Macroecology of global bryophyte invasions at different invasion stages

In this study we provide the first comprehensive assessment of the environmental and anthropogenic factors driving bryophyte invasions worldwide. We compiled data of alien bryophyte distributions from 82 regions on five continents and oceanic islands and region specific variables. For each species, we collected data on its region‐specific invasion stage, i.e. casual (ephemeral) vs naturalized (persistent) occurrences, and we differentiated between known aliens and those which are likely to be alien (cryptogenic). We used these data to test how species attributes, environmental and socio‐economic conditions of the study areas as well as introduction effort affect invasion probabilities at different invasion stages and of known alien vs cryptogenic species. We collected information on species’ attributes (native range size and location, niche breadth, habitat affiliation), and calculated variables characterising the environmental, biogeographic and socio‐economic features of the native and recepient regions. Subsequently, we related the probability of alien occurrence across different invasion stages with these species‐ and region‐wise predictor variables using generalized linear mixed effects models. Greater native range size raised the likelihood that a species becomes alien or cryptogenic. Islands are more invaded by alien (and cryptogenic) bryophytes than continental regions. Native range size and socio‐economic activity increase the likelihood that a species becomes alien or cryptogenic elsewhere. Interestingly, among alien bryophytes naturalizations occur more frequently in regions of the complementary hemisphere than in regions of their native hemisphere. In general, regions in the Southern Hemisphere have higher numbers of naturalized bryophytes. We conclude that there is a conspicuous change in factors determining bryophyte invasions at different invasion stages. Whereas alien and cryptogenic bryophyte species occurrences are more frequent on islands and depend on native range size, and hence probably propagule pressure, naturalized bryophytes are more frequent in areas which are biogeographically separated but climatically similar to the native ranges.     

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.     

Telling a different story: a global assessment of bryophyte invasions

We assess and review spatio-temporal patterns, habitat affiliations, pathways, impacts, and management experience of bryophyte invasions in extra-tropical countries and regions (n = 82) from five continents and maritime islands spanning both hemispheres. Distribution data were extracted and critically checked from a wide range of sources and supplemented with data on biology and introduction history. We identified 139 bryophytes species which we consider to be alien in at least one of our study regions (106 mosses, 28 hepatics and 5 hornworts). Numbers of average alien bryophyte species are significantly higher on islands than in continental regions of similar size, and peak on maritime islands. Cumulative numbers of first records have grown slowly until 1950 and have strongly increased since then. Accidental import as hitch-hiker (34 species) or with ornamental plants (27 species) constitute the most important introduction pathways. We found a remarkably high contribution from distant donor regions to alien bryophyte floras, especially from the complementary hemisphere. Most alien bryophytes prefer strongly modified habitats (e.g. ruderal vegetation, roadsides, lawns), and only few natural ecosystems (forests, rocks) are regularly invaded. Evidence for an ecological impact of bryophyte invasions is scarce and competitive replacement of native moss species, or vascular plant seedlings, by alien bryophytes has only been documented in a few cases. We conclude that bryophytes differ profoundly in many respects from vascular plants, and so do their invasion patterns at large scale. Our global bryophyte invasion state assessment provides the basis for future, more explicit considerations of this largely neglected taxonomic group in invasion ecology, a step we suggest to be urgently needed as studying them might provide novel insights into patterns and processes of plant invasions in general.     

