Propagule pressure and native community connectivity interact to influence invasion success in metacommunities

Mechanistic insights from invasion biology indicate that propagule pressure of exotic species and native community structure can independently influence establishment success. The role of native community connectivity via species dispersal and its potential interaction with propagule pressure on invasion success in metacommunities, however, remains unknown. Native community connectivity may increase biotic resistance to invasion by enhancing species richness and evenness, but the effects could depend upon the level of propagule pressure. In this study, a mesocosm experiment was used to evaluate the independent and combined effects of exotic propagule pressure and native community connectivity on invasion success. The effects of three levels of exotic Daphnia lumholtzi propagule pressure on establishment success, community structure and ecosystem attributes were evaluated in native zooplankton communities connected by species dispersal versus unconnected communities, and relative to a control without native species. Establishment of the exotic species exhibited a propagule dose‐dependent relationship with high levels of propagule pressure resulting in the greatest establishment success. Native community connectivity, however, effectively reduced establishment at the low level of propagule pressure and further augmented native species richness across propagule pressure treatments. Propagule pressure largely determined the negative impacts of the exotic species on native species richness, native biomass and edible producer biomass. The results highlight that native community connectivity can reduce invasion success at a low propagule dose and decrease extinction risk of native competitors, but high propagule pressure can overcome connectivity‐mediated biotic resistance to influence establishment and impact of the exotic species. Together, the results emphasize the importance of the interaction of propagule pressure and community connectivity as a regulator of invasion success, and argue for the maintenance of metacommunity connectivity to confer invasion resistance.     

Atp-bound topoisomerase ii as a target for antitumor drugs

Topoisomerase II (TOP2) poisons interfere with the breakage/reunion reaction of TOP2 resulting in DNA cleavage. In the current studies, we show that two different classes (ATP-sensitive and -insensitive) of TOP2 poisons can be identified based on their differential sensitivity to the ATP-bound conformation of TOP2. First, in the presence of 1 mm ATP or the nonhydrolyzable analog adenosine 5'-(beta,gamma-imino)triphosphate, TOP2-mediated DNA cleavage induced by ATP-sensitive TOP2 poisons (e.g. doxorubicin, etoposide, mitoxantrone, and 4'-(9-acridinylamino)methanesulfon-m-anisidide) was 30-100-fold stimulated, whereas DNA cleavage induced by ATP-insensitive TOP2 poisons (e.g. amonafide, batracylin, and menadione) was only slightly (less than 3-fold) affected. In addition, ADP was shown to strongly antagonize TOP2-mediated DNA cleavage induced by ATP-sensitive but not ATP-insensitive TOP2 poisons. Second, C427A mutant human TOP2alpha, which exhibits reduced ATPase activity, was shown to exhibit cross-resistance to all ATP-sensitive but not ATP-insensitive TOP2 poisons. Third, using ciprofloxacin competition assay, TOP2-mediated DNA cleavage induced by ATP-sensitive but not ATP-insensitive poisons was shown to be antagonized by ciprofloxacin. These results suggest that ATP-bound TOP2 may be the specific target of ATP-sensitive TOP2 poisons. Using Lac repressor-operator complexes as roadblocks, we show that ATP-bound TOP2 acts as a circular clamp capable of entering DNA ends and sliding on unobstructed duplex DNA.     

Effect of light stress from phytoplankton on the relationship between aquatic vegetation and the propagule bank in shallow lakes

