Epiphytic lichen diversity in the mediterranean beech forest in Madrid province (Spain)

Amo, G. & Burgaz, A. R.
Dpto. Biología Vegetal I, Facultad de CC. Biológicas, Universidad Complutense de Madrid, 28040-Madrid, Spain

The study area is a relictic forest called "Hayedo de Montejo", which is located in the center of the Iberian Peninsula and it represents one of the southern limits of the distribution of Fagus sylvatica in Europe.
Thirty-two relevés carried out have been analized using numerical (cluster analysis, PCA, CCA and General Lineal Models) and traditional floristic characteristics. The results indicate the presence of Nephrometum resupinati in the Lobarion pulmonariae (Fuertes & Burgaz, 1989). These Lobarion species are located in north-facing aspect on the trunk and the community is found a short distance from the ground. The location of these species is accompanied by an increase of the bryophytes cover. The lichens in other aspects of the trunk and other height belong to the parmelioid community Orthotrichetum lyellii subas. parmelietosum glabrae in Frullanion dilatatae (Fuertes et al., 1996). Our study deepens in the knowledge of the distribution on the trunk, according to the microclimatic conditions, of the present species on Fagus sylvatica.
A catalogue of 125 epiphytic lichen is presented. Some interesting species are reported, as Sclerophora peronella (first record to the Iberian Peninsula), Sclerophora nivea, Heterodermia japonica and several important extension of range.
This Natural Reserve is an old pasture woodland and, in our opinion, the changes made in the forest´s microclimate hinder the Lobarion community´s growth, except in this situation where the species find refuge. In spite of this, many species have disappeared from the area, as Nephroma parile. Other species are very rare, as Pannaria conoplea, Degelia plumbea or Lobaria scrobiculata, only found near the rivers. In the Mediterranean Region, the influence of the rivers allows, the survival of threatened species.

Contrasting fundamental and realized ecological niches of epiphytic lichens in an old-growth Pseudotsuga forest

Antoine, M. E. (1) & McCune, B. (2)
(1) Department of Biological Sciences, Humboldt State University, Arcata, California, 95521 USA; (2) Department of Botany & Plant Pathology, Oregon State University, Corvallis, Oregon, 97331 USA

Vertical stratification of forest epiphytes is well documented. While it is easy to describe epiphyte distribution, the underlying causes are more elusive. Here we consider the vertical gradient in an old-growth Pseudotsuga forest, and we examine two aspects of species' responses along this gradient: fundamental and realized niches. We measured growth rates of four epiphytic lichen species using pendant transplants. By excluding the effects of competition and establishment, we defined the vertical range in which the species were physiologically capable of living (fundamental niche). We also quantified the vertical distribution and abundance of the four species, thereby assessing their realized niches. Comparing profiles of growth rates to profiles of natural abundance showed an intriguing discrepancy. The cyanolichen Lobaria oregana peaked in abundance in the mid-canopy, but its growth was maximal in the upper canopy, where it was virtually absent. What factor(s) account for this difference between L. oregana's fundamental and realized niches? We suggest that poor dispersal, establishment, persistence, or competition may exclude L. oregana from the upper canopy, perhaps due to an inability to tolerate rare microclimatic extremes. Transplanting L. oregana into the mid and upper canopy with resident communities either removed or left intact can test these hypotheses. Long-term microclimatic monitoring can be combined with periodic visitation to assess establishment of propagules and mortality and growth of thalli. If poor dispersal precludes L. oregana in the upper canopy, then inoculated substrates will have more juveniles. If microclimate limits establishment and/or persistence in the upper canopy, then mortality peaks will be associated with microclimatic extremes. If competition excludes L. oregana from the upper canopy, then lower establishment and growth rates will occur where resident communities are intact.


Epiphytic lichens and bryophytes in Quercus: colonization, succession, and the effects of fire

Bennett, S. & Sillett, S. C.
Humboldt State University, Department of Biological Sciences, Arcata, California 95521, USA

The distribution and abundance of epiphytic lichens and bryophytes have been well studied, but juvenile recruitment in situ and community responses to low-intensity fire have not. In this study, we described the distribution and abundance of epiphytic lichens and bryophytes in eight entire Quercus garryana tree crowns, documented associations among epiphyte species, quantified juvenile recruitment of epiphytic macrolichen species, and compared epiphyte community dynamics between burned and unburned trees. We identified 94 epiphyte species (25 cyanolichens, 56 chlorolichens, and 13 bryophytes). Lobaria pulmonaria was the most abundant lichen, and Orthotrichum papillosum was the most abundant bryophyte.
Epiphyte community composition was strongly influenced by branch size/age. Lichen cover and bryophyte cover were negatively correlated. Chlorolichens were more abundant on twigs and small branches, bryophytes were more abundant on larger branches, and cyanolichens were more abundant on intermediate branches between them. Eleven macrolichen species (5 cyanolichens and 6 chlorolichens) had lower mean cover in burned than unburned trees. The cyanolichen Sticta fuliginosa occurred only once in burned trees. Bryophyte abundance was not affected by fire. 92,090 macrolichen juveniles were encountered. One chlorolichen species, Parmelia sulcata, accounted for 61% of all juveniles. The total number of juveniles was lower in burned than unburned trees. Five macrolichen species (2 cyanolichens and 4 chlorolichens) had lower average numbers of juveniles in burned trees. Fire did not affect overall species distribution patterns. Results of this study indicate that fire may have a negative effect on some species of epiphytic lichens. Lichens with flammable neighbors (e.g., tufted, fruticose species) may be especially vulnerable to fire (i.e., "flammable neighbor hypothesis").


