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 interplay of disturbance,vegetation structure,and propagule pressure contributes to Pinus kesiya invasion in Tapia woodland,Madagascar

Pinus are among the highly invasive species that have spread outside their plantation area after their introduction in the Southern Hemisphere. The case of Pinus kesiya invasion is observed in the high plateau of Madagascar, inside the sclerophyll Tapia woodland which is dominated by the endemic Uapaca bojeri tree species. The analysis of this invasion was carried out using 375 plots of 100 m² each in Tapia woodland. Data on the vegetation structure, the plot characteristics and the propagule pressure were collected. We recorded a total of 740 pines distributed in 29.8% of the plots. The generalized linear model built on P. kesiya at the three different life stages allowed us to highlight a different explicative variable on the species’ presence and abundance separately. The factors explaining pine occurrence varied according to the pine life stage. In the seedling stage, the distance of the plot from the propagule source combined with the longitudinal position of the plot explained 18% of the pine presence. In the sapling and adult stages, the vegetation structure was the main important factor (22% and 11% of variation explained regarding presence and abundance). The frequency of U. bojeri and the degree of disturbance were the most important factors characterizing this vegetation structure. Based on these results, a strategy to control pine invasion in the Tapia woodland may focus on enrichment with U. bojeri and limitation of the plantation of P. kesiya in proximity.

     

A native species does not prevent the colonization success of an introduced submerged macrophyte,even at low propagule pressure

Hydrobiologia - Invasions are a threat to biodiversity because non-native species are generally more competitive than native species. Hydrilla verticillata is an invasive macrophyte that causes...     

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.     

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.     

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.     

Fire persistence traits of plants along a productivity and disturbance gradient in mediterranean shrublands of south-east Australia

Aim To understand changes in fire persistence traits of plants along a latitudinal gradient, considering the interactions between productivity, community (fuel) structure and fire regime. Location A gradient in the south of Australia (latitude 33–37° S; longitude 140–143° E), including: Little Desert National Park (VIC), Big Desert Wilderness Park (VIC), Murray‐Sunset National Park (VIC), Danggali Conservation Park (SA) and Tarawi Nature Reserve (NSW). Methods We selected four areas along a latitudinal gradient for which information on fire history and vegetation was available. Then, we tested to what extent the four selected areas have different climate and different fire regimes. Plant cover values of different life forms provided an indication of the plant community structure and flammability, and the proportion of species with different fire persistence traits (resprouting, seedbank persistence) informed us on the trait selection. Results Precipitation decreases and temperature increases from south to north. Thus the selected sites represent a gradient from high productivity (low aridity) in the south to low productivity (high aridity) in the north. Fire statistics suggest that fire frequency parallels productivity. There is a tendency for life form dominance and community structure to shift in such a way that fuel connectivity is reduced towards the north. Resprouting species increase and obligate seeders decrease along the fire–productivity gradient. Main conclusions Changes in plant traits are difficult to understand without simultaneous consideration of both the disturbance and the productivity gradients. In our study area, fire regime and productivity interact in such a way that decreases in productivity imply changes in fuel structure that produce a reduction in fire frequency. Resprouting species are better represented at the high fire–productivity part of the gradient, while obligate seeders are better represented at the opposite end of the gradient. The results also emphasize the importance of considering not only climate changes but also changes in fuel structure to predict future fire regimes.     

The evolutionary dynamics of polyploid plants: origins, establishment and persistence

The role of polyploidy in the origin of evolutionary novelty and the maintenance of diversity in plant populations has come to be recognized as an integral component of the ecological and evolutionary dynamics of plant species populations. Recent attempts to examine the evolutionary significance of polyploidy have focused on the processes responsible for the origin of polyploid plants and the conditions that favour their establishment and persistence. The importance of these issues is not simply limited to the evolutionary dynamics of polyploidy but is, in fact, central to our understanding of the population biology processes that act on the establishment of new 'types' and the maintenance of biotic diversity at both the inter- and intraspecific levels.     

