Persoonia 35, 2015: 166–229 www.ingentaconnect.com/content/nhn/pimj RESEARCH ARTICLE http://dx.doi.org/10.3767/003158515X688000 A monograph of Otidea (Pyronemataceae, Pezizomycetes) I. Olariaga1, N. Van Vooren2, M. Carbone3, K. Hansen1 Key words Abstract The easily recognised genus Otidea is subjected to numerous problems in species identification. A number of old names have undergone various interpretations, materials from different continents have not been compared and Flavoscypha misidentifications occur commonly. In this context, Otidea is monographed, based on our multiple gene phylogenies ITS assessing species boundaries and comparative morphological characters (see Hansen & Olariaga 2015). All names ITS1 minisatellites combined in or synonymised with Otidea are dealt with. Thirty-three species are treated, with full descriptions and LSU colour illustrations provided for 25 of these. Five new species are described, viz. O. borealis, O. brunneo parva, O. ore- Otideopsis gonensis, O. pseudoleporina and O. subformicarum. Otidea cantharella var. minor and O. onotica var. brevispora resinous exudates are elevated to species rank. Otideopsis kaushalii is combined in the genus Otidea. A key to the species of Otidea is given. An LSU dataset containing 167 sequences (with 44 newly generated in this study) is analysed to place collections and determine whether the named Otidea sequences in GenBank were identified correctly. Fourty-nine new ITS sequences were generated in this study. The ITS region is too variable to align across Otidea, but had low intraspecific variation and it aided in species identifications. Thirty type collections were studied, and ITS and LSU sequences 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 char- acters are important for species identification. We conclude that to distinguish closely related or morphologically similar species, a combination of additional features are needed, i.e. the shape of the paraphyses, ectal excipulum structure, types of ectal excipulum resinous exudates and their reactions in Melzer’s reagent and KOH, tomentum and basal mycelium colours and exudates. The KOH reaction of excipular resinous exudates and basal mycelium are introduced as novel taxonomic characters. Article info Received: 26 November 2013; Accepted: 1 February 2015; Published: 10 April 2015. INTRODUCTION Our multilocus phylogenies and robust hypotheses of species limits, employing genealogical concordance phylogenetic spe- Species of Otidea produce typically ear-shaped apothecia that cies recognition (GCPSR; Taylor et al. 2000), which is the basis are unique within Pyronemataceae (Pezizomycetes). The genus for the present work, are given in Hansen & Olariaga (2015). is monophyletic based on multilocus phylogenetic analyses In the present study we present an LSU rDNA phylogeny to from a few, but broadly sampled, Otidea species (Hansen place a larger number of collections for which we have been et al. 2013). Despite being distinct at the generic level, the unable to obtain multiple genes, including several sequences species identification and nomenclature of Otidea are highly from GenBank, many of which we here re-identify. controversial. A few recent typifications have been proposed (Carbone 2009, 2010a), but many names are still subjected to Taxonomic history different interpretations. Several new species were described The first valid publication of Otidea is by Bonorden (1851), in the last decades from Europe (Harmaja 1976, 2009a) and based on Peziza (unranked) Otidea Pers., although it has some- Asia (Cao et al. 1990, Zhuang & Yang 2008), with detailed times been attributed to Fuckel (Kanouse 1949, Nannfeldt descriptions and updated identification keys. However, often 1966, Liu & Zhuang 2006, Smith & Healy 2009). Otidea species no illustrations were presented and colour photographs have were treated in a broad heterogeneous genus Peziza by early rarely been published when describing new species. Many authors. Persoon (1822) defined Peziza (unranked) Otidea names of European species currently used in North America as producing auriculate apothecia with a split, sometimes and Asia are misapplied. Multilocus phylogenetic analyses have elongated on one side, and included 10 species. Fries (1822) not been previously implemented to critically address species referred to this group as Peziza (unranked) Cochleatae, but he delimitation issues and material from different continents has included also taxa with non-split apothecia. Bonorden (1851) not