Fungal diversity in rock beneath a crustose lichen as revealed by molecular markers

Bjelland, T. & Ekman, S.
University of Bergen, Bergen, Norway

Lichen-forming fungi have been assumed to be more or less restricted to the surface of the substrate on which they grow. Conclusive identification of hyphae or an assessment of the fungal diversity inside lichen-covered rock has not been possible using methods based on direct observation. We circumvented this problem using a DNA sequencing approach. Cores were drilled from a Devonian arcosic sandstone rock harbouring the crustose lichen Ophioparma ventosa (L.) Norman on the surface. The cores were cut vertically, and DNA was extracted from the pulverized rock slices. A series of polymerase chain reactions using fungal-specific primers as well as Ophioparma ventosa specific primers were employed to amplify the internal transcribed spacer (ITS) region of the nuclear ribosomal DNA. The results show that hyphae of O. ventosa penetrate approximately 10-12 mm into the rock. Consequently, the hyphal layer formed by the lichen fungus inside the rock could be 7-12 times as thick as the symbiotic thallus at the surface of the rock. In addition, eight non-lichenized fungal species and five that could not be identified to species were encountered. One fungal species in the order Helotiales occurs in six of the eight cores. The significance of these results to the colonisation and weathering of rock by lichenised fungi is discussed.


The ecological analysis of lichens in the Teberda State Biosphere Reserve (the North-Western Caucasus, Russia)

Blinkova, O.V.
Lomonosov Moscow State University, Moscow, Russia

The Teberda State Biosphere Reserve is located on the northern side of the Great Caucasus mountain range. The total reserve's area is about 850 sq. km. The main section of the Reserve occupies the upper reaches of the Teberda river valley between 41° 35' and 41° 55' E, and 43° 13' and 43° 28' N. 24 canyons with total length about 150 km were inspected with an itinerant method. Elevation of the investigated territory ranges from 1259 m a.s.l. to 3800 m a.s.l. For an ecological summary we report lichen species by growth forms, substrate groups, and distribution of lichens on woody species.
389 species of lichens were recorded in the investigation. There are more crustose species (189 or 48,7%) than foliose species (123 or 31,4%) or fruticose species (77 or 19,8%). We distinguish 7 substrate groups of lichens. Corticolous (174 species or 44,8%), saxicolous (136 species or 34,2%) and terricolous (90 species or 32%) species form the core of lichen flora. Though some lichens regularly colonize a wide variety of substrate groups, most show a distinct preference for a single substrate type (284 species or 73,1%).
Corticolous species of lichens were found on 24 woody plant species. The greatest variety of lichen species was on Abies nordmanniana, Acer platanoides, Pinus sylvestris, and Fagus orientalis (they are the most widespread woody species in the Teberda reserve). The greatest number of corticolous species of lichens were from Lecanoraceae, Parmeliaceae and Physciaceae families.


Microclimate at the canopy of some tropical lichens

Boonpragob, K., Osatanond, N. & Polyium, W.
Department of Biology, Faculty of Science, Ramkhamhaeng University, Bangkok, Thailand

The microclimatic factors temperature, light intensity and relative humidity, were recorded for lichen growth zones and the edges of these zones in the canopies of tropical rain forest, lower montane forest and dry dipterocarp forest sites at Khao Yai National Park in Thailand. Measurements were taken every 10 minutes for 24 hours in the hot season, the rainy season and the cool season. Among the most important factors promoting lichen establishment and growth are light intensity over 200 umol m-2 s-1 in early morning and nearly saturated relative humidity during the night in every season. Temperatures range between 17–40° C, which are in the biological active zone. The maximum temperatures were recorded during the hot dry summer season, when lichens were inactive.


Contribution to the distribution of genus Ramalina in Portugal

Carvalheiro, L. (1), Figueira, R. (1, 2) & Carvalho, P. (1)
(1) Jardim Botânico da Universidade de Lisboa-Museu Nacional de História Natural, R. Escola Politécnica, 58, 1250-102 Lisboa, Portugal; (2) CVRM – Geo-Systems Centre, Instituto Superior Técnico, Av. Rovisco Pais, 1049-001 Lisboa, Portugal

The distribution of the Ramalina genus in Portugal is updated with new records of epiphytic specimens from the south-west Atlantic coast, where a natural park and a natural reserve are located. The study area is under a biomonitoring program to assess the atmospheric deposition of heavy metals and saline elements. The Ramalina genus was previously used in a section of the area to monitor the airborne salt deposition, but an inventory of the distribution of the genus is necessary to decide the better species for further biomonitoring in a larger region. An additional goal is to improve understanding of the ecology of Ramalina species, by verifying the importance of the proximity of the sea to their distribution. Seventy sampling sites were selected in the area, which comprises about 160 km of coast by 20 km inland. In each site, the total biodiversity of epiphytic Ramalina lichens was recorded, for any of the present phorophytes, and frequency releves, based on a sampling grid, were performed on sea facing (west) and opposed (east) sides of pine tree stems. The frequency of species in branches was also determined. Each site was characterized by local observation and using Geographic Information Systems, in order to fulfill a matrix of environmental variables to be cross analyzed with the frequency of the species. The presence of Ramalina elegans is for the first time referred for Portugal, in eleven sites of Alentejo and Algarve. The distribution of the species also shows new interesting sites for R. obtusata, in Algarve. The interpretation of frequency data by multivariate methods shows that some species (e.g. R. canariensis, R. lusitanica, R. pusilla) have opposite relationships with distance from the coast, while others (e.g. R. elegans) displays lower frequencies near the coast. The study also presents new interesting data on
the chemistry of R. lusitanica.


The Collema genus in the Iberian Peninsula

Carvalho, P. (1) & Figueira, R. (1, 2)
(1) Jardim Botânico da Universidade de Lisboa-Museu Nacional de História Natural, R. Escola Politécnica, 58, 1250-102 Lisboa, Portugal; (2) CVRM – Geo-Systems Centre, Instituto Superior Técnico, Av. Rovisco Pais, 1049-001 Lisboa, Portugal

Although a good taxonomic and systematic work was made by Degelius in Europe and extra-European species of Collema, a monographic study about this genus was missing in the Iberian Peninsula. The aim of this work is to present new distributional data of species of Collema genus in the Iberian Peninsula, in order to improve knowledge about the ecological range of the species belonging to this genus. Another goal is to define potential sites with special interest for lichen biodiversity and conservation, using Collema species as indicators. The indicator value is based on the number of species found and multivariate models of the relationship between taxa and ecological parameters, determined for each locality, by Geographic Information Systems. Using these tools, it is possible to build maps of potential distributions of species, that after comparison with known distributions, allows the detection of regions that still require more sampling effort.
The study in the Iberian Peninsula shows that the Collema genus is represented by 33 species (31 in Spain and 28 in Portugal), occurs on the most diverse types of substrates, and presents an high degree of morphologic and ecological diversity. Two of them are new references for the region [Collema glebulentum (Crombie) Degel. and Collema euthallinum (Zahlbr.) Degel.]. The distribution maps are based on material collected by the authors, and on several revised herbarium samples (BCC, GZU, INA, JACA, LEB, LISU, M, MA, MGC, MU, SANT, TSB, PO and VIT), with a total of about 3500 specimens.


