Reproductive traits in the tropical moss Octoblepharum albidum Hedw. differ between rainforest and coastal sites

Abstract

Octoblepharum albidum Hedw. is an autoicous moss commonly occurring in tropical savannas, dry forests, rainforests and coastal habitats. It frequently reproduces by spores and asexual structures (gemmae and protonemata or buds at leaf tips), making it a good model for understanding how reproductive traits change with respect to habitat type. Our aims were to characterize the different life-history traits in O. albidum relative to sexual and asexual cycles and to detect variations in reproductive performance among the different habitats, trade-offs among these traits, and relationships among reproductive traits and plant length. We studied colonies from two Atlantic rainforests and two coastal sites in north-eastern Brazil. Shoots in the coastal sites, compared to those of the forest sites, had higher numbers of sporophytes, male and female branches per shoot, male gametangia per sexual branch, and longer setae. Numbers of female gametangia per sexual branch did not differ between forest and coastal sites. A male-biased sex ratio of branches and gametangia occurred in all sites. Compared to gemmae, sporophytes and protonemata or buds were more likely to be found on longer shoots than on shorter ones, but this relationship was only applicable to forest sites. The abundant production of gemmae and protonemata or buds at leaf tips, and sporophytes (spores) in O. albidum are important components in explaining colonization success and maintenance in this tropical moss. Longer sporophytic setae in addition to a higher reproductive performance (especially for number of male gametangia and sporophytes per shoot) may favour spore dispersal and colonization in plants of coastal sites.     

Sexual reproduction in the phyllodioicous bryophyte Homalothecium lutescens (Hedw.) H.Rob. in relation to habitat age,growth conditions and genetic variation

Sporophyte production and female fertility were investigated in seventeen calcareous grassland demes of the moss Homalothecium lutescens (Hedw.) H.Rob. on the Baltic island of Öland, with the aim of understanding the relationships between sexual reproduction, habitat age, genetic variation and factors related to growth conditions. The overall proportion of fertile female shoots (with perichaetia) was 35%. Fertility status at the level of individual shoots was positively associated with shoot length and density, while within deme fertility was positively associated with bush cover. There was no association between female fertility and habitat age, genetic diversity (H_S) or allelic richness. Out of 1344 investigated shoots, only two were normal-sized fertile males. Dwarf males were also extremely rare, and found almost exclusively on shoots with sporophytes. Few sporophytes were observed (in the two demes with highest fertility and bush cover). No relationship between genetic variation and the frequency of sporophytes and males was found. The lack of a relationship between sexual reproduction and genetic variation suggests that sexual reproduction may not occur in the same grassland fragments as the recruitment of new clones (from spores or vegetative fragments). The majority of the dry, open grassland habitats, where H. lutescens is typically found in the study area, appear to be suboptimal for both dwarf males and fertilization. Sexual reproduction is more likely to occur in shaded (although grazed) grassland patches, where moisture levels are likely to be higher and the moss colonies are generally more vigorous.     

Notes on sporophytic details of Aneura crateriformis (Aneuraceae,Marchantiophyta)

Abstract

Pohlia nutans subsp. schimperi is recognized as a distinct taxon, differing from ssp. nutans by the pink to purple coloration of its leaves and a dioicous or polyoicous, rather than paroicous, sexual condition. We review the contrasting reports about the sexuality of these plants, as well as their nomenclatural history, and demonstrate their synonymy with P. nutans subsp. purpurascens Latzel. The subspecies shows a mainly arctic–subarctic distribution, with relict Central European occurrences in the Eastern Alps, the Sudetes and the western Carpathians, probably reflecting migratory events during the Pleistocene. The ecology and phytosociological affinities of the taxon in Central Europe are discussed in detail.     

Two new species of Distichophyllum from Malaya

Abstract

The germination and establishment abilities of different types of propagules under natural conditions is an area in bryophyte reproductive biology that is largely unknown. In some species gametophyte fragments, rather than spores, may be primarily responsible for establishment of new colonies in nature, and the presence of other mosses may have an effect on establishment processes. These ideas were tested experimentally using four species of Tortula grown in a glasshouse under semi-natural conditions. The relative germination and growth abilities of spores and gametophytic fragments were tested under each combination of the following treatments: (1) continuous hydration (using a mist system), or periodic drying; (2) substrate of bare sand, a clump of Dicranum scoparium, or (for spores only) the clump of Tortula from which sporophytes were taken. Fragments produced new shoots much more quickly than spores under either hydration condition but spores germinated at a higher rate. Germination rates of both spores and fragments were greatly reduced (usually to zero) in clumps of Tortula or Dicranum under either hydration condition. Some differences between species were noted. In a further experiment filter-sterilized water extracts of each species were applied to spores on agar. Again, strong effects on spore germination and protonemal growth were noted. Patterns of inhibition differed between species and reflected those found in the previous experiment, indicating that the inhibition effect may be due to a water soluble chemical rather than the physical properties of the moss clumps. These studies present the first experimental evidence for the chemical inhibition of spores by mature plants. When taken together with previous reports of chemical interactions among protonemal systems, considerable ecological significance of these results is suggested.     