Assemblages of native bryophytes in secondary forests with introduced Quercus rubra

We have studied the diversity of bryophytes in planted Polish post‐agricultural forests dominated by the native Scots pine Pinus sylvestris and the alien (North American) red oak Quercus rubra. The planted sites would be suitable for a mesic coniferous forest (abbreviation: CFS) or mesic broadleaved forest (abbreviation: BFS). We analysed the structure and composition of the bryophyte assemblages in relation to forest site and substrate availability. Special attention was paid to the introduced Q. rubra as a host species for native bryophytes. A total of 54 bryophyte species (9 liverworts and 45 mosses) were found in the 90 plots ( = phytosociological relevés, 10 × 10 m in area; 45 at each forest site) studied. DCA analysis showed that the bryophyte assemblages of the P. sylvestris–Q. rubra secondary forest community differed significantly between CFS and BFS sites; the similarity of the composition of bryophyte species was 36.8%. The substrate preferences (epigeic, epixylic, epiphytic), as well as the growth form and life form of the recorded bryophytes, also differed between CFS and BFS, while the proportion of bryophytes that had a particular life strategy was very similar. The introduced Q. rubra was inhabited by 28 bryophyte species, including two liverworts. This tree hosted 64% of the CFS and 47% of BFS bryophyte flora and as a host for epiphytes the species successfully fulfilled the functional role of the native oaks (Q. robur and Q. petraea). Thus, the introduction of Q. rubra may contribute to the restoration of post‐agricultural forests and to the conservation of epiphytic bryophyte species. On the other hand, the negative impact of Q. rubra observed on the ground flora (including bryophytes) puts the benefits of Q. rubra for the conservation of native biodiversity in general in question.     

Bryophyte and vascular plant species richness in boreo-nemoral moist forests and mires

We compare species richness of bryophytes and vascular plants in Estonian moist forests and mires. The material was collected from two wetland nature reserves. Bryophyte and vascular plant species were recorded in 338 homogeneous stands of approximately 1 ha in nine forest and two mire types. Regional species pools for bryophytes and vascular plants were significantly correlated. The correlations between the species richnesses of bryophytes and vascular plants per stand were positive in all community types. The relative richnesses (local richness divided by the regional species pool size) were similar for bryophyte species and for vascular plant species. This shows that on larger scales, conservation of the communities rich in species of one taxonomic plant group, maintains also the species richness of the other. The minimum number of stands needed for the maintenance of the regional species pool of typical species for the every community type was calculated using the species richness accumulation curves. Less stands are needed to maintain the bryophyte species pools (300–5300 for bryophytes and 400–35 000 for vascular plants).     

贵州乌江东风水库库区消落带苔藓植物区系分析

采用典型调查与路线调查相结合的方法,对贵州乌江东风水库库区消落带苔藓植物区系的物种组成、生活型、分布区类型和丰富性进行了调查和分析.结果表明:该区域共有苔藓植物18科58属101种,其中,藓类植物有16科56属99种,苔类植物有2科2属2种;优势科为灰藓科(Hypnaceae)、青藓科(Brachytheciaceae)和丛藓科(Pottiaceae),优势属为青藓属(Brahchythecium B.S.G.)、真藓属(Bryum Hedw.)和小曲尾藓属〔Dicranella(Müll.Hal.)Schimp.〕;单属科和单种属所占比例均较高,分别占该区域苔藓植物总科数和总属数的500%和638%.该区域分布的苔藓植物生活型可分为交织型、丛集型、垫状和平铺型4类,以丛集型种数最多(48种),占该区域苔藓植物总种数的475%.该区域分布的苔藓植物可划分为12个分布区类型,其中,温带成分种类最多,所占比例为815%;热带成分所占比例仅为185%;中国特有种所占比例也较高,为207%.该区域苔藓植物的丰富性综合系数(Si)为-03608,低于相邻的六冲河下游流域.综合分析结果表明:该区域的苔藓植物多样性较为丰富,多数种类具有较强的抗逆性,且丛集型苔藓种类最多,与库区消落带的特殊生境相适应;地理成分以东亚成分和北温带成分为主,总体属温带性质,且中国特有种较多,反映出该区域苔藓植物区系的特殊性和复杂性.     

