Cladonia Species Research Articles

Distribution and habitat preference of protected reindeer lichen species (Cladonia arbuscula, C. mitis and C. rangiferina) in the Balaton Uplands (Hungary)

AbstractThe maintenance of protected lichen species and their biodiversity in general depends on good management practices based on their distribution and habitat preferences. To date, 10 of the 17 protected lichen species of Hungary have been recorded in the Bakony Mts including the Balaton Uplands region. Habitat preferences of three protected Cladonia species (C. arbuscula, C. mitis and C. rangiferina) growing on underlying rocks of red sandstone, basalt, Pannonian sandstone and gravel were investigated by detailed sampling. We recorded aspect, underlying rock type, soil depth, pH and CaCO3 content, habitat type (as defined by the General National Habitat Classification System Á-NÉR), all species of lichen, bryophyte and vascular plants as well as percentage cover of exposed rock, total bryophytes, lichens, vascular plants and canopy, degree of disturbance and animal impacts. Sporadic populations of these species mostly exist at the top of hills and mountains in open acidofrequent oak forests, but they may occur in other habitats, such as closed acidofrequent oak forests, slope steppes on stony soils, siliceous open rocky grasslands, open sand steppes, wet and mesic pioneer scrub and dry Calluna heaths. Cladonia rangiferina was found to grow beneath higher canopy cover than either C. arbuscula or C. mitis in the Balaton Uplands. Furthermore, there were significant differences in canopy cover between occupied and unoccupied quadrats in the case of all three species. Cladonia rangiferina is a good indicator species of natural habitats in Hungary due to its restricted distribution and low ecological tolerance. These results may lead to the adoption of effective conservation methods (e.g. game exclusion, artificial dispersal) in the future.

Predicting patterns of terrestrial lichen biomass recovery following boreal wildfires

AbstractIncreased fire activity due to climate change may impact the successional dynamics of boreal forests, with important consequences for caribou habitat. Early successional forests have been shown to support lower quantities of caribou forage lichens, but geographic variation in, and controls on, the rates of lichen recovery has been largely unexplored. In this study, we sampled across a broad region in northwestern Canada to compare lichen biomass accumulation in ecoprovinces, including the Saskatchewan Boreal Shield, the Northwest Territories Taiga Shield, and Northwest Territories Taiga Plains, divided into North and South. We focused on the most valuable Cladonia species for boreal and barren‐ground caribou: Cladonia mitis and C. arbuscula, C. rangiferina and C. stygia, and C. stellaris and C. uncialis. We developed new allometric equations to estimate lichen biomass from field measurements of lichen cover and height; allometries were consistent among ecoprovinces, suggesting generalizability. We then used estimates of lichen biomass to quantify patterns of lichen recovery in different stand types, ecoprovinces, and with time following stand‐replacing fire. We used a hurdle model to account both for the heterogeneous nature of lichen presence (zero inflation) and for the range of abundance in stands where lichen was present. The first component of the hurdle model, a generalized linear model, identified stand age, stand type, and ecoprovince as significant predictors of lichen presence. With a logistic growth model, a measure of lichen recovery (time to 50% asymptotic value) varied from 28 to 73 yr, dependent on stand type and ecoprovince. The combined predictions of the hurdle model suggest the most rapid recovery of lichen biomass across our study region occurred in jack pine in the Boreal Shield (30 yr), while stands located in the Taiga Plains (North and South) required a longer recovery period (approximately 75 yr). These results provide a basis for estimating future caribou habitat that encompasses some of the large variation in fire effects on lichen abundance and vegetation types across the range of boreal and barren‐ground caribou in North America.

Morphological and Chemical Traits of Cladonia Respond to Multiple Environmental Factors in Acidic Dry Grasslands.

Terricolous lichen communities in lowlands occur especially in open dry habitats. Such communities are often dominated by species of the genus Cladonia, which are very variable in morphology, reproduction strategies, and secondary metabolites. In this work, we investigated traits-environment relationships considering vegetation dynamics, substrate pH, disturbance, and climate. A total of 122 plots were surveyed in 41 acidic dry grasslands in the western Po Plain (Northern Italy). Relationships between Cladonia traits and environmental variables were investigated by means of a model-based Fourth Corner Analysis. Thallus morphology and metabolites responded to vegetation dynamics, substrate pH, disturbance, and climate, whereas reproduction strategies responded only to vegetation dynamics. Traits’ correlations with vegetation dynamics elucidate their colonization patterns in open dry habitats or suggest biotic interactions with bryophytes and vascular plants. In addition, correlations between metabolites and environmental factors support interpretations of their ecological roles. Our results also stress the importance of studying traits’ relationships with climatic factors as an alert towards lichen reactions to climate change.

