Cyanolichens can have both cyanobacteria and green algae in a common layer as major contributors to photosynthesis

Background and Aims

Cyanolichens are usually stated to be bipartite (mycobiont plus cyanobacterial photobiont). Analyses revealed green algal carbohydrates in supposedly cyanobacterial lichens (in the genera Pseudocyphellaria, Sticta and Peltigera). Investigations were carried out to determine if both cyanobacteria and green algae were present in these lichens and, if so, what were their roles.

Methods

The types of photobiont present were determined by light and fluorescence microscopy. Small carbohydrates were analysed to detect the presence of green algal metabolites. Thalli were treated with selected strengths of Zn²⁺ solutions that stop cyanobacterial but not green algal photosynthesis. CO₂ exchange was measured before and after treatment to determine the contribution of each photobiont to total thallus photosynthesis. Heterocyst frequencies were determined to clarify whether the cyanobacteria were modified for increased nitrogen fixation (high heterocyst frequencies) or were normal, vegetative cells.

Key Results

Several cyanobacterial lichens had green algae present in the photosynthetic layer of the thallus. The presence of the green algal transfer carbohydrate (ribitol) and the incomplete inhibition of thallus photosynthesis upon treatment with Zn²⁺ solutions showed that both photobionts contributed to the photosynthesis of the lichen thallus. Low heterocyst frequencies showed that, despite the presence of adjacent green algae, the cyanobacteria were not altered to increase nitrogen fixation.

Conclusions

These cyanobacterial lichens are a tripartite lichen symbiont combination in which the mycobiont has two primarily photosynthetic photobionts, ‘co-primary photobionts’, a cyanobacterium (dominant) and a green alga. This demonstrates high flexibility in photobiont choice by the mycobiont in the Peltigerales. Overall thallus appearance does not change whether one or two photobionts are present in the cyanobacterial thallus. This suggests that, if there is a photobiont effect on thallus structure, it is not specific to one or the other photobiont.     

Midday dew--an overlooked factor enhancing photosynthetic activity of corticolous epiphytes in a wet tropical rain forest

? Additional water supplied by dew formation is an important resource for microbes, plants and animals in precipitation-limited habitats, but has received little attention in tropical forests until now. ? We evaluated the micro-environmental conditions of tree stem surfaces and their epiphytic organisms in a neotropical forest, and present evidence for a novel mechanism of diurnal dew formation on these surfaces until midday that has physiological implications for corticolous epiphytes such as lichens. ? In the understorey of a lowland forest in French Guiana, heat storage of stems during the day and delayed radiative loss during the night decreased stem surface temperatures by 6°C in comparison to the dew-point temperature of ambient air. This measured phenomenon induced modelled totals of diurnal dew formation between 0.29 and 0.69 mm d?1 on the surface of the bark and the lichens until early afternoon. ? Crustose lichens substantially benefit from this dew formation, because it prolongs photosynthetic activity. This previously unrecognized mechanism of midday dew formation contributes to the water supply of most corticolous organisms, and may be a general feature in forest habitats world-wide.     

When family matters: an analysis of Thelotremataceae (Lichenized Ascomycota: Ostropales) as bioindicators of ecological continuity in tropical forests

We analysed patterns of habitat and microhabitat preferences of 19 families (comprising 135 genera and 950 species) of crustose, corticolous lichens in Costa Rica (Arthoniaceae, Arthopyreniaceae, Coenogoniaceae, Graphidaceae, Lecanoraceae, Letrouitiaceae, Monoblastiaceae, Pertusariaceae, Physciaceae, Pilocarpaceae, Porinaceae, Pyrenulaceae, Ramalinaceae, Roccellaceae, Strigulaceae, Teloschistaceae, Thelenellaceae, Thelotremataceae, Trypetheliaceae), in order to test whether Thelotremataceae are suitable predictors of undisturbed tropical rain forest and can be used as bioindicators of ecological continuity. The dataset consisted of 12,215 specimen samples and six environmental parameters recorded for each sample (altitude, degree of seasonality, vegetation type, disturbance level, substrate nature, light exposure), which were analysed by a multivariate approach using principal component analysis (PCA). The analysis showed that three of the 19 families, Letrouitiaceae, Porinaceae, and Thelotremataceae, showed significant preferences for undisturbed primary to old growth secondary forest, fully shaded to semi-exposed microhabitats, and the bark of mature tree trunks, parameters assumed to be correlated with ecological continuity of closed rain forest habitats. Thelotremataceae had broader altitudinal range than Letrouitiaceae and Porinaceae and significantly higher genus and species diversity (16 genera, 130 species) compared to Porinaceae (4 genera, 40 species) and Letrouitiaceae (1 genus, 5 species). Our results support the hypothesis that Thelotremataceae perform best as predictors of undisturbed dry and lowland to montane rain forest and are the most suitable lichen bioindicators of ecological continuity in these ecosystems. In contrast, Lecanoraceae, Pertusariaceae, Physciaceae, and Teloschistaceae, were found to be predictors of disturbed and pioneer (micro-)habitats. We also found that, among a variety of parameters tested, the Index of Ecological Continuity (IEC), adapted to the use of Thelotremataceae in tropical forests, performs best in terms of predicting disturbance levels and site history. A semi-taxonomic approach identifying morphotypes rather than genera or species yielded the same results, making this method suitable for a broader spectrum of potential users.     

