Gypsum lichens: A global data set of lichen species from gypsum ecosystems.

Lichens are significant components of the biological soil crust communities in gypsum ecosystems and are involved in several processes related to ecosystem functioning, such as water and nutrient cycles or protection against soil erosion. Although numerous studies centered on lichen taxonomy and ecology have been performed in these habitats, global information about lichen species from gypsum substrates or their distributional ranges at a global scale is lacking. Thus, we compiled a global data set of recorded lichen species growing on gypsum. This review is based on systematic searches in two bibliographic databases (Web of Science and the more specialized database Mattick's Literature Index) using various keywords related to the substrate or ecology (i.e., gypsum, gypsiferous, semiarid, saxicolous, terricolous). In addition, we revised lichen literature from countries with gypsum soils using Mattick's, Hamburg University's Worldwide checklist, and different national lichen checklists. Ultimately the review includes a total of 321 studies. This data set included 6114 specimen records belonging to 336 recorded lichen species from 26 countries throughout the world. The results showed large differences in the number of species recorded among countries, reflecting differences in the sampling effort. We provide a table with the number of studies and species in relation to gypsum surface in order to account for the bias produced by sampling effort. The number of studies carried out per country was not related to the gypsum surface but probably to other factors, such as accessibility to field sampling, economic or political factors, or the presence of a wider community of lichenologists. Thus, Spain and Germany hosted the highest number of recorded species (160 and 114 species, respectively). Outside the European continent, only a few countries had a large number of species: Morocco (46), United States (42), and Iran (37). Remarkably, countries from the southern hemisphere (i.e., Australia, Chile, Namibia, and South Africa) showed a low number of studies from gypsum lands, supporting the stated biases observed in sampling efforts among countries. Considering the most studied countries, the results show that Teloschistaceae was the most represented family in gypsum ecosystems followed by Verrucariaceae and Cladoniaceae. Regarding particular species, Psora decipiens and Squamarina lentigera were some of the most widespread and abundant species in these habitats. This data set constitutes a basic and first step toward a much more comprehensive database, to be periodically updated in future releases, which also serves to identify countries or territories where future studies should be accomplished. There are no copyright restrictions on the data; please cite this data paper if the data are used in publications and teaching events.

Cyanobacterial Variability in Lichen Cephalodia.

The ecological success of lichens is related to both myco- and photobionts which condition the physiological limits of the lichen symbioses and thus affect their ecological niches and geographic ranges. A particular type of lichen, called cephalolichen, is characterized by housing both green algal and cyanobacterial symbionts-the latter is restricted to special structures called cephalodia. In this type of lichen, questions related to specialization within species or within individuals are still unsolved as different patterns have previously been observed. In order to study the variability at the intrathalline, intraspecific, and interspecific level, cyanobionts from different cephalodia within the same thalli and from different thalli were genetically analysed in three cephalolichen species at two different forests (18 thalli, 90 cephalodia). The results showed variability in the cephalodial Nostoc OTUs in all the studied species, both at the intrathalline and intraspecific levels. The variability of Nostoc OTUs found in different cephalodia of the same thallus suggests low specialization in this relationship. Additionally, differences in OTU diversity in the three studied species and in the two forests were found. The variability observed may confer an increased ecological plasticity and an advantage to colonize or persist under additional or novel habitats or conditions.

Specialization patterns in symbiotic associations: A community perspective over spatial scales.

