Phylogenetic significance of morphological characters in the tropical Hypotrachyna clade of parmelioid lichens (Parmeliaceae, Ascomycota)

Lichen-forming ascomycetes exhibit often complex morphologies of the vegetative thallus that are usually not found in non-lichenized fungi. This includes the thallus organization and appendical structures associated with the main thallus, such as cilia and rhizines. Such morphological characters are widely employed in the taxonomy of parmelioid lichens, especially at generic level. Within parmelioid lichens, several monophyletic groups can be distinguished, the Hypotrachyna clade being one of them, which includes mostly tropical taxa. In this first molecular study focused specifically on the Hypotrachyna clade, we used maximum parsimony and Bayesian analyses of a combined data set of nuclear ITS and mitochondrial SSU rDNA sequences to (1) test the monophyly of genera presently accepted within the clade and (2) evaluate the phylogenetic value of the morphological characters used to circumscribe genera in parmelioid lichens. Out of the 89 mtSSU and 88 nuITS sequences included in the present study, 121 sequences were newly obtained. Our results show that the taxa within the clade fall into two major groups and that the genus Hypotrachyna is polyphyletic. Everniastrum and Parmelinopsis are nested within Hypotrachyna sensu stricto, the latter being also polyphyletic. Bulbothrix is paraphyletic with Parmelinella nested within and is basal to the second major Hypotrachyna clade. Monophylies of Bulbothrix and Hypotrachyna are significantly rejected. The phylogenetic analysis demonstrates that morphological characters currently used to circumscribe genera in parmelioid lichens, such as cortical anatomy, lobe configuration, cilia, and rhizines have been overestimated and have only minor value in identifying monophyletic groups.     

Molecular phylogeny of the Laboulbeniomycetes (Ascomycota)

A first molecular-based phylogeny is presented for the Laboulbeniomycetes, a group of ascomycete fungi that utilize arthropods for nutrition and/or dispersal. Morphological diversification and life-history evolution has made it difficult to resolve relationships within the group, and to identify close relatives. Here, we infer a preliminary phylogeny based on acquisition of 51 new SSU rDNA sequences, representing a total of 65 taxa. The results of this study demonstrate that Laboulbeniomycetes is monophyletic, and related to Sordariomycetes. The class could be divided into at least 4 or 5 orders, though we refrain from formally giving names to these at this stage. Further evidence for the occurrence of asexuality within the class is provided by the inclusion of the genera Chantransiopsis and Tetrameronycha, both known only as asexual taxa with thalli consisting of linearly superposed cells. The precise placement of the genus Herpomyces (Herpomycetaceae), on cockroaches, remains unresolved in our analysis, but lies outside of the main clade of sexually reproducing Laboulbeniales. There is good support for this latter grouping, comprising taxa that are found on both aquatic and terrestrial hosts. Within this large assemblage, we recognize 5 distinct clades (clades E, F, G, H, I). Relationships among the so-called “aquatic genera” (≡ Ceratomycetaceae + some Euceratomycetaceae and Zodiomyces) are poorly resolved in our analyses, accounting for 3 of these clades (E, F, G), with the remainder of the taxa (largely equivalent to Laboulbeniaceae) split into two major groupings (clades H, I). Across all taxa, antheridial characteristics, features on which the earliest classifications were based, are shown to be homoplastic. On the other hand, features of perithecial development show an overall trend towards reduction, and appear to be phylogenetically informative. Morphological characters are identified that support the dichotomy in the Laboulbeniaceae and subclades within the two major groupings are discussed further in light of information on thallus morphology, development, and host relationships.     

Major clades of parmelioid lichens (Parmeliaceae, Ascomycota) and the evolution of their morphological and chemical diversity

Parmelioid lichens comprise about 1500 species and have a worldwide distribution. Numerous species are widely distributed and well known, including important bioindicators for atmospheric pollution. The phylogeny and classification of parmelioid lichens has been a matter of debate for several decades. Previous studies using molecular data have helped to establish hypotheses of the phylogeny of certain clades within this group. In this study, we infer the phylogeny of major clades of parmelioid lichens using DNA sequence data from two nuclear loci and one mitochondrial locus from 145 specimens (117 species) that represent the morphological and chemical diversity in these taxa. Parmelioid lichens are not monophyletic; however, a core group is strongly supported as monophyletic, excluding Arctoparmelia and Melanelia s. str., and including Parmeliopsis and Parmelaria. Within this group, seven well-supported clades are found, but the relationships among them remain unresolved. Stochastic mapping on a MC/MCMC tree sampling was employed to infer the evolution of two morphological and two chemical traits believed to be important for the evolutionary success of these lichens, and have also been used as major characters for classification. The results suggest that these characters have been gained and lost multiple times during the diversification of parmelioid lichens.     

