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Photobionts and epiphytes of Micarea and Placynthiella species in 4-weeks-old cultures: (a) cell packages of Interfilum sp.; (b) the filament of Interfilum massjukiae ; (c) schematical drawing of Radiococcus signiensis (the cells are covered with mucilage), the primary photobiont of Placynthiella icmalea and P. uliginosa ; (d) the fragment of filaments of epiphytic alga Trentepohlia umbrina ; (e) schematical drawing of Pseudochlorella sp., the primary photobiont of Placynthiella dasaea ; (f) schematical drawing of Pseudococcomyxa sp., the primary photobiont of Micarea prasina . Scale = 20 μm.
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The photobionts of 22 specimens of Placynthiella and Micarea genera were identified. The photobionts of Placynthiella dasaea (Stirt.) Khodosovtsev, P. icmalea (Ach.) Coppins & P. James, Micarea melanobola (Nyl.) Coppins and M. misella (Nyl.) Hedl. are reported for the first time. This is also the first report about Elliptochloris reniformis (Watana...
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Context 1
... seven specimens of Placynthiella uligi- nosa, as well as in two specimens of P. icmalea, the photobiont Radiococcus signiensis was discovered (Fig. 2c). The abundance of this alga in lichen thalli differed in different speci- mens. The cells of Radiococcus signiensis in the specimen of Placynthiella uliginosa (No. 10) visually presented more than 80% from a total photobiontal mass, while in the specimen of P. uliginosa (No. 6) -it was approximately 50%, and in P. uliginosa (No. 9) ...
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... in photobiont layer in smaller quantity, rather than with the primary photobiont Radiococcus signiensis. The number of these algae and their species composition varied. In most cases, additionally to Radiococ- cus signiensis the thalli of Placynthiella uliginosa also contained Elliptochloris subsphaerica (Fig. 3c, d) and Interfilum massjukiae (Fig. 2b). Rarely the members of Asterochloris and Trebouxia genera were found. One specimen (No. 10) con- tained Leptosira cf. thrombii. However, not all of the investigated species of Placynthiella were associated with Radiococcus. Both specimens of P. dasaea were associated with the photobiont Pseudochlorella sp. (Fig. 1c, ...
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... the thalli of Micarea melanobola, M. misella and M. prasina (No. 15, 16, 17, 18, 21) several additional photobionts were discovered (see Table 2). For instance, Neocystis sp. was found and recognized as an additional photobiont of M. misella; Pseudococcomyxa sp. (Fig. 2f) -as additional photobiont of Micarea melanobola, M. misella and M. prasina (No. 20). The majority of the specimens also contained Interfilum sp. (Fig. 2a), which differs from Interfilum massjukiae by the absence of distinct filaments in culture conditions. ...
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... 16, 17, 18, 21) several additional photobionts were discovered (see Table 2). For instance, Neocystis sp. was found and recognized as an additional photobiont of M. misella; Pseudococcomyxa sp. (Fig. 2f) -as additional photobiont of Micarea melanobola, M. misella and M. prasina (No. 20). The majority of the specimens also contained Interfilum sp. (Fig. 2a), which differs from Interfilum massjukiae by the absence of distinct filaments in culture conditions. ...
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... of the investigated epiphytic algae are very common terrestrial algae. The species of Apatococcus, Bracteacoccus, Parietochloris and Trentepohlia (Fig 2d) are common in aerophytic habitats: tree-bark and rocks (Ettl & Gärtner, 1995;Gärtner & Stoyneva, 2003;Mikhailyuk et al., 2003). Radiococcus signiensis is the epibryo- phyte (Ettl & Gärtner, 1995). ...
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Citations
... For the lichen genus Roccellinastrum, for example, a 'micareoid' green algal photobiont has been stated without clarifying the meaning of this term or its assignment to a green algal genus (Coppins 1983). Since 1983, this description repeatedly appears for lichen photobionts (Coppins and Spribille 2004;Yahr et al. 2015;Kantvilas 2017;Launis et al. 2019), but it has only been assumed to refer to Diplosphaera chodatii based on the morphological description of the term (Voytsekhovich et al. 2011). ...
... This genus encompasses free-living and/or lichenized algae with lobed chloroplasts and that reproduce by forming zoospores with two subapical isokont flagella that emerge symmetrically near the flattened apex (Škaloud et al. 2016). Future studies including molecular phylogenies will show if the term micareoid will always refer to Symbiochloris or to Diplosphaera chodatii as assumed earlier by Voytsekhovich et al. (2011) solely based on morphological characteristics. As only a few species of Symbiochloris have been described it can already be drawn from our phylogeny that the photobiont of R. spongoideum represents an unknown, yet to be described species. ...
