James D. Lawrey

Families of Dothideomycetes

Dothideomycetes comprise a highly diverse range of fungi characterized mainly by asci with two wall layers (bitunicate asci) and often with fissitunicate dehiscence. Many species are saprobes, with many asexual states comprising important plant pathogens. They are also endophytes, epiphytes, fungicolous, lichenized, or lichenicolous fungi. They occur in terrestrial, freshwater and marine habitats in almost every part of the world. We accept 105 families in Dothideomycetes with the new families Anteagloniaceae, Bambusicolaceae, Biatriosporaceae, Lichenoconiaceae, Muyocopronaceae, Paranectriellaceae, Roussoellaceae, Salsugineaceae, Seynesiopeltidaceae and Thyridariaceae introduced in this paper. Each family is provided with a description and notes, including asexual and asexual states, and if more than one genus is included, the type genus is also characterized. Each family is provided with at least one figure-plate, usually illustrating the type genus, a list of accepted genera, including asexual genera, and a key to these genera. A phylogenetic tree based on four gene combined analysis add support for 64 of the families and 22 orders, including the novel orders, Dyfrolomycetales, Lichenoconiales, Lichenotheliales, Monoblastiales, Natipusillales, Phaeotrichales and Strigulales. The paper is expected to provide a working document on Dothideomycetes which can be modified as new data comes to light. It is hoped that by illustrating types we provide stimulation and interest so that more work is carried out in this remarkable group of fungi.

Biological Role of Lichen Substances

The bizarre phenolic substances produced by many lichen-formingfungi are assumed to have important biological roles, an idea that has led various investigators to speculate on topics ranging from light-screening to rock mineralization. At the present time, three proposed roles- antimicrobial, allelopathic and antiherbivore-are best supported by the available evidence. Anti- microbial: The fact that lichens grow slowly and yet are resistant to decay microorganisms suggests that lichen substances, known to be clinically effective antibiotics, also have a protective role in nature. Indeed, this may be their principal role. However, there is little direct evidence from field studies. Allelopathic: Recent studies show that lichen substances can inhibit seed germination of vascular plants and spore germination of mosses and lichens, demonstrating a possible allelopathic role for these substances. The ecological significance of such inhibitory effects in natural species assemblages that include lichens, however, requires more research. Antiherbivore: Despite their availabilty as food to generalist herbivores, lichens are seldom eaten, suggesting that lichen sub- stances deter feeding. Recent research, done primarily with invertebrate consumers, indicates that some lichen compounds are capable of defending lichen thalli from predation. The secondary chemicals produced by lichens have attracted the attention of investigators for over 100 years, and information about their structure, bio- genic origin and phylogenetic significance has ac- cumulated steadily over this period. Indeed, the sec- ondary chemistry of few other groups of organisms is better known. Yet our understanding of their pos- sible biological roles is meager. This is unfortunate because the extent to which lichen secondary sub- stances can be expected to contribute to the solution of evolutionary problems depends on a better un- derstanding of their adaptive value. What do these compounds do? Most lichenolo- gists assume that they do something, since it is dif- ficult to imagine how mechanisms resulting in the production of such a wide array of energetically ex- pensive compounds could have evolved in so many groups of energetically-depauperate lichen-forming fungi. If these substances have no adaptive value at all, their relatively high energetic cost would ulti- mately lead to their disappearance over evolution- ary time. Just the opposite appears to be happening, however. In the most rapidly speciating lichen

Lichenicolous Fungi: Interactions, Evolution, and Biodiversity

Abstract The lichenicolous fungi represent an important ecological group of species that form obligate associations with lichens. They have been studied seriously for over 200 years and the European species are especially well collected. However, collecting in other areas has been far less systematic and many new species await discovery. North American species are especially under-studied and this review is intended to stimulate the collection and study of these species. To encourage this study by lichenologists, we review the sorts of interactions formed by lichenicolous fungi with their lichen hosts, discuss various aspects of host specificity, virulence, chemical ecology, and evolution of lichenicolous fungi and provide a complete taxonomic listing of lichenicolous genera arranged as far as possible into natural groupings. In each section we suggest research topics in need of further study, and provide a listing of significant literature. We hope by calling attention to the largely unexplored biodiversity of lichenicolous fungi, investigators will take up the study of these fascinating organisms.

