GENE FLOW IN LICHENS

Experimental studies of the evolutionary biology of lichen fungi have been hampered by massive difficulties of in vitro culture and artificial crosses are still not possible. Gene flow in these organisms is demonstrated here for the first time by the analysis of secondary products in the progeny of individuals from natural populations of mixed chemotypes of the Cladonia chlorophaea complex. All of the chemotypes in this study have been interpreted as distinct sibling species. In the Appalachian Mountains of North Carolina, however, the grayi and merochlorophaea chemotypes are found to belong to a single interbreeding populations that is reproductively isolated from the cryptochlorophaea chemotype. In the Coastal Plain, the cryptochlorophaea chemotype hybridizes with the local endemic perlomera chemotype. This study has major impact for species concept in lichens because consideration of neither morphologial tendencies nor biogenetic relationships of the secondary products could have predicted its result.     

Sex-specific effects of hybridization on reproductive fitness in Mytilus

Blue mussels of the genus Mytilus form extensive hybrid zones in the North Atlantic and elsewhere where the distributions of different species overlap. Mytilus species transmit both maternal and paternal mtDNA through egg and sperm, respectively, a process known as doubly uniparental inheritance (DUI), and some females produce offspring with extremely biased sex ratios. These two traits have been shown to be linked and maternally controlled, with sex determination involving nuclear–cytoplasmic interactions. Hybridization has been shown to disrupt DUI mitochondrial inheritance and sex ratio bias; however, the effect of hybridization on reproductive fitness has not previously been examined. We investigated this effect in M. edulis × M. trossulus crosses through histological examination of mature F1 progeny, and spawning of F1 hybrids to monitor survival of their progeny through to the D stage of larval development. For progeny produced from mothers with a strong bias toward female offspring (often 100%) in pure-bred crosses, there was a clear breakdown in female dominance of progeny and significantly more hermaphrodites in the hybrid crosses produced from sperm with the M-tr1 mitotype. We also found significant sex-specific differences among hybrid progeny, with females producing normal eggs while males and hermaphrodites evidenced impaired gonadal development with significantly greater numbers of Sertoli cells, phagocytic hemocytes, and degenerating germ cells, all associated with gonad resorption. Males from crosses where DUI was disrupted and where male progeny were homoplasmic for the female mtDNA were the most severely compromised. Allelic incongruity between maternal and paternal mitotypes in hybrid crosses was associated with significant disruption of male gonadal development.     

Nitrogen and carbon isotope variability in the green‐algal lichen Xanthoria parietina and their implications on mycobiont–photobiont interactions

Stable isotope patterns in lichens are known to vary largely, but effects of substrate on carbon and nitrogen stable isotope signatures of lichens were previously not investigated systematically. N and C contents and stable isotope (δ¹⁵N, δ¹³C) patterns have been measured in 92 lichen specimens of Xanthoria parietina from southern Bavaria growing on different substrates (bark and stone). Photobiont and mycobiont were isolated from selected populations and isotopically analyzed. Molecular investigations of the internal transcribed spacer of the nuclear ribosomal DNA (ITS nrDNA) region have been conducted on a subset of the specimens of X. parietina. Phylogenetic analysis showed no correlation between the symbionts X. parietina and Trebouxia decolorans and the substrate, isotope composition, or geographic origin. Instead specimens grown on organic substrate significantly differ in isotope values from those on minerogenic substrate. This study documents that the lichens growing on bark use additional or different N sources than the lichens growing on stone. δ¹⁵N variation of X. parietina apparently is controlled predominantly by the mass fraction of the mycobiont and its nitrogen isotope composition. In contrast with mycobionts, photobionts of X. parietina are much more ¹⁵N‐depleted and show less isotopic variability than mycobionts, probably indicating a mycobiont‐independent nitrogen acquisition by uptake of atmospheric ammonia.     

