Trapelioid fungi constitute a widespread group of mostly crust-forming lichen mycobionts that are key to understanding the early evolutionary splits in the Ostropomycetidae, the second-most species-rich subclass of lichenized Ascomycota. The uncertain phylogenetic resolution of the approximately 170 species referred to this group contributes to a poorly resolved backbone for the entire subclass. Based on a data set including 657 newly generated sequences from four ribosomal and four protein-coding gene loci, we tested a series of a priori and new evolutionary hypotheses regarding the relationships of trapelioid clades within Ostropomycetidae. We found strong support for a monophyletic group of nine core trapelioid genera but no statistical support to reject the long-standing hypothesis that trapelioid genera are sister to Baeomycetaceae or Hymeneliaceae. However, we can reject a sister group relationship to Ostropales with high confidence. Our data also shed light on several long-standing questions, recovering Anamylopsoraceae nested within Baeomycetaceae, elucidating two major monophyletic groups within trapelioids (recognized here as Trapeliaceae and Xylographaceae), and rejecting the monophyly of the genus Rimularia. We transfer eleven species of the latter genus to Lambiella and describe the genus Parainoa to accommodate a previously misunderstood species of Trapeliopsis. Past phylogenetic studies in Ostropomycetidae have invoked “divergence order” for drawing taxonomic conclusions on higher level taxa. Our data show that if backbone support is lacking, contrasting solutions may be recovered with different or added data. We accordingly urge caution in concluding evolutionary relationships from unresolved phylogenies.
Saxicolous, lecideoid lichenized fungi have a cosmopolitan distribution but, being mostly cold adapted, are especially abundant in polar and high-mountain regions. To date, little is known of their origin or the extent of their trans-equatorial dispersal. Several mycobiont genera and species are thought to be restricted to either the Northern or the Southern Hemisphere, whereas others are thought to be widely distributed and occur in both hemispheres. However, these assumptions often rely on morphological analyses and lack supporting molecular genetic data. Also unknown is the extent of regional differentiation in the southern polar regions. An extensive set of lecideoid lichens (185 samples) was collected along a latitudinal gradient at the southern end of South America. Subantarctic climate conditions were maintained by increasing the elevation of the collecting sites with decreasing latitude. The investigated specimens were placed in a global context by including Antarctic and cosmopolitan sequences from other studies. For each symbiont three markers were used to identify intraspecific variation (mycobiont: ITS, mtSSU, RPB1; photobiont: ITS, psbJ-L, COX2). For the mycobiont, the saxicolous genera Lecidea, Porpidia, Poeltidea and Lecidella were phylogenetically re-evaluated, along with their photobionts Asterochloris and Trebouxia. For several globally distributed species groups, the results show geographically highly differentiated subclades, classified as operational taxonomical units (OTUs), which were assigned to the different regions of southern South America (sSA). Furthermore, several small endemic and well-supported clades apparently restricted to sSA were detected at the species level for both symbionts.
Lichens are widely acknowledged to be a key component of high latitude ecosystems. However, the time investment needed for full inventories and the lack of taxonomic identification resources for crustose lichen and lichenicolous fungal diversity have hampered efforts to fully gauge the depth of species richness in these ecosystems. Using a combination of classical field inventory and extensive deployment of chemical and molecular analysis, we assessed the diversity of lichens and associated fungi in Glacier Bay National Park, Alaska (USA), a mixed landscape of coastal boreal rainforest and early successional low elevation habitats deglaciated after the Little Ice Age. We collected nearly 5000 specimens and found a total of 947 taxa, including 831 taxa of lichen-forming and 96 taxa of lichenicolous fungi together with 20 taxa of saprotrophic fungi typically included in lichen studies. A total of 98 species (10.3% of those detected) could not be assigned to known species and of those, two genera and 27 species are described here as new to science: Atrophysma cyanomelanos gen. et sp. nov., Bacidina circumpulla, Biatora marmorea, Carneothele sphagnicola gen. et sp. nov., Cirrenalia lichenicola, Corticifraga nephromatis, Fuscidea muskeg, Fuscopannaria dillmaniae, Halecania athallina, Hydropunctaria alaskana, Lambiella aliphatica, Lecania hydrophobica, Lecanora viridipruinosa, Lecidea griseomarginata, L. streveleri, Miriquidica gyrizans, Niesslia peltigerae, Ochrolechia cooperi, Placynthium glaciale, Porpidia seakensis, Rhizocarpon haidense, Sagiolechia phaeospora, Sclerococcum fissurinae, Spilonema maritimum, Thelocarpon immersum, Toensbergia blastidiata and Xenonectriella nephromatis. An additional 71 ‘known unknown’ species are cursorily described. Four new combinations are made: Lepra subvelata (G. K. Merr.) T. Sprib., Ochrolechia minuta (Degel.) T. Sprib., Steineropsis laceratula (Hue) T. Sprib. & Ekman and Toensbergia geminipara (Th. Fr.) T. Sprib. & Resl. Thirty-eight taxa are new to North America and 93 additional taxa new to Alaska. We use four to eight DNA loci to validate the placement of ten of the new species in the orders Baeomycetales, Ostropales, Lecanorales, Peltigerales, Pertusariales and the broader class Lecanoromycetes with maximum likelihood analyses. We present a total of 280 new fungal DNA sequences. The lichen inventory from Glacier Bay National Park represents the second largest number of lichens and associated fungi documented from an area of comparable size and the largest to date in North America. Coming from almost 60°N, these results again underline the potential for high lichen diversity in high latitude ecosystems.