Adaptation of epilithic lichens to high levels of UV irradiation

de la Torre Noetzel, R. (1), Sancho, L. G. (2), Horneck, G. (3), Scherer, K. (3), Facius, R. (3) & Deutschmann, U. (3)
(1) Department of Earth Observation, INTA (Spanish Aerospace Research Establishment), 28850-Torrejón de Ardoz, Madrid, Spain; (2) Universidad Complutense de Madrid, Dpm. Biología Vegetal II, Madrid, Spain; (3) DLR, Institute of Aerospace Medicine, Radiation Biology, 51170-Köln, Germany

Epilithic ecosystems in high mountains are exposed to an intense UV radiation and extreme microclimatic conditions. Rhizocarpon geographicum, an epilithic species growing at Sierra de Gredos (Central Spain), has developed a thick, pigmented cortex to protect the photobiont, which helps to adopt a latent status without metabolic activity, at summer, when temperatures, PAR and UV radiation are very high, and dryness prevails. The aim of this work was to test the resistance or survival capacity of selected samples of Rhizocarpon geographicum on the Plataforma de Gredos (2.000 m a.s.l.), with cortex or with the cortex removed, during the growing season (May, June, 2001) and in high summer (July 2003). We have used the DLR biodosimeter (Biofilm technique) to measure UV radiation (UV-A, UV-B, UV-A+B) close to the samples, and a Squirrel (Grant, U.K.) station to register microclimate. Photosynthetic activity was recorded with a MINI PAM Fluorometer, preadapting samples to dark before the first and final measurement. Results showed a high reduction of photosynthetic activity when the cortex was removed, in relation to samples with cortex. In both cases the lowest values were around 12:00 h GMT. In samples with cortex, photosynthesis recovered to initial values at 15:00 h-16:00 h GMT during the campaigns in May and June, but not in July. Recovery was less in samples without cortex.


Lichens on bare ground spots on reindeer pastures in mountain tundra of the Polar Urals

Ektova, S.
Institute of Plant and Animal Ecology of UD RAS, Ekaterinburg, Russia

Summer reindeer grazing is the most important kind of human impact to ecosystems of the Polar Urals. Lichens can be characterized as the component of plant cover most sensitive to reindeer grazing. Lichen species diversity changes, as well as decrease of height, cover, and productivity of fodder lichens were found in a quantitative survey of reindeer pastures.
In mountain tundra lichens generally are dominants and co-dominants with coverage of about 50%. In overgrazed mountain tundra, which is characterized by stony soils, decrease of total plant cover often occurs. On trails and places of intensive trampling damage to not only lichens, but also dwarf shrubs, herbs and mosses takes place. Bare ground spots are formed whose total vegetative cover is 30% or less. Mosses, crustose and foliose lichens with small lobules die back as a result of trampling. Fruticose lichens are represented by fragments of podetia, often dying, whose height is 5 mm or less. Lichens comprise less than 5-10% of the reduced vegetative cover.
We found 41 species of lichens on bare ground spots. In general, they are species characterized by high tolerance to grazing, but which have low fodder value. Frequency and cover of those species increase as the rate of stocking increases. 29 of those species belong mostly on epilithic substrates; they are not found on soil in ungrazed areas. Flavocetraria nivalis and Stereocaulon paschale dominate on bare ground sites: trails and places of intensive trampling. Moderately frequent species on bare ground sites include Allantoparmelia alpicola, Arctoparmelia centrifuga, Parmelia saxatilis, Melanelia commixta, Pseudephebe pubescens, Xanthoparmelia somloënsis, Brodoa intestiniformis, Phaeophyscia, Physcia caesia, Candelariella vitellina, Lecanora cenisia, Biatora vernalis, Pertusaria dactylina, Polyblastia terrestris, Ochrolechia inaequatula.


Influence of forest edges on Alectoria sarmentosa in a natural forest landscape

Esseen, P.-A.
Dept. of Ecology and Environmental Science, Umeå University, Sweden

Habitat loss and fragmentation profoundly affect biodiversity in managed forests. Recent studies have shown that the abundance and vitality of epiphytic lichens are strongly influenced by edge effects in fragmented forest ecosystems. In this study I examine the response of an old-growth associated lichen, Alectoria sarmentosa, to forest edges in a naturally heterogeneous boreal landscape. The study area, Granlandet, is 26 000 ha large and is located in northern Sweden. Granlandet is a mosaic of about 1000 forest islands of different size and shape dispersed in a matrix of open Sphagnum-dominated wetland. I quantified the abundance of A. sarmentosa in the lower canopy of Picea abies in relation to distance from forest edge, size of the open habitat and edge orientation. Results show that A. sarmentosa is strongly affected by both proximity to edge and exposure. The maximum length was four times lower and the biomass was ten times lower at the forest edge compared to the forest interior. The abundance of A. sarmentosa showed a monotonic decrease from the interior to the edge. Maximum distance-of-edge-influence (DEI) was estimated to over 60 m, which is beyond that reported from fragmented forests. The response of epiphytic lichens to forest ecotones is apparently due to complex interactions among several factors such as microclimate, landscape structure, forest composition, physiological characteristics, dispersal strategy, and position in canopy. It is concluded that pendulous lichens, such as A. sarmentosa, are strongly influenced by landscape mosaic structure. Results have important implications for conservation of epiphytic lichens in managed forest ecosystems.


