Epiphytic lichen is associated with species richness of gall-inducing aphids but not with niche differentiation among them

The presence of epiphytic foliose lichen amplifies the heterogeneity of habitat by creating shelters for insects living on tree bark. It thus should enhance species number and spatial niche segregation among these canopy insects. We studied this hypothesis in a field experiment using four aphid species that induce galls on Pistacia atlantica trees covered with Xanthoria parietina lichen. In autumn 2008, 3 months after aphid fundatrices were oviposited, we marked six branches on each of 29 trees. Two served as a control, whereas the other four were isolated with insect glue; two of them were scraped with sandpaper to remove epiphytic foliose lichens. We therefore obtained three treatments comprising control branches, isolated branches with lichen, and isolated branches without lichen. In summer 2009, we counted all the galls developing on five new annual shoots on each of 174 branches. We observed more cecidogenic aphid species on all the branches with lichens than without, but each species at different proportions. The different frequencies of utilization of the lichen did not lead to habitat partitioning between species. In conclusion, although habitat heterogeneity itself was associated with species richness and population abundance, it did not induce spatial niche segregation. Considering that many economically important insect species, pests and natural enemies, oviposit or spend some portion of their lives in bark cracks, it is possible that some can use lichens too for protection or/and oviposition sites. As a consequence, lichens may affect management of agrosystems and their impacts should be investigated more deeply in such contexts.     

Tree and stand-scale factors affecting richness and composition of epiphytic bryophytes and lichens in deciduous woodland key habitats

Conservation and sustainable forestry are essential in a multi-functional landscape. In this respect, ecological studies on epiphytes are needed to determine abiotic and biotic factors associated with high diversity. The aim of the present study was to evaluate relative sensitivity of conservation targets (epiphytic bryophytes and lichens) in relation to contrasting environmental variables (tree species, tree diameter at breast height, bark crevice depth, pH, tree inclination, pH, forest stand age, area and type) in boreo-nemoral forests. The study was conducted in Latvian 34 woodland key habitat (WKH) boreo-nemoral forest stands. Generalized linear mixed models and canonical correspondence analysis showed that tree species and tree bark pH were the most important variables explaining epiphytic bryophyte and lichen composition and richness (total, Red-listed, WKH indicator species). Forest stand level factors, such as stand size and habitat type, had only minor influence on epiphytic species composition and richness. The results of the present study indicate a need to maintain the diversity of tree species and large trees, particularly Acer platanoides, Carpinus betulus, Fraxinus excelsior, Populus tremula, Tilia cordata, Ulmus glabra and Ulmus laevis in conservation of epiphytic bryophyte and lichen communities in the future.     

Removal of secondary compounds increases invertebrate abundance in lichens

We investigated how lichen carbon-based secondary compounds (CBSCs) affect abundance of invertebrates in five lichen species growing on the forest floor (Cladonia rangiferina, Cladonia stellaris) or on tree trunks (Evernia prunastri, Hypogymnia physodes, Pseudevernia furfuracea). To do this, CBSCs were removed by rinsing lichen thalli in acetone (which has no adverse effects on the lichens) and the lichens were re-transplanted in their natural habitat. After 4 months there was higher abundance of mites, springtails and spiders in the three epiphytic lichens that had their CBSC concentrations reduced. The increase in predatory spiders following CBSC reduction suggests that the compounds have multitrophic consequences. The acetone treatment reduced the number of nematodes in four of the lichen species. Given that lichens serve as important habitats for a diverse range of invertebrates, increased knowledge of how lichen CBSCs may regulate their abundance helps us to better understand the role that lichens and their defence compounds play in structuring forest food webs.     

