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A. Calvitimela perlata, O-L-163770. B. C. armeniaca, O-L-170644. C. C. melaleuca, O-L-170490. D. C. talayana, O-L-126 (holotype). E. C. aglaea, O-L-173831. F. C. cuprea, O-L-179566 (holotype). G. C. livida, O-L-163835 (holotype). Bar 5 10 mm. Photo: E. Timdal.
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Contributing to the process of reassigning lecideoid lichens to natural taxa, we assessed phylogenetic relationships and species delimitation in the Calvitimela aglaea complex (Tephromelataceae) using DNA sequence data and morphological/anatomical and chemical characters. Phylogenetic analysis of nuclear (ITS, MCM7, TEF1-α) and mitochondrial (ribos...
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The former family Lobariaceae, now included in Peltigeraceae as subfamily Lobarioideae, has undergone substantial changes in its generic classification in recent years, based on phylogenetic inferences highlighting the polyphyly of the speciose genera Lobaria, Pseudocyphellaria and Sticta. Here we introduce the new genus Emmanuelia, named in honor...
Citations
... Much of our local research is based directly or indirectly on our collections (e.g., Bendiksby et al. 2014Bendiksby et al. , 2015Mienna et al. 2020;Nowak et al. 2021;Nygaard et al. 2022). These and ongoing fungarium-and herbarium-based activities include taxonomy, various herbariomics and fungariomics studies, spatial phylogenetics, functional biogeography, historical phylogeography, and assessment of insect herbivory based on photographs of herbarium sheets. ...
The Arctic has been, and is, an area of focus for the botanical and fungal (lichenized fungi included) collections at the Natural History Museum of Oslo. These collections house more than 233,000 unique Arctic specimens, the oldest dating back more than two centuries. The vascular plants account for 63 percent, lichens 30 percent, and fungi 7 percent. The Arctic collections have a circumpolar representation with emphasis on mainland Norway (48 percent) and Svalbard (13 percent), followed by Arctic America (10 percent), Greenland (9 percent), and Arctic Russia (8 percent). The Oslo herbarium and fungarium house collections from important polar expeditions like Fram-2, Gjøa, and Maud, but also of many expeditions where collecting biological specimens was the main purpose. The number of new collections was highest in the decades 1930 to 1939 and 2000 to 2009 with each around 35,000 new specimens. In the 1990s, a DNA Bank was established for DNA extracts and tissue samples, and it houses today 22,879 Arctic accessions of fungi, lichens, and plants. In times of climatic change and a tense geopolitical situation, the herbarium and fungarium at NHM-Oslo represent an invaluable source for biological information about the Arctic. We welcome the use of our collections for research-, nature management-, and teaching purposes.
... The chemistry (atranorin, norrangiformic acid, rangiformic acid, and usnic acid) also differs from other species in the genus. Bendiksby et al. (2015) showed by a phylogenetic analysis that C. talayana is the sister of the C. aglaea-complex which, in addition to C. aglaea (Sommerf.) Hafellner, includes the two recently described sorediate species Calvitimela cuprea Haugan & Timdal and C. livida Haugan & Timdal. ...
... Hafellner, includes the two recently described sorediate species Calvitimela cuprea Haugan & Timdal and C. livida Haugan & Timdal. Because of the long branches of the major clades, Bendiksby et al. (2015) divided the genus into four subgenera: Calvitimela, Severidea (the C. aglaea-complex), Paramela (to which C. talayana belongs) and Calomela. ...
... Range, Alaska (Bendiksby et al. 2015). Apart from that, all records are from Russia where it has been most extensively sampled in the Ural Mountains (GBIF.org; ...
The lichen species Calvitimela talayana (Tephromelataceae, Lecanorales) is rarely collected worldwide but seems to have a broad and northern circumpolar distribution. It is here reported as new to Fennoscandia. The collection was made on siliceous rocks in alpine to arctic boulderfields at massif Lovozerskie gory on the Kola peninsula (Murmansk oblast, Russia), extending the known distribution range of this species by almost 1200
km. An additional collection from the Priuralsky district (Yamalo-Nenets Autonomous District) was made 160 km north of the former northernmost collection in the Ural mountain. The specimens fit C. talayana in morphology (pale yellow areolae and yellow to dark green soralia), and the identification was confirmed by thin-layer chromatography and a maximum likelihood phylogenetic analysis of the barcode marker nrITS.
... Species designations are hypotheses to be tested as new evidence becomes available. Recent molecular systematic studies have repeatedly revealed evolutionary lineages within phenotypically delimited lichenized fungi (e.g., Steinová et al. 2013;Lücking et al. 2014;Bendiksby et al. 2015;Alors et al. 2016;Hawksworth and Lücking 2017;Boluda et al. 2019). Whether or not such morphologically indistinguishable, or "cryptic", evolutionary lineages should be recognized at species level may have critical implications for conservation biology and other fields of biology that use species as a fundamental unit (Lumbsch and Leavitt 2011). ...
