Recent Advances in Bryological Research

In the last two decades, all the branch fields of bryology, with classical taxonomy gradually developing into multidisciplinecourses, have witnessed a great progress. The author is attempting to give a brief view of bryology in the following five paragraphs. The first part introduces the fundamental characteristics of bryology in the recent decades. Publication of “New Manual of Bryology” Vol. I and II, edited by Schuster(1983-1984) , marked a new epoch of bryology. A series of books, “The Biology of Mosses” (Richardson, 1981), the “New Advances in Bryology “and the other bryological publications showed that bryologists were working not only on bryofloras in different regions of the world, but also deeply engaged in the micro-view fields of bryology. Evidently, the world bryological researches have entered a prosperous period. The second part gives some recent examples of investigations on taxonomy, morphology, cytology, paleobotany, ecology, reproductive biology and ecological physiology of bryophytes. In discussing the tendency of the future research, the author emphasizes that systematic and evolutionary botany of bryophytes is one of the main subjects .Following the increase of taxa, the development of chemotaxonomy, cytology, numerical taxonomy etc.will reveal the relationships between orders, families or infrafamiliar groups. However, the main problem in this respect is lacking of key fossil records of bryophytes. On the other hand, the monographs increase steadily, and they are helpful in correcting some previously reported disjunct distribution of species and distribution of regional new species. Plant geography of bryophytes can also be used for explaining the continental drift (Schuster 1969, 1972).Moss indicators, which are more sensitive than vascular plants, have been noticed. The traditional utilization of mosses will be continuous. Chinese bryology has been greatly improved in the past half a century, and about a hundred papers involving taxonomy, phytogeography, phytocoenology, morphology, cytology, numerical taxonomy and applied bryology, were published in last two decades. In China, the foundation of taxonomy of bryophytes is developing, although we already have several bryological research centers. It seems to me that too many new taxa have been described, while no sufficient attention has been paid on studies on relationships and evolution of bryophytes. Due to identification of bryophytes, hryofloras between some neighbouring regions are not comparable. Fossils of bryophytes are usually treated as the “unknown kind of plants”, so we lack the geological evidence of bryophytes. We are facing the problem in training a new generation of bryologists due to the limited funds. Finally, some suggestions are made for Chinese bryology. The ways for seeking the origin of bryophytes will be: l.Searching for primitive groups of bryophytes.2.Comprehensive studies on key genera and species.3.Expeditions to the unknown native localities of special groups of bryophytes. Further studies on the bryoflora and phytogeography are necessary in China. Studies on the distribution centers of genera endemic to China and East Asiatic endemic genera in China will provide some pieces of evidence on the origin oe the Chinese bryoflora. New records of macrofossils and microfossils of bryophytes will help to break the “neck of bottle “ of the above-mentioned problems. Experimental projects are almost unknown in China. lt is very important, therefore, to encourage people to work on them for original data on phylogeny and origin of bryophytes, although it will be a long-term task inChina.     

哀牢山湿性常绿阔叶林地生、树干及树枝 附生苔藓生活型组成及其水分特性

为探讨林内不同垂直高度生境下及不同生活型苔藓植物水分特征的差异,该文对哀牢山湿性常绿阔叶林内地生苔藓、林下树干附生和林冠层树枝附生苔藓优势或常见种的生活型组成、持水力、失水特征和水分利用效率进行了研究。结果表明:地生、树干附生和树枝附生苔藓植物类群分别以交织型、扇型、悬垂型苔藓植物占据优势;地生、树干附生、枝条附生苔藓植物的饱和持水率分别为476%DW、210.98%DW、238.95%DW;地生苔藓植物的持水率和失水速率均高于附生苔藓,树干附生苔藓植物持水率低于树枝附生苔藓,而失水速率高于树枝附生苔藓。在不同生活型苔藓的水分特性上,交织型苔藓具有较高的持水率和失水速率,保水性能较弱,其次为悬垂型苔藓,扇型苔藓的持水率最小,失水速率也较快;地生和树干附生苔藓的水分利用效率均显著高于树干附生苔藓,交织型和扇型苔藓的水分利用效率显著高于悬垂型苔藓。在三种生境下,地生苔藓持水力高,水分利用效率较高而保水能力低;枝条附生苔藓持水力低,水分利用效率低而保水能力较高;树干附生苔藓水分利用效率较高而持水力和保水能力均较差。因此,不同生境下苔藓植物生活型组成及其水分变化特性在一定程度上反映了它们对不同生境的适应策略。     