Cladonia lichens as the most effective and essential pioneers in strongly contaminated slag dumps

The epigeic lichen biota of ten post-smelting slag dumps were studied to identify the ecological importance of Cladonia species in spontaneous succession within artificial areas. Species diversity, abundance, constancy and cover indices were determined at the initial stages of vegetation establishment. Substrates under lichen vegetation were characterised in terms of the pH, organic C and total N content, C/N ratio and Zn, Pb, Cd and As concentration. Species occurrence and abundance indices were related to habitat type, enabling the ability to identify species-specific responses to environmental factors. Despite the unfavourable and artificial habitat conditions at dump sites, some Cladonia appear abundant and form swards, with Cladonia rei as a dominant species. They appear to be well-adapted for slag substrate and their special arrangement illustrative of site contamination. Certain sward-forming Cladonia lichens play a fundamental role as effective pioneers in the initial stage of succession and provide stable plant cover development.     

First evidence for seasonal fluctuations in lichen- and bark-colonising fungal communities

Endophytic fungal communities in leaves of deciduous trees usually undergo pronounced seasonal changes. We hypothesised that such compositional shifts are predominantly caused by annuality of the leaves and therefore less pronounced in fungi colonising the perennial substrates bark and corticolous lichens. To test this hypothesis, thalli of the foliose lichen-forming fungal species Xanthoria parietina and Physconia distorta, along with the adjacent bark, were sampled during spring and autumn at two sides of a single tree in southern Germany. Analysis of clone libraries by restriction fragment length polymorphism (RFLP) revealed 588 singleton and 221 non-singleton RFLP-types of non-lichenised fungi. The communities differed significantly between host lichen species. Season and exposure had only a significant impact when the two factors were combined in the analysis. Accordingly, bark- and/or the lichen-associated fungal communities change throughout the year’s course, a finding that rejects the initial hypothesis. This survey revealed valuable information for future broad-based studies, by indicating that a relatively high diversity of non-lichenised fungi is associated with corticolous lichen thalli and the adjacent bark. Furthermore, the structure of non-lichenised fungal assemblages associated with corticolous lichen communities obviously depends at least on the following factors: ‘lichen species’, ‘exposure’, and ‘season’.     

乌鲁木齐达坂城山区地衣生态位特征的研究

为揭示达坂城山区地衣对生境资源的利用状况及竞争程度,该研究在野外设置30个样方并调查其物种盖度数据,利用生态位宽度、生态位重叠指数及排序分析,探讨达坂城山区地衣的生态位特征及环境影响因子。结果表明:(1)达坂城山区生态位最宽的为丽黄鳞衣(Rusavskia elegans)和内卷野粮衣(Circinaria contorta),对不同环境具有较强的适应性,其他地衣物种生态位宽度较窄,对环境资源利用程度低。(2)物种间生态位重叠值整体偏低,生态位重叠值较高的物种对极少,地衣物种生态位分化程度高,物种间竞争不激烈。(3)生态位较宽的物种与多数物种之间都存在生态位重叠,但重叠值较低; 一些生态位较窄的物种与其他物种之间的生态位重叠值较高; 生态位重叠和生态位宽度之间并无直接的线性关系。(4)地衣物种沿海拔梯度分布具有差异,海拔、光照强度、湿度、风速和人为干扰是本地区地衣群落物种生态位特征差异的主导环境因子。综上所述,达坂城山区地衣物种由于生境资源的竞争占据各自独特的生态位,形成了生态位的分化,群落较为稳定,沿海拔梯度的生境条件差异是物种差异分布的重要原因。该研究可为地衣群落构建研究提供理论依据,并对该地区地衣物种的多样性和生境保护具有重要意义。     

Tree species and microhabitat influence the population structure of the epiphytic lichen Lobaria pulmonaria

Detailed knowledge of the habitat requirements of species is required because habitat greatly affects the persistence of species. We investigated the effects of tree species and microhabitat heterogeneity on the population of the locally threatened lichen Lobaria pulmonaria. We studied four L. pulmonaria populations in Central Spain and collected microhabitat data for individuals growing on beech and oak. The microhabitat affected the life stages of L. pulmonaria; being a phorophyte species the location of the lichen was the most important factor generating different patterns of establishment, abundance, thallus size and reproductive capacity. Although oak forests favoured the establishment and recruitment of new L. pulmonaria individuals, they apparently provided adverse environmental conditions for lichen growth, thus affecting the reproductive capacity since this is size-dependent. By contrast, beech forests offered a more favourable microclimate, because L. pulmonaria individuals reached larger sizes in these forests. In conclusion, our results indicate that habitats hosting large populations, with high rates of establishment and recruitment do not necessarily favour other life-cycle stages.     

