Relationship between propagule pressure and colonization pressure in invasion ecology: a test with ships' ballast

Increasing empirical evidence indicates the number of released individuals (i.e. propagule pressure) and number of released species (i.e. colonization pressure) are key determinants of the number of species that successfully invade new habitats. In view of these relationships, and the possibility that ships transport whole communities of organisms, we collected 333 ballast water and sediment samples to investigate the relationship between propagule and colonization pressure for a variety of diverse taxonomic groups (diatoms, dinoflagellates and invertebrates). We also reviewed the scientific literature to compare the number of species transported by ships to those reported in nature. Here, we show that even though ships transport nearly entire local communities, a strong relationship between propagule and colonization pressure exists only for dinoflagellates. Our study provides evidence that colonization pressure of invertebrates and diatoms may fluctuate widely irrespective of propagule pressure. We suggest that the lack of correspondence is explained by reduced uptake of invertebrates into the transport vector and the sensitivity of invertebrates and diatoms to selective pressures during transportation. Selection during transportation is initially evident through decreases in propagule pressure, followed by decreased colonization pressure in the most sensitive taxa.     

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.     

The more you introduce the more you get: the role of colonization pressure and propagule pressure in invasion ecology

Aim  We argue that 'propagule pressure', a key term in invasion biology, has been attributed at least three distinct definitions (with usage of a related term causing additional confusion). All of the definitions refer to fundamental concepts within the invasion process, with the result that the distinct importance of these different concepts has been at best diluted, and at worst lost.
Location  Global.
Methods  We reviewed pertinent literature on propagule pressure to resolve confusion about different uses of the term 'propagule pressure' and we introduced a new term for one variant, colonization pressure. We conducted a computer simulation whereby the introduction of species is represented as a simple sampling process to elucidate the relationship between propagule and colonization pressure.
Results  We defined colonization pressure as the number of species introduced or released to a single location, some of which will go on to establish a self-sustaining population and some of which will not. We subsequently argued that colonization pressure should serve as a null hypothesis for understanding temporal or spatial differences in exotic species richness, as the more species that are introduced, the more we should expect to establish. Finally, using a simple simulation, we showed that propagule pressure is related to colonization pressure, but in a non-linear manner.
Main conclusion  We suggest that the nature of the relationship between propagule pressure and colonization pressure, as well as the efficacy of various proxy measures of each, require more detailed exploration if invasion ecology is to continue to develop into a more predictive science.     

Towards an urban marine ecology: characterizing the drivers,patterns and processes of marine ecosystems in coastal cities

Human population density within 100 km of the sea is approximately three times higher than the global average. People in this zone are concentrated in coastal cities that are hubs for transport and trade – which transform the marine environment. Here, we review the impacts of three interacting drivers of marine urbanization (resource exploitation, pollution pathways and ocean sprawl) and discuss key characteristics that are symptomatic of urban marine ecosystems. Current evidence suggests these systems comprise spatially heterogeneous mosaics with respect to artificial structures, pollutants and community composition, while also undergoing biotic homogenization over time. Urban marine ecosystem dynamics are often influenced by several commonly observed patterns and processes, including the loss of foundation species, changes in biodiversity and productivity, and the establishment of ruderal species, synanthropes and novel assemblages. We discuss potential urban acclimatization and adaptation among marine taxa, interactive effects of climate change and marine urbanization, and ecological engineering strategies for enhancing urban marine ecosystems. By assimilating research findings across disparate disciplines, we aim to build the groundwork for urban marine ecology – a nascent field; we also discuss research challenges and future directions for this new field as it advances and matures. Ultimately, all sides of coastal city design: architecture, urban planning and civil and municipal engineering, will need to prioritize the marine environment if negative effects of urbanization are to be minimized. In particular, planning strategies that account for the interactive effects of urban drivers and accommodate complex system dynamics could enhance the ecological and human functions of future urban marine ecosystems.     

