Observed and dark diversity of alien plant species in Europe: estimating future invasion risk

Invasion by alien species is a growing concern for nature conservation. We estimated the level of invasion by alien plant species and future invasion risks at the European scale. We used a pan-European atlas and eight regional plant atlases to determine the distribution of alien and native plant richness. In addition, we estimated alien and native dark diversity (species currently absent from a site but present in the surrounding region and able to colonize the site). We used relative diversity metrics to indicate current and future risks by alien species: relative alien richness (compared to native species), alien and native completeness (log-ratio of observed to dark diversity) and completeness difference between alien and native species. Observed and relative richness of alien species were greatest in NW Europe; this suggests that sites in NW Europe could be more disturbed. Observed alien and native species richness show clear regional hotspots; the distribution of completeness values is dispersed, indicating local hotspots. Northern Europe has relatively lower alien completeness, likely because potential invaders inhabit the region but have not yet reached many localities, thereby suggesting a risk of future invasion. A greater number of potential alien species in the region increases the probability that some alien species could have detrimental impacts. Both alien richness and completeness are positively correlated with native richness and completeness, respectively, indicating that both groups share similar distribution patterns. Alien species diversity metrics in Europe are related positively to human population density and agricultural land-use. We suggest that the dark diversity concept can broaden our understanding of alien species diversity and future invasion risks.     

A multispecies approach reveals hot spots and cold spots of diversity and connectivity in invertebrate species with contrasting dispersal modes

Genetic diversity is crucial for species’ maintenance and persistence, yet is often overlooked in conservation studies. Species diversity is more often reported due to practical constraints, but it is unknown if these measures of diversity are correlated. In marine invertebrates, adults are often sessile or sedentary and populations exchange genes via dispersal of gametes and larvae. Species with a larval period are expected to have more connected populations than those without larval dispersal. We assessed the relationship between measures of species and genetic diversity, and between dispersal ability and connectivity. We compiled data on genetic patterns and life history traits in nine species across five phyla. Sampling sites spanned 600 km in the northwest Mediterranean Sea and focused on a 50‐km area near Marseilles, France. Comparative population genetic approaches yielded three main results. (i) Species without larvae showed higher levels of genetic structure than species with free‐living larvae, but the role of larval type (lecithotrophic or planktotrophic) was negligible. (ii) A narrow area around Marseilles, subject to offshore advection, limited genetic connectivity in most species. (iii) We identified sites with significant positive contributions to overall genetic diversity across all species, corresponding with areas near low human population densities. In contrast, high levels of human activity corresponded with a negative contribution to overall genetic diversity. Genetic diversity within species was positively and significantly linearly related to local species diversity. Our study suggests that local contribution to overall genetic diversity should be taken into account for future conservation strategies.     

Spreading of alien species and diversity of communities

A new species invading a new area may cause a decrease in diversity of the community already present there. Comparison of temporal changes in species diversity of the “new” community (including alien species) with those of the “original” community (including only native species) may clarify our understanding of the effect of alien species. Using a simulation-based modelling approach we considered several scenarios describing the invasion of native communities by alien species and calculated the trends in Shannon-Wiener indices and in the numbers of species of the “original” and “new” communities during the course of the invasion. We found that despite a large increase in the population size of the invasive alien species the diversity of the original community may be little affected. Native species numbers may stay relatively constant for a long time and then suddenly collapse. The results indicate some possibly still concealed consequences of the spread of the invasive ladybird Harmonia axyridis (Pallas).     

