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There is still a high diversity of lichen-forming fungi that remains undescribed, especially cryptic lineages at the species level. Integrating morphological, chemical, and DNA sequence data has proved useful in corroborating species descriptions and delimitations. Here we reviewed morphological features, secondary metabolites and the DNA sequences of ITS, mtSSU and nuLSU markers to study the diversity of Xanthoparmelia in southern Africa. A total of 37 species were recorded. Three of these appear undescribed, and we therefore describe them here as new: Xanthoparmelia nimisii, with a sorediate thallus and broad lobes, is well supported as a clade separate from X. annexa; X. pseudochalybaeizans with a white medulla is phylogenetically distinct from the otherwise similar X. chalybaeizans; and X. sipmaniana, well supported as a separate clade from the similar X. hypoprocetrarica. In addition, the separation of Xanthoparmelia capensis and X. tinctina requires further studies.
Ten new species of Trypetheliaceae are described: Astrothelium bullatothallinum from Venezuela, which is close to A. aeneum but differs by the bullate thallus with a thick cortex, intermixed in a mosaic with the prothallus; A. cayennense from French Guiana, which is similar to A. flavomegaspermum but with a yellow pigment in the pseudostroma near the ostioles; A. diaphanocorticatum from Papua New Guinea, which has a bullate thallus with a thick hyaline cortex and 3-septate ascospores of 25–28×10–12 μm; A. macroeustomum from French Guiana, with joint lateral ostioles, UV+ yellow ostiolar region and 5-septate ascospores of 50–55×12–17 μm; A. minicecidiogenum from Costa Rica, with muriform ascospores of 70–90×20–25 μm, without pseudostromata, with solitary ascomata, lateral ostioles and an inspersed hamathecium; A. palaeoexostemmatis from Thailand, which is similar to A. exostemmatis but with larger, I+ blue ascospores; A. quasimamillanum from Brazil, with muriform ascospores of 30–33×9·5–10·5 μm, without pseudostromata, with solitary ascomata, lateral ostioles and an inspersed hamathecium; A. studerae from Brazil, with astrothelioid ascomata, lichexanthone only in the pseudostromata, 3-septate ascospores of 21·5–23·0×6·5–7·5 μm; A. tanianum from Malaysia, with a bullate thallus, solitary ascomata, covered by the thallus, (9–)11(–15)-septate ascospores, 75–100×20–22 μm; and Pseudopyrenula miniflavida from Brazil, with a yellow-inspersed hamathecium, the inspersion dissolving in KOH without colour change, and 3-septate ascospores, 15–17×5·5–6·5 μm. The unusually thick, hyaline cortical layer of the thallus of Astrothelium diaphanocorticatum, through which the individual algal cells are clearly visible, is discussed. Furthermore, 30 species are reported from 34 countries in which they had previously been unrecorded; one (Astrothelium inspersaeneum) is from a new continent, Asia.
Epiphytic lichens are increasingly included in forest biodiversity monitoring schemes, but most of the standardized guidelines consider only lichens colonizing a small part of tree trunks (1·0–1·5 m) and overlook other important microhabitats, such as fallen branches and stumps. In this paper, we present results of a small-scale pilot study to evaluate the possible advantage of including four distinct microhabitats in standardized procedures for assessing epiphytic lichen diversity. Trunk bases, trunks between 100 and 150 cm above the ground, stumps, and fallen branches were each sampled with a different standardized sampling method along a forest age gradient in temperate deciduous forests of the Caucasian region. Plot-level species richness was contrasted between the standardized sampling procedures of different substrata and a non-probabilistic floristic sampling. The interactions between sampling procedure and stand age were analysed using linear mixed models, and non-metric multidimensional scaling (NMDS) and multi-response permutation procedures (MRPP) were used for comparing species composition. Overall, 97 species were recorded, their richness increasing with increasing stand age. Results were consistent across the gradient of stand age and demonstrated that the adoption of standardized sampling procedures which include stumps and fallen branches in addition to tree trunks would increase the capability of maximizing species capture. This approach would allow researchers to evaluate lichen patterns by simultaneously considering the response of different communities sensitive to different stand-related factors. Despite the likelihood that a non-probabilistic floristic survey would give a more exhaustive picture of the plot-level lichen diversity, standardized sampling procedures that include tree trunks, fallen branches and stumps are likely to represent a reasonable trade-off between exhaustiveness and cost-effectiveness.