Hotspots of plant invasion predicted by propagule pressure and ecosystem characteristics

Aim Biological invasions pose a major conservation threat and are occurring at an unprecedented rate. Disproportionate levels of invasion across the landscape indicate that propagule pressure and ecosystem characteristics can mediate invasion success. However, most invasion predictions relate to species’ characteristics (invasiveness) and habitat requirements. Given myriad invaders and the inability to generalize from single‐species studies, more general predictions about invasion are required. We present a simple new method for characterizing and predicting landscape susceptibility to invasion that is not species‐specific. Location Corangamite catchment (13,340 km²), south‐east Australia. Methods Using spatially referenced data on the locations of non‐native plant species, we modelled their expected proportional cover as a function of a site’s environmental conditions and geographic location. Models were built as boosted regression trees (BRTs). Results On average, the BRTs explained 38% of variation in occupancy and abundance of all exotic species and exotic forbs. Variables indicating propagule pressure, human impacts, abiotic and community characteristics were rated as the top four most influential variables in each model. Presumably reflecting higher propagule pressure and resource availability, invasion was highest near edges of vegetation fragments and areas of human activity. Sites with high vegetation cover had higher probability of occupancy but lower proportional cover of invaders, the latter trend suggesting a form of biotic resistance. Invasion patterns varied little in time despite the data spanning 34 years. Main conclusions To our knowledge, this is the first multispecies model based on occupancy and abundance data used to predict invasion risk at the landscape scale. Our approach is flexible and can be applied in different biomes, at multiple scales and for different taxonomic groups. Quantifying general patterns and processes of plant invasion will increase understanding of invasion and community ecology. Predicting invasion risk enables spatial prioritization of weed surveillance and control.     

Biological control as an invasion process: disturbance and propagule pressure affect the invasion success of Lythrum salicaria biological control agents

Understanding the mechanisms behind the successful colonization and establishment of introduced species is important for both preventing the invasion of unwanted species and improving release programs for biological control agents. However, it is often not possible to determine important introduction details, such as date, number of organisms, and introduction location when examining factors affecting invasion success. Here we use biological control introduction data to assess the role of propagule pressure, disturbance, and residence time on invasion success of four herbivorous insect species introduced for the control of the invasive wetland plant, Lythrum salicaria, in the Columbia River Estuary. Two sets of field surveys determined persistence at prior release sites, colonization of new sites, and abundance within colonized sites. We quantified propagule pressure in four ways to examine the effect of different measurements. These included three measurements of introduction size (proximity to introduction site, introduction size at a local scale, and introduction size at a regional scale) and one measure of introduction number (number of introduction events in a region). Disturbance was examined along a tidal inundation gradient (distance from river mouth) and as habitat (island or mainland). Statistical models and model averaging were used to determine which factors were driving invasion success. In this study we found: (1) sparse evidence for the positive influence of propagule pressure on invasion success; (2) disturbance can negatively affect the invasion success of herbivorous insects; (3) the effects of disturbance and propagule pressure are species specific and vary among invasion stages, and (4) not all measures of propagule pressure show the same results, therefore single measures and proxies should be used cautiously.     

Divergent effects of land-use,propagule pressure,and climate on woody riparian invasion

