| HOME | HELP | FEEDBACK | SUBSCRIPTIONS | ARCHIVE | SEARCH | TABLE OF CONTENTS |
|
|
|
|||||||
|
|||||||||
|
(American Journal of Botany. 2008;95:816-823.) doi: 10.3732/ajb.0800078 © 2008 Botanical Society of America, Inc. |
What's this? |
Mycology |
2 Department of Environmental Science and Policy, George Mason University, 4400 University Drive, Fairfax, Virginia 22030-4444 USA 3 Musée national d’histoire naturelle, 25 rue Munster, L-2160 Luxembourg, Luxembourg
Received for publication 27 February 2008. Accepted for publication 7 May 2008.
ABSTRACT
Fungi in the basidiomycete order Corticiales are remarkably diverse nutritionally, including a variety of saprotrophs, plant and fungal pathogens, and lichen-forming fungi. Tracing the origin of this diversity depends on a clearer understanding of the phylogenetic relationships of fungi in the order. One of its core members is the genus Marchandiomyces, originally established for lichen pathogens that form orange or coral bulbils. We describe here a new species in the genus, M. marsonii sp. nov., which is unusual in its appearance, habit, and geographic provenance. It is foliicolous on leaves of Pandanus (screw pines, Pandanaceae) and produces flattened, coral bulbils resembling apothecia of the ascomycete genus Orbilia. It is also the first member of the genus to be collected from Australia. An isolate of the new fungus and several additional cultures of related plant pathogenic fungi were obtained and investigated phylogenetically using parsimony, likelihood, and Bayesian analyses of nuclear small and large subunit ribosomal sequences. Our phylogeny makes clear that Marchandiomyces species and their close relatives contribute significantly to the ecological diversity of the Corticiales and that this diversity is derived mainly from lignicolous ancestors.
Key Words: basidiomycetes • bulbilliferous fungi • Corticiales • foliicolous • Laetisaria • lichenicolous fungi • Limonomyces • Marchandiomyces • rDNA sequence analyses
The largest and most diverse class in the phylum Basidiomycota is the Agaricomycetes, which includes a variety of mushroom-formers and crustlike resupinate forms with modes of nutrition ranging from saprotrophs to pathogens and mutualists. Recent molecular phylogenetic analyses have begun to clarify the composition of the major clades of Agaricomycetes as well as the evolutionary development of its ecological and morphological diversity (Hibbett et al., 2000
; Hibbett and Thorn, 2001; Lim, 2001; Langer, 2002; Hibbett and Binder, 2002; Larsson et al., 2004; Binder et al., 2005; Hibbett, 2006; Matheny et al., 2006, 2007; Larsson, 2007).
One of the clades of the Agaricomycetes consistently resolved in molecular phylogenies is now recognized formally as the Corticiales K.-H. Larss. (Hibbett et al., 2007), including a single family Corticiaeae Herter (Larsson, 2007). This order is almost entirely composed of resupinate species and characterized by smooth hymenophores, a monomitic hyphal system with clamps, and smooth basidiospores with pink walls, all characteristics of the sexual, basidiospore-producing forms of the fungi (teleomorphs). Asexual forms (anamorphs), named separately by convention in mycology, have only recently been recognized in the Corticiales and assigned with certainty to the group. Members of the order have a wide range of nutritional ecologies, including mutualistic and pathogenic forms as well as lignicolous saprobes (Diederich et al., 2003; Binder et al., 2005; DePriest et al., 2005; Lawrey et al., 2007).
One of the important core genera in the Corticiales is Marchandiomyces, an anamorphic genus originally established by Diederich (1990) for the coral lichen pathogen Marchandiomyces corallinus (Roberge) Diederich & D. Hawksw., commonly collected in the eastern United States and Europe. An orange, primarily European lichen pathogen [M. aurantiacus (Lasch) Diederich & Etayo] was later added to the genus by Etayo and Diederich (1996). Both species produce bulbils—small, round masses of tightly coiled hyphae that function as resting or dispersal structures (Clémençon, 2004). The relations of these asexual anamorphs to sexual teleomorphic forms were uncertain, although both species were assumed to be basidiomycetous (Hawksworth, 1979). This was confirmed unequivocally by the molecular phylogenetic study of Sikaroodi et al. (2001) and the discovery of the teleomorph of M. aurantiacus (named Marchandiobasidium aurantiacum Diederich & Schultheis by Diederich et al. [2003]).
