The importance of disturbance by fire and other abiotic and biotic factors in driving cheatgrass invasion varies based on invasion stage

Disturbances create fluctuations in resource availability that alter abiotic and biotic constraints. Exotic invader response may be due to multiple factors related to disturbance regimes and complex interactions between other small- and large-scale abiotic and biotic processes that may vary across invasion stages. We explore how cheatgrass responds to both frequency and season of prescribed burning for a 10-year period in ponderosa pine forested stands. To understand interactions of fire disturbance, other abiotic factors, biotic resistance, and propagule pressure, we use long-term data from different spatial scales representing different invasion stages (local establishment or spread and broader scale extent/impact) to model cheatgrass dynamics. We found that after 10 years, cheatgrass cover increased with fall burning regardless of burn frequency (1 burn vs. 3 burns). There was no evidence that cheatgrass invasion is decreasing through time even in areas burned only once. Factors important for explaining local fine-scale cheatgrass establishment and spread, and broader scale extent/impact varied. The spatial extent of the first burns facilitated fine-scale cheatgrass establishment while bare soil cover constrained establishment. Biotic resistance, in the form of native annual forb cover, constrained fine-scale cheatgrass spread. Initial cheatgrass abundance in 2002, a factor related to propagule pressure, was key for explaining the broader scale extent/impact of cheatgrass by 2012. Biotic resistance, in the form of native perennial grass cover, constrained extent/impact but only when initial cheatgrass abundance was low. Our findings regarding factors affecting invasion dynamics may be useful to consider for future restoration and conservation efforts in burned ponderosa pine forests.     

Manipulating propagule pressure to test the invasibility of subtidal marine habitats

Global patterns show that estuaries are more invaded than open coasts and artificial habitats are more invaded than natural ones. The contention that artificial habitats in estuaries are more invasible than other habitats may result from variation in propagule supply, however, as artificial habitats are closely linked to vectors of non-native propagules, such as ships and boats. True comparisons of habitat invasibility require manipulations of propagule pressure, which has been historically difficult in marine systems. Using in-situ larval dosing, we delivered propagules of the widespread invasive ascidian Botrylloides violaceus into field mesocosms and assessed how habitat type (floating dock vs. benthic rock), resource availability (occupied vs. unoccupied plates), and propagule number (5, 25 and 50 larvae 225 cm^?2) affected settlement (survival after 24 h) and recruitment (survival after 56 days) success. In-situ larval dosing was successful, and after eight weeks there were significant differences in recruitment due to initial dose-size, habitat type, and space availability. At the habitat scale, despite equal propagule delivery, PVC plates in natural benthic rock were not equally invasible and few propagules survived to recruitment. At the organism scale, increased habitat complexity, through facilitation by established fouling species, rather than freedom from space competitors, appears to be more important for B. violaceus to establish. Our results offer greater mechanistic understanding of broader invasion patterns between artificial and natural habitats. This work extends the possibilities for further research to counteract the confounding issue of unknown and unknowable propagule delivery when attempting to explain variation in invasion success.     

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.     

Propagule pressure governs establishment of an invasive herb

The success of plant invasions may be limited by the availability of propagules and/or of suitable microsites, with microsite availability being affected by, for example, disturbance and interspecific competition. A mechanistic understanding of the contributions of propagule pressure and microsite limitation to plant invasions is therefore required to minimise future invasions. Here, we investigated the relative roles of propagule pressure, the availability of microsites, and their interaction on the establishment of an invasive herb, Lupinus polyphyllus, in two geographic regions representing different climate and growth conditions in Finland (a more productive southern region and a harsher central region). We carried out a field experiment in 14 L. polyphyllus populations, in which we manipulated both propagule pressure and disturbance. In a complementary greenhouse experiment, we manipulated propagule pressure and interspecific competition. Seedling establishment of L. polyphyllus was higher in the more productive southern region than in the harsher central region. The number of L. polyphyllus seedlings increased with increasing propagule pressure regardless of disturbance or interspecific competition. However, the number of L. polyphyllus seedlings per sown seed (relative establishment) tended to decrease with increasing propagule pressure, indicating that the positive effect of propagule pressure on early invasion is partially counteracted by density-dependent mortality at high seed densities. Our results highlight the dominant role of propagule pressure over disturbance and interspecific competition in the establishment of L. polyphyllus, suggesting that the early stage of invasion is limited by the availability of propagules rather than the availability of suitable microsites.     