been compared. A worldwide critical revision of Otidea to elevated Otidea to generic rank with split apothecia as the key clarify species limits is highly needed. The aims of this study feature, but also referred to Fries’ (1822) Cochleatae. He did were: i) to undertake a nomenclatural and taxonomic revision not make any combinations or list any species. Fuckel (1870) of Otidea, to clarify misinterpretations and to propose pertinent refined the genus using microscopic details, namely uni- or typifications to stabilise the use of names; and ii) to provide biguttulate spores and filiform to subclaviform paraphyses, and detailed species descriptions and colour photographs of both included four species: O. abietina, O. cochleata O. leporina macro- and microscopic structures, and a key for identification. and O. onotica. Boudier (1885) notably contributed to disas- semble the large genus Peziza into smaller and more natural 1 Swedish Museum of Natural History, Department of Botany, P.O. Box 50007, genera. He placed in the genus Otidea species with entire or SE-104 05 Stockholm, Sweden; corresponding author e-mail: [email protected]. split apothecia, biguttulate spores and, importantly non-amyloid 2 36 rue de la Garde, F-69005 Lyon, France. asci and curved paraphyses, which he was the first to intro- 3 Via Don Luigi Sturzo 173 I-16148 Genova, Italy. duce. He divided Otidea into two subgenera: Otidea with split © 2015 Naturalis Biodiversity Center & Centraalbureau voor Schimmelcultures You are free to share - to copy, distribute and transmit the work, under the following conditions: Attribution: You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work). Non-commercial: You may not use this work for commercial purposes. No derivative works: You may not alter, transform, or build upon this work. For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights. I. Olariaga et al.: A monograph of Otidea 167 apothecia and Pseudotis with entire apothecia. Interestingly, A new genus Flavoscypha was erected for two species of Boudier erected the genus Wynnella (Helvellaceae) to accom- Otidea, O. concinna (as Flavoscypha cantharella) and O. phle- modate W. silvicola (as P. leporina / P. auricula), a species bophora, with strong emphasis on the ectal excipulum of textura with distinctly ear-shaped apothecia, but differing from Otidea prismatica (vs textura angularis in Otidea) (Harmaja 1974). in the uniguttulate spores and tough consistency. In Boudier’s Otidea was further emended to include a species with orna- (1907) subsequent treatment, he elevated Pseudotis to genus mented spores, O. unicisa, otherwise ‘fitting perfectly’ Otidea rank and placed here species with entire apothecia, including (Harmaja 1986). Otideopsis was published with Otideopsis O. da liensis (as P. apophysata) and O. propinquata (as P. abi- yunnanensis as the type species, distinguished from Otidea etina), both with biguttulate spores and hooked paraphyses. by having ornamented spores and paraphyses fused at the Saccardo listed O. onotica as an ‘exemplar’ species of Peziza apices (Liu & Cao 1987). Flavoscypha and Otideopsis are now subg. Otidea in his synopsis of the discomycete genera (1884; considered synonyms of Otidea based on molecular phyloge- netic analyses (Liu & Zhuang 2006, Hansen & Olariaga 2015). he listed in general only one species per genus / subgenus that appear to have been selected as typical for the genera) and Recently the circumscription of Otidea was further broadened it has since been accepted as the type species by most (Rifai when the first hypogeous species, O. subterranea, was discov- 1968, Eckblad 1968, Korf 1972, Liu & Zhuang 2006, Parslow & ered using ITS and LSU sequences (Smith & Healy 2009). All Spooner 2013), Index Nominum Genericorum (Eckblad in Farr the characters proposed so far as diagnostic for Otidea have ex- et al. 1979), NCU-3 (Greuter et al. 1993) and is prepared to be ceptions across the genus. Nevertheless, Otidea can be recog- adopted on the List of Protected generic Names for fungi (Kirk nised by the non-amyloid asci, in combination with at least et al. 2013). Clements & Shear (1931) listed O. cochleata as two of these characters (except O. subterranea): a) biguttulate spores; b) hooked or bent paraphyses; c) medium-large, split the type, but this name was not among the original species ac- apothecia; and d) a medullary excipulum of textura intricata, and cepted by Persoon (1822) and has furthermore been considered an ectal excipulum of textura angularis