Field experiments on competition between three foliose lichens using the de Wit experimental design

Competition between three foliose, saxicolous lichens common on slate rock in South Gwynedd, Wales, U.K. was studied experimentally using the de Wit design. Fragments of the three species were cut from the edges of large thalli, glued to 5 x 5 cm plots marked out on pieces of slate which were then placed on boards in the field. For each combination of pairs of species, the two species were grown either in monoculture at a density of 24 fragments per plot or together in three mixtures in differing proportions, i.e. species A:B with 16:8, 12:12 and 8:16 fragments per plot; the density remaining constant throughout. Area of the species in the plots after 3 years was used as an estimate of growth. Physcia orbicularis and Parmelia glabratula ssp. fuliginosa grew similarly in monoculture. In mixtures of the two, growth of each species was linearly related to its proportion in a mixture, suggesting little competition had occurred during three years. By contrast, the growth of Parmelia conspersa in monoculture was significantly greater than that of P. orbicularis or P. glabratula. In addition, the growth of both species was substantially reduced in mixtures with P. conspersa; P. glabratula being eliminated in the mixture in which it was the minority species. These results suggest that P. conspersa should predominate in communities with either of the other two species and, in the absence of P. conspersa, communities dominated by P.orbicularis and P. glabratula should be more stable.

Growth rates of Rhizocarpon geographicum lichens:A review with new data from Iceland

This paper reviews evidence from previous growth-rate studies on lichens of the yellow-green species of Subgenus Rhizocarpon - the family most commonly used in lichenometric dating. New data are presented from Rhizocarpon section Rhizocarpon thalli growing on a moraine in southern Iceland over a period of 4.33yr. Measurements of 38 lichen thalli, between 2001 and 2005, show that diametral growth rate (DGR, mmyr-1) is a function of thallus size. Growth rates increase rapidly in small thalli (<10 mm diameter), remain high (ca. 0.8 mm yr-1) and then decrease gradually in larger thalli (>50 mm diameter). Mean DGR in southern Iceland, between 2001 and 2005, was 0.64 mm yr-1 (SD = 0.24). The resultant growth-rate curve is parabolic and is best described by a third-order polynomial function. The striking similarity between these findings in Iceland and those of Armstrong (1983) in Wales implies that the shape of the growth-rate curve may be characteristic of Rhizocarpon geographicum lichens. The difference between the absolute growth rate in southern Iceland and Wales (ca. 66% faster) is probably a function of climate and micro-environment between the two sites. These findings have implications for previous lichenometric-dating studies, namely, that those studies which assume constant lichen growth rates over many decades are probably unreliable. © British Geological Survey/ Natural Environment Research Council copyright 2006.

Growth curves of four crustose lichens

The growth curves of four common species of crustose lichens, viz., Buellia aethalea (Ach.) Th. Fr., Lecidea tumida Massai., Rhizocarpon geographicum (L.) DC., and Rhizocarpon reductum Th. Fr. were studied at a site in south Gwynedd, north Wales, UK. Radial growth rates (RGR, mm 1.5 yr-1) were greatest in thalli of R. reductum and least in R. geographicum. Variation in RGR between thalli was greater in B. aethalea and L. tumida than in the species of Rhizocarpon. The relationship between growth rate and thallus diameter was not asymptotic; RGR increasing in smaller thalli to a maximum and then declining in larger diameter thalli. A polynomial curve was fitted to the data; the growth curves being fitted best by a second-order (quadratic) curve, the best fit to this model being shown by B. aethalea. A significant linear regression with a negative slope was also fitted to the growth of the larger thalli of each species. The data suggest that the growth curves of the four crustose lichens differ significantly from the asymptotic curves of foliose lichen species. A phase of declining RGR in larger thalli appears to be characteristic of crustose lichens and is consistent with data from lichenometric studies.