Eight new species of Pyrenulaceae are described as new to science from Brazil, Guyana and Puerto Rico. Pyrenula sanguineomeandrata Aptroot & Mercado Diaz (with a thallus with red, KOH+ purple pigmentation of lines or a reticulum, simple ascomata with vertical ostioles, a deep red inspersed, KOH+ orange hamathecium, and dark brown 3-septate ascospores 25–29×10–12 μm) and P. sanguineostiolata Aptroot & Mercado Diaz (with a thallus with deeply immersed simple ascomata with vertical ostioles, which are superficial and bright red, and 3-septate ascospores 25–28×9–12 μm) are described from submontane evergreen forests in Puerto Rico. Pyrenula biseptata Aptroot & M. Cáceres (with simple ascomata with vertical ostioles, an inspersed hamathecium and 2-septate ascospores 11–12×4·5–5·0 μm) and P. xanthinspersa Aptroot & M. Cáceres (with an ecorticate thallus containing lichexanthone, simple ascomata with vertical ostioles, not inspersed hamathecium and 3-septate ascospores 14–17×6·0–7·5 μm) are described from rainforest in Amazonian Brazil. Pyrenula subvariabilis Aptroot & Sipman (with fused ascomata with lateral ostioles and submuriform ascospores 17–20(–25)×6–9 μm) and Sulcopyrenula biseriata Aptroot & Sipman (with a thallus containing lichexanthone, simple ascomata with lateral ostioles and lozenge-shaped ascospores with 8 locules, (13–)15–17(–20)×8–10 (width)×6–7 (thickness) μm) are described from savannahs in Guyana. Special attention is paid to the genus Pyrgillus: two new species from the 3-septate core group of this small genus are described from Brazil, viz. P. aurantiacus Aptroot & M. Cáceres (with a corticate thallus containing lichexanthone, mazaedium with orange, KOH+ violet, UV+ red pruina and ascospores of 13–16×6·0–7·5 μm) and P. rufus Aptroot & M. Cáceres (with a corticate thallus containing lichexanthone, mazaedium with dark red, KOH+ orange, UV+ red pruina and ascospores of 15·0–17·5×5·0–6·5 μm). An updated key to the 3-septate species of Pyrgillus is provided.
Ramalina europaea Gasparyan, Sipman & Lücking and R. labiosorediata Gasparyan, Sipman & Lücking, two species of the R. pollinaria group, are described here as new to science. Ramalina europaea, widely distributed in Europe, can be distinguished by small, punctiform, often terminal soralia starting out on small, spine-like branchlets, whereas R. labiosorediata from North America differs from R. pollinaria s. str. and R. europaea in the almost exclusively terminal soralia formed on the tips of normal lobes, originating from the underside and becoming irregularly labriform. Morphological characters, chemistry, ecology and geographical distribution are discussed and a key to the species of the Ramalina pollinaria group is provided. The topology of a maximum likelihood tree based on ITS shows the presence of three well-supported clades, corresponding to the morphological differences of the three species. The status of several historical names variously placed in synonymy with or described as infraspecific entities of R. pollinaria is reassessed and a new neotype and an epitype are designated for Lichen pollinarius, a neotype for L. squarrosus, making it a synonym of R. farinacea, and lectotypes for R. pollinaria var. elatior, making it a synonym of R. pollinaria s. str., and for var. humilis, a taxon of yet unknown affinity.
A world key to the 21 species of Stirtonia is presented. Three new Stirtonia species are described from the Neotropics: Stirtonia ibirapuitensis Aptroot, Käffer & S. M. Martins, with whitish to cream amoeboid ascigerous areas with lichexanthone on a greenish thallus without lichexanthone and ascospores of 27–32×9·5–12·5 µm; Stirtonia punctiformis Aptroot & Sipman, with ascigerous structures of thallus colour, consisting of one or more brown asci with brown walls with surrounding tissue, in groups or irregular lines and brown ascospores 61–73×27–35 µm; and Stirtonia viridis Aptroot, L. I. Ferraro, Sipman & M. Cáceres, with ascigerous structures mostly linear, branched and anastomosing, whitish, and contrasting with the often greenish thallus and with ascospores 50–58×15–22 µm. In addition, a specimen of S. neotropica is reported that contains some patches of lichexanthone, S. curvata is reported new to the Neotropics from Brazil and the Netherlands Antilles, and S. nivea is reported new to the Northern Hemisphere from Puerto Rico. Also, Crypthonia divaricatica Aptroot & Sipman, with an irregular, thick thallus with divaricatic and usnic acids and flat white ascigerous areas and macrocephalic 5–9-septate ascospores 20–27×9·5–12·5 µm is described from Mexico. While only one Stirtonia species was known from the Neotropics as recently as 2009, the total number of Stirtonia species known from the Neotropics is now 12, an equal number to the 12 species that are known from the Palaeotropics.
