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Scanning electron micrographs of cortical features. Figs 2–3. Melanelixia glabra (MAF 10228), showing fenestrations in the epicortex. Fig. 4. Melanohalea exasperata (MAF 7636) pseudocyphella. Fig. 5. Melanelia stygia (Hafellner 51658) pseudocyhella. Bar=60 mm.
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This paper continues a revision of generic concepts in the parmelioid lichens using molecular data in order to reach a consensus among lichenologists over which segregates proposed over the last two decades should be accepted. Here we employ data from three gene portions to provide a basis for a revised generic concept of the brown parmelioid liche...
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This paper presents a taxonomic treatment of Bulbothrix and Relicina lichens (Ascomycota, Parmeliaceae) collected in the Philippines, two genera characterized by the presence of bulbate cilia. A total of five species of Bulbothrix and fifteen species of Relicina were identified, with one new record for each genus. The Philippines is a major center...
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... 2, transmitted light microscope Zeiss Primo Star, a standard set of chemical reagents for carrying out color spots reactions for identification of certain groups of lichen substances in thalli. The nomenclature of lichens mainly follows Yoshimura (1971), Elix, Hale (1987), Printzen, Tønsberg, (2003), Blanco et al. (2004), Frisch et al. (2006, Westberg (2007), Spisok… (2010), Spribille et al. (2014), Gagarina (2015), Mongkolsuk et al. (2015), Printzen et al. (2016), Lendemer, Harris (2017), Sheard et al. (2017), Tchabanenko (2018, Westberg et al. (2021), and Flora… (2022). ...
Information on lichens of Shikotan Island based on original and literature data is provided. Altogether 278 lichen species and allied fungi are documented, of which Xylographa hians is new to Eurasia, Candelariella subdeflexa is new to Russia, nine species are new to the Russian Far East, 28 species are new to the Sakhalin Region, 39 species are new to Kuril Islands, and 94 species are new to Shikotan Island. For each species, data on distribution in the Sakhalin Region and neighboring regions are given.
... Instead, the recognized lichenicolous genus in Agaricostilbomycetes is Crittendenia , which is not present among our samples. Crittendenia includes two described lichenicolous species: C. coppinsii growing on thalli of Melanelixia and Melanohalea species (Blanco et al., 2004;Arup and Sandler Berlin, 2011;Divakar et al., 2017), and C. lichenicola growing on lichen species of Micarea . Some other still undescribed Crittendenia species parasitize lichen hosts in the families Lecanoraceae, Lobariaceae, Parmeliaceae, Physciaceae, Ramalinaceae and Teloschistaceae . ...
Lichens are well-known examples of complex symbiotic associations between organisms from different Kingdoms. Microfungi in particular, establish diverse associations with the hosting lichen thallus, as species-specific parasites or transient co-inhabitants. The whole community of lichen-associated fungi constitute the ‘lichen mycobiome’ and comprises both ascomycetes and basidiomycetes, including filamentous and yeast taxa. Metabarcoding results and microscopy analyses show that in some thalli, basidiomycetes can be frequent lichen-associated fungi but so far only a few species could be axenically isolated and morphologically characterized. Within a broad project aiming at characterizing the mycobiome diversity by culture-dependent and independent approaches in two lichen species selected as reference models – Rhizoplaca melanophthalma and Tephromela atra, we succeed in isolating and culturing 76 new strains of basidiomycetous yeasts. The lichen thalli were collected in different mountain regions worldwide and at relatively high elevation. The yeast strains were isolated on different growth media and were studied for their morphological and genetic diversity. Nuclear internal transcribed spacer (ITS) and ribosomal large subunit (LSU) sequence analyses identified them to belong to ten families within the orders Agaricostilbomycetes, Cystobasidiomycetes, Microbotryomycetes, Tremellomycetes and Ustilaginomycetes. The yeasts here detected only showed patterns of host-preference in a few cases and they are potentially related to the ecological conditions.
... The brown Parmeliae (Esslinger 1977) have been an object of numerous studies (Guzow-Krzemińska and Węgrzyn 2003;Blanco et al. 2005;Crespo et al. , 2011Nelsen et al. 2011;Divakar et al. 2012;Thell et al. 2012;Leavitt et al. 2014Leavitt et al. , 2015 and, due to this exceptional attention, they are one of the best-studied assemblages in the family Parmeliaceae. These lichens are a polyphyletic group possessing foliose, a dark to medium brown thallus and usually lacking atranorin or usnic acid in the cortex (Esslinger 1977;Blanco et al. 2004). ...
... Most of the available data concern samples collected in a few regions of the world, such as Europe and North America. The North American species of this group were studied in Greenland and Canada (Leavitt et al. 2014;Leavitt et al. 2015), while samples from Europe originated mainly from the north -Iceland, Finland, Norway and Sweden (Blanco et al. 2004;Divakar et al. 2012;Xu et al. 2017). Therefore, we decided to fill in the gap in sampling and focused our study on samples collected in Central Europe. ...