1. The way light stress controls the recruitment of aquatic plants (phanerogams and charophytes) is a key process controlling plant biodiversity, although still poorly understood. Our aim was to investigate how light stress induced by phytoplankton, that is, independent from the aquatic plants themselves, determines the recruitment and establishment of plant species from the propagule bank. The hypotheses were that an increase in light stress (i) decreases abundance and species richness both of established aquatic plants and of propagules in the bank and (ii) decreases the recruitment success of plants from this bank. 2. These hypotheses were tested in 25 shallow lakes representing a light stress gradient, by sampling propagule banks before the recruitment phase and when the lakes are devoid of actively growing plants (i.e. at the end of winter), established vegetation at the beginning of the summer and phytoplankton biomass (chlorophyll a) during the recruitment and establishment phase. 3. The phytoplankton biomass was negatively correlated with the richness and abundance of established vegetation but was not correlated with the propagule bank (neither species richness nor propagule abundance). The similarity between the propagule bank and established vegetation decreased significantly with increasing phytoplankton biomass. 4. The contrast in species composition between the vegetation and the propagule bank at the highest light stress suggests poor recruitment from the propagule bank but prompts questions about its origin. It could result from dispersal of propagules from neighbouring systems. Propagules could also originate from a persistent propagule bank formerly produced in the lake, suggesting strong year‐to‐year variation in light stress and, as a consequence, in recruitment and reproductive success of plants.     

Temporal Responses of Propagule Banks during Ecological Restoration in the United Kingdom

Successful ecological restoration is expected to be accompanied by change in the propagule bank. We tested for temporal change in the soil propagule bank at two Bracken (Pteridium aquilinum)‐infested sites in the United Kingdom (acid grassland and heathland), each with replicate experiments. A combination of bracken control (cutting, spraying, and combinations) and vegetation restoration treatments (seeding, fertilizer, harrowing) were applied. Soil propagule banks were sampled in 1998, 4–5 years after the start, and in 2003 after a further 5 years. We used univariate and multivariate statistical methods to investigate the response of the propagule bank to experimental treatment in space and time. Few effects were found in 1998, but after a further 5 years, the propagule bank size and composition changed significantly, implying that propagule bank development lags behind vegetation development. The effect of treatment on the propagule bank differed within sites. Thus, ecosystem development is occurring at different speeds and directions even in closely adjacent areas. Coupling between the propagule bank and vegetation changed between sampling times with either some new coupling or decoupling. At both sites, the propagule bank contained propagules of the target community and thus propagule bank development during restoration provides increased potential for vegetation recovery. However, it can take a considerable time for management effects to be detected in the propagule bank. Moreover, the effect or speed of effect is spatially variable. Continuing application of restoration treatments is recommended at heathland where there is a deep bracken litter layer.     

Determinants of vertebrate invasion success in Europe and North America

Species that are frequently introduced to an exotic range have a high potential of becoming invasive. Besides propagule pressure, however, no other generally strong determinant of invasion success is known. Although evidence has accumulated that human affiliates (domesticates, pets, human commensals) also have high invasion success, existing studies do not distinguish whether this success can be completely explained by or is partly independent of propagule pressure. Here, we analyze both factors independently, propagule pressure and human affiliation. We also consider a third factor directly related to humans, hunting, and 17 traits on each species' population size and extent, diet, body size, and life history. Our dataset includes all 2362 freshwater fish, mammals, and birds native to Europe or North America. In contrast to most previous studies, we look at the complete invasion process consisting of (1) introduction, (2) establishment, and (3) spread. In this way, we not only consider which of the introduced species became invasive but also which species were introduced. Of the 20 factors tested, propagule pressure and human affiliation were the two strongest determinants of invasion success across all taxa and steps. This was true for multivariate analyses that account for intercorrelations among variables as well as univariate analyses, suggesting that human affiliation influenced invasion success independently of propagule pressure. Some factors affected the different steps of the invasion process antagonistically. For example, game species were much more likely to be introduced to an exotic continent than nonhunted species but tended to be less likely to establish themselves and spread. Such antagonistic effects show the importance of considering the complete invasion process.     

Invasibility of a Nutrient-Poor Pasture through Resident and Non-Resident Herbs Is Controlled by Litter, Gap Size and Propagule Pressure