Functional Analysis of the Intrathalline and Intracellular Chlorophyll Concentrations in the Lichen Family Umbilicariaceae

Intrathalline and intracellular chlorophyll concentrations togetherwith thallus structure were studied in 12 species of the lichenfamily Umbilicariaceae in order to explore pigment limitationsfor light absorption and for maximum rates of net photosynthesis(A_max). Species from high light environments tended to haverelatively low chlorophyll concentrations and vice versa. Chlorophyllconcentration on a surface area basis of all species studiedwas sufficient to absorb 80–99% of the incident light,which led us to believe that chlorophyll concentration was nota limiting factor for A_max. A_maxof Umbilicariaceae was smallerthan A_maxof leaves of vascular plants, both on a dry weightand on a surface area basis. These differences slightly decreasedbut did not disappear when referred to the chlorophyll concentrationof the photosynthetic tissues, but reference units had a confoundingeffect in the comparisons. Differences in A_maxon a dry weightbasis between Umbilicariaceae and vascular plants disappearedwhen comparisons were made with the whole plant and not simplywith the leaf. We found a significant, inverse relationshipbetween amount of alga and chlorophyll concentration withinthe algal cells in the lichens studied. This was interpretedas a mechanism to avoid pigment self-shading within the algallayer, distributing more efficiently the photosynthetic pigmentsover a given surface area, and is the first indication of sucha mechanism in lichens. It is suggested that thallus structureand physiology have a larger influence on the observed low A_maxofthe lichens studied than chlorophyll content. Algal cells; chlorophyll concentration; lichens; light harvesting; net photosynthesis; stereology; thallus structure; Umbilicariaceae     

Photosynthetic capacity in relation to nitrogen content and its partitioning in lichens with different photobionts

We tested the hypothesis that lichen species with a photosynthetic CO₂-concentrating mechanism (CCM) use nitrogen more efficiently in photosynthesis than species without this mechanism. Total ribulose bisphosphate carboxylase-oxygenase (Rubisco; EC 4.1.1.39) and chitin (the nitrogenous component of fungal cell walls), were quantified and related to photosynthetic capacity in eight lichens. The species represented three modes of CO₂ acquisition and two modes of nitrogen acquisition, and included one cyanobacterial ( Nostoc ) lichen with a CCM and N₂ fixation, four green algal ( Trebouxia ) lichens with a CCM but without N₂ fixation and three lichens with green algal primary photobionts ( Coccomyxa or Dictyochloropsis ) lacking a CCM. The latter have N₂-fixing Nostoc in cephalodia. When related to thallus dry weight, total thallus nitrogen varied 20-fold, chitin 40-fold, Chl a 5-fold and Rubisco 4-fold among the species. Total nitrogen was lowest in three of the four Trebouxia lichens and highest in the bipartite cyanobacterial lichen. Lichens with the lowest nitrogen invested a larger proportion of this into photosynthetic components, while the species with high nitrogen made relatively more chitin. As a result, the potential photosynthetic nitrogen use efficiency was negatively correlated to total thallus nitrogen for this range of species. The cyanobacterial lichen had a higher photosynthetic capacity in relation to both Chl a and Rubisco compared with the green algal lichens. For the range of green algal lichens both Chl a and Rubisco contents were linearly related to photosynthetic capacity, so the data did not support the hypothesis of an enhanced photosynthetic nitrogen use efficiency in green-algal lichens with a CCM.     