Book Review

Abstract

Most studies on population ecology of bryophytes have involved common species. However, some studies have compared life history strategies in rare and common species. We review the life history strategies (life expectancy, sexual and asexual reproduction, spore production, spore size and dispersal) for species that are rare in relation to pattern and persistency of suitable habitat patches. In particular, we discuss the dynamics on two levels, within and among localities, for different categories of rare species. We predict that most rare species will be found to have restricted dispersal capacities but higher than average life expectancies of local subpopulations. Natural rarity is distinguished from human-induced rarity and species rare for the latter reason are distinguished as 'threatened'.     

Bryophyte and lichen diversity: A comparative study

Abstract We describe the regional species richness, variation in species richness and species turnover of bryophytes and lichens from 36 sites in lowland forests of southeastern Australia. The analyses subdivided the two major taxa into their constituent sub-groups: mosses, liverworts, and crustose, fruticose and foliose lichens. They also explored correlations between selected environmental variables and patterns of diversity. On a regional scale, there were 77 species of bryophytes and 69 species of lichens, giving a total of approximately one-third of the total number of vascular plant species in the region. Mean species richness was higher for lichens than bryophytes. Also, the two taxa were negatively correlated because lichens favoured dry sites and bryophytes favoured moist ones. Species turnover was greater for bryophytes than lichens, largely due to the distribution of liverwort species. Foliose lichens showed higher levels of turnover than crustose lichens. Multiple regression and canonical correspondence analysis showed that both taxa and all sub-groups responded to the same three variables: vascular plant cover, time since last fire and topographic position. Other variables, including time since logging and intensity of logging, explained little variation in bryophyte or lichen diversity. The data suggest that the strategies for the conservation of bryophyte and lichen biodiversity will be different, to reflect the different patterns of species richness and species turnover.     

Atlantic bryophytes in Scotland

Summary

A brief overview is given of the importance of Scotland for Atlantic bryophytes within Europe and worldwide. The life strategies of Atlantic species are discussed and the main habitats of Atlantic bryophytes listed. A consideration of the use of distribution data to identify ‘hotspots’ and sites for conservation is made. Grimmia britannica A.J.E. Smith and Marsupella profunda Lindb. are added to the list of Atlantic species in Britain and Ireland.     

浙江西天目山苔藓植物物种多样性的研究

对浙江西天目山不同海拔苔藓植物进行调查,从种类组成、相似性、α多样性和β多样性等方面进行了苔藓植物物种多样性分析.结果表明,海拔1 100 m处落叶阔叶林下的苔藓植物种类最多,物种丰富度最高;它与海拔1 300 m落叶矮林下苔藓植物的相似性系数也最高;而海拔800～1 100 m之间的β多样性最大,这一区段苔藓植物的物种变化速率最快,种类更替最明显.西天目山海拔1 100 m处苔藓植物物种多样性最高,是该山体苔藓植物多样性保护的关键区段.     

A Bryophyte Flora of Berkshire and Oxfordshire

Abstract

In temperate North American forests, collections were made of myxomycetes (plasmodial slime molds) for which bryophytes served as the primary substrate for fruiting. Most associations among the 52 myxomycete species and 55 bryophyte species, represented in these collections, appeared to be coincidental but two species of myxomycetes (Barbeyella minutissima and Lepidoderma tigrinum) were evidently bryophilous, preferring leafy hepatics on rotten coniferous logs. Bryophytes obviously provide exposed surfaces convenient for myxomycete sporulation. Whether bryophytes also sustain the feeding phases of the myxomycete life-cycle (swarm cells, myxamoebae, and plasmodia) is not yet known.     

