Water Storage in the Lichen Family Umbilicariaceae

Quantitative relationships between thallus structure and water storage and retention capacities In 12 species of the lichen family Umbilicariaceae were explored using three recent techniques for plant structure analysis: stereology (3D quantification of microscopic Images), mercury Intrusion porosimetry (determination of pore size distribution of tissues) and low-temperature scanning electron microscopy (LTSEM). Water storage capacity of the thallus was related neither to thallus density nor surface area of the thallus; It was directly related to the total porosity of the thallus and inversely related to the proportion of thallus volume occupied by cell walls and gelatinous substances. Water retention capacity increased with increasing thallus density and was decreased by slight increases in the surface area of the upper side of the thallus. Water storage and retention capacities exhibited a positive correlation only when the storage capacity was expressed on a surface area basis. LTSEM study of fully hydrated specimens revealed that many intercellular spaces of the upper cortex and upper parts of the algal layer contained liquid water. Intercellular spaces of the lower part of the algal layer and medulla were in general either airfilled or partially occupied by gelatinous substances. Species with rhizinomorphs and substrate-hygrophytic (water uptake from surface run-offs) stored more water and retained it longer than aerohygrophytic species (water uptake from the atmosphere) lacking rhizinomorphs. Thallus structure of aerohygrophytic species seems to facilitate rapid gas exchange with the environment, improving water uptake and carbon gain when atmospheric moisture is available but accelerating dehydration when the atmosphere becomes dry.     

CYANOPHYTE CEPHALODIA IN THE LICHEN GENUS NEPHROMA

The lichens, Nephroma expallidum (Nyl.) Nyl. and N. arcticum (L.) Torss., consistently have at least two symbionts in a single thallus: a green alga in the algal layer and a blue-green alga in the internal cephalodia. The cephalodia originate from algal cells in contact with the lower surface of the lichen, in the zone of rhizine formation. The rhizines surround the epiphytic algal colony and form a second cortical layer; following dissociation of the original lower cortex, further growth of the two organisms results in the cyanophyte colony being enveloped by a compact layer of fungal tissue and positioned in the lichen medulla. The colony may eventually assume a superior or inferior position in relation to the lichen thallus, depending in part on the lichen species. Nephroma anticum may have two distinct morphological forms of blue-green algae in the same thallus and occasionally in the same cephalodium. It appears that the relationship that exists between the cephalodial algae and the lichen thallus is antagonistic and results, in some cases, in the exclusion of the green algal layer and death to the cephalodial cyanophytes.     

Externally held water – a key factor for hair lichens in boreal forest canopies

Lichens hold water inside (internal pool) and outside their body (external pool). Yet, external pool size is not known in hair lichens dominating boreal forest canopies. Here we quantify morphological traits and internal/external water in two widespread Bryoria species along Picea abies canopy-height gradients: Bryoria fuscescens at 5–20 m and Bryoria capillaris at 15–20 m. Dry mass and specific thallus mass (STM) of intact B. fuscescens increased with height, while STM of individual branches did not. Maximum water holding capacity (mg H₂O cm⁻²) increased with height, but did not differ between the species. Bryoria had much larger external (79–84% of total) than internal water pools, trapping water by dense clusters of thin, overlapping branches. They thus increase water storage in boreal forest canopies and influence hydrology. High external water storage extends hydration periods and improves lichen performance in upper canopies, and thereby contributes to the success of these hair lichens.     

Do secondary substances in the thallus of a lichen promote CO2 diffusion and prevent depression of net photosynthesis at high water content?

Many lichens show seriously depressed net photosynthesis (NP) at high thallus water contents due to increased carbon dioxide diffusion resistance through blockage of diffusion pathways by water. The soil lichen Diploschistes muscorum, however, shows no depression and NP is close to maximal even at the highest thallus water content. We investigated whether lichen substances (lecanoric and diploschistesic acids) in the cortex and medulla contributed to this ability to maintain high NP. Dry thalli were extracted with water-free acetone and, after this treatment, were found to be fully viable to the extent of continued growth after replanting in the field. No differences were found in the response of NP to thallus water content between the normal and extracted thalli, in fact the response curves were often nearly identical. Thus, in this species it seems that lichen substances did not maintain the water-free diffusion pathways and some other explanation, possibly structural, needs to be sought. Received: 5 April 1997 / Accepted: 26 April 1997     