Origin and diversification of major clades in parmelioid lichens (Parmeliaceae, Ascomycota) during the Paleogene inferred by Bayesian analysis

There is a long-standing debate on the extent of vicariance and long-distance dispersal events to explain the current distribution of organisms, especially in those with small diaspores potentially prone to long-distance dispersal. Age estimates of clades play a crucial role in evaluating the impact of these processes. The aim of this study is to understand the evolutionary history of the largest clade of macrolichens, the parmelioid lichens (Parmeliaceae, Lecanoromycetes, Ascomycota) by dating the origin of the group and its major lineages. They have a worldwide distribution with centers of distribution in the Neo- and Paleotropics, and semi-arid subtropical regions of the Southern Hemisphere. Phylogenetic analyses were performed using DNA sequences of nuLSU and mtSSU rDNA, and the protein-coding RPB1 gene. The three DNA regions had different evolutionary rates: RPB1 gave a rate two to four times higher than nuLSU and mtSSU. Divergence times of the major clades were estimated with partitioned BEAST analyses allowing different rates for each DNA region and using a relaxed clock model. Three calibrations points were used to date the tree: an inferred age at the stem of Lecanoromycetes, and two dated fossils: Parmelia in the parmelioid group, and Alectoria. Palaeoclimatic conditions and the palaeogeological area cladogram were compared to the dated phylogeny of parmelioid. The parmelioid group diversified around the K/T boundary, and the major clades diverged during the Eocene and Oligocene. The radiation of the genera occurred through globally changing climatic condition of the early Oligocene, Miocene and early Pliocene. The estimated divergence times are consistent with long-distance dispersal events being the major factor to explain the biogeographical distribution patterns of Southern Hemisphere parmelioids, especially for Africa-Australia disjunctions, because the sequential break-up of Gondwana started much earlier than the origin of these clades. However, our data cannot reject vicariance to explain South America-Australia disjunctions.     

东北大小兴安岭地区梅衣科岛衣类和袋衣类地衣物种多样性

东北大小兴安岭地区是中国地衣的重要分布地之一, 蕴藏着丰富的地衣生物多样性。本研究基于259份标本馆馆藏及采集自东北大小兴安岭31个保护区的新鲜地衣标本, 结合形态学、解剖学、化学和DNA片段的综合分析, 鉴定并报道了该地区梅衣科岛衣类和袋衣类地衣11属31种, 包括环北极成分13种, 东亚成分11种, 世界广布成分3种, 中国特有成分3种, 欧亚成分1种。与之前记录和报道的该地区同类地衣物种组成进行对比, 结果显示隶属于2属的6种地衣在该地区未被重新发现, 表明这些地衣在该地区的分布可能大大萎缩乃至消失, 尤其是属于东亚成分的袋衣属2种: 粉袋衣(Hypogymnia farinacea)和日光山袋衣(H. nikkoensis), 自最初以东北大小兴安岭标本为凭证在中国报道分布后, 30余年未再新增任何标本信息, 日光山袋衣在近期中国大型真菌红色名录评估中被推测处于易危状态。     

Canoparmelia scrobicularis belongs to the genus Crespoa (Parmeliaceae,lichenized Ascomycota)

Canoparmelia scrobicularis is shown to be a member of the genus Crespoa and the new combination Crespoa scrobicularis is proposed. The species is described in detail with a discussion of its relationships to other species of Crespoa, especially Crespoa carneopruinata and Crespoa crozalsiana.     

Understanding Phenotypical Character Evolution in Parmelioid Lichenized Fungi (Parmeliaceae,Ascomycota)

Parmelioid lichens form a species-rich group of predominantly foliose and fruticose lichenized fungi encompassing a broad range of morphological and chemical diversity. Using a multilocus approach, we reconstructed a phylogeny including 323 OTUs of parmelioid lichens and employed ancestral character reconstruction methods to understand the phenotypical evolution within this speciose group of lichen-forming fungi. Specifically, we were interested in the evolution of growth form, epicortex structure, and cortical chemistry. Since previous studies have shown that results may differ depending on the reconstruction method used, here we employed both maximum-parsimony and maximum-likelihood approaches to reconstruct ancestral character states. We have also implemented binary and multistate coding of characters and performed parallel analyses with both coding types to assess for potential coding-based biases. We reconstructed the ancestral states for nine well-supported major clades in the parmelioid group, two higher-level sister groups and the ancestral character state for all parmelioid lichens. We found that different methods for coding phenotypical characters and different ancestral character state reconstruction methods mostly resulted in identical reconstructions but yield conflicting inferences of ancestral states, in some cases. However, we found support for the ancestor of parmelioid lichens having been a foliose lichen with a non-pored epicortex and pseudocyphellae. Our data suggest that some traits exhibit patterns of evolution consistent with adaptive radiation.     