Diversity of saproxylic bryophytes in old‐growth and managed beech forests in the central Balkans

Abstract

The diversity of saproxylic bryophyte species in beech forest stands from the wide region of the central Balkans (i.e. Serbia and Montenegro) was studied, and this study is the first of such a type in SE Europe. Comparison of preserved old‐growth and managed forests were made. Bryophyte species diversity is higher in primeval forest stands where the spectra of dead wood in various decaying stages of its dynamics are present. The ecological group of epixylic specialists is predominant among the bryophytes recorded. Threatened bryophyte species occur in old‐growth beech stands. The dead wood as habitat together with some other factors are extremely important for the surviving of epixylic bryophyte; so these species can be used as bioindicator bryophyte species of old‐growth or managed and structured forest ecosystems.     

Reinvestigation on species richness and environmental correlates of bryophytes at a regional scale in China

Aims The aim of this article is 4-fold: (i) to update species richness of bryophytes for each of the Chinese provinces based on the most current knowledge on distributions of bryophytes in China, (ii) to provide a set of analyses based on the updated species richness data and the environmental variables used in a recent article on species richness of bryophytes in China, (iii) to expand the analysis presented in the recent article by relating species richness of bryophytes to over 15 additional climatic variables and (iv) to determine the degree to which the relationships between bryophyte species richness and environmental variables that were reported in the recent article might have been biased.Methods Over 180 literatures with national, provincial and local species lists of bryophytes in China were used in this study. Taxonomy and nomenclature of bryophytes in China were standardized according to The Plant List. Correlation and regression analyses were used to examine the relationships between species richness or species density of bryophytes in Chinese provinces and environmental variables.Important findings On average, each Chinese province possesses 700.6 species of bryophytes, which is 112.1 species more than previously reported. With the updated species richness data reported in this study, stronger relationships between species richness of bryophytes and environmental variables have been found, compared with those found in a recently published study for China. When single environmental variables were considered, precipitation-related variables were, on average, more strongly correlated with species richness and species density than were temperature-related variables. Environmental variables were on average correlated more strongly with species density than with species richness of bryophytes at the regional scale in China. Our study showed that measures quantifying the average and variation of environmental conditions within each Chinese province explained 82.7% and 71.1% of the variation in species richness of liverworts and mosses, respectively, and explained 86.5% and 70.7% of the variation in species density of liverworts and mosses, respectively.     

Bryophytes and ecological niche theory

Classical niche theory, particularly in terms of competitive exclusion, does not appear to apply as well to bryophytes as to other organisms. Bryophyte communities, as well as those of other plants and of animals, can be thought of in terms of individual species each utilizing particular portions of various resource or habitat continua. Quantitative studies carried out since 1981, particularly those involving niche breadth and overlap, are reviewed. Special attention is given to niche diversification in Sphagnum , Splachnaceae, bryophyte communities in streams, and to ephemeral bryophyte communities. Some bryophyte communities appear to have equilibrium characteristics and to contain species with relatively narrow niche breadths and with no or only partial niche overlap. In many habitats, however, bryophyte communites have non-equilibrium characteristics and diversification of species in microhabitats is opportunistic. Do any bryophyte communities persist long enough for complete saturation by species which have realized niches determined by competitive interactions? Recent studies indicate that this is the case for at least some Sphagnum communities, but that it is the exception not the rule for bryophytes.     

Effects of management and adjacent forest on the heathland bryophyte layer

Heathlands are severely threatened by decreasing habitat quantity and quality. In these habitats, bryophytes form an important component of plant diversity. Therefore, the aims of this study were to investigate the effects of management and adjacent forest land use on the heathland bryophyte layer. Furthermore, effects of these factors on invasive bryophyte species were studied.

Bryophyte communities of 11 dry heathland patches bordering forest were surveyed. In these heathlands, management turned out to be an important factor explaining community composition of bryophytes. Furthermore, significant effects of adjacent land use on bryophyte community composition and species richness were found. However, these effects were rather limited in extent, occurring only close to habitat borders, i.e., up to maximum 5 m into the heathland patch. Also for individual bryophyte species, effects of adjacent forest could only be observed within maximum 5 m from the edge. One remarkable edge effect was the increased dominance of the invasive species Campylopus introflexus near the forest edge, especially at grazed sites. Effects of adjacent land use seemed to predominate management effects. Consequently, management cannot serve as a tool to mitigate edge effects on bryophyte species in heathland patches.     