Deep convolutional neural network for preliminary in-field classification of lichen species

Lichens are unique organisms, valued for their pharmacological activity, but also well known as bioindicators of environmental pollution, key determinants for some natural ecological habitats, or just popular elements of decoration. High morphological similarity between lichen species makes their recognition complicated, especially under in-field conditions. Thus, there is a need for a quick and easy method that can help with the preliminary classification of selected lichen species. This paper presents a tool that can facilitate the recognition of Cladonia lichen species, based on a deep convolutional neural network, a model which has nowadays reached a classification level often comparable to humans. The network was trained and tested on twelve Cladonia species using a total of 1164 images, downloaded from various websites. The trained model achieved 60.94% accuracy, which is satisfactory for this novel, but still preliminary, automated classification of lichen species. • Deep convolutional neural network was designed to determine the lichen species. • The model was trained and tested on images of 12 Cladonia species, on1164 images. • Satisfactory 60.94% accuracy was achieved for the model. • This is the first use of neural network to identify lichen species. • The model can be useful for lichenologist and non-expert nature amateurs.

The genus Cladonia from Greece

Burgaz, A. R., Gutiérrez-Larruga, B., Rodríguez-Arribas, C. & Pino-Bodas, R. 2020. The genus Cladonia from Greece. – Herzogia 33: 340 –376.A study of the Cladoniaceae has been carried out in Greece, based on collections from 223 different localities. Thirty four Cladonia species were found, seven of which are new records to Greece, Cladonia conista, C. corsicana, C. digitata, C. macrophyllodes, C. peziziformis, C. polycarpoides and C. subturgida. As a result of this study the total number of Greek Cladonia species rises to 43. The currently known distribution ranges in Greece as well as information about the chemical variability of most of the species are provided. A key to all the Cladonia taxa known from Greece is included.

Global Biodiversity Patterns of the Photobionts Associated with the Genus Cladonia (Lecanorales, Ascomycota)

The diversity of lichen photobionts is not fully known. We studied here the diversity of the photobionts associated with Cladonia, a sub-cosmopolitan genus ecologically important, whose photobionts belong to the green algae genus Asterochloris. The genetic diversity of Asterochloris was screened by using the ITS rDNA and actin type I regions in 223 specimens and 135 species of Cladonia collected all over the world. These data, added to those available in GenBank, were compiled in a dataset of altogether 545 Asterochloris sequences occurring in 172 species of Cladonia. A high diversity of Asterochloris associated with Cladonia was found. The commonest photobiont lineages associated with this genus are A. glomerata, A. italiana, and A. mediterranea. Analyses of partitioned variation were carried out in order to elucidate the relative influence on the photobiont genetic variation of the following factors: mycobiont identity, geographic distribution, climate, and mycobiont phylogeny. The mycobiont identity and climate were found to be the main drivers for the genetic variation of Asterochloris. The geographical distribution of the different Asterochloris lineages was described. Some lineages showed a clear dominance in one or several climatic regions. In addition, the specificity and the selectivity were studied for 18 species of Cladonia. Potentially specialist and generalist species of Cladonia were identified. A correlation was found between the sexual reproduction frequency of the host and the frequency of certain Asterochloris OTUs. Some Asterochloris lineages co-occur with higher frequency than randomly expected in the Cladonia species.

The genus Cladonia (Lichenized Ascomycota) from Restinga Vegetation of Espírito Santo state, Brazil: Supergroups Crustaceae and Perviae

Abstract In this paper we recognize a total of 11 species of Cladonia, five belonging to Supergroup Crustaceae and six to Supergroup Perviae, occurring in the Restinga vegetation of Espírito Santo state, Brazil. We confirm the occurrence of C. sprucei and C. rangiferina to Espírito Santo state Restinga, being their most austral record in Brazil so far. Four species are new records to the state: C. consimilis, C. dendroides, C. salzmannii, and C. sphacelata. Here we expand the number of known species of Cladonia of Supergroup Perviae from three to five, and of Supergroup Crustaceae from three to six in the studied environment. An identification key, comments and illustrations are provided.

