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A polyphasic taxonomy of Daldinia (Xylariaceae)

April 2014
Studies in Mycology 77(1):1-143

April 2014
77(1):1-143

DOI:10.3114/sim0016

Source
PubMed

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CC BY-NC-ND 3.0

Authors:

Marc Stadler

Helmholtz Centre for Infection Research

Thomas Læssøe

University of Copenhagen

Jacques Fournier

Cony Decock

Université Catholique de Louvain - UCLouvain

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For a monograph based on a polythetic concept, several thousands of herbarium specimens, and several hundreds of freshly collected and cultured specimens of Daldinia and allied Xylariaceae, originating from around the world, were studied for morphological traits, including by SEM, and chemically by HPLC profiles using UV-visible and mass spectrometric detection. Emphasis was given to tropical material, and importantly, ancient specimens, including as many types as possible, were tracked and studied to review earlier taxonomic concepts. An epitype of D. eschscholtzii was selected as representative of the morphochemotype that is most widely distributed in the tropics. Six new species of Daldinia from the tropics and the southern Hemisphere are described. Daldinia asphalatum is resurrected, and D. cudonia is regarded as its synonym. In addition, the following binomials are epi-, iso-, neo- and/or lectotypified: Daldinia asphalatum, D. caldariorum, D. clavata, D. cuprea, D. durissima, D. eschscholtzii, D. grandis, D. loculata, and D. vernicosa. Annellosporium and Versiomyces are regarded as synonyms of Daldinia. Many new synonymies in Daldinia are proposed, and some previously published names are rejected. In total, 47 taxa in Daldinia are recognised and a key is provided. Their biogeography, chorology, and ecology, as well as the importance of their secondary metabolites, are also discussed. The previous definition of the genus is emended. The species concept is based mainly on morphological and other phenotype-derived characters because, despite diligent search, no molecular data or cultures of several of the accepted species could be obtained. Daldinia is segregated into five major groups, based on phenotypic characteristics. Some unnamed but aberrant specimens were not found in good condition and are therefore not formally described as new species. However, they are illustrated in detail in a hope that this will facilitate the discovery of fresh material in future. A preliminary molecular phylogeny based on 5.8S/ITS nrDNA including numerous representatives of all hitherto described taxa for which cultures are extant, was found basically in agreement with the above mentioned segregation of the genus, based on morphological and chemotaxonomic evidence. In the rDNA based phylogenetic tree, Daldinia appears clearly distinct from members of the genera Annulohypoxylon and Hypoxylon; nevertheless, representatives of small genera of predominantly tropical origin (Entonaema, Phylacia, Ruwenzoria, Rhopalostroma, Thamnomyces) appear to have evolved from daldinioid ancestors and are nested inside the Daldinia clade. Interestingly, these findings correlate with chemotaxonomic characters to a great extent, especially regarding the distribution of marker metabolites in their mycelial cultures. Hence, the current study revealed for the first time that fungal secondary metabolite profiles can have taxonomic value beyond the species rank and even coincide with phylogenetic data. Taxonomic novelties: Daldinia andina sp. nov., D. australis sp. nov., D. hausknechtii sp. nov., D. rehmii sp. nov., D. starbaeckii sp. nov., D. theissenii sp. nov., D. cahuchosa comb. nov., D. nemorosa comb. nov.

Representative HPLC profiles (210 nm) of stromatal methanol extracts of Daldinia concentrica and D. vernicosa and DAD spectra of characteristic metabolites. The profiles are rather characteristic of the respective groups as defined in the taxonomic parts. Whereas the profile of D. concentrica and other members of this species complex is dominated by BNT (1) and concentricol A (8), the profile of D. vernicosa mainly reveals BNT (1) and other compounds which are presumably binaphthyl derivatives.

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Teleomorphic characteristics of Daldinia andina A-J: Holotype, C-F-58309 (Ecuador). K-N. T. Læssøe AAU 59449 (Ecuador). A-D, K. Stromatal habit (K: Inserted: Stromatal pigments in 10 % KOH). E, L. Stroma in longitudinal section showing internal concentric zones and perithecial layer. F. Stromatal surface with ostioles (inserted: Stromatal pigments in 10 % KOH). H. Ascus top in Melzer's reagent revealing amyloid apical apparatus. I, M. Ascospores in SDS, revealing germ slits and dehiscent perispore (M). J, N. Ascospore in KOH, showing dehiscing perispore. Scale bars A-E, K, L = 1 cm; F, G = 1 mm; H-J, M, N = 10 μm.

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Teleomorphic characteristics of Daldinia sp. (cf. eschscholtzii) Martin 910 (Mexico). A, B. Stromatal habit. C, D. Stroma in longitudinal section showing internal concentric zones and perithecial layer. E. Stromatal surface with ostioles (inserted: Stromatal pigments in 10 % KOH). F. Ascospores in SDS. G. Ascospores in KOH, showing dehiscing perispore. Scale bars A-C = 5 mm; D, E = 1 mm; F, G = 10 μm.

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Teleomorphic characteristics of Daldinia palmensis (Holotype Ww3518: A-F) and D. raimundii (JF-08099 (France): G-L). A, G. Stromatal habit. B, H, J. Stroma in longitudinal section showing internal concentric zones and perithecial layer. C, I. Stromatal surface. I. (inserted): Stromatal pigments in 10 % KOH. D, K. Ascospores in SDS. E, L. Ascospores in KOH, showing dehiscing perispore. F. Ascospores by SEM (10.000×). Scale bars A, B, G, H = 1 cm; J = 2mm; C, I = 1 mm; D, E, K, L = 10 μm; F = 2 μm.

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Secondary metabolites from cultures of Daldinia and related Xylariaceae. Distribution: 10-13: Ubiquitous in Daldinia and immediate allies (Entonaema, Phylacia, Rhopalostroma, Ruwenzoria and Thamnomyces). 15: Green pigment of Hypoxylon fragiforme, also present in various cultures of Daldinia and in other Hypoxylon species. that release greenish pigments into the agar when fermented on solid media. 16-18: Not yet found in Daldinia, even after chromatographic separation of the crude extracts by HPLC, but present in most species of Hypoxylon, Annulohypoxylon, Pyrenomyxa and other hypoxyloid genera so far examined as major metabolites. 14: Ubiquitous in all the aforementioned genera, often as major metabolites.

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