Heterogeneity of Chlorophyll Fluorescence over Thalli of Several Foliose Macrolichens Exposed to Adverse Environmental Factors: Interspecific Differences as Related to Thallus Hydration and High Irradiance

Spatial heterogeneity of chlorophyll (Chl) fluorescence over thalli of three foliose lichen species was studied using Chl fluorescence imaging (CFI) and slow Chl fluorescence kinetics supplemented with quenching analysis. CFI values indicated species-specific differences in location of the most physiologically active zones within fully hydrated thalli: marginal thallus parts (Hypogymnia physodes), central part and close-to-umbilicus spots (Lasallia pustulata), and irregulary-distributed zones within thallus (Umbilicaria hirsuta). During gradual desiccation of lichen thalli, decrease in Chl fluorescence parameters (F_O - minimum Chl fluorescence at point O, F_P - maximum Chl fluorescence at P point, ₂ - effective quantum yield of photochemical energy conversion in photosystem 2) was observed. Under severe desiccation (>85 % of water saturation deficit), substantial thalli parts lost their apparent physiological activity and the resting parts exhibited only a small Chl fluorescence. Distribution of these active patches was identical with the most active areas found under full hydration. Thus spatial heterogeneity of Chl fluorescence in foliose lichens may reflect location of growth zones (pseudomeristems) within thalli and adjacent newly produced biomass. When exposed to high irradiance, fully-hydrated thalli of L. pustulata and U. hirsuta showed either an increase or no change in F_O, and a decrease in F_P. Distribution of Chl fluorescence after the high irradiance treatment, however, remained the same as before the treatment. After 60 min of recovery in the dark, F_O and F_P did not recover to initial values, which may indicate that the lichen used underwent a photoinhibition. The CFI method is an effective tool in assessing spatial heterogeneity of physiological activity over lichen thalli exposed to a variety of environmental factors. It may be also used to select a representative area at a lichen thallus before application of single-spot fluorometric techniques in lichens.     

In situ development of the foliicolous lichen Phyllophiale (Trichotheliaceae) from propagule germination to propagule production

The vegetative cycle of the foliicolous lichen Phyllophiale, from propagule germination to propagule production, was studied by light microscope observation of thalli colonizing plastic cover slips placed within a lowland tropical forest. Discoid propagules germinated by growth of radially arranged fungal cells and developed directly into lichen thalli. The young lichen comprised a single disc of closely branched, radiating filaments of the algal symbiont Phycopeltis, covered by a network of fungal hyphae extending onto the substrate as a prothallus. The prothallic hyphae incorporated additional Phycopeltis thalli encountered on the substrate. The phycobiont formed a single layer, with individual algal thalli clearly distinguishable within the lichen. Radial growth ceased at points of contact between adjacent phycobiont thalli. The visible shape of the crustose lichen thallus corresponded to the perimeter of the phycobiont thalli within. Propagules were initiated at points corresponding to the margins of the phycobiont thalli, by vertical reorientation of horizontal algal filaments surrounded by fungal hyphae. The lichenized alga produced intercalary gametangia. Degeneration of propagules unsuccessful in lichen establishment sometimes resulted in free growth of the phycobiont. The alga generally maintained its shape, growth pattern, and reproductive independence within the lichen, while also participating in the formation of unique symbiotic propagules.     

CHANGES IN WATER ECONOMY IN RELATION TO ANATOMICAL AND MORPHOLOGICAL CHARACTERISTICS DURING THALLUS DEVELOPMENT INPARMELIA ACETABULUM

The influence on uptake and water loss of the structural changes experienced by Parmelia acetabulum during thallus development were investigated. Small specimens were characterized by flat lobes and a thin thallus and cortex. Large specimens appeared strongly rugose, imbricate and irregularly folded, and had a significantly thicker cortex and medulla than small thalli. Maximum water storage capacity did not differ between large and small thalli, although water retention was much higher in large thalli, presumably due to the interaction of structural characteristics and a higher boundary layer resistance. This translated into a longer duration of the period of thallus hydration in large thalli compared to small thalli. Incubation of thalli in water-vapour-saturated atmospheres induced full recovery of photosynthetic electron transport to the values before desiccation in small thalli but only induced a partial recovery in large thalli. The close correlation between photosynthetic electron transport and net photosynthesis during desiccation found in this species suggested that carbon-fixation activity could be regained to a larger extent by incubation of thalli in water vapour in small compared to large thalli. The higher ability for water vapour uptake of small thalli might allow them to efficiently use small amounts of intermittently available water or periods of high relative humidity. The significance of this differential ability to utilize water is discussed with regard to the known ecological preferences of the species.     