Twenty-three species of Pyrenula from Latin America are treated here. Several species show characters that were not previously reported in the genus and are rare or new to lichenized fungi, viz. yellow, orange or red (KOH+ green) oil inspersion in the hymenium, yellow oil in young ascospores or longitudinal ridges on the ascospore wall. Two taxonomically significant types of over-mature spores are illustrated. The following new species are described: Pyrenula aggregataspistea Aptroot & M. Cáceres, P. aurantioinspersa Aptroot & Sipman, P. cornutispora Aptroot & M. Cáceres, P. flavoinspersa Aptroot & Sipman, P. guyanensis Sipman & Aptroot, P. infraleucotrypa Aptroot & M. Cáceres, P. inframamillana Aptroot & M. Cáceres, P. infrastroidea Aptroot & Sipman, P. maritima Sipman & Aptroot, P. mattickiana Sipman & Aptroot, P. minoides Aptroot & Sipman, P. monospora Aptroot & Sipman, P. paraminarum Aptroot & M. Cáceres, P. perfecta Aptroot & Sipman, P. plicata Sipman & Aptroot, P. rubroinspersa Aptroot & Sipman, P. rubronitidula Aptroot & M. Cáceres, P. rubrostigma Aptroot & M. Cáceres, P. tetraspora Aptroot & Sipman, P. triangularis Aptroot & Sipman, P. viridipyrgilla Aptroot & M. Cáceres. Pyrenula seminuda (Müll. Arg.) Sipman & Aptroot is a new combination.
The lichen Pyrenula minutissima is described as new to science from the Hyrcanian forests in northern Iran. In addition, three further, essentially tropical, Pyrenula species are reported for the first time from Iran. An identification key is provided for all eight Pyrenula species now known from Iran.
The lichen genus Sporodochiolichen is installed to accommodate a group of four, so far undescribed, tropical corticolous lichen species with arthric conidia in discrete sporodochia. Three species are so far only known from Papua New Guinea; one is known from five countries on two continents and is probably pantropical.
The species of the genus Immersaria in Iran are reviewed. Immersaria iranica is described as a new species in the Lecideaceae, characterized by pycnoconidia opening by stellate, winding cracks and 2′-O-methylsuperphyllinic acid as major secondary metabolite. The characters separating it from other taxa of the Lecideaceae are discussed. Immersaria athroocarpa and I. cupreoatra are new records for Iran. An identification key and a table with diagnostic characters of the species hitherto known from Iran are provided.
