Geography, not host identity, shapes bacterial community in reindeer lichens

Background and AimsTremendous progress have been recently achieved in host-microbe research, however, there is still a surprising lack of knowledge in many taxa. Despite its dominance and crucial role in boreal forest, reindeer lichens have until now received little attention. We characterize, for the first time, the bacterial community of four species of reindeer lichens from Eastern North America’s boreal forests. We analysed the effect of two factors (host-identity and geography) in the bacterial community composition, we verified the presence of a common core bacteriota and identified the most abundant core taxa.MethodsMorphological and molecular lichen species delimitation was performed based on the ITS region. The bacterial community of around 200 lichen samples was characterised using the 16S rRNA gene.Key ResultsOur results showed that host-lichen identity does not determine bacterial community composition in reindeer lichens, but we confirmed the influence of geography in shaping the diversity and abundance of bacteria associated to the species Cladonia stellaris from lichen woodlands. We also revealed that reindeer lichens share a reduced common core bacteriota composed exclusively by Proteobacteria.ConclusionsThe bacterial community in reindeer lichens is not host-selective. Northern lichen woodlands exhibit a significant higher diversity and abundance of bacteria associated to Cladonia stellaris. Nevertheless, the specific role of those bacteria as well as the process of host colonization remains to be determined. Elucidating these two aspects would be key to have a better understanding of the whole boreal ecosystems. The reduced and not diverse core bacteriota of reindeer lichens might be due to the larger size of our study area. The presence of the species Methylorosula polaris in the core bacteriota is evident and might have a particular importance for reindeer lichens.

A review of reindeer lichen (Cladonia subgenus Cladina) linear growth rates

Cladonia subgenus Cladina (the reindeer lichens) can be a dominant part of terrestrial ecosystems worldwide. They are particularly abundant in arctic-alpine and boreal regions, where they are a primary food source for woodland caribou/reindeer in winter months. Determining the growth rates of reindeer lichen is important for understanding and managing lichen regeneration following disturbances such as timber harvesting, mining, grazing, and wildfire. Regeneration and rehabilitation rates can be calculated with greater accuracy when growth rates are well understood. We provide a summary of 17 studies from 6 countries that determined the linear growth rates of three reindeer lichen groups, Cladonia arbuscula/mitis (mean = 4.7 mm/yr.), C. rangiferina/ C. stygia (mean = 5.1 mm/yr.), and C. stellaris (mean = 4.8 mm/yr.). We use linear growth rates as a proxy for over-all growth and biomass. Variables found to influence lichen growth rates are also discussed, which include light, moisture, temperature, air pollution, acid rain, precipitation, snow accumulation, substrate, age of individuals, and type of disturbance. These results can assist land managers in developing more accurate strategies for restoring lichens in disturbed areas.

Factors influencing the persistence of reindeer lichens (Cladonia subgenus Cladina) within frequent-fire environments of the Mid-Atlantic Coastal Plain, USA

Background Prescribed fire is increasingly used to accomplish management goals in fire-adapted systems, yet our understanding of effects on non-target organisms remains underdeveloped. Terricolous lichens in the genus Cladonia P. Browne, particularly cushion-forming reindeer lichens belonging to Cladonia subgenus Cladina Nyl., fit into this category, being characteristic of fire-adapted ecosystems, yet highly vulnerable to damage or consumption during burns. Moreover, inherently slow dispersal and growth rates raise questions about how to conserve these taxa in the context of fire-mediated restoration management. This research was undertaken to identify factors that contribute to Cladonia persistence within areas subject to repeated burning and involved tracking the fate of 228 spatially isolated individuals distributed across seven sites previously burned zero to two times. Site selection was determined by edaphic factors associated with a rare inland dune woodland community type known to support relatively high densities of Cladonia. Results Evaluated across all sites, the post-burn condition of Cladonia subtenuis (Abbayes) Mattick samples, categorized as intact (32%), fragmented (33%), or consumed (36%) individuals, approximated a uniform distribution. However, their status was highly variable at the different sites, where from 0 to 70% were assessed as intact and 11 to 60% consumed. Machine-learning statistical techniques were used to identify the factors most strongly associated with fire damage, drawing from variables describing the proximate fuel bed, growth substrate, and fire weather. The final descriptive model was dominated by variables characterizing the understory fuel matrix. Conclusions Areas with highly contiguous fuels dominated by pyrogenic pine needles were most likely to result in consumption of individual Cladonia, whereas those growing in areas with low fuel continuity or in areas dominated by hardwood litter were more likely to persist (intact or as fragments). Further, substrates including bare soil and moss mats afforded more protection than coarse woody debris or leaf litter in settings where fuels were both contiguous and highly flammable. Our findings describe the characteristics of within-site fire refugia, the abundance of which may be enhanced over time through restoration and maintenance treatments including thinning, promotion of mixed-species overstory composition, and periodic burning. Because lichens contribute to, and are considered reliable indicators of forest health, fire-based restoration management efforts will benefit from improved understanding of how these vulnerable organisms are able to persist.