CO2 exchange and thallus nitrogen across 75 contrasting lichen associations from different climate zones

Aiming to investigate whether a carbon-to-nitrogen equilibrium model describes resource allocation in lichens, net photosynthesis (NP), respiration (R), concentrations of nitrogen (N), chlorophyll (Chl), chitin and ergosterol were investigated in 75 different lichen associations collected in Antarctica, Arctic Canada, boreal Sweden, and temperate/subtropical forests of Tenerife, South Africa and Japan. The lichens had various morphologies and represented seven photobiont and 41 mycobiont genera. Chl a, chitin and ergosterol were used as indirect markers of photobiont activity, fungal biomass and fungal respiration, respectively. The lichens were divided into three groups according to photobiont: (1) species with green algae, (2) species with cyanobacteria, and (3) tripartite species with green algal photobionts and cyanobacteria in cephalodia. Across species, thallus N concentration ranged from 1 to 50 mg g^-1 dry wt., NP varied 50-fold, and R 10-fold. In average, green algal lichens had the lowest, cyanobacterial Nostoc lichens the highest and tripartite lichens intermediate N concentrations. All three markers increased with thallus N concentration, and lichens with the highest Chl a and N concentrations had the highest rates of both P and R. Chl a alone accounted for ca. 30% of variation in NP and R across species. On average, the photosynthetic efficiency quotient [K_F=(NP_max+R)/R)] ranged from 2.4 to 8.6, being higher in fruticose green algal lichens than in foliose Nostoc lichens. The former group invested more N in Chl a and this trait increased NP_max while decreasing R. In general terms, the investigated lichens invested N resources such that their maximal C input capacity matched their respiratory C demand around a similar (positive) equilibrium across species. However, it is not clear how this apparent optimisation of resource use is regulated in these symbiotic organisms.     

Fungal farmers or algal escorts: lichen adaptation from the algal perspective

Domestication of algae by lichen‐forming fungi describes the symbiotic relationship between the photosynthetic (green alga or cyanobacterium; photobiont) and fungal (mycobiont) partnership in lichen associations ( Goward 1992 ). The algal domestication implies that the mycobiont cultivates the alga as a monoculture within its thallus, analogous to a farmer cultivating a food crop. However, the initial photobiont ‘selection’ by the mycobiont may be predetermined by the habitat rather than by the farmer. When the mycobiont selects a photobiont from the available photobionts within a habitat, the mycobiont may influence photobiont growth and reproduction ( Ahmadjian & Jacobs 1981 ) only after the interaction has been initiated. The theory of ecological guilds ( Rikkinen et al. 2002 ) proposes that habitat limits the variety of photobionts available to the fungal partner. While some studies provide evidence to support the theory of ecological guilds in cyanobacterial lichens ( Rikkinen et al. 2002 ), other studies propose models to explain variation in symbiont combinations in green algal lichens ( Ohmura et al. 2006 ; Piercey‐Normore 2006 ; Yahr et al. 2006 ) hypothesizing the existence of such guilds. In this issue of Molecular Ecology, Peksa & ?kaloud (2011) test the theory of ecological guilds and suggest a relationship between algal habitat requirements and lichen adaptation in green algal lichens of the genus Lepraria. The environmental parameters examined in this study, exposure to rainfall, altitude and substratum type, are integral to lichen biology. Lichens have a poikilohydric nature, relying on the availability of atmospheric moisture for metabolic processes. Having no known active mechanism to preserve metabolic thallus moisture in times of drought, one would expect a strong influence of the environment on symbiont adaptation to specific habitats. Adaptation to changes in substrata and its properties would be expected with the intimate contact between crustose lichens in the genus Lepraria. Altitude has been suggested to influence species distributions in a wide range of taxonomic groups. This is one of the first studies to illustrate an ecological guild, mainly for exposure to rainfall (ombrophiles and ombrophobes), with green algal lichens.     

