Endolithic Research Articles

Endolithic invertebrate communities and bioerosion rates in southwestern Atlantic intertidal consolidated sediments

Fil: Bagur Creta, Maria. Consejo Nacional de Investigaciones Cientificas y Tecnicas. Oficina de Coordinacion Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales ; Argentina

The effects of lichen cover upon the rate of solutional weathering of limestone

The contribution of lichens to the biomodification of limestone surfaces is an area of conflict within bioweathering studies, with some researchers suggesting a protective effect induced by lichen coverage and others a deteriorative effect induced by the same organisms. Data are reported demonstrating the potential role of endolithic lichen, in particular of Bagliettoa baldensis, in the active protection of Carboniferous limestone surfaces from rainfall-induced solutional weathering. During a 12-month microcatchment exposure period in the west of Northern Ireland, average dissolutional losses of calcium are greater from a lichen-free limestone surface compared with a predominantly endolithic lichen-covered surface by just under 1.25 times. During colder winter months, the lichen-free surface experiences calcium loss almost 1.5 times greater than the lichen-covered surface. Using extrapolation to upscale from the micro-catchment sample scale, for the year of sample exposure, the rate of calcium loss is 1.001gm−2a−1 from lichen-covered limestone surfaces and 1.228gm−2a−1 from lichen-free bare limestone surfaces. This research has implications for our understanding of karst environments, the contribution of lichens to karren development and the conservation of lichen-colonised dimension stone within a cultural setting.

Spatial ecology goes to space: Metabiospheres

Lithopanspermia is increasingly accepted as a possible mechanism for the exchange of organisms between planets. Meteoritic impacts on inhabited planets can generate ejecta reaching escape velocity, which could carry endolithic ecosystems from the deep biosphere into space. If the ejecta travel long and fast enough they could be captured by another planet. Organisms inside the rock might survive re-entry, reaching the planet lithosphere. These processes re-enact spatial ecology at an astronomical level, for example mirroring metapopulation ecology: empty (of life) and occupied patches (planets and large moons) are separated by an inadequate matrix (space), being colonised (turned into biospheres) and perhaps suffering extinctions. As in metapopulations, we can predict this colonisation/extinction dynamics will lead to higher probabilities of life persisting longer times in a Solar System given adequate conditions. This system may be called a metabiosphere, and this ecological dynamics at an astronomical level astroecology, a new scientific branch mingling ecology and astrobiology.

The perks of being endolithic

The perks of being endolithic

Lichen-induced biomodification of calcareous surfaces: Bioprotection versus biodeterioration

Studies demonstrate the active and passive capability of lichens to inhibit or retard the weathering of calcareous surfaces. Lichen coverage may actively protect a surface through shielding by the thallus and the binding and waterproofing of the rock surface and subsurface by fungal hyphae. Passive protection of rock surfaces may be induced by the formation of an insoluble encrustation, such as calcium oxalate, at the lichen-rock interface. Recent research suggests that the decay of hyphae, induced by changes in microenvironmental conditions, necrosis, parasitism or the natural physiological traits of particular lichen species, may expose a chemically and physically weakened substrate to dissolution, triggering relatively rapid weathering-related surface lowering. Consequently, certain epilithic crustose and endolithic lichens may induce a period of surface stability throughout the course of their lifespan, followed by a phase of instability and rapid episodic microtopographical evolution after death and decay. A series of conceptual models is proposed to illustrate this idea over short (single lichen lifespan) and long (multiple lichen lifespans) timescales. The models suggest that the microscale biogeomorphological system of lichen-rock interaction is underpinned by non-linear dynamical system theory as it exhibits dynamical instability and is consequently difficult to predict over a long timescale. Dominance by biodeterioration or bioprotection may be altered by changes in lichen species or in environmental conditions over time.

