A comparison of the growth curves of the foliose lichen Parmelia conspersa determined by a cross-sectional study and by direct measurement

Changes in the radial growth rate (RGR mm/yr) through life were studied in thalli of the foliose lichen Parmelia conspersa by two methods: (1) a cross-sectional study (Study A) in which the RGR was measured in 60 thalli from 0.2 to 13 cm in diameter, and (2) by radial growth measurements over 4.5 years of fragments, consisting of a single major lobe, which were removed from large thalli and glued to pieces of slate (Study B). Both studies suggested there was a phase of increasing RGR in small thalli followed by a more constant phase, the latter beginning at approximately a thallus radius of 6-8 mm. However, in Study B significantly increased RGR was observed during the second 6-month growth period. This phase of growth was more likely to be due to an increase in lobe width than to an effect of climate. In addition, a lobe in a large thallus with both adjacent lobes removed significantly increased in width over 1 year compared with control lobes. These results suggest that (1) mean lobe width in a thallus may be determined by the intensity of marginal competition between adjacent lobes, and (2) changes in lobe width during the life of a lichen thallus may be a factor determining the establishment of the linear phase of growth in foliose lichens. © 1992.

The response of lichen growth to additions of distilled water, rainwater and water from a rock surface

Three lichen species were wetted in the field with distilled water, rainwater or water which had run off a rock surafce, during July 1974 to February 1975. The radial growth rate of Parmelia glabratula ssp. fuliginosa was not influenced by the wetting treatments. The radial growth rate of P. conspersa with the distilled water was greater than the control, rainwater and runoff treatments. The radial growth rate of Physcia orbicularis was lower with rainwater and runoff treatmentss than the control and distilled water treatment. These results may be explained by the effect of wetting on the carbon balance of the lichens and by the influence of water chemistry.

The reponse of lichen growth to transplantation to rock surfaces of different aspect

Thalli of four saxicolous lichens on slate rock fragments were transplanted from rock surafces to horizontal boards and then to south-east-facing and north-west-facing rock surfaces. The radial growth rate of Physcia orbicularis and Parmelia conspersa thalli declined after transplatation to north-west-facing rock surfaces and was unchanged after transplantation to south-east-facing rock surfaces cmpared with growth rates on the boards. The radial growth rate of P. glabratula ssp. fuliginosa thalli declined after transplantation to south-east-facing rock surfaces and was unchanged after transplantation to north-west-facing rock surfaces compared with grwoth rate on the boards. The radial growth rate of P. saxatilis thalli was similar on the horizontal boards, south-east-facing and north-west-facing rock surfaces. These results are dsicussed in relation to the aspect distribution of the four lichens in South Gwynedd, Wales.

Growth phases in the life of a lichen thallus

The growth rates of thalli of foliose saxicolous lichens before and after the linear phase of growth were measured in 1973. Changes in the radial growth rate (measured as mm/year) with thallus size in the prelinear phase (thalli less than approximately 1.5 cm in diameter) were consistent with the hypothesis that early growth of these lichens is loagarithmic. When growth in the prelinear phase was measured as a relative growth rate (measured as sq cm/sq cm/year) there was a rapid rise in growth rate until about 3 mm thallus diameter and then a decline in growth rate. The radial growth rate of non-fragmenting thalli when compared with fragmenting thalli at different stages of fragmentation suggested that radial growth rate does not significantly decline after fragmentation of the thallus. This result is not consistent with a postlinear phase in the radial growth of a lichen thallus.

Lichen competition: two-dimensional warfare in slow motion

Lichens, a symbiotic association between a filamentous fungus and an alga, are often dominant in stressful environments such as the surfaces of rock and tree bark. Under these conditions, lichens experience extremes of temperature, moisture supply, and low availability of nutrients. As a consequence, lichens sequester a high proportion of their carbon production for stress resistance rather than for growth. Hence, as a group lichens are particularly slow growing organisms with many species growing at less than 2mm per year and some at less than 0.5mm per year. Whether or not competition occurs between lichen thalli in these communities is controversial. This article discusses the evidence that competition occurs between lichens on rock and tree bark and assesses whether competitive effects are likely to be important in structuring these communities.

