Effect of water vapour condensation on the radon content in subsurface air in a hypogeal inactive-volcanic environment in Galdar cave, Spain

Fluctuations of trace gas activity as a response to variations in weather and microclimate conditions were monitored over a year in a shallow volcanic cave (Painted Cave, Galdar, Canary Islands, Spain). 222Rn concentration was used due to its greater sensitivity to hygrothermal variations than CO2 concentration. Radon concentration in the cave increases as effective vapour condensation within the porous system of the rock surfaces inside the cave increases due to humidity levels of more than 70%. Condensed water content in pores was assessed and linked to a reduction in the direct passage of trace gases. Fluctuations in radon activity as a response to variations in weather and microclimate conditions were statistically identified by clustering entropy changes on the radon signal and parameterised to predict radon concentration anomalies. This raises important implications for other research fields, including the surveillance of shallow volcanic and seismic activity, preventive conservation of cultural heritage in indoor spaces, indoor air quality control and studies to improve understanding of the role of subterranean terrestrial ecosystems as reservoirs and/or temporary sources of trace gases.

Competitive interactions between four foliose lichen species with and without nutrient enrichment

Competition between four foliose lichen species, common on slate rock surfaces in South Gwynedd, Wales, UK, was studied in experimental plots with and without nutrient enrichment by bird droppings. Fragments of the four lichens were glued to pieces of slate on horizontal boards in monoculture and in two-, three- and four-species mixtures in a factorial experimental design. In monoculture, nutrient enrichment increased thallus area of Parmelia conspersa (Ehrh. ex. Ach.) Ach., decreased thallus areas of Parmelia saxatilis (L.) Ach. and Parmelia glabratula ssp. fuliginosa (Fr. ex. Duby) Laundon, and did not affect thallus area of Phaeophyscia orbicularis (Necker) Moberg compared with untreated thalli. In the mixtures, P. conspersa and Ph. orbicularis were equally effective competitors in plots with and without nutrient enrichment. Addition of bird droppings, however, altered the ability of P. saxatilis and P. glabratula ssp. fuliginosa, to compete with the other species, the competitive ability of both species being reduced in some mixtures but increased in others. The results suggest that nutrient enrichment may alter the competitive balance between the four lichen species and this may be a factor determining their relative abundance on rock surfaces in South Gwynedd.

Competition between three lichen species using a factorial experimental design

Competition between three foliose lichens common on slate rock in south Gwynedd, Wales was studied in the field using a factorial experimental design. The lichens were grown as fragments glued to pieces of slate in two- and three-species mixtures. In the two-species mixtures, Parmelia conspersa (Ehrh. Ex Ach.) Ach. outcompeted Parmelia glabratula ssp. fuliginosa (Fr. ex Duby) Laund. strongly and Physcia orbicularis (Neck.) Poetsch less strongly, while P. orbicularis outcompeted P. glabratula weakly. Significant two-factor interactions indicated that the results from the three-species mixture could not be predicted from the two-species mixtures. Parmelia glabratula and P. orbicularis grew better in the presence of two competitors than one. This result suggests that the three species may co-occur on well-lit rock surfaces at the site.

The biology of the crustose lichen Rhizocarpon geographicum

The crustose lichen Rhizocarpon geographicum (L.) DC. comprises yellow-green lichenized areolae which develop and grow on the surface of a non-lichenized fungal hypothallus, the latter extending beyond the edge of the areolae to form a marginal ring. The hypothallus advances very slowly and the considerable longevity of R. geographicum, especially in Arctic and Alpine environments, has been exploited by geologists in dating the exposure age of rock surfaces (lichenometry). This review explores various aspects of the biology of R. geographicum including: (1) structure and symbionts, (2) lichenization, (3) development of areolae, (4) radial growth rates (RaGR), (5) growth physiology, (6) changes in RaGR with thallus size (growth ratesize curve), (7) maturity and senescence, and (8) aspects of ecology. Lichenization occurs when fungal hyphae become associated with a compatible species of the alga Trebouxia, commonly found free-living on the substratum. Similarly, 'primary' areolae develop from free-living algal cells trapped by the advancing hypothallus. The shape of the growth rate-size curve of R. geographicum is controversial but may exhibit a phase of decreasing growth in larger thalli. Low rates of translocation of carbohydrate to the hypothallus together with allocation for stress resistance results in very slow RaGR, a low demand for nutrients, hence, the ability of R. geographicum to colonize more extreme environments. Several aspects of the biology of R. geographicum have implications for lichenometry including early development, mortality rates, the shape of the growth-rate size curve, and competition. © The Author(s) 2012.

