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.

Studies on the growth rates of lichens

Progress in the field of lichen growth rates is briefly reviewed. The application of a new method of measuring growth rate to thalli of different size has led to the conclusion that there are changes in the radial growth rate during the life of a lichen thallus. For most of the life of a lichen thallus the radial growth rate is constant and the thallus radius increases linearly. Preceeding the linear phase the radial growth rate increases with time and the thallus radius increases logarithmically. There is no evidence for a postlinear phase in the radial growth of a lichen thallus. Studies on the growth rate of lichens are applied both to the problems of determining the age of a lichen thallus on an undated substratum and to an ecological investigation in the field.

Could lichens grow on Mars?

Lichens meet some but not all of the criteria that must be fulfilled by inhabitants of Mars. They could withstand many aspects of the hostile environment especially if they live within the rocks as they do in the dry valleys of Antarctica. Lichens, however, are dual organisms and we have to presuppose the successful establishment of a variety of microorganisms on Mars and especially algae and fungi. To date, the evidence for the existence of microorganisms in Martian meteorites is controversial and there is no conclusive evidence of present life on the surface. In addition, if endolithic lichens have evolved on Mars and are alive today they would be subjected to a considerably more hostile environment than the extreme environments on Earth, which are regarded as at the limit of tolerance of present day lichens. The lack of liquid water over most of the surface and the problem of obtaining sufficient nitrogen resources are particular problems for Martian lichens. Further landings on Mars, scheduled for 2005 and future missions are likely to increase substantially our knowledge of the Martian surface and the possibilities for life by attempting to bring back samples of rock and minerals. In addition, the use of techniques such as Laser Raman technology and the development of gas chromatographic methods for use in space increase the probability that an answer to the question of whether lichens have existed on Mars will be obtained in the near future.

Lichens, lichenometry and global warming

Lichenometry is one of many techniques now available for estimating the elapsed time since the exposure of a substratum. Its advantages include an ability to date surfaces during the last 500 years, a time interval in which radiocarbon dating is least efficient, and provides a quick, cheap, and relatively accurate date for a substratum.

Studies on the growth rates of lichens

Progress in the field of lichen growth rate studies is briefly reviewed. The application of a new method of measuring growth rate to thalli of different size has led to the conclusion that there are changes in the radial growth rate during the life of a lichen thallus. For most of the life of a lichen thallus the radial growth rate is constant and the thallus radius increases linearly. Preceding the linear phase the radial growth rate increases with time and the thallus radius increases logarithmically. There is no evidence for a postlinear phase in the radial growth of a lichen thallus. Studies on the growth rate of lichens are applied both to the problems of determining the age of a lichen thallus on an undated susbtratum and to an ecological investigation in the field.

Growth of crustose lichens: a review

Crustose species are the slowest growing of all lichens. Their slow growth and longevity, especially of the yellow-green Rhizocarpon group, has made them important for surface-exposure dating (‘lichenometry’). This review considers various aspects of the growth of crustose lichens revealed by direct measurement including: 1) early growth and development, 2) radial growth rates (RGR, mm yr-1), 3) the growth rate-size curve, and 4) the influence of environmental factors. Many crustose species comprise discrete areolae that contain the algal partner growing on the surface of a non-lichenised fungal hypothallus. Recent data suggest that ‘primary’ areolae may develop from free-living algal cells on the substratum while ‘secondary’ areolae develop from zoospores produced within the thallus. In more extreme environments, the RGR of crustose species may be exceptionally slow but considerably faster rates of growth have been recorded under more favourable conditions. The growth curves of crustose lichens with a marginal hypothallus may differ from the ‘asymptotic’ type of curve recorded in foliose and placodioid species and the latter are characterized by a phase of increasing RGR to a maximum and may be followed by a phase of decreasing growth. The decline in RGR in larger thalli may be attributable to a reduction in the efficiency of translocation of carbohydrate to the thallus margin or to an increased allocation of carbon to support mature ‘reproductive’ areolae. Crustose species have a low RGR accompanied by a low demand for nutrients and an increased allocation of carbon for stress resistance; therefore enabling colonization of more extreme environments.

Growth of foliose lichens: a review

This review considers various aspects of the growth of foliose lichens including early growth and development, variation in radial growth rate (RaGR) of different species, growth to maturity, lobe growth variation, senescence and fragmentation, growth models, the influence of environmental variables, and the maintenance of thallus symmetry. The data suggest that a foliose lichen thallus is essentially a ‘colony’ in which the individual lobes exhibit a considerable degree of autonomy in their growth processes. During development, recognisable juvenile thalli are usually formed by 15 months to 4 years while most mature thalli exhibit RaGR between 1 and 5 mm yr-1. RaGR within a species is highly variable. The growth rate-size curve of a foliose lichen thallus may result from growth processes that take place at the tips of individual lobes together with size-related changes in the intensity of competition for space between the marginal lobes. Radial growth and growth in mass is influenced by climatic and microclimatic factors and also by substratum factors such as rock and bark texture, chemistry, and nutrient enrichment. Possible future research topics include: (1) measuring fast growing foliose species through life, (2) the three dimensional changes that occur during lobe growth, (3) the cellular changes that occur during regeneration, growth, and division of lobes, and (4) the distribution and allocation of the major lichen carbohydrates within lobes.