Little,but increasing evidence of impacts by alien bryophytes

Based on data of bryophyte invasions into 82 regions on five continents of both hemispheres, we aim here at a first comprehensive overview of the impacts that bryophytes may have on biodiversity and socio-economy. Of the 139 bryophytes species which are alien in the study regions seven cause negative impacts on biodiversity in 26 regions, whereas three species cause negative impacts on socio-economic sectors in five regions. The vast majority of impacts stem from anecdotal observations, whereas only 14 field or experimental studies (mostly on Campylopus introflexus in Europe) have quantitatively assessed the impacts of an alien bryophyte. The main documented type of impact on biodiversity is competition (8 alien bryophytes), with native cryptogams being most affected. In particular, C. introflexus (9 regions) and Pseudoscleropodium purum (7 regions) affect resident species composition. The few socio-economic impacts are caused by alien bryophytes which form dense mats in lawns and are then considered a nuisance. Most negative impacts on biodiversity have been recorded in natural grasslands, forests, and wetlands. Impacts of alien bryophytes on biodiversity and socio-economy are a recent phenomenon, with >85 % of impacts on biodiversity, and 80 % of impacts on socio-economy recorded since 1990. On average, 40 years (impacts on biodiversity) and 25 years (impacts on socio-economy) elapsed between the year a bryophyte species has been first recorded as alien in a region and the year impacts have been recorded first. Taking into account the substantial time lag between first record and first recorded impact in a region, it seems to be likely that the currently moderate impacts of alien bryophytes will continue to increase. As quantitative studies on impacts of alien bryophytes are rare and restricted to few environments and biogeographic regions, there is a need for addressing potential impacts of alien bryophytes in yet understudied settings.     

云南真菌学与苔藓植物学研究百年记———回顾与展望

A general introduction to the mycological and bryological researches in Yunnan during the last about 100 years was made . Although several contributors such as Lan Mao (1397 - 1476) and Li Shi-zhen (1518 - 1593) paid their attention to fungi and bryophytes for medical purposes in as early as the Ming Dynasty, much of scientific researches were done in the last one hundred years .Western missionaries and biological explorers, and domestic experts, such as F. L. Tai, T. N. Liou, S. T. Teng , W. F. Chiu. D. F. Yu and J. C. Chou, traveled and collected from different localities of Yunnan and its adjacent areas . Collections of fungi and bryophytes were then studied by mycologists and bryologists from both home and abroad .Many new species of fungi and bryophytes from Yunnan have been discovered and described. The bryophytes in Yunnan are now relatively well-known. However, much mycological work should be done in the future before the diversity of fungi in the region can be recognized and understood.     

干旱与半干旱地区苔藓植物生态学研究综述

苔藓植物一般生活在阴湿的地方,但也有不少种类具有极强的耐旱性,在干旱和半干旱地区也有分布,在干旱和半干旱地区,影响苔藓植物分布的环境因子包括降水,pH值,CaCO3含量,植被盖度,有机质含量以及土壤质地等,其中起沙,固沙和生态小环境的改善中发挥着不可替代的作用。     

