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Recent studies have demonstrated that species boundaries among the lichen-forming fungi are in need of revision with the discovery of cryptic species in numerous clades, especially in parmelioid lichens. Here we focus on addressing the species boundaries in Canoparmelia texana, a sorediate species with a pantropical distribution that extends into temperate regions. We extracted DNA sequences of the nuclear ribosomal internal transcribed spacer region (ITS), large subunit (nuLSU) and mitochondrial small subunit (mtSSU) from samples mostly collected in Kenya, and analyzed them in a phylogenetic framework. We illustrate that our samples of the species as currently circumscribed do not form a monophyletic group but fall into two distinct clades, with the apotheciate C. nairobiensis nested within. Both of the discovered lineages have a wide distributional range and are common in Kenya, and Parmelia albaniensis C. W. Dodge is resurrected to accommodate one of the clades; consequently a new combination, Canoparmelia albaniensis (C. W. Dodge) Divakar & Kirika comb. nov., is proposed.
The widespread species Parmotrema crinitum (Ach.) M. Choisy and Parmotrema perlatum (Huds.) M. Choisy are mainly distinguished by their reproductive strategies. While P. crinitum propagates by isidia, P. perlatum produces soredia. In this study, we aim to evaluate the phylogenetic relationship between both species and to critically examine their species boundaries. To this purpose, 46 samples belonging to P. crinitum and P. perlatum were used in our analysis, including 22 for which we studied the morphology and chemistry, before extracting their DNA. We used 35 sequences of the internal transcribed spacer region of nuclear ribosomal DNA (ITS) of Parmotrema perlatum from Europe and Africa (20 of which were newly generated), and 11 of Parmotrema crinitum from Europe, North America and North Africa (two newly generated). Additionally, 28 sequences of several species from Parmotrema were included in the ITS dataset. The ITS data matrix was analyzed using different approaches, such as traditional phylogeny (maximum likelihood and Bayesian analyses), genetic distances, automatic barcode gap discovery (ABGD) and the coalescent-based method poisson tree processes (PTP), in order to test congruence among results. Our results indicate that all samples referred to P. crinitum and P. perlatum nested in a well-supported monophyletic clade, but phylogenetic relationships among them remain unresolved. Delimitations inferred from PTP, ABGD and genetic distance analyses were comparable and suggested that P. crinitum and P. perlatum belong to the same lineage. Interestingly, two samples of P. perlatum separate in a different monophyletic clade, which is supported as a different lineage by all the analyses.
The species of the Parmelia saxatilis complex occurring in the Iberian Peninsula were revised. Eight species are accepted, including a new species found in southern Spain, described as P. rojoi A. Crespo, V. J. Rico & Divakar. The new species, which forms a sister-group relationship with P. saxatilis s. str., is rare in the Iberian Peninsula and is restricted to higher altitudes of northern and central Spain. Parmelia rojoi differs from P. saxatilis by generally narrower isidia and a more fragile thallus. The segregation of the new species is also supported by ITS (rDNA) and Mcm7 (MS456) phylogeny and multispecies coalescent-based approaches, including StarBEAST and BP&P. Furthermore, the divergence of P. rojoi is dated back to the Pleistocene, c. 2.13 Ma. A key to the identification of species from the P. saxatilis complex with their diagnostic features is provided. All species of the complex known from Europe are also found in the Iberian Peninsula. We hypothesize that P. rojoi is a relict species that survived the Pleistocene glaciations in refugia in Spain and has been unable to extend its distributional range in postglacial periods.
