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.     

A new comprehensive database of alien plant species in Chile based on herbarium records

There is an urgent need for comprehensive national databases on alien plant species, especially in developing countries. Despite the fact that plant invasions are considered a major threat to biodiversity, they have been poorly studied or not considered a conservation priority in South America. We aim to assess alien plant distribution in Chile, using the first comprehensive public alien plant database, and discuss the implications of using herbarium records to develop national databases of alien plants. We used herbarium records to assemble a comprehensive national database of alien plants. We calculated the number of alien and native species and specimens recorded in each 10 × 10 km cell. We evaluated sampling efforts and tested for relationships between alien and native species collections, as well as other spatial patterns along the latitudinal gradient. Alien and native species richness was positively correlated. Alien plants were mostly collected in central Chile, with few species collected in both the extreme north and south. However, native plants were strongly collected in central Chile, as well as in both extremes of the country. Alien and native plants followed the same pattern of accumulation along the latitudinal gradient, with native plants being relatively more collected than alien plants. Herbarium records provide valuable baseline information to evaluate plant species distribution. However, there are important gaps in this database, (e.g. variable sampling effort for alien and native plants, incomplete information on life-history traits). Given scientists and land managers increasing demand for baseline information and the high cost of collecting such data in developing countries, herbarium records should be used more frequently for research and management of plant invasions.     

Invasive aliens on tropical East Asian islands

Tropical East Asia (TEA) has numerous islands, both continental and oceanic. This study uses information on invasive aliens in terrestrial habitats on these islands to test the generality of the continental-oceanic contrast in invasibility, assess the conservation impacts of invasive species, and suggest ways to mitigate these. The continental islands of Hong Kong and Singapore are worst-case scenarios for continental invasibility and alien species often dominate in chronically disturbed sites, but very few have successfully invaded closed forests, with the exception of birds in Hong Kong. On other, less densely populated, continental islands, closed-canopy forests appear to resist invasions by all taxa, with few known exceptions. Forests on oceanic islands isolated by <100 km during the last glacial maximum appear no more susceptible to plant and invertebrate invasions than those on continental islands, but invasions by mammals are widespread. Snake invasions may be under-recognized. The remote oceanic Ogasawara (Bonin) Islands, >1000 km from the nearest continent, have a native biota of largely tropical East Asian origin and are suffering from alien forest invasions across the taxonomic spectrum. These patterns of invasibility are consistent with the idea that alien invasion is facilitated by the absence of native species in the same functional group. Alien invasives are not yet a major conservation problem in TEA, except on remote islands, but their dominance on disturbed sites may slow or prevent recovery of native biodiversity. Strict quarantine is impractical in TEA, although some major introduction routes could be blocked. Management efforts should focus on early recognition and immediate control of potential problem species.     

Alien plants introduced by different pathways differ in invasion success: unintentional introductions as a threat to natural areas

Background

Understanding the dimensions of pathways of introduction of alien plants is important for regulating species invasions, but how particular pathways differ in terms of post-invasion success of species they deliver has never been rigorously tested. We asked whether invasion status, distribution and habitat range of 1,007 alien plant species introduced after 1500 A.D. to the Czech Republic differ among four basic pathways of introduction recognized for plants.

Principal Findings

Pathways introducing alien species deliberately as commodities (direct release into the wild; escape from cultivation) result in easier naturalization and invasion than pathways of unintentional introduction (contaminant of a commodity; stowaway arriving without association with it). The proportion of naturalized and invasive species among all introductions delivered by a particular pathway decreases with a decreasing level of direct assistance from humans associated with that pathway, from release and escape to contaminant and stowaway. However, those species that are introduced via unintentional pathways and become invasive are as widely distributed as deliberately introduced species, and those introduced as contaminants invade an even wider range of seminatural habitats.

Conclusions

Pathways associated with deliberate species introductions with commodities and pathways whereby species are unintentionally introduced are contrasting modes of introductions in terms of invasion success. However, various measures of the outcome of the invasion process, in terms of species'' invasion success, need to be considered to accurately evaluate the role of and threat imposed by individual pathways. By employing various measures we show that invasions by unintentionally introduced plant species need to be considered by management as seriously as those introduced by horticulture, because they invade a wide range of seminatural habitats, hence representing even a greater threat to natural areas.     

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.     

How do bryophytes govern generative recruitment of vascular plants?