Factors associated with degeneration of the thallus centre in foliose lichens

Degeneration of the older parts of foliose lichen thalli often lead to the formation of a space or 'window' in the centre of the colonies. The percentage of thalli of different size which exhibited 'windows' was studied in twenty saxicolous lichen populations in south Gwynedd, Wales. The proportion of thalli with 'windows' increased with thallus size. The size class at which 50% and 100% of thalli exhibited 'windows' varied between populations. Differences between populations were not correlated with distance from the sea, aspect, slope or porosity of the substrate or the total number of lichen species present. However, a higher percentage of smaller thalli had 'windows' on rock surfaces with a greater lichen cover. There were no significant differences in the levels of Ca, Mg, Cu or Zn in large (>4 cm) and small (<2 cm) Parmelia conspersa (Ehrh. ex Ach.) Ach. thalli or in the centres and marginal lobes of these thalli. The concentration of ribitol, arabitol and mannitol was significantly reduced in the centre of large thalli compared with the margin of large thalli and the centre of small thalli. However, carbohydrate levels were similar in the centre of large thalli and the margin of small thalli. The data suggest that loss of the thallus centre is a degenerative process related to thallus size. In the field, the formation of 'windows' may be related to the intensity of competition on a substrate. Central degeneration was not associated with a deficiency or an accumulation of Ca, Mg, Cu and Zn in the thallus centre. However, degeneration may be associated with a reduction in carbohydrates in the centre compared with the marginal lobes.

The levels of ribitol, arabitol and mannitol in individual lobes of the lichen Parmelia conspersa (Ehrh. ex Ach.) ACH.

The levels of the soluble carbohydrates ribitol, arabitol and mannitol were measured in individual lobes of the lichen Parmelia conspersa (Ehrh. ex Ach.) Ach. Lobes were collected from a north and a south facing slate rock surface in South Gwynedd, Wales, U.K. on 4 days during 1990-1991. On each day sampled, the most significant variation in soluble carbohydrate levels was between the individual lobes of a thallus. In addition, carbohydrate levels were significantly greater on the south facing rock surface on 2 of the 4 days sampled. Factorial analyses of variance suggested that the levels of individual carbohydrates varied significantly between days but not between north and south facing rock surfaces. Mannitol levels varied less between days than arabitol levels. Levels of ribitol, arabitol and mannitol were positively correlated in individual lobes. A stepwise multiple regression suggested that on the north facing rock surface, arabitol and mannitol levels could be explained by variations in the level of ribitol. By contrast, on the south facing rock surface, the levels of fungal carbohydrates were less dependent on the level of ribitol and there was evidence of a relationship between arabitol and mannitol. Variations in carbohydrate production, allocation and metabolism could help to explain lobe growth variation in foliose lichens and the radial growth of lobes over a longer period of time. Greater carbohydrate production rather than differences in allocation and metabolism may explain the increased growth and frequency of P. conspersa on south facing rock surfaces in South Gwynedd. © 1994.

The influence of the frequency of wetting and drying on the radial growth of three saxicolous lichens in the field

Three lichen species were wetted with distilled water at different frequencies during August 1973 to July 1974. The radial growth rates of Parmelia glabratula ssp. fuliginosa and Physcia orbicularis thalli declined with increased wetting while the radial growth rate of Parmelia conspersa thalli increased with wetting frequency until ten experimental wettings per month but at fifteen wettings per month fell to a value near to the control. In the summer months, wetting resulted in a decline in the radial growth of P. glabratula ssp fuliginosa compared with the control but had little influence on the growth of P. conspersa and Physcia orbicularis. In the winter months, wetting had no significant influence on the radial growth of Parmelia glabratula ssp. fuliginosa, while the radial growth of P. conspersa increased and Physcia orbicularis declined compared with controls. These results are interpreted physiologically and in relation to the aspect distribution of the three lichens on rock surfaces.