Antifungals: mechanism of action and resistance, established and novel drugs

Serious fungal infections, caused mostly by opportunistic species, are increasingly common in immunocompromised and other vulnerable patients. The use of antifungal drugs, primarily azoles and polyenes, has increased in parallel. Yet, established agents do not satisfy the medical need completely: azoles are fungistatic and vulnerable to resistance, whereas polyenes cause serious host toxicity. Drugs in clinical development include echinocandins, pneumocandins, and improved azoles. Promising novel agents in preclinical development include several inhibitors of fungal protein, lipid and cell wall syntheses. Recent advances in fungal genomics, combinatorial chemistry, and high-throughput screening may accelerate the antifungal discovery process.     

Population structure, propagule pressure, and conservation biogeography in the sub-Antarctic: lessons from indigenous and invasive springtails

The patterns in and the processes underlying the distribution of invertebrates among Southern Ocean islands and across vegetation types on these islands are reasonably well understood. However, few studies have examined the extent to which populations are genetically structured. Given that many sub‐Antarctic islands experienced major glaciation and volcanic activity, it might be predicted that substantial population substructure and little genetic isolation‐by‐distance should characterize indigenous species. By contrast, substantially less population structure might be expected for introduced species. Here, we examine these predictions and their consequences for the conservation of diversity in the region. We do so by examining haplotype diversity based on mitochondrial cytochrome c oxidase subunit I sequence data, from two indigenous (Cryptopygus antarcticus travei, Tullbergia bisetosa) and two introduced (Isotomurus cf. palustris, Ceratophysella denticulata) springtail species from Marion Island. We find considerable genetic substructure in the indigenous species that is compatible with the geological and glacialogical history of the island. Moreover, by employing ecological techniques, we show that haplotype diversity is likely much higher than our sequenced samples suggest. No structure is found in the introduced species, with each being represented by a single haplotype only. This indicates that propagule pressure is not significant for these small animals unlike the situation for other, larger invasive species: a few individuals introduced once are likely to have initiated the invasion. These outcomes demonstrate that sampling must be more comprehensive if the population history of indigenous arthropods on these islands is to be comprehended, and that the risks of within‐ and among‐island introductions are substantial. The latter means that, if biogeographical signal is to be retained in the region, great care must be taken to avoid inadvertent movement of indigenous species among and within islands. Thus, quarantine procedures should also focus on among‐island movements.     

Recruitment in species-rich grasslands: the effects of functional traits and propagule pressure

Aims The assembly of plant communities is a complex process which combines impacts from the species pool, dispersal and propagule pressure, niche requirements of colonizing species and the niche structure of the community. Recent theory development has incorporated all these aspects, e.g. in 'stochastic niche theory'. We investigated recruitment into a species-rich grassland community, using an experimental approach where we manipulated the trait composition of the community and examined the success of colonizing species entering with various propagule pressure. Specifically, we examined two predictions: (i) colonization success increases with increasing difference between traits of the colonizing species and the trait profile of the community and (ii) colonization success increases with increasing propagule pressure.Methods The examined communities were species-rich semi-natural grasslands located in southern Sweden. After a careful documentation of the composition of the plant communities at the experimental sites, we manipulated the trait profile of species-rich grassland plots based on the plant functional trait specific leaf area (SLA), which is correlated with several key life history functions. In addition to SLA, seed mass was also used to describe the trait profile of grassland plots. Seeds of 12 plant species from the regional species pool, varying in SLA and seed mass, were sown into plots using four different levels of propagule pressure. Recruitment was examined after 1 year. We also planted juvenile 'plug plants' of the same species which allowed us to examine survivorship and growth beyond the seedling stage.Important findings Overall we found very limited evidence for relationships between the traits of the colonizing species and the trait profile of the community and for recruitment after sowing these relationships were contrary to the prediction. Survival of plug plants after two seasons of growth was high irrespective of the trait profile of the community, but growth of plug plants was affected by the trait profile of the surrounding community. For four of the species there was a positive effect of increased propagule pressure on colonization. The results suggest that species assembly in species-rich grasslands is not strongly dependent on the niche structure of the community. However, the finding that colonization of only a third of the species responded positively to increased propagule pressure indicates that there might be niche-related effects that were not captured by our treatments. Overall, our results indicate that the factors determining colonization in this community are species specific. Some species are able to colonize irrespective of niche relationships, provided that the propagule pressure is sufficiently high to overcome stochastic mortality after seed arrival. For other species, however, we cannot exclude that niche assembly occurred, but we failed to identify the relevant niche factor.     

