Corticolous lichens as potential bioindicators of fire history: a study in the cerrado of the Distrito Federal, central Brazil

Abstract. The aim of this study was to investigate the potential use of epiphytic corticolous lichens as indicators of fire history in the cerrado (savannas) of central Brazil. Work was carried out at the Reserva Ecológica do IBGE and the Jardim Botânico de Brasilia, 33 km outside Brasilia D.F., in ten plots of cerrado denso within the ‘Fire Project’ area. Each plot was subjected to a specific prescribed burning regime, with study sites varying from a plot protected from fire for over 20 years to a plot burned every 2 years. Research was carried out in two stages: (1) a preliminary survey of plots with different fire histories, measuring variables about the lichen habitat and the lichen communities present in the habitats; (2) lichen sampling in plots with different fire histories, where collection and identification of lichen species took place. Field techniques used included plotless sampling, and identification of lichens was carried out using taxonomic keys, both in the field and in the laboratory. The results show that fire is a major determinant of epiphytic corticolous lichen communities in cerrado denso vegetation. The abundance, distribution and recolonization of lichen communities can clearly be correlated with the frequency and behaviour, in terms of homogeneity and flame heights, of the fires that have occurred in each of the plots surveyed. Particular lichen species also show differential sensitivities to fire frequency and behaviour. This study shows that responses of corticolous lichens, at both the community and species level, can be used as bioindicators of fire history in areas of cerrado denso vegetation in central Brazil.     

Vertical distribution of epiphytic lichens on Quercus laurina Humb. & Bonpl. in a remnant of cloud forest in the state of Veracruz,México

In the tropics, corticolous lichen richness and cover tend to increase from the trunk base to the top of the crown of trees. In this study we calculated the total beta diversity of the lichen community along a vertical gradient on Quercus laurina in Mexican cloud forest. By comparing the richness and cover of the lichens by zone, we show that foliose and fruticose lichens are a minor component of the total lichen species richness, but have a higher cover than the crustose lichens. Five zones were identified along each phorophyte (n = 15) with a diameter at breast height >40 cm. A total of 92 species were identified. Of these, 38% were found only in a single zone, 51% were shared between the different zones and 11% occurred across all zones. Species richness and cover increased from the lowest to the highest zones of the phorophytes. Dissimilarity in species composition between the zones could be explained by species replacement. An indicator species analysis revealed that only a few species, e.g. Hypotrachyna vexans, H. cf. sublaevigata and Ramalina cf. sinaloensis prefer a particular zone. The results show that the lichen community associated with Quercus laurina phorophytes is highly diverse and suggest that species richness and cover are related to the zone and the various growth forms.     

POST-FIRE COLONIZATION OF A MEDITERRANEAN FOREST STAND BY EPIPHYTIC LICHENS

The post-fire colonization of a Quercus ilex forest by epiphytic lichens has been studied in Catalonia (NE Spain), eleven years after a fire. Specific richness and lichen biomass have been studied separately on Quercus ilex , Erica arborea , Rosmarinus officinalis . and Cistus albidus , to reveal possible differences among phorophytes in facilitating lichen establishment. Shrubs play an important role in colonization by common species whereas the stools of Quercus ilex offer a suitable substratum for rarer species. Cistus albidus , with the highest lichen diversity and 98·2% of the total biomass of macrolichens, is the most suitable phorophyte for lichen establishment.     

Population dynamics of the lichen genus Bulbothrix Hale as potential bioindicators of 'time-since-last-fire' in the cerrado of the Distrito Federal, central Brazil

This paper investigates the potential use of the corticolous lichen genus Bulbothrix Hale as a bioindicator of ‘time-since-last-fire’ in the cerrado (savannas) of central Brazil. Work was carried out at the Reserva Ecológica do IBGE and the Jardim Botãnico de Brasilia, 33 km outside Brasilia D.F., in plots of cerrado denso with known fire histories within the ‘Fire Project’ area. Measurements of abundance and thallus size were taken of all Bulbothrix individuals encountered up to 2 m on the trunks of three phorophyte species. The results show that abundances and thalli sizes of the Bulbothrix populations can be correlated with the ‘time-since-last-fire’ that had occurred in each of the plots. This study shows that is is possible to use characteristics of specific lichen populations as bioindicators of fire history.     