Janzen–Connell effects partially supported in reef-building corals: adult presence interacts with settler density to limit establishment

In many plant and sessile marine invertebrate (SMI) taxa, population and community dynamics are heavily influenced by processes occurring during the dispersal and establishment phases. The Janzen–Connell (J–C) hypothesis predicts increased survival of early life stages with decreasing conspecific density and increased distance from conspecific adults. Evidence of J–C effects in maintaining diversity is common in plant communities, but its importance in SMI communities remains unclear. Under controlled aquarium conditions, we examined the effect of density-dependence and adult conspecific water treatments (absent/present) on propagule settlement success and settler post-settlement survival, along with associated spatial patterns, for six broadcast-spawning, reef-building coral species from three families. We also tested if settlement success was linked to increasing propagule species diversity for three coral species from two families. We found that the probability of settlement was density independent and not influenced by adult present water treatments. Yet, adult present water treatments and settler density did have a synergistic negative effect on the probability of short-term settler survival for all species examined. Settlers also showed greater spatial aggregation as their numbers increased, but were less aggregated in adult present water treatments compared to those in adult absent water treatments. We further show evidence of significant species interactions among propagules, as settlement in single-species trials was four-fold higher compared to mixed-species trials. Our findings from controlled experimental arenas indicate that the early establishment of corals was predominantly limited by density-dependent settler–adult interactions among conspecifics and propagule–propagule interactions among heterospecifics. Thus, the proximity to established conspecific adults, settler density and species diversity of propagules are relevant drivers of local coral community diversity and structure. Based on these outcomes, we suggest that the J–C hypothesis, with demonstrated importance for plants, is partially upheld for reef corals.     

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.     

Disturbance affects short-term facilitation, but not long-term saturation, of exotic plant invasion in New Zealand forest

We investigate the spread of an exotic herb, Hieracium lepidulum, into a New Zealand Nothofagus forest with the aim of understanding how stand-development of tree populations, propagule pressure and invader persistence, affect invasion across the landscape and within communities. Using data repeatedly collected over 35 years, from 250 locations, we parametrize continuous-time Markov chain models and use these models to examine future projections of the invasion under a range of hypothetical scenarios. We found that the probability of invasion into a stand was relatively high following canopy disturbance and that local abundance of Hieracium was promoted by minor disturbances. However, model predictions extrapolated 45 years into the future show that neither the rate of landscape-level invasion, nor local population growth of Hieracium, was affected much by changing the frequency of canopy disturbance events. Instead, invasion levels were strongly affected by the ability of Hieracium to persist in the understorey following forest canopy closure, and by propagule supply from streams, forest edges and plants already established within the stand. Our results show that disturbance frequency has surprisingly little influence on the long-term trajectory of invasion, while invader persistence strongly determines invasion patterns.     

Invasion resistance and persistence: established plants win,even with disturbance and high propagule pressure

Disturbances and propagule pressure are key mechanisms in plant community resistance to invasion, as well as persistence of invasions. Few studies, however, have experimentally tested the interaction of these two mechanisms. We initiated a study in a southwestern ponderosa pine (Pinus ponderosa Laws.)/bunch grass system to determine the susceptibility of remnant native plant communities to cheatgrass (Bromus tectorum L.) invasion, and persistence of cheatgrass in invaded areas. We used a 2 × 2 factorial design consisting of two levels of aboveground biomass removal and two levels of reciprocal seeding. We seeded cheatgrass seeds in native plots and a native seed mixture in cheatgrass plots. Two biomass removal disturbances and sowing seeds over 3 years did not reverse cheatgrass dominance in invaded plots or native grass dominance in non-invaded native plots. Our results suggest that two factors dictated the persistence of the resident communities. First, bottlebrush squirreltail (Elymus elymoides (Raf.) Swezey) was the dominant native herbaceous species on the study site. This species is typically a poor competitor with cheatgrass as a seedling, but is a strong competitor when mature. Second, differences in pretreatment levels of plant-available soil nitrogen and phosphorus may have favored the dominant species in each community. Annual species typically require higher levels of plant-available soil nutrients than perennial plants. This trend was observed in the annual cheatgrass community and perennial native community. Our study shows that established plants and soil properties can buffer the influences of disturbance and elevated propagule pressure on cheatgrass invasion.     