Since inference concerning the relative effects of propagule pressure, biotic interactions, site conditions and species traits on the invasibility of plant communities is limited, we carried out a field experiment to study the role of these factors for absolute and relative seedling emergence in three resident and three non-resident confamilial herb species on a nutrient-poor temperate pasture. We set up a factorial field experiment with two levels each of the factors litter cover (0 and 400 g m(-2)), gap size (0.01 and 0.1 m(2)) and propagule pressure (5 and 50 seeds) and documented soil temperature, soil water content and relative light availability. Recruitment was recorded in spring and autumn 2010 and in spring 2011 to cover initial seedling emergence, establishment after summer drought and final establishment after the first winter. Litter alleviated temperature and moisture conditions and had positive effects on proportional and absolute seedling emergence during all phases of recruitment. Large gaps presented competition-free space with high light availability but showed higher temperature amplitudes and lower soil moisture. Proportional and absolute seedling recruitment was significantly higher in large than in small gaps. In contrast, propagule pressure facilitated absolute seedling emergence but had no effects on proportional emergence or the chance for successful colonisation. Despite significantly higher initial seedling emergence of resident than non-resident species, seed mass and other species-specific traits may be better predictors for idiosyncratic variation in seedling establishment than status. Our data support the fluctuating resource hypothesis and demonstrate that the reserve effect of seeds may facilitate seedling emergence. The direct comparison of propagule pressure with other environmental factors showed that propagule pressure affects absolute seedling abundance, which may be crucial for species that depend on other individuals for sexual reproduction. However, propagule batch size did not significantly affect the chance for successful colonisation of disturbed plots.     

Propagule pressure and environmental conditions interact to determine establishment success of an invasive plant species,glossy buckthorn (<Emphasis Type="Italic">Frangula alnus</Emphasis>), across five different wetland habitat types

Many invasive plant species are able to establish within a wide range of community types. This establishment success depends on high propagule pressure and successful recruitment of seedlings following propagule dispersal into receptive environments. This study aimed to investigate interactions between propagule pressure and environmental resistance to seedling recruitment of the invasive shrub, glossy buckthorn (Frangula alnus Mill.), over a range of wetland habitat types. We measured propagule deposition using seed traps and recruitment success using sown plots, while characterizing vegetation and abiotic environmental conditions in five adjacent wetland habitat types. Drier habitats, which included Cedar Swamp, Shrub Carr, and String, had lower resistance to buckthorn establishment than the wetter Flark and Cattail Marsh. The drier habitats supported more woody species and provided more raised hummock surfaces essential for successful buckthorn recruitment and establishment. Propagule pressure was also higher in dry habitats that supported higher densities of adult glossy buckthorn, while long-distance dispersal into areas with low adult density was uncommon. Natural recruitment was highest in sites with intense propagule pressure, but experimental sowing of seeds demonstrated that buckthorn establishes in wet sites with higher resistance if propagule pressure is increased and seeds are deposited on hummocks. This study demonstrates the affinity of glossy buckthorn for drier wetland sites, and provides empirical evidence that environmental resistance can be overcome by higher propagule pressure.     

Climate change can reduce the risk of biological invasion by reducing propagule size

Climate change has been conclusively linked to species extinctions, and to expansion and contractions and shifts of species ranges. Climate change is exerting similarly profound pressures on the individual stages of biological invasion which can significantly impact the biodiversity and ecology of invaded areas. Propagule pressure is perhaps the single most important determinant of invasion success, but the effects of climate change on propagule pressure are still largely uncertain because we have few observations of introduction events (or their size) that can be analyzed together with climate records. The common surrogate variables for propagule pressure do not logically respond to climate. Here I use a process-based simulation model to examine the potential effects of climate change (specifically temperature) on propagule size of a common invading insect species by estimating in-transit survivorship rate of propagules using historical and future (projected) temperatures and two common trade routes between a donor and a recipient location (Yokohama, Japan and Sydney, Australia). Propagule size (=the number of individuals in an introduction event) was lower under climate change temperatures than under historical temperatures in both routes. The route had significant effects on propagule size through its influence on the duration (and also the timing) of exposure to temperature conditions that are of time-sensitive importance to the development of the invasive species. Under historical temperatures propagule size was higher and less variable in the direct than the indirect route in 20 independent iterations. Under the future temperatures propagule size was also higher in the direct route but it was more variable than in the indirect route. Increased trade is increasing the opportunities for introductions, but the results reported here suggest that climate change will have inconsistent effects on biological invasion because of the complex relationship between temperature and insect ontogeny.     