Naming and outline of Dothideomycetes-2014 including proposals for the protection or suppression of generic names

Article 59.1, of the International Code of Nomenclature for Algae, Fungi, and Plants (ICN; Melbourne Code), which addresses the nomenclature of pleomorphic fungi, became effective from 30 July 2011. Since that date, each fungal species can have one nomenclaturally correct name in a particular classification. All other previously used names for this species will be considered as synonyms. The older generic epithet takes priority over the younger name. Any widely used younger names proposed for use, must comply with Art. 57.2 and their usage should be approved by the Nomenclature Committee for Fungi (NCF). In this paper, we list all genera currently accepted by us in Dothideomycetes (belonging to 23 orders and 110 families), including pleomorphic and non-pleomorphic genera. In the case of pleomorphic genera, we follow the rulings of the current ICN and propose single generic names for future usage. The taxonomic placements of 1261 genera are listed as an outline. Protected names and suppressed names for 34 pleomorphic genera are listed separately. Notes and justifications are provided for possible proposed names after the list of genera. Notes are also provided on recent advances in our understanding of asexual and sexual morph linkages in Dothideomycetes. A phylogenetic tree based on four gene analyses supported 23 orders and 75 families, while 35 families still lack molecular data.

Lichen Secondary Compounds: Evidence for a Correspondence Between Antiherbivore and Antimicrobial Function

Laboratory tests were done to determine the antimicrobial activity of lichen secondary substances known to influence the feeding behavior of the lichen herbivore, Pallifera varia. Antibiotic test disks were prepared using acetone extracts of two lichens preferred (Aspicilia gibbosa and Lasallia papulosa) and two lichens avoided (Flavoparmelia baltimorensis and Xanthoparmelia cumberlandia) by the herbivore. These disks were placed on nutrient agar plates inoculated with various bacteria. The lichen extracts consistently inhibited gram-positive bacteria used in the assays, and the order ofeffectiveness was the same as it had been against the herbivore. Pure lichen substances were also found to vary in effectiveness in these assays. These results suggest that lichen secondary substances can defend lichens against attack by both herbivores and microorganisms; however, they are apparently not designed to defend specifically against either.

A single macrolichen constitutes hundreds of unrecognized species

Significance Macrolichens are considered to be well known, including the tropical montane basidiolichen fungus Dictyonema glabratum, also known as Cora pavonia, an important component of threatened paramo ecosystems, where it acts as a biological fertilizer due to its ability to fix atmospheric nitrogen. This lichen was long believed to represent a single species, but after revising this number to 16 in two genera (Cora and Corella), here we show that at least 126 phylogenetically and morphologically distinct species are contained within this group, with statistical analysis predicting more than 400. Our findings underline the importance of accurately documenting species richness for conservation purposes and support the notion of neotropical paramos as hotspots of recent diversification in plants, animals, and fungi. The number of Fungi is estimated at between 1.5 and 3 million. Lichenized species are thought to make up a comparatively small portion of this figure, with unrecognized species richness hidden among little-studied, tropical microlichens. Recent findings, however, suggest that some macrolichens contain a large number of unrecognized taxa, increasing known species richness by an order of magnitude or more. Here we report the existence of at least 126 species in what until recently was believed to be a single taxon: the basidiolichen fungus Dictyonema glabratum, also known as Cora pavonia. Notably, these species are not cryptic but morphologically distinct. A predictive model suggests an even larger number, with more than 400 species. These results call into question species concepts in presumably well-known macrolichens and demonstrate the need for accurately documenting such species richness, given the importance of these lichens in endangered ecosystems such as paramos and the alarming potential for species losses throughout the tropics.

High concentration of basidiolichens in a single family of agaricoid mushrooms (Basidiomycota: Agaricales: Hygrophoraceae).

The Agaricales is the largest and most diverse order of mushroom-forming Basidiomycota, with over 100 natural groups recognized in recent Fungal Tree of Life studies. Most agarics are either saprotrophic or ectomycorrhizal fungi, but the family Hygrophoraceae is in part characterized by a unique and remarkable diversity of lichenized forms. The most familiar of these is the chlorolichen genus Lichenomphalia, whose phylogenetic position in the Agaricales has been established. Recent limited evidence suggested that Hygrophoraceae also contains cyanolichens in the genus Dictyonema, which indicates a remarkable concentration and diversity of lichen-formers in a single family of agarics. To demonstrate the relationships of lichen-formers to other fungi in the family, we assembled ribosomal sequences from 52 species representing recognized groups within the Hygrophoraceae, among them new sequences representing Acantholichen and most species and forms of Dictyonema. The molecular data were evaluated using parsimony, likelihood, Bayesian, and distance analyses, including coding of ambiguous regions by means of INAASE and ARC, all of which indicate that Dictyonema and Acantholichen form a monophyletic clade derived from the primarily bryophilous genus Arrhenia and sister to the enigmatic Athelia pyriformis, a species unrelated to the Atheliales for which we are proposing a new genus name Eonema. The chlorolichen genus Lichenomphalia may be polyphyletic. Fungi in the Dictyonema-Acantholichen clade are typically tropical, entirely lichenized, and associate with cyanobacterial photobionts. Our data indicate a transition from agaricoid-omphalinoid basidiomes observed in Arrhenia to stereoid-corticioid forms in Dictyonema, and also support a previous suggestion of a connection between loss of clamp connections and lichenization. The diverse basidiome and thallus morphologies and nutritional ecologies of these fungi indicate a remarkable evolutionary flexibility that appears to have developed in part as a consequence of symbiosis.