Specialization, contextualized in a resource axis of an organism niche, is a core concept in ecology. In biotic interactions, specialization can be determined by the range of interacting partners. Evolutionary and ecological factors, in combination with the surveyed scale (spatial, temporal, biological, and/or taxonomic), influence the conception of specialization. This study aimed to assess the specialization patterns and drivers in the lichen symbiosis, considering the interaction between the principal fungus (mycobiont) and the associated Nostoc (cyanobiont), from a community perspective considering different spatial scales. Thus, we determined Nostoc phylogroup richness and composition of lichen communities in 11 Nothofagus pumilio forests across a wide latitudinal gradient in Chile. To measure specialization, cyanobiont richness, Simpson's and d' indices were estimated for 37 mycobiont species in these communities. Potential drivers that might shape Nostoc composition and specialization measures along the environmental gradient were analysed. Limitations in lichen distributional ranges due to the availability of their cyanobionts were studied. Turnover patterns of cyanobionts were identified at multiple spatial scales. The results showed that environmental factors shaped the Nostoc composition of these communities, thus limiting cyanobiont availability to establish the symbiotic association. Besides, specialization changed with the spatial scale and with the metric considered. Cyanolichens were more specialized than cephalolichens when considering partner richness and Simpson's index, whereas the d' index was mostly explained by mycobiont identity. Little evidence of lichen distributional ranges due to the distribution of their cyanobionts was found. Thus, lichens with broad distributional ranges either associated with several cyanobionts or with widely distributed cyanobionts. Comparisons between local and regional scales showed a decreasing degree of specialization at larger scales due to an increase in cyanobiont richness. The results support the context dependency of specialization and how its consideration changes with the metric and the spatial scale considered. Subsequently, we suggest considering the entire community and widening the spatial scale studied as it is crucial to understand factors determining specialization.

Contrasting Environmental Drivers Determine Biodiversity Patterns in Epiphytic Lichen Communities along a European Gradient.

Assessing the ecological impacts of environmental change on biological communities requires knowledge of the factors driving the spatial patterns of the three diversity facets along extensive environmental gradients. We quantified the taxonomic (TD), functional (FD), and phylogenetic diversity (PD) of lichen epiphytic communities in 23 beech forests along Europe to examine their response to environmental variation (climate, habitat quality, spatial predictors) at a continental geographic scale. We selected six traits related to the climatic conditions in forest ecosystems, the water-use strategy and the nutrient uptake, and we built a phylogenetic tree based on four molecular markers. FD and climate determined TD and PD, with spatial variables also affecting PD. The three diversity facets were primarily shaped by distinct critical predictors, with the temperature diurnal range affecting FD and PD, and precipitation of the wettest month determining TD. Our results emphasize the value of FD for explaining part of TD and PD variation in lichen communities at a broad geographic scale, while highlighting that these diversity facets provide complementary information about the communities' response under changing environmental conditions. Furthermore, traits such as growth form, photobiont type, and reproductive strategy mediated the response of lichen communities to abiotic factors emerging as useful indicators of macroclimatic variations.

Intraspecific variability drives functional changes in lichen epiphytic communities across Europe.

Traditional approaches in trait-based community ecology typically expect that trait filtering across broad environmental gradients is largely due to replacement of species, rather than intraspecific trait adjustments. Recently, the role of intraspecific trait variability has been largely highlighted as an important contributor mediating the ability of communities to persist under changing conditions and determining the community-level trait variation, particularly across limited environmental gradients. Unfortunately, few studies quantify the relative importance of species turnover versus intraspecific variability mediating the response of communities different from vascular plants. Here, we studied the functional changes in epiphytic lichen communities within 23 beech forests across large latitudinal (ca. 3,000 km) and environmental gradients in Europe to quantify the relative contribution of species turnover and intraspecific variability and the role of climate controlling community-level trait changes. For 58 lichen species, we focused on a set of 10 quantitative functional traits potentially affected by climatic conditions and related to photosynthetic performance (n = 1,184 thalli), water use strategy (n = 1,018 thalli), and nutrient uptake (n = 1,179 thalli). Our results showed that intraspecific trait variability explained most of the functional changes in lichen communities in response to the latitudinal gradient. Further, such functional changes were determined by the covariation between intraspecific trait variability and species turnover, which varied in sign depending on the trait considered. Finally, different climatic predictors explained functional variation due to both intraspecific trait variability and species turnover. We propose that lichen communities cope with contrasting climatic conditions by adjusting the functional trait values of the most abundant species within the communities rather than by the replacement of the species. Consequently, intraspecific variability should be explicitly incorporated to understand the effect of environmental changes on lichen communities, even over large environmental variations, better. Our results challenge the universality of the hypothesis that species turnover chiefly drives functional trait changes across large environmental gradients and call for a wider test of such important assumptions in trait ecology in different organism types and ecosystems.

How much matching there is in functional, phylogenetic and taxonomic optima of epiphytic macrolichen communities along a European climatic gradient?