An account of thelotremoid species of Graphidaceae in India is provided, which includes 124 species in 24 genera. Ocellularia and Thelotrema are the most diverse genera represented by 34 and 18 species, respectively. Type specimens were re-examined and additional samples studied morphologically and chemically. One new species, Ocellularia upretii S. Joshi, Divakar, Lumbsch & Lücking, is described; it is characterized by a greyish green thallus, porinoid ascomata, brown proper exciple, simple, carbonized columella, clear hymenium, transversely septate, amyloid ascospores of 110–125×15–20 µm and an absence of secondary metabolites. Asteristion australianum, Astrochapsa mirabilis, Cruentotrema cruentatum, C. kurandense, Ocellularia violacea and Thelotrema adjectum are reported as new to the country, and Astrochapsa mirabilis, Melanotrema submicrosporoides, Ocellularia annuloelevata, O. subkeralensis and Rhabdodiscus verrucoisidiatus are proposed as new combinations. Diploschistes awasthii, Ocellularia gupeti, O. leucina, O. mahabalei, Thelotrema confertum and T. verrucorugosum are synonymized under D. scruposus, O. neomasonhalei, O. urceolaris, O. thelotremoides, Chapsa leprocarpoides and T. rugatulum, respectively, with Ocellularia canariana and O. verrucomarginata reduced to synonymy with O. allosporoides.
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.
Species richness is not evenly distributed across the tree of life and a limited number of lineages comprise an extraordinarily large number of species. In lichen-forming fungi, only two genera are known to be ‘ultradiverse’ (>500 species), with the most diverse genus, Xanthoparmelia, consisting of c. 820 species. While Australia and South Africa are known as current centres of diversity for Xanthoparmelia, it is not well known when and where this massive diversity arose. To better understand the geographical and temporal context of diversification in this diverse genus, we sampled 191 Xanthoparmelia specimens representing c. 124 species/species-level lineages from populations worldwide. From these specimens, we generated a multi-locus sequence data set using Sanger and high-throughput sequencing to reconstruct evolutionary relationships in Xanthoparmelia, estimate divergence times and reconstruct biogeographical histories in a maximum likelihood and Bayesian framework. This study corroborated the phylogenetic placement of several morphologically or chemically diverse taxa within Xanthoparmelia, such as Almbornia, Chondropsis, Karoowia, Namakwa, Neofuscelia, Omphalodiella, Paraparmelia, Placoparmelia and Xanthomaculina, in addition to improved phylogenetic resolution and reconstruction of previously unsampled lineages within Xanthoparmelia. Our data indicate that Xanthoparmelia most likely originated in Africa during the early Miocene, coinciding with global aridification and development of open habitats. Reconstructed biogeographical histories of Xanthoparmelia reveal diversification restricted to continents with infrequent intercontinental exchange by long-distance dispersal. While likely mechanisms by which Xanthoparmelia obtained strikingly high levels of species richness in Australia and South Africa remain uncertain, this study provides a framework for ongoing research into diverse lineages of lichen-forming fungi. Finally, our study highlights a novel approach for generating locus-specific molecular sequence data sets from high throughput metagenomic reads.
Macro-morphological features traditionally used to segregate genera in Parmeliaceae have been shown to be highly plastic, placing limits on their taxonomic value. Here we aim to elucidate the evolutionary relationships of the genera Relicina and Relicinopsis and reassess the phenotypic features traditionally used to separate these genera. To this end, we gathered ribosomal DNA sequences of ITS, nuLSU and mtSSU and analyzed them in a phylogenetic framework. Relicina was recovered as paraphyletic, with Relicinopsis nested within, and three different clades were identified within Relicina. Alternative hypothesis tests significantly rejected the monophyly of Relicina. Our results indicate that the presence or absence of bulbate cilia is of limited taxonomic value in this clade. Based on differences in conidia, however, we propose to accept Relicinopsis as a subgenus within Relicina as Relicina subgen. Relicinopsis (Elix & Verdon) Kirika, Divakar & Lumbsch. It is proposed that five new combinations of species previously classified in Relicinopsis be placed in Relicina.
The results of the first molecular phylogenetic study of Pseudephebe are presented; a three-locus phylogeny. The genus is confirmed as monophyletic within the alectorioid clade of Parmeliaceae. Two major clades were recovered, which can be assigned to the two traditional taxa, P. minuscula and P. pubescens, with modifications of the species delimitation, especially the variable P. minuscula. These species are cryptic and cannot be confidently distinguished morphologically due to phenotypic convergence. Therefore, the use of P. pubescens aggr. is recommended for samples not molecularly analyzed. Contrary to previous studies, specimens of both species might have indistinct pseudocyphellae and also contain lichen substances; norstictic acid was detected in c. 60% of specimens tested. An SSU 1516 Group I intron is usually present in P. minuscula but always absent in P. pubescens. The species-level nomenclature is summarized and sequenced reference specimens (RefSpec) for both Pseudephebe species are selected. Sequences from Bryoria mariensis established that this name was a synonym of P. minuscula.