Molecular phylogeny of Heteroplacidium, Placidium, and related catapyrenioid genera (Verrucariaceae, lichen-forming Ascomycota)

? Premise of the study: Verrucariaceae is a fascinating lineage of lichenized fungi for which generic and species delimitation is problematic due to the scarcity of discriminating morphological characters. Members of this family inhabit rocks, but they further colonize soils, barks, mosses, and other lichens. Our aim is to contribute to the DNA-based inference of the Verrucariaceae tree of life and to investigate characters that could be useful for proposing a more natural classification. We focused on catapyrenioid genera, which are often part of biological soil crusts, a cryptogam-dominated ecosystem contributing to soil formation and stabilization in arid environments. Understanding their evolution and taxonomy is essential to assess their roles in these fragile and important ecosystems. ? Methods: A multigene phylogeny of Verrucariaceae including catapyrenioid genera is presented. We further examined the phylogenetic relationships among members of Heteroplacidium and Placidium. The evolution of selected characters was inferred using the latter phylogeny. ? Key results: Anthracocarpon and Involucropyrenium were closely related to Endocarpon. Placidium comprised two monophyletic clades sister to Heteroplacidium. Inferred ancestral states of diagnostic characters revealed that the type of medulla and the pycnidia location were homoplasious within the Placidium clade. In contrast, the presence of rhizines was a synapomorphy for Clavascidium. ? Conclusions: Our results provide new information on the usefulness of characters for delineating groups in Verrucariaceae. Taxonomic changes are proposed to reflect more natural groupings: Heteroplacidium podolepis is transferred to Placidium, and Clavascidium is recognized as a different genus. Eight new combinations are proposed for Clavascidium.     

A new sorediate species of Menegazzia (Parmeliaceae,lichenized Ascomycota) from Chile

Abstract:Menegazzia wandae Bjerke from Chile is newly described. It is characterized by lacerate soralia that are formed on laminar pustules; the sinuous, rather imbricate lobes with brown-black margins, and the presence of thamnolic acid in the medulla. Apothecia are unknown, and pycnidia are rare. The species is known from Chile between latitudes 38°30′S and 46°40′S.     

Remototrachyna, a newly recognized tropical lineage of lichens in the Hypotrachyna clade (Parmeliaceae, Ascomycota), originated in the Indian subcontinent

Biogeographical studies of lichens used to be complicated because of the large distribution ranges of many species. Molecular systematics has revitalized lichen biogeography by improving species delimitation and providing better information about species range limitations. This study focuses on the major clade of tropical parmelioid lichens, which share a chemical feature, the presence of isolichenan in the cell wall, and a morphological feature, microscopic pores in the uppermost layer. Our previous phylogenetic studies revealed that the largest genus in this clade, Hypotrachyna, is polyphyletic with a clade mainly distributed in South and East Asia clustering distant from the core of the genus. To divide the Hypotrachyna clade into monophyletic groups and to reevaluate morphological and chemical characters in a phylogenetic context, we sampled ITS, nuclear large subunit (nuLSU) and mitochondrial small subunit (mtSSU) rDNA sequences from 77 species. We are erecting the new genus Remototrachyna for a core group of 15 former Hypotrachyna species. The segregation of Remototrachyna from Hypotrachyna receives support from morphological and chemical data, as well from maximum parsimony, maximum likelihood, and Bayesian phylogenetic analyses of the DNA. We used a likelihood approach to study the geographic range evolution of Remototrachyna and Bulbothrix, which are sister groups. This analysis suggests that the ancestral range of Remototrachyna was restricted to India and that subsequent long-distance dispersal is responsible for the pantropical occurrence of two species of Remototrachyna.     