The fall and rise of the Oblong woodsia in Britain

Summary

The vegetation of a small spoil heap at Jerah mine near Stirling was classified as species-rich acid grassland, an open barren-soil community with a high frequency of the moss Pohlia nutans, and a group which showed intermediate characteristics. Copper was present in the soil in potentially toxic concentrations and when the vegetation data were ordinated the axes showed highly significant correlations with soil copper and loss-on-ignition. The results suggest that copper is a major determinant of the spoil-heap vegetation, and contrast sharply with those reported for the Burn of Sorrow mine, a site with many superficial similarities.     

Deposition of moss spores in relation to distance from parent gametophytes

Abstract

Daily counts were made of spores trapped on microscope slides coated with petroleum jelly and placed on the ground at distances up to 200 em from isolated colonies of Atrichum undulatum and Bryum argenteum. The observations continued for 30–34 days during the annual period of spore release in these species. Estimates of the number of spores released by the colonies were obtained as the product of the numbers of capsules and the difference between the mean numbers of spores present in undehisced capsules at the beginning of the experiment and remaining in dehisced capsules at the end.

Spore deposition over the experimental periods showed a steep gradient, deposition per unit area falling from 4740—14, 230 spores cm_?2 in the centre of the colonies to less than 10 spores cm_?2 200 cm from the edge of the colonies. Despite the steep deposition gradients, however, it was estimated that more than 85% of the spores in A. undulatum, and more than 95% of those in B. argenteum, were dispersed to unknown distances beyond the trapping areas.     

Large‐scale changes in abundance of terricolous bryophytes and macrolichens in Finland

Abstract. Major changes in forest floor vegetation were identified on the basis of three nationwide surveys conducted as part of national forest inventories in 1951–1953, 1985–1986 and 1995. These surveys provided objectively selected, statistically representative samples of all forested land in Finland. The 1951–1953 data consist of over 10000 sample plots, while the later surveys were conducted on ca. 3000 permanent plots. Changes in relative abundance of dominant species (i.e. in the proportions of species of the total cover of forest floor vegetation) were analysed across biogeographical provinces. Spatial correlation, systematic sampling, partial re‐measurement and multiple testing were taken into account in assessment of the statistical significance of the observed changes. The most notable changes in forest floor vegetation were a decrease in the relative abundance of Hylocomium splendens and an increase in Dicranum polysetum. In N Finland, where forests are grazed by semi‐domestic reindeer, we observed a decline in the abundance of Cladina lichens and an increase in Dicranum mosses. Peltigera aphthosa declined throughout the country. Polytrichum juniperinum, Pohlia nutans, and Brachythecium species, which occupy disturbed sites or grow on litter, increased in abundance. The relative abundance of Sphagnum species decreased, particularly in W Finland, where Pleurozium schreberi was favoured. A major decline in S. fuscum was also recorded in C and E Finland. Many of the changes detected in this study are apparently related to intensified forest management; but solely on the basis of this study, its effects cannot be distinguished from those of other large‐scale environmental changes.     

Effects of reindeer grazing on understorey vegetation in dry Pinus sylvestris forests

Abstract. Data on floristic composition and environmental variables were collected in floristically homogeneous oligotrophic pine (Pinus sylvestris) forests with heath-like under- storey vegetation in eastern Fennoscandia, and ordinated by non-linear multidimensional scaling (NMDS) in order to study the effect of lichen grazing by reindeer on the understorey vegetation. The study sites included areas with varying grazing pressure, as well as 50-yr old grazing exclosures. Sites rich in respectively bryophytes and lichens were placed at opposite ends of the ordination axes, and heavily grazed sites were placed in between them. Reindeer grazing increased the abundance of bryophytes, especially Dicranum spp. and Pleurozium schreberi. Grazing changed the vegetation to the extent that it resembled more mesotrophic sites, but this did not show any relationship with tree volume or other site productivity indicators. This was observed both in the ordination and, in a more compelling way, when exclosures with adjacent grazed areas were compared. No such signs were evident at ungrazed sites, where especially Cladina spp. spatially replace Cladonia spp. and tiny bryophytes like Barbilophozia spp., Polytrichum spp. and Pohlia nutans during succession. Cladina stellaris had almost disappeared from the most intensively grazed sites. The soil at ungrazed sites was characterized by high Al and Fe concentrations and bryophyte-rich sites by high Mn concentrations. Shannon's diversity index, depth of humus layer and proportion of bare ground also increased in sites getting richer in bryophytes.     