Secondary chemistry, stress experiments and DNA-analyses of the Australian lichen Heterodea muelleri (Hampe) Nyl. and culture of the mycobiont

Hager, A. & Stocker-Wörgötter, E.
Institute of Plant Physiology, University of Salzburg, Salzburg, Austria

The study aimes to figure out suitable culture conditions and nutrient requirements for the biosynthesis of secondary metabolites in the cultured mycobionts of Heterodea muelleri. The production of biologically active substances like usnic acid and diffractaic acid in the cultured mycobionts is of high interest. The voucher specimens of Heterodea muelleri contain diffractaic acid as the major substance, barbatic acid as a minor compound and 4-O-demethyldiffractaic acid in traces.
The cultured Heterodea-mycobionts produce secondary metabolites only under stress-conditions, e.g. shortwave UV-light and low temperature treatment. Both stress parameters initiate the production of the same substances as in the voucher specimens, but in different concentrations. Moreover, the stressed mycobionts produce 4-O-demethylbarbatic acid in minor quantity. All secondary compounds were analysed by HPLC, TLC and an additional microcrystallisation test. DNA-analyses confirmed the identity of the aposymbiotically grown mycobionts.


Successional stages of the lichen component in an dry acidic grassland (Spergulo-Corynephoretum)

Hasse, T. & Daniëls, F. J. A.
Institute of Plant Ecology, Hindenburgplatz 55, 48143 Münster, Germany

This poster presents the first results of a detailed permanent plot study on the dynamics of the lichen component in Corynephorus canescens vegetation in the lowland of the Netherlands and North Germany. A total of 186 permanent plots were established in ten vegetation types with different composition of cryptogamic plants. Frequency and cover of species were recorded and analysed by means of tabular comparison, correspondence analysis, and spectra of different species attributes. Most of the vegetation units are linked by progressive succession accompanied by humus accumulation, whereas the others differ in macroclimate, mineral substrate, or by invasion of Campylopus introflexus. The succession is characterised by a cumulative arrival of new species. Due to frequent small-scale disturbances most pioneer species survive in gaps throughout the later successional stages. As a result species diversity is highest in the final successional stage. Thallus fragmentation seems to be the most important dispersal strategy in terricolous lichen vegetation. Repeated measurements of the plots as well as experimental treatments (mechanical disturbance, nitrogen eutrophication, sand and litter deposition) in some of these plots are supposed to contribute to a more detailed understanding of the dynamics in lichen vegetation on sandy soil.


The environmental factors determing the lichen species richness on islets in the West-Estonian Archipelago

Jüriado, I., Suija, A. & Liira, J.

Institute of Botany and Ecology, University of Tartu 38 Lai str. Tartu 51005

Estonia is a country rich in islands, over 1500 islands and islets are situated on its western and northern coasts. The islets are higher parts of earlier sea ground risen above sea level as a result of the constant and rather fast neotectonic uplift. The investigated islets are relatively young, the rise of them started about 2000 years ago. The islets formation and disappearance, joining together or with mainland continues also nowadays. The lichen flora of the Estonian islands and islets has been investigated quite intensively, but the environmental factors, which could affect the lichen species richness, are statistically evaluated for the first time. This study deals with the lichens of the Hiiumaa Islets Landscape Reserve which is part of the West-Estonian Archipelago. We investigated 25 islets during the summer period in 2001 (Suija & Jüriado 2002). The islets vary by size (0.1 to 140 ha), age, distance from mainland (the farthermost islet is ca 18 km from mainland) and vegetation. We studied all biotopes with different kind of substrate types, suitable for growth of lichens, thoroughly and compiled the species lists for every islets. The total number of species registered on the islets is 297 (lichenicolous and allied fungi are included as well). For every lichen species we calculated the relative abundance on a four-point scale. A general linear model (GLM) analysis (Statistica 6.0) was applied to study the dependence of the species richness on heterogenity of biotopes on islets, distance of the islets from mainland, area of the islets and human influence. The heterogenity of biotopes is related to the size of the studied islets. The total species richness on islets is influenced by the heterogenity of biotopes and also by the distance from the point of mainland. The area of the islets is additionally important factor for the lichen species richness on different substrate types.


Effect of wildfire on lichens in grass-encroached dunes at Terschelling (The Netherlands)

Ketner-Oostra, R. (1), Van der Peijl, M. J. (2) & Sykora, K. V. (3)
(1) Freelance ecologist, Algemeer 42, 6721 GD Bennekom, The Netherlands; (2) ESM-EcoSystem Modelling, Roghorst 132, 6708 KR Wageningen, The Netherlands; (3) Department of Environmental Sciences, Nature Conservation and Plant Ecology Group, Wageningen University, Bornsesteeg 69, 6708 PD Wageningen, The Netherlands

The effect of fire on soil lichens and their re-establishment in grassland vegetation could give basic information for a possible future management regime to regain the former lichen diversity in calcium-poor dunes. In 1993 an opportunity for new information presented itself after a wildfire devastated the vegetation on a coastal dune. The rather steep 32 m long southern and the 100 m long gently sloping western slope had a monitoring history going back to 1966. Inside a contiguous grid of blocks of 4 x 4 m, quantities of living and dead higher plants, mosses, lichens, bare sand and ashes (after the fire) were estimated in 1966, 1990 and every year from 1993 until 2001. Vegetation relevés from a pre-fire survey in 1990 served as permanent plots (PQs) for presence and quantity of all plant species, including the cryptogams. The data of the blocks and PQs were analysed with multivariate analysis.
The results made clear that fire alone does not restore the open lichen-rich grassland from 1966. Re-growth of graminoids and rosette plants was favoured by a surplus of minerals from the ashes. After two years, an expanding moss cover grew rampantly over the bare and ash-covered sand. Lichen species arrived after 3 years, and increased in number after 5 years. After 10 years the lichen diversity of 11, including several pioneer species, was high compared to 5 species in the grass-encroached stage of 1990, but still lower than the 17 species in the open grassland of 1966. However, like in 1990, the vegetation was dominated by higher plants and had a complete moss cover without bare sand and a total lichen cover of only 1%.
The almost flat part of the western slope was an exception. Sand with a relatively high base content was blown-in from nearby foredunes there. Lichens of sub-neutral sand were promoted to grow on and between the damaged mosses, to a mean cover of 12%.
If fire should be used for management in calcium-poor dunes, additional measures are recommended, such as promoting in-blowing sand.


