Functional ecology of rare and common epigeic lichens in alvar grasslands

The autecology of rare species can be derived using similarities among functional traits and environmental conditions observed for common species, i.e. we employed the ‘matching analogy approach’ with the analytical scheme ‘common species → driver → trait → driver → rare species’. We addressed the driver–trait relationship for common epigeic lichens of thin-soil calcareous grasslands, which are endangered by cessation of traditional land use. Common lichens were suppressed by encroaching herbs and shrubs, and were supported by ground disturbances. The lichens of open low-productivity alvars are predominantly calciphilous, epibryic, crustose-squamulose, sexually reproducing and contain UV-protective pigments. Lichens of encroached alvars are soil-type generalists, fruticose, reproduce vegetatively and contain herbivore-deterring compounds. Rare lichens resemble the species of open low-productivity alvars, except their more limited niche space, i.e. they are restricted to arctic-alpine habitats. The conservation practices on alvars should support the formation of crust-forming communities by suppressing the growth of shrubs and herbs, and by promoting recurrent small-scale soil disturbances.     

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.     

Environmental factors and ground disturbance affecting the composition of species and functional traits of ground layer lichens on grey dunes and dune heaths of Estonia

A unique, species‐rich and endangered lichen biota can be found on European coastal and inland sand dunes. However, it is increasingly affected by natural succession as well as by anthropogenic disturbances. We studied lichen diversity on the grey dunes and dune heaths of coastal and inland regions of Estonia. A total of 28 study plots were investigated; in each 0.1 ha study plot general environmental variables and anthropogenic disturbances were described and all epigeic lichen species were identified. We found 66 lichenized fungus (lichen) species, including several rare and ten red‐listed lichens. Multivariate analysis (DCA, CCA) was performed to examine gradients in species composition and to relate variation in species data to environmental factors. In addition, we used redundancy analysis (RDA) to relate variation in species’ trait composition to environmental factors. Species composition on grey dunes differed significantly from that on dune heaths. The characteristic species for grey dunes are, besides several Cladonia species, foliose lichens, e.g. Hypogymnia physodes, Parmelia sulcata and Peltigera spp. Also species’ traits composition was different for either habitat, indicating that sorediate lichens, foliose lichens, lichens with cyanobacterium as the main photobiont, and sparsely branched Cladonia species dominate on grey dunes, while esorediate, green‐algal, crustose and richly branched fruticose lichens are common on dune heaths. Soil pH was the most essential environmental variable for determining both species composition and species’ traits composition. The composition of lichen species was also significantly influenced by forest closeness, soil Mg content and cover of bare sand; the effect of ground disturbances was low compared to the effect of these environmental factors. To protect and conserve the species‐rich lichen biota, it is necessary to protect the dune habitats from building activity, to avoid overtrampling in recreation areas and to regularly remove shrubs and trees.     

Vegetation,environmental characteristics and ideas on the maintenance of alvars on the Bruce Peninsula,Canada

Abstract. Despite international recognition that alvar habitats are important reservoirs of biodiversity, they remain little studied in North America. In this paper, the results are reported on an investigation of alvars in the central portion of their known distribution on this continent. 210 plots were distributed among seven sites and were quantitatively sampled for vascular plants, lichens, bryophytes and a suite of environmental variables. Detrended and Canonical Correspondence Analyses and other methods were used to investigate differences among alvars, within alvars and between alvars and adjacent forested habitats. The plant communities and environmental conditions were highly similar among sites in the study region, yet very different from surrounding habitats. There were abrupt changes in vegetation and environmental conditions from alvar to forest, without the presence of transition zones in the vegetation or environmental gradients as the forest was approached. The environmental factors associated with the change from alvar to forest and with variation within alvar habitat were examined. Some alvars in the study were found to contain stunted, slow-growing trees reaching ages of 524 yr. These same sites appear to have remained unburned for several centuries, while other sites likely burned 90 yr ago. The plant communities were very similar between the alvars that lacked a major, biomass-removing disturbance in centuries and alvars that had experienced catastrophic fire relatively recently. Maintenance of the plant communities and open nature of alvars appears site-specific rather than habitat specific.     