HAPLOID PARTHENOGENETIC SPOROPHYTES OF LAMINARIA JAPONICA (PHAEOPHYCEAE)1

Parthenogenetic sporophytes were obtained from three strains of Laminaria japonica Areschoug. These sporophytes grew to maturity in the sea, producine spores that all grew into female gametophytes. These female gametophytes gave rise to another generation of parthenogenetic sporophytes during the next year, so that by the year 1990 parthenogenetic sporophytes had been cultivated for 12, 9, and 7 generations, respectively, for the three strains. When female gametophytes from parthenogenetic sporophytes were combined with normal male gametophytes, normal sporophytes that reproduced and gave rise to both female and male gametophytes were obtained. The parthenogenetic sporophytes were shorter and narrower than the normal sporophytes of the same strain. Chromosome counts on mature sporophytes showed that normal sporophytes (from fertilized eggs) were diploid (2n = approximately 40) and that the spores they produced were haploid (n = approximately 20), while nuclei from both somatic and sporangial cells in parthenogenetic sporophytes were haploid. All gametophytes were haploid. Young sporophytes derived from cultures with both female and male gametophytes were diploid, while young, sporophytes obtained from female gametophytes from parthenogenetic sporophytes had haploid, diploid, or polyploidy chromosome numbers. Polyploidy was associated with abnormal cell shapes. The presence of haploid parthenogenetic sporophytes should be use in breeding kelp strains with useful characteristics, since the sporophyte phenotype is expressed from a haploid genotype which can be more readily selected.     

Genetic diversity in the moss Pohlia nutans on geothermal ground of Mount Rittmann, Victoria Land, Antarctica

The only known population of the moss Pohlia nutans in continental Antarctica occurs on geothermally heated ground of volcanic Mt. Rittmann in northern Victoria Land. Colonisation by this bryophyte is due to peculiar environmental characteristics of the geothermal ground, because mosses do not normally grow in Antarctica at such elevations. Specimens from several moss patches within two sites of 80 m² total area were analysed genetically to determine whether they all originated from a single colonisation event, whether the population is genetically diverse, and whether the temperature range of geothermal ground (17-35°C under moss colonies) affects rates of mutation. Both the RAPD technique and DNA sequencing of the conserved nuclear ribosomal RNA 18S-26S ITS region were used to compare this extremely isolated population with specimens of P. nutans from elsewhere in Antarctica. Like the moss Campylopus pyriformis on volcanic Mt. Melbourne, the Pohlia population exhibits low levels of genetic diversity and appears to be derived from a single immigration event followed by vegetative growth, mutation and dispersal.     

Pohlia integra (Card.) Shaw a neglected species,recorded from southern Africa and South America

Abstract

Mniobryum bracteatum Bartr. from Ecuador is reduced to a synonym of Pohlia integra (Card.) Shaw, previously known only from Mexico. The species, characterized by the occurrence of rhizoidal tubers, is also recorded from Bolivia and southern Africa.     

Reproductive modes in the epixylic hepatic Anastrophyllum hellerianum

The frequency and dynamics of sexual and asexual reproduction were investigated in a dioecious epixylic hepatic, Anastrophyllum hellerianum, which has declined in recent decades in Finland as a consequence of forestry practices. In our investigation asexual reproduction by gemmae was the dominant mode of reproduction and specialised gemmiparous shoots were present in all colonies studied. The proportions of dead shoots were considerably higher among sex-expressing than among non-sexexpressing shoots. Our results suggest that lower reproductive investment is required for asexual than for sexual reproduction. For instance, no trade-off is detected between asexual reproduction and survival of the gemmiparous shoots in A. hellerianum. Sexual reproduction occurred only in 12% of the colonies and it was promoted by the following factors: medium shoot density, high proportion of sex-expressing shoots, an even sex ratio and very short distances between individuals representing opposite sexes. The ratio of dead males to dead females was significantly female-biased, which suggests higher mortality among female shoots. At the level of individual shoots, more spores than gemmae were produced. However, as a consequence of the low frequency of sporophyte-bearing shoots, gemma production highly exceeded spore production at the colony level. Furthermore, cultivation tests of the propagules showed that gemmae germinate faster than spores.     