Distribution of lichen species and biomass in pine forests in Poland

Lesnianski, G. Z.
Departmant of Biosystematics, University of Opole, Oleska 48, PL-45-052 Opole, Poland

The goal of the study, the results of which are to some extent shown on the poster, is to describe the current state of the lichen biota in pine forests in Poland. Experimental sites (in total 150) are being established in the following communities: Cladonio-Pinetum, Leucobryo-Pinetum, Peucedano-Pinetum, Molinio-Pinetumi Vaccinio uliginosi-Pinetum. Some research questions are: What are the species of lichen biota in the above five communities of pine forests in Poland? What is the allocation of lichen biomass in those pine forests? In which one of those communities does biomass reaches the highest values? What is the pine forest development stage when lichen biomass reaches its maximum amount? What are the common features and what are the differences concerning development of lichen biotas in pine forest communities under examination? Is there any, and if so, what is the correspondence between development stage of the given pine forest community and the age structure of lichen populations that occur in them?
Locations of experimental sites have been chosen at random, after dividing the range of the given community in Poland into blocks (each with 100 km side). The chosen blocks vary considerably from each other concerning the share of the given pine forest association in its total vegetation, as well as the state of the natural environment, including, mainly, the level of air pollution. Some experimental sites were established in patches of forests formed as substitute communities.
At this time the following facts are established: 1. Cladonio-Pinetum community has the biggest registered lichen biomass (200 g/m2); 2.The most balanced biomass distribution in three dimensional space of pine forest patches was registered in Leucobryo-Pinetum and Peucedano-Pinetum communities. In Cladonio-Pinetum lichens are most common in the ground-layer while in Vaccinio uliginosi-Pinetum almost 100% of lichen biomass consists of epiphytic lichens. 3. Thirty lichen species have been found so far in the area covered by the research. 4. The explored areas in southern and central Poland, in comparison with northern Poland, are characterised by significantly smaller total lichen biomass. Fifty experimental sites have been established to date, out of one hundred and fifty sites planned for the whole research.


Composition and substrata of forest lichens in Estonia: a meta-analysis

Lõhmus, P.
Institute of Botany & Ecology, University of Tartu, Lai St. 38, 51005 Tartu, Estonia

Species richness, composition and substratum use of "forest lichens" in Estonia were studied by re-analysing published and unpublished reports (incl. 13 quantitative studies). A total of 481 species of lichenised, lichenicolous and systematically allied fungi were confirmed; their probable number reached 599 species (195 macrolichens, and 404 microlichens and lichenicolous fungi). The eight most frequent species were present in all but 1–2 studies, whereas 47% of species occurred in only 1–2 studies. Calicioid, cetrarioid, cladonioid, parmelioid, pendulous, sorediate crustose and lichenicolous and parasitic species totalled nearly half (46 %) the lichen species in Estonian forests. The highest number of unique species has been recorded on living trees, but different substrata have not been equally researched. Ground and coarse woody debris (e.g., snags with exposed wood) were found to be inhabited by a relatively high number of species. According to species composition, there were five distinct substratum groups for the Estonian "forest lichens" – broad-leaved trees, common deciduous and coniferous trees, branches and undergrowth, windthrows, and other ground-level substrata. In future research, 1) compilation of a list of obligate "forest lichens" could be valuable for practical purposes in conservation management, 2) a more detailed analysis of the causes of rarity among "forest lichens" should be possible after additional research of inadequately studied forest types and substrata.


Altitudinal distribution of lichens and lichenindication of altitudinal belts

Magomedova, M.A.
Institute of Plant and Animal Ecology UD RAS, Ekaterinburg, Russia

Altitude above sea level plays a great role in distribution of lichens in highlands. Differences in composition and structure of lichen communities had been studied along altitudinal ecological profiles on the eastern slope of the Polar, Prepolar and Northern Urals on the base of traditional geobotanical methods adapted to special features of lichen communities. All lichens, even widespread, have altitudinal coenotic optimums. Coenotic optimums of boreal species are concentrated in a lower part of the alpine tundra altitudinal belt, and of arctic-alpine species in central and upper parts of this belt. The upper limit of wide distribution of boreal lichen species can be used as an indicator of position of the climatic boundary between elfin-wood and alpine tundra altitudinal belts. Decreases in species diversity and number of coenotic optimums were discovered in the transitional zone between alpine tundra and cold desert altitudinal belts. Changes in correlation of arctic-alpine and other species, abundance of species and their coenotic role, composition of dominant species, total cover and predominant communities were used as indicators of this transitional zone. Positions of this boundary at different latitudes in the Urals are different: 1450 (1430-1485) m at Northern, 850 (800-1000) m at Prepolar, 500 (400-700) m at Polar Urals. The Polar Urals boundary is less distinct in connection with strong determination of lichen distribution by habitats. Changes of composition and structure of lichen communities as well as distribution and coenotic role of species along altitudinal (and zone) gradients demonstrate sensitivity of lichens to temperature and humification. Altitudinal differences in lichen species diversity, individual distribution of species, composition and structure of communities, and distribution of communities can be used as bases for indication of altitudinal belts in highlands where the role of higher plants is limited.