黄土高原北部水蚀风蚀交错区苔藓多样性及苔藓结皮发育的微生境特征

微生境因子对苔藓结皮的物种分布和发育有重要调控作用。为探明其作用途径和影响程度,以黄土高原水蚀风蚀交错区的苔藓结皮为研究对象,通过样线法调查了8种微生境中苔藓结皮的物种分布和发育特征,结合结构方程模型剖析了植物冠层覆盖、坡向和坡度对苔藓多样性和苔藓结皮发育的影响途径并量化了其影响系数。结果表明: 1)与无冠层相比,冠层覆盖使苔藓的Patrick、Shannon、Pielou和Simpson指数分别降低了63.4%、66.6%、91.0%和68.3%,但使苔藓结皮的厚度、生物量和叶绿素含量分别提高了0.5、0.2和1.3倍;2)北坡样地的4种苔藓物种多样性指数较南坡分别提高了0.6、0.9、5.6和0.9倍,而3种苔藓结皮发育指标分别提高了0.3、0.3和0.6倍;3)坡度从14°增加到34°,4种苔藓多样性指数分别降低了59.8%、84.1%、57.3%和68.0%,而3种苔藓结皮发育指标分别降低了15.2%、25.0%和16.5%;4)3种微生境因子对苔藓多样性和苔藓结皮发育的影响程度为冠层覆盖＞坡向＞坡度,其主要影响途径因微生境因子不同而存在差异。综上,植物冠层覆盖、坡度和坡向分异通过直接和间接途径显著影响了苔藓的物种分布和苔藓结皮发育,因此,在利用苔藓结皮进行荒漠化治理时应充分考虑微生境条件。     

Predicting bryophyte diversity in grassland and eucalypt-dominated remnants in subhumid Tasmania

Aim To determine the environmental factors associated with bryophyte diversity in remnants in a fragmented, agricultural landscape. Location Eighty‐two remnants of tussock grassland, eucalypt woodland and eucalypt forest in the subhumid Midlands region of Tasmania, Australia. Methods Remnants were surveyed for bryophytes and predictor variables, such as vascular plant cover, climate, and topography. Management histories for each remnant were compiled using both site observation and landowner surveys. Bryophyte cover, richness, and composition were related to the independent variables using simple correlation and general linear models. Results We found weak relationships between the dependent variables and the fragmentation variables (remnant area, remnant perimeter to area ratio, distance to nearest remnant, distance to nearest larger remnant, and remnant age). Instead, climatic variables were important in predicting bryophytes, in particular those affecting humidity (minimum temperature of the coldest month, precipitation). Despite extensive sheep grazing in this landscape, grazing was not correlated with bryophyte diversity. Bryophyte diversity was not explained by vascular plant richness and was only weakly explained by composition, but was predicted by the cover of vascular plants. There was greater bryophyte cover and richness and different composition where the cover of native vascular plants was lower. Main conclusions The implications of our results are that all remnants, regardless of area, age and isolation, appear to be valuable for bryophyte conservation in this highly altered landscape. Our results also suggest that the cover of the vascular plant community, rather than its diversity, holds promise as a guide to bryophyte diversity. Bryophyte composition was similar between sites and a focus on the most species‐rich sites may be the best conservation strategy in this ecosystem.     