Effect of ploughing on plant species abundance and diversity in the northwestern coastal desert of Egypt

This study focuses on the effect of ploughing on plant abundance,vegetation cover, species richness, and taxonomic diversity during the growingseasons (winter and spring) of 1992 and 2000 in the habitat of inland plateau(natural habitat), 21 km south of Mersa-Matrouh (Egypt).Ninety-five species belonging to 27 families were recorded. High percentages oflife-forms and a large number of species were recorded in ploughed andunploughed stripes in the winter and spring of 2000. Higher averages of importancevalues (IVs) and absolute frequencies were recorded for most perennial andannual species in the unploughed stripes compared to the ploughed ones. This may beattributed to crop failure and consequently unfavourable soil conditions. On theother hand, some shrubby species (e.g. Noaea mucronata andHaloxylon scoparium) and perennial herbs (e.g.Gynandriris sisyrinchium) attained higher IVs in theploughed stripes compared to unploughed ones. This may be attributed to thecultivation of Prosopis juliflora trees in the elevatedpart of the ploughed stripes, which have an ecological role in protecting andenriching the soil with organic matter, thus favouring the growth of theseshrubs and perennial herbs. Higher species richness and diversity wereassociated with low concentration of dominance and low taxonomic diversity inthe spring of 2000 in ploughed and unploughed stripes compared to the winter of1992, for both perennials and annuals. The lowest taxonomic diversities wereexhibited in the spring of 2000 for ploughed and unploughed stripes where thevegetation had the largest number of congeneric species and confamilial genera.Higher species richness and diversity characterized the vegetation of theunploughed stripes, especially in winter and spring 2000, as compared to those ofploughed ones. The present study also reveals low species richness and diversityof therophytes in winter for both ploughed and unploughed stripes.     

Invasive alien plant species in China: regional distribution patterns

Plant invasions have been attracting increasing attention from ecologists because of their worldwide environmental impacts and huge economic costs. Research on the characteristics of the recipient regions is essential for understanding the process of plant invasion. However, few previous studies on invasibility of habitats include social factors, although human activities are critical in the process of plant invasion. China is a vast country with high plant species diversity and a long history of introduction of exotic plant species and is particularly vulnerable to invasive plant species. Alien plant species are widespread in the country. Therefore, the study of invasive plants in China is urgent in practice and theoretically important for developing invasion ecology. For the present study, 126 species were selected to represent the major invasive plant species in China. We then collected data on their species richness in 31 provincial administrative units of China and performed Spearman rank correlations between species richness and possible natural and socio‐economic factors. We found that socio‐economic factors, such as human density and GDP, correlated positively with the species richness of invasive plants in China. In conjunction with the natural and socio‐economic correlations in the study of regional distribution pattern of the major invasive plants, we discussed the factors influencing the regional distribution pattern of the major invasive plants in China. We suggest that native plant species richness was mainly determined by the natural conditions of the regions, while invasive species richness was influenced by natural conditions and human disturbance together.     

不受欢迎的生物多样性：香港的外来植物物种

香港早在19世纪中叶开始就有外来植物入侵的记录,迄今为止,已发现多达238种已归化的外来或怀疑为外来的植物,其中又以薇甘菊（Mikania micrantha)、五爪金龙（Ipomoea cairica)、假臭草（Eupatorium catarium)、大黍（Panicum maximum)等最常见,外来植物最常见于受人为干扰的生境,例如荒废农田及路旁等,而较少在天然林地生境及贫瘠的灌草丛中发现,外来植物的对本地生态系统的影响主要局限于低地生境,它们常形成单优种群,减少了生境及贫瘠的灌草丛中发现,外来植物对本地生态系统的影响主要局限于低地生境,它们常形成单优种群,减少少了生境及动植物的多样性,外来动物对香港原生植物影响最大的是于20世纪70年代入侵的松树线虫（Bur-saphelenchus xylophilus)。外来的脊椎动物也有可能对香港的植物被演替产生影响,目前香港的外来植物当中,有些在大陆较少分布或没有记录,作为华南最大的港口,香港对外来物种的引入扮演着重要的角色,因此制定控制外来种在香港及华南地区的引入及传播的政策及措施非常重要。     