Bryoria araucana sp. nov. is described from Chile on the basis of morphological, chemical and molecular data. It has a grey to dark greyish brown pendent thallus with the base usually black, branching angles mainly obtuse, terminal branches with few lateral branchlets acutely inserted, fumarprotocetraric acid, and often protocetraric and confumarprotocetraric acids. It is morphologically similar to the Northern Hemisphere B. trichodes, but lacks soralia and has inconspicuous concolorous or slightly darker pseudocyphellae. Bryoria glabra is also reported for the first time from the Southern Hemisphere. New phylogenetic data based on ITS, mtSSU and MCM7 analyses suggest that Bryoria sect. Bryoria is polyphyletic and needs revision.
In order to confirm and investigate the extent of reported mismatches between chemotypes and molecular sequence data in Bryoria fuscescens s. lat., we examined 15 morphologically similar thalli from each of three Pinus forest sites in the Sistema Central of central Spain. Three thalli were rejected due to infections by Phacopsis huuskonenii (not previously published from Spain). The remaining 42 thalli represented nine ITS rDNA haplotypes and four chemotypes (by TLC): fumarprotocetraric and protocetraric acids; norstictic and connorstictic acids; psoromic acid; and fumarprotocetraric, protocetraric and psoromic acids. The molecular phylogenetic tree was characterized by extremely short branch lengths, often only with a single mutational difference, and a single haplotype could have different chemical products. In some cases, adjacent specimens represented different chemotypes, and three thalli appeared to be mixed individuals. Consistency of both molecular and chemical data within individual specimens was demonstrated by examining four different parts of each thallus, which showed only a difference in the location of psoromic acid in some. This is the first population-level study of this taxon, and so it is premature to propose taxonomic changes at this time. Further populations in different parts of the geographical range of this widespread complex now need to be analyzed, and more sensitive chemical analyses conducted, in order to understand the basis of the variability and determine the appropriate taxonomic treatment.
Recently, the number of cryptic species known has increased considerably, showing that species diversity has in many cases been underestimated in the past. Parmelia sulcata is a widely distributed species and one of the most common taxa in temperate Europe. The first intra-specific molecular studies on P. sulcata showed an unexpectedly high genetic variability. In the present work, we study the biodiversity of this taxon including specimens from four continents and using three molecular markers (nuITS, nuIGS rDNA, and partial β-tubulin gene). Two monophyletic groups of P. sulcata were encountered; one of these is epitypified as P. sulcata s. str and the other one is segregated as the new cryptic species P. encryptata sp. nov. Issues surrounding the lectotypification of Parmelia sulcata have also been elucidated.
This article traces the changing systems in the classification of the parmelioid lichens from the early 19th century to the present day. Molecular phylogenetic approaches have enabled the relative importance of traditionally used characters to be objectively re-assessed, and led to the realization of the significance of others that had previously been passed over or dismissed; for example, differences in ascospore and conidium types or the cell-wall constituents. Information on the different characters employed is presented and assessed. It is concluded that characters related to thallus form or chemical products prove to be less informative at the generic level than has sometimes been assumed. Features requiring further study are identified, and in the light of experience in this group, lichenologists should be cautious in translating their results into formal taxonomies.
We used a genetic distance approach in conjunction with molecular phylogeny to establish species boundaries and detect cryptic lineages in the Parmotrema reticulatum – P. pseudoreticulatum complex. The phylogeny of specimens from a broad geographic distribution was reconstructed from the internal transcribed spacer region. Pairwise genetic distances were calculated and compared to an intraspecific range defined for the parmelioid lichens to circumscribe species-level groups. Our results showed that P. reticulatum and P. pseudoreticulatum are polyphyletic, being comprised of at least seven well-supported lineages. In contrast, the genetic distance approach revealed ten cryptic lineages within the P. reticulatum – P. pseudoreticulatum complex. Neither morphology nor geography was conclusive in attempting to corroborate these genetic lineages. However FST indices suggest significant genetic differentiation between these lineages. Our results suggest that the morphology-based circumscriptions underestimated species in Parmotrema and that, in some cases, genetic distances may be used as an additional tool to determine species boundaries in morphologically cryptic species complexes. The most significant contribution of the present study is the application of a fast and accurate method to identify problematic groups and candidate species using the ITS locus with a genetic distances approach.
