Alexander Khodosovtsev

Taxonomy and phylogeny of the Caloplaca cerina group in Europe

Using ITS nrDNA sequence data, the Caloplaca cerina group (Teloschistaceae) is defined here as a monophyletic, but internally richly branched lineage. The group is also characterized by a combination of morphological and anatomical characters. Its internal lineages are supported by phenotypic characters in addition to ecology and distribution. Within the large C. cerina group, we have found at least 20 phylospecies in the temperate zone of the Northern Hemisphere. Two species complexes do not produce any vegetative diaspores: the polyphyletic, corticolous Caloplaca cerina s. lat. (six separated cryptic or semi-cryptic species) and the monophyletic C. stillicidiorum s. lat. that grows mainly on plant debris, small shrubs and bryophytes and consists of at least four internal lineages. All lineages producing vegetative diaspores (soredia, blastidia, isidia or lobules) are phenotypically characteristic and represent fairly easily distinguishable species: C. chlorina, C. isidiigera, C. monacensis, C. subalpina, C. thracopontica, C. turkuensis and C. ulmorum. Only the North American sorediate C. pinicola possibly represents an aggregate of species. Caloplaca sterilis is described as a new species. A key to the phenotypically distinguishable species is provided. Lectotypes are designated for C. albolutea, Caloplaca cerina f. coronulata and for C. monacensis. The Australian C. hanneshertelii belongs to this group, but this and other possible species from the Southern Hemisphere are not treated here in detail. Some species traditionally placed in the C. cerina group due to their similar morphology are excluded here on the basis of our phenotype examinations and molecular data. Caloplaca albolutea, C. mydalaea and C. virescens are uncertain taxa and their identities still remain unclear. (Less)

Two new European species from the heterogeneous Caloplaca holocarpa group ( Teloschistaceae )

The Caloplaca holocarpa group contains members of the family Teloschistaceae with a strongly reduced thallus and conspicuous yellow, orange or red apothecia. In the absence of well-defined thallus characters, taxa of this group must be identified mainly by apothecial characters and are as a result often difficult to separate. The species of this group have been shown not to form a monophyletic entity, with representatives of other Teloschistaceae with more complex thalli intermixed among them. Caloplaca skii and C. syvashica are recognized here as two homogeneous clades with Caloplaca holocarpa -like phenotypes. Caloplaca skii , which is widespread in southern Europe, resembles C. cerinelloides but is distinguished by its smaller and narrower ascospores and by growing predominantly on xerophilous shrubs. Caloplaca syvashica is restricted to shrubs in salt marshes in the northern Black Sea region. It is similar to the British Caloplaca suaedae and Australian C. yarraensis , but differs from both, mainly in ascospore characters. Caloplaca yarraensis is closely related to the new C. syvashica but arguments against their conspecificity are emphasised. A key for epiphytic C. holocarpa -like Teloschistaceae from Europe is provided.

The Caloplaca crenulatella species complex; its intricate taxonomy and description of a new species

The Caloplaca crenulatella species complex is monophyletic, based on analysis of ITS nrDNA sequences. It is characterized mainly by its ascospores and a simplified thallus. Caloplaca aquensis, C. borysthenica, C. interfulgens, C. lactea, C. marmorata and C. tominii represent well-defined monophyletic lineages within it. Caloplaca diffusa, described here as a new species, is a well-supported lineage with a distinctive phenotype. Caloplaca crenulatella s. lat. and C. ferrarii s. lat. are heterogeneous taxa appearing in four and two places respectively within our tree. Caloplaca ochracea also belongs to the group, although it has different ascospore characters. Caloplaca gyalolechiaeformis and C. pseudo- citrina are later synonyms of C. tominii. Arctic and North American sorediate specimens morphologi- cally very close to C. tominii represent a separate lineage recently recognized as C. erichansenii. The North American Caloplaca nashii has a C. crenulatella-like phenotype but does not belong to the C. crenulatella complex. The identities of Caloplaca lacteoides and Caloplaca epigaea are not clear, but the latter may be a terricolous ecotype of one of the entities within C. crenulatella s. lat. A key to the European species of the group is provided.