Late Pleistocene eolian activity linked to permafrost degradation in the Upper Mississippi River basin, USA

The contribution of lichens to the biomodification of limestone surfaces is an area of conflict within bioweathering studies, with some researchers suggesting a protective effect induced by lichen coverage and others a deteriorative effect induced by the same organisms. Data are reported demonstrating the potential role of endolithic lichen, in particular of Bagliettoa baldensis, in the active protection of Carboniferous limestone surfaces from rainfall-induced solutional weathering. During a 12-month microcatchment exposure period in the west of Northern Ireland, average dissolutional losses of calcium are greater from a lichen-free limestone surface compared with a predominantly endolithic lichen-covered surface by just under 1.25 times. During colder winter months, the lichen-free surface experiences calcium loss almost 1.5 times greater than the lichen-covered surface. Using extrapolation to upscale from the micro-catchment sample scale, for the year of sample exposure, the rate of calcium loss is 1.001 g m− 2 a− 1 from lichen-covered limestone surfaces and 1.228 g m− 2 a− 1 from lichen-free bare limestone surfaces. This research has implications for our understanding of karst environments, the contribution of lichens to karren development and the conservation of lichen-colonised dimension stone within a cultural setting.

Heat Shock Treatments: A New Safe Approach against Lichen Growth on Outdoor Stone Surfaces

The control of lichen growth, particularly important in the field of stone conservation of outdoor monuments, largely depends on the use of biocides, that may be dangerous for the users, the environment and the substratum. A new, alternative approach is proposed, which makes the most of a poorly known peculiarity of poikilohydrous organisms: they are thermo-tolerant (up to 65-70 °C) when dry, but thermo-sensitive when wet. The efficacy of thermal treatments (range: 20-55 °C), in parallel to the application of three biocides, was verified in the laboratory with six epi- and endolithic lichens. Chlorophyll a fluorescence emission was checked in treated and nontreated samples of all the species, whereas histochemical observations with a dead cell stain were carried out on one of them. The feasibility of the thermal treatments in the field was verified with a seventh species. The results confirm that a 6 h treatment at 55 °C is sufficient to kill the lichens if they are kept fully hydrated. At 40 °C the organisms are damaged: in this case biocides at concentrations 10× lower than in normal applications can profitably be used. The new protocol is simple, the field equipment cheap, and the negative effects associated with standard biocide treatments are absent.

Microscale biopitting by the endolithic lichen Verrucaria baldensis and its proposed role in mesoscale solution basin development on limestone

Data are reported demonstrating the potential role of microscale morphologies, induced by endolithic lichen communities, specifically Verrucaria baldensis, in the initiation and development of mesoscale solution basin formation on limestone in the Burren, County Clare. A biophysical model is proposed outlining the different microscale stages leading to solution basin initiation with a progression from initial lichen colonization and growth, associated perithecial biopitting followed by biopit coalescence to form biotroughs, their subsequent enlargement and eventual incipient solution basin formation. This model provides one explanation for solution basin development as this end state may also be achieved through simple solutional means without biological input. The complexity of interactions at the lichen–rock interface are identified with emphasis on the spatial and temporal variability of these underlining the point that, as with macrotopographies at the landscape scale, rock surface microtopographies also reflect historical weathering legacies. Copyright © 2011 John Wiley & Sons, Ltd.

Physical and Chemical Deterioration of Silicate and Carbonate Rocks by Meristematic Microcolonial Fungi and Endolithic Lichens (Chaetothyriomycetidae)

Physicochemical deterioration processes driven by lithobiontic microcolonial fungi (MCF) and endolithic lichens (EL) are still mostly unresolved. Here, the millimetric penetration of MCF strains within silicate and carbonate lithotypes was quantified. The influence of petrographic features in determining hyphal passageways satisfies a model of physical penetration during the early stages of colonization, already described for EL. The MCF and EL secretion of iron-chelating metabolites accounts for iron mobilization in desert-varnish formation, often putatively related to fungal colonization. Increased dissolution of limestone by the model iron chelator desferrioxamine indicates the possible involvement of these MCF and EL secretes in pitting carbonates.