Additions and corrections to the checklist published in 2008 are presented. Ten additional species are recognized: Allantoparmelia almquistii, Asahinea chrysantha, Hypogymnia bryophila, Hypotrachyna afrorevoluta, Melanelixia glabratula, Parmelia encryptata, Parmelina atricha, P. cryptotiliacea, Parmotrema cinereopruinata, and Punctelia rudecta. Myelochroa subaurulenta is deleted, and Xanthoparmelia perezdepazii is also noted as newly described from the Canary Islands. In addition, Cavernularia hultenii is now placed in Hypogymnia, the Canoparmelia crozalsiana group is included within Parmotrema, and Parmelinopsis is treated as a synonym of Hypotrachyna. The typification of the name Lichen quercinus is also corrected with the designation of the original figure as lectotype, and the new combination Hypotrachyna cryptochlora comb. nov. (syn. Parmelia cryptochlora) is made.
The cetrarioid core group has been the focus of numerous taxonomic and phylogenetic studies in recent years, yet the phylogenetic resolution and support among these clades remains unclear. Here we use four commonly employed loci to estimate if their use increases phylogenetic resolution and support. The present study largely confirms the topologies of previous studies, but with increased support. Approximately half of the genera in the cetrarioid core were not monophyletic. Melanelia sorediella was clustered within Cetrariella, and the combination Cetrariella sorediella (Lettau) V. J. Rico & A. Thell comb. nov. is made. Additionally, the genus Flavocetrariella was supported as part of Nephromopsis and is considered to be a synonym of the latter. Finally, a comparison of genetic distances shows that the maximum intrageneric genetic distance encompassed by many cetrarioid genera is lower than that of many other genera in Parmeliaceae.
Parmelina tiliacea is a common, widely distributed species in south-western Europe, easily identifiable by morphology and much used as an air pollution bioindicator in many regions. A molecular phylogenetic survey of samples from many geographical areas, using Maximum Parsimony and Bayesian inference of nuITS and mtLSU rDNA regions, revealed a group of samples geographically restricted to a small region of the Iberian Peninsula and genetically separated from the other P. tiliacea specimens studied. These samples are morphologically indistinguishable from P. tiliacea, apart from subtle anatomical characters in the ascomata (hyphae of the exciple and ascospore width), which are frequently absent. Although geographically different, the two taxa occupy similar habitats and are even sympatric in some areas, indicating that they do not exchange genetic material. This previously overlooked, and apparently endemic lineage, is described as P. cryptotiliacea sp. nov., and the name Lichen tiliaceus is epitypified by a sequenced specimen to fix the application of Parmelina tiliacea to the widespread genotype. A second unexpected result was the discovery that the morphologically distinct P. pastillifera was nested within P. tiliacea. These two cases stress the need to use molecular tools to elucidate species concepts even within widespread morphologically well-characterized macrolichens. Such investigations are necessary to improve our understanding and estimation of biodiversity, and to facilitate the development of sound biodiversity conservation strategies for lichens.
Twenty-six species recently described from Thailand are revised. Eleven taxa are reduced to synonymy with previously described species, including Leptotrema phaeosporum var. vainiona Räsänen. The following new combinations are proposed: Chapsa calathiformis (Vain.) Lumbsch & Papong, C. laemensis (Homchantara & Coppins) Lumbsch & Papong, Melanotrema melanophthalmum (Homchantara & Coppins) Lumbsch & Papong, Ocellularia albocincta (Hale) Divakar & Mangold, O. guianensis (Sipman) Divakar & Mangold, O. khunantensis (Homchantara & Coppins) Lumbsch & Papong, O. percolumellata (Sipman) Divakar & Mangold, O. subcalvescens (Nyl.) Divakar & Mangold and Ocellularia subgranulosa (Homchantara & Coppins) Lumbsch & Papong.