Estimating epiphytic macrolichen biomass from topography, stand structure, and lichen community data in Oregon, USA

McCune, B. (1) & Berryman, S. D. (2)
(1) Department of Botany and Plant Pathology, Cordley 2082, Oregon State University, Corvallis, OR, 97331-2902, USA; (2) Department of Forest Science, 321 Richardson Hall, Oregon State University, Corvallis, OR, 97331, USA

We modeled lichen epiphyte biomass in relation to topography, stand structure, and lichen community composition in the central Cascades of western Oregon. Sampling was stratified by stand structure. Lichen biomass was estimated for three functional groups: cyanolichens, forage lichens, and matrix lichens. We used lichen litter to estimate epiphytic lichen biomass in forest stands. Lichen biomass changed along an elevation gradient, with highest cyanolichen biomass at low elevations (470 – 950 m) and highest forage lichen biomass at higher elevations (950 – 1470 m). Lichen biomass was related to stand structure; lowest lichen biomass in even-aged young stands and highest in old-growth and mature stands with remnant trees (trees that survived the most recent logging or fire). Lichen biomass was higher in stands with more remnant trees as compared to similar even-aged stands. We modeled epiphytic macrolichen biomass (oven-dried; 10x kg/ha) using stepwise linear regression and nonlinear regression. We followed a three-step approach to develop the predictive models, each step demanding an additional class of variables. First, we created models from topographic predictors that are easily extracted from GIS data. We then developed models based on both topographic and stand structure variables. Finally, we developed models based on topography, stand structure, and lichen community data. Models with the greatest explanatory power were: cyanolichen biomass predicted as a function of elevation, stand age index, the sum of abundance for Lobaria oregana and L. pulmonaria, and cyanolichen species richness (R2 = 0.85); forage lichen biomass predicted as a function of stand age index and the abundance of Alectoria sarmentosa (R2 = 0.55); and matrix lichen biomass predicted as a function of stand age index and the sum of abundances for matrix lichen species in the stand (R2 = 0.58).


Influence of environmental factors on spatial distribution of several epiphytic lichen species in the North Urals

Mikhailova, I. (1), Trubina, M. (1), Vorobeichik, E. (1) & Scheidegger, C. (2)
(1) Institute of Plant and Animal Ecology UD RAS, Ekaterinburg, Russia; (2) WSL Swiss Federal Research Institute, CH-8903 Birmensdorf, Switzerland

Spatial distribution and abundance of ten lichen species on Populus tremula trunks were studied in the mountain taiga of the North Urals. Within an area of 150x200 m, each aspen trunk was mapped, and a range of environmental variables (including characteristics of tree stand, shrub and grass layers; 30 variables in total) were registered within a circle of 6 m diameter around each tree. Factor analysis was applied to reduce the total variation to three independent factors. Extracted factors can be interpreted as follows: Factor 1 - the portion of tall grasses in the grass layer; Factor 2 - soil fertility; Factor 3 - insolation of the site. The factors accounted for about 45% of the total variation. Abundance of seven lichen species was found to correlate significantly with at least one of the factors though correlations were very weak (Lobaria pulmonaria, Leptogium saturninum, Nephroma parile and N.resupinatum negatively correlated with Factor 1, Pannaria pezizoides and Vulpicida pinastri positively correlated with Factor 2, and L. saturninum, V. pinastri and Phlyctis argena negatively correlated with Factor 3). Step-wise regression revealed a low probability to predict presence or absence of the species using the collected variables. Even for "the most predictable" N. resupinatum the factors explained only 15% of the variance. Analysis of spatial patterns of extracted factors revealed the moderate degree of their structuring. However, lichen species showed almost random spatial distribution over the study area. Only L.pulmonaria showed signs of spatial aggregation, and the size of clusters is smaller than that of the environmental factors. It can be a sign of lower dispersal ability of this species compared to others.
This study suggests that the environmental variables are of small importance for spatial distribution of the lichens on a local scale. However, the abundance of the species was shown to depend, at least in a certain extent, on size and vitality of their host trees.


Influence of environmental variables and forest management on the lichen flora of eutrophic paludified forests in Estonia

Nõmm, M., Jüriado, I. & Kohv, K.
Institute of Botany & Ecology, University of Tartu, Lai St. 38, 51005 Tartu, Estonia

Estonia is situated in hemiboreal vegetation zone and forests cover 51,5 % of the land. Intensive forest management is defined as severe threat for the forest lichens in Estonia; however so far, the forest lichen flora has not been studied along the management intensity gradient. Altogether, 12 forest stands of different management intensities from eutrophic paludifying forests type group were studied in NE, E and SW Estonia. The epiphytic lichen flora was described following the method developed by Scheidegger et al. (2002). In addition, the lichen species from other available substrates were registered and the lists of lichen species at stand-level were compiled. Forest stands were characterised by the proportion of different tree species, number of snags, logs and stumps, canopy cover, age of forest trees etc., and the index of management intensity was calculated. The index was based on traces of concrete management activities e.g. cutting stumps, ditches, roads etc. registered at the 30 m and 60 m radius from the stand center. Different ordination methods were used to describe the overall structures of epiphytic and stand-level lichen communities and to evaluate the relationships between lichen species composition and environmental parameters (Canoco 4.0). A total of 132 lichen taxa were recorded, 61% of them were crustose lichens. Two species, Fellhaneropsis vezdae and Arthopyrenia fraxini, were found new to Estonia. In the studied eutrophic paludifying forest stands the present level of management intensity seems to have smaller effect on the composition of lichen flora than the other observed environmental variables. However, some of the environmental variables important for the lichen flora could be indirect result of different management activities, that our management index did not detect, e.g. tree species composition and stand age. For future, the development of more lichens specific management index and analyses on bigger datasets are needed.