Interactions between propagule pressure and seed traits shape human-mediated seed dispersal along roads

Human-mediated dispersal along the road network is a crucial process in the population dynamics of roadside vegetation and during plant invasions. The potential for a species to be dispersed by vehicles is, however, difficult to quantify. The predictive power of categorical classification schemes of human-mediated dispersal is limited as many species that are usually attributed to particular primary dispersal vectors may become subject to very different secondary dispersal vectors owing to human activity. Analysing seed traits that promote seed transport by human dispersal vectors could overcome these limitations. However, the analysis has to account for the divergent chance of seed transport that results from different propagule pressures along the transport corridor.To reveal the effects of seed traits and their interplay with propagule pressure on the chance and magnitude of human-mediated dispersal by vehicles, we compared traits and regional frequencies of a set of species that were dispersed by vehicles to a control set not dispersed but present in the same study area. We then used the same traits for a comparison of intentionally and unintentionally introduced species with the flora of Berlin.Different traits influenced the chance of vehicle dispersal and its magnitude. While propagule pressure was most important for determining the magnitude of seed transport, small seed mass and size best predicted the absolute chance of species dispersal by vehicles. The dispersal of nonnative species was least dependent on propagule pressure.Seed traits that were important in vehicle dispersal were similarly reflected in unintentionally introduced species in the Berlin flora. Mean seed size of these species was lower compared to the entire Berlin flora, whereas it was higher for intentionally introduced species. This suggests that unintentional introduction of nonnative plant species pre-selects for seed traits that promote further spread by human-mediated adhesive dispersal.Probability and magnitude of adhesive seed transport by vehicles can be predicted by dispersal-related plant traits. However, the effect size of plant traits on dispersal strongly depends on regional propagule pressure. This highlights the need to analyse interactions between species traits and propagule pressure.     

Decomposing propagule pressure: the effects of propagule size and propagule frequency on invasion success

Propagule pressure quantifies the inflow of individuals to a location and appears to be a key driver of invasion success. It is often defined as the average number of individuals introduced per time unit, or equivalently as the product of the average number of individuals introduced per introduction event (propagule size) and the frequency of introduction events (propagule frequency). Here we study how the influence of propagule size, frequency, and their product depends on the underlying ecological conditions. While previous studies have focused on introductions under environmental heterogeneity or a strong Allee effect, we examine a range of ecological scenarios that differ in the type of density dependence and in the sign of per capita growth rate. Our results indicate that the relative influence of propagule size and frequency depends mainly on the sign of per capita growth rate. Given a certain average number of individuals introduced per time unit, a high propagule frequency accelerates invasions under ecological scenarios with positive average per capita growth rate throughout the invasion process (‘easy’ scenarios). If per capita growth rate is negative throughout the invasion process (‘difficult’ scenarios) or if there is both an easy and a difficult stage (‘mixed scenarios’), a high propagule size leads to a faster invasion than a high propagule frequency. To explain this finding, we argue that for a fixed value of the product of propagule size and frequency, an increase in propagule size leads to an increase in demographic variance, which promotes invasion success in difficult and mixed but not in easy scenarios. However, we also show that in many of these cases, the product of propagule size and frequency still correlates more strongly with invasion success than either of the single components. Finally, we illustrate our approach with empirical examples from the literature.     

Colonization in metapopulations: a review of theory and observations

In metapopulation dynamics turnover of populations in isolated patches may be frequent. Regional survival of a species in such a system with frequent extinctions hinges on its colonization ability. Colonization is more than just dispersal; when a propagule reaches a new patch it faces higher extinction probabilities than does an established population. Extinction models as well as empirical data suggest that a large propagule with a potential for rapid increase in a varying environment, or with a low mortality rate in an environment perceived as constant, has a higher probability of successful colonization. Large variation in population size when it is still small increases the risk of failure. Factors introducing such variation are demographic stochasticity and environmental variation. It is hard to single out demographic traits that ensure good colonizing ability, since colonization can be achieved in many different ways, but generalists and species with self-fertilization seem to be superior.     