GROWTH FROM THE CENTRE IN AN UMBILICATE LICHEN

The pustules on the surface of the umbilicate lichenLasallia pustulataare moved away from the centre and towards the thallus margins with an average speed of 1·8 mm per year, revealing a growth pattern organized radially and flowing from the centre of the thallus. Pustules close to the margin move substantially faster than those close to the centre; for each 10 mm added to the initial distance from the centre, the speed of outward movement is about doubled. This suggests that intercalary growth over the entire thallus is displaced outwards in an accumulating manner, accelerating the pustules. Moving away from the centre the pustules expand, often collapse in the middle, and mature by developing isidia. In the less active (senescent ?) margins the pustules are eroded away and the supplementary intercalary growth seems insufficient to keep the outflowing thallus intact. Thus the margins are disrupted into irregular lobes.     

Dew as a key factor for the distribution pattern of the lichen species Teloschistes lacunosus in the Tabernas Desert (Spain)

The role of different sources of water (rain, dew and water vapor) has been investigated under natural conditions in order to explain the activity and the distribution patterns of Teloschistes lacunosus (Rupr.) Sav. in the Tabernas Desert (Almeria, Spain). This field work was carried out at two neighboring sites: a pediment where T. lacunosus is well developed and an east-facing slope where only few small thalli are developed. Diurnal courses of photosynthetic activity were assessed by the use of chlorophyll a fluorescence measurements, at each site for a total of 12 days distributed among different seasons over the year. Microclimatic data (thallus temperature, relative humidity (RH) and light intensity) were recorded continuously for a period of 1 year including all the days on which fluorescence measurements were made. Dried T. lacunosus in its natural habitat only became photosynthetically active after re-hydration with liquid water (dew or rain). In contact with an atmosphere of high RH (higher than 90%) but without dew condensation, thalli were not able to obtain sufficient water to become physiologically active. The microclimatic study showed notably differences between the two studied expositions. After dawn, thalli from the east-facing slope were exposed to higher temperatures and light intensity (PPFD) levels than thalli from the pediment. This was reflected in the length of time that the air remained saturated and the lichen remained wet and active. The high incident PPFD and the resultant increased temperatures at the east-facing slope led to short dew duration and, therefore to shorter periods of morning photosynthetic activity than on the pediment as fluorescence measurements showed. Additionally, the microclimatic differences between the two sites indicated a high frequency of dew fall events on the thalli from the pediment. The time periods of thallus dew imbibition vary strongly with the exposure of the lichens.     

Lichen respiration in relation to active time, temperature, nitrogen and ergosterol concentrations

1. Respiration in eight lichen species was related to thallus hydration status, external temperature and to total nitrogen, chitin and ergosterol concentrations. Chitin is a nitrogenous and major compound of the fungal cell wall, and ergosterol is a sterol of the plasma membrane in fungi and sometimes in algae.
2. Hydration of previously dry thalli resulted in an initially high rate of respiration. Both the amplitude of this resaturation respiration and the time required to reach steady state varied among species. Generally, peak rates were one to three times higher than steady-state rates, which were reached 3–7 h after hydration.
3. Increases in external temperature also resulted in transient bursts in respiration. Again, both the amplitude of the burst and the time required to reach steady state varied among species. Also depending on species, a temperature increase from 5 to 15 °C resulted in two- to fivefold increases in steady-state respiration.
4. Steady-state respiration, at optimal thallus hydration and a given temperature, varied three- to sixfold among the species, when related to thallus dry mass. This difference correlated best ( r ² = 0·89) with their ergosterol concentration, where a doubling in ergosterol resulted in more than a doubling in respiration. Respiration correlated less well to total nitrogen or chitin.
5. The chitin to ergosterol ratio varied more than one order of magnitude between the species, where species with high nitrogen concentrations had the highest ratio. This implies that species with access to ample amounts of nitrogen can make more fungal cell walls in relation to plasma membrane surface area.     