Infraspecific variation of the ITS rDNA region of some brown Parmeliae occurring in Poland is studied and compared with non-molecular characters. Haplotype networks are used to illustrate the variability within the species. Both newly-produced sequences from Central Europe and from all over the world, downloaded from the GenBank, are used.
The number of haplotypes found for each taxon ranged from five in Melanelia stygia to 12 in Melanelia hepatizon and Montanelia disjuncta ; however, their numbers correlate with the number of specimens tested. New haplotypes for Melanelia agnata , M. hepatizon and Cetraria commixta are found. Based on our 169-sample dataset, we could not infer any geographical correlation, either locally or world-wide. Many of the analysed haplotypes were widely distributed and the same haplotype was often shared between temperate and polar populations. A comparison of molecular, morphological, anatomical and chemical characters also shows no correlation.
... Our results show that the species delimitation in the new genus will need further investigation. Crittendenia coppinsii is a well-delimited species according to our phylogenetic studies and microscopical observations, and apparently grows on Melanelixia and Melanohalea only, two closely related genera in the Parmeliaceae (Blanco et al. 2004;Arup & Sandler Berlin 2011;Divakar et al. 2017). In contrast, herbarium specimens currently assigned to C. lichenicola probably represent a heterogeneous assemblage of several species, some of which are possibly host-specific. ...
The lichenicolous ‘heterobasidiomycetes’ belong in the Tremellomycetes (Agaricomycotina) and in the Pucciniomycotina. In this paper, we provide an introduction and review of these lichenicolous taxa, focusing on recent studies and novelties of their classification, phylogeny and evolution. Lichen-inhabiting fungi in the Pucciniomycotina are represented by only a small number of species included in the genera Chionosphaera , Cyphobasidium and Lichenozyma . The phylogenetic position of the lichenicolous representatives of Chionosphaera has, however, never been investigated by molecular methods. Phylogenetic analyses using the nuclear SSU, ITS, and LSU ribosomal DNA markers reveal that the lichenicolous members of Chionosphaera form a monophyletic group in the Pucciniomycotina, distinct from Chionosphaera and outside the Chionosphaeraceae . The new genus Crittendenia is described to accommodate these lichen-inhabiting species. Crittendenia is characterized by minute synnemata-like basidiomata, the presence of clamp connections and aseptate tubular basidia from which 4–7 spores discharge passively, often in groups. Crittendenia , Cyphobasidium and Lichenozyma are the only lichenicolous lineages known so far in the Pucciniomycotina, whereas Chionosphaera does not include any lichenicolous taxa.
... (Fig. 1). Air dried samples were studied with a Nikon SMZ1500 stereomicroscope and a Nikon Eclipse 80i light microscope with standard identification methods for lichenized and lichenicolous fungi (Arup et al. 2013, Blanco et al. 2004, Brodo et al. 2001, Calatayud et al. 2002, Darmostuk 2016, Dobson 2005, Esslinger 1997, Etayo and Calatayud 1998, Galloway and Moberg 2005, Giralt 2001, Hawksworth 1983, Kocakaya et al. 2011, Marina et al. 2005, Navarro-Rosinés et al. 2009, Nordin et al. 2010, Smith et al. 2009). Specimens are deposited in the herbarium of the Biology Department, Faculty of Science, Karadeniz Technical University, Turkey (KTUB). ...
A contribution to the lichen flora of Turkey is presented. A total of 282 lichen taxa and 20 lichenicolous fungi, of which 4 are varieties, are determined from 87 different localities in Muş province (Turkey). Lichenostigmagracile, a lichenicolous fungus, is new to Turkey, and 274 lichen species and 20 lichenicolous fungi are new for Muş.
... The lichen nomenclature follows Nordin et al. (2011a) and some other works (Darbishire 1909;Oxner 1933;Harada 1993;Moberg 1995;Makarova 2004;Blanco et al. 2004;Sipman 2006;Hafellner & Türk 2016). Description. ...
Fifty-six species of saxicolous lichens are reported for the first time from the Kodar Range. Circinaria scyphulifera is described as new to science. Aspicilia nikrapensis and Fuscidea submollis are new for Russia; Aspicilia sublapponica , Lepra monogona and Thelignya lignyota are new for southern Siberia; and 35 species of saxicolous lichens are reported for the first time for the Stanovoye Nagor’e highlands. Fuscopannaria ahlneri alredy appears in the Red Data Book of the Trans-Baikal Territory.