Fire does not facilitate invasion by alien annual grasses in an infertile Australian agricultural landscape

Plant invasions are a significant threat to fragmented native plant communities in many agricultural regions. Fire potentially facilitates invasions, but in landscapes historically subject to recurrent fires, exclusion of fire is also likely to result in loss of biodiversity. We investigated the relationship between fire, fragmentation and alien plant invasion in mallee communities of the Western Australian wheatbelt. We hypothesized that invasion is limited by lack of propagules and the low soil nutrient levels of this old, infertile landscape, but that fire and/or fragmentation disrupt these limits. We tested the effects of three factors on establishment and abundance of alien annuals: ± fire, ± post-fire seeding with the locally invasive Avena barbata (propagule availability) and three landscape contexts. The three landscape contexts, exploring site limitations, were reserve interiors, perimeter edges adjacent to agricultural land and internal reserve roadside edges. Our first hypothesis was supported: Avena establishment was consistently greater in seeded plots, but away from perimeter edges, growth was poor. Our second hypothesis was supported only for perimeter edges: neither fire nor fragmentation by interior roads enhanced invasive plant establishment or biomass. At perimeter edges, invasive plant biomass was significantly greater. This was associated with higher propagule availability and elevated soil nutrient levels but was not enhanced by fire. We conclude that fire is unlikely to promote invasion by alien annuals in low-nutrient ecosystems such as mallee, hence is a viable disturbance strategy for biodiversity conservation away from nutrient-enriched edges.     

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.     

Relative Roles of Disturbance and Propagule Pressure on the Invasion of Humid Tropical Forest by Cordia alliodora (Boraginaceae) in Tanzania

Current understanding of the vulnerability of tropical forests to plant invasion is limited but is widely believed to increase where forests: (1) suffer marked natural or man-made disturbance; and/or (2) are exposed to high propagule pressure of alien species. This study aimed, for the first time, to address the importance of propagule pressure and disturbance by examining the spread of an introduced tree, Cordia alliodora , from a single plantation into a surrounding mosaic of humid forest in the East Usambara Mountains, Tanzania. By assessing vulnerability to invasion along transects radiating from the plantation, the effects of distance (measure of propagule pressure), and disturbance could be discerned. For all life stages, distance from source population was the strongest correlate of density. A marked influence of disturbance was only evident for C. alliodora seedlings. Spatial variation in the densities of later life stages may be a function of past disturbances, less easy to assess from current surveys, especially following the marked self-thinning between seedling and adult densities. Nevertheless, the evidence suggests that propagule pressure is a more important determinant of Cordia density than disturbance. If this is true for other alien tree species in tropical forests, controlling for introduction effort is essential to assess the drivers of plant invasion. Given an annual population growth rate of ca 3.5 percent, equivalent to the population doubling every 20 yr, C. alliodora poses a significant threat to the East Usambaras as well as other humid forests where it is promoted for agroforestry.     

Antagonistic effects of large- and small-scale disturbances on exotic tree invasion in a native tussock grassland relict

It is generally accepted that disturbances increase community invasibility. Yet the role of disturbance in plant invasions may be less predictable than often assumed, due to the influence of environmental stochasticity and interactions between disturbance regimes. We evaluated the single and interactive effects of prescribed burning (large-scale, infrequent event) and animal diggings (small-scale, frequent events) on the invasion success of Gleditsia triacanthos L. in a tussock grassland relict of the Inland Pampa, Argentina. Tree seedling emergence and survival were monitored over 4 years, after adjusting for propagule pressure through copious seed addition to all disturbance treatments. Burning altered community structure by suppressing tussock grasses and promoting exotic forbs, whereas simulated, armadillo-like diggings had little impact on herbaceous composition. Overall, seedling emergence rather than survival represented the main demographic bottleneck for tree invasion. Tree establishment success varied among seedling cohorts emerged in different climatic years. In a dry year, emergence was only slightly affected by disturbances. In contrast, for two consecutive wet years, initial burning and armadillo-like diggings exerted strong, antagonistic effects on tree recruitment. Whereas fire alone increased recruitment, the simulated burrowing regime prevented seedling emergence in both burned and unburned plots. The latter effect might be explained by reduced soil moisture, and increased seed burial and predation in excavated patches. Thus, the impact of a single, large-scale perturbation promoting woody plant invasion was overridden by a regime of small-scale, frequent disturbances. Our results show that grassland invasibility was contingent on inter-annual climatic variation as well as unexpected interactions between natural and anthropogenic disturbance agents.     