Competitive behaviour of umbilicate lichens – an experimental approach

The lichens Lasallia pustulata and Umbilicaria spodochroa grow in dense monospecific or mixed populations on the coastal cliffs of southern Scandinavia. Attached to the substrate by only a thin central holdfast, their shield-shaped thalli compete for light and space for growth by overlapping each other. Matched pair experiments in the laboratory and field observations of interacting pairs show that different behavioural responses to precipitation tend to result in the margins of U. spodochroa overlapping those of L. pustulata within a few minutes. The behaviour is apparently caused by different capacities for water absorption in the upper and lower cortices of the species. An initial period of repeated encounter caused by thallus expansion and contraction during precipitation will be followed by a period in which U. spodochroa grows to overlap L. pustulata more and more. When the overlapping lichens are wet, flexible and photosynthetically active, the thallus above rests directly on the upper surface of the one below. Very little light is transmitted through thalli of U. spodochroa, and the shaded parts of L. pustulata are retarded in their growth and die off. Received: 29 August 1996 / Accepted: 19 April 1997     

Habitat overlap/niche segregation in two Umbilicaria lichens: a possible mechanism

Summary Umbilicaria muhlenbergii and Umbilicaria papulosa are two similar lichens which show extensive habitat overlap. Current theory predicts that for those resources that are limited, and of critical importance to both species, segregation is demand for or use of those resources ought to be present. Controlled experiments showed that the upper and lower cortical surfaces of the two species functioned in very different ways so that U. muhlenbergii is wetted most rapidly by runoff and mist-like precipitation whereas U. papulosa is wetted most rapidly by large raindrops. An analysis of the meteorological conditions in the natural habitat for the two lichens shows that the four months providing significant productivity in U. papulosa are the same months with a very high frequency of thunderstorms (which produce large drops). Conversely, the months which produce the greatest production in U. muhlenbergii are spread out over most of the year. When water is supplied after periods of dryness, the two species absorb it at different times and from different sources. This difference supports current ideas about niche segregation in plants but represents the first time it has been demonstrated for lichens.     

Vertical distribution of epiphytic lichens on Quercus laurina Humb. & Bonpl. in a remnant of cloud forest in the state of Veracruz,México

In the tropics, corticolous lichen richness and cover tend to increase from the trunk base to the top of the crown of trees. In this study we calculated the total beta diversity of the lichen community along a vertical gradient on Quercus laurina in Mexican cloud forest. By comparing the richness and cover of the lichens by zone, we show that foliose and fruticose lichens are a minor component of the total lichen species richness, but have a higher cover than the crustose lichens. Five zones were identified along each phorophyte (n = 15) with a diameter at breast height >40 cm. A total of 92 species were identified. Of these, 38% were found only in a single zone, 51% were shared between the different zones and 11% occurred across all zones. Species richness and cover increased from the lowest to the highest zones of the phorophytes. Dissimilarity in species composition between the zones could be explained by species replacement. An indicator species analysis revealed that only a few species, e.g. Hypotrachyna vexans, H. cf. sublaevigata and Ramalina cf. sinaloensis prefer a particular zone. The results show that the lichen community associated with Quercus laurina phorophytes is highly diverse and suggest that species richness and cover are related to the zone and the various growth forms.     

Anatomical and histochemical differentiation in lobes of the lichen Lobaria pulmonaria

The growth pattern of the thallus of Lobaria pulmonaria, a foliose lichen, has been described as differentiated into upwards and downwards growing lobes. The former show meristematic properties, whereas the latter, owing to the formation of soralia inactivating apical meristems, become senile lobes. A simple and sensitive histochemical procedure, the TBO test performed in this study, shows co-distribution of zones of active growth and polyphosphates accumulation in L. pulmonaria and gives evidence of the central role of phosphate in lichen metabolism. Upwards and downwards growing lobes mainly differentiate in pattern of accumulation of histochemically detectable polyphosphates. In the former, actively growing zones correspond to the algae, cortical and medullary hyphal cells adjacent to the algal layer of the pseudomeristematic marginal rim and of the adjacent elongation zone. In the downwards sorediate lobes, lacking apical growth, the actively growing zones correspond to medullary hyphae and algae occurring in areas where soredia are formed.     