DISPERSAL PATTERNS OF TERRICOLOUS LICHENS BY THALLUS FRAGMENTS

Dispersal patterns from seven terricolous lichen species, with a high capacity for asexual reproduction by fragmentation (Cetraria muricata, Cladonia species), were studied in differing vegetation types in north-eastern Germany. Marked lichen thalli were fragmented by trampling. After 15 days the spread of thallus fragments was monitored. Whereas most of the fragments that were dispersed by wind remained within a 20-cm radius from the source, the maximal dispersal distance was 57 cm in a dry sand grassland and 68 cm in an open pioneer pine forest. Dispersal was negligible in a closed old-growth pine forest. Several fragmented lichen cushions were disturbed and removed by animals, and led to a maximal dispersal distance of 9·70 m. These results suggest that: (a) thallus fragments provide good short-distance dispersal in open vegetation, but are inefficient for long-distance dispersal, and (b) wind and animals are important factors for the dispersal of thallus fragments. For restoration management of man-made substrata, artificial introduction of lichen thalli is proposed.     

ENVIRONMENTAL FACTORS AFFECTING THE DISTRIBUTION OF LICHENS ON GRANITIC MONUMENTS IN THE IBERIAN PENINSULA

The phytosociology of lichens colonizing granitic churches and dolmens in Galicia (N.W. Spain) and Alentejo (central Portugal) was studied so as to identify (a) a group of species whose frequency on these structures would make them potentially useful as test species for the evaluation of biocides used for conservation, (b) environmental factors likely to control species distribution, and (c) species that may be useful as bioindicators of environmental factors in this kind of construction in the region studied. Chief among the factors correlating with species distribution is the class of structure (church or dolmen), which is attributed to the probable influence of this factor on environmental parameters affected by structural complexity, the presence of non-granitic materials and the influence of man in the immediate surroundings. The bioindicator species most sensitive to this complex of parameters appear to be three species found only on churches (Caloplaca citrina , Sarcogyne clavus and Verrucaria macrostoma) and the most widespread of the species found only on dolmens (Lasallia pustulata). Other important factors, probably affecting the influence of class of structure, are pH and the availability of nitrogen and moisture.     

When habitat management can be a bad thing: effects of habitat quality,isolation and climate on a declining grassland butterfly

The conservation of most temperate grassland habitats and their characteristic fauna and flora requires regular low-intensive forms of land-use to counteract natural succession. Although many species tolerate moderate disturbance regimes, some are known to be susceptible to grazing or mowing, thereby causing a management dilemma. One of these species is the Woodland Ringlet butterfly, Erebia medusa. In this study, we analysed which environmental factors determine the occurrence of E. medusa in the Diemel Valley (Central Germany). Furthermore, we conducted microclimatic measurements during the winter months to investigate the role of the litter layer as a microclimatic buffer. Patch occupancy in the Diemel Valley was well explained by the amount of litter present in a patch and connectivity to other inhabited patches. The role of local climatic conditions could not be clarified, due to inter-correlations with connectivity. During the winter, the air temperature inside the litter layer was significantly less variable than above it. We conclude that the current distribution of E. medusa in the Diemel Valley is caused by the combined effect of habitat quality and connectivity, and perhaps also by climatic factors. The importance of the litter layer reflects the dependence of E. medusa on low-intensive or absent land-use. In addition, the litter layer possibly constitutes an essential habitat element, as it buffers temperature fluctuations and thus probably reduces the energy consumption of overwintering larvae. Given the species’ preference for abandoned grasslands, the conservation of E. medusa requires a low-intensity habitat management, for example, by rotational grazing or mowing of small parts of the sites. On the landscape level, the preservation of well-connected habitat networks is important.     