Epigeic lichen communities of taiga and tundra regions

The major physiognomic and ecological categories of the lichen-rich, epigeic communities in the boreal (taiga) and arctic (tundra) zones are defined and their syntaxonomy and ecology in Europe, Asia and North America is reviewed. In the boreal and hemiarctic areas open, dry, acidophytic lichen woodlands are widespread, especially on sandy habitats. Their epigeic lichen synusiae are usually dominated by four fruticoseCladina species, being extremely homogeneous in species composition and structure throughout the boreal zone, while the dominant trees and the other vascular plant flora of the woodlands are geographically more variable. No phytosociological classification system exists that would cover most of these communities over the circumpolar regions. Very similar communities, though much more poorly known, are found on thin soils over Precambrian rock outcrops. Other sites to produce epigeic lichen communities include open sand dunes, treeless heathlands, drier bogs and many seral stages, like those on road banks. Boreal lichen-rich communities on eutrophic soils may be developed in semiarid regions, in particular. In the Arctic, lichens are common in most communities, and the driest ones are regularly lichen-dominated, whether acidophytic or eutrophytic, chionophytic or achionophytic. Detailed syntaxonomic systems for their classification have been developed, especially in Greenland and Scandinavian mountains (in oroarctic zones in the latter regions). The richest fruticose lichen areas are in continental, hemiarctic timberline regions in northern Siberia and Canada. The southern and middle arctic subzones are also characterized by many macrolichens, such asCetraria cucullata, C. nivalis, Alectoria ochroleuca, andThamnolia vermicularis, but everywhere also small, crustose lichens are common on soil, such asRinodina turfacea andLopadium pezizoideum, which are often overlooked in vegetation analyses. The presence of microlichens and the formation of mosaic micropatterns of soil lichen communities is particularly typical of the northern arctic subzone. The conservation problems of the boreal and arctic lichen communities include overgrazing by reindeer or caribou, which has caused delichenization in some regions, extensive forest and tundra fires, use of heavy transport vehicles in forestry and tundra operations, and, locally, heavy industrial air pollution.     

Life History Traits Reflect Changes in Mediterranean Butterfly Communities Due to Forest Encroachment

The biodiversity of the Southern Balkans, part of the Mediterranean global biodiversity hot-spot, is threatened by land use intensification and abandonment, the latter causing forest encroachment of formerly open habitats. We investigated the impact of forest encroachment on butterfly species richness, community species composition and the representation of life history traits by repeated seasonal visits of 150 one-hectare sites in five separate regions in three countries—Greece, Bulgaria, and the Republic of Macedonia (FYROM—the Former Yugoslav Republic of Macedonia)— 10 replicates for each habitat type of grasslands, open formations and scrub forest within each region. Grasslands and open formations sites hosted in average more species and more red-listed species than scrub forest, while no pattern was found for numbers of Mediterranean species. As shown by ordination analyses, each of the three habitat types hosted distinct butterfly communities, with Mediterranean species inclining either towards grasslands or open formations. Analysing the representation of life history traits revealed that successional development from grasslands and open formations towards scrub forest shifts the community composition towards species overwintering in earlier stages, having fewer generations per year, and inhabiting large European or Eurosiberian (e.g. northern) ranges; it decreases the representation of Mediterranean endemics. The loss of grasslands and semi-open formations due to forest encroachment thus threatens exactly the species that should be the focus of conservation attention in the Mediterranean region, and innovative conservation actions to prevent ongoing forest encroachment are badly needed.     