Changes in the ground environment under the influence of fires in the territory of the State Natural Reserve «Olekminsky»

This paper discusses the chronology and cause of forest fires on the territory of the State Reserve “Olekminsky”. Forest fires on the territory of the reserve are low-level, caused by dry thunderstorms. The overgrowth of burnt areas occurs at the expense of specific groups of pyrophyte plants. For 2-3 years after the fires, Camenerion angustifolium (L.) Scop., Corydalis sibirica (L.) Pers., Plantago canescens Adams, Crepis tectorum L. and others were found. In waterlogged areas, Tephroseris palustris (L.) Reichenb appear and subsequently do not occur, and mosses: Marshantia polymorpha L., Ceratodon purpureum (Hedw.) Brid. It was found that the restoration of ground cover is mainly due to species characteristic only for intermediate stages: Lichens - Baeomyces carneus, Dibaeis baeomyces and Trapeliopsis granulosa, as well as Arctous erythrocarpa (Small) M. Ivanova, Chamaenerion angustifolium (L.) Scop., Calamagrostis obtusata Trin, Сarex spp., cereals and green mosses. More than 40 species of lichens participate in the formation of the ground cover of the studied larch forests, mainly bushy species of Cladonia, Stereocaulon, Peltigera. Forest fires of a natural nature occurring on the territory of the reserve are the main limiting factor affecting bio-resources and theirrestoration.

The first survey of Cystobasidiomycete yeasts in the lichen genus Cladonia; with the description of Lichenozyma pisutiana gen. nov., sp. nov.

The view of lichens as a symbiosis only between a mycobiont and a photobiont has been challenged by discoveries of diverse associated organisms. Specific basidiomycete yeasts in the cortex of a range of macrolichens were hypothesized to influence the lichens' phenotype. The present study explores the occurrence and diversity of cystobasidiomycete yeasts in the lichen genus Cladonia. We obtained seven cultures and 56 additional sequences using specific primers from 27 Cladonia species from all over Europe and performed phylogenetic analyses based on ITS, LSU and SSU rDNA loci. We revealed yeast diversity distinct from any previously reported. Representatives of Cyphobasidiales, Microsporomycetaceae and of an unknown group related to Symmetrospora have been found. We present evidence that the Microsporomycetaceae contains mainly lichen-associated yeasts. Lichenozyma pisutiana is circumscribed here as a new genus and species. We report the first known associations between cystobasidiomycete yeasts and Cladonia (both corticate and ecorticate), and find that the association is geographically widespread in various habitats. Our results also suggest that a great diversity of lichen associated yeasts remains to be discovered.

Metabolite profiling of the Cladonia lichens using gas chromatography-mass spectrometry

A metabolite analysis of 52 lichen samples belonging to 13 species of the genus Cladonia was carried out. Data of metabolite profiling were in good agreement with the results obtained using anatomical and morphological methods for lichens identification. The genetic heterogeneity of the studied groups is discussed.

The Lichens of Point Pleasant Park, Halifax, Nova Scotia

The peninsula on which the city of Halifax is located ends in a park that has remained mostly wooded since 1749, despite being periodically disturbed and partially cleared by military activities and storms. This first detailed study of the lichens of Point Pleasant Park is based on collections from almost 300 survey sites, which showed a remarkably diverse flora of 164 species, varying from pollution-tolerant lichens such as Lecanora conizaeoides at the northern end of the park, to members of the Lobarion community at the southern end. In 2003, Hurricane Juan felled a large number of the larger, older trees, which explains the current high proportion of crustose species established on the smaller, younger, trees. The baseline data reported in this study will be of value to follow the succession of lichens on trees as the bark surfaces change from smooth to ridged, with age, over the next few decades. The rich lichen flora of the park also reflects the fact that there are rock outcrops and vertical rock faces. These substrates support a lichen flora of 43 species, and the terricolous habitats are colonized by a further 23 species, including 18 species of Cladonia.