Determinants of native and alien species richness in the urban flora of Rome

Until now, analytical studies on European urban floras have mostly concentrated on the central and north‐western parts of the continent. In this paper, factors determining species richness of urban flora were studied for the city of Rome, Italy, based on a comprehensive floristic survey carried out between 1985 and 1994, and updated in 2005. All species were recorded in grid cells of 1.6 km² and classified into native and alien (the latter divided into archaeophytes and neophytes). The grids were classified with respect to the prevailing habitat type, area available to vegetation, level of disturbance and geographical position within the city. Data were analysed using minimal adequate models. Total species number was determined by habitat and its interaction with position on the north‐west gradient; other variables explained much less variance. Holding other variables constant, the average species number per grid cell was highest in archaeological sites and parks, followed by woodlands and rivers, and grasslands and recent developments. Residential areas and the historical centre were poorest in species number. Towards the north of the city, species richness in corresponding habitats increases because of higher landscape heterogeneity and closer association with diaspore pools in the surroundings. Native species make up on average 84% of the total species numbers, and trends opposite to those for the total number of species were found for the proportional representation of aliens. The occurrence of alien and native species in the flora of Rome is driven by similar factors, but factors that increase representation of aliens decrease that of natives and vice versa. The representation of neophytes and native species in grid cells was easier to explain (74% of variation accounted for) than that of archaeophytes (27%); this result reflects that in terms of ecology and response to factors, archaeophytes take an intermediate position between native plants and neophytes. Proportional representation of neophytes decreased with increasing area available to vegetation, reflecting that semi‐natural vegetation is better developed where less fragmented.     

Forest vascular plants as indicators of plant species richness: A data analysis of a flora atlas from northwestern Germany

Abstract

Our study had the objective to examine whether the number of forest vascular plants in a forest-poor region may be indicative of total plant species richness and of the number of threatened plant species. We also related forest plant species richness to geological and soil variables. The analysis was based on a regional flora atlas from the Weser-Elbe region in northwestern Germany including incidence data of species in a total of 1109 grid cells (each ca. 2.8 × 2.8 km²). All taxa were classified either as forest or non-forest species. Total species richness in the grid cells ranged from 65 to 597, with a mean value of 308. The number of forest species varied between 20 and 309 (mean 176). Grid cells with or without particular geological units differed in total and forest species richness, with those containing peatland and marshland being particularly species-poor. Indicator value analysis showed that both total and forest species richness in the grid cells were related to soil acidity and nitrogen in a hump-backed manner, with the highest number of species found at moderately low values for nitrogen and at intermediate values of pH. Forest species richness was highly positively correlated with the number of non-forest species and threatened non-forest species. Indicators for high species richness were primarily those species that are confined to closed semi-natural forests with a varied topography and relatively base- and nutrient-rich soils. Grid cells including historically ancient forest exhibited a higher species richness than grid cells lacking ancient forest, indicating the importance of a long habitat continuity for a high phytodiversity. The “habitat coincidence” of high species richness is best explained by similar responses of forest species and species of other habitats to the main environmental gradients. It is suggested that the regional patterns found for the Weser-Elbe region can be transferred also to other forest-poor regions in Central Europe.     

Altitudinal distribution of alien plant species in the Swiss Alps

In summer 2003 we recorded the presence and abundance of alien plant species at 232 sites (107 railway stations and 125 road sites) along mountain passes in the Swiss Alps. The altitudinal distribution of species was related to the current abundance of the species in Switzerland and time since introduction. A total of 155 alien taxa were recorded. Numbers of species per site declined exponentially with altitude, and only a few species were found in the alpine zone (>2000 m). In contrast, species richness among comparable native taxa appeared to be nearly independent of altitude over the range investigated. Maximum altitude reached by alien species was related positively to both total area occupied in Switzerland and to time since introduction. A comparison of the results with earlier records suggests that many species, particularly those previously restricted to low or intermediate altitudes, have advanced their altitudinal limits over the past few decades. Various hypotheses are presented to explain the declining abundance of alien species with altitude: low-altitude filter effects, low propagule pressure, and genetic swamping of peripheral populations at higher altitudes. However, at present we do not have sufficient evidence to determine the relative importance of these effects. We conclude that invasion into mountain areas such as the Swiss Alps tends to proceed rather slowly, though the process may be accelerated by climatic warming. For this reason, further research to investigate the processes determining how plants invade mountain areas is urgently needed. And more generally, investigations into the distribution of alien species along strong altitudinal gradients may provide valuable insights into the mechanisms driving the spread of alien organisms.     