Slow response of plant species richness to habitat loss and fragmentation

We examined the response of vascular plant species richness to long-term habitat loss and fragmentation of Estonian calcareous grasslands (alvars). The current number of habitat specialist species in 35 alvars was not explained by their current areas and connectivities but it was explained by their areas and connectivities 70 years ago ( R ² = 0.27). We estimated the magnitude of extinction debt in local communities by assuming an equilibrium species richness in 14 alvars that had lost only a small amount of area and by applying this model to the remaining alvars, in which the average area has declined from 3.64 km² in the 1930s to 0.21 km² at present. The extinction debt estimated for individual alvars was around 40% of their current species number. Our conclusions are applicable to temperate grasslands in general, which have lost much area because of agricultural intensification and cessation of traditional management.     

Carabid beetles as indicators for shrub encroachment in dry grasslands

Semi-natural grasslands are key habitats for biodiversity conservation in Central Europe. Shrub encroachment is one of the most threatening drivers of grassland degradation and affects soil properties, microclimate, and vegetation with possible impacts on higher trophic levels. We aimed to analyse the impact of shrub encroachment with broom (Cytisus scoparius) on carabid beetle diversity, species composition, and functional traits. In a field study on dry grasslands on the island of Hiddensee (Germany) we studied 15 sites along a gradient of increasing broom encroachment and classified them into three dry grassland types with low, medium, and high shrub cover. Our results provide evidence that shrub encroachment initially has positive effects on species richness and activity densities of dry grassland carabids. Carabid species composition differed among differently shrub-covered dry grassland types, and sites with low and high shrub cover were each characterised by unique carabid assemblages. The species composition of sites with a medium shrub biomass had a transitional character and contained species which are typical for open dry grassland, but also shared species with sites with a high shrub cover. Among functional trait parameters investigated, especially the body size of carabid beetles was related to environmental parameters associated with shrub encroachment. Body size was positively correlated to shrub biomass and soil humidity, but negatively to temperature. Eurytopy values of carabids were related to high litter cover, i.e. habitat generalist (eurytopic) species mainly occurred in densely shrub-encroached sites. In order to preserve unique carabid assemblages of open dry grasslands with stenotopic and smaller species, it is most important to prevent a shrub encroachment higher than about 60% cover. For management we suggest extensive grazing (by cattle, sheep or horses) to prevent shrub encroachment on dry grasslands. In areas with high shrub cover additionally the use of goats or mechanical removal of shrubs might be necessary.     

Soil fungal composition changes with shrub encroachment in the northern Chihuahuan Desert

Woody species encroachment of grasslands globally causes many socioecological impacts, including loss of grazing pastures and decreased biodiversity. Soil microbial communities may partially regulate the pace of shrub encroachment, as plant-microbial interactions can strongly influence plant success. We measured fungal composition and activity under dominant plant species across a grassland to shrubland transition to determine if shrubs cultivate soil microbial communities as they invade. Specifically, soil microbial communities, abiotic soil properties, and extracellular enzyme activities were quantified for soils under four common Chihuahuan Desert plant species (three grasses, one shrub) in central New Mexico, U.S.A. Extracellular enzyme activity levels were fairly consistent under different plant species across the grassland to shrubland transition. Activity levels of two enzymes (alkaline phosphatase and beta-N-acetyl-glucosaminidase) were lower in the ecotone, presumably because soil organic matter content was also lower in ecotone soils. Community composition of soil fungi mirrored patterns in the plant community, with distinct plant and fungal communities in the shrubland and grassland, while grassland-shrubland ecotone soils hosted a mix of taxa from both habitats. We show that shrubs cultivate a distinct microbial community on the leading edge of the invasion, which may be necessary for shrub colonization, establishment, and persistence.     