Field experiments on the dispersal, establishment and colonization of lichens on a slate rock surface

A sample of run-off water from a vertical, slate rock surface in Wales, U.K. contained abundant fragments of the lichen Parmelia glabratula ssp. fuliginosa from about 0.6 to 8.0 mm in diameter, a few fragments of Parmelia conspersa from 0.6 to 4.0 mm in diameter and a large number of unidentified propagules from 0.2 to 0.5 mm in diameter. The colonization of permanent plots on the rock surface was studied over six years. At the end of the experiment relatively few thalli of Parmelia conspersa, Parmelia glabratula ssp. fuliginosa and Buellia aethalea had established in plots on undisturbed and newly-exposed slate. Fragments (2 mm in diameter) of Parmelia conspersa placed on horizontal pieces of slate survived up to 120 days in cracks, 20 days on a thin smear of bird droppings but only 2-3 days on smooth slate, against small joints in the rock or in small holes. Isidia of Parmelia conspersa placed on horizontal pieces of slate established equally in plots on smooth undisturbed slate and in plots on the surface exposed after the removal of large Parmelia conspersa thalli, but less well on newly-exposed slate. These results suggest that lichen propagules are abundant in run-off water but establishment is a hazardous process. This may be attributable to a shortage of suitable sites on the substratum for attachment of propagules.

The descriptive ecology of saxicolous lichens in an area of South Merionethshire, Wales

Saxicolous lichen vegetation on Ordovician rock at the mouth of the River Dovey, South Merionethshire, is examined in relation to aspect, slope angle, light intensity, rock porosity, rock microtopography and rock stability. A number of characteristic groups of species are recognized. The environmental factors measured are discussed in some detail. In addition, the wide tolerance of most saxicolous species is emphasized.

Seasonal growth and growth rate-colony size relationships in six species of saxicolous lichens

The pattern of seasonal growth and the relation of growth rate to colony size were studied in four foliose and two crustose species of saxicolous lichens. A new method of measuring growth was used whereby the advance of a sample of lobes along millimetres marked on the substrate was measured under a magnification of x10. Three peaks of growth were found(in March, June and November) for the foliose species and a single peak (in May to August) for the crustose species. THe peaks of growth corresponded approximately to peaks of rainfall. Growth rate in relation to increasing colony size fell in a smooth exponential curve when expressed on a cm squared/ cm squared/ unit time basis. The result is consistent with a linear radial rate for most of the thallus sizes for the six species. There is also evidence for an exponential incresae in growth rate initially until about 1.5 cm thallus diameter in two of the sepcies when the linear radial rate is achieved.

Are metal ions accumulated by saxicolous lichens growing in a rural environment?

The levels of Mn, Cu, zn, Mg and Ca were measured in the lichens Xanthoparmelia conspersa (Ach.) Hale and Parmelia glabratula ssp. fuliginosa (Fr. ex Duby): Laund. growing on a steep slate rock surface in south Gwynedd, Wales, UK. The objective was to test the following hypotheses: 1) that foliose lichens growing in a rural environment concentrate metal ions relative to the substratum, 2) that the concentration of metal ions increases significantly with thallus size and 3) that individual ions accumulate preferentially either in the marginal lobes or thallus centre. Mg and Ca were present in rainfall whereas all ions were present in rock surface runoff and in the substratum. Levels of Mn, Mg and Ca were increased in runoff collected from the bottom compared with the top of the rock surface. In P. glabratula ssp. fuliginosa thalli, ions were present as follows, Mg > Ca=Mn=Zn>Cu, and there were no significant differences in thalli collected from the top and bottom of the rock surface. With the exception of Mg and Ca, ion levels in thalli were similar to or less than those in the substratum. The levels of Ca, Cu and Zn were similar in thalli from under 1 to over 4 cm in diameter. However, the level of Mg increased with thallus size in X. conspersa while the level of Mn decreased with thallus size in both species. Ion levels were similar in the marginal lobes and centres of large diameter (over 4 cm) and small diameter (under 2 cm) X. conspersa thalli. With the exception of Mg, there was no evidence for a significant accumulation of ions relative to the substratum or with thallus size. In addition, there was no evidence to suggest preferential accumulation of ions either in the thallus centre or marginal lobes.

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.

Do the lichens Xanthoparmelia conspersa (Ach.) Hale and Rhizocarpon Ram. Em. Th. Fr. subgenus Rhizocarpon utilise exogenous carbohydrates for radial growth?