Costs of sporophyte production in the moss,Dicranum polysetum

We investigated the cost of sporophyte production in the moss Dicranum polysetum both by examining patterns of growth and reproduction in unmanipulated shoots and by experimentally manipulating sexual reproduction.The estimated proportion of total carbon investment allocated to sexual reproduction in sporophyte-producing shoots over the study period was 74.8%. Unmanipulated shoots that aborted all sporophytes had a significantly higher growth in the top shoots than shoots that produced sporophytes. In sporophyte-producing shoots, total apical growth decreased proportionately with the number of sporophytes.Experimental prevention of sporophyte development resulted in significantly higher total apical growth of the gametophytes. Shoots where current perichaetia were lacking when marked had a mass increase in the top shoots similar to manipulated sporophytic shoots whereas sporophytic control shoots grew significantly less than these two categories.The difference between control shoots and manipulated shoots in the mass of vegetative apical growth was mainly because of different length increments whereas mass per unit length was similar between groups. The probability to reproduce sexually in the year after the manipulation, and the biomass allocated to this reproduction, were not affected by the experimental treatment.     

The occurrence of polysety in relation to the number of archegonia in the female inflorescences of Phascum cuspidatum Hedw.

Abstract

In Phascum cuspidatum the number of sporophytes per inflorescence was found to increase, up to a certain limit, with increase in the number of archegonia present. The number of archegonia present varies with the age of the inflorescence, and fertilization terminates the production of archegonia. The common occurrence of solitary sporophytes is due to fertilization at a time when only a small number of archegonia is present in most inflorescences. Delayed fertilization, leading to an increase in the average number of archegonia per inflorescence, is a necessary precondition for increased occurrence of polysety. As many as five sporophytes in a single inflorescence formed capsules with spores without significant reduction in the size of capsules. Variation in the frequency of abortive embryos gave evidence of competition amongst sporophytes and this was most severe in the smaller inflorescences. The occurrence of polysety in Tortula muralis appears to be governed by the same rules.     

A Bryophyte flora of Kent. I

Abstract

Gender expression and sporophyte frequency of Frullania tamarisci were compared between epiphytic woodland and exposed coastal colonies in an Atlantic region of western Britain. Corticolous colonies exhibited very high sporophyte frequency in mixed-sex colonies. Samples from well-grown epiphytic patches commonly had intimate mixtures of highly fertile female and male shoot networks, and fertilisation reduced gynoecia number. In marked contrast, colonies in exposed coastal habitats were found to be sub-fertile or non sex-expressing and sporophytes were rare. There was a distinct female bias among colonies on the coast where spatial segregation of the sexes and male scarcity appear to limit sporophyte frequency. It is suggested that the close association of males and sporophytic females in epiphytic colonies may be influenced by within-colony sporophyte frequency.     

念珠丝瓜藓(缺齿藓科)形态特征及其在中国的新分布

通过对中国产丝瓜藓属Pohlia(缺齿藓科Mielichhoferiaceae)植物系统地分类学研究,对本属一具有假根生念珠状排列的芽胞的种——念珠丝瓜藓的形态解剖学特征和地理分布进行了修订,并讨论了念珠丝瓜藓的生境,以及与其相近种泛生丝瓜藓、明齿丝瓜藓和黄丝瓜藓等进行了比较分析。根据标本信息,确定念珠丝瓜藓在中国的分布由原来的5省区扩大到10省区,并绘制了该种在中国的分布图。     

Three propaguliferous species of Pohlia (Mielichhoferiaceae,Musci) in China

Abstract

As a result of a literature survey and examination of specimens, we believe that the identification of three species of propaguliferous Pohlia in China, Pohlia flexuosa, Pohlia proligera, and Pohlia leucostoma needs to be clarified. Pohlia flexuosa has two differing types of propagule: one short and oblong and the other long and thread-like; both propagules have inconspicuous leaf primordia, and can occur on the same stem. Propagules of P. proligera are shortly clavate to long vermicular, with one to three unicellular or multicellular prong-like leaf primordia arising at the apex of the propagule, and the leaf primordia are approximately one-fourth of the total length of the propagules. Pohlia leucostoma has long filiform propagules, with two or three short peg-like leaf primordia about one-tenth of total propagule length.     