Heat tolerance of two Cladonia species and Campylopus praemorsus in a hot steam vent area of Hawaii

Summary Temperatures were measured in soil, Cladonia skottsbergii, Cl. oceanica, and Campylopus praemorsus growing in the almost barren geothermal area at Puhimau, Hawaii. The measurements were made in the early morning in winter when insolation and air temperatures were minimal and the geothermal effects were predominant. Measurements were made on healthy, dew moistened plants. Close to steam vents Campylopus praemorsus forms thick cushions on hot soil and temperatures up to 29.8°C are recorded in the active parts of the moss. Cladonia oceanica grows exclusively on moss in this area, but not as close to steam vents as the moss itself. Maximum temperatures were 27.2°C in stunted and 23°C in ramified growth forms. In this area Cl. skottsbergii normally colonizes tree stumps of Metrosideros only where the steam is already cool. Maximum temperatures were 23°C in normal thalli, through higher temperatures were measured in partly damaged or killed thalli overhanging the stump where they are immersed in hot steam. With respect to heat tolerance only Campylopus can be considered as adapted to the hot environment. Therefore it is able to colonize the hot dry soil while deriving its moisture from adjacent steam vents. The lichens, particularly Cl. skottsbergii, are not adapted and are as sensitive to heat as most other lichens. Therefore they can only survive where there is at most a small geothermal impact yet they are obviously dependent on moisture from the steam vents.     

Features of the biomass distribution of epiphytic lichens on scotch pine (Lower Angara Region)

The biomass of epiphytic lichens growing on Scots pine varies from 130 to 1090 g and is composed mainly of lichens from three genera: Bryoria (45%), Hypogymnia (34%), and Evernia (12%). The majority of lichens (66%) grow on tree branches (96%) in the zone of maximum development, located at a height of 9–13.5 m. The lichen biomass on tree trunks is insignificant (4%) and located mainly at their bottom part (70%).     

Eine diözische Flechte

The lichenLecidea verruca, growing parasitically on other crustaceous lichens, is, contrary to most other lichenized ascomycetes, dioecious. The male thalli bearing spermogonia are mostly smaller than the female ones.

     

Estimating plant abundance using inflated beta distributions: Applied learnings from a lichen–caribou ecosystem

Quantifying abundance and distribution of plant species can be difficult because data are often inflated with zero values due to rarity or absence from many ecosystems. Terrestrial fruticose lichens (Cladonia and Cetraria spp.) occupy a narrow ecological niche and have been linked to the diets of declining caribou and reindeer populations (Rangifer tarandus) across their global distribution, and conditions related to their abundance and distribution are not well understood. We attempted to measure effects related to the occupancy and abundance of terrestrial fruticose lichens by sampling and simultaneously modeling two discrete conditions: absence and abundance. We sampled the proportion cover of terrestrial lichens at 438 vegetation plots, including 98 plots having zero lichens. A zero‐inflated beta regression model was employed to simultaneously estimate both the absence and the proportion cover of terrestrial fruticose lichens using fine resolution satellite imagery and light detection and ranging (LiDAR) derived covariates. The probability of lichen absence significantly increased with shallower groundwater, taller vegetation, and increased Sphagnum moss cover. Vegetation productivity, Sphagnum moss cover, and seasonal changes in photosynthetic capacity were negatively related to the abundances of terrestrial lichens. Inflated beta regression reliably estimated the abundance of terrestrial lichens (R² = .74) which was interpolated on a map at fine resolution across a caribou range to support ecological conservation and reclamation. Results demonstrate that sampling for and simultaneously estimating both occupancy and abundance offer a powerful approach to improve statistical estimation and expand ecological inference in an applied setting. Learnings are broadly applicable to studying species that are rare, occupy narrow niches, or where the response variable is a proportion value containing zero or one, which is typical of vegetation cover data.     