Vegetative Propagule Pressure and Water Depth Affect Biomass and Evenness of Submerged Macrophyte Communities

Vegetative propagule pressure may affect the establishment and structure of aquatic plant communities that are commonly dominated by plants capable of clonal growth. We experimentally constructed aquatic communities consisting of four submerged macrophytes (Hydrilla verticillata, Ceratophyllum demersum, Elodea nuttallii and Myriophyllum spicatum) with three levels of vegetative propagule pressure (4, 8 and 16 shoot fragments for communities in each pot) and two levels of water depth (30 cm and 70 cm). Increasing vegetative propagule pressure and decreasing water level significantly increased the growth of the submerged macrophyte communities, suggesting that propagule pressure and water depth should be considered when utilizing vegetative propagules to re-establish submerged macrophyte communities in degraded aquatic ecosystems. However, increasing vegetative propagule pressure and decreasing water level significantly decreased evenness of the submerged macrophyte communities because they markedly increased the dominance of H. verticillata and E. nuttallii, but had little impact on that of C. demersum and M. spicatum. Thus, effects of vegetative propagule pressure and water depth are species-specific and increasing vegetative propagule pressure under lower water level can facilitate the establishment success of submerged macrophyte communities.     

Ecological considerations in producing and formulating fungal entomopathogens for use in insect biocontrol

Insect pests persist in a wide-variety of agricultural, arboreal and urban environments. Effective control with fungal entomopathogens using inundation biocontrol requires an understanding of the ecology of the target insect, fungal pathogen, and the insect-pathogen interaction. Historically, the development of production and formulation processes for biocontrol fungi has primarily focused on reducing costs by maximizing the yield of infective propagules, increasing storage stability, and improving product form for ease of application. These goals are critical for commercialization but are often in conflict with environmental and ecological considerations. Critical parameters for selecting a fungal pathogen for use in inundation biocontrol include the cost-effective production of a stable, infective propagule that is suited for use in the environment where the insect must be controlled. Production processes can be manipulated nutritionally and environmentally to produce efficacious propagules or to direct fungal differentiation to propagule forms that may be better suited for use in specific environments. Formulation development must also consider ecological and environmental factors to maximize biocontrol efficacy. A basic understanding of the surface chemistries of the fungal propagule and insect, the interactions between a fungal propagule and the insect cuticle that lead to infection, and the impact of the environment on this interaction can aid in the development of effective formulations.     

Strength in size not numbers: propagule size more important than number in sexually reproducing populations

Propagule pressure has consistently been identified as a primary factor in invader success, and reducing it can be one of the most effective methods for preventing the establishment of non-native species. However, when policy is implemented to reduce propagule pressure it almost exclusively focuses on the size of individual introduction events (‘propagule size’), with little confirmation that controlling this single aspect of propagule pressure is the most effective strategy. The number of introduction events (‘propagule number’) can play as much, or more, of a role in invader success, yet only a small portion of propagule pressure research has studied the relative importance of size and number. We investigated the relative roles of propagule size and number in the establishment of a sexually reproducing species using a field mesocosm experiment that introduced Hemimysis anomala (a non-native mysid) across a range of propagule sizes and numbers. We found that single, large introductions had higher abundances and probabilities of survival than smaller, more frequent additions. This experiment illustrated that, for sexual reproducers, focusing on lowering propagule size can be the most effective method for reducing non-native establishment.     

Differentiation of adult rat bone marrow stem cells into epithelial progenitor cells in culture

We have previously obtained monoclonal bone marrow stem cells from adult rats (rMSCs) and induced them into phenotypic neurons. In the present study, we aimed to induce rMSCs into epithelial cells by culturing them onto compartmentalized permeable supports, which have been used for growing a variety of polarized epithelia in culture. Hematoxylin staining showed that after 4 days grown on permeable supports, rMSCs formed an epithelial-like monolayer. Immunofluorescence of the permeably-supported monolayers, but not the rMSCs grown in culture flasks, showed positive signals for epithelial markers, cytokeratin 5 & 8. RT-PCR results also showed the mRNA expression of epithelial sodium channel (ENaC) and cystic fibrosis transmembrane conductance regulator (CFTR) as well as tight junction protein ZO-1 in the rMSC-derived monolayers grown on permeable supports but absent from those grown in culture flasks. However, western blot only detected protein expression of ZO-1 but not ENaC nor CFTR. The short-circuit current measurements showed that the rMSC-derived monolayers grown on permeable supports exhibited a trans-monolayer resistance of 30-50 Omega cm(2); however, the monolayers did not respond to activators or blockers of CFTR or ENaC. The results suggest that compartmentalized or polarized culture conditions provide a suitable environment for rMSCs to differentiate into epithelial progenitor cells with tight junction formation; however, this condition is not sufficient for functional expression of epithelial ion channels associated with well-differentiated epithelia.     

