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 influence of bird droppings on the growth of lichen fragments transplanted to slate and cement substrates

The influence of bird droppings on the growth and fragmentation of five lichen species transplanted to slate and cement substrates was studied over a period of 15 months in South Gwynedd, Wales. The results suggested that at 15 months (1) thallus areas of Parmelia conspersa (Ehrh. Ex Ach.)Ach. were greater on both substrates with the addition of bird droppings with a greater increase on cement; (2) In Parmelia saxatilis (L.)Ach. And Parmelia glabratula ssp. fuliginosa (Fr. ex Duby)Laund., thallus areas were greatest on slate alone and least on cement with bird droppings; (3) in Physcia orbicularis (Neck.)Poetsch, thallus area was significantly reduced on cement alone compared with slate and cement treated with bird droppings; and (4) in Xanthoria parietina (L.)Th.Fr., thallus area was significantly greater on cement with bird droppings compared with slate and cement alone. These responses were attributable to the effect of the substrate and bird droppings on radial growth and the degree of fragmentation of the thalli. The results suggested that nutrient enrichment was more important than the substrate in determining the distribution of P. conspersa and Ph. orbicularis. However, the substrate and bird droppings were important in the remaining species, the data suggesting that P. saxatilis and P. glabratula ssp. fuliginosa would prefer nutrient-poor, siliceous rocks and X. parietina calcareous, nutrient enriched rocks 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 colonization of a slate rock surface by a lichen

Thalli of Parmelia glabratula ssp. fuliginosa on a vertical slate rock surafce in South Gwynedd were signifiacntly larger at the top of the face than at the bottom. The radial growth rates of the thalli measured over 1 year were not correlated with height on the face or with variation in thallus diameter down the face. These results suggest that the lichen thalli may be older at the top than at the bottom are are consistent with the hypothesis that the lichen colonized the top of the face first and then gradually spread downwards.

Diagenetic processes in ore formation with special reference to the Zambian copperbelt and permian marl slate.

This thesis is concerned with the role of diagenesis in forming ore deposits. Two sedimentary 'ore-types' have been examined; the Proterozoic copper-cobalt orebodies of the Konkola Basin on the Zambian Copperbelt, and the Permian Marl Slate of North East England. Facies analysis of the Konkola Basin shows the Ore-Shale to have formed in a subtidal to intertidal environment. A sequence of diagenetic events is outlined from which it is concluded that the sulphide ores are an integral part of the diagenetic process. Sulphur isotope data establish that the sulphides formed as a consequence of the bacterial reduction of sulphate, while the isotopic and geochemical composition of carbonates is shown to reflect changes in the compositions of diagenetic pore fluids. Geochemical studies indicate that the copper and cobalt bearing mineralising fluids probably had different sources. Veins which crosscut the orebodies contain hydrocarbon inclusions, and are shown to be of late diagenetic lateral secretion origin. RbiSr dating indicates that the Ore-Shale was subject to metamorphism at 529 A- 20 myrs. The sedimentology and petrology of the Marl Slate are described. Textural and geochemical studies suggest that much of the pyrite (framboidal) in the Marl Slate formed in an anoxic water column, while euhedral pyrite and base metal sulphides formed within the sediment during early diagenesis. Sulphur isotope data confirm that conditions were almost "ideal" for sulphide formation during Marl Slate deposition, the limiting factors in ore formation being the restricted supply of chalcophile elements. Carbon and oxygen isotope data, along with petrographic observations, indicate that much of the calcite and dolomite occurring in the Marl Slate is primary, and probably formed in isotopic equilibrium. A depositional model is proposed which explains all of the data presented and links the lithological variations with fluctuations in the anoxicioxic boundary layer of the water column.

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.

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.

Factors associated with lobe division in the lichen Parmelia conspersa (Ehrh. ex. Ach.)Ach.

The factors associated with lobe division were studied in thalli of the lichen Parmelia conspersa (Ehrh. ex Ach.)Ach. Lobe division was studied in sequences of adjacent lobes using spatial pattern analysis. In five large thalli, lobe division within the thallus margin was randomly distributed. Correlations between the degree of lobe division, the radial growth of the lobe and lobe morphology were studied in six thalli. Lobe division was positively correlated with either lobe width or area in four thalli. Correlations were observed with radial growth or morphology of the adjacent lobes in two thalli. Dividing and non-dividing lobes were removed from large thalli and glued to pieces of slate with their tips either at the same level or in front of neighbouring lobes. Dividing lobes divided more rapidly when their tips were glued in front of their neighbours. The levels of ribitol, arabitol and mannitol were measured within a 2 mm region of the tip in dividing and non-dividing lobes on four occasions in 1994. Carbohydrate levels were significantly increased in dividing compared with non-dividing lobes. In addition, the mean size of the algal cells was greater in non-dividing compared with dividing lobes especially at the lobe base. However, the percentage of zoosporangia and aplanosporangia did not vary significantly in dividing and non-dividing lobes. These results suggest that: 1) the pattern of lobe division within the thallus margin may be random, 2) lobe division may be determined by lobe size and the location of the lobe tip relative to the neighbouring lobes and 3) there may be an increase in the productivity of lobes associated with lobe division.

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.

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 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.

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.

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.