Later molecular studies (DePriest et al., 2005; Lawrey et al., 2007) made clear that Marchandiomyces species and their close relatives are not always associated with lichens. Several species in the genus are lignicolous (M. lignicola Lawrey & Diederich and M. nothofagicola Diederich & Lawrey), a common and apparently basal habit in the Corticiales (DePriest et al., 2005). Even more surprising was the discovery that the basidiolichen formerly called Omphalina folicacea P.M.Jørg., originally assumed to be an omphalinoid agaric but subsequently placed in the Hymenochaetales (Palice et al., 2005), is instead a member of the Corticiales closely related to Marchandiobasidium aurantiacum, renamed Marchandiomphalina foliacea (P.M.Jørg) Diederich, Lawrey & Binder (Diederich and Lawrey, 2007).
The wide range of nutritional modes observed in Marchandiomyces-like species, combined with the close affinities they have with numerous plant pathogens [Laetisaria fuciformis (McAlpine) Burds., Waitea circinata Warcup & P.H.B. Talbot] and saprobes (Corticium roseum Pers.), suggests an unusually high level of evolutionary flexibility in these fungi (DePriest et al., 2005; Lawrey et al., 2007). Understanding the origin of this diversity will require clarification of the phylogenetic relationships of the fungi in the order.
We recently had the opportunity to study a foliicolous fungus from Australia with bulbils similar in color to those of Marchandiomyces corallinus, but differing in size and shape. We were able to isolate the fungus and obtain from it nuclear small (nuc-SSU) and large (nuc-LSU) subunit rDNA sequences, which we analyzed to determine its phylogenetic position and possible relationship to Marchandiomyces. In addition to the unknown culture, we obtained cultures of three plant pathogens representing genera closely related to Marchandiomyces, and their DNA was also sequenced. Preliminary analysis made use of a core data set from two recent comprehensive phylogenetic studies of the Agaricomycetes (Binder et al., 2005; Lawrey et al., 2007). A second data set extensively sampled the Corticiales, using 36 terminals. Our objectives were (1) to determine the phylogenetic position of the unknown Marchandiomyces-like fungus in relation to existing species (2) and to consider possible nutritional transitions that have taken place in lineages containing these fungi.
MATERIALS AND METHODS
Specimens collected and anatomical methods
Specimens of the Marchandiomyces-like unknown were collected by Guy Marson from dead, hanging leaves of Pandanus oblatus H.St.John (screw pine, Pandanaceae) near Cairns, Australia.
Dry herbarium specimens were examined and measured using a binocular microscope Leica (Wetzlar, Germany) MZ 7.5 (magnification up to 50x) and photographed using a Nikon (Melville, New York USA) Coolpix 4500. Entire unsectioned and sectioned bulbils were studied in water, KOH, or lactophenol cotton blue either with or without pressure on the coverslip. Microscopic photographs of bulbils were prepared using a Zeiss (Düsseldorf, Germany) Photomikroskop III with a Canon (Lake Success, New York, USA) PowerShot G5.
Isolation of fungal culture
A culture of the Marchandiomyces-like fungus was isolated from bulbils from herbarium material as discussed in Lawrey (2002). Bulbils germinated within two days on potato dextrose agar (PDA, Difco, Detroit, Michigan, USA) without antibiotics, and outgrowths were isolated for liquid culture in either potato dextrose or Sabouraud’s (Difco) medium with dextrose. Approximately 2 µg dry mycelial mass was harvested after two weeks and extracted for DNA analysis.
In addition to the unknown culture, we obtained cultures of Limonomyces culmigenus (R. K. Webster & D. A. Reid) Stalpers & Loer. (ATCC 22523), L. roseipellis Stalpers & Loer. (CBS 299.82), and Laetisaria arvalis Burds. (type strain CBS 131.82) for DNA extraction and analysis. Previous studies (Andjic et al., 2005; Lawrey et al., 2007) suggested that these species may have phylogenetic affinities with the Marchandiomyces clade. Isolates were maintained in liquid culture (Sabouraud’s with dextrose) until approximately 2 µg dry mycelial mass had accumulated, and this mycelium was extracted for DNA analysis.