Ecological preferences of alien plant species in North-Eastern Germany

The large, comprehensive vegetation database of Mecklenburg-Vorpommern/NE Germany with 51,328 relevés allowed us to study an entire regional flora of 133 non-native plants (NNP, immigration after 1492 AD) with regard to their preferences to all kinds of habitats and along different ecological gradients. For each relevé, we computed average Ellenberg indicator values (EIV) for temperature, light, moisture, reaction, nutrients and salt as well as plant strategy type weights. We partitioned the dataset into relevés with and without occurrences of NNP and compared them with respect to the relative frequencies of EIVs and strategy type weights. We identified deviations from random differences by testing against permuted indicator values. To account for bias in EIV between community types, NNP preferences were differentiated for 34 phytosociological classes. We tested significance of preferences for the group of NNP as a whole, as well as for single NNP species within the entire dataset, as well as differentiated by phytosociological classes and formations. NNP as a group prefer communities with high EIVs for temperature and nutrients and low EIVs for moisture. They avoid communities with low EIV for reaction and high EIV for salt. NNP prefer communities with high proportions of ruderal and low proportion of stress strategists. The differentiation by phytosociological classes reinforces the general trends for temperature, nutrients, moisture, R and S strategy types. Nevertheless, preferences of single species reveal that NNP are not a congruent group but show individualistic ecological preferences.     

Protein hot spots: the islands of stability

Understanding the structural basis of protein-protein interactions (PPIs) may shed light on the organization and functioning of signal transduction and metabolic networks and may assist in structure-based design of ligands (drugs) targeting protein-protein interfaces. The residues at the bimolecular interface, designated as the hot spots, contribute most of the free binding energy of PPI. To date, there is no conclusive atomistic explanation for the unique functional properties of the hot spots. We hypothesized that backbone compliance may play a role in protein-protein recognition and in the mechanism of binding of small-molecule compounds to protein surfaces. We used a steered molecular dynamics simulation to explore the compliance properties of the backbone of surface-exposed residues in several model proteins: interleukin-2, mouse double minute protein 2 and proliferating cell nuclear antigen. We demonstrated that protein surfaces exhibit distinct patterns in which highly immobile residues form defined clusters ("stability patches") alternating with areas of moderate to high mobility. These "stability patches" tend to localize in functionally important regions involved in protein-protein recognition. We propose a mechanism by which the distinct structural organization of the hot spots may contribute to their role in mediating PPI and facilitating binding of structurally diverse small-molecule compounds to protein surfaces.     

Threats and benefits of invasive alien plant species on pollinators

Invasive alien plant species are usually disliked due to their high pressure on native communities. However, their ecological effects on pollinators are complex: some species provide abundant floral resources, boosting the number of pollinators, while they often disrupt plant-pollinator interactions by outcompeting native plants. Our direct knowledge is mainly based on single-species studies, while understanding the mechanism of these complex ecological interactions needs multi-species field-based approaches. It is also imperative to clarify the pros and cons of invasive plants and drivers of invasion from the perspective of pollinators. We conducted a standard protocol-driven regional study in Central and Eastern Europe, comparing 6-7 invaded and non-invaded sites of 12 herbaceous invasive plant species. We sampled floral resources, bees, and hoverflies before and during the flowering of the invasive plants. We analysed the effects of plant invasion at the invasive plant species level and in combined analyses, and tested whether the life span (perennial vs. annual) and flowering time (early-, middle-, and late-flowering) of invasive plants affect the abundance, species richness, diversity and species composition of native plants and pollinators. The combined analyses showed lower abundance and species richness of flowering plants and pollinators before, and higher abundance of both during the flowering of invasive plants in invaded sites. However, invasive plants had significant species-specific effects. Perennial invasive plants had a stronger negative impact on floral resources and pollinators already before their flowering compared to annuals. Flowering time of invasive plants affected the pollinator guilds differently. We suggest that in certain critical time periods of the year, invasive plants might provide the dominant foraging resources for pollinators in an invaded ecosystem. But, they also often cause significant losses in native floral resources over the year. Instead of simple eradication, careful preparation and consideration might be needed during removal of invasive plants.     