Is local distribution of the epiphytic lichen <Emphasis Type="Italic">Lobaria pulmonaria</Emphasis> limited by dispersal capacity or habitat quality?

By surveying and re-surveying 12 forest sites in southern Sweden for the epiphytic lichen Lobaria pulmonaria with a 9-year interval, and measuring tree-related habitat quality variables, we have investigated whether the local distribution of the lichen is limited by poor dispersal capacity or by habitat quality. Dispersal distances were measured indirectly as the distances between colonised trees and the nearest trees occupied by L. pulmonaria in both 1992 and 2001. To compare habitat quality between trees occupied by L. pulmonaria and neighbouring control trees, we recorded tree species and measured age and growth rate of trees, light conditions, bark structure and bryophyte cover. The estimated mean dispersal distance was 35 m, with a recorded maximum of 75 m. Occupied trees were larger and had a larger cover of bryophytes than unoccupied trees of similar size. The results indicate that dispersal capacity probably is the most important factor in limiting the local distribution of L. pulmonaria, but habitat-quality factors may be important on a smaller spatial scale.     

Differential herbivore damage to calcicolous lichens by snails

Abstract: A laboratory experiment was conducted to determine the damage caused by snail grazing to 35 calcicolous lichen species and cyanobacteria from the Baltic island of Öland, Sweden. Individuals of four species of land snails (Balea perversa, Chondrina clienta, Clausilia bidentata and Helicigona lapicida) were allowed to graze on identified lichen species growing on pieces of limestone. Snail grazing on thalli and ascocarps was classified into four categories ranging from no damage to completely eaten. In general, some lichen species were heavily grazed, whereas others were only slightly damaged or not injured. Aspicilia calcarea, Tephromela atra and Verrucaria nigrescens were preferred by all snail species except Chondrina clienta. In general, lichen thalli that were not immersed in the calcareous rock and cyanobacteria were preferred, whereas ascocarps were avoided by one of the snail species (C. clienta). Immersed perithecia with a carbonized outer layer were avoided by all snail species except C. bidentata. Possibilities of chemical and mechanical defence properties in calcicolous lichens are discussed.     

Extremotolerance and Resistance of Lichens: Comparative Studies on Five Species Used in Astrobiological Research II. Secondary Lichen Compounds

Lichens, which are symbioses of a fungus and one or two photoautotrophs, frequently tolerate extreme environmental conditions. This makes them valuable model systems in astrobiological research to fathom the limits and limitations of eukaryotic symbioses. Various studies demonstrated the high resistance of selected extremotolerant lichens towards extreme, non-terrestrial abiotic factors including space exposure, hypervelocity impact simulations as well as space and Martian parameter simulations. This study focusses on the diverse set of secondary lichen compounds (SLCs) that act as photo- and UVR-protective substances. Five lichen species used in present-day astrobiological research were compared: Buellia frigida, Circinaria gyrosa, Rhizocarpon geographicum, Xanthoria elegans, and Pleopsidium chlorophanum. Detailed investigation of secondary substances including photosynthetic pigments was performed for whole lichen thalli but also for axenically cultivated mycobionts and photobionts by methods of UV/VIS-spectrophotometry and two types of high performance liquid chromatography (HPLC). Additionally, a set of chemical tests is presented to confirm the formation of melanic compounds in lichen and mycobiont samples. All investigated lichens reveal various sets of SLCs, except C. gyrosa where only melanin was putatively identified. Such studies will help to assess the contribution of SLCs on lichen extremotolerance, to understand the adaptation of lichens to prevalent abiotic stressors of the respective habitat, and to form a basis for interpreting recent and future astrobiological experiments. As most of the identified SLCs demonstrated a high capacity in absorbing UVR, they may also explain the high resistance of lichens towards non-terrestrial UVR.     