Molecular techniques
Genomic DNA was extracted from fungal tissue using the Bio 101 Fast DNA Spin Kit for tissue (Qbiogen, Illkirch, France) according to the manufacturer’s protocol with slight modifications. About 10 ng of extracted DNA were subjected to a standard PCR in a 50-µL reaction volume. We amplified approximately 1700 bp of portions of the nuc-LSU and ITS2 rDNA using primers (LR0R, LR3R, LR8R, LR5, LR7, LR9) available from the Vilgalys laboratory web site (http://www.biology.duke.edu/fungi/mycolab/primers.htm). The nuc-SSU with an approximate length of 1750 bp for most species was completely sequenced using NS17UCB, NS19UCB, NS3, NS21UCB, NS23UCB, NS24UCB, NS22UCB, NS20UCB, NS2, and CNS26 (White et al., 1990; Gargas and Taylor, 1992). After confirming the PCR product using ethidium bromide following separation on a 1% agarose gel, the products were purified with magnetic beads (Agencourt Biosciences, Beverly, Massachusetts, USA).
The purified PCR products were used in standard sequencing reactions with BigDye Terminator Ready Reaction Mix (Applied Biosystems, Foster City, California, USA). The sequencing reactions were then purified using Sephadex G-50 (Sigma-Aldrich, St. Louis, Missouri, USA), dried in a speed vacuum, denatured in HiDi Formamide (Applied Biosystems) and run on a SCE-9610 capillary machine (SpectruMedix, Reedsville, Pennsylvania, USA). The data collected were analyzed using the program BaseSpectrum (SpectruMedix), and about 600 bases were collected for each primer used. These sequences were then transferred to the program Sequencher (GeneCodes, Ann Arbor, Michigan, USA) for manual base calling and to make contiguous alignments of overlapping fragments.
Phylogenetic analyses
Initial placement of the unknown was done using a core data set of nuclear small and large subunit (nuc-SSU, nuc-LSU) rDNA sequences from 142 species (see the supplement in Binder et al. [2005] for strain information and GenBank accession numbers), with sequences added from a study of lichen-associated and sclerotial taxa to yield 268 terminals (Lawrey et al., 2007). The core data set was tested for positive conflict by bootstrapping nuc-SSU and nuc-LSU partitions separately in PAUP* version 4.0b10 (Swofford, 2002), using the neighbor-joining (NJ) nonparametric bootstrap (1000 replicates) with a maximum likelihood distance. Likelihood models were selected and parameters estimated using the Akaike information criterion (with the program MODELTEST version 3.7; Posada and Crandall, 1998). We found no evidence of topological conflict between partitions >70% and therefore combined the data. Missing data at the terminal ends of sequences were trimmed, but no other sequence data were excluded. Parsimony analysis of this core alignment indicated the unknown was a member of the Corticiales (Binder et al., 2005), closely related to Marchandiomyces spp. and other related fungi. We then assembled a Corticiales alignment containing nuc-SSU and nuc-LSU sequences from the core data set plus additional sequences obtained from GenBank or AFTOL (Assembling the Fungal Tree of Life; http://www.aftol.org). The Corticiales data set was aligned by eye in the program MacClade version 4.08 (Maddison and Maddison, 2005) and submitted to the TreeBASE database (S2053; http://www.treebase.org).
The Corticiales data set (Table 1) consisted of 36 terminals including five outgroup sequences representing the Gloeophyllales [Gloeophyllum sepiarum (Wulfen) P. Karst. and Heliocybe sulcata (Berk.) Redhead & Ginns] and the Thelephorales [Sarcodon imbricatus (L.) P. Karst., Thelephora sp. and Bankera fuligineoalba (J. C. Schmidt) Coker & Beers]. The data were tested for conflict using NJ bootstrap values as before and analyzed in PAUP* using maximum likelihood under the GTR+
+I model (estimated with MODELTEST 3.7, Posada and Crandall, 1998) with nucleotide frequencies estimated (A = 0.25, C = 0.21, G = 0.28, T = 0.26), a rate matrix of substitutions (A-C = 1.05, A-G = 3.85, A-T = 1.54, C-G = 0.75, C-T = 7.87, G-T = 1.0), proportion of invariable sites = 0.67, and 
= 0.55. A maximum likelihood bootstrap analysis was performed under the same settings using 1000 replicates with MAXTREES set to 1000. In addition, Bayesian phylogenetic analyses were carried out using the Metropolis-coupled Markov chain Monte Carlo method (MCMCMC) in MrBayes version 3.0b4 (Ronquist and Huelsenbeck, 2003). Analyses were run using a model with six categories of base substitution, a gamma-distributed rate parameter, and a proportion of invariant sites (GTR++I). Two parallel MCMCMC runs were performed each using four chains and 2000000 generations, sampling trees every 100th generation. The proportion of burn-in trees sampled before reaching equilibrium was estimated by plotting likelihood scores as a function of the number of generations. Posterior probabilities (PP) were determined by calculating a 50% majority-rule consensus tree in PAUP* with the proportion of trees gathered after convergence of likelihood scores was reached, and clades with PP 
0.95 were considered to be significantly supported.