The use of lichen growth rings in lichenometry: some preliminary findings

Certain species of crustose lichens have concentrically zoned margins which probably represent yearly growth rings. These marginal growth rings offer an alternative method of studying annual growth fluctuations, establishing growth rate-size curves, and determining the age of thalli for certain crustose species. Hence, marginal growth rings represent a potentially valuable, unexploited, tool in lichenometry. In a preliminary study, we measured the widths of the successive marginal rings in 25 thalli of Ochrolechia parella (L.) Massal., growing at a maritime site in north Wales. Mean ring widths of all thalli varied from a minimum of 1.02 mm (the outermost ring) to a maximum of 2.06 mm (the third ring from the margin). There is some suggestion that marginal ring width and thallus size are positively correlated; and hence that growth rates increase in larger thalli in this small population. In a further study on recently exposed bedrock adjacent to Breidalon, SE Iceland, we examined the potential for using marginal growth rings to estimate thallus age of a lichen tentatively identified as a Rhizocarpon (possibly R. concentricum (Davies) Beltram.) and thus confirm the timing of surface exposure (c. 50 years). Collectively, these results suggest: 1) the measurement of marginal rings is a possible alternative method of studying the growth of crustose lichens; 2) O. parella may grow differently to other crustose species, exhibiting a rapidly increasing radial growth rate in thalli >40 mm; 3) where lichens with marginal rings grow on recently exposed surfaces (<60 yrs), minimum age estimates can be made using growth rings as an in situ indication of lichen growth rate; 4) it is suggested that this phenomenon could provide a valuable, previously unexploited, in situ lichenometric-dating tool in areas lacking calibration control.

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.

Statnote 32 : the partial correlation coefficient

In previous statnotes, the application of correlation and regression methods to the analysis of two variables (X,Y) was described. The most important statistic used to measure the degree of correlation between two variables is Pearson’s ‘product moment correlation coefficient’ (‘r’). The correlation between two variables may be due to their common relation to other variables. Hence, investigators using correlation studies need to be alert to the possibilities of spurious correlation and the methods of ‘partial correlation’ are one method of taking this into account. This statnote applies the methods of partial correlation to three scenarios. First, to a fairly obvious example of a spurious correlation resulting from the ‘size effect’ involving the relationship between the number of general practitioners (GP) and the number of deaths of patients in a town. Second, to the relationship between the abundance of the nitrogen-fixing bacterium Azotobacter in soil and three soil variables, and finally, to a more complex scenario, first introduced in Statnote 24involving the relationship between the growth of lichens in the field and climate.

The influence of environment on foliose lichen growth and its ecological significance

This chapter considers various aspects of the influence of the environment on the growth of foliose lichens and its significance in determining the ecology of individual species. Radial growth (RaG) and growth in mass of foliose lichens is influenced by climate and microclimate and also by substratum factors such as rock and bark texture, substrate chemistry, and nutrient enrichment. Seasonal fluctuations in growth, as measured by radial growth rate (RaGR) per month, often correlate best with average or total rainfall, the number of rain days, or rainfall in a specific season. Temperature has also been identified to be an important climatic factor influencing growth in some studies. Interactions between microclimatic factors and especially light intensity, temperature, and moisture status are important in determining differences in growth in relation to aspect and slope of the substratum. The physical and chemical nature of the substratum has a profound influence on the growth of foliose lichens. Hence, the effects of texture, porosity, rate of drying, and the physical changes of the substratum on growth are likely to influence lichen distributions. Bird droppings may influence growth and survival by smothering the thalli, altering the pH, or adding inhibitory and stimulatory compounds. Nitrogen and phosphate availability may also influence growth. Chemical factors also have an important influence on lichens of maritime rocks, the effect of salinity and calcium ions being of particular importance. Effects of environmental factors on growth influence the competitive ability of a lichen and ultimately its ecology and distribution.

Polyols in the crustose lichen Rhizocarpon geographicum

A lichen is an intimate association between an alga and a fungus and is regarded as one of the best examples of ‘mutualism’ or ‘symbiosis’ involving microorganisms. In lichens which have Trebouxia as the algal partner, photosynthesis by the algae results in the production of the soluble polyol ribitol which is then transported to the fungus where it is converted to arabitol and mannitol. Within the fungus, arabitol may act as a short-term carbohydrate reserve while mannitol may be involved in stress resistance. The crustose lichen Rhizocarpon geographicum (L.) DC., has an unusual thallus structure consisting of discrete granules (areolae) containing the algal component growing in association with a non-lichenised fungal hypothallus that extends beyond the areolae to form a marginal ring. The concentrations of ribitol, arabitol, and mannitol were measured, using gas chromatography, in the central areolae and marginal hypothallus of the crustose lichen Rhizocarpon geographicum (L.) DC. growing on slate rocks in north Wales, UK. The concentrations of all three polyols were greater in the central areolae than in the marginal hypothallus. In addition, the ratios of polyols in the marginal hypothallus to that in the central areolae varied through the year. The concentration of an individual poyol in the hypothallus was correlated primarily with the concentrations of the other polyols in the hypothallus and not to their concentrations in the areolae. Low concentration of ribitol, arabitol, and mannitol in the marginal hypothallus compared with the central areolae suggests either a lower demand for carbohydrate by the hypothallus or limited transport of polyols from areolae to hypothallus, and may explain the low growth rates of this species. In addition, polyols appear to be partitioned differently through the year with an increase in mannitol compared with arabitol in more stressful periods.