The influence of aspect on the pattern of saesonal growth in the lichen Parmelia glabratula ssp. fuliginosa (Fr. ex Duby) Laund

The radial growth of samples of thalli of Parmelia glabratula ssp. fuliginosa were measured in situ on a south-facing and a northwest-facing rock surface each month from August 1973 to July 1974. In the periods August to October 1973 and March to July 1974 the radial growth of thalli in the northwest population was greater than in the south population. In the period November 1973 to February 1974 the radial growth of thalli in the south population was greater than in the northwest population. A physiological basis for the differences in seasonal growth in the two populations was suggested. The mean annual radial growth rate (in units of mm/year) was not significantly different in the two populations. However, the variability in radial growth rate between thalli was signifiacntly larger in the northwest than in the south population. These results may be explained by genetic difference between the populations and environmental differences between the rock surfaces.

The biology of the lichen Rhizocarpon geographicum: symbionts, growth, ecology, and lichenometry

Rhizocarpon geographicum (L.) DC. is one of the most widely distributed species of crustose lichens. This unusual organism comprises yellow-green ‘areolae’ that contain the algal symbiont which develop and grow on the surface of a non-lichenized, fungal ‘hypothallus’ that extends beyond the margin of the areolae to form a marginal ring. This species grows exceptionally slowly with annual radial growth rates (RGR) as low as 0.07 mm yr-1 and its considerable longevity has been exploited by geologists in the development of methods of dating the age of exposure of rock surfaces and glacial moraines (‘lichenometry’). Recent research has established some aspects of the basic biology of this important and interesting organism. This chapter describes the general structure of R. geographicum, how the areolae and hypothallus develop, why the lichen grows so slowly, the growth rate-size curve, and some aspects of the ecology of R. geographicum including whether the lichen can inhibit the growth of its neighbours by chemical means (‘allelopathy’). Finally, the importance of R. geographicum in direct and indirect lichenometry is reviewed.

Within-site variation in lichen growth rates and its implications for direct lichenometry

Variation in lichen growth rates poses a significant challenge for the application of direct lichenometry, i.e. the construction of lichen dating curves from direct measurement of growth rates. To examine the magnitude and possible causes of within-site growth variation, radial growth rates (RaGRs) of thalli of the fast-growing foliose lichen Melanelia fuliginosa ssp. fuliginosa (Fr. ex Duby) Essl. and the slow-growing crustose lichen Rhizocarpon geographicum (L.) DC. were studied on two S-facing slate rock surfaces in north Wales, UK using digital photography and an image analysis system (Image-J). RaGRs of M. fuliginosa ssp. fuliginosa varied from 0.44 to 2.63 mmyr-1 and R. geographicum from 0.10 to 1.50 mmyr-1.5. Analysis of variance suggested no significant variation in RaGRs with vertical or horizontal location on the rock, thallus diameter, aspect, slope, light intensity, rock porosity, rock surface texture, distance to nearest lichen neighbour or distance to vegetation on the rock surface. The frequency distribution of RaGR did not deviate from a normal distribution. It was concluded that despite considerable growth rate variation in both species studied, growth curves could be constructed with sufficient precision to be useful for direct lichenometry. © 2014 Swedish Society for Anthropology and Geography.

Lichen growth and lichenometry

Lichenometry is one of the most widely used methods available for dating the surface age of various substrata including rock surfaces, boulders, walls, and archaeological remains. It depends on the assumption that if the lag time before colonisation of a substratum by a lichen is known and lichen age can be estimated, then a minimum date can be obtained by measuring the diameter (or another property related to size) of the largest lichen at the site. Lichen age can be determined by variety of methods including calibrating lichen size against surfaces of known age (‘indirect lichenometry’), by constructing a growth rate-size curve from direct measurement of lichen growth (‘direct lichenometry’), using radio-carbon (RC) dating, and from lichen ‘growth rings’. This chapter describes: (1) lichen growth rates and longevity, (2) methods of estimating lichen age, (3) the methodology of lichenometry and (4) applications of lichenometry. Despite its limitations, lichenometry is likely to continue to play an important role in dating a variety of surfaces and also in providing data that contribute to the debate regarding global warming and climate change.