RESULTS
Phylogenetic placement of the new species in the corticioid clade
Maximum likelihood (ML) analysis of an alignment of nuclear small (nuc-SSU) and large (nuc-LSU) subunit rDNA sequences from the Corticiales and closely related Gloeophyllales and Thelephorales resulted in one optimal tree (Fig. 1) with a score of –lnL = 9607.902. Bayesian runs converged after 200000 generations and 24402 trees were used to compute posterior probability (PP) values. The Corticiales as a whole and the clade containing described Marchandiomyces spp. are both strongly supported by PP values and ML bootstrap support, a result that has been observed before (DePriest et al., 2005; Lawrey et al., 2007). The new species Marchandiomyces marsonii forms a weakly supported clade with the turfgrass pathogen Laetisaria fuciformis (the generitype), which is sister to another strongly supported clade containing two recently described Marchandiomyces species (M. nothofagicola from southern Chile and M. buckii from the United States) and the grass pathogen Limonomyces culmigenus. The plant pathogen L. roseipellis (generitype) is basal to the entire clade, which is strongly supported (100%) by PP and moderately supported (79%) by ML bootstrap. This fungus is sister to another well-supported clade containing two additional Marchandiomyces species, the M. corallinus (the generitype) and M. lignicola. The entire Marchandiomyces clade appears to be sister to the type species of the Corticiales, Corticium roseum, and the Marchandiomyces + Corticium clade is sister to a weakly supported and diverse clade containing the basidiolichen Marchandiomphalina foliacea; the lichen pathogen Marchandiobasidium aurantiacum; Erythricium laetum, a salmon lignicolous fungus that grows on decayed wood, moist leaves, and possibly living mosses (Binder et al., 2005); Laetisaria arvalis, a facultative fungal parasite that has been studied for possible use as a biocontrol agent against Rhizoctonia and Pythium spp. (Burdsall et al., 1980; Conway et al., 2000; Bobba and Conway, 2003); and two plant pathogens, Waitea circinata, a soilborne saprobe and pathogen of cereals, turf grasses and legumes; and Erythricium salmonicolor (Berk. & Broome) Burds., a fungus causing pink crust of citrus, coffee, and rubber trees (Burdsall, 1985).
|
), we decided to retain the name E. salmonicolor in this paper. It is probable that neither name is appropriate.
We propose to describe the unknown here as a species of Marchandiomyces s.l. within the clade containing two other Marchandiomyces spp., and Laetisaria fuciformis and Limonomyces roseipellis. Further taxonomic resolution of these groups will require additional study of more species. A species we hope to include soon is Laetisaria agaves Burds. & Gilb., which was described from Arizona and produces coral, cartilaginous basidiocarps on leaf bases of Agave species (Burdsall and Gilbertson, 1982).
New species description
Marchandiomyces marsonii Diederich & Lawrey, sp. nov. (Figs. 2–11)
|
Type: Australia: Queensland: Cairns, on trees along road near Convention Centre, 16°55'40''S, 145°46'45''E, alt. 6 m a.s.l. (coordinates obtained by GPS), on dead, hanging leaves of Pandanus oblatus (Pandanaceae), 22 August 2006, G. Marson & M.-L. Wu s.n. (BRI–holotype; herb. Diederich–isotype). Type culture ATCC MYA-4210.