We have studied the highly oceanic genus Coniocarpon in Norway. Our aim has been to delimit species of Coniocarpon in Norway based on an integrative taxonomic approach. The material studied comprises 120 specimens of Coniocarpon , obtained through recent collecting efforts (2017 and 2018) or received from major fungaria in Denmark, Finland, Norway and Sweden, as well as from private collectors. We have assessed (1) species delimitations and relationships based on Bayesian and maximum likelihood phylogenetic analyses of three genetic markers (mtSSU, nucITS and RPB2 ), (2) morphology and anatomy using standard light microscopy, and (3) secondary lichen chemistry using high-performance thin-layer chromatography. The results show three genetically distinct lineages of Coniocarpon , representing C. cinnabarinum , C. fallax and C. cuspidans comb. nov. The latter was originally described as Arthonia cinnabarina f. cuspidans and is herein raised to species level. All three species are supported by morphological, anatomical and chemical data.
... Historically, lichens were classified using a mixture of traits assumed only in symbiosis and other, purely fungal characteristics such as spore size; the totality was called a lichen species. Assumed evolutionary groupings, such as the genus Lecidea, included dozens of interdigitated species that were specialized for rock, bark or wood [20][21][22]. In 1950, in order to rectify the nomenclatural instability arising from the recognition of lichens as multi-domain symbioses, the name of a lichen was anchored to that of its presumed single fungus by a change to the code of nomenclature [23]. ...
... The second change is another by-product of fungal molecular phylo- genetics: species once thought closely related have often turned out not to be [21]. We now know that the rock- dwelling species of Lecidea, for instance, are only distantly related to those found on bark and wood, placed in other genera and families altogether [22,26]. The third change concerns the nature of the symbioses themselves. ...
Lichens exhibit varying degrees of specialization with regard to the surfaces they colonize, ranging from substrate generalists to strict substrate specialists. Though long recognized, the causes and consequences of substrate specialization are poorly known. Using a phylogeny of a 150-200 Mya clade of lichen fungi, we asked whether substrate niche is phylogenetically conserved, which substrates are ancestral, whether specialists arise from generalists or vice versa and how specialization affects speciation/extinction processes. We found strong phylogenetic signal for niche conservatism. Specialists evolved into generalists and back again, but transitions from generalism to specialism were more common than the reverse. Our models suggest that for this group of fungi, 'escape' from specialization for soil, rock and bark occurred, but specialization for wood foreclosed evolution away from that substrate type. In parallel, speciation models showed positive diversification rates for soil and rock dwellers but not other specialists. Patterns in the studied group suggest that fungal substrate specificity is a key determinant of evolutionary trajectory for the entire lichen symbiosis.
... In these two studies, the species of interest were shown to be conspecific. In contrast, Bendiksby et al. (2015) showed that sterile, sorediate specimens of the Calvitimela aglea complex were two distinct lineages, impossible to distinguish morphologically but differentiated in chemistry and ecology. ...
The two corticolous species Fuscidea lightfootii (Sm.) Coppins & P. James and F . pusilla Tønsberg are morphologically and chemically similar and it has been suggested that they are conspecific. We investigated the interspecific relationship between F. lightfootii and F . pusilla using ITS, LSU and mtSSU rDNA. The combined multigene phylogeny shows that these species are genetically distinct. They are similar in ascocarp anatomy but in thallus morphology and substratum preferences there may be slight differences between them. Moreover, F. pusilla displays a broader ecological range than F . lightfootii . Even though some morphotypes appeared distinct and may be assigned to one of the two species with some degree of certainty, the use of DNA sequencing is recommended for their identification. Epitypes are designated for both species.
... Alignments were established in BIOED-IT 7.2.3 (Hall 1999) using the "ClustalW/Multiple alignment" option with subsequent manual adjustments. We analysed and summarized the data with parsimony and Bayesian phylogenetic methods, including model testing, as described in Bendiksby et al. (2015). As configuration settings in PartitionFinder v.1.1.1 (Lanfear et al. 2012), we used linked branch lengths, data blocks according to named genetic region (i.e. ...
Herein we describe the new species, Psora taurensis, from two localities in the Taurus Mountains in Turkey at ca. 1000 m altitude. Investigations of anatomy, secondary chemistry and DNA sequences (ITS and mtSSU) of P. taurensis and presumed close relatives suggest that P. taurensis is a distinct evolutionary line-age with P. tenuifolia as its sister, although it is morphologically more similar to P. russellii and P. vallesiaca.
... The species grew on ultramafic rocks and produced atranorin and substances of the stictic acid aggregate together with accessory norstictic acid. Initial attempts to identify the specimen were unsuccessful, and as such it was filed away in the undetermined sterile crustose lichens at NY. Subsequently when indexing the recent contribution on Calvitimela by Bendiksby et al. (2015) the first author made the connection between C. cuprea, which was newly described therein, and the specimen collected in Newfoundland. Further study confirmed that the material was conspecific with C. cuprea and thus represents the first report from North America. ...