不同水分条件下两种叶状地衣的光合活性对冻融交替的响应

冻融交替是我国北方常见气候现象,其对地衣光合作用的影响尚不清楚。研究了采自雾灵山的卷叶点黄梅Flavopunctelia soredica和平盘软地卷Peltigera elisabethae的光合活性(以净光合速率表示,net photosynthetic rate,Pn)对冻融处理的响应及其与地衣体含水量(干冻组:含水量<20%干重;湿冻组:含水量>200%)和物种的关系。结果显示卷叶点黄梅的干冻组Pn经5次冻融后下降至对照的21%,湿冻组经3次冻融后下降至负值,平盘软地卷的干冻组和湿冻组在5次冻融后Pn均为负值;相对净光合速率(relative net photosynthetic rate,Rpn)与冻融次数的线性回归分析表明,卷叶点黄梅湿冻组的斜率绝对值(58.06)>平盘软地卷湿冻组(41.01)>平盘软地卷干冻组(32.27)>卷叶点黄梅干冻组(11.44)。结果表明冻融胁迫可显著抑制两种地衣的光合活性,这种抑制作用具有物种差异且和地衣体内水分含量有关:水分含量的增高将增强冻融胁迫对地衣光合活性的抑制作用;干燥状态下,卷叶点黄梅的低温耐性远高于平盘软地卷,但在湿润状态下则低于后者。两种地衣对冻融循环的光合响应的物种差异与其微生境气候有关:生长于较干燥开阔地带的卷叶点黄梅与生于阴湿生境中的平盘软地卷相比,可能已形成了更强的低温干燥适应能力,其低温湿润适应能力则较弱。全球气候变化可能会通过冻融事件的时空格局的改变而对地衣的光合作用和分布造成负面影响。     

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.     

Diversity and vertical distribution of epiphytic macrolichens in lowland rain forest and lowland cloud forest of French Guiana

Recent work on bryophyte diversity in lowland forests of northern South America has suggested the existence of a new type of cloud forest, the “tropical lowland cloud forest” (LCF). LCF occurs in river valleys with high air humidity and radiation fog, and is rich in epiphytes. We explored the lichenological characteristics of putative LCF in a lowland area (200–400 m a.s.l.) near Saül, central French Guiana, using macrolichens (including large crustose species) as indicator taxa. We analyzed macrolichen diversity on 16 trees in two 1 ha plots, in LCF and in lowland rain forest without fog (LRF). Sampling efficiency was ca. 80% in both forest types. Canopies of both LRF and LCF were richer in lichen species than understory trunks. Species richness of macrolichens was rather similar in the two forest types but species composition was significantly different. Cyanolichen richness in LCF was ca. 2.5 times higher than in LRF; in contrast, LRF had 4 times more species of green-algal Parmeliaceae. Our study suggests that cyanolichens except for Coccocarpiaceae serve as indicators of LCF. We explain the detected diversity patterns by differences in water availability due to fog precipitation and higher humidity. This is indicated by the higher relative air humidity in the lowland cloud forest, which was >6% higher than in the rain forest.     

As thick as three in a bed

During the evolution of the lichen symbiosis, shifts from one main type of photobiont to another were infrequent (Miadlikowska et al. 2006 ) but some remarkable transitions from green algal to diazotrophic cyanobacterial photobionts are known from unrelated fungal clades within the ascomycetes. Cyanobacterial, including tripartite, associations (green algal and cyanobacterial photobionts in one lichen individual) facilitate these holobionts to live as C‐ and N‐autotrophs. Tripartite lichens are among the most productive lichens, which provide N‐fertilization to forest ecosystems under oceanic climates (Peltigerales) or deliver low, but ecologically significant N‐input into subarctic and alpine soil communities (Lecanorales, Agyriales). In this issue of Molecular Ecology, Schneider et al. (2016) mapped morphometric data against an eight‐locus fungal phylogeny across a transition of photobiont interactions from green algal to a tripartite association and used a phylogenetic comparative framework to explore the role of nitrogen‐fixing cyanobacteria in size differences in the Trapelia–Placopsis clade (Agyriales). Within the group of tripartite species, the volume of cyanobacteria‐containing structures (cephalodia) correlates with thallus thickness in both phylogenetic generalized least squares and phylogenetic generalized linear mixed‐effects analyses, and the fruiting body core volume increased ninefold. The authors conclude that cyanobacterial symbiosis appears to have enabled lichens to overcome size constraints in oligotrophic environments such as rock surfaces. The Trapelia–Placopsis clade analyzed by Schneider et al. (2016) is an exciting example of interactions between ecology, phylogeny and lichen biology including development – from thin crustose green algal microlichens to thick placodioid, tripartite macrolichens: as thick as three in a bed (Scott 1820 ).     