赤水河上游主要森林植被中苔藓物种多样性研究

对赤水河上游典型森林植被中苔藓植物多样性调查分析。(1)苔藓植物有15科19属28种(含变种、亚种);生活型有3种：交织型(64.29%)、矮丛集型(28.57%)和平铺型(7.14%)。(2)丰富度以竹叶林中最大(2.131),残存阔叶林中次之(0.687),针叶林中最小(-1.444)。(3)α多样性呈现一定的规律,竹叶林（3.170）＞残存阔叶林（3.000）＞人工植被地（2.807）＞灌木林（2.322）＞针叶林=谷顶区(1.585),α多样性的排序规律与丰富度基本保持一致,均出现在竹叶林中最大。(4)β多样性出现灌木林—残存阔叶林生境中最高（1.000）,竹叶林—残存阔叶林中次之（0.938）,灌木林—针叶林（0.857）中最小的变化差异趋势。分析得出了植被型、湿度和荫蔽度是制约着该地区苔藓植物多样性的主要生态因素。通过苔藓植物多样性的探究,为保护该流域生态环境和生物多样性提供参考价值。     

Conservation of Portuguese red-listed bryophytes species in Portugal: Promoting a shift in perspective on climate changes

Knowledge of bryophyte diversity can be an important tool for identifying overall biodiversity hotspots. The distribution of red-listed species is an essential data for biodiversity conservation actions, and the assessment of species' response to climate change scenarios is also a key tool in future conservation strategies. In this study, we examine the response of four phytogeographic assemblages of all Portuguese red-listed bryophytes whose distributions are well documented in Portugal. The red-listed species were selected based on their vulnerability as listed in the new Atlas and Red Data book of Portuguese bryophytes according to the IUCN criteria. The main purpose of this study is to develop predictive distributions of threatened bryophytes grouped according to phytogeographic trends aiming to conserve this bryoflora in future. This is achieved by the identification of relationships between specimens' distributions and environmental ecologically meaningful data, which is known to influence different phytogeographic assemblages. Significant differences were found in all distribution models based on future climate scenarios. Several variables play a vital role in the species' distribution models in present and future environmental conditions.     

The bryophyte paradox: tolerance of desiccation,evasion of drought

Vascular plants represent one strategy of adaptation to the uneven and erratic supply of water on land. Desiccation-tolerant (DT) bryophytes represent an alternative, photosynthesising and growing when water is freely available, and suspending metabolism when it is not. By contrast with vascular plants, DT bryophytes are typically ectohydric, carrying external capillary water which can vary widely in quantity without affecting the water status of the cells. External water is important in water conduction, and results in bryophyte leaf cells functioning for most of the time at full turgor; water stress is a relatively brief transient phase before full desiccation. All bryophytes are C3 plants, and their cells are essentially mesophytic in important physiological respects. Their carbohydrate content shows parallels with that of maturing embryos of DT seeds. Initial recovery from moderate periods of desiccation is very rapid, and substantial elements of it appear to be independent of protein synthesis. Desiccation tolerance in effect acts as a device that evades the problems of drought, and in various adaptive features DT bryophytes are more comparable with (mesic) desert ephemerals or temperate winter annuals (but on a shorter time scale, with DT vegetative tissues substituting for DT seeds) than with drought-tolerant vascular plants.     

Plant colonisation in small forest–floor patches: importance of plant group and disturbance traits

It has been proposed that the interaction between life–history attributes of different organisms and distrbance characteristics play an important role in determining the successional pattern following a disturbance event. We compared the responses of vascular plants and bryophytes (mosses and hepatics) to variation in disturbance size and severity in an old-growth boreal forest during a four–year period. The experiment included two patch sizes (0.25 and 2.5 m²) and two levels of severity: humus patches (removal of vegetation) and mineral soil patches (removal of both vegetation and humus layer). Treatments were chosen to reflect some aspects of disturbance by uprooting. Species richness was significantly affected by both disturbance size and severity but the response differed among plant groups. In vascular plants, species numbers were highest in humus patches while mosses were more numerous in mineral soil patches, the most severe disturbance. In contrast, severity had no effect on hepatics. Plant recovery was more rapid in bryophytes than in vascular plants. Species richness of bryophytes had exceeded that of adjacent, undisturbed vegetation after 2-3 yr. We attribute the contrasting response of the plant groups to differences in regeneration strategies. As a group, bryophytes had a greater variety of regeneration methods than vascular plants, with several types of asexual propagules and abundant production of spores in some species. In contrast, clonal ingrowth dominated in vascular plants while seedlings were rare. Thus, our analysis supports the view that plant response to patchy disturbance is strongly dependent on the interplay among disturbance traits and specific attribtites of different plant groups.     