Reproductive and dispersal strategies shape the diversity of mycobiont-photobiont association in Cladonia lichens

Ecological preferences, partner compatibility, or partner availability are known to be important factors shaping obligate and intimate lichen symbioses. We considered a complex of Cladonia species, traditionally differentiated by the extent of sexual reproduction and the type of vegetative propagules, to assess if the reproductive and dispersal strategies affect mycobiont-photobiont association patterns. In total 85 lichen thalli from 72 European localities were studied, two genetic markers for both Cladonia mycobionts and Asterochloris photobionts were analyzed. Variance partitioning analysis by multiple regression on distance matrices was performed to describe and partition variance in photobiont genetic diversity. Asexually reproducing Cladonia in our study were found to be strongly specific to their photobionts, associating with only two closely related Asterochloris species. In contrast, sexually reproducing lichens associated with seven unrelated Asterochloris lineages, thus being photobiont generalists. The reproductive mode had the largest explanatory power, explaining 44% of the total photobiont variability. Reproductive and dispersal strategies are the key factors shaping photobiont diversity in this group of Cladonia lichens. A strict photobiont specialisation observed in two studied species may steer both evolutionary flexibility and responses to ecological changes of these organisms, and considerably limit their distribution ranges.

New or Otherwise Interesting Records ofCladoniaSpecies from Turkey

Kocakaya, M., Halici, M. G., Ahti, T. & Kocakaya, Z. 2018. New or otherwise interesting records of Cladonia species from Turkey. – Herzogia 31: 327–331.Six Cladonia species (C. conista, C. graeca, C. magyarica, C. peziziformis, C. scabriuscula and C. subturgida) are reported from Turkey; C. graeca, C. peziziformis and C. subturgida for the first time, for which comments on their habitats, substrates and distribution are provided.

TwoCladoniaSpecies New to Italy from Dry Habitats in the Po Plain

Gheza, G., Nascimbene, J., Mayrhofer, H., Barcella, M. & Assini, S. 2018. Two Cladonia species new to Italy from dry habitats in the Po Plain. - Herzogia 31: 293–303.Cladonia conista and C. pulvinata occur at several sites in dry lowland habitats in the Po Plain (northern Italy). Vegetation releves are provided in order to characterize the main lichen associations (Cladonietum rei, Cladonietum foliaceae, Pycnothelio-Cladonietum cervicornis) and vascular plant communities in which the two species were found in Thero-Airion dry acidic grasslands and in dry open Calluna heathlands.

Diversity loss of lichen pine forests in Poland

In Central Europe, deciduous forests are the dominant community type and lichen pine forests are restricted to certain areas with extremely nutrient-poor and xeric soil types. In recent decades, a retreat of vegetation of oligotrophic habitats has been observed in Central Europe. In this study, we assessed changes of lichen pine forests in Poland: within the main area of the range in Central Europe. We used two sets of data collected at a local and regional (nation-wide) scale. On the basis of data from semi-permanent plots, we examined changes in the structure and species composition of lichen pine forests over 33 years at the local scale (between 1975 and 2008). To compare trends at the regional scale, we used data collected in the Polish Vegetation Database (PVD). For identification of lichen pine forests we determined a group of co-occurring Cladonia species. We analyzed differences in species richness and vegetation structure at the regional scale in tree time periods (1) between 1951 and 1969, (2) 1970 and 1989, and (3) 1990 and 2011. We found that changes in lichen pine forests are primarily quantitative at both scales. Our results indicate that the abundance of Cladonia species is limited by strong competitors, i.e., vascular plants and bryophytes, which may be explained by eutrophication and climate warming. Only pine forests with a minor abundance of lichens have chances to persist in the vegetation of Central Europe, while the most valuable communities with high abundance of indicators will disappear. Though an assessment of the total decrease in the area of lichen pine forests is not possible with the available regional data, local observations indicate a large decline in the area of lichen pine forests in Central Europe. Their conservation seems to be a serious challenge, because it is difficult to provide optimal conditions for all indicators.