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.     

Fire in tropical dry forests: corticolous lichens as indicators of recent ecological changes in Thailand

The distribution of lichens in lowland deciduous and evergreen forests in Thailand is used to interpret recent changes in the distribution of these forests. The role of fire in changing the forest structure, microclimate and species content is discussed. Characteristic corticolous lichen communities of dry deciduous and moist evergreen forests are described, as well as changes in the composition of the flora following fire events. Where frequent fires have altered the forest rates of change in forest type are suggested using lichen data from randomly selected trees in forest plots, and growth rates of sampled species in quadrats. The disjunct nature of the lichen floras in lowland deciduous and evergreen forests is discussed, their origin and use in interpreting changes in forest types in monsoon climates over long periods of time.     

Nitrate sequestration by corticolous macrolichens during winter precipitation events

Nitrogen is an essential nutrient in the biogeochemistry of forested ecosystems. The influence of canopy lichens on the winter biogeochemistry of nitrate in broadleaved deciduous forests is examined and it is hypothesized that nitrate sequestration will not differ between winter precipitation events. Rejection of this hypothesis would mean that meteorological conditions of winter precipitation events have a detectable influence on nitrate sequestration by canopy lichens and nitrate input to the forest floor. Canopy lichens of the genus Parmelia were found to influence winter nitrate stemflow inputs to forest soils differentially. Epiphytic lichens on an individual Carya glabra Mill. (pignut hickory) canopy tree, centrally located within the stand of an open deciduous forest, actively sequestered nitrate leached from the tree's woody frame, lowering aqueous stemflow inputs at the tree base. The quantities of nitrate sequestered by corticolous lichens during the 2 February 1999 mixed-precipitation event were significantly greater than those during all other precipitation events examined. Greater rates of nitrate uptake during the 2 February 1999 event may be attributed to (1) its intermediate rain intensity, which would have soaked the lichen thalli in a nutrient-rich bath, and (2) an air temperature range between -2 degrees C and 8 degrees C that would have increased viscosity and surface tension of stemflow drainage, thereby decreasing stemflow velocity and increasing the contact time of stemflow water on the lichen thalli. Other precipitation events were either too cold to promote metabolic activity by canopy lichens or too warm and intense for an optimal contact time of stemflow with lichen thalli, resulting in lower quantities of nitrate sequestered. Meteorological conditions of winter precipitation events have been documented to influence sequestration of nitrate by corticolous lichens and decrease aqueous stemflow inputs to the forest floor of broadleaved deciduous forests.     

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.     

Small Canopy Gaps Influence Plant Distributions in the Rain Forest Understory1

This study tested three hypotheses regarding how plants respond to the spatial heterogeneity in light availability in the rain forest understory: (1) understory plants occur preferentially in the lighter parts of the understory; (2) under–story palms are more shade tolerant than other understory plants; (3) rain forest plants differ in their ontogenetic response to understory light conditions. The study was carried out in old–growth rain forest in the Yasuní National Park, Amazonian Ecuador. The hypotheses were tested by comparing the distributions of 20 plant species (1454 individuals) over microsites with differing degrees of exposure to canopy gaps to the background distribution of these microsites in the forest. The gap exposure of a given microsite was described by an index based on the number and size of gaps in the canopy to which the site was exposed. Two plant height classes were studied: 0.80–2.49 and 2.50–5 m. The first and third hypotheses were accepted, while the second hypothesis was rejected. The results for the individual species corresponded well with what is known from earlier studies about the ecology of these species or close relatives, suggesting that the patterns observed can be generalized for Neotropical rain forests. Notably, the most abundant species in the study represent several different life history strategies. Thus, abundance in the rain forest understory can be achieved by several different strategies. This suggests that niche differentiation in terms of response to small changes in understory light conditions may be an important factor in the maintenance of the high local plant species richness of tropical rain forests.     