The GenusSpheniaTurton, 1822 (Bivalvia: Myidae) from Shallow Waters of Argentina

Fil: Pastorino, Roberto Santiago Guido. Consejo Nacional de Investigaciones Cientificas y Tecnicas. Oficina de Coordinacion Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”; Argentina

An Investigation of the Role of Water (Rain and Dew) in Controlling the Growth Form of Lichens on Cobbles in the Negev Desert

Dew and rain were measured along the north facing slope (NFS, with epilithic lichens predominating) and the south-facing slope (SFS, with endolithic lichens predominating) in the Negev (P = 95 mm) in order to evaluate the relationships between the abiotic conditions and the lichen growth form. Although insignificant differences characterize the rain and dew amounts with the epilithic growth form, high correlation was obtained between dew duration (r2 = 0.73) and the combined duration of dew and rain (r2 = 0.79). The data imply that daylight wetness duration rather than the amount of precipitation may explain the predominance of epilithic lichens at the shaded NFS.

Snail herbivory decreases cyanobacterial abundance and lichen diversity along cracks of limestone pavements

Herbivores are known to decrease plant species diversity in ecosystems with low productivity. Limestone pavements are low-productive habitats harboring specialized communities of cyanobacteria, and endo- and epilithic lichens exposed to extreme temperature and humidity fluctuations. Pavements of the Great Alvar (O¨ land, Sweden) are covered by free-living cyanobacteria giving the rock surface a dark color. Based on cyanobacterial abundance along the edges, two types of cracks intersecting the pavements have been described: Type one with abundant cyanobacteria and type two without cyanobacteria resulting in light-colored edges. Erosion and different lengths of inundation by melt water have been suggested to cause the conspicuous differences in community composition and hence color between cracks. We hypothesized that this pattern results from the grazing activity of the cyanobacteria- and lichen-feeding snail Chondrina clienta, which reduces cyanobacterial cover along light-colored cracks and facilitates endolithic lichens. Three dark and three light-colored cracks were investigated at each of three localities. Crack characteristics (i.e., aspect, width, depth and erosion) and snail density were assessed at the crack level. Cyanobacterial cover and lichen diversity were recorded in 1-cm sections, sampled every 5 cm along eight 160-cm-long transects per crack. Model selection was applied to estimate effects of snail density and distance from crack edges on cyanobacterial abundance and lichen diversity. Crack characteristics explained no differences in cyanobacterial cover or lichen diversity. However, cyanobacterial cover decreased towards the edges of cracks with high snail densities. A transplant experiment supported the correlational evidence. The abundant cyanobacterial cover on pieces of stone placed close to cracks with high snail densities was completely grazed within 19 months. By contrast, cyanobacteria recolonized initially completely grazed pieces of stone when fixed near cracks without snails. Abundance and diversity of endolithic lichens increased along cracks with high compared to low snail densities but decreased in epilithic lichens and lichens with cyanobacterial symbionts. However, the presence of the gastropod herbivore decreased overall lichen diversity. Comparing presence-absence matrices with null models revealed that species co-occurred less frequently than expected. Taken together, we provide evidence that herbivory indirectly released endolithic lichens from competition for light by reducing cyanobacterial cover. (Less)

Respiration-induced weathering patterns of two endolithically growing lichens

The two endolithic lichen species Hymenelia prevostii and Hymenelia coerulea were investigated with regard to their thallus morphology and their effects on the surrounding substrate. The physiological processes responsible for the observed alterations of the rock were identified. Whereas the thallus surface of H.coerulea was level, H.prevostii formed small depressions that were deepest in the thallus center. In a cross-section, both species revealed an algal zone consisting of algal cavities parallel to the substrate surface and a fungal zone below. However, H.prevostii revealed significantly larger cavities with more than twice the cell number and a denser pattern of cavities than H.coerulea, resulting in a biomass per surface area being more than twice as large. Below H.prevostii the layer of macroscopically visibly altered rock material was about twice as deep and within this layer, the depletion of calcium and manganese was considerably higher. In simultaneous measurements of the oxygen uptake/oxygen release and pH shift, the isolated algal strains of both lichens revealed respiration-induced acidification of the medium in the dark. At higher light intensities, H.coerulea and to a lesser extent also H.prevostii alkalized the medium which may lessen the acidification effect somewhat under natural conditions. In a long-term growth experiment, the isolated algal strains of both lichens revealed acidification of the medium to a similar extent. Neither acidic lichen substances nor oxalic acid was identified. The significant differences between the weathering patterns of both species are based on the same respiration-induced acidification mechanism, with H.prevostii having a greater effect due to its higher biomass per area.