Landscape-scale analyses of biological invasion are needed to understand the relative importance of environmental drivers that vary at larger scales, such as climate, propagule pressure, resource availability, and human disturbance. One poorly understood landscape-scale question is, how does human land-use influence riparian plant invasion? To evaluate the relative importance of land-use, climate, propagule pressure, and water availability in riparian invasion, we examined tamarisk (Tamarix ramosissima, T. chinensis, hybrids), Russian olive (Elaeagnus angustifolia), and Siberian elm (Ulmus pumila) occurrence, abundance, and dominance in 238 riparian sites in developed, cultivated, and undeveloped areas of four western USA river basins (281,946 km²). Temperature and propagule pressure from individuals planted nearby largely drove invasive species occurrence, whereas factors likely to affect resource availability (e.g., land-use, precipitation, streamflow intermittency) were more important to abundance and dominance, supporting the argument that species distribution models based on occurrence alone may fail to identify conditions where invasive species have the greatest impact. The role of land-use varied among taxa: urban and suburban land-use increased Siberian elm occurrence, abundance, and dominance, and urban land-use increased Russian olive occurrence, whereas suburban land-use reduced tamarisk dominance. Surprisingly, Siberian elm, which has received scant prior scientific and management attention, occurred as or more frequently than tamarisk and Russian olive (except in undeveloped areas of the Colorado River headwaters) and had higher density and dominance than tamarisk and Russian olive in developed areas. More research is needed to understand the impacts of this largely unrecognized invader on riparian ecosystem services, particularly in urban and suburban areas.     

Spatially variable propagule pressure and herbivory influence invasion of chaparral shrubland by an exotic grass

Although numerous studies have identified mechanisms that either resist or facilitate biological invasions, few studies have explicitly tested how resisting and facilitating mechanisms interact to drive invasion success. In California, USA, undisturbed Mediterranean-type shrublands have resisted invasion by the perennial tussock grass Cortaderia jubata. In some cases, however, this resistance has been spectacularly breached even in the absence of large-scale disturbance. I tested the hypothesis that these invasions are facilitated by local reductions in the strength of biotic resistance. I evaluated invasive success using C. jubata seed and seedling additions at different microhabitats: the edge of a chaparral stand, under shrub canopy at different distances from the stand edge, and in canopy gaps within the stand. When left exposed to mammalian herbivores, seedling survivorship decreased sharply from nearly 40% on the stand edge to zero just 10 m into the stand. When transplants were protected from herbivory, however, distance from the edge had no significant influence on transplant survivorship. Seedling emergence was also greater on the edge and in canopy gaps than under the canopy, but these differences were not caused by differences in herbivory. The flux of invasive propagules reaching the soil surface was immense and greater along the edge and within gaps than under the stand canopy. Mirroring these patterns, naturally occurring seedling abundance declined dramatically with distance from the stand edge, and seedlings were far more common within stand gaps than would be expected given gap frequency within the stand. Despite strong biotic resistance to invasion within the stand, the cover of C. jubata has increased 20% over the last 9 years. These results suggest that the relative amount of susceptible edge habitat and the supply of invasive propagules can facilitate invasion even in the face of strong local biotic resistance.     

The vulnerability of habitats to plant invasion: disentangling the roles of propagule pressure,time and sampling effort

Aim To quantify the vulnerability of habitats to invasion by alien plants having accounted for the effects of propagule pressure, time and sampling effort. Location New Zealand. Methods We used spatial, temporal and habitat information taken from 9297 herbarium records of 301 alien plant species to examine the vulnerability of 11 terrestrial habitats to plant invasions. A null model that randomized species records across habitats was used to account for variation in sampling effort and to derive a relative measure of invasion based either on all records for a species or only its first record. The relative level of invasion was related to the average distance of each habitat from the nearest conurbation, which was used as a proxy for propagule pressure. The habitat in which a species was first recorded was compared to the habitats encountered for all records of that species to determine whether the initial habitat could predict subsequent habitat occupancy. Results Variation in sampling effort in space and time significantly masked the underlying vulnerability of habitats to plant invasions. Distance from the nearest conurbation had little effect on the relative level of invasion in each habitat, but the number of first records of each species significantly declined with increasing distance. While Urban, Streamside and Coastal habitats were over‐represented as sites of initial invasion, there was no evidence of major invasion hotspots from which alien plants might subsequently spread. Rather, the data suggest that certain habitats (especially Roadsides) readily accumulate alien plants from other habitats. Main conclusions Herbarium records combined with a suitable null model provide a powerful tool for assessing the relative vulnerability of habitats to plant invasion. The first records of alien plants tend to be found near conurbations, but this pattern disappears with subsequent spread. Regardless of the habitat where a species was first recorded, ultimately most alien plants spread to Roadside and Sparse habitats. This information suggests that such habitats may be useful targets for weed surveillance and monitoring.     

