Multigene Phylogeny of the Coronophorales: Morphology and New Species in the Order
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Mycologia, 102(1), 2010, pp. 185–210. DOI: 10.3852/09-043 # 2010 by The Mycological Society of America, Lawrence, KS 66044-8897 Multigene phylogeny of the Coronophorales: morphology and new species in the order George K. Mugambi1 an important character in defining the Scortechinia- Botany Department, Field Museum of Natural History, ceae, while taxa within the group show a mixture of Chicago, Illinois 60605–2496, Department of morphological characteristics of varying phylogenetic Biological Sciences, University of Illinois at Chicago, importance. The presence of smooth versus spinulose 845 W. Taylor Street (MC 066), Chicago, IL 60607, and National Museums of Kenya, Botany Department, subiculum aids in separating Tympanopsis and P.O. Box 45166, 00100, Nairobi, Kenya Scortechinia, and erumpent ascomata distinguish Cryptosphaerella species. Taxa within the Bertiaceae Sabine M. Huhndorf vary along the lines of robust, tuberculate, collapsing Botany Department, Field Museum of Natural History, Chicago, Illinois 60605–2496 ascomata and large, hyaline to pigmented, septate ascospores. Key words: Ascomycota, Coronophorales, LSU Abstract: The phylogenetic relationships within Cor- rDNA, phylogeny, quellko¨rper, rpb2, tef1 onophorales have been debated because of uncer- tainty over the taxonomic usefulness of characteristics INTRODUCTION such as quellko¨rper, number of ascospores per ascus, presence of ascospore appendages, presence of Members of the Coronophorales are common wood- subiculum and ascomatal vestiture. The phylogenetic inhabiting Ascomycete fungi with a worldwide distri- relationships are examined with DNA sequence data bution. Coronophorales is characterized by taxa with from three nuclear genes targeting 69 taxa and 130 mostly superficial ascomata, sometimes with an new sequences representing collections from Africa extensive hyphal subiculum or well developed basal and the Americas. Analyses recovered monophyletic stroma that often becomes cupulate or collapsed, and Bertiaceae, Chaetosphaerellaceae and Scortechinia- in some cases an ostiolar opening is either indistinct ceae and a paraphyletic Nitschkiaceae. A single or lacking. In many taxa a quellko¨rper is present in collection of Coronophora gregaria is included and the centrum, and this structure is unique to this Coronophoraceae is accepted. Bertiaceae is expanded group of ascomycetes. The quellko¨rper is a subcy- to include Gaillardiella, and Thaxteria is synonymized lindrical to inverted-conical structure attached to the with Bertia with a new combination, B. didyma. Three roof of the centrum, composed of thick-walled, new species of Bertia are described: B. ngongensis concentrically arranged, hyaline cells that functions from Kenya, B. orbis from Kenya and Costa Rica and by enlarging and forcibly breaking open the ascoma B. triseptata from Ecuador and Puerto Rico. Bertia and allowing ascospore release. Other important gigantospora is transferred from Nitschkia. Scortechi- characteristics of the group include Munk pores niaceae is confirmed for the quellko¨rper-bearing taxa that are found in the ascomatal wall cells, thin-walled including monotypic Biciliospora, Coronophorella, asci with a long stipe and in most cases without an Neofracchiaea, Scortechiniella and Scortechiniellopsis. apical ring and ascospore numbers often greater than Tympanopsis is reinstated for T. confertula and T. eight. uniseriata, while Scortechinia is more narrowly cir- Coronophorales are found on woody substrates in cumscribed to include S. acanthostroma and the new many parts of the world. Collections from temperate species, S. diminuspora from Ecuador. Cryptosphaer- European localities form the majority of the early ella is accepted in Scortechiniaceae including six new names in the group with some additional collections species from Kenya and Costa Rica, C. celata, C. made from North America (von Ho¨hnel 1907; costaricensis, C. cylindriformis, C. elliptica, C. globosa Fitzpatrick 1923; Nannfeldt 1975a, b). Over the years and C. malindensis. Spinulosphaeria is accepted in new species and collections have been added from Coronophorales with uncertain family placement. tropical locations such as French Guiana, India, Thenumberofascosporesintheascusisnot Singapore and western Africa (Sivanesan 1974, phylogenetically useful in distinguishing genera Nannfeldt 1975b). Until now Coronophorales have within the order. The quellko¨rper continues to be not been seriously studied in eastern Africa, although a few species were described from Congo Submitted 18 Mar 2009; accepted for publication 22 Jun 2009. and Kenya (von Arx and Mu¨ller 1954, Nannfeldt 1 Corresponding author. E-mail: [email protected] 1975b). 