Temperature-dependent performance of competitive native and alien invasive plant species

To assess the likely impacts of environmental change, the responses of two well-known invasive plant species, native Pueraria lobata and alien Humulus japonicus, to differences in growth temperature were studied in South Korea. Habitat preferences, physiological responses such as photosynthetic rates and chlorophyll contents, growth rates, and nutrient contents were quantified for each species. A competition experiment was conducted to evaluate the temperature preferences of the two species. All results indicated that the alien species H. japonicus can take advantage of elevated temperatures (35 °C) to enhance its competitive advantage against the native species P. lobata. While H. japonicus took advantage of elevated temperatures and preferred high-temperature areas, P. lobata showed reduced performance and dominance in high-temperature areas. Therefore, in future, due to global warming and urbanization, there are possibilities that H. japonicus takes advantage of elevated temperature against P. lobata that could lead to increased H. japonicus coverage over time. Therefore, consistent monitoring of both species especially where P. lobata is dominated are required because both species are found in every continents in the world. Controlling P. lobata requires thorough inspection of H. japonicus presence of the habitat in advance to prevent post P. lobata management invasion of H. japonicus.     

Exotic plant species of the St Lawrence River wetlands: a spatial and historical analysis

Aim To evaluate the importance (number of species, plant cover) of the exotic flora in seven well‐defined sectors of one of the most important transportation waterways in North America. To determine the impact of exotic species on wetland plant diversity and reconstruct the spread of some invasive species. Location St Lawrence River, southern Québec. Methods The exotic flora (vascular plants) of wetlands bordering the St Lawrence River was studied using 713 sampling stations (25 m²) along a 560‐km long corridor. Results Exotic species represent 13.7% of the vascular flora of the St Lawrence wetlands. The relative plant cover occupied by exotic species is high in some of the fluvial sectors (42–44%), but low (6–10%) in the estuarine sectors. Wetlands (marshes) surrounding islands were particularly susceptible to invasion by exotic plants. Historical, abiotic and landscape factors may explain the differences observed between sites. Purple loosestrife (Lythrum salicaria L.) is the most common exotic species of the St Lawrence wetlands, but other species, namely flowering‐rush (Butomus umbellatus L.) and reed canary grass (Phalaris arundinacea L.) are much more invasive. There is no linear relationship between the exotic species cover and the diversity of wetland plants; low diversity sites can be dominated by either exotic or native plant species. In the other sites, exotic species generally have little impact on plant communities and can contribute to increase diversity. Common reed (Phragmites australis (Cav.) Trin. ex Steudel) and reed canary grass, both considered as exotic species in this study, clearly have a stronger impact on plant diversity than flowering‐rush and purple loosestrife. Main conclusions This study shows that the global impact of an invader cannot be adequately evaluated with only a few highly invaded sites. While nationwide strategies have been developed to control exotic species, large surveys are essential to adapt them to regional particularities.     