The interaction between propagule pressure,habitat suitability and density‐dependent reproduction in species invasion

Seedling recruitment limitations create a demographic bottleneck that largely determines the viability and structure of plant populations and communities, and pose a core restriction on the colonization of novel habitat. We use a shade‐tolerant, invasive grass, Microstegium vimineum, to examine the interplay between seed and establishment limitations – phenomena that together determine recruitment success but usually are investigated individually. We add increasing amounts of seed to microhabitats containing variable levels of leaf litter thickness – with reduced leaf litter simulating disturbance – to investigate whether reduced seed limitation overcomes the establishment limitation posed by litter cover. We do this across gradients in understory light, moisture and temperature, and quantify germination, survival, and then per capita adult biomass and reproduction in order to understand the implications for invasion across the landscape. We find that the combined effects of seed and establishment limitation influence recruitment; however, propagule pressure overwhelms the inhibitory effects of leaf litter thickness. Leaf litter reduces germination by 22–57% and seedling survival by 13–15% from that observed on bare soil. However, density‐dependent reproduction compensates as 1–3 plants can produce far more seeds (approx. 525) than are required for persistence. As such, just a few plants may establish in understory forest habitat and subsequently overwhelm establishment barriers with copious propagule production. These results, for a widespread, invasive plant, are consistent with the emerging perspective for native plants that seed and establishment limitation jointly influence recruitment. The ability for an exotic plant species to compensate for low population densities with high per capita seed production, that then overrides establishment limitations, makes its invasive potential daunting. Further work is required to test if this is a common mechanism underlying plant invasions.     

Sulfate reduction in coastal ecosystems

The diversity and activity of dissimilatory Fe(III)-reducing bacteria was investigated in acidic, ochre-precipitating springs on Mam Tor, East Midlands, UK. The springs at this acid rock drainage site are located below a 3000 year old landslip, where biooxidation of exposed pyrite-containing minerals has resulted in the production of metal-laden acidic waters. A diverse microbial community was found downstream in the sediments dominated by Fe(III) minerals, and included close relatives to known acidophilic (Acidimicrobium and Acidiphilium) and neutraphilic (Geobacter and Pelobacter) Fe(III)-reducing bacteria. Analysis by XRD and TEM confirmed the presence of both amorphous and well-defined Fe(III) mineral phases in the sediments including lepidocrocite, goethite and schwertmannite. Microcosm-based experiments demonstrated that the bioavailable Fe(III) was reduced under anaerobic conditions, concomitant with sulphate release. XRD analysis suggested that schwertmannite (an iron sulphate hydroxide) was utilized preferentially by the Fe(III)-reducing bacteria, leading to the release of sulphate. Although the microcosms contained sufficient concentrations of naturally occurring electron donor to sustain significant levels of Fe(III) reduction, this process was stimulated by the addition of glycerol and complex electron donors. Thus, the acidic Fe(III)-containing sediments contain a diversity of DIRBs that can be stimulated by the addition of electron donor as a first step in the reversal of acid rock and acid mine drainage contamination.     

Seasonality in coastal benthic ecosystems

For historical reasons, knowledge about seasonality in the dynamics of marine benthic suspension feeders from temperate areas comes mainly from studies of cold temperate seas. Recent surveys of Mediterranean taxa show different patterns from those observed in cold temperate seas, which are characterized by winter dormancy. In the Mediterranean, summer dormancy predominates among taxa and appears to be related to energetic constraints. Temperature and food availability are crucial to the dynamics of benthic suspension feeders. However, because these factors tend to be positively correlated in cold temperate seas, it is difficult to distinguish between their effects. Such correlation does not occur in Mediterranean ecosystems. The contrast between recent studies in the Mediterranean and in other areas can help to disentangle confounded environmental controls.     