Environmental context shapes the bacterial community structure associated to Peltigera cyanolichens growing in Tierra del Fuego,Chile

The structure of the associated bacterial community of bipartite cyanolichens of the genus Peltigera from three different environmental contexts in the Karukinka Natural Park, Tierra del Fuego, Chile, was assessed. The sampling sites represent different habitat contexts: mature native forest, young native forest and grassland. Recently it has been determined that the bacterial community associated to lichens could be highly structured according to the mycobiont or photobiont identities, to the environmental context and/or to the geographic scale. However, there are some inconsistencies in defining which of these factors would be the most significant on determining the structure of the microbial communities associated with lichens, mainly because most studies compare the bacterial communities between different lichen species and/or with different photobiont types (algae vs. cyanobacteria). In this work bipartite lichens belonging to the same genus (Peltigera) symbiotically associated with cyanobacteria (Nostoc) were analyzed by TRFLP to determine the structure of the bacterial community intimately associated with the lichen thalli and the one present in the substrate where they grow. The results indicate that the bacterial community intimately associated differs from the one of the substrate, being the former more influenced by the environmental context where the lichen grows.     

Further evidence that activation of net photosynthesis by dry cyanobacterial lichens requires liquid water

Abstract: In contrast to green algal lichens, cyanobacterial species of different families, growth forms and habitats proved to be unable to attain positive net CO₂ assimilation when the dry thalli were treated with air of high relative humidity; they needed liquid water for the reactivation of their photosynthetic apparatus. Identical behaviour is shown by all of the 47 lichen species with cyanobacterial photobionts, from six different genera, studied so far. This suggests a widely distributed, if not general, characteristic of cyanobacterial lichens. The difference in performance between both groups of photobionts was maintained when the lichen thallus was macerated. Furthermore, cultures of Chroococcidiopsis were unable to make use of water vapour hydration for positive net photosynthesis, and were similar in this respect to some free-living aerophilic cyanohacteria tested earlier. Possible physiological implications as well as ecological consequences for water-relation-dependent habitat selection of green-algal and cyanobacterial lichens are discussed.     

中国蜈蚣衣科地衣多样性及区系研究

该研究通过对山西管涔山和五鹿山等区域采集的蜈蚣衣科地衣300余份标本的鉴定分析,结合已有报道,从物种多样性、空间分布特点、生境类型及区系成分等方面对蜈蚣衣科地衣进行研究,以明确中国蜈蚣衣科地衣物种组成及区系特点。结果表明：(1)中国蜈蚣衣科地衣共有7属197种54变种,饼干衣属(Rinodina)占绝对优势,约占蜈蚣衣科地衣总种数的30.5%。(2)中国蜈蚣衣科地衣多分布于新疆,蜈蚣衣属(Physcia)是该科中分布最广泛的属,但其中有26.4%的物种仅分布于单一地区。(3)蜈蚣衣科地衣生境类型多样,有树生、石生、藓丛生、土生和多生境5种,以多生境地衣为主(48.5%)。(4)中国蜈蚣衣科地衣的地理成分复杂,同时伴随多种区系成分,但以温带性质显著(占43.1%),东亚特色明显。(5)仅分布于中国的蜈蚣衣科地衣共4属10种,且主要分布于中国东北、西北和西南地区。     

Natural peat bog remnants promote distinct spider assemblages and habitat specific traits

Peat bogs are very sensitive and highly endangered ecosystems. They were once typical landscape elements in northern Germany, but today only a few remnants exist. On-going habitat degradation has alarming adverse effects on biodiversity, and, from a conservation viewpoint, it is imperative to evaluate the current ecological quality of the remaining peat bog remnants to assess the intensity of degradation and to suggest reasonable management strategies. In 2007, spiders were sampled in 23 study sites representing typical peat bog habitat types in the northern parts of Westphalia (NW Germany). In all, 214 species from 18,413 adult individuals have been collected. Multivariate analyses showed that different peat bog successional stages harbour distinct spider assemblages with succession being the main driver for species separation. Pardosa sphagnicola, Pirata piscatorius and P. uliginosus can be considered as flagship species for near-natural or natural peat bogs. Trait analyses showed that habitat specialisation for high moisture is negatively affected by succession. Peat bogs are now only small remnants, but, nevertheless, they have a high conservation value as they still harbour a distinct species assemblage and specialised species.     