Bryophytes in perennially moist forests of Papua New Guinea: ecological orientation and predictions of disturbance effects

Water relations of bryophytes must be understood along at least four dimensions: hydration/dehydration frequency; hydration duration; dehydration duration; degree of water loss. All these are biomass-dependent functions such that large colonies of bryophytes may maintain hydration longer than smaller colonies. The spread of a bryophyte colony allows lateral movement of capillary water; contiguous clones will thus allow lateral conduction of that water. In contrast, separated tufts or cushions may store water but will contribute very little to its transfer over the surface of the phorophyte.
Many of the tropical rainforests of New Guinea have a larger mass of epiphytic bryophytes than any temperate forest. The primary effect of disturbance in these forests is a reduction in bryomass, presumably due to desiccation because of increased insolation and wind movement. The reduction of bryomass will decrease water and mineral retention in the bryophyte clones; and the reduction in the size of those clones will reduce their contiguity and thus interfere with bryophyte-mediatcd water and mineral transfer. Disturbance may alter bryophyte species frequency but, under present agricultural and forestry practices in New Guinea, I have seen no evidence of significant loss of bryophyte species.     

Productivity–diversity patterns in arctic tundra vegetation

Productivity has long been argued to be a major driver of species richness patterns. In the present study we test alternative productivity–diversity hypotheses using vegetation data from the vast Eurasian tundra. The productivity–species pool hypothesis predicts positive relationships at both fine and coarse grain sizes, whereas the productivity–interaction hypothesis predicts unimodal patterns at fine grain size, and monotonic positive patterns at coarse grain size. We furthermore expect to find flatter positive (productivity–species pool hypothesis) or more strongly negative (productivity–interaction hypothesis) relationships for lichens and bryophytes than for vascular plants, because as a group, lichens and bryophytes are better adapted to extreme arctic conditions and more vulnerable to competition for light than the taller‐growing vascular plants. The normalised difference vegetation index (NDVI) was used as a proxy of productivity. The generally unimodal productivity–diversity patterns were most consistent with the productivity–interaction hypothesis. There was a general trend of decreasing species richness from moderately to maximally productive tundra, in agreement with an increasing importance of competitive interactions. High richness of vascular plants and lichens occurred in moderately low productive tundra areas, whereas that of bryophytes occurred in the least productive tundra habitats covered by this study. The fine and coarse grain richness trends were surprisingly uniform and no variation in beta diversity along the productivity gradient was seen for vascular plants or bryophytes. However, lichen beta diversity varied along the productivity gradient, probably reflecting their sensitivity to habitat conditions and biotic interactions. Overall, the results show evidence that productivity–diversity gradients exist in tundra and that these appear to be largely driven by competitive interactions. Our results also imply that climate warming‐driven increases in productivity will strongly affect arctic plant diversity patterns.     

XET activity is found near sites of growth and cell elongation in bryophytes and some green algae: new insights into the evolution of primary cell wall elongation

