On the evolution of the Hysteriaceae and Mytilinidiaceae (Pleosporomycetidae, Dothideomycetes, Ascomycota ) using four nuclear genes
Eric W.A. BOEHM 1, Conrad L. SCHOCH 2, Joseph W. SPATAFORA 2
1 Department of Biological Sciences, Kean University, 1000 Morris Ave., Union, New Jersey
2 Department of Botany and Plant Pathology, Oregon State University, Corvallis, Oregon
Hysterium Lophium Tree of Life web project: Hysteriaceae & Mytilinidiaceae
Subphylum Pezizomycotina
Ascomycota Class Dothideomycetes pseudoparaphyses absent pseudoparaphyses present
Classification history
Zogg (1962) Luttrell (1973) Barr (1987) Kirk et al. (2001) Eriksson (2006) Lumbsch & Huhndorf (2007) Loculoascomycetes Dothideomycetes Dothideomycetes Dothideomycetes Dothideomycetes
Dothideales Hysteriales Pleosporales Hysteriales Hysteriales Dothideomycetes orders incertae sedis Hysteriaceae Hysteriaceae Hysteriaceae Hysteriaceae Hysteriaceae Hysteriales Hysterioideae Hysteriaceae Melanommatales Dothideomycetes Pleosporomycetidae fam. incertae sedis Pleosporales Lophiaceae Lophioideae Mytilinidiaceae Mytilinidiaceae Mytilinidiaceae Mytilinidiaceae The Hysteriaceae Chevall.: The ascoma
• Fungi classified in the Hysteriaceae are defined by a specialized ascocarp, termed the hysterothecium. • Hysterothecia are dense, persistent carbonaceous structures, distinctly navicular in outline, and bear a pronounced longitudinal slit running the length of the long axis of the fruitbody.
Bar = 1 mm The Hysteriaceae Chevall.: Early images...
• Early authors provided remarkably accurate figures of hysteriaceous fungi (e.g., Bolton 1789). • Greville (1825) was the first to provide illustrations of asci and ascospores.
left: Bolton J, 1789. An history of fungusses growing about Halifax, Vol III. right: Greville RK, 1825. Scottish Cryptogamic Flora, III. Edinburgh, Scotland. The Hysteriaceae Chevall.: Peridium & hamathecium
• In vertical section, hysterothecia are globose to obovoid, typically with a thick three-layered peridium. • Hamathecium composed of persistent cellular pseudoparaphyses, often borne in a gel matrix, tips darkened or branched at maturity to form an epithecium.
Zogg, 1962 The Hysteriaceae Chevall.: Spore septation defines genera
pedicillate amerospores: Farlowiella Sacc. pigmented phragmospores: Hysterium Pers. : Fr. hyaline phragmospores: Gloniella Sacc. pigmented dictyospores: Hysterographium Corda hyaline dictyospores: Gloniopsis De Not. hyaline didymospores: Glonium Muhl. : Fr. pigmented didymospores: Actidiographium Vasilyeva
White bar = 1 mm; Black bar = 10 um The Mytilinidiaceae Kirschst.: The ascoma
• Obovoid to strongly laterally compressed, erect oyster- or bivalve shell-shaped (conchate) or hatchet-shaped (dolabrate) pseudothecia, standing on edge, with a prominent longitudinal keel or cristate apex. • Peridium: Unlike Hysteriaceae, mytilinidiaceous fungi possess a thin-walled, fragile prosenchymatous peridium. • Hamathecium: Unlike Hysteriaceae, narrow non-cellular, trabeculate pseudoparaphyses, sparse to lacking at maturity.
Ostreichnion Mytilinidion Lophium The Mytilinidiaceae Kirschst.: Spore septation defines genera
pigmented phragmospores: Mytilinidion Duby subgenus Eu Mytilinidion Lohman pigmented dictyospores: Ostreichnion Duby, Ostreola Darker pigmented didymospores: Actidium Fr. pigmented scolecospores: Mytilinidion Duby subgenus Lophiopsis Lohman hyaline scolecospores: Lophium Fr.
Ostreola
Mytilinidion Mytilinidion subgenus subgenus Ostreichnion Lophiopsis Eu Mytilinidion Actidium
Lophium
credits: Barr, 1990; Ellis & Ellis 1985; Lohman, 1934; Zogg, 1962 Morphological character states versus molecules Barr (1987) established a number of morphological character states used to differentiate the Hysteriaceae from the Mytilinidiaceae. Question: Will morphological character states prove to be phylogenetically informative in the context of sequence based phylogenies ?
Character states Hysteriaceae Mytilinidiaceae
Pseudothecium elongate navicular globoid, conchate, dolabrate Hysterium Dehiscence invaginated sulcus connivent with cristate apex (keel) Peridium thick-walled, thin-walled, multilayered prosenchymatous Centrum elongate, cylindrical elongate, laterally compressed Hamathecium persistent cellular evanescent narrow pseudoparaphyses trabeculate type Ascospore asymmetric symmetric symmetry Lophium Taxon sampling strategy: single ascospore cultures
Hysteriaceae : 35 isolates, representing five of the eight currently accepted genera. Mytilinidiaceae : 11 isolates, representing three of the currently accepted seven genera.
Hysteriaceae Chevall. Mytilinidiaceae Kirschst. * Hysterium Pers. : Fr. (6) * Mytilinidion Duby (7) Gloniella Sacc. * Ostreichnion Duby (2) * Hysterographium Corda (4) Ostreola Darker Hysterocarina Zogg Actidium Fr. * Gloniopsis De Not. (2) Quasiconcha Barr & Blackw. * Glonium Muhl. : Fr. (5) * Lophium Fr. (1) Actidiographium Vasil. Zoggium Vassil. * Farlowiella Sacc. (1) Four gene phylogeny: nuSSU, nuLSU, RPB2 & TEF1α
Contiguous data set consisted of four genes (3400+ nucleotides per taxon ).
RPB2 RPB2 5F
123 4 5 6 7891011
fRPB2 7cR
Vilgalys
526F 983FEFcf EFdf 1577F
Elongation Factor 1 α
EFjrEFir 1567R EFgr 1953R2218R
Vilgalys Hysteriaceae & Mytilinidiaceae
Hysteriaceae Multigene phylogenies provide strong support for:
• Monophyly for the Hysteriaceae. • Monophyly for the Mytilinidiaceae. Pezizomycotina • Both in the Pleosporomycetidae. Mytilinidiaceae • Both families not closely related within the subclass . The Hysteriales Lindau 1897 We propose that the Hysteriaceae be retained under the original ordinal designation, Hysteriales Lindau 1897 :
• High support (90/100) as monophyletic entity.
Pezizomycotina • Strong support for its separation from the Pleosporales. Mytilinidiaceae • Elongated hysteriaceous locule, capable of relatively indeterminate linear growth , as distinct from the strict Pleospora