Genomic insights into the mitochondria of 11 eastern North American species of Cladonia

Cladonia is among the most species-rich genera of lichens globally. Species in this lineage, commonly referred to as reindeer lichens, are ecologically important in numerous regions worldwide. In some locations, species of Cladonia can comprise the dominant groundcover, and are a major food source for caribou and other mammals. Additionally, many species are known to produce substances with antimicrobial properties or other characteristics with potentially important medical applications. This exceptional morphological and ecological variation contrasts sharply with the limited molecular divergence often observed among species. As a new resource to facilitate ongoing and future studies of these important species, we analyse here the sequences of 11 Cladonia mitochondrial genomes, including new mitochondrial genome assemblies and annotations representing nine species: C. apodocarpa, C. caroliniana, C. furcata, C. leporina, C. petrophila, C. peziziformis, C. robbinsii, C. stipitata, and C. subtenuis. These 11 genomes varied in size, intron content, and complement of tRNAs. Genes annotated within these mitochondrial genomes include 15 protein-coding genes, the large and small ribosomal subunits (mtLSU and mtSSU), and 23–26 tRNAs. All Cladonia mitochondrial genomes contained atp9, an important energy transport gene that has been lost evolutionarily in some lichen mycobiont mitochondria. Using a concatenated alignment of five mitochondrial genes (nad2, nad4, cox1, cox2, and cox3), a Bayesian phylogeny of relationships among species was inferred and was consistent with previously published phylogenetic relationships, highlighting the utility of these regions in reconstructing phylogenetic history.

Contribution of selected lichens species of the genus Cladonia on the heathlands in Toruń (N, Poland)

The study presents data on the distribution of selected species of lichens – Cladonia furcata (Huds.) Schrad. , C. rangiferina (L.) Weber ex F.H. Wigg. and C. uncialis (L.) Weber ex F.H. Wigg. in Torun and along its southern boundary - on a heathland on the military training ground. A comparison was carried out of the research findings on the occurrence of the analysed species against variable habitat conditions over a period of nearly 40 years. The occurrence of Cladonia uncialis was most frequently observed within the city, in the habitats of Calluna vulgaris . A tendency was shown of selected lichen species to spread in the investigated area.

Genetic variation and factors affecting the genetic structure of the lichenicolous fungus Heterocephalacria bachmannii (Filobasidiales, Basidiomycota).

Heterocephalacria bachmannii is a lichenicolous fungus that takes as hosts numerous lichen species of the genus Cladonia. In the present study we analyze whether the geographical distance, the host species or the host secondary metabolites determine the genetic structure of this parasite. To address the question, populations mainly from the Southern Europe, Southern Finland and the Azores were sampled. The specimens were collected from 20 different host species representing ten chemotypes. Three loci, ITS rDNA, LSU rDNA and mtSSU, were sequenced. The genetic structure was assessed by AMOVA, redundance analyses and Bayesian clustering methods. The results indicated that the host species and the host secondary metabolites are the most influential factors over the genetic structure of this lichenicolous fungus. In addition, the genetic structure of H. bachmannii was compared with that of one of its hosts, Cladonia rangiformis. The population structure of parasite and host were discordant. The contents in phenolic compounds and fatty acids of C. rangiformis were quantified in order to test whether it had some influence on the genetic structure of the species. But no correlation was found with the genetic clusters of H. bachmannii.

Cladoniaceae of the Azores

Abstract: Pino-Bodas, R., Ahti, T. & Stenroos, S. 2017. Cladoniaceae of the Azores. — Herzogia 30: 445 — 462. Based on material collected from 40 localities on four islands of the Azores, three species of Cladonia are new to Macaronesia: C. conista, C. mauritiana and C. novochlorophaea and four are new to the Azores: C. bellidiflora, C. dimorpha, C. rei and Cladonia sp. In addition, several species are new to individual islands of the archipelago. Cladonia dactylota is new to continental Europe (SW France). Cladonia stereoclada, described from the Azores, is lectotypified. These data are complemented with annotations on species distribution and chemical variations. Uncertain specimens were sequenced to confirm the species identities.

The genus Cladonia (Lichenized Ascomycota) from Restinga vegetation of Espírito Santo state, Brazil: Supergroups Cladonia and Cocciferae

Abstract This paper deals with 14 species of the genus Cladonia, occurring at Restinga vegetation of Espírito Santo state, with eight belonging to the Supergroup Cladonia and six to the Supergroup Cocciferae. Cladonia corallifera, C. crustacea, C. subminiata, C. pityrophylla, and C. polyscypha are new records for the state. An identification key, comments and illustrations are also provided.