The importance of pre-mating barriers and the local demographic context for contemporary mating patterns in hybrid zones of Eucalyptus aggregata and Eucalyptus rubida

The frequency of hybridization in plants is context dependent and can be influenced by the local mating environment. We used progeny arrays and admixture and pollen dispersal analyses to assess the relative importance of pre‐mating reproductive barriers and the local demographic environment as explanations of variation in hybrid frequency in three mapped hybrid zones of Eucalyptus aggregata and E. rubida. A total of 731 open‐pollinated progeny from 36 E. aggregata maternal parents were genotyped using six microsatellite markers. Admixture analysis identified substantial variation in hybrid frequency among progeny arrays (0–76.9%). In one hybrid zone, hybrid frequency was related to pre‐mating barriers (degree of flowering synchrony) and demographic components of the local mating environment (decreasing population size, closer proximity to E. rubida and hybrid trees). At this site, average pollen dispersal distance was less and almost half (46%) of the hybrid progeny were sired by local E. rubida and hybrid trees. In contrast, at the other two sites, pre‐mating and demographic factors were not related to hybrid frequency. Compared to the first hybrid zone where most of the E. rubida (76%) and all hybrids flowered, in the remaining sites fewer E. rubida (22–41%) and hybrid trees (0–50%) flowered and their reproductive success was lower (sired 0–23% of hybrids). As a result, most hybrids were sired by external E. rubida/hybrids located at least 2–3 km away. These results indicate that although pre‐mating barriers and local demography can influence patterns of hybridization, their importance can depend upon the scale of pollen dispersal.     

Differential utilization of space and food by the inshore rockfishes (Scorpaenidae: Sebastes) of Carmel Bay,California

Synopsis Six rockfish species were found to be common in the Carmel Bay kelp forests. Five of these occupy spatial zones that are more or less distinct from one another. Three species (S. atrovirens, S. chrysomelas, and S. carnatus) occupy species-specific spatial zones while two others (S. serranoides and S. melanops), although spatially distinct from each other, occupy zones that overlap completely with a sixth species, S. mystinus. Food overlap values (PSI) indicate that S. mystinus has low competitive overlap with all other species, and hence can coexist with two species using the same habitat. Dietary arrays reflect the distinct spatial zones occupied by most of the rockfishes. Structural characteristics associated with feeding (maxillary, intestine, and gill raker length) suggest that these species are differentially adapted in regard to feeding morphology. The coexistence of these rockfishes appears to be the result of the partitioning of food and space resources, rather than the result of stochastic events which appear to be important in the structuring of some coral-reef fish communities. S. mystinus juveniles represent a major trophic link between adult piscivores and the planktonic organisms upon which the juveniles feed. A sustained crash in S. mystinus numbers might have a negative effect on abundance of kelp forest piscivorous species.     

Distribution and floristics of moss- and lichen-dominated soil crusts in a patterned Callitris glaucophylla woodland in eastern Australia

The distribution and abundance of soil crust lichens and bryophytes was examined in a patterned Callitris glaucophylla woodland in eastern Australia. Twenty-one lichen species and 26 bryophyte species were collected within thirty quadrats along a sequence of runoff, interception and runoff zones. Crust cover was significantly greatest in the interception zones (79.0 %), followed by the runoff zones (24.0 %), and lowest in the groved, runon zones (6.6 %). Lichens and bryophytes were distributed across all geomorphic zones, and, although there were significantly more moss species in the interception zones (mean = 9.1) compared with either the runoff (4.2) or runon (3.2) zones, the number of lichen species did not vary between zones. Ordination of a reduced data set of 32 species revealed a separation of taxa into distinct groups corresponding to the three geomorphic zones. Canonical correspondence analysis (CCA) of the 32 species and thirteen environmental variables revealed that the most important factors associated with the distribution of species were sheet and scarp erosion, soil stability and coherence, litter cover and crust cover. Surface cracking, microtopography and plant cover were of intermediate importance. The CCA biplot revealed that the timbered runon zones (groves) were dominated by `shade-tolerant' mosses Fissidens vittatus and Barbula hornschuchiana, whilst the heavily eroded runoff zones supported sparse populations of `erosion tolerant' lichens (Endocarpon rogersii) and mosses (Bryum argenteum and Didymodon torquatus). Interception zones supported a rich suite of `crust forming' mosses and lichens capable of tolerating moderate inundation by overland flow. Two other groups of taxa were identified by this analysis: the `pioneer' group, comprising mainly nitrogen-fixing lichens which occupy the zone of active erosion at the lower edge of the groves, and the `opportunists' dominated by liverworts, occupying the shallow depressions or bays at the margins of the groves and the interception zones. This study confirms that the non-vascular lichens and bryophytes in these arid soil crusts, are, like the vascular plants, strongly patterned according to geomorphic zone, being most strongly associated with soil surface and erosional features.     