Some deserts on Earth such as the Namib or the Atacama are influenced by fog which can lead to the formation of local fog oases-unique environments hosting a great diversity of specialized plants and lichens. Lichens of the genera Ramalina, Niebla or Het-erodermia have taxonomically been investigated from fog oases around the globe but not from the Atacama Desert, one of the oldest and driest deserts. Conditioned by its topography and the presence of orographic fog, the National Park Pan de Azúcar in the Atacama Desert is considered to be such a lichen hotspot. Applying multi-gen loci involving phylogenetic analyses combined with intense morphological and chemical characterization, we determined the taxonomic position of five of the most abundant epiphytic lichens of this area. We evaluated Roccellinastrum spongoideum and Hetero-dermia follmannii which were both described from the area but also finally showed that the genus Cenozosia is the endemic sister genus to Ramalina, Vermilacinia, Namibialina and Niebla. As a result, we have described the species Heterodermia adunca, C. cava and C. excorticata as new lichen species. This work provides a comprehensive dataset for common fog lichen genera of the Coastal Range of the Atacama Desert that can be used as a baseline for monitoring programs and environmental health assessments.
... The surface of mature blastidia/ isidia in Placynthiella shows a paraplectenchymatic cellular pattern in outer view unlike the hyphal pattern in vegetative propagules of Japewia. The photobiont in Placynthiella dasaea forms regular, densely packed colonies of pairs/tetrads of algal cells, with the youngest daughter cells often closely attached to each other and belongs to the genus Pseudochlorella (Voytsekhovich et al. 2011). This photobiont is different from the variously sized and less-organized Trebouxia-like algal partner in Japewia gyrophorica, where the algal daughter cells are soon visibly separated by hyphae of the mycobiont. ...
Molecular identification of organisms is now a common practice and, increasingly, species are identified from environmental samples. However, for most organisms, we still lack comprehensive reference databases of DNA barcodes to identify the sequences produced. We present a near-complete database of ITS and mtSSU barcodes, named Martin7, for accurate molecular identification of epiphytic lichens (mycobionts) of central Europe. New data were obtained by Sanger and PacBio sequencing. We obtained 907 ITS sequences from 603 species and 844 mtSSU sequences from 546 species and supplemented our dataset with sequences from other reliable sources. In total, 1,172 species are included in the database, 1,004 for the ITS barcode and 906 for mtSSU. ITS was newly sequenced for 224 species and mtSSU for 234 species. For 45 genera these are the first ITS or mtSSU (or both) barcodes ever obtained. In most cases, these barcodes distinguish species as currently circumscribed, but we detected 82 groups or pairs of species where at least one of the barcodes (mostly mtSSU) does not clearly discriminate between species. We revealed diverging genotypes, possibly representing cryptic taxa, within 37 traditionally conceived species. By sequencing phenotypically unidentifiable lichens, we detected numerous “known-unknowns” (presumed undescribed species), especially in the genera Bacidina and Micarea. Five species of sorediate crustose lichens are newly described in the genera Bacidina (two species), Chrysothrix, Japewia and Lecanora. We provide a number of taxonomic novelties, for example that Lecidea betulicola and L. coriacea are teleomorphs of Cheiromycina, and Dictyocatenulata is an anamorph of Thelenella.
... If acquisition of additional photobionts is indeed a common occurrence in the course of lichen development, lichen thalli may be expected to contain a heterogeneous photobiont population, at least at certain stages. Some authors have observed and illustrated quite different chlorobionts occurring together within single thalli (Voytsekhovich et al. 2011). Data from molecular markers have also addressed this question. ...
... It is sister to clades of the widely distributed Klebsormidium (Rindi et al. 2011). Interfilum was reported by Voytsekhovich et al. (2011) as a secondary photobiont within the algal layer of Micarea and Placynthiella thalli collected in Ukraine, based on light microscopic examination of thalli and cultured isolates. The principal photobionts in those lichens were reported to be Elliptochloris and Radiococcus, respectively. ...
... Cultures assigned to Neocystis as well as other genera were recently reviewed with molecular sequence analyses, revealing considerable taxonomic redundancy assigned to only two closely related, genetically distinct but morphologically plastic species (Eliáš et al. 2013). An alga identified as Neocystis sp. was cited as 'additional photobiont' of Micarea misella, in thalli having Elliptochloris bilobata as principal photobiont (Voytsekhovich et al. 2011 Phycopeltis Millardet -Members of this trentepohliaceous genus are most often seen as coppery orange discs a few mm in diameter on leaf surfaces in humid subtropical and tropical regions, with one or two species extending to cooler regions such as oceanic Europe (Rindi et al. 2004). Thallus discs consist of a monostromatic layer of closely appressed, bifurcating filaments (Fig. 6A). ...