Additions to the diversity of rare or overlooked lichens and lichenicolous fungi in Ukrainian Carpathians

Наводяться дані щодо 103 рідкісних, маловідомих та спорадично поширених видів лишайників та ліхенофільних грибів з Українських Карпат. Вперше для України виявлено 29 видів ліхенізованих, три види ліхенофільних та один альгофільний гриб: Absconditella sphagnorum, Adelolecia kolaensis, Arthonia muscigena, Arthrorhaphis aeruginosa, Biatora albohyalina, Brodoa atrofusca, Bryodina rhypariza, Calicium pinastri, Caloplaca fuscorufa, C. isidiigera, Carbonea invadens, Catillaria croatica, Cryptodiscus gloeocapsa, Cystocoleus ebeneus, Epigloea medioincrassata, Gyalidea fritzei, Lecidea pullata, Lecidella patavina, Melaspilea granitophila, Micarea turfosa, Monodictys epilepraria, Opegrapha corticola, Phaeographis inusta, Polyblastia schaereriana, Protothelenella sphinctrinoides, Psilolechia clavulifera, Pycnora leucococca, Rinodina orculata, Sclerococcum griseisporodochium, Thelocarpon robustum auct. brit., non Eitner, Trapeliopsis glaucolepidea and Vezdaea stipitata. Усі гербарні зразки під назвою Brodoa intestiniformis, які були зібрані з території Українських Карпат відносятьсяся до B. atrofusca. Повідомлення щодо зростання Chaenotheca cinerea на території Українських Карпат є недостовірними. Наводяться перші реальні місцезнаходження Caloplaca conversa, Chaenotheca cinerea та Lecidea sphaerella для України. Multiclavula mucida та Schaereria fuscocinerea у зведеннях щодо лишайників Східних Карпат та України відсутні, однак повідомлення про ці види знайдені в українських та чеських джерелах. Pertusaria ophthalmiza був невірно наведений для України під назвою P. multipuncta і тому повинен бути виключеним із чекліста лишайників України. Phaeographis dendritica та Biatora meiocarpoides також виключаються із списку українських лишайників, тому що перший був некоректно наведений з Українських Карпат, а другий є синонімом до Micarea lithinella.

New lichenized and lichenicolous fungi for the Crimean peninsula (Ukraine)

Data on 25 species of lichenized and lichenicolous fungi new to the Crimean Peninsula, collected during Czech and Ukrainian-Czech lichenological excursions in June 2006 and May 2007 respectively, are provided. Among them, Arthonia lecanorina (Almq.) R. Sant., A. nideri (J. Steiner) Clauzade, Diederich & Cl. Roux, A. punctella Nyl., Caloplaca furax Egea et Llimona, C. soralifera Vondrák & Hrouzek, C. veneris Cl. Roux & Nav.-Ros., Carbonea assimilis (Körber) Hafelner & Hertel, Catillaria aff. scotinodes (Nyl.) Coppins, Endococcus macrosporus (Arnold.) Nyl., E. ramalinarius (Lind.) D. Hawksw., Fuscidea gothoburgensis (H. Magn.) V. Wirth & Vĕzda, Lichinella cribellifera (Nyl) P.P. Moreno & Egea, Milospium graphideorum (Nyl.) D. Hawksw., Phaeospora lecanorae Eitner, Placidium pilosellum (Breuss) Breuss, Verrucaria biatorinaria Zehetl, and V. poeltiana Clauzade & Cl. Roux are new to Ukraine. Caloplaca irrubescens (Nyl. ex Arnold) Zahlbr., Lecanora rouxii S. Ekman & Tønsberg, Lichenothelia scopularia (Nyl.) D. Hawksw., Melanelia hepatizon (Ach.) A. Thell, Peltigera collina (Ach.) Schrad., Staurothele areolata (Ach.) Lettau, Stigmidium rouxianum Calatayud & Triebel and Zwackhiomyces coepulonus (Norm.) Grube & R. Sant. are new to Crimea.

Absence of anthraquinone pigments is paraphyletic and a phylogenetically unreliable character in the Teloschistaceae