Correlations Between Lichen Secondary Chemistry and Grazing Activity by Pallifera varia

Individuals of the slug Pallifera varia were observed under field conditions in a talus slope habitat in Shenandoah National Park during late sum- mer, 1979. Slugs were most frequently observed feeding on the crustose lichen Aspicilia gibbosa. A quantitative analysis of saxicolous cryptogam coverage and frequency in the study area demonstrated that Aspicilia spp. and several other lichen species exhibited relatively high importance values in the community. Thin- layer chromatography of fecal material obtained from field-collected slugs was done to ascertain the feeding preferences of slugs under field conditions. Aspicilin, a lichen secondary compound produced by Aspicilia spp., was extracted from slug fecal material most frequently. Laboratory experiments designed to determine those lichen species that were preferred by slugs established a preference gradient with Aspicilia spp. most preferred and Xanthoparmelia cumberlandia and Huilia albocaerulescens actively avoided. Slugs apparently do not feed on those lichen species most frequently encountered in the community. Rather, they appear to make food choices that are based at least in part on lichen chemistry. Stictic acid and protocetraric acid appear to function as anti-herbivore compounds in observed slug-lichen interactions. Despite the enormous amount of published information available on the biosynthesis, structure and taxonomic significance of lichen secondary compounds, the biological role of these compounds is virtually unknown. Some early lichen chemical investigations have suggested that lichen compounds serve to protect slow-growing lichen thalli from predation (Bachman 1890, Zukal 1895, Stahl 1904). There are a number of anecdotal reports of lichen avoidance by herbivores that would tend to support this suggestion (reviews by Gerson & Seaward 1977 and Rundel 1978). The actual protective mechanisms are not known. However, it has been suggested that reduced palatability, direct toxicity, or indirect effects through antibiosis of gut microflora may all be involved. If one considers the known chemical associations between invertebrate herbivores and lichens (Gerson & Seaward 1977), it appears that lichen material is ingested by many herbivores without regard to chemistry. However, in most studies, accurate information on the chemistry of the lichen material ingested is not available. Controlled experiments are usually not designed to determine if food choices by herbivores are made on the basis of chemistry. Because there are few experimental field studies on lichen-chemical inter- actions with naturally occurring herbivores, it is difficult to formulate general concepts of the evolutionary significance. This paper reports on food preferences of the lichen-grazing slug Pallifera varia under field and laboratory conditions, and the role that lichen secondary compounds play in determining observed preferences.

Briancoppinsia, a new coelomycetous genus of Arthoniaceae (Arthoniales) for the lichenicolous Phoma cytospora, with a key to this and similar taxa

Morphological, anatomical, chemical and molecular data suggest that a relatively common lichenicolous coelomycete on Lecanora conizaeoides is conspecific with Phoma cytospora, previously known only from parmelioid lichens, and that further populations on Cladonia and Pertusaria belong to the same species. This species is distinguished from Phoma by several taxonomically important characters and obviously represents a previously unrecognized genus, for which the name Briancoppinsia is introduced. Phylogenetic analyses using nuLSU and mtSSU sequences of isolates obtained in pure culture suggest that the new genus belongs to the Arthoniaceae (Arthoniales). This is the first obligate lichenicolous, non-lichenized anamorph confirmed to belong to the Arthoniales based on molecular data.

New lichenicolous, muscicolous, corticolous and lignicolous taxa of Burgoa s. l. and Marchandiomyces s. l. (anamorphic Basidiomycota), a new genus for Omphalina foliacea, and a catalogue and a key to the non-lichenized, bulbilliferous basidiomycetes

A catalogue and a key to the non-lichenized, bulbilliferous basidiomycetes are given. The new genus Burgella is described for the lichenicolous B. flavoparmeliae, phylogenetically close to Sistotrema oblongisporum and Multiclavula. The genera Pneumatospora and Tricellulortus are placed in synonymy of Minimedusa, the new combination M. obcoronata is proposed, and the new facultative lichenicolous M. pubescens is described. The new facultative lichenicolous Burgoa angulosa is phylogenetically close to the generic type B. verzuoliana, whilst the new B. moriformis and B. splendens are provisionally included in the genus Burgoa. A Burgoa-like species in the Ceratobasidiaceae is left unnamed. Two new species of Marchandiomyces, M. buckii and M. nothofagicola, are described. As Marchandiomyces aurantiacus is phylogenetically more close to Erythricium than to Marchandiomyces, it is proposed to exclude it from that genus and to use the holomorphic generic name Marchandiobasidium for both anamorph and teleomorph of this species. The new genus Marchandiomphalina is introduced for the lichenized Omphalinafoliacea, a taxon phylogenetically close to Marchandiobasidium.