Role of propagule pressure in colonization success: disentangling the relative importance of demographic,genetic and habitat effects

High propagule pressure is arguably the only consistent predictor of colonization success. More individuals enhance colonization success because they aid in overcoming demographic consequences of small population size (e.g. stochasticity and Allee effects). The number of founders can also have direct genetic effects: with fewer individuals, more inbreeding and thus inbreeding depression will occur, whereas more individuals typically harbour greater genetic variation. Thus, the demographic and genetic components of propagule pressure are interrelated, making it difficult to understand which mechanisms are most important in determining colonization success. We experimentally disentangled the demographic and genetic components of propagule pressure by manipulating the number of founders (fewer or more), and genetic background (inbred or outbred) of individuals released in a series of three complementary experiments. We used Bemisia whiteflies and released them onto either their natal host (benign) or a novel host (challenging). Our experiments revealed that having more founding individuals and those individuals being outbred both increased the number of adults produced, but that only genetic background consistently shaped net reproductive rate of experimental populations. Environment was also important and interacted with propagule size to determine the number of adults produced. Quality of the environment interacted also with genetic background to determine establishment success, with a more pronounced effect of inbreeding depression in harsh environments. This interaction did not hold for the net reproductive rate. These data show that the positive effect of propagule pressure on founding success can be driven as much by underlying genetic processes as by demographics. Genetic effects can be immediate and have sizable effects on fitness.     

A new method for cell permeabilization reveals a cytosolic protein requirement for Ca2+ -activated secretion in GH3 pituitary cells

Ca2+ is a major regulator of exocytosis in secretory cells, however, the biochemical mechanisms underlying regulation remain to be identified. To render the secretory apparatus accessible for biochemical studies, we have developed a cell permeabilization method (cell cracking) which utilizes mechanical shear. GH3 pituitary cells subjected to cracking were permeable to macromolecules but retained a normal cytoplasmic ultrastructure including secretory granules. Incubation of the permeable cells at 30-37 degrees C with 0.1-1.0 microM Ca2+ and millimolar MgATP resulted in the release of the secretory proteins, prolactin (PRL) and a proteoglycan, but not lysosomal enzymes. Extensively washed permeable cells were incapable of releasing PRL in response to Ca2+ and MgATP addition. However, addition of cytosol was found to restore Ca2+-activated, MgATP-dependent PRL release. The cytosolic factor responsible for activity was thermolabile and protease sensitive. The protein was partially purified, and its molecular mass was estimated to be equivalent to that of a globular protein of 200-350 kDa by molecular sieve chromatography. Inhibitors of calmodulin or protein kinase C (trifluroperazine, calmidazolium, H-7) failed to inhibit Ca2+-activated PRL release, and the required cytosolic protein could not be replaced by purified calmodulin, calmodulin-dependent protein kinase II, protein kinase C, or calpactin I. Further purification and characterization of the cytosolic protein should reveal the nature of biochemical events involved in regulated secretory exocytosis.     

Recovery of ΔF508-CFTR function by analogs of hyaluronan disaccharide

We recently discovered that hyaluronan was exported from fibroblasts by MRP5 and from epithelial cells by cystic fibrosis (CF) transmembrane conductance regulator (CFTR) that was known as a chloride channel. On this basis we developed membrane permeable analogs of hyaluronan disaccharide as new class of compounds to modify their efflux. We found substances that activated hyaluronan export from human breast cancer cells. The most active compound 2-(2-acetamido-3,5-dihydroxyphenoxy)-5-aminobenzoic acid (Hylout4) was tested for its influence on the activity of epithelial cells. It activated the ion efflux by normal and defective ΔF508-CFTR. It also enhanced the plasma membrane concentration of the ΔF508-CFTR protein and reduced the transepithelial resistance of epithelial cells. In human trials of healthy persons, it caused an opening of CFTR in the nasal epithelium. Thus compound Hylout4 is a corrector that recovered ΔF508-CFTR from intracellular degradation and activated its export function.     