185 186 MYCOLOGIA TABLE I. Coronophoralean genera accepted in selected works since 1973 Subramanian and Mu¨ller and von Arx 1973, Sekar 1990, one family Nannfeldt 1975a, b, one family von Arx 1981, one family two families Coronophoraceae, Nitschkiaceae, Sordariales Coronophoraceae, Coronophoraceae, Coronophorales Coronophorales Coronophorales Acanthonitschkea Speg. Acanthonitschkea (syn. Euacanthe Theiss., Acanthonitschkea Acanthonitschkea Neotrotteria Sacc.) Bertia de Not. Bertia Bertia Biciliospora Biciliospora Petr. Coronophora (syn. Cryptosphaerella Sacc.) Biciliospora Biciliosporina Subram. & Sekar Calyculosphaeria Gaillardiella Coronophora Cryptosphaerella Coronophora Fuckel Nitschkia (syn. Biciliospora, Coronophorella Fracchiaea Euacanthe Ho¨hn., Fracchiaea, Neofracchiaea Teng, Scortechinia Sacc., Scortechiniella, Scortechiniellopsis Sivan., Tympanopsis Fracchiaea Sacc. Gaillardiella Fracchiaea Gaillardiella Pat. Lasiobertia Sivan. Gaillardiella Nitschkia G. H. Otth Nitschkia Janannfeldtia Subram. & Sekar Scortechiniella Arx & E. Mu¨ll. Spinulosphaeria Sivan. Neotrotteria Thaxteria Sacc. Sydowinula Petr. Nitschkia (syn. Thaxteria) Tympanopsis Starba¨ck Tympanopsis Schizocapnodium Fairman Scortechinia Bertiaceae, Coronophorales Bertia Spinulosphaeria Members of the Coronophorales have been treated over the taxonomic usefulness of distinguishing in their own order (Nannfeldt 1932, Mu¨ller and von characterisitics, such as quellko¨rper, number of Arx 1973, Subramanian and Sekar 1990) or in order ascospores per ascus, presence of ascospore append- Sordariales (Nannfeldt 1975a, b; Barr 1990; Hawks- ages, presence of subiculum and ascomatal vestiture. worth et al 1995) under either one or two families, Nannfeldt (1975b) accepted five genera in the Coronophoraceae and Nitschkiaceae (Nannfeldt Nitschkiaceae and also published detailed morphol- 1932; Nannfeldt 1975a, b) (TABLE I). Huhndorf et ogy, taxonomy and nomenclature of the many taxa. al (2004) used molecular data to show that the group He considered most of the morphological characters is not related to Sordariales and to demonstrate the of value in the delimitation of the taxa at specific monophyly of the Coronophorales in subclass Hypo- levels only and therefore adopted broad generic creomycetidae. They suggested the separation of the concepts, which resulted in drastic reduction in the taxa into four families (family Coronophoraceae was number of genera recognized (TABLE I). Von Arx not sampled). However their molecular study was (1981) followed the generic circumscriptions of based on limited taxon sampling and therefore did Mu¨ller and von Arx (1973) (TABLE I), adding a few not adequately address various conflicting taxonomic more taxa including Lasiobertia, which since has been treatments previously proposed for taxa within the shown to belong outside the group (Huhndorf et al group by different taxonomists. 2004). Subramanian and Sekar (1990) also adopted Narrow generic circumscriptions for taxa in the narrow generic circumscriptions, differing from Coronophorales were adopted by Mu¨ller and von Arx Nannfeldt (1975b) and chose to recognize two (1973) (TABLE I). Nannfeldt (1975a, b) and Subra- families for the taxa (TABLE I). manian and Sekar (1990) presented detailed histor- Huhndorf et al (2004) in molecular studies of ical perspectives of the Coronophorales and its Coronophorales used large subunit ribosomal nucle- components. The phylogenetic relationships within ar DNA sequence data that resulted in their segrega- group have long been debated because of uncertainty tion of four families in the group, including two new MUGAMBI AND HUHNDORF:CORONOPHORALES PHYLOGENY 187 families, Chaetosphaerellaceae for Chaetosphaerella 40 taxa from which at least two genes were available was phaeostroma (Durieu & Mont.) E. Mu¨ll. & C. Booth used in the combined analyses allowing for some missing and Crassochaeta nigrita (Berk. & Broome) Re´blova´ data. All voucher specimens are deposited in F and the and Scortechiniaceae for Scortechinia conferta Kenyan specimens also are deposited in EA. (Schwein.) Subram. & Sekar, Neofracchiaea callista DNA extraction, PCR amplification, sequencing and (Berk. & M.A. Curtis) Teng and Euacanthe foveolata sequnce alignment.—Total DNA was extracted with the (Berk. & M.A. Curtis ex Berk.) Subram. & Sekar. In Dneasy Plant Mini Kit (QIAGEN, Hilden, Germany) their analyses Bertiaceae was represented by Bertia following the instructions of the manufacturer. Phylogenet- tropicalis Huhndorf, A.N. Mill. & F.A. Fern. and B. ic analyses were conducted with partial sequences of three moriformis (Tode) de Not. and a paraphyletic Nitsch- genes: translation elongation factor 1 alpha (tef1), nuclear kiaceae by Nitschkia meniscoidea Huhndorf, A.N. Mill. ribosomal large subunit (LSU rDNA) and RNA polymerase & F.A. Fern.,