Do co-occurring plant species adapt to one another? The response of <Emphasis Type="Italic">Bromus erectus</Emphasis> to the presence of different <Emphasis Type="Italic">Thymus vulgaris</Emphasis> chemotypes

Local modification of the soil environment by individual plants may affect the performance and composition of associated plant species. The aromatic plant Thymus vulgaris has the potential to modify the soil through leaching of water-soluble compounds from leaves and litter decomposition. In southern France, six different thyme chemotypes can be distinguished based on the dominant monoterpene in the essential oil, which is either phenolic or non-phenolic in structure. We examine how soils from within and away from thyme patches in sites dominated by either phenolic or non-phenolic chemotypes affect germination, growth and reproduction of the associated grass species Bromus erectus. To do so, we collected seeds of B. erectus from three phenolic and three non-phenolic sites. Seeds and seedlings were grown on soils from these sites in a reciprocal transplant type experiment in the glasshouse. Brome of non-phenolic origin performed significantly better on its home soil than on soil from a different non-phenolic or a phenolic site. This response to local chemotypes was only observed on soil collected directly underneath thyme plants and not on soil in the same site (<5 m away) but where no thyme plants were present. This is preliminary evidence that brome plants show an adaptive response to soil modifications mediated by the local thyme chemotypes. Reproductive effort was consistently higher in brome of phenolic origin than in brome of non-phenolic origin (on both thyme- and grass-soil), indicating that life-history variation may be related to environmental factors which also contribute to the spatial differentiation of thyme chemotypes. Moreover, we found that brome growing on thyme-soil in general was heavier than when growing on grass-soil, regardless of the origin of the brome plants. This is concordant with thyme-soil containing higher amounts of organic matter and nitrogen than grass-soil. Our results indicate that patterns of genetic differentiation and local adaptation may modify competitive interactions and possible facilitation effects in natural communities.     

Differential effects of reindeer on high Arctic lichens

Abstract. We studied the effects of Svalbard reindeer on the abundance of lichens in Spitsbergen. A survey was carried out in 14 areas with contrasting reindeer densities. Separate cover estimates for crustose, fructose and foliose lichens were taken in each area, and related to the density of reindeer pellet groups, a measure of reindeer density. Dominant macro lichen families were identified in 10 areas, and a full record of macrolichen species was taken in four additional areas. Variation in reindeer density is partially due to past overhunting, and subsequent incomplete recovery, releasing some areas from reindeer grazing for 100–200 yr. The cover of fruticose lichens was negatively related to reindeer pellet group density, indicating suppression by Svalbard reindeer. This makes their impact comparable to other members of the Rangifer genus around the northern hemisphere. The generally recorded low abundance of lichens in the diet of Svalbard reindeer compared to other Rangifer species, therefore, was interpreted as the depletion of fruticose lichens in Spitsbergen, and a subsequent switch to alternative foods. Of all fruticose lichens, Stereocaulon spp. appeared least sensitive to grazing. Crustose and foliose lichen cover was independent of reindeer pellet group density. The cover of crustose lichens was significantly related to latitude, with greater cover in more northern areas. Foliose lichens were more abundant in places where moss cover was high. We conclude that the impact of Svalbard reindeer on lichens is dependent on growth form, with fruticose lichens suffering from grazing, whereas foliose lichens might indirectly benefit from higher densities of reindeer or, like crustose lichens, be controlled by other factors.     