Neogene diversification in the temperate lichen-forming fungal genus Parmelia (Parmeliaceae,Ascomycota)

The lichen-forming genus Parmelia Acharius occurs worldwide but its centre of distribution is in the northern hemisphere and it is widespread in boreal-temperate Eurasia and North America. Recent molecular work on Parmelia has identified phylogenetic relationships within two major groups of the genus: P. saxatilis s. lat. and P. sulcata s. lat. However, little is known about the diversification and historical biogeography of these groups. Here we have used a dataset of two genetic markers and 64 samples to estimate phylogenetic relationships within Parmelia. The dated phylogeny provides evidence for major diversification during the Neogene and Pleistocene. These diversification events are probably correlated with climatic changes during these periods. Evidence of gene flow within species between populations from North America and Europe has been found in three species: P. sulcata Taylor, P. saxatilis (L.) Acharius and P. barrenoae Divakar, M.C. Molina & A. Crespo. Cryptic species recently segregated on the basis of molecular differences (P. encryptata A. Crespo, Divakar & M.C. Molina vs. P. sulcata and P. saxatilis vs. P. mayi Divakar, A. Crespo & M.C. Molina) do not share a common ancestor. Moreover, the P. saxatilis complex is remarkably diverse. Two morphotypes of P. saxatilis s. lat. were shown to represent independent monophyletic lineages. Consequently, two species (P. sulymae Goward, Divakar, & M.C. Molina & A. Crespo and P. imbricaria Goward, Divakar, M.C. Molina & A. Crespo) are newly described here.     

Molecular phylogeny of Acarosporaceae (Ascomycota) with focus on the proposed genus Polysporinopsis

The molecular phylogeny of Acarosporaceae with a focus on the recently proposed genus Polysporinopsis was investigated using maximum parsimony and Bayesian analyses, using nuITS-LSU and mtSSU rDNA sequence datasets. A well-supported monophyletic clade corresponding to Acarospora (including the type species A. schleicheri, A. fuscata, A. nitrophila, A. rugulosa, A. bullata, A. sinopica, A molybdina and A. peliscypha) was present in all analyses. Acarospora as currently delimited is not monophyletic; neither A. smaragdula nor A. badiofusca belongs to the genus in the restricted sense. Polysporinopsis, which comprises three species previously classified in Acarospora (P. sinopica-type species, P. smaragdula, and P. rugulosa) is not a monophyletic group separate from Acarospora s. str. Acarospora sinopica and A. smaragdula are not closely related; A. sinopica belongs to Acarospora s. str., but A. smaragdula is one of the most basal taxa currently known in Acarosporaceae.     

同岛衣属一新种—粉头同岛衣(梅衣科,子囊菌门)（英文）

描述了采自中国四川的同岛衣属新种粉头同岛衣,提供了新种的特征提要与描述并附有新种原型的形态及解剖图片共4张。新种与该属所有已知种之区别在于其裂片末端具头状粉芽堆。     

New species and new records of Xanthoparmelia (lichenized Ascomycota,Parmeliaceae) from Western Australia

The following new species of Xanthoparmelia (Parmeliaceae) are described from Western Australia: X. cranfieldii Elix, X. elevata Elix, X. kimberleyensis Elix, X. monadnockensis Elix,X. nanoides Elix, X. nortegeta Elix, X. sleei Elix and X. toolbrunupensis Elix. In addition, X. luminosa (Elix) Hale and X. victoriana Elix & J. Johnst. are reported from Western Australia for the first time.     

Surviving climate changes: high genetic diversity and transoceanic gene flow in two arctic–alpine lichens,Flavocetraria cucullata and F. nivalis (Parmeliaceae,Ascomycota)

Aim We examined genetic structure and long‐distance gene flow in two lichenized ascomycetes, Flavocetraria cucullata and Flavocetraria nivalis, which are widespread in arctic and alpine tundra. Location Circumpolar North. Methods DNA sequences were obtained for 90 specimens (49 for F. cucullata and 41 for F. nivalis) collected from various locations in Europe, Asia and North America. Sequences of the nuclear internal transcribed spacer (ITS) + 5.8S ribosomal subunit gene region were generated for 89 samples, and supplemented by beta‐tubulin (BTUB) and translation elongation factor 1‐alpha gene (EF1) sequences for a subset of F. cucullata specimens. Phylogenetic, nonparametric permutation methods and coalescent analyses were used to assess population divergence and to estimate the extent and direction of migration among continents. Results Both F. cucullata and F. nivalis were monophyletic, supporting their morphology‐based delimitation, and had high and moderately high intraspecific genetic diversity, respectively. Clades within each species contained specimens from both North America and Eurasia. We found only weak genetic differentiation among North American and Eurasian populations, and evidence for moderate to high transoceanic gene flow. Main conclusions Our results suggest that both F. cucullata and F. nivalis have been able to migrate over large distances in response to climatic fluctuations. The high genetic diversity observed in the Arctic indicates long‐term survival at high latitudes, whereas the estimated migration rates and weak geographic population structure suggest a continuing long‐distance gene flow between continents that has prevented pronounced genetic differentiation. The mode of long‐distance dispersal is unknown, but wind dispersal of conidia and/or ascospores is probably important in the open arctic landscapes. The high genetic diversity and efficient long‐distance dispersal capability of F. cucullata and F. nivalis suggest that these species, and perhaps other arctic lichens as well, will be able to track their potential niche in the changing Arctic.     