Lichen Colonization on Cobbles in the Negev Desert Following 15 Years in the Field

Rock slabs were placed from 1991 through 2006 at the bottom of a north- (NFS) and a south-facing (SFS) slope of a first order drainage basin in the Negev Desert Highlands, Israel (P = 95 mm) where natural cobbles have >95% cover of endolithic and epilithic lichens. Periodic measurements of rain, dew and chlorophyll content took place. Lichen cover and the regeneration time were aspect-dependent. Lichen cover was 98% and 61% at NFS and SFS, respectively, while the lichen's recovery time was respectively ∼45 and ∼80 years. The fast regeneration time may have important implications regarding weathering and soil formation.

Chlorophyll a fluorescence as a practical tool for checking the effects of biocide treatments on endolithic lichens

The efficacy of three biocides (Benzalkonium chloride, Koretrel, and Rocima 110) on two endolithic lichens ( Acrocordia conoidea and Bagliettoa marmorea) was monitored in the laboratory and in the field with portable pulse-amplitude-modulated (PAM) equipment. For comparison, epifluorescence microscope observations were carried out in the laboratory on cross-sections of treated and untreated thalli. The PAM equipment revealed excellent sensitivity for quantifying variations that could not be appreciated in epifluorescence. The two lichens showed a differential resistance to the biocides, probably related to the different photobionts and absorption of the active principle. The contribution of Chl a F techniques to the field of stone conservation is discussed.

In vitro receptivity of carbonate rocks to endolithic lichen-forming aposymbionts

Sterile cultured isolates of lichen-forming aposymbionts have not yet been used to investigate lichen–rock interactions under controlled conditions. In this study mycobionts and photobiont of the endolithic lichens Bagliettoa baldensis and Bagliettoa marmorea were isolated and inoculated with coupons of one limestone and four marbles commonly employed in the Cultural Heritage framework. After one year of incubation, microscopic observations of polished cross-sections were performed to verify if the typical colonization patterns observed in the field may be reproduced in vitro and to evaluate the receptivity of the five lithotypes to endolithic lichens. The mycobionts of the two species developed both on the surface of and within all the lithotypes, showing different penetration pathways which depend on mineralogical and structural features and highlight different receptivity. By contrast, algae inoculated with the coupons did not penetrate them. Observations suggest that the hyphal penetration along intrinsic discontinuities of rocks is a relatively fast phenomenon when these organisms are generally considered as slow-growing. Samples from limestone outcrops and abandoned marble quarries, colonized by the same species or other representatives of Verrucariaceae, showed penetration pathways intriguingly similar to those reproduced in vitro and highlighted that lichen-driven erosion processes only increase the availability of hyphal passageways after a long-term colonization. These results show that in vitro incubation of sterile cultured lichen-forming ascomycetes with rock coupons is a practicable experimental system to investigate the lichen–rock interactions under controlled conditions and, together with analysis in situ, may support decisions on conservative treatments of historical and cultural significant stone substrata.

Endolithic lichens, rapid biological weathering and schmidt hammer r‐values on recently exposed rock surfaces: storbreen glacier foreland, jotunheimen, norway

.The endolithic lichen Lecidea auriculata is known to enhance rock surface weathering on the Little Ice Age moraines of the glacier Storbreen in Jotunheimen, central southern Norway. This study demonstrates the reduction in Schmidt hammer Rvalues that followed the rapid colonization by this lichen of pyroxene‐granulite boulders on terrain deglaciated over the last 88 years. In the absence of this lichen, the characteristic mean R‐value of boulder surfaces is 61.0 ± 0.3; where this lichen is present, R‐values are lower by at least 20 units on surfaces exposed for 30–40 years. A similar reduction in rock hardness on rock surfaces without a lichen cover requires about 10 ka. The rapid initial weakening of the rock surfaces is indicative of rates of biological weathering by endolithic lichens that may be two orders of magnitude (200–300 times) faster than rates of physico‐chemical weathering alone. If not avoided, the effects of this type of lichen are likely to negate the effectiveness of the Schmidt hammer and other methods for exposure‐age dating, including cosmogenic‐nuclide dating, in severe alpine and polar periglacial environments. The results also suggest a new method for dating rock surfaces exposed for <50 years.