EVIDENCE FOR COMPETITIVE RELEASE IN SIMPLIFIED SAXICOLOUS LICHEN COMMUNITIES

Saxicolous cryptogam communities were sampled quantitatively on two Potomac River island habitats in June of 1980. As a result of air pollution stress, the community on one island was found to be much simpler than that found on the other island. Two relatively pollution-tolerant foliose lichen species, Xanthoparmelia conspersa and Pseudoparmelia baltimorensis, were found to dominate the communities on both islands. An examination of niche breadth and position along a light intensity gradient demonstrated that P. baltimorensis was most frequent at low light intensities and X. conspersa was most frequent at high light intensities. However, X. conspersa exhibited a niche shift toward intermediate light intensities in the species-poor community, likely the result of reduced competitor diversity in this community since the intermediate light intensities supported the greatest number of species in the species-rich community. Niche overlap between the two species was found to occur at lower light intensities in the species-poor community, suggesting that X. conspersa is more likely to appropriate intermediate light intensities in this community. The significance of competition as a regulatory factor in lichen communities has not been fully appreciated. Since lichens and other cryptogams are frequently used as indicators of environmental quality, competitive processes in naturally occurring lichen communities must be thoroughly understood before changes in species distribution that apparently result from disturbance can be properly interpreted.     

Nivalenol and 15‐acetyldeoxynivalenol Chemotypes of Fusarium graminearum Clade Species are Prevalent on Maize throughout China

Fusarium graminearum clade species are among the main causative agents of Gibberella ear rot (GER) in maize and responsible for the various trichothecene mycotoxins accumulated in contaminated maize grains. In this study, a total of 620 isolates from diseased maize ears collected from 59 districts in 19 provinces throughout China, previously identified morphologically as Fusarium graminearum clade, was genetically characterized at the species level based on SCAR (Sequence Characterized Amplified Region) and for their potential capability of mycotoxin production using the genetic chemotyping assay. The results showed that 359 isolates were F. asiaticum (SCAR 5), which consisted of 97% nivalenol (NIV)‐chemotypes, 0.8% 3‐acetyldeoxynivalenol (3‐ADON)‐producing isolates and 2.2% 15‐acetyldeoxynivalenol (15‐ADON) producers, whereas the remaining 261 isolates were identified as F. graminearum sensu stricto (SCAR 1), all of which produced 15‐ADON mycotoxins. This high proportion of NIV producers present in F. asiaticum is different from the chemotype patterns in F. asiaticum populations isolated from wheat and barley, where DON and its acetylated chemotypes were the predominant mycotoxins. Moreover, the majority of NIV producers (59.1%) and all the 3‐ADON‐producing strains were derived from the warmer regions in southern China, whereas most of the 15‐ADON‐producing strains (78.4%) were isolated from the colder regions in northern China. Our study is the first report of NIV chemotypes of F. asiaticum and 15‐ADON chemotypes of F. graminearum sensu stricto that were associated with the GER of maize in China.     

Identification of the Chemotypes of Ocimum forskolei and Ocimum basilicum by NMR Spectroscopy

The chemotypes of Ocimum forskolei Benth and Ocimum basilicum L. growing wild in Oman have been established by ¹³C‐NMR analyses of the vegetative and floral oils of the plants. The chemotypes, estragole for O. forskolei and linalool for O. basilicum, suggested by ¹³C‐NMR fingerprinting were also confirmed by GC‐FID and GC/MS analyses. The oil of O. forskolei demonstrated better activities against bacteria and dermatophytes. The significance of the presence of estragole and linalool in the volatile oils of plants whose fragrances are traditionally inhaled, added to food, or rubbed on the skin are discussed.     

Genetic control of geranial formation inPerilla frutescens

A novel chemotype (C type) having a lemon-like odor segregated out in the F₂ progeny of a cross between PK and PL chemotypes ofPerilla frutescens. Chemical analysis of C-type plants revealed that geranial was the major component of essential oils in the leaves. Genetic analysis suggested that geranial is accumulated by individuals homozygous for two pairs of recessive, polymeric genes,fr ₁ andfr ₂, which are incapable of converting geranial into perillene.