The use of the lichen genus Rhizocarpon in lichenometric dating with special reference to Holocene glacial events

Lichenometry is one of the most widely used methods of dating the surface age of substrata including rock surfaces, boulders, walls, and archaeological remains and has been particularly important in dating late Holocene glacial events. Yellow-green species of the crustose genus Rhizocarpon have been the most useful lichens in lichenometry because of their low growth rates and longevity. This review describes: (1) the biology of the genus Rhizocarpon, (2) growth rates and longevity, (3) environmental growth effects, (4) methods of estimating lichen age, (5) the methodology of lichenometry, (6) applications to dating glacial events, and (7) future research. Lichenometry depends on many assumptions, most critically that if the lag time before colonisation of a substratum is known and lichen age can be estimated, then a minimum surface age date can be obtained by measuring the size of the largest Rhizocarpon thallus. Lichen age can be estimated by calibrating thallus size against surfaces of known age (‘indirect lichenometry’), by constructing a growth rate-size curve from direct measurement of growth (‘direct lichenometry’), using radio-carbon (RC) dating, or from lichen ‘growth rings’. Future research should include a more rigorous investigation of the assumptions of lichenometry, especially whether the largest thallus present at a site is a good indicator of substratum age, and further studies on the establishment, development, growth, senescence, and mortality of Rhizocarpon lichens.

The structure and dynamics of saxicolous lichen communities

The saxicolous lichen vegetation on Ordovician slate rock at the mouth of the River Dovey, South Merionethshire, Wales was described in relation to several environmental variables which include aspect, slope angle, light intensity, rock porosity, rock microtopography and rock stability. Each of the measured environmental variables was shown to influence the lichen vegetation. A number of groups of species which were characteristic of certain environments were described. The data from the saxicolous lichen communities were analysed using multivariate analysis. Qualitative and quantitative data were ordinated, the qualitative data being easier to interpret ecologically, and site number (which reflects distance from the sea and altitude), rock porosity and light intensity were shown to be important environmental variables. A classification of the data was also carried out. The results of the ordination and classification were combined together and a model constructed which describes saxicolous lichen vegetation. A method which uses the model as an aid to the design and interpretation of field experiments is described. The model is applied to an experiment which investigates the effect on growth of transplanting four saxicolous lichens to different aspects. Growth was inhibited in Physcia orbicularis and Parmelia conspersa on rock surfaces of northwest aspect compared with growth on rock surfaces of southeast aspect. Growth was inhibited in Parmelia glabratula ssp. fuliginosa on rock surfaces of southeast aspect compared with rock surfaces of northwesr aspect. The growth of Parmelia saxatilis was similar at both southeast and northwesr aspects. Growth inhibition or stimulation in thalli of Physcia orbicularis, Parmelia conspersa and Parmelia glabratula ssp. fuliginosa after transplantation was consistent with the predictions of the model while the results for Parmelia saxatilis were not as expected. There was evidence that the frequency of Parmelia conspersa and Parmelia glabratula at a site is related to an effect of the environment on the growth of the thalli. There was also evidence that the frequency of Physcia orbicularis at a site is related to an effect of the environment on the establishment phase of the thalli and for the competitive exclusion of Parmelia saxatilis thalli from southeast facing rock surfaces. The distribution of lichens in relation to height on nine rock surfaces was investigated. It was suggested that the distribution of the lichens was influenced by microclimatic factors which are related to height on the rock, environmental variables which are associated with the rock substratum (e.g. rock porosity and rock microtopography) and by historical factors. The pattern of one crustose and one foliose lichen on four rock surfaces of different aspect and slope was investigated. On the vertically inclined surface the density of small thalli of Buellia aethalea and Parmelia glabratula ssp fuliginosa was correlated with the microtopography of the surface in transects horizontally across the rock surface but not in transects vertically down the rock surface. there were consitent differences in the scale and intensity of pattern horizontally and vertically and also a decrease in the intensity of pattern vertically as the slope of the rock surface decreased. These results were consistent with the suggestion of a gradient of microclimatic factors up the rock. The differences in the scale and intensity of pattern in different size classes in the population were consistent with the changes in pattern with time which have been shown to occur during succession in sand dune and salt marsh vegetation. The relationship between thallus size and height on a rock surface and between the radial growth rate and location of a thallus on a rock surface were investigated. Thalli of Parmelia glabratula ssp. fuliginosa were larger at the top of the rock surface than at the bottom and the data were consistent with the suggestion that the colonisation of the rock surface began at the top and, in time, spread downwards. The radial growth rate of the thalli could not be related to variation in slope, porosity, microtopography or directly to height on the rock but could be related to the horizontal location of the thalli on the rock. These results were consistent with the suggestion that here is a gradient of microclimatic factors across the rock surface which is also modified by height on the rock surface. The succession of lichen communities was described by relating the vegetation to rock porosity, rock microtopography, species diversity and rock stability. An initial stage dominated by crustose lichens leads to communities dominated by crustose, foliose and fruticose species. In the late stages of the succession on some rock surfaces crustose species again become dominant. The occurrence of the climax state and cyclic vegetation change in lichen communities are discussed. A mthod of estimating the age structure of a lichen population by relating thallus size to growth rate is described. The sources of error in the method are discussed in some detail and several refinements suggested to increase the accuracy of the method. The population dynamics of Parmelia glabratula ssp. fuliginosa was investigated by applying life tables to the age structures of eight different populations. The data were consistent with a period of relatively constant recruitment of thalli into the populations. Mortality in lichen populations was divided into deaths which occur after fragmentation of the thallus and deaths which occur after catastrophic environmental events. THe data suggest that the rate of fragmenting death is dependent on the age of the thallus while the rate of catastrophic death is dependent on the number of thalli established in an age class. A comparison of the numbers of thalli in each age class in the eight populations suggested that population density is controlled firstly, by climate and secondly, by variables related to the local rock surface environment. The rate of fragmenting death is related to the diversity of the community and the influence of diversity together with environmental variables in fluctuating or cyclic changes in population number.