A review of algal (including cyanobacterial) symbionts associated with lichen-forming fungi is presented. General aspects of their biology relevant to lichen symbioses are summarized. The genera of algae currently believed to include lichen symbionts are outlined; approximately 50 can be recognized at present. References reporting algal taxa in lichen symbiosis are tabulated, with emphasis on those published since the 1988 review by Tschermak-Woess, and particularly those providing molecular evidence for their identifications. This review is dedicated in honour of Austrian phycologist Elisabeth Tschermak-Woess (1917–2001), for her numerous and significant contributions to our knowledge of lichen algae (some published under the names Elisabeth Tschermak and Liesl Tschermak).
... Micarea melanobola is confirmed from Finland (Launis et al. 2019a), Sweden (Kantelinen et al. 2021), and possibly also found in Estonia and Ukraine (Czarnota 2007, Voytsekhovich et al. 2011). ...
We report new records of 19, predominantly rare, Micarea species, mostly from dead wood in mixed montane forests characterized mainly by Norway spruce, European beech and silver fir in the Bavarian Forest National Park on the German-Czech border. Their ecology and key morphological features are discussed. Micarea contexta, M. fallax, M. melanobola, M. pseudomicrococca, M. pusilla, M. soralifera and M. tomentosa are reported for the first time from Germany. Micarea anterior, M. byssacea, M. elachista, M. laeta, M. micrococca, M. nigella and M. nowakii, in addition to the aforementioned, are reported as new for the Bavarian Forest National Park.
... Colony-or packet-forming algae, like Interfilum massjukiae, create layered structures where upper layers, usually exposed to harsh environmental conditions, protect underlying cells [66]. I. massjukiae was originally described from Crimea Mountains as an epilith [67] and later found as a phycobiont of lichens [68]. Colony-or packet-like morphotype is considered as an adaptation of algal cells to retain cellular water under dry conditions [69], which corresponds to the desert habitat of PA. ...
Biocrusts are associations of various prokaryotic and eukaryotic microorganisms in the top millimeters of soil, which can be found in every climate zone on Earth. They stabilize soils and introduce carbon and nitrogen into this compartment. The worldwide occurrence of biocrusts was proven by numerous studies in Europe, Africa, Asia and North America, leaving South America understudied. Using an integrative approach, which combines morphological and molecular characters (small subunit rRNA and ITS region), we examined the diversity of key biocrust photosynthetic organisms at four sites along the latitudinal climate gradient in Chile. The most northern study site was located in the Atacama Desert (arid climate), followed by open shrubland (semiarid climate), a dry forest region (Mediterranean climate) and a mixed broad leaved-coniferous forest (temperate climate) in the south. The lowest species richness was recorded in the desert (18 species), whereas the highest species richness was observed in the Mediterranean zone with (40 species). Desert biocrusts were composed exclusively of single-celled Chlorophyta algae, followed by cyanobacteria. Chlorophyta, Streptophyta and cyanobacteria dominated semiarid biocrusts, whereas Mediterranean and temperate Chilean biocrusts were composed mostly of Chlorophyta, Streptophyta and Ochrophyta. Our investigation of Chilean biocrust suggests high biodiversity of South American biocrust phototrophs.
... The Elliptochloris sequences generated here grouped with a sequence from E. perforata (Fig. 1), a species known as a photobiont in Micaria prasina (Yahr et al., 2015). Other species of Elliptochloris associate with lichenized fungi in genera such as Catillaria, Micarea and Verrucaria, which all form crustose lichens (Thüs et al., 2011;Voytsekhovich et al., 2011;Dal Grande et al., 2014), and some have even been found as symbionts in sea anemones (Letsch et al., 2009;Gustavs et al. 2017). Species of Elliptochloris are also known to occur free-living (Darienko et al., 2016). ...
... JM 8962). Since the thallus of Placynthiella is known to provide a suitable niche for rich algal communities (both symbiotic and epiphytic algae;Voytsekhovich et al. 2011) and it may also be overgrown by other Epigloea species(Czarnota & Hernik 2013, Döbbeler 1994), we suggest Epigloea urosperma may not be necessarily an obligate lichenicolous fungus, exclusively associated with Placynthiella spp. Bohemia, Český les Mts, Bělá nad Radbuzou, Rybník: Malý Zvon Nature Reserve, old-growth beech forest on steep E-facing slope of Mt Malý Zvon, 49°32'06''N, 12°38'40''E, alt. ...