Interactions between vascular plants and bryophytes determine plant community composition in many ecosystems. Yet, little is known about the importance of interspecific differences between bryophytes with respect to their effects on vascular plants. We compared the extent to which species-specific bryophyte effects on vascular plant generative recruitment depend on the following underlying mechanisms: allelopathy, mechanical obstruction, soil moisture and temperature control. We sowed 10 vascular plant species into monospecific mats of six chemically and structurally diverse bryophytes, and examined 1-yr seedling recruitment. Allelopathic effects were also assessed in a laboratory phyto-assay. Although all bryophytes suppressed vascular plant regeneration, there were significant differences between the bryophyte species. The lack of interactions indicated the absence of species-specific adaptations of vascular plants for recruitment in bryophyte mats. Differences between bryophyte species were best explained by alterations in temperature regime under bryophyte mats, mostly by reduced temperature amplitudes during germination. The temperature regime under bryophyte mats was well predicted by species-specific bryophyte cushion thickness. The fitness of established seedlings was not affected by the presence of bryophytes. Our results suggest that climatically or anthropogenically driven changes in the species' composition of bryophyte communities have knock-on effects on vascular plant populations via generative reproduction.     

植物激素在苔藓生长发育与逆境响应过程中的作用机制研究进展

植物激素是由植物自身代谢产生的一类从产生部位移动到作用部位发挥调控功能的微量小分子有机物质,在植物生长发育、响应环境胁迫过程中起到关键作用.苔藓植物作为早期登陆的非维管植物,处于陆生植物进化早期的阶段,具有许多不同于维管植物的形态和生理特征.大部分苔藓中普遍存在8种主要的植物激素及其衍生物(包括ABA、JA、ET、SA...     

Historical introduction,geographical distribution,and biological characteristics of alien plants in China

Using the published literature and records, this study investigated the historical introduction, geographical distribution, and biological characteristics of 404 alien plants in 32 provinces (including municipalities) in China. The results showed that three fifths of the alien plants was introduced for intentional purposes, and the introduction of alien plants increased with time from 139 BC to the current time, especially during historical events that played an important part in the process of introduction. The geographical distribution of alien plant species tended to decrease from the south and the east to the north and the west. Annual average precipitation (Y = 7.51X + 7.05, R² = 0.67) and temperature (Y = 0.07X + 41.54, R² = 0.67) had a linear relationship with alien plant introduction throughout China. Furthermore, population density and economic conditions also affected and promoted plant invasions in China. The life forms were well adapted to regional precipitation and temperature, but different life forms of alien plants were not consistent with the distribution tendency based on the quantity of alien plants. There was a higher percentage of annual and biennial plants in the northern provinces (e.g. 66.7 and 15.7% in Inner Mongolia), whereas there were more perennial and woody plant species in the southern provinces (e.g. 37.0 and 21.2% in Hainan). Family characteristics could not accurately predict plant invasions; however, predictions based on genera were acceptable. In addition, the origin of alien plants can also predict the distribution and potential risk of invasions. Thus, our study investigated the factors affecting plant invasions in order to provide information on the prevention and control of alien plants.     

Vascular Plants Facilitated Bryophytes in a Grassland Experiment

In grassland communities vascular plants and bryophytes form two distinct layers. In order to understand the factors responsible for plant community structure, more information about interactions between these plant groups is needed. Often negative correlations between vascular plant and bryophyte covers have been reported, suggesting competition. Here we tested experimentally whether different grassland vascular plant species (Trifolium pratense, Festuca pratensis, Prunella vulgaris) had different influences on the cover of two bryophyte species (Rhytidiadelphus squarrosus, Brachythecium rutabulum). In a two-year garden pot experiment one bryophyte species and one vascular plant species were planted per pot. Bryophytes were planted at a constant density, vascular plants in four densities. The cover of both bryophyte species increased with increasing vascular plant cover, showing the facilitative effect of vascular plants through creating better microclimate, e.g., optimising temperature. Bryophyte responses to vascular plant species were species-specific. Festuca had significantly positive effects on both bryophyte species in the second year, and Trifolium on Brachythecium in both years, whereas Prunella had no significant effect on bryophytes. The facilitative effect of vascular plants was stronger at the second experimental year. In summary, the biotic effects between bryophytes and grassland vascular plants are species-specific and positive interactions are prevailing at low vascular plant densities.     

苔藓植物对植物天然更新的影响

种子是天然更新的基础,幼苗和幼树的生长是天然更新过程中最敏感的阶段。苔藓植物作为许多生态系统的主要地被物,在维管植物天然更新过程中发挥着不容忽视的作用。在许多植被类型中,都有发育良好的苔藓群落,大量的研究证明它们的存在影响到维管植物的萌发、建成和生长,并且可能进一步影响到物种的共生,有一些种类甚至可能指示树种的更新。本文就苔藓植物对物种更新早期阶段的作用和影响进行了综述。包括苔藓植物的生理特性造成的微气候变化;苔藓群落对种子传播、萌发及幼苗建成的影响。并讨论了苔藓植物与更新物种的种间关系及其对幼苗生长的化感作用。     

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).     