... Examples include the genera Melanelixia Blanco et al. (2004a: 881), Melanohalea Blanco et al. (2004a: 882) and Montanelia Divakar et al. (2012: 2022 as segregates of Melanelia Esslinger (1978: 46) s. lat. (Blanco et al. 2004a;Divakar et al. 2012); Austroparmelina Crespo et al. (2010a: 209) segregated from Parmelina Hale (1974a: 481). Lastly, Remototrachyna Divakar et al. (2010: 584) was segregated from Hypotrachyna Hale (1974b: 340) s. lat. ...
... GenBank accession numbers and details of studied material are shown in Table 1. The data sets include 122 sequences from previous publications by our group (Blanco et al. 2004a;Crespo et al. 2007;Divakar et al. 2006;Divakar et al. 2010;Lumbsch et al. 2008), five downloaded from GenBank and 118 newly generated sequences. ...
Hypotrachyna is a speciose genus of primarily tropical and oceanic lichen-forming fungi. It includes species with distinct distribution patterns, such as pantropical, restricted and disjunct species. We used a dataset of mitochondrial SSU, nuclear ITS and LSU ribosomal DNA from 89 specimens to study the historical biogeography of the genus. We employed Bayesian and maximum likelihood approaches for phylogenetic analyses, a likelihood-based approach to ancestral area estimation, and a Bayesian approach to estimate divergence times of major lineages within the genus based on molecular evolutionary rates for ITS and a secondary calibration point at the Hypotrachyna clade – Parmeliopsis split. Our analyses suggest that the genus might have originated in the Neotropics during the Eocene and that the split of major lineages happened primarily during the Eocene and Oligocene. The major diversification within those clades is estimated to have occurred during the Miocene. Pantropical species distributions are explained by long-distance dispersal. A number of currently accepted species were found to be non-monophyletic, illustrating that the delimitation of species in the genus needs attention.
... Sequential Elution Technique (SET) In order to evaluate whether Fe and Zn are bound to the melanized hyphae, a SET experiment was carried out on two species: the two-side heavily melanized M. glabratula and the one-side lightly melanized P. subrudecta, selected for their uniform cell wall chemistry (Krog 1982;Blanco et al. 2004;Smith et al. 2009). SET protocols call for the use of a well-defined chemical compound to extract the extracellular fraction of a specific element. ...
Lichens belonging to Parmeliaceae are highly diversified, but most of them share an extremely conserved morpho-chemical trait: the lower cortex is heavily melanized. The adaptive value of this character is still uncertain. Melanins are ubiquitous compounds found in most organisms since they fulfil several biological functions including defense against UV radiation, oxidizing agents, microbial stress, and metal complexation. This work aims to establish whether melanization can affect the elemental content of lichen thalli. The relative abundance of macro- (Ca, K and S) and micro- (Fe, Mn and Zn) nutrients in melanized and non-melanized pseudotissues of nine species was first evaluated by a non-destructive micro-X-ray fluorescence elemental analysis on either the upper and lower cortex, and on the internal medulla, which was artificially exposed to the mechanical removal of the lower cortex. Afterwards, the total concentration of the same elements was measured in composite samples by inductively coupled plasma atomic emission spectroscopy after acidic digestion. In order to verify whether Fe and Zn are chemically bound to the melanized pseudotissues, a sequential elution experiment was performed on two species: the two-side heavily melanized Melanelixia glabratula and the one-side lightly melanized Punctelia subrudecta. The content of Fe and Zn was higher in the melanized species than in the non-melanized ones. Species deprived of their melanized lower cortex showed a sharp decrease in Fe but not in Zn, suggesting that the melanized lower cortex is involved in Fe complexation, whereas Zn is homogeneously distributed throughout the thallus.
... This genus was reported to contain ca 40 taxa with M. stygia as the type species [1]. However, according to numerous extensive systematic revisions, the number of species in this genus was reduced to four: M. stygia, M. hepatizon, M. agnata and M. pseudoglabra [3][4][5][6][7][8][9][10][11][12][13]. The rest of the formerly assigned Melanelia species are circumscribed into other genera using molecular data, including Melanohalea [3,10,12], Melanelixia [3,10], Cetrariella [4], Montanelia [8][9][10] and Nephromopsis [11]. ...
... However, according to numerous extensive systematic revisions, the number of species in this genus was reduced to four: M. stygia, M. hepatizon, M. agnata and M. pseudoglabra [3][4][5][6][7][8][9][10][11][12][13]. The rest of the formerly assigned Melanelia species are circumscribed into other genera using molecular data, including Melanohalea [3,10,12], Melanelixia [3,10], Cetrariella [4], Montanelia [8][9][10] and Nephromopsis [11]. A recent phylogenetic study [4] supports the placement of the genus Melanelia in the cetrarioid group, instead of the parmelioid group where the genus was originally placed. ...