In the course of an ongoing systematic and taxonomic revision of the lichen family Graphidaceae (including Thelotremataceae), we present world-wide keys to the currently accepted thelotremoid genera and species, excluding the columellate taxa and their relatives of the Ocellularia-Myriotrema-Stegobolus clade (Melanotrema, Myriotrema, Ocellularia, Ocellularia clandestina group, Redingeria, Stegobolus), which will be treated in a forthcoming paper. The keys include all genera and species with chroodiscoid, lepadinoid, and topeliopsidoid apothecia and other taxa featuring periphysoids or fibrils, and their relatives. Taxa keyed out to genus and species level are Acanthotrema, Chapsa, Chroodiscus, Diploschistes, Fibrillithecis, Gyrotrema, Leptotrema, Leucodecton, Melanotopelia, the ‘Ocellularia’ cruentata group, Pseudoramonia, Reimnitzia, Schizotrema, Thelotrema, Topeliopsis and Wirthiotrema. Over 260 species are treated, including a few yet unnamed taxa. The following taxonomic and nomenclatural novelties are introduced: Acanthotrema frischii Lücking sp. nov., Chapsa aggregata (Hale) Sipman & Lücking comb. nov., C. albida (Nyl.) Lücking & Sipman comb. nov.; C. albomaculata (Sipman) Sipman & Lücking comb. nov., C. boninensis (Tat. Matsumoto) Rivas Plata & Mangold comb. nov., C. elabens (Müll. Arg.) Rivas Plata & Mangold comb. nov., C. imperfecta (Hale) Rivas Plata & Lücking comb. nov., C. laceratula (Müll. Arg.) Rivas Plata & Lücking comb. nov., C. magnifica (Berk. & Broome) Rivas Plata & Lücking comb. nov., C. meghalayensis (Patw. & Nagarkar) Lumbsch & Divakar comb. nov., C. meridensis (Kalb & Frisch) Lücking, Lumbsch & Rivas Plata comb. nov., C. mirabilis (Zahlbr.) Lücking comb. nov., C. neei (Hale) Mangold & Lücking comb. nov., C. paralbida (Riddle) Rivas Plata & Lücking comb. nov., C. pseudoexanthismocarpa (Patw. & C. R. Kulk.) Rivas Plata & Lücking comb. nov., C. pulvereodisca (Hale) Rivas Plata & Mangold comb. nov., C. scabiomarginata (Hale) Rivas Plata & Lücking comb. nov., C. waasii (Hale) Sipman & Lücking comb. nov., Fibrillithecis argentea (Müll. Arg.) Rivas Plata & Lücking comb. nov., F. carneodisca (Hale) Rivas Plata & Lücking comb. nov., F. confusa Lücking, Kalb & Rivas Plata spec. nov., F. diminita (Hale) Rivas Plata & Lücking comb. nov., F. eximia (R. C. Harris) Rivas Plata & Lücking comb. nov., F. fissurata (Nagarkar & Hale) Rivas Plata & Lücking comb. nov., F. gibbosa (H. Magn.) Rivas Plata & Lücking comb. nov., Leucodecton desquamescens (Vain.) Lücking comb. nov., L. oxysporum (Redinger) Lücking comb. nov., Schizotrema cryptotrema (Nyl.) Rivas Plata & Mangold comb. nov., Thelotrema patwardhanii (Hale) Rivas Plata & Mangold comb. nov., Topeliopsis guaiquinimae (Sipman) Rivas Plata & Mangold comb. nov., and T. tuberculifera (Vain.) Rivas Plata & Mangold comb. nov. Using the examples of Fibrillithecis halei s. lat., Leucodecton compunctellum s. lat., and Thelotrema monosporum s. lat., we show how difficult species complexes can be flexibly treated in a key, allowing for either a broad concept or the distinction of several individual taxa.
In a recent treatment of the Pannariceae of Ecuador, Jørgensen & Arvidsson (2004: 131) pointed out the surprising absence of the genus Fuscopannaria in Ecuador, a genus which is rather common, with many species, in the Pacific Northwest of North America (Jørgensen 2001). The authors claim that, on the western side of the Americas, the genus appears to stop north of Mexico, only to reappear in southern Chile and Argentina. While going through his material, the second author discovered two specimens belonging in this genus from Mexico and Ecuador, which contradicts that statement. Both are typical Fuscopannaria leucosticta specimens (Fig. 1), the type species of the genus originally described from eastern North America (Jørgensen 2001: 684), and fairly widely distributed in warm temperate parts there, and extending as far south as Cuba (Fig. 2).
The new genus Aptrootia Lücking & Sipman is described for Thelenella terricola, an enigmatic terricolous and muscicolous, pyrenocarpous taxon known from Papua New Guinea and Costa Rica, and the new combination Aptrootia terricola (Aptroot) Lücking, Umaña & Chaves comb. nov. is introduced. Aptrootia is characterized by completely immersed perithecia with brown-black ostiolar region, surrounded by a white, cartilaginous thallus resembling that of Gomphillaceae. The hamathecium is typical of Trypetheliaceae, with thin, anastomosing paraphysoids embedded in a gelatinous matrix, but the dark brown ascospores are otherwise unknown within the family. The only known species was tentatively described in Thelenella, but hamathecium type and molecular data place Aptrootia within Trypetheliaceae.