Basidiomata and conidiomata unknown. Colonies foliicolous, appearing as numerous bulbils. Mycelium not observed. Bulbils developing superficially over leaves of Pandanus oblatus, dispersed, rarely touching each other, roundish, strongly applanate, 160–260 µm in diameter, pastel red (Kornerup and Wanscher, 1984: 8A4–5) (Figs. 3, 4); surrounded by a thin, necrotic, hyaline layer 3–4.5 µm thick that does not stain with lactophenol cotton blue (Fig. 9) and that remains as an extremely thin, white, subspherical envelope after the disappearance of the interior of old bulbils (Fig. 5), sometimes giving the impression of white, empty "bulbils" (Fig. 8); external portions of bulbils with dispersed crystalline granules best visible in polarized light (Fig. 7) but without specialized cells (Fig. 10); bulbils composed of a dense agglomeration of thin-walled, smooth, hyaline, irregular, elongate, occasionally branched and septate hyphae (Fig. 11), cells mainly 1.5–3.5 µm diameter, clamps not observed in squash preparations.
Colonies in liquid culture pale pinkish orange, with aerial hyphae from which dispersed bulbils occasionally develop. Hyphae hyaline, septate, straight or curved, frequently branched, 2–3(–3.5) µm diameter, septa with clamps. Bulbils rare, leaving white, empty "bulbils" when disappearing (Fig. 8).
Marchandiomyces marsonii named after Guy Marson (Luxembourg) who discovered and collected the new taxon while searching for new Orbilia species in Australia.
Additional specimen examined—Australia: Queensland: Cairns, 4 km north of city center, Flecker Botanical Gardens, 16°54'01''S, 145°44'57''E, 14 m a.s.l. (coordinates obtained by global positioning system), on dead, hanging leaves of Pandanus oblatus, August 2006, G. Marson & M.-L. Wu s.n. (herb. Diederich).
DISCUSSION
The new species Marchandiomyces marsonii differs from others in the genus by its foliicolous habit and its large, reddish, flattened bulbils (Fig. 3). These resemble apothecia of species in the ascomycete genus Orbilia so closely that they were initially collected and labeled as such, and only subsequent microscopic examination revealed them to be basidiomycete bulbils. These bulbils presumably develop superficially from mycelium growing on Pandanus leaves without indication that they invade or degrade the leaves. Because the two known collections are from dead leaves still hanging on the tree (Fig. 2), it is not known if they are also present on living leaves. We therefore assume the species is saprobic/foliicolous.
The large, flattened bulbils of M. marsonii are similar to those of other members of the genus in color; production of pink, red or coral-colored pigments appears to be a common characteristic of many members of the clade. Formation of bulbils (or sclerotia) is also relatively common among the corticioid fungi and is a characteristic of all members of the genus Marchandiomyces. These are best regarded as resting or dispersal structures capable of surviving unfavorable conditions for long periods of time. In the case of M. marsonii, 5-month-old herbarium specimens were still viable, and other bulbilliferous species remain viable even longer. For example, the lichenicolous fungus Burgoa angulosa Diederich, Lawrey & Etayo (Cantharellales) could be cultured after five years in a herbarium.
Bulbils may themselves sometimes disintegrate and disappear from the surface of the host. We have repeatedly observed that in older herbarium specimens of Marchandiomyces corallinus and Marchandiobasidium aurantiacum the fungus apparently disappeared, even in rich specimens originally containing numerous bulbils. Some of this change in appearance can be attributed to mechanical loss because bulbils can be dislodged from the host lichen relatively easily. However, we believe that some loss takes place because bulbils remain viable for extended periods of time and eventually disintegrate in the herbarium. Our observations of Marchandiomyces marsonii provide unexpected support for this idea. Unlike other Marchandiomyces species, bulbils of M. marsonii are surrounded by a thin, hyaline, necrotic layer (Fig. 9). As bulbils age, they entirely disintegrate, leaving only this outer layer. In specimens from Pandanus leaves, older bulbils appear to get paler, turn white, and eventually disappear, leaving only the outer, necrotic layer (Fig. 5). Similarly, bulbils in isolated cultures entirely disappear as they age, leaving a surrounding layer in the form of a white, ball- or cuplike, empty structure with a large irregular opening (Fig. 8). The disintegration of bulbils in M. marsonii, and by implication other Marchandiomyces species as well, indicates these structures may remain viable for long periods under highly unfavorable conditions, but once they exhaust their resources they die and disintegrate, leaving little evidence of their presence on the host.