Epiphytic lichens on Juniperus communis – an unexplored component of biodiversity in threatened alvar grassland

Semi‐natural grasslands, among them thin‐soil calcareous grasslands (alvars), have great conservation value but have become increasingly rare in Europe. The main threat to alvar grasslands is the encroachment by juniper Juniperus communis and therefore it is usually removed during the restoration practice. Juniper can also be a host plant for many epiphytic lichens, but its role as a phorophyte is poorly known. We studied epiphytic lichen diversity on 126 junipers in 17 sites in western Estonia and found 140 lichenized taxa including several rare and red‐listed species. Using indirect and direct multivariate analyses (DCA, pCCA) and general linear models we revealed that both habitat and phorophyte properties affect lichen assemblies on juniper. Lichen species richness per site showed a unimodal relationship with compound factors of site productivity and juniper characteristics (stem circumference and juniper width). Lichen species richness per phorophyte was increasing with its size and with the proportion of dead branches, and was twice higher in plate alvars than in ryhk alvars. Also, the species composition in plate alvars differed from ryhk alvars by having 42 characteristic lichen species in plate alvars vs three indicators of ryhk alvars. The composition of lichens was significantly influenced by encroachment of alvars, e.g. by high juniper cover and shrub layer height, as well as by the proportion of dead branches and stem circumference of juniper. We conclude that the epiphytic lichen assemblies on junipers are threatened by grassland encroachment similarly to ground layer lichen assemblies. We suggest that some old and scencent junipers should be preserved during the restoration of alvar grasslands.     

Phorophyte specificity and environmental parameters versus stochasticity as determinants for species composition of corticolous crustose lichen communities in the Atlantic rain forest of northeastern Brazil

A transect of 47 mature trees was studied within an Atlantic rain-forest plot in northeastern Brazil to determinate effects of phorophyte specificity and environmental parameters vs. stochasticity on the structure of corticolous, crustose microlichen communities. A total of 150 lichen species was found, most being rare to extremely rare. Multivariate analysis of sample plots indicated subtle phorophyte preferences among certain lichen species, corresponding to differences in bark pH, degree of bark shedding, density and size of bark lenticels, and presence of milk sap. Individual and multiple regressions revealed correlations between lichen species richness; respectively, area cover and bark pH (negative); density and size of bark lenticels (negative); degree of bark shedding (negative); presence of milk sap (positive); and diffuse site factor (positive). No strongly delimited lichen communities were detected, but cluster analysis revealed three main groups and six subgroups with slightly different lichen species composition, each one with characteristic indicator species but with highly variable overall species composition. Beta diversity was high among samples and lacked spatial structure. However, beta diversity was significantly lower among samples belonging to the same tree species, independent of their spatial arrangement. It was concluded that community formation in tropical rain-forest understory lichens subtly correlates with two main environmental factor complexes—phorophyte bark characteristics and microclimate—but is to a large extent determined by the stochastic effects of species dispersal, especially of rare species.     

Content of heavy metals in thalli of the lichen <Emphasis Type="Italic">Hypogymnia physodes</Emphasis>: Sources of heterogeneity

Size and reproductive traits of thalli of the epiphytic lichen Hypogymnia physodes (L.) Nyl. and concentrations of heavy metals (Cu, Fe, Cd, Zn, Pb) in them were determined in the region of the Middle Ural Copper Smelting Plant, in two contrast zones of atmospheric pollution. The nonuniformity of populations in both size and reproductive traits and metal concentration was demonstrated. A high heterogeneity of subpopulations growing on different phorophytes was revealed. It is recommended to optimize data collection by taking samples from a larger number of trees (20–30) at a smaller number of thalli (10–20) from one phorophyte.     