Adopting an integrative approach that explicitly includes the different facets of biodiversity is crucial to assess the response of biological communities to changing environments. The identification of the optimal climatic conditions where communities maximize their functional, phylogenetic and taxonomic diversity is useful to compare whether the optima of the different facets of biodiversity match. Using a wide climatic gradient across Europe, we quantified the functional, phylogenetic and taxonomic diversity of epiphytic macrolichen communities, which are valuable early-warning ecological indicators. We ordinated 22 environmental variables and simultaneously illustrated non-parametric regressions of the diversity metrics against the climatic space using the 'hilltop plot' method to detect the climatic conditions in which the different diversity facets peaked and to compare the match between them. Functional diversity predicted at least part of the peaks of phylogenetic and taxonomic diversity, but phylogenetic and taxonomic hotspots did not overlap. Epiphytic macrolichen communities maximized their functional and phylogenetic diversity in the southernmost forests, with the Mediterranean region appearing as a biodiversity hotspot. Regarding the studied traits, photobiont type and growth form showed clearly defined optima while the quantitative physiological traits and families' optima did not show this pattern in response to climate. The different facets of biodiversity were not surrogates of each other highlighting the need for an integrative approach to assess the effect of environmental changes on communities and to establish conservation priorities. As functional traits mediated the response of lichen communities to climate, preserving high functional diversity might indirectly preserve high phylogenetic and taxonomic diversity. Relevant ecological indicators useful to develop rapid assessment methods to evaluate the effects of climatic changes include the photobiont type and growth form. The lack of relation between quantitative traits and climate call for further research to unveil their role as ecological indicators of small-scale variables or as effect traits.

Intraspecific life history variation in contrasting habitats: Insights from an obligate symbiotic organism.

PREMISE OF THE STUDY: Life history theory predicts that plants in unfavorable habitats for juvenile growth and survival will commence reproduction at smaller sizes and exhibit higher reproductive allocations than those in favorable habitats. The scope of life history theory will increase if these predictions apply to a broad range of organisms. Populations of organisms in contrasting habitats may experience different demographic rates. Thus, we compared the demography and life history traits of a lichen species in contrasting habitats. METHODS: We compared the abundance, growth, mortality, and reproductive strategy (threshold size for reproduction and reproductive allometry) of epiphytic and saxicolous populations of the asexually reproducing lichen Lobarina scrobiculata in two oak forests in central Spain. KEY RESULTS: The growth rates of saxicolous individuals were two times faster than those of epiphytic individuals. Epiphytic specimens exhibited a smaller threshold size for reproduction and a higher reproductive allocation than their saxicolous counterparts. The populations hosted by trees were two times larger than those on rocks (13,788 vs. 6629 individuals, respectively). The mortality rate did not vary between habitats. CONCLUSIONS: Our results demonstrate that contrasting habitats selected for differences in the demography and life history traits of a lichen species. Consistent with life history theory predictions, in the habitat with slower growth, L. scrobiculata started to reproduce at a smaller size and its reproductive allocation was higher. This study extends the scope of life history theory and improves our understanding of life history patterns and variations in overlooked taxa such as lichens.

Variation in the reproductive strategy of a lichenized fungus along a climatic gradient.

Background and aims: Onset of reproduction and reproductive allocation patterns are key components of plant reproductive strategies. Life history theory predicts that plants in adverse environments for juvenile performance start reproduction at smaller sizes and exhibit higher reproductive allocation compared to their counterparts in favourable environments. Life history theory will gain in generality if its predictions are shown to apply to a broad range of organisms and modes of reproduction. This study tested whether the asexual reproductive strategy of a lichenized fungus changed along a climatic gradient. Methods: The variation in threshold size for asexual reproduction and asexual reproductive allocation of the lichen Lobarina scrobiculata was assessed in 18 populations (9665 individuals) along a climatic gradient spanning 800 km in latitude in Southern Europe. Using generalized linear models and standardized major axis regressions, the allometric relationships and the associated variation in climatic factors according to the changes in the threshold size for reproduction and reproductive allocation patterns were assessed. Key Results: The onset of reproduction was size-dependent and the reproductive allocation increased with individual size. Both the threshold size for reproduction and the reproductive allocation varied along the rainfall gradient. A lower threshold size for reproduction and higher reproductive allocation in drier, adverse locations were found. Therefore, populations in drier locations fitted the predictions of life history theory for sexually reproducing organisms in adverse environments for juvenile performance. Conclusions: This study highlights the applicability of the life history theory to fungi and to modes of reproduction other than sexual reproduction. Based on the intraspecific variation in the asexual reproductive strategy of a fungal organism with climatic factors, these findings expand the scope of life history theory predictions and increase our understanding of life history diversity and reproductive strategies across environments.