Nomenclatural type definitions are one of the most important concepts in biological nomenclature. Being physical objects that can be re-studied by other researchers, types permanently link taxonomy (an artificial agreement to classify biological diversity) with nomenclature (an artificial agreement to name biological diversity). Two proposals to amend the International Code of Nomenclature for algae, fungi, and plants (ICN), allowing DNA sequences alone (of any region and extent) to serve as types of taxon names for voucherless fungi (mainly putative taxa from environmental DNA sequences), have been submitted to be voted on at the 11th International Mycological Congress (Puerto Rico, July 2018). We consider various genetic processes affecting the distribution of alleles among taxa and find that alleles may not consistently and uniquely represent the species within which they are contained. Should the proposals be accepted, the meaning of nomenclatural types would change in a fundamental way from physical objects as sources of data to the data themselves. Such changes are conducive to irreproducible science, the potential typification on artefactual data, and massive creation of names with low information content, ultimately causing nomenclatural instability and unnecessary work for future researchers that would stall future explorations of fungal diversity. We conclude that the acceptance of DNA sequences alone as types of names of taxa, under the terms used in the current proposals, is unnecessary and would not solve the problem of naming putative taxa known only from DNA sequences in a scientifically defensible way. As an alternative, we highlight the use of formulas for naming putative taxa (candidate taxa) that do not require any modification of the ICN.
Although several lichen inventories exist for European ultramafic sites, only four surveys of serpentine lichens for North America have been published to date. Of those, only one has been conducted in California. We conducted a survey of saxicolous lichens from ultramafic rocks (including nephrite, partially serpentinized peridotite, and serpentinite) and non-ultramafic rocks (including silica-carbonate, shale, and sandstone) at the New Idria serpentinite mass, San Benito County, California. X-ray Fluorescence Analysis of the rocks from which the lichens were collected revealed significant elemental differences between the ultramafic and non-ultramafic rocks for 26 of the 32 major and trace elements analyzed. We identified a total of 119 species of lichenized and lichenicolous fungi; 60 species were restricted to ultramafic substrata, 19 to silica-carbonate, and 15 to shale and sandstone. Only 4 species were shared in common. A permutational multivariate analysis of variance (perMANOVA) test revealed significant differences in lichen assemblages between ultramafic and non-ultramafic rocks at the species level but not at the generic level, with species richness (alpha-diversity) significantly greater at the ultramafic sites. We suggest that, although differences in geochemistry clearly influence the lichen community composition, other factors, especially substratum age and the physical characteristics of the rock, are of equal, if not greater, importance. Of all the species collected, six, Buellia aethalea, B. ocellata, Caloplaca oblongula, Rhizocarpon saurinum, Thelocarpon laureri, and Trapelia obtegens, are reported new to California, along with an apparently previously undescribed Solenopsora sp. The rest of the species encountered are relatively frequent in the lichen flora of southern and central California, except Aspicilia praecrenata, a rare California endemic that we collected on both ultramafic and non-ultramafic rocks.
Ten new species in nine different genera are described from the Falkland Islands (Islas Malvinas): Bryonora granulata with a finely granular thallus containing perlatolic acid; Bryoria mariensis, a terricolous species with norstictic acid and unusual cortex cells; Carbonea hypopurpurea with a K+ purple hypothecium and a thallus containing confluentic and 2′-O-methylperlatolic acids; Caloplaca megalariicola lichenicolous on Megalaria grossa; Cladonia flammea with a red-orange coloration on the lower side of the primary squamules; Cliostomum falklandicum, on rocks and with a dispersed thallus containing only atranorin; Lepraria malouina with usnic and stictic acids; Rimularia andreaeicola, over bryophytes and lacking lichen substances; R. subpsephota, similar to R. psephota but with a discrete white thallus lacking norstictic acid; and Usnea austrocampestris, a straggling species in sect. Neuropogon from the mountain tops. Rimularia andreaeicola is also known from Tierra del Fuego and R. subpsephota from Tierra del Fuego and South Georgia, but the other species are known only from the Falkland Islands. The new combinations Carbonea agellata, C. subdeclinans, Cliostomum aeruginascens and C. violascens are also made; Lecidea interrupta Darb. and Lecidea protracta Darb. are reduced to synonymy with Lecanora xantholeuca (Müll. Arg.) Hertel; Rhizocarpon simillimum is reported for the first time from the Southern Hemisphere, from the Falkland Islands and New Zealand; and Bryoria chalybeiformis is reported for the first time from the Falkland Islands.
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