ERUMPENT CEPHALODIA,AN APPARENT CASE OF PHYCOBIAL INFLUENCE ON LICHEN MORPHOLOGY1,2,3

On the upper surface of some specimens of the lichen Lobaria cf. crosa (Eschw.) Nyl. there are shrub-like growths which have a morphology similar to that of a free-living fruticose lichen, Polychidium umhausense (Auersw.) Henss., and contain a cyanophyte phycobiont, Nostoc. A few growths also contain scattered colonies of a chlorophyte phycobiont, in which case the lichen tissue locally assumes the foliose form characteristic of the parent Lobaria thallus. The differentiation of dorsiventral lichen cortices and the formation of a lax medulla and distinct algal layer are correlated with the presence of the green phycobiont. The lichen substances atranorin, gyrophoric acid, and 4-0-methylgyrophoric acid occur in both, the foliose L. erosa thallus and the fruticose tissue. It is suggested that the fruticose structures are erumpent cephalodia which are derived from the outgrowth of internal, cephalodia and should not be considered, to be epiphytic colonies of the lichen Polychidium.     

The significance of thallus size for the water economy of the cyanobacterial old-forest lichen Degelia plumbea

Rosette-formed, circular thalli of Degelia plumbea were studied in the laboratory. Regardless of thallus size, the optimal quantum yield of photosystem II (F _V/F _M) remained at a high, constant level during a drying cycle starting with fully hydrated thalli until the thallus water content reached about 200%. Net photosynthesis reached a maximum level at this hydration level. Thereafter, both F _V/F _M and net photosynthesis fell rapidly to zero at a water content of somewhat less than 100%. There was a highly significant, positive relationship between thallus size and the water-holding capacity, as well as a strong, negative correlation between size and water loss per thallus area. Consequently, an increase in thallus size from 1 to 36 cm² lead to a tenfold prolongation of the photosynthetically active period during a drying cycle at a low radiation regime. The improved water-holding capacity in larger thalli is mainly a result of a thicker hypothallus. The fast desiccation of small thalli suggests that the regeneration of D. plumbea could be severely hampered by nearby logging that raises the evaporative demand by increasing radiation loads and wind exposure at remaining lichen sites. Received: 12 December 1997 / Accepted: 20 March 1998     

DISPERSAL PATTERNS OF TERRICOLOUS LICHENS BY THALLUS FRAGMENTS

Dispersal patterns from seven terricolous lichen species, with a high capacity for asexual reproduction by fragmentation (Cetraria muricata, Cladonia species), were studied in differing vegetation types in north-eastern Germany. Marked lichen thalli were fragmented by trampling. After 15 days the spread of thallus fragments was monitored. Whereas most of the fragments that were dispersed by wind remained within a 20-cm radius from the source, the maximal dispersal distance was 57 cm in a dry sand grassland and 68 cm in an open pioneer pine forest. Dispersal was negligible in a closed old-growth pine forest. Several fragmented lichen cushions were disturbed and removed by animals, and led to a maximal dispersal distance of 9·70 m. These results suggest that: (a) thallus fragments provide good short-distance dispersal in open vegetation, but are inefficient for long-distance dispersal, and (b) wind and animals are important factors for the dispersal of thallus fragments. For restoration management of man-made substrata, artificial introduction of lichen thalli is proposed.     

DIATOMS LIVING INSIDE THE THALLUS OF THE GREEN ALGAL LICHEN COENOGONIUM LINKII IN NEOTROPICAL LOWLAND RAIN FORESTS1

Coenogonium linkii Ehrenb. is a very common filamentous lichen, growing on stems, hanging roots, and lianas in the understory of neotropical lowland rain forests. We investigated several thalli of this species from locations in Panama and French Guiana. All thalli were inhabited by various species of terrestrial diatoms, which were found between the thallus filaments on extracellular material of the mycobiont. We identified 18 species of diatoms belonging to nine genera: Diadesmis, Eunotia, Hantzschia, Luticola, Melosira, Nitzschia, Orthoseira, Pinnularia, and Stauroneis. The potential benefits both diatoms and lichens could derive from symbiosis in relation to water, irradiance, and nitrogen availability are discussed.     

Multiple-scale environmental modulation of lichen reproduction

It is necessary to understand how environmental changes affect plant fitness to predict survival of a species, but this knowledge is scarce for lichens and complicated by their formation of sexual and asexual reproductive structures. Are the presence and number of reproductive structures in Lobaria pulmonaria, a threatened lichen, dependent on thallus size, and is their formation sequential? Does any size-dependence and sequential formation vary along a climate gradient? Generalized linear mixed models were used to explore the effect of environmental predictors on the size and presence/abundance of each reproductive structure and to determine the probability of a given-sized thallus to develop any reproductive structure. The largest individuals are more likely to develop reproductive structures, and the lichen uses a mixed strategy of early asexual reproduction and late sexual. Macro and microclimatic variables also influenced reproductive capacity. Relationships among climate conditions and lichen size and reproductive capacity can compromise the future viability of the species in the most southern populations of Europe.     