A species on a tightrope: Establishment limitations of an endangered lichen in a fragmented Mediterranean landscape.

PREMISE OF THE STUDY: Habitat loss and forest fragmentation affect the dispersal and establishment of species. Furthermore, populations growing far from the species' optimal climate might be less viable because good-quality habitat can be scarce and easily altered by smaller changes. The lichen Pectenia plumbea has oceanic climatic requirements, so in the Mediterranean region it needs the humidity provided by well-preserved forests to thrive, but most of this habitat has disappeared and the remnants are fragmented. In central Spain, this species occupies only a small proportion of the existing forests, so we aimed to determine whether this scattered distribution is due to limitations on dispersal or establishment. METHODS: We selected a Mediterranean fragmented forest surface in central Spain and extracted environmental variables from 371 plots. We modeled the presence and abundance of P. plumbea and developed species distribution models (SDMs) to detect all the suitable habitats inside the Cabañeros National Park area. KEY RESULTS: Pectenia plumbea was present in most of the habitats predicted as good-quality and was generally absent from the poor-quality zones (85.9% overall success). The abundance correlated fairly well with that predicted by the SDM (67%). Both models show that P. plumbea is linked to high temperature and precipitation. CONCLUSIONS: Good-quality habitat requirements for P. plumbea that are similar to oceanic conditions are found only in specific forested, stony slopes derived from historical land management. This habitat is scarce, but P. plumbea has successfully tracked all of these scattered areas via its high dispersal capacity.

Species delimitation and phylogeography of the Pectenia species-complex: A misunderstood case of species-pairs in lichenized fungi, where reproduction mode does not delimit lineages.

The main goal of this work was to study species boundaries in the genus Pectenia and elucidate the biogeographic history of the four currently accepted species. To accomplish this, we included 92 specimens across the range of Pectenia in Europe and northern Africa. We used three nuclear loci and assessed species circumscription using two Bayesian coalescent-based methods and the Bayes Factor approach. We reviewed the value of reproductive mode and other morphological features as predictors of monophyletic groups. Our results suggest that the production of asexual propagules and sexual structures are not characterizing monophyletic groups. The genus includes two morphologically well-characterized main lineages, where one lineage is composed by two sub-lineages that are with a case of cryptic speciation explained by a biogeographic pattern. We suggest treating the two lineages as two species, which are characterized by lobe structure: Pectenia plumbea and P. atlantica. Both of these species include samples with asexual propagules and apothecia, and thus do not correspond to any of the earlier morphologically defined species. The results of the biogeographic analysis indicate that the Mediterranean basin is the most likely ancestral distribution area of P. plumbea, whereas P. atlantica probably originated in Macaronesia.

The threatened epiphytic lichen Lobaria pulmonaria in the Iberian Peninsula: genetic diversity and structure across a latitudinal gradient.

The current genetic diversity and structure of a species plays a marked role in the species' future response to environmental changes. Identification of the factors that might ensure the long-term viability of populations along its distribution area is therefore important for conserving biodiversity. In this work, infraspecific genetic diversity and structure of the threatened lichen Lobaria pulmonaria was investigated along a latitudinal gradient, spanning the Spanish latitudinal range of L. pulmonaria. Eighteen populations in Northern, Central, and Southern Spain were analysed using six specific fungal microsatellites of L. pulmonaria. Genetic diversity indices were calculated and compared among populations. Genetic differentiation was assessed using AMOVA and Bayesian methods. Additionally, a redundancy analysis was used to estimate the relative importance of environmental factors on the genetic variation among populations. Annual precipitation was the only factor affecting the genetic diversity probably through its influence on population and thallus size of L. pulmonaria, and significantly higher levels of genetic diversity were detected in southern populations. Isolation by distance was not significant, being environmental variables most important factors controlling genetic variation in L. pulmonaria populations.