The genus Haematomma is treated as part of the Ticolichen biodiversity inventory in Costa Rica. Together with literature reports, fourteen taxa are recognized as occurring in the country (plus three species to be expected), and three of them are new to science: Haematomma nicoyense Nelsen, Lücking & Chaves, with a sorediate thallus, russulone and atranorin, and zeorine apothecia with a split between proper excipulum and thalline margin, Haematomma staigeriae Nelsen, Lücking & Umaña, with a sorediate thallus, russulone and lichexanthone, and submuriform ascospores, and Haematomma fluorescens var. longisporum Nelsen, Lücking & Navarro, with russulone and lichexanthone and ascospores up to 120 μ m in length with up to 27 septa. A key is presented for the 17 taxa known or expected from Costa Rica.
Thirty-one taxa of Gyalideopsis and its recent segregates (Ferraroa, Jamesiella, Lithogyalideopsis) are reported in a first assessment of the group from Costa Rica. Six species are described as new, all originating from perhumid montane rainforest: Gyalideopsis altamirensis Lücking & Umaña, with broadly sessile, brown apothecia and single-spored asci with muriform ascospores (hyphophores unknown); G. macarthurii Lücking, Umaña & Aptroot, with dark greyish brown apothecia having their margin covered by triangular thalline lobules and shortly bristle-shaped hyphophores producing moniliform diahyphae; G. pseudoactinoplaca Lücking & Chaves, with sessile, globose diahyphal bunches similar to those of Actinoplaca and resembling isidia (apothecia unknown), G. wesselsii Lücking, Sipman & Chaves, with verrucose thallus, dark greyish brown, applanate apothecia with single-spored asci producing muriform ascospores, and minutely spathulate, dark brown hyphophores with moniliform diahyphae, and Jamesiella chaverriae Chaves, Umaña & Lücking, with isidioid hyphophores (‘thlasidia’) which are flask-shaped and apically thinly ciliate (apothecia unknown). A further species, Gyalideopsis sp., with distinctly stipitate, umbelliform hyphophores, is not formally described due to the lack of apothecia and hyphophores. A further nine species are reported as new to Costa Rica: G. buckei, G. capitata, G. confluens, G. giganteoides, G. megalospora, G. napoensis, G. nepalensis, G. palmata, and Lithogyalideopsis zeylandica. A world-wide key to all presently known 94 taxa of Gyalideopsis and its recent segregates is presented, as well as a checklist.
Descriptions are provided for two new species, Mycomicrothelia lateralis Sipman from Norfolk Island and M. triangularis Aptroot from Puerto Rico, both characterized by lateral ostioles and uniseriate ascospores. Based on the examination of additional fresh material, the genus Ornatopyrenis is included in Mycomicrothelia and the new combination Mycomicrothelia queenslandica (Müll. Arg.) Sipman & Aptroot is made. Ornatopyrenis muriformis Aptroot is proposed as a synonym of M. decipiens (Müll. Arg.) R. C. Harris.
The taxonomy of lichenized fungi (lichens) is relatively well-known compared to that of other fungi because lichens are long-lived and possess a thallus that requires exposure to light. Therefore they are easily observed on their natural substratum and sampling can be carried out much more thoroughly than in other fungal groups. Comprehensive catalogues of lichen taxa known worldwide are available and currently about 13500 species are accepted.
An analysis of selected, recently revised groups suggests that there has been a mean increase in species number of about 25% since 1931. One third of the species recognized before 1931 were withdrawn, and almost half of the currently recognized species have been described after 1931. Some 3000 names seem to be ‘orphaned’, not included in currently accepted genera and not appearing in recent species counts. New species are being described at an increasing rate, so that large numbers of ‘missing lichens’ seem to be present. However, there is no clear indication how numerous they are, only an estimation that they amount to some 25% of all species.
A geographical analysis of checklists suggests that most missing species will be found in the tropics and the southern hemisphere. A taxonomic analysis of recently published species suggests that there is no particular group yielding more novelties than others. Our own fieldwork impressions suggest that tropical primary forests are the most productive in revealing undescribed species. These are found in particular on leaves and on bark high up on the tree trunks. In well-studied areas like western Europe the new discoveries tend to be cryptic species or species rarely producing ascocarps.
In conclusion, the ‘missing’ lichens are estimated at about 4000, and are to be found everywhere and in all taxonomic groups, but predominantly in primary tropical forests. This would result in a total of about 18000 lichen species.
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