Growth and vitality of epiphytic lichens

 We tested the hypothesis that changed microclimate at induced forest edges causes reduced growth of epiphytic lichens. Two foliose, green algal lichens were transplanted to the lower canopy of a mature Picea abies forest at six distances (2, 6.25, 12.5, 25, 50 and 100 m) from a clearcut. The biomass growth in Platismatia glauca (6.2% in 16 months) was 41% higher than in Lobaria pulmonaria (4.4%). We found no growth reduction near the forest edge. In contrast, the highest growth in both species occurred within 12 m from the edge. Further, fluorescence and chlorophyll measurements showed that lichen vitality was unaffected by distance from edge. The light intensity was 4.3 times higher at the edge than in the interior during the growing season, but there were only minor differences in air temperature and relative humidity. Monitoring of thallus water content revealed clear differences in both number and length of wetting and drying cycles. However, the total time with water content sufficient for photosynthetic activity was only slightly higher at the edge. The data thus indicate that our gradient in microclimate was too small to significantly affect lichen growth, and that lichens are largely metabolically inactive when large edge-interior contrasts in microclimate occur. Lichen response to forest edge microclimate results from intricate interactions among several biotic and abiotic factors. Linking data on lichen growth, microclimate and thallus water content with physiological measurements provides a framework for future studies of the mechanisms behind abiotic edge effects. Received: 15 April 1996 / Accepted: 21 June 1996     

A NEW FOSSIL ALGA FROM THE SILURIAN OF MICHIGAN

The record of Paleozoic algal diversity is strongly biased toward calcareous and stromatolitic types due to the general paucity of well preserved compression fossils of other taxa. A new genus, Thalassocystis (type species T. striata) is described on the basis of compression fossils in middle Silurian dolomite from the northern peninsula of Michigan. The plant is characterized by a short branched thallus, each branch terminated by an enlarged bladder-like structure. The plants were deposited in a tropical, shallow water marine environment but whether they were benthic, adventive to the pelagic zone, or completely pelagic cannot be determined from the evidence available.     

Copper toxicity to five Parmelia lichens in vitro

Treatment of Parmelia caperata, P. perlata, P. subrudecta, P. sulcata and P. tiliacea with CuSO₄ resulted in a time- and copper-concentration-dependent decrease in the total and intracellular potassium concentrations of the thallus, indicating that copper damaged the cytoplasmic membrane. Treatment with copper also resulted in a time-dependent increase in the total copper concentration of the thallus. After 4 h of exposure to copper, the process of potassium efflux was essentially completed but the absorption of copper was still increasing; moreover, the amount of copper bound to the thallus exceeded twice the amount of potassium released from the thallus, suggesting that cupric ions reached intracellular sites in the thallus, and K⁺/Cu²⁺ exchange was not electroneutral. After 5 h of exposure to copper, the extent of decrease in the total and intracellular potassium concentrations of the thallus was positively correlated with copper absorption levels, but only at 0.05<P<0.10, suggesting that membrane damage was proportional to the amount of bound copper, but other factors could have been operative, namely binding of copper to the cell wall. Acetone extracts of untreated thalli contained low concentrations of amino acids, polyols, and sugars, but considerable amounts of lichen substances: atranorin, caperatic, constictic, lecanoric, menegazziaic, protocetraric, salazinic, stictic, and usnic acids. Titration of the extracts with copper and assay of the free Cu²⁺ concentration revealed the presence of copper-binding ligands, and several successive absorption cycles, most probably corresponding to the binding of Cu²⁺ to each of the lichen substances detected in the extracts. However, no significant correlation (P>0.10) was found between the Cu²⁺-complexing capacity of acetone extracts and copper-induced membrane damage. It was concluded that in the studied Parmelia species, and in the experimental conditions used in this work, copper toxicity was not a simple function of the Cu²⁺-binding properties of the lichen substances present in the thallus. Several hypotheses were formulated to interpret the results.     