Environmental factors that drive the distribution and abundance of a threatened cyanolichen in Southern Europe: a multi-scale approach.

PREMISE OF THE STUDY: High-quality information about threatened species is required to prevent current global biodiversity losses. Lichens are important components of forest biodiversity and help to maintain ecosystem functioning. The epiphytic cyanolichen Lobarina scrobiculata is red-listed in Europe and North America, but knowledge of its ecology and distribution in Southern Europe is scarce. METHODS: We used a multispatial scale design to investigate the effects of macroclimate, forest structure, tree features, and microhabitat on the occurrence, abundance, spatial distribution, and performance of Lobarina scrobiculata in the Iberian Peninsula at plot and tree scales. Generalized linear models and mixed models were used for analysis. KEY RESULTS: We recorded ca 14000 individuals of the threatened species Lobarina scrobiculata from 22 populations in the Iberian Peninsula. Our results suggest that L. scrobiculata thrives mainly in oak forests with highly variable annual precipitation levels. At the plot scale, the L. scrobiculata abundance increased with annual precipitation and tree density (habitat quantity). At the tree scale, our models highlighted the importance of tree size and bark roughness (habitat quality) as the main drivers of species occurrence and abundance. We detected a marked spatial pattern on tree trunks, i.e., L. scrobiculata occurred preferentially on north-facing surfaces and close to the ground where humidity is higher. CONCLUSIONS: By integrating multiscale modeling, we analyzed a unique large dataset and these results are essential for understanding the ecology of this threatened cyanolichen. There is an urgent need to preserve the forests that this species currently inhabits as well as potential colonization sites.

Cardinal characters on a slippery slope--a re-evaluation of phylogeny, character evolution, and evolutionary rates in the jelly lichens (Collemataceae s. str).

Recent molecular systematic studies have indicated that the traits currently used for generic delimitation in the jelly lichens (Collemataceae s. str.), may not characterize monophyletic groups. Here we reconstruct the phylogeny of Collemataceae using Bayesian and maximum likelihood analyses based on mitochondrial (mtSSU rDNA) and nuclear (nuLSU rDNA, Beta-tubulin and MCM7) markers of 70 Collemataceae species. We studied the evolution of four morphological and ecological characters traditionally used to delimit genera and infra-generic groups. Finally, we tested if differences in branch-lengths between clades are due to differences in rates of molecular evolution. Eleven strongly supported groups were recovered in the resulting well-resolved and well-supported phylogeny. The presence/absence of a eucortex, which is currently used as the cardinal character to define genera in the group, does not characterize monophyletic groups corresponding to the genera as currently circumscribed. Ancestral state reconstruction indicates that the most recent common ancestor of the jelly lichens most likely was saxicolous/terricolous, lacked a tomentum, and had transversally septate ascospores. Although the cortex state could not be reconstructed for the ancestor of the family, our observations indicate that a lack of cortex may have an evolutionary advantage in saxicolous/terricolous species in semi-arid environments, as non-corticate species tends to be larger and occur in higher frequency and abundance in such regions, compared to corticate species. A significant evidence for faster evolutionary rates was found in a lineage mainly including taxa that occur in the wet tropics and humid temperate regions, compared to other lineages. We suggest that this can explain the greater diversity of Collemataceae in tropical and humid areas.

Does the reproductive strategy affect the transmission and genetic diversity of bionts in cyanolichens? A case study using two closely related species.

Observed levels of population genetic diversity are often associated with differences in species dispersal and reproductive strategies. In symbiotic organisms, the genetic diversity level of each biont should also be highly influenced by biont transmission. In this study, we evaluated the influence of the reproductive strategies of cyanolichen species on the current levels of population genetic diversity of bionts. To eliminate any phylogenetic noise, we selected two closely related species within the genus Degelia, which only differ in their reproductive systems. We sampled all known populations of both species in central Spain and genotyped the fungal and cyanobacterial components of lichen samples using DNA sequences as molecular markers. We applied population genetics approaches to evaluate the genetic diversity and population genetic structure of the symbiotic components of both lichen species. Our results indicate that fungal and cyanobiont genetic diversity is highly influenced by the reproductive systems of lichen fungus. We detected higher bionts genetic diversity values in the sexual species Degelia plumbea. By contrast, the levels of fungal and cyanobiont genetic diversity in the asexual species Degelia atlantica were extremely low (almost clonal), and the species shows a high specificity towards its cyanobiont. Our results indicate that reproduction by vegetative propagules, in species of the genus Degelia, favors vertical transmission and clonality, which affects the species' capacity for resources and competition, thereby limiting the species to restricted niches.