Inferring process from pattern in plant invasions: a semimechanistic model incorporating propagule pressure and environmental factors

Propagule pressure is intuitively a key factor in biological invasions: increased availability of propagules increases the chances of establishment, persistence, naturalization, and invasion. The role of propagule pressure relative to disturbance and various environmental factors is, however, difficult to quantify. We explored the relative importance of factors driving invasions using detailed data on the distribution and percentage cover of alien tree species on South Africa's Agulhas Plain (2,160 km2). Classification trees based on geology, climate, land use, and topography adequately explained distribution but not abundance (canopy cover) of three widespread invasive species (Acacia cyclops, Acacia saligna, and Pinus pinaster). A semimechanistic model was then developed to quantify the roles of propagule pressure and environmental heterogeneity in structuring invasion patterns. The intensity of propagule pressure (approximated by the distance from putative invasion foci) was a much better predictor of canopy cover than any environmental factor that was considered. The influence of environmental factors was then assessed on the residuals of the first model to determine how propagule pressure interacts with environmental factors. The mediating effect of environmental factors was species specific. Models combining propagule pressure and environmental factors successfully predicted more than 70% of the variation in canopy cover for each species.     

Propagule Pressure: A Null Model for Biological Invasions

Invasion ecology has been criticised for its lack of general principles. To explore this criticism, we conducted a meta-analysis that examined characteristics of invasiveness (i.e. the ability of species to establish in, spread to, or become abundant in novel communities) and invasibility (i.e. the susceptibility of habitats to the establishment or proliferation of invaders). There were few consistencies among invasiveness characteristics (3 of 13): established and abundant invaders generally occupy similar habitats as native species, while abundant species tend to be less affected by enemies; germination success and reproductive output were significantly positively associated with invasiveness when results from both stages (establishment/spread and abundance/impact) were combined. Two of six invasibility characteristics were also significant: communities experiencing more disturbance and with higher resource availability sustained greater establishment and proliferation of invaders. We also found that even though ‘propagule pressure’ was considered in only ~29% of studies, it was a significant predictor of both invasiveness and invasibility (55 of 64 total cases). Given that nonindigenous species are likely introduced non-randomly, we contend that ‘propagule biases’ may confound current paradigms in invasion ecology. Examples of patterns that could be confounded by propagule biases include characteristics of good invaders and susceptible habitats, release from enemies, evolution of ‘invasiveness’, and invasional meltdown. We conclude that propagule pressure should serve as the basis of a null model for studies of biological invasions when inferring process from patterns of invasion. An erratum to this article can be found at     

群落可侵入性及其影响因素

 可侵入性用于评价群落易遭受生物入侵的程度,受外来种死亡率、区域气候、干扰水平、生态系统抵抗入侵的能力、本地种竞争和抗干扰能力等因素的影响。当前对群落或区域间可侵入性的比较常以外来种数量或丰度为据,然而这两者仅代表了群落内单一的动态过程,不足以作为衡量群落可侵入性的广泛标准。借助一个描述外来种数量的简单模型阐明,由于影响可侵入性因素的复杂性,各地之间的可侵入性几乎不可比较。并从入侵过程、入侵种特性及本地种、本地群落对入侵的抵抗性几方面对群落可侵入性进行了阐述分析,其中着重介绍入侵生态中几个重要的概念,如可侵     

Snowpack,fire, and forest disturbance: interactions affect montane invasions by non‐native shrubs