Population size and fire intensity determine post‐fire abundance in grassland lichens

Questions: What is the variability in abundance of lichens on grassland soil between and within fields after prescribed fire? Is post‐fire lichen abundance an effect of pre‐fire population size? Location: Cedar Creek Natural History Area, Minnesota, USA. Methods: Lichen abundance, estimated as ground cover and dominated by Cladonia spp., was mapped in plots in two fields before prescribed burning on 06.10.2003 and 15.10.2003 for the first time since abandonment in the 1950s. The plots were resurveyed one year post‐fire. Results: Post‐fire cover of Cladonia spp. varied strongly between the fields, most likely due to different weather conditions between the burn events, which resulted in different fire intensities, one of low and one of high intensity. In the field that experienced the low intensity fire, post‐fire cover of Cladonia spp. was still relatively high, and showed a positive relationship with pre‐fire cover, while no such relationship was found after the high intensity fire. In that field Cladonia spp. experienced high mortality rates irrespective of pre‐fire cover. Conclusions: This study provides an example of how species response to disturbance can be a function of population size, but that this relationship can be non‐linear; lichens in grassland can survive a low intensity fire proportionally to pre‐fire population size, but experience high mortality rates above a fire intensity threshold. The applications of these results are that fire intensity matters to species response to prescribed fire, and that the persistence of climax lichen communities and biodiversity in the study system needs a broad range of fire intervals.     

Fate of epiphytes on phorophytes with different architectural characteristics along the perturbation gradient of Sabal mexicana forests in Veracruz,Mexico

Question: Vascular epiphytes and hemiepiphytes (E/HE) in neotropical forests account for a large fraction of plant richness, but little is known of how the interplay between phorophyte architectural characteristics and habitat perturbation affect communities of E/HE. Location: Sabal mexicana forests in a coastal area of Veracruz, Mexico. Methods: We compared communities of E/HE on phorophytes with different architectural characteristics – the palm S. mexicana and non‐palm phorophytes – in three environments: conserved sites, perturbed sites and small regenerated forest fragments. We combined traditional (abundance, species richness, similarity and complementarity indices) and more recent (phylogenetic diversity) metrics to describe the communities of E/HE. Results: Overall, we recorded 924 E/HE individuals (nine families, 16 genera and 21 species). The abundance and species richness of E/HE was higher on palms than on non‐palm phorophytes. Abundance‐based complementarities between phorophytes and sites were high. We detected clear changes in community structure of E/HE with habitat perturbation, but there were no effects on the phylogenetic diversity of the E/HE community. Palm phorophytes hosted a more phylogenetically diverse community of E/HE than did non‐palm phorophytes. Conclusions: Palm phorophytes are key elements supporting the conservation of resilient communities of E/HE in S. mexicana forest. Habitat fragmentation has a strong effect on the structure of the E/HE community in S. mexicana forests. Ferns are the group of epiphytes most severely affected by habitat perturbation, but we detected no significant effect on the phylogenetic diversity of the community.     

Different effects of rain and artificial watering on element zonation patterns in lichen thalli and bark: A study on Physcia biziana (Massal.) Zahlbr. v. leptophylla Ve˘zda

Translocation processes between the inner and outer rings of lichen thalli, as well as between lichens and bark substratum, induce zonation patterns, with a dynamic stratification of pollution history memory which is often affected by the intensity of the atmospheric events. Two experiments of passive biomonitoring are presented, aimed to measure metal concentrations and zonation patterns in lichens and bark, before and after raining events or after a water sprinkling treatment. Ten elements were monitored in the lichen Physcia biziana (Massal.) Zahlbr. v. leptophylla Ve?zda in a moderately polluted area of Campania region (Italy). Element concentrations changed with lichen age, with generally higher concentrations in the older than the younger tissues. Rain and sprinkling had differential effects on lichen and bark: a significant decrease of the overall concentrations was observed after rain in bark, whereas a slight increase was observed in lichen thalli; however, when lichen thalli were just rehydrated with a sprinkling treatment, some elements were found to increase in the younger portions of the thalli and decrease in the older portions. The extent to which zonation patterns account for site “pollution memory” or are affected by the short-term local atmospheric variability is discussed.     

Post-fire recolonization of dominant epiphytic lichen species on Quercus hypoleucoides (Fagaceae)

Following a forest fire (27 500 ha) in 1994, post-fire recolonization of Quercus hypoeleucoides by epiphytic lichens was documented as changes in lichen cover, number of small thalli, specific factors that affected reestablishment of lichens, and modes of dispersal. Three sites in the Chiricahua Mountains (Arizona, USA) were chosen according to the severity of fire damage—unburned, moderately burned, and severely burned. From 1994 through 1997, the amount of dead lichen cover significantly increased at the moderately burned site. For the same time period, the amount of live lichen cover significantly increased at the severely burned site. Numbers of new thalli increased significantly at the severely burned site each year but only in the last year (1996–1997) for the moderately burned site. Bark texture and proximity to trees with lichens were among the most important physical factors for recolonization. The most important means of dispersal for Flavopunctelia praesignis was fragmentation. For Punctelia hypoleucites, the primary means of dispersal was spores. Increases in live lichen cover and numbers of new thalli occur faster in severely burned areas probably due to the loss of lichens on tree trunks, which provides space and a lack of competition.     