To field test the hypothesis that lichen thalli can use environmental sources of carbon, solutions of ribitol, arabitol and mannitol were added to intact thalli of Xanthoparmelia conspersa (Ach.) Hale and a yellow species of Rhizocarpon (Rhizocarpon Ram. Em. Th. Fr. subgenus Rhizocarpon). In addition, ribitol and an arabitol/mannitol mixture were added to the marginal hypothalli of Rhizocarpon thalli after removal of the areolae. Carbohydrates were added at the beginning of 2- or 3-month growth periods for up to 15 months at concentrations approximately three times the levels estimated to be in the thalli. Addition of carbohydrates to intact thalli of both species had no effect on total radial growth but addition of mannitol significantly increased growth of X. conspersa thalli in the September/October growth period in one experiment. However, this effect was not repeated in a subsequent experiment in which different concentrations of mannitol were added to intact thalli. Addition of ribitol to hypothalli of Rhizocarpon resulted in significantly increased growth in the first few months of the experiment, growth then declining to levels below that of untreated thalli. The data suggest that although hypothalli of Rhizocarpon may have the ability to utilise exogenous carbohydrates for growth, there was little evidence that intact thalli of either species utilise environmental sources of carbon in the field.

The growth of six saxicolous lichens transplanted to lime-rich and lime-poor substrates in south Gwynedd, Wales

The objectives of this study were to investigate: (1) whether foliose lichen thalli could be transplanted from one substrate to another and (2) whether such transplants could be used to study the influence of the substrate on growth. Hence, six saxicolous lichens, with contrasting distributions on lime-rich and lime-poor substrates in South Gwynedd, Wales, were transplanted onto slate, granite, asbestos and cement. Fragments of the perimeters of thalli were glued to the different substrates using Bostic adhesive. Parmelia conspersa (Ehrh. Ex Ach.)Ach. and Parmelia saxatilis (L.)Ach., fragments increased in area over 15 months on slate and granite but decreased in area or did not survive on asbestos and cement. Fragments of Xanthoria parietina (L.)Th.Fr. and Physcia tenella (Scop.)DC. em Bitt. did not survive on slate and granite while some fragments survived but grew poorly on asbestos and cement. Parmelia glabratula ssp. fuliginosa (Fr. ex Duby)Laund. fragments decreased in area on all substrates and especially on cement and asbestos while Physcia orbicularis (Neck.)Poetsch fragments increased in area on granite and cement, decreased on asbestos and did not change significantly on slate. The results suggested that the distribution of P. conspersa and P. saxatilis was determined primarily by physico-chemical properties of the substrate. By contrast, P. glabratula ssp. fuliginosa may have responded to the transplant procedure while X. parietina, Ph. tenella and Ph. orbicularis may require nutrient enrichment to grow successfully on a substrate.

Seasonal growth of foliose lichens in successive years in South Gwynedd, Wales

The aim of this study was to test the hypothesis that differences in the pattern of seasonal growth in foliose lichens from year to year were determined by yearly differences in the distribution of rainfall, shortwave radiation and temperature. Hence, the radial growth of Parmelia conspersa (Ehrh. Ex Ach.) Ach. , P. glabratula ssp. fuliginosa (Fr. ex Duby) Laund. and Physcia orbicularis (Neck) Poetsch. was studied on slate fragments over 34 successive months in an area of South Gwynedd, Wales. U.K. Similarities and differences were observed in the pattern of seasonal growth in the three species. Periods of maximum growth of a species occurred in different seasons in successive years. Correlation and multiple regression analysis suggested that total rainfall per month was the most important climatic variable positively correlated with monthly growth. Significant positive correlations were found in some growth periods with number of raindays per month, average wind speed and maximum and minimum temperature. Total number of sunshine hours per month and the frequency of ground frosts were negatively correlated with monthly growth in some growth periods. For each species, monthly radial growth was correlated with different climatic variables in each growth period. Hence, the results support the hypothesis in that periods of maximum growth can occur in any season in South Gwynedd and depend on (1) the distribution of periods of high total rainfall and (2) whether or not these periods coincide with periods of maximum sunlight.