The impact of forest encroachment after agricultural land abandonment on passerine bird communities: The case of Greece

Agricultural land abandonment is one of the main drivers of land use change, leading to various responses of farmland ecological communities. In an effort to better understand the effect of agricultural land abandonment on passerine bird communities, we sampled 20 randomly selected sites [1 km × 1 km] in remote Greek mountains, reflecting an abandonment gradient, in terms of forest encroachment. We sampled 169 plots using the point count method of fixed distance (47 passerine species), and we investigated bird diversity and community structure turnover along the gradient. We found that grazing intensity has a beneficial effect hampering forest encroachment that follows progressively land abandonment. Habitat composition changes gradually with forests developing at the expense of open meadows and heterogeneous grasslands. Forest encroachment has a significant negative effect on bird diversity and species richness, affecting in particular typical farmland and Mediterranean shrubland species. Birds form five distinct ecological clusters after land abandonment: species mostly found in pinewoods and cavity-dwelling species; species that prefer open forests forest edges or ecotones; species that prefer shrubland or open habitats with scattered woody vegetation; Mediterranean farmland birds that prefer semi-open habitats with hedges and/or woodlots; and, generalist forest-dwelling or shrubland species. We extracted a set of 22 species to represent the above ecological communities, as a new monitoring tool for agricultural land use change and conservation. We suggest that the maintenance of rural mosaics should be included in the priorities of agricultural policy for farmland bird diversity conservation.     

Lichenicolous fungi are more specialized than their lichen hosts in primeval forest ecosystems,Białowieża Forest,northeast Poland

Taxonomy and diversity of symptomatic lichenicolous fungi (visible as fruitbodies on lichen thalli, their discoloration, and/or deformation) and their specificity to lichen hosts is becoming more and more studied. However, information on their ecology is still scarce. We assess how large the specialization of these fungi towards their hosts and microhabitat is. Epiphytic, epixylic and epigeic lichens and associated lichenicolous fungi were studied on 144 permanent plots in Białowieża Forest in relation to forest communities, species of tree phorophyte and substrates. On all these three studied levels lichenicolous fungi were more specialized than their lichen hosts. Our study provides the first estimation of ecological dependences between associations of lichenicolous fungi and their hosts, microhabitats and forest communities in a primeval forest ecosystem representative of lowland Europe.     

Spontaneous restoration of epiphytic lichen biota in managed forests planted on habitats typical for temperate deciduous forest

The study examines the changes of epiphytic lichen diversity in differently aged stands developing after clear cutting and pine planting on fertile habitats typical for deciduous forests. The study was conducted within one large complex consisting of pine, mixed pine-hornbeam and typical old oak-linden-hornbeam forests in northern Poland. Epiphytic lichens were recorded in 50 study plots randomly selected within 5 forest stand classes of a different structure and age, ranging from 80 to over 220 years. Altogether 143 lichen species were recorded, of which only 41 were entirely nonspecific, and were occurring in all the studied forest stand classes. Significant differences in lichen species richness between stand classes were found and the number of species increases with the forest age. Lichen species composition also differs and its changes progress towards restoration of lichen biota typical for deciduous forest consistent with the habitat. The age of the forest has the most significant effect on the biodiversity of lichen biota. Microhabitat space provided by oaks is highly desirable since it greatly enriches lichen biota in forests. Phorophyte specificity of particular lichens were assessed. Hornbeam and oak have the greatest number of species mostly confined to them and constitute a main refuge for lichens with a high conservation value. The changes of lichen biota are basically parallel with the changes of the forest stand structure. The selection of some parts within managed pine forests that should not be assigned for cutting in the future can be a simple procedure which helps to restore and preserve forest biodiversity.     