... However, according to numerous extensive systematic revisions, the number of species in this genus was reduced to four: M. stygia, M. hepatizon, M. agnata and M. pseudoglabra [3][4][5][6][7][8][9][10][11][12][13]. The rest of the formerly assigned Melanelia species are circumscribed into other genera using molecular data, including Melanohalea [3,10,12], Melanelixia [3,10], Cetrariella [4], Montanelia [8][9][10] and Nephromopsis [11]. A recent phylogenetic study [4] supports the placement of the genus Melanelia in the cetrarioid group, instead of the parmelioid group where the genus was originally placed. ...
Taxa in the genus Melanelia (Parmeliaceae, Ascomycota) belong to a group of saxicolous lichens with brown to black foliose thalli, which have recently undergone extensive changes in circumscription. Taxa belonging to Parmeliaceae are prolific producers of bioactive compounds, which have also been traditionally used for chemotaxonomic purposes. However, the chemical diversity of the genus Melanelia and the use of chemical data for species discrimination in this genus are largely unexplored. In addition, identification based on morphological characters is challenging due to few taxonomically informative characters. Molecular identification methods, such as DNA barcoding, have rarely been applied to this genus. This study aimed to identify the Melanelia species from Iceland using DNA barcoding approach, and to explore their chemical diversity using chemical profiling. Chemometric tools were used to see if lichen metabolite profiles determined by LC-MS could be used for the identification of Icelandic Melanelia species. Barcoding using the fungal nuclear ribosomal internal transcribed spacer region (nrITS) successfully identified three Melalenlia species occurring in Iceland, together with Montanelia disjuncta (Basionym: Melanelia disjuncta). All species formed monophyletic clades in the neighbor-joining nrITS gene tree. However, high intraspecific genetic distance of M. stygia suggests the potential of unrecognized species lineages. Principal component analysis (PCA) of metabolite data gave a holistic overview showing that M. hepatizon and M. disjuncta were distinct from the rest, without the power to separate M. agnata and M. stygia due to their chemical similarity. Orthogonal partial least–squares to latent structures–discriminate analysis (OPLS-DA), however, successfully distinguished M. agnata and M. stygia by identifying statistically significant metabolites, which lead to class differentiation. This work has demonstrated the potential of DNA barcoding, chemical profiling and chemometrics in identification of Melanelia species.
... For example, nine genera were synonymized within Xanthoparmelia, four within Parmotrema, three within Hypotrachyna, and more recently, Bulborrhizina within Bulbothrix (reviewed by Crespo et al. 2011; Thell et al. 2012; and Divakar et al. 2013a; Kirika et al. 2015 ). At the same time, molecular phylogenies have helped to uncover previously unrecognized genuslevel lineages such as: Melanelixia Blanco et al. (2004: 881), Melanohalea Blanco et al. (2004) and Montanelia Divakar et al. ( : 2022), each segregated from Melanelia Esslinger (1978: 46) s. lat. (Blanco et al. 2004;). ...
... At the same time, molecular phylogenies have helped to uncover previously unrecognized genuslevel lineages such as: Melanelixia Blanco et al. (2004: 881), Melanohalea Blanco et al. (2004) and Montanelia Divakar et al. ( : 2022), each segregated from Melanelia Esslinger (1978: 46) s. lat. (Blanco et al. 2004;). Other examples include: Austroparmelina Crespo et al. (2010a: 209) which was segregated from Parmelina Hale (1974a: 481); remototrachyna Divakar et al. (2010: 584) segregated from Hypotrachyna Hale (1974b: 340) s. lat. ...
Many phenotypical features traditionally used to classify genera in Parmeliaceae and in lichens in general have evolved several times independently, potentially limiting their taxonomic utility. Here, we aim to elucidate evolutionary relationships of Canoparmelia s. lat. among other parmotremoid taxa. A multilocus dataset (ITS, nuLSU and mtSSU rDNA sequences) was gathered and analyzed within a phylogenetic framework. Canoparmelia s. lat. was recovered as highly polyphyletic within the parmelioid clade, and three divergent lineages representing Canoparmelia s. lat. were identified in addition to the previously segregated Crespoa group. Of these, two formed a sister relationship with Parmotrema. However, no apparent diagnostic morphological features were found distinguishing the distinct Canoparmelia s. lat. clades reconstructed in the phylogenetic analyses. As a consequence, we propose to restrict the circumscription of Canoparmelia to clade 1 (i.e. the C. texana group) and to include clades 2 and 3 in Parmotrema. We propose to recognize these well-supported monophyletic clades at subgeneric level. Consequently, the new subgeneric name Parmotrema subgen. Africanae is proposed for clade 3 recovered in this study. Since clade 4, which clusters with the genera Nesolechia and Punctelia, is only represented by a single sequenced specimen, we refrain from proposing any taxonomic changes. The new combinations Parmotrema epileu-cum, and P. zimbabwense are proposed.