This contribution presents new records of lichenized and “lichen-allied” fungi for the Czech Republic and a list of all recently published species missing in the last national checklist (Liška & Palice 2010). Lecanora tephraea is supposed to be synonymous with L. cenisia and the lectotype is designated here. Polyblastia brunnensis is synonymized with Thelidium zwackhii. Caloplaca fiumana, lectotypified here, was found to be an older name of the recently described taxon Caloplaca substerilis subsp. orbicularis. Candelariella subdeflexa is replaced by C. blastidiata in the national checklist; Lecanora reagens is excluded from the Czech lichen biota. Twenty nine species are published as new to the Czech Republic: Absconditella rubra, Alyxoria ochrocheila, Aspicilia verrucigera, Blastenia hungarica, Carbonicola anthracophila, Chaenothecopsis montana, C. savonica, Epigloea pleiospora, E. urosperma, Gyalecta ophiospora, Lecanora epibryon, L. flavoleprosa, L. silvae-nigrae, L. stenotropa, Leptorhaphis maggiana, Micarea tomentosa, Myriolecis perpruinosa, Ochrolechia mahluensis, Parmelia serrana, Peltigera ponojensis, Pertusaria borealis, Placynthium caesium, Protoblastenia lilacina, Ramalina europaea, Rinodina trevisanii, Strigula glabra, Verrucaria subcincta, Xanthomendoza huculica and Xylographa soralifera. Including the cited taxa, the lichen biota of the Czech Republic currently comprises 1691 taxa. Key words: Biodiversity, boreal lichens, checklist, lichen-forming fungi, microlichens.
... Lichen phycobionts have been studied in only a limited number of lichen associations (Honegger 2008;Voytsekhovich et al. 2011;Yahr et al. 2015), particularly relative to estimated lichen diversity (Lumbsch et al. 2011;Lücking et al. 2016). In many cases, studies of algal diversity in lichens are constrained by limited availability of lichen material, and research is often focused on those associations represented by the most conspicuous macrolichen lineages (e.g. ...
... The authors, however, stressed that morphological features within a single species-level lineage may vary based on environmental and culture conditions and phenotypic data, and must be based on a wide sampling(Darienko et al. 2015).Members of Elliptochloris, with seven currently accepted species (Guiry & Guiry 2017), generally have spherical vegetative cells and two types of autospores, and consistently lack mucilage layers (Tschermak-Woess 1980b). Members of this genus have been reported to associate with a few species in the lichenised mycobiont genera Catolechia Flotow, Placynthiella Elenkin and Micarea E.M.Fries, as well as with the species Verrucaria sublobulata Eitner ex Servít(Tschermak-Woess 1980b; Thüs et al. 2011;Voytsekhovich et al. 2011;Yahr et al. 2015).Coccomyxa/Pseudococcomyxa and Elliptochloris all bear characteristic cell walls rich in sporopollenin-like polymers; the cell walls are three-layered in Coccomyxa, conferring high resistance to degradation(Honegger 1982). Due to the type of cell wall, fungal haustoria cannot penetrate the algal cells in the symbiotic stage(Brunner & Honegger 1985;Muggia et al. 2011), and it has been speculated that this is why very few fungi lichenise with these microalgae.APATOCOCCUS: Recent phylogenetic and morphological analyses confirmed the identity of Apatococcus (Chodat) G.B.Petersen as the phycobiont of the lichen-forming fungal genus Fuscidea V.Wirth & Veˇzda (Zahradníkova´et al. 2017). ...
Phycologia: 2018, Vol. 57, No. 5, pp. 503-524. www.phycologia.org
https://doi.org/10.2216/17-134.1
The class Trebouxiophyceae is comprised of coccoid to ellipsoid unicells, filaments, blades and colony-forming species of green algae occurring in diverse terrestrial and aquatic environments. Some representatives have evolved parasitic heterotrophic lifestyles, others have been investigated for their biotechnological potential and others have evolved as integral components of lichen symbioses. In this review, we provide an overview of the current understanding of diversity, taxonomy and evolutionary context for the important lichen-forming algal class Trebouxiophyceae (Chlorophyta). In particular, we focus on members of the family Trebouxiaceae (Trebouxiales), the best-known, most widespread and most species-rich group of terrestrial, lichenised green algae. Recent investigations on the diversity of lichen phycobionts demonstrate the importance of implementing integrative taxonomic approaches. Therefore, combining analyses of morphological and anatomical traits with genetic data has improved our perspective of diversity in lichenised algae. More accurate recognition of diversity in Trebouxiophyceae will enhance our understanding of phylogenetic relationships and trait evolution, specimen identification in genomic and meta–bar-coding studies and patterns of specificity and selectivity among the lichen symbionts. We conclude with a discussion of the roles and transformative potential of high-throughput sequencing in research related to lichen-associated algae. http://joionline.org/
... Furthermore, the presence of Heveochlorella in thalli of may partner with these photobionts as well. Preference at the family level will not be absolute; the Pilocarpaceae also include the non-foliicolous genus Micarea, whose species lichenize algae of Elliptochloris and Coccomyxa (Voytsekhovich et al. 2011a, Yahr et al. 2015). Yet if further sampling worldwide among these fungal families, and others that share their habitats, continues to reveal partnership with Heveochlorella, this algal taxon may prove to rank among the major eukaryotic photobiont genera. ...