Terrestrial Bryophyte and Lichen Responses to Canopy Opening in Pine-Moss-Lichen Forests

Pinus contorta-dominated montane forests of western Canada with relatively dense tree canopies have ground layers with abundant bryophytes, especially the feather mosses (Pleurozium schreberi and Hylocomium splendens), while those with more open canopies are dominated by species of reindeer lichens, especially Cladonia arbuscula s.l. and C. rangiferina s.l. Woodland caribou (Rangifer tarandus caribou), which are a threatened species in Alberta, prefer open, Cladonia-dominated forests for their winter food supply. This study investigated if opening the canopy by thinning mature montane forests of the Canadian Rocky Mountain foothills would change the abundance of lichens and bryophytes. In 1997, forests were thinned by removing 20%, 40%, and 60% by volume. In 2016, 19 years after treatment, we re-surveyed a subset of these plots (n = 97) for lichen and bryophyte abundance and species richness by utilizing the amount of canopy opening at the plot level as our prime gradient. We then used ordination to determine the relationship of control plots to treatment plots. In uncut forest, the control plots were highly variable, but were mostly dominated by feather mosses, with little or no bare ground. Feather moss abundance was lower in treatment plots when compared to control plots, while cover of bare ground was greater. Overall, 19 years after treatment, we found that, in treatment plots, lichen abundance remained stable or slightly increased, feather mosses decreased markedly, and unoccupied space was double that of the control plots. We conclude that the canopy opening had little effect on understory and ground layer diversity, but considering species abundance (1) bryophytes have not recovered after canopy opening, (2) populations of reindeer lichens increased marginally, but have not colonized areas left bare from bryophyte dieback, and (3), after 19 years there, remains unoccupied areas of bare ground in plots with a reduced canopy cover. Our study demonstrated that, with canopy cover reduction resulting from forest thinning operations, the ground layer diversity is maintained, but recovery of ground layers in old-growth pine-dominated forests is not promoted. Therefore, timber harvest that partially opens the tree canopy is unlikely to benefit caribou by augmenting or accelerating winter food availability and habitat suitability for caribou.

Phylogenetic relationships among reindeer lichens of North America

Cladonia is one of the largest lichen-forming ascomycete genera. It was formerly divided into ten sections, three of which, Crustaceae (Cladina), Tenues, and Impexae, are called the reindeer lichens. While previous studies have elucidated the relationships between species and sections, they often examined only one or a few specimens of each species in the analysis. This study examined the monophyly of selected members of sections Crustaceae, Tenues, and Impexae and their relationships in the genus Cladonia using the internal transcribed spacer region of the nuclear ribosomal DNA (ITS rDNA) and the mitochondrial small subunit gene of the mitochondrial ribosomal DNA (mtSSU). The phylogenetic tree contained four clades, two representing species in section Impexae, one representing species that belong to sections Crustaceae and Tenues, and one clade with C. arbuscula and related species. Five of 22 species, C. pycnoclada, C. stellaris, C. evansii, C. ciliata and C. subtenuis, showed monophyly in the phylogenetic tree; some of these 5 species have been shown previously to be monophyletic. The thallus branching pattern was interpreted as an important heritable character using the mtSSU network. Three duplets of paraphyletic species were further examined using ITS rDNA haplotype networks and AMOVA analysis. The results for the species duplets showed some mixing of haplotypes but the AMOVA analysis provided support for species separation within the duplets. While the evidence supports distinct species, further study is needed to conclusively show separate species in these duplets.

Phylogeny of Cladonia uncialis (Cladoniaceae, Lecanoromycetes) and its allies

The species from Cladonia section Unciales are characterized by the absence of squamules and soredia on the corticate podetia and the presence of usnic acid. Different subspecies, varieties and forms have been distinguished in the type species C. uncialis. In this study, a molecular phylogeny of Cladonia uncialis and members of the traditionally recognized section Unciales, along with additional potentially allied species, was constructed. DNA sequences from three gene loci, namely ITS rDNA, IGS rDNA and ß-tubulin, were analyzed using Maximum Parsimony, Maximum Likelihood, and Bayesian methods. Eleven species were analyzed for the first time using DNA sequence data. Cladonia uncialis subsp. uncialis and subsp. biuncialis were recognized as distinct taxa. The recognition of C. pseudostellata as a species was not supported by the analyses, but it represents a hypothamnolic acid chemotype, which is reported here as new to Europe (Scotland). The presence of subsp. biuncialis in North America (Newfoundland) was substantiated. The subsp. uncialis usually lacks squamatic acid, but in the eastern United States a morph referred to as subsp. uncialis does normally contain that acid. However, this morph did not attain taxonomic recognition based on phylogenetic analyses. All the other taxa formerly included in sect. Unciales turned out to belong to other groups of Cladonia, mainly Amaurocraeae, Borya, Divaricatae, and Perviae. The formerly recognized genus Cladina (reindeer lichens) is non-monophyletic, consisting of three groups within Cladonia, making the concept Cladina even nomenclaturally useless. Alternative topology tests rejected the monophyly of C. pseudostellata, section Unciales and Cladina.