PROTOPARMELIA HYPOTREMELLA,A NEW STERILE CORTICOLOUS SPECIES FROM EUROPE,AND ITS LICHENICOLOUS FUNGI

Protoparmelia hypotremellavan Herk, Spier & Wirth, a sterile corticolous lichen, is described as a species new to science, so far known from Austria, SW Germany and the Netherlands. It is rapidly spreading in Germany and the Netherlands. It is apparently close toP. oleagina, and is often found growing together with that species on oak trees along roads in the Netherlands. Both species are host to the lichenicolous fungiSphinctrina anglicaandTremella wirthii.     

How Many Wolves (Canis lupus) Fit into Germany? The Role of Assumptions in Predictive Rule-Based Habitat Models for Habitat Generalists

Due to legislative protection, many species, including large carnivores, are currently recolonizing Europe. To address the impending human-wildlife conflicts in advance, predictive habitat models can be used to determine potentially suitable habitat and areas likely to be recolonized. As field data are often limited, quantitative rule based models or the extrapolation of results from other studies are often the techniques of choice. Using the wolf (Canis lupus) in Germany as a model for habitat generalists, we developed a habitat model based on the location and extent of twelve existing wolf home ranges in Eastern Germany, current knowledge on wolf biology, different habitat modeling techniques and various input data to analyze ten different input parameter sets and address the following questions: (1) How do a priori assumptions and different input data or habitat modeling techniques affect the abundance and distribution of potentially suitable wolf habitat and the number of wolf packs in Germany? (2) In a synthesis across input parameter sets, what areas are predicted to be most suitable? (3) Are existing wolf pack home ranges in Eastern Germany consistent with current knowledge on wolf biology and habitat relationships? Our results indicate that depending on which assumptions on habitat relationships are applied in the model and which modeling techniques are chosen, the amount of potentially suitable habitat estimated varies greatly. Depending on a priori assumptions, Germany could accommodate between 154 and 1769 wolf packs. The locations of the existing wolf pack home ranges in Eastern Germany indicate that wolves are able to adapt to areas densely populated by humans, but are limited to areas with low road densities. Our analysis suggests that predictive habitat maps in general, should be interpreted with caution and illustrates the risk for habitat modelers to concentrate on only one selection of habitat factors or modeling technique.     

Seasonal variation in the diet of the serotine bat (Eptesicus serotinus): A high-resolution analysis using DNA metabarcoding

Bats play an important role as predators of insect populations but are threatened by a variety of factors, including the loss of foraging habitat and insect declines. Knowledge on trophic interactions, foraging strategies, and hunting areas is key to understanding the ecology of bat species, to assess their impact on ecosystems and to optimize conservation strategies. We investigated seasonal trends in the diet of two nursery colonies of the serotine bat, Eptesicus serotinus, from an intensively farmed agricultural landscape in Germany. Using DNA-metabarcoding of food remains in bat droppings collected from May to July 2018, we identified 254 taxa of 13 arthropod orders to species or genus level, including numerous pest species. Our results indicate an equal use of Coleoptera, Diptera, and Lepidoptera, contradicting previous morphological dietary analyses that had shown beetles to be the most frequent prey. The dietary composition was seasonally highly variable and mainly determined by prey phenology. Dietary richness significantly increased throughout the sampling period, reflecting increasing insect activity with progressing season. Our findings demonstrate that E. serotinus is a generalist forager, linking different habitat types through trophic interactions.