Life in extreme environments: survival strategy of the endolithic desert lichen Verrucaria rubrocincta

Verrucaria rubrocincta Breuss is an endolithic lichen that inhabits caliche plates exposed on the surface of the Sonoran Desert. Caliche surface temperatures are regularly in excess of 60 degrees C during the summer and approach 0 degrees C in the winter. Incident light intensities are high, with photosynthetically active radiation levels typically to 2,600 micromol/m(2) s(-1) during the summer. A cross-section of rock inhabited by V. rubrocincta shows an anatomical zonation comprising an upper micrite layer, a photobiont layer containing clusters of algal cells, and a pseudomedulla embedded in the caliche. Hyphae of the pseudomedulla become less numerous with depth below the rock surface. Stable carbon and oxygen isotopic data for the caliche and micrite fall into two sloping, well-separated arrays on a delta(13)C-delta(18)O plot. The delta(13)C(PDB) of the micrite ranges from 2.1 to 8.1 and delta(18)O(SMOW) from 25.4 to 28.9, whereas delta(13)C(PDB) of the caliche ranges from -4.7 to 0.7 and delta(18)O(SMOW) from 23.7 to 29.2. The isotopic data of the micrite can be explained by preferential fixing of (12)C into the alga, leaving local (13)C enrichment and evaporative enrichment of (18)O in the water. The (14)C dates of the micrite range from recent to 884 years b.p., indicating that "dead" carbon from the caliche is not a significant source for the lichen-precipitated micrite. The endolithic growth is an adaptation to the environmental extremes of exposed rock surfaces in the hot desert. The micrite layer is highly reflective and reduces light intensity to the algae below and acts as an efficient sunscreen that blocks harmful UV radiation. The micrite also acts as a cap to the lichen and helps trap moisture. The lichen survives by the combined effects of biodeterioration and biomineralization. Biodeterioration of the caliche concomitant with biomineralization of a protective surface coating of micrite results in the distinctive anatomy of V. rubrocincta.

The Mycobiont Role in Crustose Lichen Expansion on Cobbles in the Negev Desert

Whereas the role of the photobiont within microlichen symbiosis is indisputable, the role of the mycobiont in the symbiosis is still questionable. Here we report a unique expansion of the endolithic lichen Caloplaca alociza on cobbles in the Negev Desert due to structural change induced by the mycobiont burrowing activity. By increasing the water-holding capacity of the rock it facilitates lichen expansion from the cobble margins (where they establish due to water capillary rise) to its sides. And thus, while rare at the cobble top, lichen expansion is taking place to the sides of the cobble albeit the fact that dew (the main water source) is significantly lower at the cobble sides than at its top.

Endolithic Microbial Ecosystems

The endolithic environment, the pore space in rocks, is a ubiquitous microbial habitat and an interface between biology and geology. Photosynthesis-based endolithic communities inhabit the outer centimeters of rocks exposed to the surface, and offer model systems for microbial ecology, geobiology, and astrobiology. Endolithic ecosystems are among the simplest microbial ecosystems known and as such provide tractable models for testing ecological hypotheses. Such hypotheses have been difficult to test because microbial ecosystems are extraordinarily diverse. We review here recent culture-independent, ribosomal RNA-based studies that evaluate hypotheses about endolithic ecosystems, and provide insight for understanding general principles in microbial ecology. Comparison of endolithic communities supports the principle that patterns of microbial diversity are governed by similar principles observed in macroecological systems. Recent results also explore geobiological processes that shape the current biosphere and potentially provide clues to life's history on Earth and where to seek life elsewhere in the Solar System.