Threatened Alvar Grasslands in NW Russia and their Relationship to Alvars in Estonia

Alvar grasslands are thin-soil calcareous grasslands on Silurian and Ordovician limestone bedrock. They are considered unique plant communities for northern Europe and America with a high conservation value. Until now, European alvar grasslands have only been described in Sweden and Estonia. We described species-rich alvar grasslands on the Izhora plateau, NW Russia and compared this vegetation with that of Estonian alvar grasslands. Floristic composition and environmental conditions of Russian alvars were similar to Estonian alvar grasslands on thicker dry soil. Russian alvars are isolated from similar communities in Estonia, but extensive transport of domestic animals from Estonia to NW Russia in past centuries has probably enhanced species dispersal. Some characteristic Estonian alvar species were missing in NW Russia because it lies outside their natural ranges. Other differences in vegetation were explained by higher atmospheric nitrogen deposition and younger age of Russian sites. In NW Russia the alvar grasslands were small, but several red-listed plant species were present. In order to preserve European alvar vegetation effectively, Russian alvar grasslands need further study and conservation as an outpost of this threatened vegetation type.     

亚热带森林附生地衣压力-体积曲线分析及其适用性

附生地衣是森林生态系统中的重要组成部分, 在维护森林物种多样性以及水分和养分循环等方面发挥着重要作用。地衣类群的水势特征以及压力-体积(PV)曲线及相关参数是否适用于探索地衣等变水性生物类群应对水分胁迫机制, 仍有待阐明。该研究以中国西南地区云南哀牢山亚热带山地森林中5个类群15种常见附生地衣为对象, 探讨了其PV曲线及水势参数的功能群和物种水平的变化。结果显示, 蓝藻型地衣的内部含水量(WC_internal)和共质体水含量(Rs)显著高于绿藻型地衣, 是其1倍以上, 但其他参数表现出较高的一致性。功能群水平上, 地衣仅WC_internal、Rs和膨压损失点的相对含水量(RWC_TLP)具有显著性差异; 但物种间差异均非常显著。结合主成分分析进一步发现, PV曲线及相关参数在评估地衣整体应对水分胁迫以及揭示生境水分条件选择策略时具有较大的限制性, 受到共生藻和生长型的显著影响; 但内部最大持水力可以解释蓝藻型地衣的生境适应策略, 而部分参数如饱和渗透势(Ψsat)和RWC_TLP可以分别用于解释狭叶地衣、阔叶地衣和枝状地衣对生境水分条件的适应。研究表明PV曲线及水势参数不适用于地衣类群整体的抗旱性评价, 和其他生物类群进行抗旱性比较时更要慎用。     

How lichens impact on terrestrial community and ecosystem properties

Lichens occur in most terrestrial ecosystems; they are often present as minor contributors, but in some forests, drylands and tundras they can make up most of the ground layer biomass. As such, lichens dominate approximately 8% of the Earth's land surface. Despite their potential importance in driving ecosystem biogeochemistry, the influence of lichens on community processes and ecosystem functioning have attracted relatively little attention. Here, we review the role of lichens in terrestrial ecosystems and draw attention to the important, but often overlooked role of lichens as determinants of ecological processes. We start by assessing characteristics that vary among lichens and that may be important in determining their ecological role; these include their growth form, the types of photobionts that they contain, their key functional traits, their water‐holding capacity, their colour, and the levels of secondary compounds in their thalli. We then assess how these differences among lichens influence their impacts on ecosystem and community processes. As such, we consider the consequences of these differences for determining the impacts of lichens on ecosystem nutrient inputs and fluxes, on the loss of mass and nutrients during lichen thallus decomposition, and on the role of lichenivorous invertebrates in moderating decomposition. We then consider how differences among lichens impact on their interactions with consumer organisms that utilize lichen thalli, and that range in size from microfauna (for which the primary role of lichens is habitat provision) to large mammals (for which lichens are primarily a food source). We then address how differences among lichens impact on plants, through for example increasing nutrient inputs and availability during primary succession, and serving as a filter for plant seedling establishment. Finally we identify areas in need of further work for better understanding the role of lichens in terrestrial ecosystems. These include understanding how the high intraspecific trait variation that characterizes many lichens impacts on community assembly processes and ecosystem functioning, how multiple species mixtures of lichens affect the key community‐ and ecosystem‐level processes that they drive, the extent to which lichens in early succession influence vascular plant succession and ecosystem development in the longer term, and how global change drivers may impact on ecosystem functioning through altering the functional composition of lichen communities.     