Impact of Land-Use Intensity and Productivity on Bryophyte Diversity in Agricultural Grasslands

While bryophytes greatly contribute to plant diversity of semi-natural grasslands, little is known about the relationships between land-use intensity, productivity, and bryophyte diversity in these habitats. We recorded vascular plant and bryophyte vegetation in 85 agricultural used grasslands in two regions in northern and central Germany and gathered information on land-use intensity. To assess grassland productivity, we harvested aboveground vascular plant biomass and analyzed nutrient concentrations of N, P, K, Ca and Mg. Further we calculated mean Ellenberg indicator values of vascular plant vegetation. We tested for effects of land-use intensity and productivity on total bryophyte species richness and on the species richness of acrocarpous (small & erect) and pleurocarpous (creeping, including liverworts) growth forms separately. Bryophyte species were found in almost all studied grasslands, but species richness differed considerably between study regions in northern Germany (2.8 species per 16 m²) and central Germany (6.4 species per 16 m²) due environmental differences as well as land-use history. Increased fertilizer application, coinciding with high mowing frequency, reduced bryophyte species richness significantly. Accordingly, productivity estimates such as plant biomass and nitrogen concentration were strongly negatively related to bryophyte species richness, although productivity decreased only pleurocarpous species. Ellenberg indicator values for nutrients proved to be useful indicators of species richness and productivity. In conclusion, bryophyte composition was strongly dependent on productivity, with smaller bryophytes that were likely negatively affected by greater competition for light. Intensive land-use, however, can also indirectly decrease bryophyte species richness by promoting grassland productivity. Thus, increasing productivity is likely to cause a loss of bryophyte species and a decrease in species diversity.     

Impact of invasions by alien plants on soil seed bank communities: Emerging patterns

Much of our current understanding of the impact of invasive species on plant communities is based on patterns occurring in the above-ground vegetation, while only few studies have examined changes in soil seed banks associated with plant invasions, despite their important role as determinants of vegetation dynamics. Here, we reviewed the literature on the impact of plant invasions on the seed bank and we provide a quantitative synthesis using a meta-analysis approach. Specifically, (1) we quantified the impact of 18 invasive alien plants on (i) species richness and (ii) density of the seed banks of invaded communities, based on 58 pair-wise invaded-uninvaded comparisons (cases); we identified (2) the invasive taxa that are responsible for the largest changes in the seed bank; and (3) the habitats where substantial changes occur. Our study showed three major findings: (1) species richness (68% of cases) and density (58% of cases) were significantly lower in native seed banks invaded by alien plants; (2) species richness and density of native and alien species were remarkably lower in seed banks invaded by large, perennial herbs compared to uninvaded sites; and (3) invaded seed banks were often associated with a larger richness and/or abundance of alien species. This study indicates a need for additional seed bank data in invasion ecology to characterize species-specific and habitat-specific impacts of plant invasions, and to determine whether changes in the seed banks of native and alien species are a symptom of environmental degradation prior to a plant invasion or whether they are its direct result. The findings of this study help improve our capacity to predict the long-term implications of plant invasions, including limitations in the recruitment of native species from the seed bank and the potential for secondary invasions by seeds of other alien species.     

Bryophyte vegetation in a wooded meadow: relationships with phanerogam diversity and responses to fertilisation

In the Laelatu wooded meadow in Estonia, famous for its phanerogam diversity, the bryophyte community has been investigated in order to compare its flora and diversity relationships with those of the vascular plant community. Ninety-six bryophyte species were found, 13 of them are hepatics; the majority of the bryophytes are epigeic species common to meadows and forests, including many calciphilous species. Vascular plants and bryophytes display opposite responses to fertilisation. For vascular plants, fertilisation increases the coverage and diminishes the number of species, while for bryophytes it diminishes coverage and increases the number of species. The relationship between the number of species in small plots and the total number of species in the area is similar for vascular plants and bryophytes. No significant changes in the bryophyte community in Laelatu wooded meadow has been detected during the last 30 years.     

Residence time and human-mediated propagule pressure at work in the alien flora of Galapagos

Introduced species present the greatest threat to the unique terrestrial biodiversity of the Galapagos Islands. We assess the current status of plant invasion in Galapagos, predict the likelihood of future naturalizations and invasions from the existing introduced flora, and suggest measures to help limit future invasions. There has been a 1.46 fold increase in plant biodiversity in Galapagos due to alien plant naturalizations, reflecting a similar trend on islands elsewhere. There are 870 alien plant species recorded in the archipelago. Of evaluated species, 34% species have naturalized. Within this group are the invasive species (16% of evaluated) and the transformers (3.3% of evaluated). We show that, as expected, naturalized species have been present in the archipelago longer than non-naturalized species. We also find that a higher human-mediated propagule pressure is associated with a greater human population and with properties that have been settled longer. This, combined with the relatively recent introduction of most species, leads us to the conclusion that Galapagos is at an early stage of plant invasion. We predict that more species from the existing alien flora will find an opportunity to naturalize and invade as propagule pressure increases alongside rapid human population growth associated with immigration to serve the booming tourism industry. In order to reduce future invasion risk, we suggest reviewing inter-island quarantine measures and continuing community education efforts to reduce human-mediated propagule pressure.     