Concentration of mannitol and other soluble carbohydrates in the crustose lichen Rhizocarpon geographicum

In symbiotic lichens which have Trebouxia as the algal partner, photosynthesis by the algae results in the production of the soluble carbohydrate ribitol which is then transported to the fungus where it is converted to arabitol and mannitol. Within the fungus, arabitol may act as a short-term carbohydrate reserve while mannitol may have a more protective function and be important in stress resistance. The concentrations of ribitol, arabitol, and mannitol were measured, using gas chromatography, in the central areolae and marginal hypothallus of the crustose lichen Rhizocarpon geographicum (L.) DC. growing on slate rocks in north Wales, UK. The concentrations of all three soluble carbohydrates were greater in the central areolae than in the marginal prothallus. In addition, the ratio of mannitol in the prothallus to that in the areolae was least in July. The concentration of an individual carbohydrate in the prothallus was correlated primarily with the concentrations of the other carbohydrates in the prothallus and not to their concentrations in the areolae. Low concentration of ribitol, arabitol, and mannitol in the marginal prothallus compared with the central areolae suggests either a lower demand for carbohydrate by the prothallus or limited transport from areolae to prothallus and may explain the low growth rates of this species. In addition, soluble carbohydrates appear to be partitioned differently through the year with an increase in mannitol compared with arabitol in more stressful periods.

The influence of environment on foliose lichen growth and its biological significance

Radial growth and growth in mass of lichens is influenced by climatic and microclimatic factors and also by substratum factors such as rock and bark texture, chemistry, and nutrient enrichment. Seasonal fluctuations in growth, as measured by radial growth rate (RaGR) per month, often correlate best with average or total rainfall, the number of rain days, or rainfall in a specific season. Temperature is also considered to be an important climatic factor in some studies. Interactions between microclimatic factors and especially light intensity, temperature, and moisture are the most important in determining local annual growth rates. The physical and chemical nature of the substratum has a profound influence on the growth of foliose lichens. Hence, the effects of texture, porosity, rate of drying, and the physical changes of the substratum on growth are likely to influence lichen distributions. Bird droppings may influence growth and survival by smothering the thalli, altering the pH, or adding inhibitory and stimulatory compounds. Nitrogen and phosphate availability may also influence growth. Chemical factors may also have an important influence on lichens of maritime rocks, the effect of salinity and calcium ions being of particular importance. Zinc, copper, and mercury may also be important in lichen growth as they have been shown to affect the chlorophyll content of lichen algae. Effects of environmental factors on growth influence the competitive ability of lichens thus influencing their ecology and distribution.

Polyols in the crustose lichen rhizocarpon geographicum

A lichen is an intimate association between an alga and a fungus and is regarded as one of the best examples of ‘mutualism’ or ‘symbiosis’ involving microorganisms. In lichens which have Trebouxia as the algal partner, photosynthesis by the algae results in the production of the soluble polyol ribitol which is then transported to the fungus where it is converted to arabitol and mannitol. Within the fungus, arabitol may act as a short-term carbohydrate reserve while mannitol may be involved in stress resistance. The crustose lichen Rhizocarpon geographicum (L.) DC., has an unusual thallus structure consisting of discrete granules (areolae) containing the algal component growing in association with a non-lichenised fungal hypothallus that extends beyond the areolae to form a marginal ring. The concentrations of ribitol, arabitol, and mannitol were measured, using gas chromatography, in the central areolae and marginal hypothallus of the crustose lichen Rhizocarpon geographicum (L.) DC. growing on slate rocks in north Wales, UK. The concentrations of all three polyols were greater in the central areolae than in the marginal hypothallus. In addition, the ratios of polyols in the marginal hypothallus to that in the central areolae varied through the year. The concentration of an individual poyol in the hypothallus was correlated primarily with the concentrations of the other polyols in the hypothallus and not to their concentrations in the areolae. Low concentration of ribitol, arabitol, and mannitol in the marginal hypothallus compared with the central areolae suggests either a lower demand for carbohydrate by the hypothallus or limited transport of polyols from areolae to hypothallus, and may explain the low growth rates of this species. In addition, polyols appear to be partitioned differently through the year with an increase in mannitol compared with arabitol in more stressful periods.

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.