Competitive interactions between four foliose lichens on north and south facing rock surfaces

Competition between four foliose lichen species which have distinct aspect distributions on slate rock in South Gwynedd, Wales, U.K. was studied in the field using a factorial experimental design. The lichens were grown as fragments glued to pieces of slate in monoculture and in two-, three- and four-species mixtures. The pieces of slate were placed to face a northerly or southerly direction. Growth in area (mm2) was used as a measure of performance in the experiment. The growth in area of Parmelia conspersa in south facing plots was not reduced in the presence of any of its competitors but its growth was reduced in the presence of Parmelia saxatilis in north facing plots. The growth of Parmelia glabratula ssp. fuliginosa was reduced in the presence of P. conspersa and P. saxatilis in south and north facing plots. Physcia orbicularis was reduced by P. conspersa in south facing plots and by both P. glabratula ssp. fuliginosa and P. saxatilis in north facing plots. The growth of P. saxatilis was increased by P. glabratula ssp. fuliginosa in south facing plots but was not reduced by any of its competitors in north facing plots. Significant two and three factor interactions suggested that the results from the three- and four-species mixtures were not always predictable from the results of the two-species mixtures. The results of the experiment may help to explain the existing aspect distribution of the four species on slate rock in South Gwynedd.

The influence of calcium and magnesium on the growth of the lichens Parmelia saxatilis and Xanthoria parietina on slate substrates

In South Gwynedd, Wales, U.K., the calcicole lichen Xanthoria parietina occurs not only on alkaline substrates at inland sites but also on siliceous rock at coastal martimie sites while the calcifuge species Parmelia saxatilis occurs only at inland sites and on slate rocks. Samples of maritime and inland slate did not differ significantly in their calcium or magnesium content. Thalli of X. parietina on pieces of slate did not survive when transplanted from maritime rocks to a site inland. Thalli of maritime X. parietina and P. saxatilis on slate were then transplanted to a site inland and were treated at intervals during 1 year either with calcium carbonate applied as a thick paste or a 0.25 mM solution of calcium chloride. Treatment of X. parietina with calcium carbonate enabled the thalli to survive and grow. However, addition of calcium carbonate to P. saxatilis resulted in low growth rates and fragmentation of the centres of the thalli. The calcium chloride solution had no statistically significant effects on the growth of either species. In addition, thalli of both species were treated with calcium or magnesium carbonates or wetted with an alkaline buffer at intervals over 12-14 months. Thalli of X. parietina survived and grew rapidly when treated with either carbonate but the growth of the buffer-treated thalli gradually declined over the experimental period. Thalli of P. saxatilis fragmented and disappeared after 8-10 months after treatment with either carbonate but normal growth occurred in the buffer treatment. Xanthoria parietina may occur on siliceous maritime rocks at the site because of the presence of calcium or magnesium in sea spray combined with the spray’s alkaline pH. By contrast, P. saxatilis may be confined to siliceous rocks inland because the thalli grow poorly in the presence of calcium and magnesium.

The influence of bird droppings and uric acid on the radial growth of five species of saxicolous lichens

Bird droppings were applied over 1 year as a thick paste and as a suspension in deionized water to five species of lichens with different distributions on and off bird perching stones. The paste and suspension increased the radial growth of Parmelia conspersa while the paste increased the growth of Xanthoria parietina and reduced the growth and caused loss of colour in Parmelia glabratula ssp. fuliginosa. There were no statistically significant effects of paste or suspension on the growth of Physcia grisea or Parmelia saxatilis. In P. conspersa and X. parietina the growth responses were similar through the year but in P. glabratula the inhibitory effect of the paste was significant after 8 months growth. Application of a suspension of uric acid over 1 year had no statistically significant effects on the growth of P. conspersa, P. glabratula or X. parietina and was unlikely to be responsible for the effects of bird droppings on growth. The growth responses of the five species agreed well with their distributions in the field.

Competition between three saxicolous species of Parmelia (lichens)

Fragments of three foliose, saxicolous lichens were glued in 5 x 5 cm plots on pieces of slate in the field, either in monoculture or paired in 1:1 mixtures with each of the other two species. A preliminary experiment suggested that glueing did not influence the radial growth of the lichen fragments. No species eliminated another after 3 years but the growth (total area in sq mm) of Parmelia saxatils and P. glabratula ssp. fuliginosa was reduced significantly in mixtures with P. conspersa; the growth of P. glabratula ssp. fuliginosa was reduced significantly in the mixture with P. saxatilis compared with their growth in monoculture. The results suggest that the three lichens show interference by competition for space and light in the following order of competitive ability: P. conspersa > P. saxatilis > P. glabratula ssp. fuliginosa. A high radial growth rate and the ability to overgrow a thallus may be important competitive attributes in foliose lichens and the results also suggest that competition can reduce the abundance of a lichen and lead to distribution patterns in the field.