Exploring Phenotypic Plasticity in the LichenRamalina capitata: Morphology, Water Relations and Chlorophyll Content in North- and South-facing Populations

The present work analyses the morphology, anatomy, water relationsand chlorophyll content of thalli of the lichenRamalina capitatavar.protectafromtwo different populations exposed to contrasting microclimaticconditions due to differences in the orientation of the rocksurface. The population on the north-facing rock surface (NFS)was exposed to lower photosynthetic photon flux densities (PPFD),remained at high relative humidities for longer periods of timeand was exposed to lower temperatures than the population onthe south-facing surface (SFS). We proposed the hypothesis thatthe shadier the habitat the greater the ecological advantagefor enhanced light harvesting. Thalli from the SFS had shorterand wider lacinia, thicker thalli, mostly due to increased medullathickness, a higher water-retention capacity, a higher percentageof thallus volume occupied by the algal cells and a higher chlorophyllcontent than thalli from NFS. The phenotypic plastic responseof the traits studied inR. capitatavar.protectawas not directlyrelated to differences in the light availability, at least forthe range of PPFD experienced by the two populations studied,since the population exposed to higher PPFD exhibited largeramounts of light harvesting pigments. Both populations exhibitedthe same intrathalline distribution of algal cells and chlorophylls,which were more abundant in the apical than in the basal zonesof all thalli studied. Periods of water-induced metabolic activitywere shorter in the SFS than in the NFS, and structural andchlorophyll data indicated that thalli from the SFS were betterprepared for the photosynthetic exploitation of these brieferperiods and for maintaining thallus hydration into dry periods.These results suggest that differences in selective pressurebetween the two populations ofR. capitatavar.protectastudiedinvolved maximization of the photosynthetic exploitation ofthe periods of metabolic activity when they are brief, as hasbeen described for certain vascular plants from xeric environments. Ramalina capitatavar.protecta; algal cells; chlorophylls; water relations; microclimate; morphology; intrathalline variation; lichen; phenotypic plasticity     

Surface hydrophobicity causes SO2 tolerance in lichens

BACKGROUND AND AIMS: The superhydrophobicity of the thallus surface in one of the most SO(2)-tolerant lichen species, Lecanora conizaeoides, suggests that surface hydrophobicity could be a general feature of lichen symbioses controlling their tolerance to SO(2). The study described here tests this hypothesis. METHODS: Water droplets of the size of a raindrop were placed on the surface of air-dry thalli in 50 lichen species of known SO(2) tolerance and contact angles were measured to quantify hydrophobicity. KEY RESULTS: The wettability of lichen thalli ranges from strongly hydrophobic to strongly hydrophilic. SO(2) tolerance of the studied lichen species increased with increasing hydrophobicity of the thallus surface. Extraction of extracellular lichen secondary metabolites with acetone reduced, but did not abolish the hydrophobicity of lichen thalli. CONCLUSIONS: Surface hydrophobicity is the main factor controlling SO(2) tolerance in lichens. It presumably originally evolved as an adaptation to wet habitats preventing the depression of net photosynthesis due to supersaturation of the thallus with water. Hydrophilicity of lichen thalli is an adaptation to dry or humid, but not directly rain-exposed habitats. The crucial role of surface hydrophobicity in SO(2) also explains why many markedly SO(2)-tolerant species are additionally tolerant to other (chemically unrelated) toxic substances including heavy metals.     

Development of areolae and growth of the peripheral prothallus in the crustose lichen Rhizocarpon geographicum: an image analysis study

Areolae of the crustose lichen Rhizocarpon geographicum (L.) DC., are present on the peripheral prothallus (marginal areolae) and also aggregate to form confluent masses in the centre of the thallus (central areolae). To determine the relationships between these areolae and whether growth of the peripheral prothallus is dependent on the marginal areolae, the density, morphology, and size frequency distributions of marginal areolae were measured in 23 thalli of R. geographicum in north Wales, UK using image analysis (Image J). Size and morphology of central areolae were also studied across the thallus. Marginal areolae were small, punctate, and occurred in clusters scattered over the peripheral prothallus while central areolae were larger and had a lobed structure. The size-class frequency distributions of the marginal and central areolae were fitted by power-law and log-normal models respectively. In 16 out of 23 thalli, central areolae close to the outer edge were larger and had a more complex lobed morphology than those towards the thallus centre. Neither mean width nor radial growth rate (RaGR) of the peripheral prothallus were correlated with density, diameter, or area fraction of marginal areolae. The data suggest central areolae may develop from marginal areolae as follows: (1) marginal areolae develop in clusters at the periphery and fuse to form central areolae, (2) central areolae grow exponentially, and (3) crowding of central areolae results in constriction and fragmentation. In addition, growth of the peripheral prothallus may be unrelated to the marginal areolae.     