Low modularity and specialization in a commensalistic epiphyte–phorophyte network in a tropical cloud forest

Species interactions can shape the structure of natural communities. Such sets of interactions have been described as complex ecological networks, an example of which is the commensal network formed by epiphyte–phorophyte interactions. Vascular epiphytes germinate and grow on phorophytes (support trees), assuming a horizontal distribution (among the phorophyte species) and a vertical distribution (from the base of the tree trunk to the crown of phorophytes, i.e., through ecological zones). Here, we investigated the organization of these structural dimensions of the epiphyte–phorophyte network in a Brazilian tropical montane cloud forest. The analyzed network, comprising 66 epiphyte species and 22 phorophyte species, exhibited a nested structure with a low degree of specialization, a typical pattern for epiphyte–phorophyte networks in forests. The network was slightly modular, with 65% of the species common to three modules, and had vertical structure corresponding to the vertical organization of the phorophytes. The size (diameter at breast height) of phorophyte individuals influenced the network structure, possibly due to the increase in habitat area, the time available for colonization by epiphytes, and a greater number of microenvironments. We found that the distribution of the epiphyte species differed between the phorophyte ecological zones, with greater richness in the lower portions and greater abundance in the upper portions of the phorophytes. The results provide relevant guidance for future research on the characteristics and the vertical and horizontal organization of vascular epiphyte and phorophyte networks. Abstract in Portuguese is available with online material.     

The weight of the crust: Biomass of crustose lichens in tropical dry forest represents more than half of foliar biomass

In recent years, our ecological knowledge of tropical dry forests has increased dramatically. However, the functional contributions of whole ecosystem components, such as lichens, remain mostly unknown. In these forests, the abundance of epiphyte crustose lichens is responsible for the characteristic white bark on most woody plants, conspicuous during the dry season, but the amount of resources that the lichen component represents remains unexplored. We estimated lichen biomass in a Mexican tropical dry forest using the bark area of trees, the dry mass of lichens per unit area and the percentage of bark covered by lichens, together with previously known tree densities. The lowest 2.5 m of the forests main trunks contained 188 kg/ha of lichen biomass, with lichens covering 85% of the available bark for trees <12 cm DBH and 38% for trees >12 cm. Total epiphytic lichen biomass was 1.34–1.99 Mg/ha. Lichen biomass represented 61% of the foliar biomass in the forest. To our knowledge, this is the first time that a lichen biomass estimate is provided for an ecosystem in which crustose lichens are the dominant lichen growth form. Crustose lichens are typically considered to contribute little to the total lichen biomass and to be difficult to include in ecological analyses. The high lichen biomass in this ecosystem implies a significant ecological role which so far is unexplored. We suggest the crustose lichen component should not be underestimated a priori in ecological studies, especially in ecosystems with abundant lichen cover.     

Fire,grazing history,lichen abundance,and winter distribution of caribou in Alaska's taiga

In the early 1990s the Nelchina Caribou (Rangifer tarandus) Herd (NCH) began a dramatic shift to its current winter range, migrating at least an additional 100 km beyond its historic range. We evaluated the impacts of fire and grazing history on lichen abundance and subsequent use and distribution by the NCH. Historic (prior to 1990) and current (2002) winter ranges of the NCH had similar vascular vegetation, lichen cover (P = 0.491), and fire histories (P = 0.535), but the former range had significantly less forage lichen biomass as a result of grazing by caribou. Biomass of forage lichens was twice as great overall (P = 0.031) and 4 times greater in caribou selected sites on the current range than in the historic range, greatly increasing availability to caribou. Caribou on the current range selected for stands with >20% lichen cover (P < 0.001), greater than 1,250 kg/ha (P < 0.001) forage lichen biomass and stands older than 80 yr postfire (P < 0.001). After fires, forage lichen cover and biomass seldom recovered sufficiently to attract caribou grazing until after ≥60 yr, and, as a group, primary forage lichen species did not reach maximum abundance until 180 yr postfire. Recovery following overgrazing can occur much more quickly because lichen cover, albeit mostly fragments, and organic substrates remain present. Our results provide benchmarks for wildlife managers assessing condition of caribou winter range and predicting effects of fires on lichen abundance and caribou distribution. Of our measurements of cover and biomass by species, densities and heights of trees, elevation, slope and aspect, only percentage cover by Cladonia amaurocraea, Cladina rangiferina, Flavocetraria cuculata, and lowbush cranberry (Vaccinium vitis-idaea) were necessary for predicting caribou use of winter range. © 2011 The Wildlife Society     