Multiple-scale environmental modulation of lichen reproduction.

It is necessary to understand how environmental changes affect plant fitness to predict survival of a species, but this knowledge is scarce for lichens and complicated by their formation of sexual and asexual reproductive structures. Are the presence and number of reproductive structures in Lobaria pulmonaria, a threatened lichen, dependent on thallus size, and is their formation sequential? Does any size-dependence and sequential formation vary along a climate gradient? Generalized linear mixed models were used to explore the effect of environmental predictors on the size and presence/abundance of each reproductive structure and to determine the probability of a given-sized thallus to develop any reproductive structure. The largest individuals are more likely to develop reproductive structures, and the lichen uses a mixed strategy of early asexual reproduction and late sexual. Macro and microclimatic variables also influenced reproductive capacity. Relationships among climate conditions and lichen size and reproductive capacity can compromise the future viability of the species in the most southern populations of Europe.

Effects of tropical montane forest disturbance on epiphytic macrolichens.

The high diversity of epiphytes typical of undisturbed montane tropical forests has been negatively affected by continuous deforestation and forest conversion to secondary vegetation. Macrolichens are an important component of these epiphytes. Because their physiology is strongly coupled to humidity and solar radiation, we hypothesized that microclimatic changes derived from forest clearing and logging can affect the diversity of these poikilohydric organisms. In southern Ecuador, we examined three types of forests according to a disturbance gradient (primary forests, secondary forests, and monospecific forests of Alnus acuminata) for the presence/absence and coverage of epiphytic macrolichens that we identified on 240 trees. We found that total richness tended to decrease when the range of the disturbance increased. The impoverishment was particularly drastic for "shade-adapted lichens", while the richness of "heliophytic lichens" increased in the drier conditions of secondary growth. Epiphytic composition also differed significantly among the three types of forests, and the similarity decreased when the range of the disturbance was greater. We concluded that a span of 40 years of recovery by secondary vegetation was not enough to regenerate the diversity of epiphytic macrolichens that was lost due to forest disturbances.

Loss of epiphytic diversity along a latitudinal gradient in southern Europe.

Latitudinal gradients that involve macroclimatic changes can affect the diversity of several groups of plants and animals. Here we examined the effect of a latitudinal gradient on epiphytic communities on a single host species (Fagus sylvatica) to test the core-periphery theory. The latitudinal span considered, covering two biogeographic regions, is associated with major changes in rainfall during the dry season. Because bryophytes and lichens are poikilohydric, we hypothesized that their species richness and composition might vary at different latitudes. We also speculated how epiphytic communities may respond to future climate change. The present study was carried out in Spain, and three latitudes that cover the distributional range of F. sylvatica were selected. The presence/absence and coverage of epiphytic lichens and bryophytes were identified on 540 trees (180 in each zone). We found consistent south to north change in the total richness and in the richness of bryophytes and of lichens separately, all of which tend to increase at higher latitudes due to the presence of several hygrophytic species. Epiphytic composition also differed significantly among the three latitudes, and the similarity decreased when the latitudinal span was greater. In addition, high species turnover was driven by the increased rainfall at higher latitudes. We conclude that epiphytic communities have a similar pattern to the predictors of the core-periphery theory from populations, and they suffer a great impoverishment in species richness at lower latitudes, coincident with the southern boundary of the F. sylvatica distribution.

Molecular phylogeny of Heteroplacidium, Placidium, and related catapyrenioid genera (Verrucariaceae, lichen-forming Ascomycota).

