bull, H. elastica Bull, H. lacunosa Afzel, H. subspadicea Q. Zhao, Zhu L. Yang & K.D. Hyde and H. taiyuanensis B. Liu, Du & J.Z. Cao (Liu et al. 1985, 1988, Zhao et al. 2015, Wang et al. 2016b. ...
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During investigations into fungal diversity in the Tibetan Plateau, two Helvella collections were made. Morphological characteristics and phylogenetic analyses of combined ITS and LSU sequence data revealed one collection represents a distinct new species that is introduced as Helvella cystidiata. Helvella cystidiata is characterized by having loosely saddle-shaped pileus, greyish to grey brown hymenium, lacunose stipe and inflated ectal excipulum cells. Furthermore, a known species Helvella lacunosa is reported in Tibetan Plateau. Detailed descriptions, illustrations and a phylogenetic tree to show the placement of two taxa are provided.
... 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). ...
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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.
... The large, narrowly fusiform spores with multiple guttules are unique within Helvel laceae. It has been suggested that the closest relatives of Pindara are to be found among the Helvella subgenera Cupuli formae or Macropodes sensu Abbott & Currah 1997(Landvik et al. 1999, Van Vooren 2014 based on the stipitate-cupulate ascomata without external ribs and grooves (Fig. 7a -c), and the verruculose, fusoid ascospores (Fig. 7i), similar to characters observed in H. macropus (Fig. 4d, 5b). Helvella rivularis was also compared to Pindara because of its small ascoma size (3 -9 mm). ...
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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.
Article
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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.
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In the present study, current information based on morphological characters and DNA sequence (nrITS) data is given on some uncommon or noteworthy Ascomycetes species collected from different geographic regions of Turkey. First phylogenetic analysis based on nrITS data of Helvella fibrosa and H. macropus the rare and little-known species was performed in Turkey by this study. Also, the nrITS sequence of H. fibrosa was uploaded to current databases for the first time. Detailed morphological description, macro photographs and detailed drawings of micro structures of the two species studied have been presented, and the ecology and distributions of each species has been given.
Article
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Helvella is a widespread, speciose genus of large apothecial ascomycetes (Pezizomycete:Pezizales) that are found in terrestrial biomes of the Northern and Southern Hemispheres. This study represents a beginning on assessing species limits and applying correct names forHelvellaspecies based on type material and specimens in the university herbaria (fungaria) of Copenhagen (C), Harvard (FH) and Oslo (O). We use morphology and phylogenetic evidence from four loci - heat shock protein 90 (hsp), translation elongation factor alpha (tef), RNA polymerase II (rpb2) and the nuclear large subunit ribosomal DNA (LSU) - to assess species boundaries in an expanded sample ofHelvellaspecimens from Europe. We combine the morphological and phylogenetic information from 55Helvellaspecies from Europe with a small sample ofHelvellaspecies from other regions of the world. Little intraspecific variation was detected within the species using these molecular markers;hspandrpb2markers provided useful barcodes for species delimitation in this genus, while LSU provided more variable resolution among the pertinent species. We discuss typification issues and identify molecular characteristics for 55 EuropeanHelvellaspecies, designate neo- and epitypes for 30 species, and describe sevenHelvellaspecies new to science, i.e.,H. alpicola,H. alpina,H. carnosa,H. danica,H. nannfeldtii,H. pubescensandH. scyphoides.
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During the annual foray 2014 of the FMBDS, organized in the National Park of Vanoise (Savoy, France), several pezizalean species have been collected and studied. Some of them are registered for the first time in the Rhône-Alpes region such as Helvella macrosperma and Iodophanus hyperboreus. Lamprospora leptodictya, only known from a single collection in this national park, has been found again. They are presented and illustrated in this note
Book
The Rhône-Alpes region, by its surface (the second French region) and the diversity of its natural environments, offers an interesting territory for the study of ascomycetes. This work provides an overview of the diversity of discomycetes belonging to the order of Pezizales, in this region, through the detailed presentation and illustration of many species. This first part includes 94 species
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Discina parma J. Breinteb. & Mass Gest. est une espèces nuvelle pour la flore mycologique du Monténégro. Dans cet article les auteurs proposent une description macroscopique, des photographies prises in situ et en laboratoire, des photos et dessins d’éléments de la microscopie et des commentaires écologiques et taxonomiques.
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Two Helvella species, Helvella macropus (Pers.) P. Karst and H. pezizoides Afzel. are given as new records for Turkey. Short descriptions and photographs related to macro and micromorphologies of the species are provided and discussed briefly.
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Monographie der Gattung Helvella
Article
Octospora villosa was described and illustrated by Hedwig in 1789, and has been reassigned first as a variety in Peziza, and later at species rank in Fuckelina, Cyathipodia, and Helvella. Hedwig's species was sanctioned (at varietal level) by Fries, who treated it as Peziza macropus [var.] β hirta, an unintentional error in attributing to this taxon the epithet of a different Hedwig species. Error upon error has plagued Hedwig's species name, and an attempt here is made to untangle the most flagrant of these. Names in current use are Helvella villosa, H. chinensis, and H. dissingii. None turns out to be correct under the International Code of Botanical Nomenclature. Overlooked by all recent authors is the species name Peziza fibrosa proposed by Wallroth in 1833 as a new name to be substituted for O. villosa since that epithet could not be transferred to the genus Peziza, it being preoccupied there by Persoon's P. villosa of 1801. The new combination Helvella fibrosa is formally proposed, and an epitype specimen for O. villosa is designated.
Article
Epigeous genera of the Helvellaceae (Pezizales, Ascomycota) are separated into tribes Helvelleae and Gyromitreae on the basis of excipulum structure, while hypogeous genera are included as incertae sedis at the tribe level. Ornamentation of ascospores (SEM) is valuable in the delimitation of genera, subgenera, and species. New subgenera proposed are Gyromitra subgenus Melaleucoides subgen. nov., with warted, nonapiculate, biguttulate ascospores; Gyromitra subgenus Caroliniana subgen. nov., with coarsely reticulate ascospores with multiple spicules at the poles; Helvella subgenus Cupuliformae subgen. nov., with regularly cupulate apothecia, solid terete stipes, and broadly ellipsoidal, finely rugose ascospores; and Hydnotrya subgenus Cerebriformae subgen. nov., with globose, echinate ascospores. Gyromitra subgenus Discina is emended. New combinations include Helvella subgenus Leucomelaenae comb. nov, Helvella subgenus Silvicolae comb. nov., Helvella subgenus Macropodes comb. nov., and Helvella subgenus Elasticae comb. nov. Epigeous taxa and hypogeous taxa with chambered ascocarps were studied in detail. Collections of 44 species in six genera (Gyromitra, Rhizina, Pseudorhizina, Helvella, Underwoodia, and Hydnotrya) from northern and northwestern North America are described. One extralimital genus (Gymnohydnotrya) and 15 species outside the study area are included with systematic notes. The known geographic range for many species is extended and there are several new records for North America (Helvella rivularis, H. verruculosa, H. unicolor). Maps and keys for taxa in northern and northwestern North America are provided.