Epiphytic lichen synusiae and functional trait groups in boreo‐nemoral deciduous forests are influenced by host tree and environmental factors

Deciduous forests with temperate broad‐leaved tree species are particularily important in terms of biodiversity and its protection, but are threatened habitats in northern Europe. Using multivariate analyses we studied the effect of forest site type, environmental variables and host tree properties on epiphytic lichen synusiae as well as on the composition of species‐specific functional traits. Epiphytic lichens were examined on Acer platanoides, Fraxinus excelsior, Quercus robur, Tilia cordata, Ulmus glabra and U. laevis in two types of forests: Humulus‐type floodplain forests and Lunaria‐type boreo‐nemoral forests on the talus slopes of limestone escarpment (klint forests). Klint forests located near the seashore were under greater maritime influence compared to floodplain forests located in inland Estonia which experience stronger air temperature contrasts. In addition to stand level and climatic variables, tree level factors (bark pH, trunk circumference and cover of bryophytes) considerably affected the species composition of the lichen synusiae. Overall, 137 lichen species were recorded, including 14 red‐listed species characteristic of deciduous trees. We defined 13 lichen societies and showed their preference to forests of a specific site type and/or host tree properties. In forests of both types, most of the epiphytic lichens were crustose, and had apothecia as the fruit bodies and chlorococcoid algae as the photobiont. However, the proportion of lichens with a foliose or fruticose growth form, as well as the proportion of lichens with vegatative diaspores, were higher in floodplain forests. In klint forests with a stronger influence from the wind, crustose species completely dominated, while species with vegetative diaspores were rare and most species dispersed sexually. Lichens with Trentepohlia as the photobiont were characteristic of these forests, and lichens with lirellate ascomata were prevailing, indicating the great uniqueness of the kint forests for epiphytic lichens in the boreo‐nemoral region.     

The effect of traditional management burning on lichen diversity

Question: How does regular management burning of a northern, Calluna vulgaris‐dominated heathland affect the lichen diversity at the patch and landscape scale? Location: Mar Lodge Estate, Scottish Highlands, United Kingdom. Methods: 26 fire sites of different ages and 11 long‐term unburnt stands were surveyed to create a chronosequence of changing lichen diversity following burning. Data were analysed graphically, with a GLM and using a CCA. Results: Though the immediate effect of fire was to significantly reduce lichen diversity, it generally recovered within 20 years. There was a significant difference in the population dynamics between wet and dry moorland areas with terricolous lichens in the former site being replaced by pleurocarpous mosses. Older stands, unburnt for 25 years or more, generally had lower diversity than stands 10 to 15 years old. Changes in lichen diversity and community composition can be attributed to the development of Calluna stand structure following burning. Conclusions: Fire can be seen to play an important role in maintaining the diversity of lichens in heathland areas by providing a variety of stand‐structures and ages across the landscape that favours the development of greater beta‐diversity.     

Conservation of Vascular Epiphyte Diversity in Shade Cacao Plantations in the Chocó Region of Ecuador