Effect of high light on the efficiency of photochemical energy conversion in a variety of lichen species with green and blue-green phycobionts

Exposure to high light induced a quantitatively similar decrease in the rate of photosynthesis at limiting photon flux density (PFD) and of photosystem II (PSII) photochemical efficiency, F_V/F_M, in both green and blue-green algal lichens which were fully hydrated. Such depressions in the efficiency of photochemical energy conversion were generally reversible in green algal lichens but rather sustained in blue-green algal lichens. This greater susceptibility of blue-green algal lichens to sustained photoinhibition was not related to differences in the capacity to utilize light in photosynthesis, since the light-and CO₂-saturated rates of photosynthetic O₂ evolution were similar in the two groups. These reductions of PSII photochemical efficiency were, however, largely prevented in lichen thalli which were fully desiccated prior to exposure to high PFD. Thalli of green algal lichens which were allowed to desiccate during the exposure to high light exhibited similar recovery kinetics to those which were kept fully hydrated, whereas bluegreen algal lichens which became desiccated during a similar exposure exhibited greatly accelerated recovery compared to those which were kept fully hydrated. Thus, green algal lichens were able to recover from exposure to excessive PFDs when thalli were in either the hydrated or desiccated state during such an exposure, whereas in blue-green algal lichens the decrease in photochemical efficiency was reversible in thalli illuminated in the desiccated state but rather sustained subsequent to illumination of thalli in the hydrated state.Abbreviations and Symbols F_o yield of instantaneous fluorescence - F_M maximum yield of fluorescence induced by pulses of saturating light - F_V variable yield of fluorescence - PFD photon flux density (400–700 nm) - PSII photosystem II This work was supported by the Deutsche Forschungsgeneinschaft (Forscherguppe Ökophysiologic and Sonderforschungsbereich 251 of the University of Würzburg) and the Fonds der Chemischen Industrie. W.W.A. gratefully acknowledges the support of a fellowship from the Alexander von Humboldt Foundation. We thank Professor T.G.A. Green for identifying and supplying all of the New Zealand lichen material and Professor F.-C. Czygan for advice concerning the chlorophyll analyses which were performed by Johanna Leisner.     

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.     

THALLUS ORGANIZATION AND DEVELOPMENT IN THE FRUTICOSE LICHENASPICILIA CALIFORNICA,WITH COMPARISONS TO OTHER TAXA

Thallus organization is examined inAspicilia californicaRosentreter, a fruticose lichen known from several localities in central and southern California. The sprawling, terete thallus branches possess a dense central medulla of thick-walled, longitudinally oriented fungal cells. This central tissue emerges at branch apices to form a darkly pigmented fungal tip. Thallus development involves the apical extension of the tip to produce a fungal tissue over which a cylindrical algal layer and cortex will eventually be formed. Apical branches are initiated by furcation entirely within the fungal tip. Lateral branches, emerging from the lichenized thallus, arise as a divergent bundle of elongate fungal cells originating in the medulla. The photobiont appears to play no direct role in initiation of apical or lateral branches. It is concluded that thallus development inA. californicaoccurs with a relatively low degree of synchrony between mycobiont and photobiont growth, similar to the pattern observed in crustose lichens with prothallic growth. A rather similar type of thallus organization is observed inA. hispida, although in that species mycobiont growth and branch initiation appear to be somewhat more closely associated with algal cell proliferation. A squamuloseAspiciliafrom central Spain produces rhizomorphs that may sometimes become invested with an algal layer and cortex, resembling the thallus axes ofA. californica.     

Do leaf‐cutting ant nests make “bottom–up” gaps in neotropical rain forests?: a critical review of the evidence

We discuss the role of leaf cutting ant nests ( Atta sp.) on the formation of understory "bottom–up" gaps in tropical rain forests. We (i) describe the characteristics of Atta nests that qualify them as promoters of understory gaps; (ii) review the effects of Atta nests on the dynamics of systems other than tropical forests; and (iii) discuss the hypotheses that in neotropical rain forests Atta nests affect plant productivity, and also influence distribution and community composition of the flora. Finally, we discuss the importance of Atta nests in the dynamics of forests and compare their effects with the better-known effects of treefall gaps. Atta nests modify soil structure, soil nutrient content, light intensity at ground level, seed bank composition and understory structure. The presence of the nest and associated refuse also accelerates the soil nutrient cycles. The size, high frequency, turnover rate and duration of these nests could make them a good substrate for particular species of plants. In spite of their potential importance, little information exists on the effects of Atta nests on plant productivity and/or plant composition in neotropical rain forests.