The effect of fertilization and irrigation on the vegetation dynamics of a pine-heath ecosystem

The year-to-year changes in the field-and bottom layers were studied by non-destructive recording of cover percentage on permanent plots in a young Scots pine (Pinus sylvestris) stand, subjected to experimental treatment. The treatments were: control (0), daily irrigation (I), fertilization once a year (F) and daily irrigation plus fertilization five days a week (IF). The cover of the total field layer of the 0-, I-, F-and IF-plots increased during the period of study from 51, 40, 47 and 50% in 1973 to 70, 79, 86 and 92% in 1980. The bulk of this increase consisted of Vaccinium vitis-idaea with the exception of the IF-plots, where the increase especially during the later part of the period of study was due to Chamaenerion angustifolium and and to some extent, to Rubus idaeus. The cover of the total bottom layer of the 0-, I-, F-and IF-plots changed from 63, 59, 54 and 41% in 1973 to 68, 90, 37 and 36% in 1980. The most drastic change took place in the IF-plots, where the composition changed from almost exclusively lichens to bryophytes. Pohlia nutans and Pleurozium schreberi were recorded in substantial amounts filling the gap left by the vanishing lichens. There were no conclusive changes on the I-plots as compared with the 0-plots. The cover estimates, however, were generally more substantial. The overall decline in the bottom layer on the F-plots was mainly due to a decrease in the cover of Pleurozium schreberi.The work was carried out within the Swedish Coniferous Forest Project and supported by the Swedish Natural Research Council, the Swedish National Environmental Protection Board, the Swedish Council of Forestry and Agricultural Research and the Wallenberg Foundation.Nomenclature follows Lid (1974) for vascular plants, Nyholm (1954–69) and Arnell (1956) for bryophytes, Dahl & Krog (1973) for lichens, except for Lecidia granulosa, and the collective group Cladonia silvatica, where Magnusson (1929, 1952) was followed.I wish to thank J. G. K. Flower-Ellis for taking an active part in the planning of this work; S. Bråkenhielm for help and co-operation in the field; H. Sjörs, C. O. Tamm and many colleagues in the Swedish Coniferous Forest Project, among them F. Andersson, A. Aronsson & B. Axelsson for encouragement and support.     

Induced apogamous sporophytes inPteris dispar andP. semipinnata,and the meiotic behavior in their sporocytes

Spores ofPteris dispar andP. semipinnata were aseptically cultured in flasks for apogamous sporophyte induction. Calli or cell colonies similar to calli were induced in cultures supplemented with hormones. Sporophytic leaves subsequently developed from them in hormone-free medium and the young sporophytes were raised into plants with sporangia. Since the wild-type plants having 116 chromosomes are tetraploid, the sporophytic plants originating from spores would appear to be diploid (dihaploid). In induced sporophytes ofP. semipinnata, non-homologous chromosomes (58 univalents) were found during the meiotic process in sporocytes. InP. dispar, however, the meiotic cells showed many bivalent chromosomes (maximum 29ll). These results suggest thatP. semipinnata is allotetraploid, whereasP. dispar is autotetraploid.     

Reproductive biology of Syzygiella rubricaulis (Nees) Steph. (Adelanthaceae,Marchantiophyta), a liverwort disjunctly distributed in high‐altitude Neotropical mountains

Syzygiella rubricaulis is a dioecious leafy liverwort disjunctly distributed and restricted to high‐altitude mountains in the Neotropics and the Azores. This study is part of a larger project examining the phylogeography of S. rubricaulis in the Neotropics, and our main goals were to understand its reproductive biology, where sex expression occurs, if vegetative propagules are frequently found, how the sexes are distributed in populations, how frequently sporophytes are formed and what environmental conditions influence sexual expression. S. rubricaulis patches are mostly female, but all patches also contain non sex‐expressing shoots. Out of 42 patches examined, 29 (69%) were sex‐expressing: 25 were unisexual (21 female and four male) and four of mixed sex (two male‐biased and two unbiased). At shoot level, out of 4200 shoots 18% were female and 7% male; among sex‐expressing shoots, 73% were female, representing a sex ratio of 0.8 (female‐biased). We encountered a total of 33 sporophytes in six patches (in Brazil, Venezuela and Ecuador). Leaf regenerants were found in one patch in Mexico. Low rates of sporophytes were likely related to low frequencies of male shoots and large distances between the sexes. As 25% of S. rubricaulis shoots expressed sex (occasionally producing sporophytes), we suggest that short‐distance (and rarely long‐distance) spore dispersal events occur in mountainous areas on a short‐term basis. On a long‐term basis, however, these events likely contribute to dynamic exchanges among populations in the Neotropics.