The influence of environmental factors on the growth of lichens in the field

The majority of studies of the effects of environmental factors on lichen growth have been carried out in the field. Growth of lichens in the field has been measured as absolute growth rate (e.g., length growth, radial growth, diameter growth, area growth, or dry weight gain per unit of time) or as a relative growth rate, expressed per unit of thallus area or weight, e.g., thallus specific weight. Seasonal fluctuations in growth in the field often correlate best with changes in average or total rainfall or frequency of rain events through the year. In some regions of the world, temperature is also an important climatic factor influencing growth. Interactions between microclimatic factors such as light intensity, temperature, and moisture are particularly important in determining local differences in growth especially in relation to aspect and slope of rock surface, or height on a tree. Factors associated with the substratum including type, chemistry, texture, and porosity can all influence growth. In addition, growth can be influenced by the degree of nutrient enrichment of the substratum associated with bird droppings, nitrogen, phosphate, salinity, or pollution. Effects of environmental factors on growth can act directly to restrict species distribution or indirectly by altering the competitive balance among different species in a community.

'Growth rings' in crustose lichens:comparison with directly measured growth rates and implications for lichenometry

Some species of crustose lichens, such as Ochrolechia parella (L.) Massal., exhibit concentric marginal rings, which may represent an alternative technique of measuring growth rates and potentially, a new lichenometric dating method. To examine this hypothesis, the agreement and correlation between ring widths and directly measured annual radial growth rates (RaGR, mm a^-1) were studied in 24 thalli of O. parella in north Wales, UK, using digital photography and image analysis. Variation in ring width was observed at different locations around a thallus, between thalli, and from year to year. The best agreement and correlation between ring width and lichen growth rates was between mean width of the outer two rings (measured in 2011) and mean RaGR (in 2009/10). The O. parella data suggest that mean width of the youngest two growth rings, averaged over a sample of thalli, is a predictor of recent growth rates and therefore could be used in lichenometry. Potential applications include as a convenient method of comparing lichen growth rates on surfaces in different environmental settings; and as an alternative method of constructing lichen growth-rate curves, without having to revisit the same lichen thalli over many years. However, care is needed when using growth rings to estimate growth rates as: growth ring widths may not be stable; ring widths exhibit spatial and temporal variation; rings may not represent 1-year's growth in all thalli; and adjacent rings may not always represent successive year's growth.

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 in lichen communities

Lichens are symbiotic organisms that often dominate stressful environments such as the surfaces of rock and tree bark. Whether or not competition occurs between lichens in these environments, however, is controversial. This review considers various aspects of the competitive interactions between lichens including the observational studies that suggest competitive effects may be important, the methods that have been used to study lichen competition in the field, the result of marginal contacts between lichen thalli, the attributes that may give a species a competitive advantage, and the role of competition in structuring lichen communities. These studies suggest that competition for space and light does occur in lichen communities and that individual lichen species can be excluded from a substratum as a result of competition. Moreover, competitive interactions in multi-species communities can also lead to stable assemblages of species. Future research should consider those aspects of the lichen symbiosis that may confer a competitive advantage and the factors that may promote stability in multi-species communities. Studies of competition in lichen communities may have implications for other stressful environments in which symbiotic organisms play a significant role. ©2007 Balaban.

Lobe growth variation and the maintenance of symmetry in foliose lichen thalli

The aim of this study was to determine how thallus symmetry could be maintained in foliose lichens when variation in the growth of individual lobes may be high. Hence, the radial growth of a sample of lobes was studied monthly, over 22 months, in 7 thalli of Parmelia conspersa (Ehrh. Ex Ach.) Ach. And 5 thalli of P. glabratula ssp fuliginosa (fr. ex Duby) Laund. The degree of variation in the total radial growth of different lobes within a thallus over 22 months varied between thalli. Individual lobes showed a fluctuating pattern of radial growth from month to month with alternating periods of fast and slow growth. Monthly variations in radial growth of different lobes were synchronized in some but not in all thalli. Few significant correlations were found between the radial growth of individual lobes and total monthly rainfall or shortwave radiation. The levels of ribitol, arabitol and mannitol were measured in individual lobes. All three polyols varied significantly between lobes within a thallus suggesting that variations in algal phostosynthesis and in the partitioning of fungal polyols may contribute to lobe growth variation. The effect on thallus symmetry of lobes which grew radially either consistently faster or slower than average was studied. Slow growing lobes were overgrown, and gaps in the perimeter were eliminated by the growth of neighbouring lobes, in approximately 7 to 9 months. However, a rapidly growing lobe, with its neighbours removed on either side, continued to grow radially at the same rate as rapidly growing control lobes. The results suggested that lobe growth variation results from a combination of factors which may include the origin of the lobes, lobe morphology and the patterns of algal cell division and hyphal elongation in different lobes. No convincing evidence was found to suggest that exchange of carbohydrate occurred between lobes which would tend to equalize their radial growth. Hence, the fluctuating pattern of lobe growth observed may be sufficient to maintain a degree of symmetry in most thalli. In addition, slow growing lobes would tend to be overgrown by faster growing neighbours thus preventing the formation of indentations in the thallus perimeter.

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.

Growth and regeneration of lichen thalli with the central portions artificially removed

The 12-month radial growth of Parmelia conspersa thalli with isidia or with apothecia and isidia was not influenced by removal of the thalli centres. When large thalli had their centres removed and the thallus perimeter was divided into fragments of about 1.0cm in diameter, growth of the fragments was less than the controls, but recovered to near control values after four or five months growth. These results suggest first, that fixed carbon for radial growth may be made in a narrow annulus at the perimeter and second, that there may be little transfer of fixed carbon between the annulus and the centre of the thallus ar around the annulus. Fragments of the centre and the perimeter regenerated growing points, suggesting that fragmentation may be an important method of vegetative reproduction in some lichens.

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.