Lichenometric dating (lichenometry) and the biology of the lichen genus rhizocarpon:challenges and future directions

Lichenometric dating (lichenometry) involves the use of lichen measurements to estimate the age of exposure of various substrata. Because of low radial growth rates and considerable longevity, species of the crustose lichen genus Rhizocarpon have been the most useful in lichenometry. The primary assumption of lichenometry is that colonization, growth and mortality of Rhizocarpon are similar on surfaces of known and unknown age so that the largest thalli present on the respective faces are of comparable age. This review describes the current state of knowledge regarding the biology of Rhizocarpon and considers two main questions: (1) to what extent does existing knowledge support this assumption; and (2) what further biological observations would be useful both to test its validity and to improve the accuracy of lichenometric dates? A review of the Rhizocarpon literature identified gaps in knowledge regarding early development, the growth rate/size curve, mortality, regeneration, competitive effects, colonization, and succession on rock surfaces. The data suggest that these processes may not be comparable on different rock surfaces, especially in regions where growth rates and thallus turnover are high. In addition, several variables could differ between rock surfaces and influence maximum thallus size, including rate and timing of colonization, radial growth rates, environmental differences, thallus fusion, allelopathy, thallus mortality, colonization and competition. Comparative measurements of these variables on surfaces of known and unknown age may help to determine whether the basic assumptions of lichenometry are valid. Ultimately, it may be possible to take these differences into account when interpreting estimated dates.

Molecular identification and localization of filamentous symbiotic bacteria associated with the hydrothermal vent annelid Alvinella pompejana

Alvinella pompejana is a polychaetous annelid that inhabits high temperature environments associated with active deep-sea hydrothermal vents along the East Pacific Rise. A unique and diverse epibiotic microflora with a prominent filamentous morphotype is found associated with the worm's dorsal integument. A previous study established the taxonomic positions of two epsilon proteobacterial phylotypes, 13B and 5A, which dominated a clone library of 16S rRNA genes amplified by PCR from the epibiotic microbial community of an A. pompejana specimen. In the present study deoxyoligonucleotide PCR primers specific for phylotypes 13B and 5A were used to demonstrate that these phylotypes are regular features of the bacterial community associated with A. pompejana. Assaying of other surfaces around colonies of A. pompejana revealed that phylotypes 13B and 5A are not restricted to A. pompejana. Phylotype 13B occurs on the exterior surfaces of other invertebrate genera and rock surfaces, and phylotype 5A occurs on a congener, Alvinella caudata. The 13B and 5A phylotypes were identified and localized on A. pompejana by in situ hybridization, demonstrating that these two phylotypes are, in fact, the prominent filamentous bacteria on the dorsal integument of A. pompejana. These findings indicate that the filamentous bacterial symbionts of A. pompejana are epsilon Proteobacteria which do not have an obligate requirement for A. pompejana.