Diverse bryophyte mesofossils from the Triassic of Antarctica

Compared with the fossil record of vascular plants, bryophyte fossils are rare; this circumstance is probably related to a lower preservation potential compared with that of vascular plants. We searched for bryophyte remains in extensive collections of plant‐fossil assemblages from the Triassic of Antarctica and identified three assemblages with surprisingly well‐preserved bryophyte fossils. Although most bryophyte remains are too fragmented to conclusively place them in a detailed systematic context, they exhibit features sufficient to suggest the presence of at least four types of leafy bryophytes and two orders of thallose liverworts (Pallaviciniales and Metzgeriales) in the high‐latitude Triassic ecosystems of Antarctica. The leafy bryophytes exhibit combinations of morphological features (e.g. keeled and entire‐margined, ecostate leaves with elongated cells) that today occur in only a few small, systematically isolated groups, but were common among Palaeozoic and especially Mesozoic bryophytes. The diverse morphologies of the bryophyte fossils add further support to previous hypotheses that during warmer periods in the Earth's history, bryophyte vegetation may have been particularly rich and diverse in high‐latitude regions. Through analysis of the sedimentology and taphonomy of these assemblages, we identify a combination of key factors that may explain the preservation of bryophyte fossils in these deposits: (1) punctuated, high‐energetic sedimentary events causing traumatic removal and incorporation of bryophytes into sediment‐laden flood waters; (2) limited transport distance, and short period of suspension, followed by rapid settling and burial as a result of a rapidly decelerating flow discharging into a floodplain environment; and (3) early‐diagenetic cementation with iron hydroxides in locally anoxic zones of the organic‐rich, muddy substrate.     

Birds as key vectors for the dispersal of some alien species: Further thoughts

W. Solarz, K. Najberek, A. Pociecha & E. Wilk‐Wo?niak ( 2017 , Diversity and Distributions, 23 , 113–117) published a letter in Diversity and Distributions debating our view that waterbirds are important vectors of alien species (C. Reynolds, N. A. F. Miranda & G. S. Cumming, 2015 Diversity and Distributions, 21 , 744–754; A. J. Green, 2016 Diversity and Distributions, 22 , 239–247) and question whether future research into the mechanisms under‐pinning this phenomenon can be advantageous for the practical management of alien species. Additionally, Solarz et al. suggest that human activities are the primary source of all alien species introductions and that waterbirds may only act as vectors of secondary dispersal. In this letter, we respond to several arguments raised by the authors surrounding the relevance of waterbird‐mediated dispersal in the introduction and spread of alien species. We emphasize the partly deterministic nature of waterbird dispersal and the significance of long‐distance dispersal events (and hence the potential for primary introductions of new alien species across political boundaries). Finally, we reaffirm the importance of further research into dispersal by birds to improve our capacity to foresee and manage invasions of those alien species with strong capacity to spread via avian vectors.     

BryForTrait – A life‐history trait database of forest bryophytes

In ecological research, plant functional trait analyses are widely applied to understand to what extent the inter‐specific variation in trait attributes has an adaptive value or to predict ecosystem processes and changes. Compared to vascular plants, trait studies using bryophytes are scarce, which is likely due to missing trait information for bryophyte species. With the BryForTrait database, we want to reduce this deficit. Our database represents a compilation of autecological information and morphological and regenerative trait data on different stages of the life cycle of bryophytes occurring in forest ecosystems. The database contains information for 35 traits and 721 Central European bryophyte species; in total more than 23,000 trait values. The BryForTrait database will enable future trait studies, providing new insights into bryophyte‐dominated ecosystems.     

Bryophyte diversity and distribution along an altitudinal gradient on a lava flow in La Réunion