The role of propagule pressure in the invasion success of bluegill sunfish, Lepomis macrochirus, in Japan

The bluegill sunfish, Lepomis macrochirus, is a widespread exotic species in Japan that is considered to have originated from 15 fish introduced from Guttenberg, Iowa, in 1960. Here, the genetic and phenotypic traits of Japanese populations were examined, together with 11 native populations of the USA using 10 microsatellite markers and six meristic traits. Phylogenetic analysis reconfirmed a single origin of Japanese populations, among which populations established in the 1960s were genetically close to Guttenberg population, keeping high genetic diversity comparable to the ancestral population. In contrast, genetic diversity of later-established populations significantly declined with genetic divergence from the ancestral population. Among the 1960s established populations, that from Lake Biwa showed a significant isolation-by-distance pattern with surrounding populations in which genetic bottlenecks increased with geographical distance from Lake Biwa. Although phenotypic divergence among populations was recognized in both neutral and adaptive traits, P(ST)-F(ST) comparisons showed that it is independent of neutral genetic divergence. Divergent selection was suggested in some populations from reservoirs with unstable habitats, while stabilizing selection was dominant. Accordingly, many Japanese populations of L. macrochirus appear to have derived from Lake Biwa population, expanding their distribution with population bottlenecks. Despite low propagule pressure, the invasion success of L. macrochirus is probably because of its drastic population growth in Lake Biwa shortly after its introduction, together with artificial transplantations. It not only enabled the avoidance of a loss in genetic diversity but also formed a major gene pool that supported local adaptation with high phenotypic plasticity.     

Niche based distribution modelling of an invasive alien plant: effects of population status, propagule pressure and invasion history

Forecasting the spatial spread of invasive species is important to inform management planning. Niche-based species distribution models offer a well-developed framework for assessing the potential range of species. However, these models assume equilibrium between the species’ distribution and its ecological requirements. During range expansion, invasive species are not in such equilibrium due to both dispersal limitation and frequent casual occurrence in sites unsuitable to persistent populations. In this article we use the example of the invasive annual plant Ambrosia artemisiifolia in Austria to evaluate if model accuracy can be enhanced in such non-equilibrium situations by taking account of propagule pressure and by restricting model calibration to naturalized populations. Moreover, we test if model accuracy increases during invasion history using distribution data from 1984 to 2005. The results suggest that models calibrated with naturalized populations are much more accurate than those based on the total set of records. Proxies of propagule pressure slightly but significantly improve goodness of fit, accuracy, and Type I and II error rates of models calibrated with all available records but have less consistent effects on models of naturalized populations. Model accuracy did not increase during the recent invasion history, probably because the species is still far from an equilibrium distribution. We conclude that even a coarse assessment of population status with records of invasive species delivers important information for predictive modelling and that proxies of propagule pressure should be included into such models at least during early to intermediate stages of the invasion history.     

Nitrogen cycling in coastal marine ecosystems

It is generally considered that nitrogen availability is one of the major factors regulating primary production in temperate coastal marine environments. Coastal regions often receive large anthropogenic inputs of nitrogen that cause eutrophication. The impact of these nitrogen additions has a profound effect in estuaries and coastal lagoons where water exchange is limited. Such increased nutrient loading promotes the growth of phytoplankton and fast growing pelagic macroalgae while rooted plants (sea-grasses) and benthic are suppressed due to reduced light availability. This shift from benthic to pelagic primary production introduces large diurnal variations in oxygen concentrations in the water column. In addition oxygen consumption in the surface sediments increases due to the deposition of readily degradable biomass. In this review the physico-chemical and biological factors regulating nitrogen cycling in coastal marine ecosystems are considered in relation to developing effective management programmes to rehabilitate seagrass communities in lagoons currently dominated by pelagic macroalgae and/or cyanobacteria.