Our results revealed an unanticipated link between Marchandiomyces and certain members of the genera Laetisaria and Limonomyces, most of which are pathogens or endophytes of grasses that form orange, red, or coral sclerotia (Burdsall, 1979; Burdsall et al., 1980; Stalpers and Loerakker, 1982). The genera Laetisaria and Limonomyces have been assumed to be related, based largely on disease symptoms and anatomical characters of isolated cultures (Burdsall, 1979; Burdsall et al., 1980; Stalpers and Loerakker, 1982), but support of this relationship from genetic studies has been sparse. A recent phylogenetic study of the sterile red fungus pathogen of grasses (Andjic et al., 2005) using nuc-ITS1 sequences indicated that Limonomyces roseipellis, L. culmigenus, Laetisaria arvalis, and L. fuciformis were all closely related, a result partially (but not completely) supported by our analysis. Because we were not able to align GenBank ITS1 sequences from Andjic et al. (2005) with ITS sequences that we obtained from our cultures of Limonomyces or Laetisaria, we could not replicate their results. Nevertheless, our results using nuc-SSU and nuc-LSU sequences indicate a close relationship among Limonomyces spp., certain Marchandiomyces spp. (M. buckii, M. nothofagicola, M. marsonii) and Laetisaria fuciformis, with Limonomyces roseipellis occupying a basal position in the clade. The group has diverse substrate ecologies, with the use of monocot vascular plants a notable theme, suggesting an important influence on the evolution of the group. Contrary to Andjic et al. (2005), our results do not support the hypothesis that L. arvalis is closely related to either Limonomyces or to the generitype Laetisaria fuciformis. Given the apparent relationship between Erythricium salmonicolor and L. arvalis, taxonomic reevaluation of these species is needed.
The Corticiales is one of the most ecologically diverse groups of Agaricomycetes, containing saprobes, plant and fungal pathogens, and lichens. Members of the genus Marchandiomyces s.l. and their close relatives are especially diverse ecologically, with lichenicolous (M. corallinus, M. buckii, Marchandiobasidium aurantiacum), lignicolous (Marchandiomyces lignicola, M. nothofagicola), lichen-forming (Marchandiomphalina foliacea), and foliicolous (Laetisaria and Limonomyces spp., Marchandiomyces marsonii) members represented in the group. Because of the wide range of ecological conditions in the group, determining the ecology of the common ancestor would be of interest. Many members of the Corticiales form fruiting structures on recently deceased branches or grow on decorticated wood. These include Corticium roseum, Galzinia incrustans (Höhn. & Litsch.) Parmasto, Punctularia strigosozonata (Schwein.) P.H.B. Talbot, Dendrocorticium spp., Vuilleminia spp., strongly suggesting a lignicolous and possibly saprotrophic ancestor, an idea mentioned before (DePriest et al., 2005). Because many species have been described only recently and additional species are likely to be discovered in the future, discussions about ancestral ecology may be premature at this point. In Marchandiomyces s.l., however, the close association of lichenized, lichenicolous, and various plant-associated species indicates an unusually marked tendency for ecological transitions among these particular fungi.
|
FOOTNOTES
1 The authors thank G. Marson and H.-O. Baral for making available their specimens and photographs of Marchandiomyces marsonii and M. Binder for graciously providing advice and assistance with the phylogenetic analyses. 
4 Author for correspondence (e-mail: jlawrey{at}gmu.edu)
LITERATURE CITED
Andjic, V., A. L. Cole, AND J. D. Klena. 2005. Taxonomic identity of the Sterile Red Fungus inferred using nuclear rDNA ITS1 sequences. Mycological Research 109: 200–204.[CrossRef][Web of Science][Medline]
Binder, M., D. S. Hibbett, K.-H. Larsson, E. Larsson, E. Langer, AND G. Langer. 2005. The phylogenetic distribution of resupinate forms across the major clades of mushroom-forming fungi (Homobasidiomycetes). Systematics and Biodiversity 3: 113–157.[CrossRef][Web of Science]
Bobba, V., AND K. E. Conway. 2003. Competitive saprophytic ability of Laetisaria arvalis compared with Sclerotium rolfsi. Proceedings of the Oklahoma Academy of Science 83: 17–22.