BACKGROUND AND AIMS: In angiosperms xyloglucan endotransglucosylase (XET)/hydrolase (XTH) is involved in reorganization of the cell wall during growth and development. The location of oligo-xyloglucan transglucosylation activity and the presence of XTH expressed sequence tags (ESTs) in the earliest diverging extant plants, i.e. in bryophytes and algae, down to the Phaeophyta was examined. The results provide information on the presence of an XET growth mechanism in bryophytes and algae and contribute to the understanding of the evolution of cell wall elongation in general. METHODS: Representatives of the different plant lineages were pressed onto an XET test paper and assayed. XET or XET-related activity was visualized as the incorporation of fluorescent signal. The Physcomitrella genome database was screened for the presence of XTHs. In addition, using the 3' RACE technique searches were made for the presence of possible XTH ESTs in the Charophyta. KEY RESULTS: XET activity was found in the three major divisions of bryophytes at sites corresponding to growing regions. In the Physcomitrella genome two putative XTH-encoding cDNA sequences were identified that contain all domains crucial for XET activity. Furthermore, XET activity was located at the sites of growth in Chara (Charophyta) and Ulva (Chlorophyta) and a putative XTH ancestral enzyme in Chara was identified. No XET activity was identified in the Rhodophyta or Phaeophyta. CONCLUSIONS: XET activity was shown to be present in all major groups of green plants. These data suggest that an XET-related growth mechanism originated before the evolutionary divergence of the Chlorobionta and open new insights in the evolution of the mechanisms of primary cell wall expansion.     

Bryophytes as experimental models for the study of environmental stress tolerance: Tortula ruralis and desiccation-tolerance in mosses

The development of a complete understanding of how plants interact with the environment at the cellular level is a crucial step in advancing our ability to unravel the complexities of plant ecology particularly with regard to the role that many of the less complex plants (i.e., algae, lichens, and bryophytes) play in plant communities and in establishing areas for colonization by their more complex brothers. One of the main barriers to the advancement of this area of plant biology has been the paucity of simple and appropriate experimental models that would enable the researcher to biochemically and genetically dissect the response of less complex plants to environmental stress. A number of bryophytes model systems have been developed and they have been powerful experimental tools for the elucidation of complex biological processes in plants. Recently there has been a resurgent interest in bryophytes as models systems due to the discovery and development of homologous recombination technologies in the moss Physcomitrella patens (Hedw.) Brach & Schimp. In this report we introduce the desiccation-tolerant moss Tortula ruralis (Hedw.) Gaert., Meyer, and Scherb, as a model for stress tolerance mechanisms that offers a great deal of promise for advancing our efforts to understand how plants respond to and survive the severest of stressful environments. T. ruralis, a species native to Northern and Western North America, has been the most intensely studied of all bryophytes with respect to its physiological, biochemical, and cellular responses, to the severest of water stresses, desiccation. It is our hope that the research conducted using this bryophyte will lay the foundationfor not only the ecology of bryophytes and other less complex plants but also for the role of desiccation-tolerance in the evolution of land plants and the determination of mechanisms by which plant cells can withstand environmental insults. We will focus the discussion on the research we and others have conducted in an effort to understand the ability of T. ruralis to withstand the complete loss of free water from the protoplasm of its cells.     

Relationships between distribution and abundance vary with spatial scale and ecological group in stream bryophytes

1. We examined whether the local abundance of stream bryophytes in a boreal drainage basin (Koutajoki system in northeastern Finland) correlated with their: (i) regional occupancy; (ii) provincial distribution in northwestern Europe; and (iii) global range size. We specifically tested whether aquatic and semi‐aquatic species differ in their distribution–abundance relationships. We also analysed the frequency distributions of occupancy at two spatial scales: within the focal drainage system and across provinces of northwestern Europe. 2. Regional occupancy and mean local abundance of stream bryophytes were positively correlated, and the relationship was rather strong in aquatic species but very weak in semi‐aquatic species. Local abundance was related neither to provincial distribution nor global distribution. 3. Species frequency distributions differed between regional occupancy and provincial distribution. While most species were rare with regard to their regional occupancy within the focal drainage system, most of the same set of species were common and occurred in most provinces in northwestern Europe. 4. The results indicate the presence of dominants (core species) and transients/subordinates (satellite species) among stream bryophytes, highlighting marked differentiation in life‐history strategies and growth form. The observed abundance–occupancy relationships suggest that dispersal limitation and metapopulation processes may govern the dynamics of obligatory aquatic stream bryophytes. In semi‐aquatic species, however, habitat availability may be more important in contributing to regional occupancy.