Invasibility of a coastal strip in NE Spain by alien plants

Abstract. The alien plant species and the ecological factors that facilitate their invasion to a coastal strip in the Baix Camp region (Tarragona, NE Spain) were studied. A detailed inventory of the area showed that 20% of the plant species, most of them from the American Continent, were aliens, many of which were strongly invasive. At the habitat level, the relationships between the invasive behaviour and a number of autecological, ecological and habitat variables were analysed by means of logistic analyses. In the best model five variables were related to alien plant invasions: climatic affinity, disturbance of the invaded habitat, route of introduction, soil moisture during summer, and life form. To elucidate the environmental factors that could be responsible for alien plant invasions at a regional scale, the number of alien taxa and the environmental features at 13 localities of the western Mediterranean area were analysed. The warm lowlands, subjected to low annual rainfall and high population density, exhibited the greatest reception capacity for alien plants. It is concluded that various specific environmental features – high mean temperatures, the abundance of riparian systems, the existence of biotopes with favourable water balance which harbour a great richness of alien species and the high human pressure to which the enclave is subjected, combined with the ecological requirements of the alien species and certain species attributes (biotype) can interact at different scales and have caused the over‐representation of alien plants.     

Analysis of introduction history of invasive plants in Brazil reveals patterns of association between biogeographical origin and reason for introduction

Knowledge of historical factors associated with biological invasions in a region can help identify source regions, vectors and pathways more likely to originate potential invaders as well as prioritize resource allocation for selective prevention and early detection strategies. In Brazil, little is known about the introduction history of many invasive plant species, and analyses of historical factors associated with invasions are lacking. To fill this gap, I used a dataset of 117 invasive alien plants across 13 habitats in Brazil to identify potential patterns of continent of origin, reason for introduction, and to test the hypotheses that (i) more Eurasian species are invasive in Brazil than species native from other continents, that (ii) more horticultural species are invasive in Brazil than species introduced for other reasons, and that (iii) continent of origin and reason for introduction are associated. I found that significantly more invasive plant species in Brazil are native to Africa and Asia, were introduced for horticulture and forage, and are part of the families Poaceae, Fabaceae, and Pinaceae. I also found a significant association between continent of origin and reason for introduction, with more invasive species than average being African forage grasses and Asian agroforestry or ornamental plants. In conclusion, the results suggest that the current invasive alien flora of Brazil results from the combination of patterns of recent human migration waves and deliberate species introductions for technological and commercial reasons. These results can help prioritize invasive species and vectors in prevention, early detection, and control strategies.     

Snakes on the Balearic Islands: An Invasion Tale with Implications for Native Biodiversity Conservation

Biological invasions are a major conservation threat for biodiversity worldwide. Islands are particularly vulnerable to invasive species, especially Mediterranean islands which have suffered human pressure since ancient times. In the Balearic archipelago, reptiles represent an outstanding case with more alien than native species. Moreover, in the last decade a new wave of alien snakes landed in the main islands of the archipelago, some of which were originally snake-free. The identification of the origin and colonization pathways of alien species, as well as the prediction of their expansion, is crucial to develop effective conservation strategies. In this study, we used molecular markers to assess the allochthonous status and the putative origin of the four introduced snake species (Hemorrhois hippocrepis, Malpolon monspessulanus, Macroprotodon mauritanicus and Rhinechis scalaris) as well as ecological niche models to infer their patterns of invasion and expansion based on current and future habitat suitability. For most species, DNA sequence data suggested the Iberian Peninsula as the potential origin of the allochthonous populations, although the shallow phylogeographic structure of these species prevented the identification of a restricted source-area. For all of them, the ecological niche models showed a current low habitat suitability in the Balearic, which is however predicted to increase significantly in the next few decades under climate change scenarios. Evidence from direct observations and spatial distribution of the first-occurrence records of alien snakes (but also lizards and worm lizards) suggest the nursery trade, and in particular olive tree importation from Iberian Peninsula, as the main pathway of introduction of alien reptiles in the Balearic islands. This trend has been reported also for recent invasions in NE Spain, thus showing that olive trees transplantation may be an effective vector for bioinvasion across the Mediterranean. The combination of molecular and ecological tools used in this study reveals a promising approach for the understanding of the complex invasion process, hence guiding conservation management actions.