It has been suggested that the absence of anthraquinones is not a synapomorphic character, but appears independently in unrelated lineages of Teloschistaceae . We analyzed ITS nrDNA regions in species of the genus Caloplaca and present evidence for five such examples: the Caloplaca cerina group, C. obscurella , the C. servitiana group, the C. xerica group and the C. variabilis group ( Pyrenodesmia ). In some cases, loss of anthraquinones is observed only in individuals within ordinarily pigmented populations, but sometimes the loss covers whole lineages containing one or more species. Both situations are observed in the C. servitiana group. Loss of anthraquinones is always followed by the synthesis of ‘alternative’ pigments (often Sedifolia -grey). In the specimens with anthraquinone-containing apothecia studied, these pigments are not visible in apothecial sections after dissolving anthraquinones in K. Fully unpigmented apothecia have not been observed. The Caloplaca xerica group is a newly established, infraspecific grouping of species related to, and similar to, C. xerica . The Caloplaca servitiana group is also newly established and represents an isolated lineage covering two rather different, but related species. Caloplaca neotaurica is described here as a new species with apothecia of two colour variants; orange-red (with anthraquinones) and grey (with Sedifolia -grey). The genus Huea represents another taxon lacking anthraquinones within Teloschistaceae . The genera Apatoplaca and Cephalophysis , which lack anthraquinones, are tentatively placed in Teloschistaceae , but their phylogenetic identity has not been recognized. Hueidea is reported to have no anthraquinones, but its secondary metabolites should be studied further and its possible placement in Teloschistaceae assessed. We suggest that Caloplaca abbreviata var. lecideoides and C. celata represent variants of C. stillicidiorum lacking anthraquinones.

Methods for phenotypic evaluation of crustose lichens with emphasis on Teloschistaceae

We present the methods for phenotype evaluation, which give good chances to identify taxa larger according to the anatomical and morphological features. The basic idea is to evaluate characters having regard to completeness and reproducibility. We composed a list of about 100 basic characters and propose standardized ways of measuring them. We also present a list of morphological and anatomical terms with their standardized meaning. Basic rules for collecting, preparing and measuring lichen material are described. The proposed phenotypic evaluation process has two levels: (1) a pilot study is carried out on a restricted number of samples, but all available characters are evaluated; (2) a detailed study of a large number of samples that includes only potentially diagnostic features that are chosen for the pilot study according to their discriminatory weight. (Less)

Three New, Seemingly-Cryptic Species in the Lichen Genus Caloplaca (Teloschistaceae) Distinguished in Two-Phase Phenotype Evaluation

We describe three new, seemingly-cryptic species in the lichen genus Caloplaca (Teloschistaceae) with black apothecia. Those species, separated in nrITS and β-tubulin DNA phylogenies, appeared to be phenotypically indistinguishable. We looked for their phenotypic differences using a two-phase method comprised of a preliminary examination in which diagnostic value of all available characters was evaluated using a small number of samples and potentially-diagnostic characters were selected, and a subsequent detailed study in which characters selected in the first phase were tested using more samples. We found 19 diagnostic characters (continuous and discrete) of which four continuous and three discrete characters could be considered “fully diagnostic”, i.e. allowing for correct identification of at least one species. Hence, the three species are not cryptic, but can be distinguished phenotypically. Here, they are formally described as Caloplaca micromarina Frolov, Khodos. & Vondrák sp. nova, C. micromontana Frolov, Wilk & Vondrák sp. nova and C. microstepposa Frolov, Nadyeina, Khodos. & Vondrák sp. nova.

The identity of type specimens in BP of some names in Caloplaca

Type materials held in BP of 16 specific and intraspecific names now placed in Caloplaca are appraised here. The names Caloplaca balatonica, C. cerinella f. aggregata, C. lactea var. subimmersa, C. lactea f. densa, C. lallavei f. fulva, C. vitellinaria, C. vitellinoides, and Gasparrinia granulosa f. flavovirens are reduced into synonymy of older names. We did not find any older synonyms for the names Caloplaca brachyspora, C. flavovirescens var. persica, C. gyalolechiiformis, C. hungarica, and C. servitiana. The identity of the names C. lojkae, C. variabilis f. densa, and C. variabilis f. geographica is presently unclear. Caloplaca pseudocitrina is reduced into synonymy with C. gyalolechiiformis. Caloplaca servitiana is considered to be different from another “black-fruiting” corticolous species C. oleicola. Lectotypes are designated for Caloplaca brachyspora, C. lojkae, and C. servitiana.

Three new Pronectria species in terricolous and saxicolous microlichen communities (Bionectriaceae, Ascomycota)

Abstract: Three new species of Bionectriaceae are described; (1) Pronectria algicola grows within Gloeocystis-like algal crusts with Steinia geophana and Sarcosagium campestre; (2) P. caloplacae is lichenicolous on areoles of Caloplaca austrocitrina; (3) P. diplococca grows in squamules of Collema spp. The latter species has early dissolving asci and 1-septate ascospores disintegrating into simple globose mature ascospore fragments. Fragmentation of ascospores appears in three additional species of Pronectria. A key to the known Pronectria species is provided.