PREMISE OF THE STUDY: Verrucariaceae is a fascinating lineage of lichenized fungi for which generic and species delimitation is problematic due to the scarcity of discriminating morphological characters. Members of this family inhabit rocks, but they further colonize soils, barks, mosses, and other lichens. Our aim is to contribute to the DNA-based inference of the Verrucariaceae tree of life and to investigate characters that could be useful for proposing a more natural classification. We focused on catapyrenioid genera, which are often part of biological soil crusts, a cryptogam-dominated ecosystem contributing to soil formation and stabilization in arid environments. Understanding their evolution and taxonomy is essential to assess their roles in these fragile and important ecosystems. METHODS: A multigene phylogeny of Verrucariaceae including catapyrenioid genera is presented. We further examined the phylogenetic relationships among members of Heteroplacidium and Placidium. The evolution of selected characters was inferred using the latter phylogeny. KEY RESULTS: Anthracocarpon and Involucropyrenium were closely related to Endocarpon. Placidium comprised two monophyletic clades sister to Heteroplacidium. Inferred ancestral states of diagnostic characters revealed that the type of medulla and the pycnidia location were homoplasious within the Placidium clade. In contrast, the presence of rhizines was a synapomorphy for Clavascidium. CONCLUSIONS: Our results provide new information on the usefulness of characters for delineating groups in Verrucariaceae. Taxonomic changes are proposed to reflect more natural groupings: Heteroplacidium podolepis is transferred to Placidium, and Clavascidium is recognized as a different genus. Eight new combinations are proposed for Clavascidium.

Fine spatial pattern of an epiphytic lichen species is affected by habitat conditions in two forest types in the Iberian Mediterranean region.

Persistence and abundance of species is determined by habitat availability and the ability to disperse and colonize habitats at contrasting spatial scales. Favourable habitat fragments are also heterogeneous in quality, providing differing opportunities for establishment and affecting the population dynamics of a species. Based on these principles, we suggest that the presence and abundance of epiphytes may reflect their dispersal ability, which is primarily determined by the spatial structure of host trees, but also by host quality. To our knowledge there has been no explicit test of the importance of host tree spatial pattern for epiphytes in Mediterranean forests. We hypothesized that performance and host occupancy in a favourable habitat depend on the spatial pattern of host trees, because this pattern affects the dispersal ability of each epiphyte and it also determines the availability of suitable sites for establishment. We tested this hypothesis using new point pattern analysis tools and generalized linear mixed models to investigate the spatial distribution and performance of the epiphytic lichen Lobaria pulmonaria, which inhabits two types of host trees (beeches and Iberian oaks). We tested the effects on L. pulmonaria distribution of tree size, spatial configuration, and host tree identity. We built a model including tree size, stand structure, and several neighbourhood predictors to understand the effect of host tree on L. pulmonaria. We also investigated the relative importance of spatial patterning on the presence and abundance of the species, independently of the host tree configuration. L. pulmonaria distribution was highly dependent on habitat quality for successful establishment, i.e., tree species identity, tree diameter, and several forest stand structure surrogates. For beech trees, tree diameter was the main factor influencing presence and cover of the lichen, although larger lichen-colonized trees were located close to focal trees, i.e., young trees. However, oak diameter was not an important factor, suggesting that bark roughness at all diameters favoured lichen establishment. Our results indicate that L. pulmonaria dispersal is not spatially restricted, but it is dependent on habitat quality. Furthermore, new spatial analysis tools suggested that L. pulmonaria cover exhibits a distinct pattern, although the spatial pattern of tree position and size was random.

Effects of Mediterranean dehesa management on epiphytic lichens.

Spanish holm oak (Quercus ilex subsp. ballota) open woodlands (dehesas) maintain a high diversity of plants and animals compared to other forested Mediterranean habits, but little is known about the responses of epiphytic lichens to different management regimes that are applied to this woodland type. The present study was carried out in central-southern Spain and included four management regimes: agriculture, grazing of sheep, grassland grazed by wild ungulates (deer), and abandoned dehesas covered by shrubs. Total species richness and cover exhibited considerable variation among management regimes. Both parameters tended to decrease with the intensity of management, abandoned dehesas maintaining a higher number of species than more intensively managed habitats. Lichen composition also significantly differed among the four regimes. Nitrophytic species were clearly associated with more intensive management regimes (farming or livestock management), whereas non-nitrophytic species favored abandoned dehesas.