Burdsall, H. H. Jr. 1979. Laetisaria, a new genus for the teleomorph of Isaria fuciformis. Transactions of the British Mycological Society 72: 419–422.[Web of Science]
Burdsall, H. H. Jr. 1985. A contribution to the taxonomy of the genus Phanerochaete (Corticiaceae, Aphyllophorales). Mycological Memoirs 10. J. Cramer, Braunschweig, Germany.
Burdsall, H. H. Jr., AND R. L. Gilbertson. 1982. New species of Corticiaceae (Basidiomycotina, Aphyllophorales) from Arizona. Mycotaxon 15: 333–340.[Web of Science]
Burdsall, H. H. Jr., H. C. Hoch, M. G. Boosalis, AND E. C. Setliff. 1980. Laetisaria arvalis (Aphyllophorales, Corticiaceae): A possible biological control agent for Rhizoctonia solani and Pythium species. Mycologia 72: 728–736.[CrossRef][Web of Science]
Clémençon. H. [with the assistance of V. Emmet and E. Emmet]. 2004. Cytology and plectology of the Hymenomycetes. Bibliotheca Mycologica 199, viii + 488 pp.
Conway, K. E., D. A. Gerken, M. A. Sandburg, AND D. L. Martin. 2000. Population dynamics of Laetisaria arvalis and Burkholderia cepacia, potential biocontrol agents in soil cores and thatch of creeping bent grass (Agrostis palustris). Proceedings of the Oklahoma Academy of Science 80: 39–46.
DePriest, P. T., M. Sikaroodi, J. D. Lawrey, AND P. Diederich. 2005. Marchandiomyces lignicola sp. nov. shows recent and repeated transition between a lignicolous and a lichenicolous habit. Mycological Research 109: 57–70.[CrossRef][Web of Science][Medline]
Diederich, P. 1990. New or interesting lichenicolous fungi. 1. Species from Luxembourg. Mycotaxon 37: 297–330.[Web of Science]
Diederich, P., AND J. D. Lawrey. 2007. 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. Mycological Progress 6: 61–80.[CrossRef][Web of Science]
Diederich, P., B. Schulteis, AND M. Blackwell. 2003. Marchandio-basidium aurantiacum gen. sp. nov., the teleomorph of Marchandiomyces aurantiacus (Basidiomycota, Ceratobasidiales). Mycological Research 107: 523–527.[CrossRef][Web of Science][Medline]
Etayo, J., AND P. Diederich. 1996. Lichenicolous fungi from the western Pyrenees, France and Spain. II. More deuteromycetes. Mycotaxon 60: 415–428.[Web of Science]
Gargas, A., AND J. W. Taylor. 1992. Polymerase chain reaction (PCR) primers for amplifying and sequencing 18S rDNA from lichenized fungi. Mycologia 84: 589–592.[CrossRef][Web of Science]
Hawksworth, D. L. 1979. The lichenicolous hyphomycetes. Bulletin of the British Musuem of Natural History. Botany 6: 183–300.
Hibbett, D. S. 2006. A phylogenetic overview of the Agaricomycotina. Mycologia 98: 917–925.
Hibbett, D. S., AND M. Binder. 2002. Evolution of complex fruiting-body morphologies in homobasidiomycetes. Proceedings of the Royal Society of London, B, Biological Sciences 269: 1963–1969.
Hibbett, D. S., M. Binder, J. F. Bischoff, M. Blackwell, P. F. Cannon, O. E. Erikssone, S. Huhndorff et al. 2007. A higher-level phylogenetic classification of the Fungi. Mycological Research 111: 509–547.[CrossRef][Web of Science][Medline]
Hibbett, D. S., L.-B. Gilbert, AND M. J. Donoghue. 2000. Evolutionary instability of ectomycorrhizal symbioses in basidiomycetes. Nature 407: 506–508.[CrossRef][Medline]
Hibbett, D. S., AND R. G. Thorn. 2001. Basidiomycota: Homobasidiomycetes. In D. J. McLauglin, E. G. McLauglin, and P. A. Lemke [eds.], The Mycota, vol. VII, part B. Systematics and Evolution, 121–167. Springer-Verlag, Berlin, Germany.
Kornerup, A., AND J. H. Wanscher. 1984. Methuen handbook of colour, 3rd ed. Methuen, London, UK.
Langer, E. 2002. Phylogeny of non-gilled and gilled basidiomycetes: DNA sequence inference, ultrastructure and comparative morphology. Habilitationsschrift, Tübingen University, Tübingen, Germany.