GROWTH AND DEVELOPMENT OF THE RETICULATE THALLUS IN THE LICHEN RAMALINA MENZIESII

The thallus of the lichen Ramalina menziesii Tayl. is comprised of subunits which resemble planar nets. The nets arise upon older nets as buds which develop perforations and expand. Microphotographic study of thallus development in the field shows several features which differ from a recently proposed model of development for this species. New perforations develop continually in new tissue produced at the apical margin of the net. The apical margin is inrolled, and may furcate. This type of branching is developmentally distinct from the formation of new nets from buds borne upon expanded parts of the net. Anatomical study of net apices appears to support assertions that programmed tissue necrosis contributes to perforation formation at sites where gaps are present in the algal layer.     

Moisture content and CO2 exchange of lichens

Summary Net photosynthesis (10 klx light intensity, 150 E m^-2 s^-1 PAR) and dark respiration of the lichen Ramalina maciformis at different temperatures are measured in relation to thallus water content. Both first increase with increasing hydration. Dark respiration then remains constant with increased water content until thallus saturation. In contrast, a further increase in water content leads to a depression of net photosynthesis, as shown in previous studies, after a maximum of CO₂ uptake has been attained. However, the extent of this depression depends strongly on temperature. In saturated thalli (160% water content in relation to lichen dry weight) the depression amounts to about 15% and 63% of the maximum unsaturated rate at 5°C and 25°C thallus temperature, respectively. The moisture compensation-point of net photosynthesis is also decisively determined by temperature (for 0°C at 20% water content; for 25°C at 15%), and the water content that allows maximum rates of CO₂ uptake (for 0°C at 80%; for 25°C at less than 40% water content). An electrical analogue of CO₂ exchange in a lichen thallus is presented, and it is suggested that the experimental results may be interpreted in terms of temperature-dependent CO₂ diffusion resistances in imbibed lichen thalli.     

Carbon dioxide exchange in Cladina lichens from subarctic and temperate habitats

Summary The survival potential of lichens in a given habitat is determined by the response of CO₂ exchange to photosynthetically active radiation (PhAR), thallus temperature, and thallus relative water content (RWC). Therefore morphologically similar lichens from contrasting climatic environments 1) should differ in their CO₂ exchange responses, and 2) these differences should reflect adaptations to their climatic regimes. The CO₂ exchange responses of a subarctic (55°N, 67°W) Cladina stellaris (Opiz) Brodo population and a temperate (29°N, 82°W) Cladina evansii (Abb.) Hale and W. Culb, population were used to test these two related hypotheses.Infrared gas analysis with lichens collected in September–October 1975 established that the two populations differed in their responses to incident PhAR, thallus temperature, and thallus RWC. Net photosynthesis in C. stellaris had an optimum at a lower temperature and a greater relative photosynthetic capacity at low temperatures than did C. evansii. Cladina evansii maintained net photosynthesis above 35°C thallus temperature; C. stellaris did not. In both species the optimum temperature for net photosynthesis increased with increasing irradiance. The C. stellaris light saturation point was consistently lower than that of C. evansii. Both species had maximal rates of net photosynthesis at 70–80% relative water content. In C. evansii the CO₂ exchange rates, expressed as percentages of the maximum rate, declined more rapidly under suboptimal conditions. The absolute CO₂ exchange rates of C. evansii were greater than those of C. stellaris. At 20°C and 90–95% RWC, resaturation respiration occurred in both species and continued until 6–7 h after wetting.Contrasts in the temporal patterns of thallus condition at each collection site suggest that not all differences in the two response surfaces reflect climatic adaptation. The two populations appear well adapted to incident PhAR and thallus temperature regimes but the 70–80% RWC optimum for net photosynthesis common to both species is puzzling since their water regimes differ markedly. The overall adaptedness of the CO₂ exchange responses in the two species cannot be judged without a comprehensive quantitative analysis of carbon balance under differing climatic regimes.     

Escape from the cryptic species trap: lichen evolution on both sides of a cyanobacterial acquisition event