Propagule banks, herbivory and nutrient supply control population development and dominance patterns in macroalgal blooms

Destructive macroalgal mass blooms threaten estuarine and coastal ecosystems worldwide. We asked which factors regulate macroalgal bloom intensity, distribution and species composition. In field experiments in the Baltic Sea, we analyzed the relative effects of nutrients, herbivores and algal propagule banks on population development and dominance patterns in two co-occurring bloom-forming macroalgae, Enteromorpha intestinalis and Pilayella littoralis . Both species were highly affected by the combined effects of a propagule bank, herbivory and nutrients. The magnitude of effects varied with season. The propagule bank was an important overwintering mechanism for both algae, and allowed for recruitment two months earlier than recruitment via freshly dispersed propagules. This provided a seasonal escape from intense herbivory and nutrient limitation later in the year. Favored by massive recruitment from the propagule bank, Enteromorpha was the superior space occupier in early spring, thereby reducing recruitment of Pilayella . Elimination of the propagule bank and recruitment via freshly dispersed propagules favored Pilayella . Strong and selective herbivory on Enteromorpha supported Pilayella in the presence, but not in the absence of the propagule bank. Nutrient enrichment in summer counteracted herbivore pressure on Enteromorpha , thereby negatively affecting Pilayella . Herbivore and nutrient effects were more pronounced for early life stages than adult algae. These results show that recruitment processes and forces affecting early life stages at the beginning of the vegetation period determine development and dominance patterns of macroalgal blooms. Herbivores naturally suppress blooms but increasing nutrient enrichment can override this important control mechanism. The propagule bank plays a previously unrecognized role for population and community dynamics.     

Changes along a disturbance gradient in the density and composition of propagule banks in floodplain aquatic habitats

This study used germination methods to examine the density, species composition and functional composition of propagule banks in a series of riverine wetland aquatic habitats subject to varying degrees of hydrological and management-related disturbance. Under permanent inundation (the conditions prevailing at most sites during the growing season) propagule germination and species richness was low, with floodplain perennials and helophytes particularly affected. Densities of floodplain annuals were largely maintained through continued germination of a few flooding tolerant species. On damp mud (conditions associated with hydrological instability) total seedling number and species richness increased significantly, but species richness of germinating hydrophytes declined. Mean seedling density at 0–0.1m depth was 15450 ± 4400 m–2, reaching a maximum (162 050 m–2) in temporary backwaters. Annual (e.g., Lindernia dubia, Cyperus fuscus) and facultative ruderal species (e.g., Lythrum salicaria and Alisma plantago-aquatica) predominated. Vertical zonation of the propagule bank was weakly developed. The numbers of individuals and species germinating varied significantly between sites. The seasonal, most intensely disturbed sites (temporary backwaters) supported a numerically large, species-rich propagule bank based on floodplain annuals, while the permanent, less disturbed sites (ditches and an oxbow pond) had a small, species-poor propagule bank composed of hydrophytes and helophytes supplemented by allochthonous seed inputs. Sites intermediate on the gradient had a propagule bank dominated by facultative amphibious, ruderal hydrophytes. The composition of the seed bank and the established vegetation was most similar at the heavily disturbed sites where the seed bank was maintained by vigorously fruiting annuals and supplemented by inputs from temporary habitats upstream. At permanent sites much of the propagule bank composition could be accounted for by inputs of floodborne seed from the immediately adjacent floodplain. The established vegetation at such sites appeared to be maintained mainly by vegetative propagation with recruitment from the propagule bank likely only after severe disturbance. The potential contribution of functionally diverse propagule banks to sucessional processes within fluvially dynamic floodplain aquatic habitats is emphasised.