Propagule pressure and disturbance interact to overcome biotic resistance of marine invertebrate communities

Propagule pressure is fundamental to invasion success, yet our understanding of its role in the marine domain is limited. Few studies have manipulated or controlled for propagule supply in the field, and consequently there is little empirical data to test for non-linearities or interactions with other processes. Supply of non-indigenous propagules is most likely to be elevated in urban estuaries, where vessels congregate and bring exotic species on fouled hulls and in ballast water. These same environments are also typically subject to elevated levels of disturbance from human activities, creating the potential for propagule pressure and disturbance to interact. By applying a controlled dose of free-swimming larvae to replicate assemblages, we were able to quantify a dose–response relationship at much finer spatial and temporal scales than previously achieved in the marine environment. We experimentally crossed controlled levels of propagule pressure and disturbance in the field, and found that both were required for invasion to occur. Only recruits that had settled onto bare space survived beyond three months, precluding invader persistence in undisturbed communities. In disturbed communities initial survival on bare space appeared stochastic, such that a critical density was required before the probability of at least one colony surviving reached a sufficient level. Those that persisted showed 75% survival over the following three months, signifying a threshold past which invaders were resilient to chance mortality. Urban estuaries subject to anthropogenic disturbance are common throughout the world, and similar interactions may be integral to invasion dynamics in these ecosystems.     

Periodic local disturbance in host-parasitoid metapopulations: host suppression and parasitoid persistence

Within many agricultural systems, insect pests and their natural enemies are forced to persist as a metapopulation, continuously recolonizing patches following disturbance through harvesting or spraying with insecticides. Despite the need to understand factors influencing biocontrol success, few theoretical studies of host-parasitoid interactions have examined the potential impact of local disturbance within a metapopulation framework. Here, we add periodic local mortality to series of classical host-parasitoid models to examine its effect on host suppression and parasitoid persistence. Using a deterministic lattice model, we show that despite the wide range of complex dynamics generated at the patch level, the region wide pattern of disturbance is the key factor influencing host suppression. The level of host suppression achieved can be understood in terms of both the strength of density dependent parasitism, and the relative amounts of host and parasitoid mixing amongst patches of different ages. Local dispersal among patches is sufficient to ensure coexistence of the host and parasitoid, though persistence is not necessarily associated with the formation of self-organized spatial structures reported in previous studies. Finally, a stochastic version of the model is developed, in order to highlight how the effects of demographic stochasticity may influence biocontrol success in highly disturbed agricultural systems.     

Parsing propagule pressure: Number,not size,of introductions drives colonization success in a novel environment

Predicting whether individuals will colonize a novel habitat is of fundamental ecological interest and is crucial to conservation efforts. A consistently supported predictor of colonization success is the number of individuals introduced, also called propagule pressure. Propagule pressure increases with the number of introductions and the number of individuals per introduction (the size of the introduction), but it is unresolved which process is a stronger driver of colonization success. Furthermore, their relative importance may depend upon the environment, with multiple introductions potentially enhancing colonization of fluctuating environments. To evaluate the relative importance of the number and size of introductions and its dependence upon environmental variability, we paired demographic simulations with a microcosm experiment. Using Tribolium flour beetles as a model system, we introduced a fixed number of individuals into replicated novel habitats of stable or fluctuating quality, varying the number of introductions through time and size of each introduction. We evaluated establishment probability and the size of extant populations through seven generations. We found that establishment probability generally increased with more, smaller introductions, but was not affected by biologically realistic fluctuations in environmental quality. Population size was not significantly affected by environmental variability in the simulations, but populations in the microcosms grew larger in a stable environment, especially with more introduction events. In general, the microcosm experiment yielded higher establishment probability and larger populations than the demographic simulations. We suggest that genetic mechanisms likely underlie these differences and thus deserve more attention in efforts to parse propagule pressure. Our results highlight the importance of preventing further introductions of undesirable species to invaded sites and suggest conservation efforts should focus on increasing the number of introductions or reintroductions of desirable species rather than increasing the size of those introduction events into harsh environments.