Large, architecturally complex lichen symbioses arose only a few times in evolution, increasing thallus size by orders of magnitude over those from which they evolved. The innovations that enabled symbiotic assemblages to acquire and maintain large sizes are unknown. We mapped morphometric data against an eight‐locus fungal phylogeny across one of the best‐sampled thallus size transition events, the origins of the Placopsis lichen symbiosis, and used a phylogenetic comparative framework to explore the role of nitrogen‐fixing cyanobacteria in size differences. Thallus thickness increased by >150% and fruiting body core volume increased ninefold on average after acquisition of cyanobacteria. Volume of cyanobacteria‐containing structures (cephalodia), once acquired, correlates with thallus thickness in both phylogenetic generalized least squares and phylogenetic generalized linear mixed‐effects analyses. Our results suggest that the availability of nitrogen is an important factor in the formation of large thalli. Cyanobacterial symbiosis appears to have enabled lichens to overcome size constraints in oligotrophic environments such as acidic, rain‐washed rock surfaces. In the case of the Placopsis fungal symbiont, this has led to an adaptive radiation of more than 60 recognized species from related crustose members of the genus Trapelia. Our data suggest that precyanobacterial symbiotic lineages were constrained to forming a narrow range of phenotypes, so‐called cryptic species, leading systematists until now to recognize only six of the 13 species clusters we identified in Trapelia.     

The Physiology of the Lichen Peltigera aphthosa, with Special Reference to the Blue-green Phycobiont (Nostoc sp.)

Variations in the morphology and physiology were noted when parts of the Peltigera aphthosa Willd. thallus differing in age were examined. The many small cephalodia on the growing apex of the lichen showed a lower heterocyst frequency (14%) than those on the rest of the thallus (21–22%), which was reflected in the nitrogenase activity. In contrast, highest levels of photosynthesis (¹⁴CO₂ uptake and O₂ evolution) were noted at the growing apex. while respiration rates were fairly stable over the thallus. The water-holding capacity was greatest in the midparts. Cephalodial biomass represented an average of 2.6% of total thallus biomass. while the number and size of these structures varied considerably. A minor part of the total carbon fixation (¹⁴CO₂) and net oxygen evolution (O₂ electrode) was performed by the blue-green phycobiont (Nostoc) in light. A rapid excretion of ammonia from isolated cephalodia was noticed, which together with a comparatively constant C:N ratio throughout the thallus indicated a rapid transport of metabolites facilitated by close physical contact (electron microscopy).     

Lichenizing Rhizomorphs and Thallus Development in the Squamulose Lichen Aspicilia crespiana Rico ined. (Lecanorales,Ascomycetes)

This work describes the rhizomorphs and their relation to thallus development in the squamulose lichen Aspicilia crespiana Rico ined. The rhizomorphs capture algal cells, giving rise to new squamules terminally or laterally. The thallus thus consists of a network of lichenized squamules interlinked by mycobiontic rhizomorphs. A previously proposed comparison of lichen rhizomorphs to the prothallus (hypothallus) of crustose lichens is applied and expanded upon. Other less organized prothallic structures sometimes produced by A. crespiana are also described. The importance of lichenizing rhizomorphs as an essential feature of thallus growth in this species is emphasized and a competitive role in substrate occupation and thallus expansion is suggested.     

Die parasitische FlechteLecidea insidiosa und ihre Biologie

Lecidea insidiosa, up to now regarded as a lichenicolous fungus without thallus, is a parasitic lichen, growing exclusively onLecanora varia. It kills the plectenchyma of the host, but presumably takes over part of the host-algae to build its own thallus, first inside, then outside of the host thallus. Host and parasite thallus are chemically different.Lecidea insidiosa is not related toLecidea vitellinaria, even though the two were thought to be synonymous for a long time. The species is reported for the first time from the Alps (Steiermark). It is the first known parasitic lichen to occur on a normally wood inhabiting host.

     

Caloplaca erodens [sect. Pyrenodesmia], a new lichen species from Italy with an unusual thallus type

Caloplaca erodens is a new species of sect. Pyrenodesmia, characterised by an orbicular, sorediate, bluish-grey thallus which is endolithic but emerges at the periphery with a white, K−, obscurely lobate prothallus. The species is frequent on calcareous outcrops and walls of isolated churches and ruins of the Central Apennines (Sibillini, Gran Sasso), where it may occur in large monospecific populations, from 1000 to 2500 m asl., and is also known from dry sites of the southern Alps. It has been found with apothecia only in the type locality, being predominantly sterile. The reproduction of this lichen is evidently linked to the release of fragments of clusters of photobiont cells and mycobiont hyphae which are continuously exposed with the dissolution of the substratum. Readily distinguished from the apparently similar endolithic C. alociza (which is characterised by numerous apothecia, black, K+ purple prothallus, and esorediate thallus), C. erodens probably belongs to the C. circumalbata complex, whose taxa are always epilithic but have a white, K− prothallus. The phylogenetic position of the new species within sect. Pyrenodesmia as inferred by ITS sequences of the nuclear ribosomal DNA is shortly discussed.