Larsson, K.-H. 2007. Re-thinking the classification of corticioid fungi. Mycological Research 111: 1040–1063.[Web of Science][Medline]
Larsson, K.-H., E. Larsson, AND U. Kõljalg. 2004. High phylogenetic diversity among corticioid homobasidiomycetes. Mycological Research 108: 983–1002.[CrossRef][Web of Science][Medline]
Lawrey, J. D. 2002. Isolation and culture of lichenicolous fungi. In I. Kranner, R.P. Beckett, and A. Varma [eds.], Protocols in lichenology: Culturing, biochemistry, physiology and use in biomonitoring, 75–84. Springer-Verlag, Berlin, Germany.
Lawrey, J. D., M. Binder, P. Diederich, M. C. Molina, M. Sikaroodi, AND D. Ertz. 2007. Phylogenetic diversity of lichen-associated homobasidiomycetes. Molecular Phylogenetics and Evolution 44: 778–789.[CrossRef][Web of Science][Medline]
Lim, Y. W. 2001. Systematic study of corticioid fungi based on molecular sequence analyses. Ph.D. dissertation, Seoul National University, Seoul, South Korea.
Maddison, D. R., AND W. P. Maddison. 2005. MacClade 4. 08: Analysis of phylogeny and character evolution. Sinauer, Sunderland, Massachusetts, USA.
Matheny, P. B., J. M. Curtis, V. Hofstetter, M. C. Aime, J.-M. Moncalvo, Z.-W. Ge, Z.-L. Yang et al. 2006. Major clades of Agaricales: A multilocus phylogenetic overview. Mycologia 98: 982–995.
Matheny, P. B., Z. Wang, M. Binder, J. M. Curtis, Y. W. Lim, R. H. Nilsson, K. W. Hughes et al. 2007. Contributions of rpb2 and tef1 to the phylogeny of mushrooms and allies (Basidiomycota, Fungi). Molecular Phylogenetics and Evolution 43: 430–451.[CrossRef][Web of Science][Medline]
Palice, Z., I. Schmitt, AND H. T. Lumbsch. 2005. Molecular data confirm that Omphalina foliacea is a lichen-forming basidiomycete. Mycological Research 109: 447–451.[CrossRef][Web of Science][Medline]
Posada, D., AND K. A. Crandall. 1998. MODELTEST: Testing the model of DNA substitution. Bioinformatics (Oxford, England) 14: 817–818.[CrossRef]
Ronquist, F., AND J. P. Huelsenbeck. 2003. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics (Oxford, England) 19: 1572–1574.[CrossRef]
Sikaroodi, M., J. D. Lawrey, D. L. Hawksworth, AND P. T. DePriest. 2001. The phylogenetic position of selected lichenicolous fungi: HobsoniaIllosporium, and Marchandiomyces. Mycological Research 105: 453–460.[CrossRef][Web of Science]
Stalpers, J. A., AND W. M. Loerakker. 1982. Laetisaria and Limonomyces species (Corticiaceae) causing pink diseases in turf grasses. Canadian Journal of Botany 60: 529–537.
Swofford, D. L. 2002. PAUP*: Phylogenetic analysis using parsimony (*and other methods), version 4.0b10. Sinauer, Sunderland, Massachusetts, USA.
White, T. J., T. D. Bruns, S. B. Lee, AND J.W. Taylor. 1990. Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In M. A. Innis, D. H. Gelfand, J. J. Sninsky, and T. J. White [eds.], PCR protocols, a guide to methods and applications, 315–322. Academic Press, San Diego, California, USA.
CiteULike Complore Connotea Del.icio.us Digg Facebook Reddit Technorati Twitter What's this?
This article has been cited by other articles:
|
|
 |
|
  D. Ertz, J. D. Lawrey, M. Sikaroodi, P. M. Gillevet, E. Fischer, D. Killmann, and E. Serusiaux A new lineage of lichenized basidiomycetes inferred from a two-gene phylogeny: The Lepidostromataceae with three species from the tropics Am. J. Botany, December 1, 2008; 95(12): 1548 - 1556. [Abstract] [Full Text] [PDF]
|
|
| ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
| HOME | HELP | FEEDBACK | SUBSCRIPTIONS | ARCHIVE | SEARCH | TABLE OF CONTENTS |