Content of Gibberellic Acid in Apical Parts of Male and Female Thalli of Chara Tomentosa in Relation to the Content of Sugars and Dry Mass

The contents of gibberellic acid (GA₃), glucose and starch, and dry mass in four nodes of apical parts of male and female thalli of Chara tomentosa were measured. The GA₃ concentration was about 4 times higher in male than in female thallus and about 10 times higher in the first node than in the next three nodes of both thalli. The apical part of male thallus contains 2.5 times less glucose and 1.7 times less starch than female one. The average amount of dry mass of male and female thallus was similar: 9.9 and 9.8 % of fresh mass. The findings of this investigation demonstrate the interdependence of gibberellic acid and sugars in development of male and female Chara tomentosa thalli.     

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     

QUANTITATIVE VARIATION IN VULPINIC AND PINASTRIC ACIDS PRODUCED BY TUCKERMANNOPSIS PINASTRI (LICHEN-FORMING ASCOMYCOTINA,PARMELIACEAE)

Tuckermannopsis pinastri is a lichen species found commonly on rocks and tree branches in boreal and alpine habitats in the northern United States. Members of this species produce three yellow-pigmented phenolic compounds: usnic, pinastric, and vulpinic acids. The objective of our study was to quantify the variation in concentration of the latter two of these compounds in relation to substrate factors, chlorophyll content, and thallus size. Using high-performance liquid chromatography, we analyzed 120 thalli of T. pinastri collected randomly from a single large population located at Spruce Knob, West Virginia. Although individuals were sampled from both tree and rock substrates that differed markedly in light intensity, these environmental factors were not correlated with observed variations in vulpinic or pinastric acid concentrations. Instead, compound concentrations were correlated most closely with thallus size, with small rather than large thalli having the highest concentrations of the two compounds. Small thalli did not have higher concentrations of chlorophylls than large thalli, however, which suggests that the rate of production of secondary compounds by the fungus in T. pinastri is independent of algal biomass. Inasmuch as lichen secondary compounds serve a defensive role against microorganisms and herbivores, our results suggest that small, juvenile thalli are better defended than more mature thalli.     

Lobe formation and division in the foliose lichen <Emphasis Type="Italic">xanthoparmelia conspersa</Emphasis>

New lobe development and lobe division was studied in the foliose lichen Xanthoparmelia conspersa (Ehrh. ex. Ach.) Hale. In thalli with either the centre or margin removed, the inside edge of the perimeter, the outer edge of the reproductive centre, and fragments derived from the thallus perimeter all regenerated growing points (‘lobe primordia’) within a year. Thalli possessing isidia had the greatest ability to regenerate growing points. In reproductive thalli, there was a positive correlation between the density of new growing points and thallus size. When fragments were cut from the perimeters of mature X. conspersa thalli and glued to pieces of slate, the ratio of growing points to mature lobes increased over 54 months. Lobes within a thallus exhibited different degrees of bifurcation. In some bifurcating lobes, the point of origin of the bifurcation advanced at the same rate as the lobe tips over 4 months but in most lobes, the bifurcation point either advanced less rapidly than the lobe tips or retreated from its original location. Removing adjacent lobes had no significant effect on the radial growth of a lobe over 4 months or on the location of the bifurcation point but it increased the number of growing points. These results suggest that for X. conspersa: 1) all portions of of thalli can regenerate growing points, 2) few growing points actually develop into mature lobes, 3) individual lobes within a thallus grow and divide differently, and 4) adjacent lobes inhibit the development of growing points on their neighbours.     

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.     

The Localization of Lead in the Lichen Ramalina duriaei (De Not.) Bagl.