Non‐vascular plant distribution patterns were examined in three microhabitats along an altitudinal gradient on a recent lava flow of the Piton de la Fournaise volcano (La Réunion, Mascarene archipelago). The uniform nature of the lava flow provides an excellent system to study the relationship between altitude and species diversity and distribution, and at the same time avoiding confusing multiple effects of substrate and vegetation heterogeneity. Non‐vascular plants were surveyed with quadrats within an altitudinal range from 250 m to 850 m a.s.l. Fine‐scale variations in bryophyte communities between three ecological microhabitats (the ground and on the rachises of two fern species) were investigated. Three specific questions were addressed: (1) What is the species diversity of bryophyte communities on a 19‐year‐old lava flow? (2) How does altitude influence the diversity and distribution of bryophytes on a lava flow? (3) Does microhabitat variation control bryophyte diversity? In our study, bryophyte diversity increased with altitude. Unexpectedly, species richness was very high; 70 species of bryophytes were recorded including nine new records for the island. Diversity was also controlled by ecological microhabitats. Bryophyte species were structured into six categories according to altitude and microhabitat preferences. Results suggested that the high diversity of these cryptic organisms on this lava flow is fostered in part by their host substrate and their adaptative strategies on new substrates. On a broader scale, it was concluded that lava flows as primary mineral environments are important to conserve, as they support a high diversity of pioneer organisms that constitute the early stages of the development of La Réunion's remnant lowland rainforest.     

Bryophytes as indicators of ancient woodlands in Schleswig-Holstein (Northern Germany)

Ancient woodlands, with their long ecological continuity, frequently harbor a high number of typical, rare and threatened species, and are therefore of particular importance for nature conservation. To pinpoint these habitats, a common application is the use of plants as “ancient woodland indicators”. The occurrence of these particular species allows for evaluating the continuity of woodland cover in time. While lists of ancient woodland vascular plants have been derived for many regions, the identification and use of bryophytes as ancient woodland indicators has been widely neglected. This is a bit surprising because certain woodland bryophytes are very sensitive to varying environmental conditions or changes in land management. It therefore appeared promising to compile an ecologically grounded list of ancient woodland indicator bryophytes for practical use.In this study, we present a set of ancient woodland indicator bryophytes based on the analysis of datasets from the North German federal state of Schleswig-Holstein. To compile this list, we systematically evaluated the bryophyte distribution data from floristic surveys in relation to ancient woodland cover data from state-wide inventories. In this way, we were able to determine ancient woodland bryophytes using consistent and repeatable statistical methods.The presented list of 31 ancient woodland indicator bryophytes is ecologically sound and corresponds well with data from the sparse literature. We could distinguish two groups of ancient woodland indicator bryophytes. The first group is linked to base-rich, semi-natural deciduous woodlands with high soil and air humidity. The second group comprises acidophilic bryophytes that occur not only in acidic beech and oak woods, but also in acidic mixed or coniferous forests on ancient woodland sites. Apart from the ancient woodland indicator bryophytes, we could identify one group of recent woodland bryophytes and four groups of bryophytes that are more or less indifferent with respect to woodland continuity.Finally, we provide recommendations for the application of ancient woodland indicator bryophytes in nature conservation practice. Management suggestions for the conservation of the typical bryophyte diversity of ancient semi-natural woodlands are also given.     

Is vascular plant species diversity a predictor of bryophyte species diversity in Mediterranean forests?

This study aimed to (i) investigate the congruence among the species composition and diversity of bryophytes and vascular plants in forests; (ii) test if site prioritization for conservation aims by the maximization of the pooled number of vascular plant species is effective to maximize the pooled number of bryophyte species. The study was performed in six forests in Tuscany, Italy. Four-hundred and twenty vascular plant species (61 of which were woody) and 128 bryophyte species were recorded in 109 plots. Despite the good predictive value of the compositional patterns of both woody plants and total vascular with respect to the compositional pattern of bryophytes, the species richness of the latter was only marginally related to the species richness of the former two. Bryophyte rare species were not spatially related to rare plant species and neither coincided with the sites of highest plant species richness. The species accumulation curves of bryophytes behaved differently with respect to those of woody plants or total vascular plants. Reserve selection analysis based on the maximization of the pooled species richness of either woody plants or total vascular plants were not effective in maximizing the pooled species richness of bryophytes. This study indicates that species diversity of vascular plants is not likely to be a good indicator of the bryophyte species diversity in Mediterranean forests.     

Determinants of stream bryophyte community structure: bringing ecology into conservation

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北京百花山自然保护区不同植被地面生苔藓植物物种多样性

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