To assess the contributions of rustic shade cacao plantations to vascular epiphyte conservation, we compared epiphyte species richness, abundance, composition, and vertical distributions on shade trees and in the understories of six plantations and adjacent natural forests. On three phorophytes and three 10 × 10 m understory plots in each of the agroforestry plantations and natural forests, 54 and 77 species were observed, respectively. Individual-based rarefaction curves revealed that epiphyte species richness was significantly higher on forest phorophytes than on cacao farm shade trees; detailed analyses showed that the differences were confined to the inner and outer crown zones of the phorophytes. No differences in epiphyte species richness were found in understories. Araceae, Piperaceae, and Pteridophyta were less species-rich in plantations than in forests, while there were no differences in Orchidaceae and Bromeliaceae. Regression analysis revealed that epiphyte species richness on trunks varied with canopy cover, while abundance was more closely related to soil pH, canopy cover, and phorophyte height. For crown epiphytes, phorophyte diameter at breast height (dbh) explained much of the variation in species richness and abundance. There were also pronounced downward shifts in the vertical distributions of epiphyte species in agroforests relative to natural forests. The results confirm that epiphyte diversity, composition, and vertical distributions are useful indicators of human disturbance and showed that while the studied plantations serve to preserve portions of epiphyte diversity in the landscape, their presence does not fully compensate for the loss of forests.     

Adverse Effects of Epiphytic Crustose Lichens upon Stem Photosynthesis and Chlorophyll of Populus tremula L.

Dry cork layer (phellem) in stems of Populus tremula transmitted 35–55 percent of incident irradiation, depending upon moisture content. A cover of crustose Lecanora lichens reduced transmission through phellem to 10 percent or less of incident irradiation. The bark contains photosynthetically active cells. Apparent quantum yield for photosynthetic O₂-evolution was 0.017 in bark covered with dry Lecanora compared with 0.070 in naked bark. The capacity for gross photosynthesis in high light (1090 μmol photons m^?2 s^?1) was reduced by 50 percent in Lecanora-covered bark. Lecanora did not reduce the ratio between variable and maximal chlorophyll a fluorescence (F_v/F_m). Chlorophyll content per unit area was similar in leaves and naked bark of Populus tremula. The chlorophyll content in the bark decreased with increasing chlorophyll content in Lecanora. Chlorophyll a/b ratio was 2.5 in the bark compared with 4.0 in leaves and in Lecanora, and the ratio decreased down the stems. The a/b ratio was 2.3 in Lecanora covered bark compared with 2.6 in naked bark. The changes in bark photosynthesis below a Lecanora crust were probably due to acclimation of bark photosynthesis to shade, since the lichen acids in the measured lichens neither suppressed photosynthetic O₂-evolution nor changed the F_v/F_m in bark disks.     

Detection probability and abundance estimation of epiphytic lichens based on height‐limited surveys

Questions: What are the detection probabilities of epiphytic crustose lichens on oak (Quercus robur) when only the lowest 2 m of the trunk are surveyed? How does the abundance of lichen species change with height above the ground, and is the change related to tree age? How well can total abundance (0‐6 m) be predicted based on data from the lowest 2 m? Which tree characteristics explain the vertical distribution of the study species? Location: Southeast Sweden. Methods: The occurrence and abundance of eight crustose lichen species were recorded on 35 oaks, 0‐6 m from the ground. Results: The detection probability was high (>96%) for seven out of the eight species. The abundance of six species declined significantly with increasing height. For five species, >69% of the total abundance (0‐6 m) was recorded on the lowest 2 m. The proportion of the total abundance present above 2 m increased significantly with age for three species. Models predicting total abundance based on data from the lowest 2 m and diameter explained >80% of the deviance for all except one species. The vertical distribution of the study species was explained by bark fissure depth, area and cover of macrolichens. Conclusions: For crustose lichens associated with old oaks, surveying only the lowest 2 m of the trunk yields reliable occurrence data and fairly good estimates of total abundance. However, before interpreting data from the lowest 2 m, knowledge of species vertical distribution, and how the distribution changes with tree age is essential.     

Lichens on branches of Siberian fir (Abies sibirica Ledeb.) as indicators of atmospheric pollution in forests

The abundance distribution of different ecological groups of lichens depending on bark pH has been studied on 1- to 24-year shoots of Siberian fir in the mountains of southern Siberia. Along with acidophytic lichens commonly found on the Siberian fir (Usnea sp., Bryoria sp., etc.), its young shoots are also colonized by nitrophytic species (Physcia tenella, Melanelia exasperatula, etc.), which is evidence for the increasing pH of shoot bark. The proportion of thalli of nitrophytic lichen species shows a significant positive correlation with the pH of the upper (dusted) bark layer and is greater in the Eastern Sayan (at bark pH averaging 5.4) than in the Western Sayan (pH 4.7). The trends revealed in this study may be used for indication of pollution and ecological monitoring of forest ecosystems.