In an attempt to ascertain whether the lichen Ramalina duriaei could be employed to biomonitor atmospheric lead pollution, specimens of this lichen were exposed to Pb (NO₃)₂ solutions and a buffered solution (tartaric acid/sodium bitartrate, pH 2.8) of sodium rhodizonate (C₆O₆Na₂) was used to locate lead in their thalli. The procedure entailed exposure of the lichen to 0, 5, 50 and 100 ppm Pb for 5 min and 1 h and the subsequent determination of the lead contents from photographs of the thalli. Distribution of lead in different parts of the thallus was assessed after exposure of the lichens to 2 ppm Pb (9 h or three d), 50 ppm (45 min) or 200 ppm (4d). Cross sections of vegetative parts of the thallus and of the apothecia revealed that lead penetrated into the cortical cells of the thallus but not into the algal cells of the phycobiont nor the ascospores or medullary cells. The observed massive penetration of lead into cortical cells supports the notion that Ramalina duriaei is sensitive to atmospheric lead pollution.     

Developmental morphology of the thalloid Hydrobryum japonicum (Podostemaceae)

We describe the unique development and branching of lobed thalli in Hydrobryum japonicum. Lobe formation begins with meristem initiation at random sites near the thallus margin fringed by protective tissues. As the protective tissues are successively peeled off particularly in the growing new lobes, the lobes become naked and then become fringed again by new protective tissues that develop from the marginal part of the new meristems. Subsequently the meristems become less active and are differentiated into parenchymatous ground tissue at maturity. The random pattern of meristem formation during the sporadic development gives rise to a nonorderly branching pattern of the thalli. Some other lobes (～10%) are regenerated from injured parts of the thalli. The vegetative shoots arise endogenously near the thallus margin and are enclosed by the nonvascular strand nets. The rudimentary shoot apices remain embedded in the thalli. The thalli, though remarkably different from typical roots of other angiosperms, might be extremely transformed roots.     

Induction of urease by urea analogues in Evernia prunastri thallus

Urease activity in Evernia prunastri (L.) Ach. thallus is induced by incubation of lichen samples on 20 m M N,N-dimethylformamide and 20 m M N-formylurea or 40 m M thiourea although, in these two last cases, activity subsequently decreases again. The induction of enzyme activity is repressed by including 40 μ M cycloheximide in the medium. Filtration through Sepharose 6B of cell-free extracts from thalli incubated on 20 m M N,N-dimethylformamide shows a main peak of urease activity which has a molecular weight of about 560000 dalton. However, those extracts from thalli floated on 20 m M N-formylurea and 40 m M thiourea show several peaks of similar enzyme activity, which have molecular weights of about 1 100000, 670000, 260000 and 140000 dalton and 1 100000, 670000 and 140000 dalton respectively.
A time-course of urease activity could be related to the accumulation of lichen phenols in the thallus for samples incubated on N,N-dimethylformamide and thiourea.     

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.     

Fungal melanins as a sun screen for symbiotic green algae in the lichen<Emphasis Type="Italic"> Lobaria pulmonaria</Emphasis>

The mycobiont of the high-light-susceptible forest lichen Lobaria pulmonaria was shown to deposit brown, melanic compounds in the outer layer of the upper cortex, depending on the long-term level of solar radiation in its natural habitat. Furthermore, pale thalli from a shady habitat produced melanic compounds after transplantation to a sunny habitat. This browning of the cortex appeared to be a physiologically active process, taking place only during periods with frequent hydration. Melanin production was slow. After transplantation, more than 1 year was needed for a shade-adapted thallus to reduce the cortical transmittance (230-1000 nm) to a similar level to that of naturally sun-exposed specimens. Melanic compounds acted as a sun screen, especially reducing UVB and UVA wavelengths, but also visible wavelengths, at the photobiont level. In the near infrared range, there was only a small difference in transmittance between shade- and sun-adapted cortices. A negative correlation was found between the natural light level and the cortical transmittance of wavelengths below 700 nm. However, previous studies have shown that even photobionts of melanic L. pulmonaria thalli are relatively susceptible to high-light exposure. Since melanins also increase the absorbance of solar energy for the whole thallus, it appears that what is gained in terms of UV- and light protection in melanic L. pulmonaria specimens may be offset by increased exposure to excess temperatures for this highly heat-susceptible lichen.     

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.