Heterogeneity fosters biodiversity: Linking history and ecology of dry calcareous grasslands

Calcareous grasslands in Central Europe harbour a high diversity of plant and animal species. However, as man-made habitats, they need to be managed in order to maintain high species diversity. Conservation efforts often aim at reintroducing historical management regimes, such as regular grazing or mowing. Despite such efforts, the diversity and number of species of calcareous grasslands is still decreasing. We propose that, besides fragmentation and eutrophication, a lack of structural heterogeneity within and around calcareous grasslands as created by historic management is causing species loss as well. Here, we review the literature on the history of calcareous grassland management in northern Switzerland, the heterogeneity that it created and the relevance of this heterogeneity for biodiversity at three spatial scales: (1) within grasslands, (2) in their close surroundings and (3) at the landscape scale. Considering that historic management has created heterogeneity at all three scales and that many species do indeed depend on this structural diversity, we conclude that in order to conserve the full range of biodiversity associated with calcareous grasslands, conservation management should aim at increasing heterogeneity in, around, and between grasslands.     

Shrub encroachment alters composition and diversity of ant communities in abandoned grasslands of western Carpathians

Invasions of woody species into grasslands abandoned by agriculture are a global phenomenon, but their effects on diversity of other taxa have been rarely investigated across both regional and local scales. We quantified how shrub encroachment affected the activity, composition, and diversity of ant communities in managed and abandoned grasslands in western Carpathians of Central Europe across four regions and four shrub encroachment stages in each region. We surveyed ant communities on 48 sites in total, with each encroachment stage replicated three times in each region and twelve times overall. We used pitfall traps to sample ants over three years (2008, 2009, 2011) and identified 9,254 ant workers belonging to 33 species in total. Although the epigaeic activity and composition of ant communities varied with region, abandoned grasslands supported a greater species richness of ants than managed grasslands regardless of the region, and especially so in more advanced shrub encroachment stages. Since the woody colonization within grasslands was moderate even in the advanced encroachment stages (on average ~40 % of grassland colonized by woody species), it allowed coexistence of forest specialists (e.g. Temnothorax crassispinus) with species typical of open grasslands, thus increasing overall ant diversity. Managed grasslands were not only less species rich compared to abandoned grasslands, but they were characterized by different species (e.g. Lasius niger, Myrmica rugulosa). The differences in ant communities between managed and abandoned grasslands are likely to cause differences in ecological functions mediated by ants (e.g. predation of arthropods or plant seed dispersal).     

Bacterial diversity across individual lichens

Symbioses are unique habitats for bacteria. We surveyed the spatial diversity of bacterial communities across multiple individuals of closely related lichens using terminal restriction fragment length polymorphism (T-RFLP) and pyrosequencing. Centers of lichens house richer, more consistent assemblages than species-poor and compositionally disparate lichen edges, suggesting that ecological succession plays a role in structuring these communities.     

The taxocene of lichen-feeding darkling beetles (Coleoptera,Tenebrionidae: Helopini) in a forest-steppe ecotone

The taxocene of lichen-feeding darkling beetles from the tribe Helopini is studied in the steppes of the south of Rostov Province. The species of lichens grazed on by the representatives of this tribe are identified for the first time and dependence of circadian activity and spatial distribution of adults on temperature and air humidity is shown. Global warming and human activities may have caused degradation of steppe lichen communities that formed the dietary base for the lichen-feeding darkling beetles in natural ecosystems. As a result, 3 out of 4 beetle species completely left steppe biotopes for tree shelterbelts. This caused a change in the trophic relationships (transition from epigeic to epiphytic lichens) and in the spatial distribution of darkling beetles, promoted the establishment of competitive interactions, and increased the duration of circadian and seasonal activity due to the lengthening of the periods with optimum temperature and air humidity for each species.