Intersexual spatial relationships in a lekking species: blue-crowned manakins and female hot spots

Leks offer an intriguing evolutionary problem: why do malesaggregate when this apparently leads to fitness costs? Aggregationcosts can be balanced if males settle on patches where theyare more likely to encounter females (hot-spot hypothesis).We evaluated whether female hot spots can account for patternsof lek structure in the blue-crowned manakin (Lepidothrix coronata)by modeling female distribution patterns relative to lek locationsin two 100-ha plots. Individual females were mapped based onnest locations and capture points and had their home ranges(HRs) modeled based on radiotelemetry data. The number of femalesthat lekking males can be expected to encounter was estimatedas the number of individual female HRs overlapping each maleterritory; hot spots were defined as patches where more femalesare found than average. We investigated how changes in femaleHR size and devaluation effects (decrease in female availabilitydue to the presence of neighboring males) influence male accessto females. Both factors strongly influenced the expected ratesof female encounter, but the hot-spot hypothesis was not supported:most male territories consistently overlapped fewer or justas many female HRs as expected by chance. Leks were not closerto hot spots than similar-sized nonlek sites. A proportion ofmales were, indeed, settled at hot spots, but, contrary to predictionsof the hot-spot hypothesis, they belonged to smaller leks thanmales located outside hot spots. Our results indicate that thislack of spatial correlation between males and females resultspartly from differences in sex-specific habitat preferences.     

Evolution of diversity: the Cape flora

Although the environmental correlates of plant species richness have long received attention, research into the genesis of this diversity is in its infancy. The recent development of molecular dating methods and the inference of robust phylogenetic hypotheses have made it possible to investigate this problem. I use the Cape flora as a model to show that much modern diversity could be the result of recruiting diverse lineages over the entire Cenozoic into this flora, followed by in situ diversification (thus increasing species richness) of at least some of these lineages over a long period in an environmentally heterogeneous area.     

Invasive alien plant control: The priority to save one of the most rapidly declining island-endemic plant species worldwide

Up to 6,800 plant species endemic to oceanic islands are highly threatened with extinction. Although habitat destruction and fragmentation have greatly contributed to this, it is generally recognised that invasive alien species currently pose the single most important threat to island plants. Most studies exploring the role of novel interspecific interactions in driving declines of island plants, focus on threats mediated by animals, be it direct (e.g. browsing, seed predation, mutualism disruption) or indirect (e.g. extinction of seed dispersal or pollination mutualists). Relatively few studies have investigated the specific role of plant-plant interactions, particularly in-situ. We studied a threatened island endemic plant in rapid decline to evaluate the short (1–2 years) and medium-term (about 1–2 decades) influence of invasive alien plants (IAPs) on individuals and a variety of proxies of plant fitness. We compared mortality of traceable individuals that were recorded 12–20 years previously between habitats that are invaded with IAPs and habitats where IAPs are absent, or have been removed decades ago. We also carried out an in-situ manipulative experiment using 14 randomly chosen plants from around which IAPs were removed, paired with controls, at two sites. Canopy cover change before and after IAPs’ removal was quantified along with above ground biomass of IAPs removed for use as potential explanatory variables of change in proxies of plant fitness. Ten branches were randomly selected per plant and branch dynamics, leaves’ sizes and reproductive structure production were monitored quarterly for two years. Over the medium term, plant mortality was recorded only in presence of IAPs (X² = 4.80, df = 1, p < 0.05). Over the short term, at the plant level, IAPs’ removal triggered overall weak to moderate improvements in the number of surviving and new branches as well as change in number of branches at one of the sites. At the leaf and branch levels, we found weak evidence for positive effects of IAPs removal on surviving leaves, flower buds produced and difference in leaf surface area per branch in one site. We therefore provide some experimental evidence of negative effects of alien plants on overall fitness of the threatened species in-situ presumably through competitive interactions. We posit that these effects were found to be weak to moderate due to the short experimental period over which they could develop (1–2 years). Overall, IAPs stand out as the most severe threat from among all documented threats to the species, for being the only one capable of causing mortality of adult plants. Results hence highlight island plants’ vulnerability to IAPs, and how their timely control would improve the survival and fitness of threatened plants, even at the scale of single individuals. Such a strategy could be more often employed. Our study stresses on prioritising IAPs’ control for rescuing long-lived threatened plants that grow in habitats invaded by alien plants (itself a very common situation on oceanic islands) before addressing other subtler, slower-acting threats, like disrupted pollination or seed dispersal mutualisms, florivory or seed predation.