Corticolous lichen flora on Pinus patula from the pinus-oak forests in Sierra de Juárez, Oaxaca, Mexico

Pérez Pérez, R. E. (1), Herrera-Campos, M. A. (2), Quiroz Castelán, H. (3) Garcia Barrios, R. (4)
(1) Facultad de Ciencias Biológicas, UAEM, Av. Universidad 1001, Col. Chamilpa, 62210 Cuernavaca, Morelos, Mexico; (2) Instituto de Biología, UNAM, Departamento de Botánica, Apdo. Postal 70-233, Coyoacán 04510, México, D. F.; (3) Centro de Investigaciones Biológicas, UAEMor, Av. Universidad 1001, Col. Chamilpa, 62210 Cuernavaca, Morelos, Mexico; (4) Centro Regional de Investigaciones Multidisciplinarias, UNAM,, Apdo. Postal 4-106, C.P. 62431, Cuernavaca, Morelos, Mexico

Corticolous lichen communities have different functions in the forest ecosystems. They increase structural complexity, influence nutrient cycles, and provide habitat, food, and nesting material for some animals. Forest management has direct effects on these organisms (Dettki & Esseen, 2003); therefore, it is necessary to implement silvicultural methods that allow for their abundance and diversity. In the Sierra de Juárez of Oaxaca, well organized local indigenous communities maintain diverse strategies of sustainable forest management that allow them to take economic advantage of the forests' products and environmental services while maintaining their biodiversity.
We present a comparative study of the corticolous macrolichen communities on Pinus patula Schl. et Cham. between the timber-producing forest of Ixtlán de Juárez and the non-timber-producing forest of Santa María Yavesía. Forest structure was significately different for the two localities particularly in DBH and height of the tree species mentioned. However, no statistically significant differences were detected in the macrolichen population richness, which allows us to suggest that the management techniques used in this region, such as the rotation period of 50 years, do not produce irreversible alterations to the processes of dispersion, establishment and growth of the corticolous macrolichens on this phorophyte. A total of 52 species were collected, 49 from the managed forest and 41 from the non-managed forest. Hypotrachyna was the most frequent genus with 12 species and Heterodermia the most uncommon, found only in one tree.


Functional ecology of the biological soil crust in semiarid SE Spain: sun and shade ecotypes of Diploschistes diacapsis

Pintado, A., Sancho, L. G. & Blanquer, J. M.
Departamento de Biología Vegetal II, Facultad de Farmacia, Universidad Complutense, Madrid, Spain
The Tabernas badlands in semiarid southeastern Spain is one of the driest regions in Europe. The mean annual precipitation is below 200 mm, most of which falls as intense storm rainfall specially in September and October. The landscape shows deep dissection, with canyons, ramblas and sparsely vegetated eroded badlands slopes. The biological soil crust that covers this area consists of different types of lichen-dominated communities. One of the most conspicuous is dominated by Diploschistes diacapsis (Ach.) Lumbsch. Its distribution is mainly restricted to the north-facing slopes, where it forms extensive whitish carpets. This thick biological soil crust appears to play a decisive role in preventing erosion of the slopes and allowing plant colonization. South-facing slopes are much more eroded and lack of vegetation.
The aim of this work was to establish the photosynthetic performance of north and south-facing populations of D. diacapsis, in order to determine if these different populations showed any adaptation or limitation of the photosynthetic response to the prevailing microclimatic conditions on slopes with different exposure. For this purpose, and as a first step, we determined the CO2 exchange response to light intensity, temperature and water content under controlled conditions in the laboratory. Results showed that the optimum temperature for net photosynthesis in the population with northern exposure ranged between 15 and 25 ºC, while the population with southern exposure showed a narrow peak around 25 ºC. Dry weigh-based net photosynthetic rates were higher in the southern-exposed population. The physiological differences observed between both populations are discussed in terms of their distribution patterns.


Why do lichens like dead trees and fir more than living trees and spruce? Results of a study on Whiteface Mt., NY, USA

Schmull, M. & Hauck, M.
Albrecht von Haller Institute for Plant Sciences, Dept. of Systematic Botany, University of Goettingen, 37073 Goettingen, Germany

Epiphytic lichen distribution on the trunks of living and dead Picea rubens and Abies balsamea and its dependency on chemical and microclimatic site factors were studied in a spruce-fir forest on Whiteface Mountain, New York, USA.
Epiphytic lichen diversity was considerably higher on dead compared to living trees and on fir compared to spruce. Diversity differed more between living and dead spruce than between living and dead fir.
Cover of most lichen species decreased with increasing mean concentration of NO3- in stem flow. Concentrations of NO3- were higher on living spruce compared to dead spruce and to living and dead fir. Additionally, NO3- was found experimentally to decrease chlorophyll contents a and b of Hypogymnia physodes compared to the controls, whereas ergosterol concentrations remained unaffected. This result indicates that a dependence of lichen cover on NO3- concentrations in stemflow may be the cause for negative correlations. Mn in the stemflow, known to damage soredia of Hypogymnia physodes, negatively correlated with cover of Mycoblastus sanguinarius and Parmelia saxatilis. However, the present work shows that concentrations measured in the stem flow were considerably lower than in experimental studies.
Mn concentration of A. balsamea bark was considerably higher than in the stemflow. Mn is known to limit epiphytic lichen abundance on Picea abies in the Harz Mountains, Germany, where total Mn concentrations in the bark are much lower than on Whiteface Mountain. Total Mn concentrations in the bark of P. rubens on Whiteface Mountain were in the same range as in the bark of P. abies in Germany. X-ray microanalysis showed that A. balsamea immobilized most Mn in crystals in spongy cork and in sclerotic phelloid cells of the outer bark and explains the high lichen abundance on A. balsamea despite the high total Mn concentrations.
Microclimatic parameters did not differ between fir and spruce, nor between living and dead trees.


The lichen flora of Aegean Islands (Greece) compared

Sipman, H. J. M.
Botanic Garden & Botanical Museum Berlin-Dahlem, Free University of Berlin, Berlin, Germany

In an ongoing project to inventory the lichens of selected islands in the Aegean Sea (Greece) in a standardized way, 4 islands and island groups have been studied: Santorini, Paros/Antiparos, Kalimnos/Kos and Ikaria. They differ in size, altitudinal range, substrate availability and vegetation continuity. The encountered species numbers per island range between 182 and 332.The lichen flora of the region appeared poorly known and several species known so far only from the western Mediterranean were found. A few species seem to have an Eastern Mediterranean distribution. They are common on the investigated islands, and not known west of the Adriatic coast of Italy. Some species or chemotypes show discontinuous distributions, being present on one island but absent from suitable habitats on a nearby island, or even on a nearby mountain ridge.