Arthothelium lichenale is placed in synonymy with Mycoporum compositum. The morphological distinctions between Collema pustulatum and Leptogium apalachense are discussed and the North American distributions of the species are revised. The distribution of Lecidea roseotincta in North America is extended to include the central and southern Appalachian Mountains. The distribution of Lecidella subviridis is expanded in northeastern North America. Pyrenula reebiae is placed in synonymy with P. leucostoma and both species are illustrated and discussed. Pyrenula shirabeicola is removed from synonymy with P. pseudobufonia and both species are illustrated and discussed. The following taxa are newly reported from North America: Calvitimela cuprea (Canada, Newfoundland & Labrador), Hypotrachyna consimilis (U.S.A., North Carolina), Schismatomma graphidioides (U.S.A., Alabama and New Jersey).
... Nonetheless, these studies have yielded inconsistent results as to whether the chemotypes were monophyletic and should be recognized as distinct taxa (not monophyletic not recognized. e.g., Hodkinson 2009, Lendemer et al. 2015; monophyletic and recognized, e.g., Leavitt et al. 2013, Bendiksby et al. 2015. Given the many chemically defined species in lichens, particularly in macrolichen groups, such as the Parmeliaceae, it is hoped that the present study serves as further motivation for others to examine such taxa with studies that combine investigations of molecular data with those from other avenues of inquiry, such as biogeography, ecology, and morphology. ...
Species delimitation in foliose macrolichens has long been dominated by chemotaxonomy, resulting in many examples of seemingly morphologically identical species that differ only in the secondary metabolites they produce. This study examines two such species that are widespread in southeastern North America: Parmotrema madagascariaceum (Hue) Hale and Parmotrema xanthinum (Müll. Arg.) Hale. Drawing on data from biogeography, chemistry, morphology, and molecular phylogenetic analyses of nuclear ribosomal internal transcribed spacer sequence data, the two species are treated as conspecific. A taxonomic treatment is presented along with a discussion of previous studies that have examined the delimitation of species that differ only in the presence or absence of compounds that are accessories in addition to a primary set of substances present in both species.
... Alignments were established in BIO-EDIT 7.2.3 (Hall 1999) using the "ClustalW/Multiple alignment" option with subsequent manual adjustments. We analysed and summarized the data with parsimony and Bayesian phylogenetic methods, including model testing, as described in Bendiksby et al. (2015). The nuclear and mitochondrial datasets were analysed separately and in combination (concatenated) with indels treated as missing data. ...
In the present study, we describe the new species, Psora altotibetica, from nine localities in China (Tibetan area) and Nepal. The study includes analyses of anatomy, secondary chemistry, and DNA sequence data of P. altotibetica and presumed close relatives. Psora altotibetica resembles P. indigirkae morphologically, but is phylogenetically closer to P. tenuifolia and P. vallesiaca. It differs from P. indigirkae in the colour of the apothecia, the size of the ascospores, and in the secondary chemistry. The species is terricolous and was collected in the alpine zone of the Great Himalayas between 4230 and 5000 m altitude. Psora tenuifolia and P. vallesiaca are here reported as new to China and the Himalayas. Copyright Einar Timdal et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
... Only a handful of basic apothecial growth types have been described and this, combined with a finite number of basic ascospore schemes, led early researchers to assume that rock-dwelling lichens arose from only a handful of fungal lineages. In the most extreme case, more than 1300 species, including non-rock-inhabiting species, were placed in a single genus, Lecidea, only to be subsequently found to represent dozens of different convergent lineages (Hedlund 1892;Hertel 1977;Coppins 1983;Printzen 1995;Bendiksby et al. 2015). The most recent extension of this trend has been in the discovery, using molecular tools, that many long-recognized species names are themselves merely fronts for even more genetically distinct but phenotypically virtually indistinguishable species (Ruprecht et al. 2010;Orange 2014;Westberg et al. 2015), so-called 'cryptic species'. ...
Large, architecturally complex lichen symbioses arose only a few times in evolution, increasing thallus size by orders of magnitude over those from which they evolved. The innovations that enabled symbiotic assemblages to acquire and maintain large sizes are unknown. We mapped morphometric data against an eight-locus fungal phylogeny across one of the best-sampled thallus size transition events, the origins of the Placopsis lichen symbiosis, and used a phylogenetic comparative framework to explore the role of nitrogen-fixing cyanobacteria in size differences. Thallus thickness increased by >150% and fruiting body core volume increased nine-fold on average after acquisition of cyanobacteria. Volume of cyanobacteria-containing structures (cephalodia), once acquired, correlates with thallus thickness in both phylogenetic generalized least squares (PGLS) and phylogenetic generalized linear mixed-effects (pGLMM) analyses. Our results suggest that the availability of nitrogen is an important factor in the formation of large thalli. Cyanobacterial symbiosis appears to have enabled lichens to overcome size constraints in oligotrophic environments such as acidic, rain-washed rock surfaces. In the case of the Placopsis fungal symbiont this has led to an adaptive radiation of more than 60 recognized species from related crustose members of the genus Trapelia. Our data suggest that pre-cyanobacterial symbiotic lineages were constrained to forming a narrow range of phenotypes, so-called cryptic species, leading systematists until now to recognize only six of the 18 species clusters we identified in Trapelia. This article is protected by copyright. All rights reserved.