Foliicolous lichens are formed by diverse, highly specialized fungi that establish themselves and complete their life cycle within the brief duration of their leaf substratum. Over half of these lichen-forming fungi are members of either the Gomphillaceae or Pilocarpaceae, and associate with Trebouxia-like green algae whose identities have never been positively determined. We investigated the phylogenetic affinities of these photobionts to better understand their role in lichen establishment on an ephemeral surface. Thallus samples of Gomphillaceae and Pilocarpaceae were collected from foliicolous communities in southwest Florida and processed for sequencing of photobiont marker genes, algal cultivation and/or TEM. Additional specimens from these families and also from Aspidothelium (Thelenellaceae) were collected from a variety of substrates globally. Sequences from rbcL and nuSSU regions were obtained and subjected to Maximum Likelihood and Bayesian analyses. Analysis of 37 rbcL and 7 nuSSU algal sequences placed all photobionts studied within the provisional trebouxiophycean assemblage known as the Watanabea clade. All but three of the sequences showed affinities within Heveochlorella, a genus recently described from tree trunks in East Asia. The photobiont chloroplast showed multiple thylakoid stacks penetrating the pyrenoid centripetally as tubules lined with pyrenoglobuli, similar to the two described species of Heveochlorella. We conclude that Heveochlorella includes algae of potentially major importance as lichen photobionts, particularly within (but not limited to) foliicolous communities in tropical and subtropical regions worldwide. The ease with which they may be cultivated on minimal media suggests their potential to thrive free-living as well as in lichen symbiosis. This article is protected by copyright. All rights reserved.
... We use the term Bprimary photobiont^for the main and predominant alga that forms a constant, steady and reproducible association with a certain lichen-forming fungus. In contrast, Badditional photobionts^(sometimes called or accessory photobionts) are algae that are different from the main photobiont morphologically, taxonomically and/or phylogenetically, that might have been trapped from the outside, and temporarily included in the lichen thallus as an possible additional source of nutrients (Voytsekhovich et al. 2011a). Additional photobionts should not be confused with a secondary photobionta term used for cyanobacterialcephalodiate lichens. ...
... The presence of additional photobionts in nontrebouxioid lichens might be caused by the relatively lower selectivity of the mycobiont towards its photobiont as well as its ability to use an additional source of carbon. As is shown in Voytsekhovich et al. (2011a) for the lichen genera Micarea and Placynthiella, some lichens with non-trebouxioid photobionts may be associated with several non-specific additional photobionts simultaneously. In the present study different specimens of Candelariella medians were associated with Trebouxia decolorans, T. crenulata and T. gigantea (Table 1). ...
Despite the fact that lichen algae have been explored for over100 years, their diversity, taxonomy and ecology still remains understudied. In present study we analyzed 114 specimens representing 72 lichen taxa collected on the Crimean Peninsula, culturing and sequencing their algal partners. In total we identified 26 taxa of photobionts representing 10 algal genera (Asterochloris, Chloroidium, Dilabifilum, Diplosphaera, Gloeocapsa, Myrmecia, Nostoc, Printzina, Stichococcus, Trebouxia). As most of the studied lichens (more than 80 % of studied lichen species) contained Trebouxia, for which morphology-based identification is often difficult and insufficient, we additionally used molecular methods. According to the molecular phylogeny of internal transcribed spacer (ITS) rDNA, Trebouxia species were related to three main clades: “Arboricola” (88.9 % of the investigated Trebouxia-containing lichens), “Impressa” (4.4 %) and “Simplex” (6.7 %). Based on results of molecular phylogeny and morphology three new species of Trebouxia, namely T. solaris, T. vagua and T. cretacea were described. In several lichens, two coexisting Trebouxia species were revealed in the same thallus.