Ammonia as a key factor for the composition of epiphytic lichen communities

Sparrius, L. B.
BIO.DIV consultancy, Gouda, The Netherlands

Lichens are useful study objects for nutrient-limited communities due to their quick response to changes in the environment. This paper presents ammonia-dependent optimum curves for epiphytic lichens and show that some ammonia pollution increases the species number and abundance of epiphytes. Decreasing the local ammonia air concentration in fairly polluted areas may lead to a decline of especially macrolichens.
Three ecological groups can be distinguished:
– acidophytes are lichens growing on acid bark. This group is negatively correlated with nutrient availability and bark pH, e.g. Evernia prunastri and Pseudevernia furfuracea.
– nitrophytes are lichens that are positively correlated with nutrient availability and pH, e.g. Xanthoria parietina and Candelaria concolor.
– neutrophytes are lichens which are generally considered as indifferent to low and moderate levels of acidification, e.g. macrolichens in the genera Parmelia s.l. and Ramalina. Their ecological optimum is in between the acidophytes and the nitrophytes.
A case study at 241 monitoring sites with oak trees revealed that neutrophytes decreased in areas where the ammonia levels became lower. The largest decline could be observed in areas where the ammonia levels were already low and decreasing. The largest increase of neutrophytes could be observed in areas where the ammonia levels are very low, but increasing. This suggests that acidophilous epiphytic communities are most vulnerable to hypertrophication.


The results of transplantation of Lobaria pulmonaria in southern part of Sakhalin Island

Taran, A. A.
Sakhalin Botanical Garden, Russia

In southern part of Sakhalin island in 2001 a population of Lobaria pulmonaria in the Madera River Valley (Makarovsk District) was described. The centre of population is situated in riparian forest with different species of Salix. Lobaria pulmonaria had been investigated near this place in mixed and coniferous forest too. There are five species of trees in this site (Salix sachalinensis, S. schwerinii, Picea ajanensis, Abies sachalinensis, Alnus hirsuta). We described 102 trees with Lobaria pulmonaria. More often the thallus of Lobaria was found on various species of Salix (59 trees) and Picea (41 trees); these dominate or co-dominate on this site. As rule, the thalli are well developed, with marginal and laminal soredia and 23,5 % with apothecia. More often fertile thallus were found on northern and northeastern exposures of trunks. For this experimental site we used Picea ajanensis as acceptor trees. These were growing on a northeast slope. We transplanted to these trees 225 fragments of Lobaria pulmonaria from 5 clones in 4 sites. These transplants were collected on a transect from Europe to Asia (Ukraine Carpathian Mountains, near Syktyvkar City, north Asian Ural Mountains, Sakhalin). We used the method developed by J-C. Wasler and C. Scheidegger (2002) for transplanting lichen fragments for provenance-clone tests. During 3 years, 18,2 % fragments died or bleached, 20,9 % were partially necrotic, and 60,9 % were healthy (vivid green). More often dead and necrotic fragments were found among transplants from the Carpathians (37 fragments).


Comparison of lichen community response to environmental variables at regional and subregional geographic scales

Will-Wolf, S. (1), Geiser, L. H. (2) & Neitlich, P. (3)
(1) University of Wisconsin-Madison, Madison, WI, USA; (2) USDA Forest Service, Corvallis, OR, USA; (3) National Park Service, Winthrop, WA, USA

We compared correlation of environmental variables with axes of Nonmetric Multidimensional Scaling ordinations based on composition of epiphytic macrolichen communities of forests at two different spatial scales. Data for two USA regions, 1) Midatlantic states and 2) Washington and Oregon west of the Cascades divide, from the USA Forest Service Forest Inventory and Analysis program were used for the larger scale analyses. Data from the Allegheny National Forest, embedded in 1), and the Willamette National Forest, embedded in 2), were used for the smaller scale analyses. Major axes of community variation at the larger scales were correlated with regional air quality, climate variables, and large scale topographic variables such as elevation. At the smaller scales, forest vegetation variables such as stand age and tree species composition, and local environmental variables such as topographic position were important correlates of variation in lichen species composition. Elevation in the western data sets was the only variable significantly correlated with major axes at both scales. Few lichen species were equally good indicators of gradients in community composition and response to environmental variables at both scales. Good indicators of gradient response at larger scales were often insensitive to gradients important at smaller scales.


Epiphyte microhabitat and substrate preferences on Sequoia sempervirens (redwood) in northwestern California, USA

Williams, C. & Sillett, S. C.
Department of Biological Sciences, Humboldt State University, Arcata, California, USA

Epiphytic lichens and bryophytes on Earth's tallest trees have not been explored until now. Specimens of Sequoia sempervirens that combine massive size and structural complexity with towering height provide an incredibly diverse array of microhabitats for epiphytes. To what extent do epiphytes display microhabitat and substrate specificity? Nine study trees were each divided into seven strata, and each stratum was then further separated into microhabitat-substrate combinations (plots). These plots were extensively scoured for species richness to reveal microhabitat and substrate affinities. We encountered 193 lichen taxa, including 82 macrolichens and 111 microlichens, 53 bryophyte taxa, including 31 mosses and 22 liverworts, as well as 50 vascular plant taxa in 978 plots. Upper crown strata were richest, carrying 74% of epiphyte taxa and over 98% of epiphytic macrolichen species. Richness diminished toward trunk flare strata, where only 9% of epiphytic lichen taxa occurred. Redwood bark and xylem in upper crown strata supported many lichen species, while epiphytic tree, shrub, and fern leaves as well as positions closer to the ground harbored fewer epiphytes. Interestingly, dead redwood foliage supported 44% of overall epiphyte species richness compared to live foliage, which supported only 5%. Cyanolichens on redwood substrates accounted for merely 0.4% of all epiphyte encounters, while neighboring, non-redwood trees (e.g., Picea sitchensis) were often festooned with cyanolichens. Litterfall directly beneath these tree crowns is a poor representation of redwood's rich epiphyte communities. Despite a relatively low abundance of epiphytic lichens and bryophytes, redwoods support far more species richness than other conifers that have been surveyed, primarily because of their high microlichen diversity.

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