Biogeography and Genetic Structure in Populations of a Widespread Lichen (Parmelina tiliacea,Parmeliaceae, Ascomycota)

The genetic diversity and population structure of the foliose lichenized fungus Parmelina tiliacea has been analyzed through its geographical range, including samples from Macaronesia (Canary Islands), the Mediterranean, and Eurosiberia. DNA sequences from the nuclear ribosomal internal transcribed spacer, the mitochondrial large subunit ribosomal RNA gene, and the translation elongation factor 1-α were used as molecular markers. The haplotypes of the three markers and the molecular variance analyses of multilocus haplotypes showed the highest diversity in the Canary Islands, while restricted haplotypes occurred at high frequencies in Mediterranean coastal samples. The multilocus haplotypes formed three unevenly distributed clusters (clusters 1-3). In the Canary Islands all the haplotypes were present in a similar proportion, while the coastal Mediterranean sites had almost exclusively haplotypes of cluster 3; cluster 2 predominated in inland Mediterranean sites; and cluster 1 was more abundant in central and northern Europe (Eurosiberian area). The distribution of clusters is partially explained by climatic factors, and its interaction with local spatial structure, but much of the variation remains unexplained. The high frequency of individuals in the Canary Islands with haplotypes shared with other areas suggests that could be a refugium of genetic diversity, and the high frequency of individuals of the Mediterranean coastal sites with restricted haplotypes indicates that gene flow to contiguous areas may be restricted. This is significant for the selection of areas for conservation purposes, as those with most genetic variation may reflect historical factors and biological properties of the species.     

中国大陆的梅衣科地衣V.球针叶属和球针黄叶属

球针叶属Bulbothrix和球针黄叶属Relicina地衣区别于其它梅衣的主要特征是裂片边缘具有膨大似葱头状的缘毛.球针叶属地衣的上表面灰色,含有黑茶渍素(atranorin),而球针黄叶属地衣的上表面为黄绿色含有松萝酸(usnic acid).经过形态学与化学研究,承认中国大陆有球针叶属地衣5种和球针黄叶属地衣2种.其中戈氏球叶B.goebelii,烟草球针叶B. tabacina,玄球针黄叶R.abstrusa和悉尼球针黄叶R.sydneyensis等4种为中国大陆首次记载.云南球针叶B.yunnana为中国特有种.本文报道的7个种均出现在中国长江以南地区,这与球针叶属和球针黄叶属地衣呈热带,亚热带的世界分布格局相吻合.     

Lichen chemistry is concordant with multilocus gene genealogy in the genus Cetrelia (Parmeliaceae,Ascomycota)

The lichen genus Cetrelia represents a taxonomically interesting case where morphologically almost uniform populations differ considerably from each other chemically. Similar variation is not uncommon among lichenized fungi, but it is disputable whether such populations should be considered entities at the species level. Species boundaries in Cetrelia are traditionally delimited either as solely based on morphology or as combinations of morpho- and chemotypes. A dataset of four nuclear markers (ITS, IGS, Mcm7, RPB1) from 62 specimens, representing ten Cetrelia species, was analysed within Bayesian and maximum likelihood frameworks. Analyses recovered a well-resolved phylogeny where the traditional species generally were monophyletic, with the exception of Cetrelia chicitae and Cetrelia pseudolivetorum. Species delimitation analyses supported the distinction of 15 groups within the studied Cetrelia taxa, dividing three traditionally identified species into some species candidates. Chemotypes, distinguished according to the major medullary substance, clearly correlated with clades recovered within Cetrelia, while samples with the same reproductive mode were dispersed throughout the phylogenetic tree. Consequently, delimiting Cetrelia species based only on reproductive morphology is not supported phylogenetically. Character analyses suggest that chemical characters have been more consistent compared to reproductive mode and indicate that metabolite evolution in Cetrelia towards more complex substances is probable.