Montane regions worldwide have experienced relatively low plant invasion rates, a trend attributed to increased climatic severity, low rates of disturbance, and reduced propagule pressure relative to lowlands. Manipulative experiments at elevations above the invasive range of non‐native species can clarify the relative contributions of these mechanisms to montane invasion resistance, yet such experiments are rare. Furthermore, global climate change and land use changes are expected to cause decreases in snowpack and increases in disturbance by fire and forest thinning in montane forests. We examined the importance of these factors in limiting montane invasions using a field transplant experiment above the invasive range of two non‐native lowland shrubs, Scotch broom (Cytisus scoparius) and Spanish broom (Spartium junceum), in the rain–snow transition zone of the Sierra Nevada of California. We tested the effects of canopy closure, prescribed fire, and winter snow depth on demographic transitions of each species. Establishment of both species was most likely at intermediate levels of canopy disturbance, but at this intermediate canopy level, snow depth had negative effects on winter survival of seedlings. We used matrix population models to show that an 86% reduction in winter snowfall would cause a 2.8‐fold increase in population growth rates in Scotch broom and a 3.5‐fold increase in Spanish broom. Fall prescribed fire increased germination rates, but decreased overall population growth rates by reducing plant survival. However, at longer fire return intervals, population recovery between fires is likely to keep growth rates high, especially under low snowpack conditions. Many treatment combinations had positive growth rates despite being above the current invasive range, indicating that propagule pressure, disturbance, and climate can all strongly affect plant invasions in montane regions. We conclude that projected reductions in winter snowpack and increases in forest disturbance are likely to increase the risk of invasion from lower elevations.     

Traits linked with species invasiveness and community invasibility vary with time,stage and indicator of invasion in a long‐term grassland experiment

Much uncertainty remains about traits linked with successful invasion – the establishment and spread of non‐resident species into existing communities. Using a 20‐year experiment, where 50 non‐resident (but mostly native) grassland plant species were sown into savannah plots, we ask how traits linked with invasion depend on invasion stage (establishment, spread), indicator of invasion success (occupancy, relative abundance), time, environmental conditions, propagule rain, and traits of invaders and invaded communities. Trait data for 164 taxa showed that invader occupancy was primarily associated with traits of invaders, traits of recipient communities, and invader‐community interactions. Invader abundance was more strongly associated with community traits (e.g. proportion legume) and trait differences between invaders and the most similar resident species. Annuals and invaders with high‐specific leaf area were only successful early in stand development, whereas invaders with conservative carbon capture strategies persisted long‐term. Our results indicate that invasion is context‐dependent and long‐term experiments are required to comprehensively understand invasions.     

Plant invasions along mountain roads: the altitudinal amplitude of alien Asteraceae forbs in their native and introduced ranges

Studying plant invasions along environmental gradients is a promising approach to dissect the relative importance of multiple interacting factors that affect the spread of a species in a new range. Along altitudinal gradients, factors such as propagule pressure, climatic conditions and biotic interactions change simultaneously across rather small geographic scales. Here we investigate the distribution of eight Asteraceae forbs along mountain roads in both their native and introduced ranges in the Valais (southern Swiss Alps) and the Wallowa Mountains (northeastern Oregon, USA). We hypothesised that a lack of adaptation and more limiting propagule pressure at higher altitudes in the new range restricts the altitudinal distribution of aliens relative to the native range. However, all but one of the species reached the same or even a higher altitude in the new range. Thus neither the need to adapt to changing climatic conditions nor lower propagule pressure at higher altitudes appears to have prevented the altitudinal spread of introduced populations. We found clear differences between regions in the relative occurrence of alien species in ruderal sites compared to roadsides, and in the degree of invasion away from the roadside, presumably reflecting differences in disturbance patterns between regions. Whilst the upper altitudinal limits of these plant invasions are apparently climatically constrained, factors such as anthropogenic disturbance and competition with native vegetation appear to have greater influence than changing climatic conditions on the distribution of these alien species along altitudinal gradients.     

Dealing with scarce data to understand how environmental gradients and propagule pressure shape fine‐scale alien distribution patterns on coastal dunes

Questions: On sandy coastal habitats, factors related to substrate and to wind action vary along the sea–inland ecotone, forming a marked directional disturbance and stress gradient. Further, input of propagules of alien plant species associated to touristic exploitation and development is intense. This has contributed to establishment and spread of aliens in coastal systems. Records of alien species in databases of such heterogeneous landscapes remain scarce, posing a challenge for statistical modelling. We address this issue and attempt to shed light on the role of environmental stress/disturbance gradients and propagule pressure on invasibility of plant communities in these typical model systems. Location: Sandy coasts of Lazio (Central Italy). Methods: We proposed an innovative methodology to deal with low prevalence of alien occurrence in a data set and high cost of field‐based sampling by taking advantage, through predictive modelling, of the strong interrelation between vegetation and abiotic features in coastal dunes. We fitted generalized additive models to analyse (1) overall patterns of alien occurrence and spread and (2) specific patterns of the most common alien species recorded. Conclusion: Even in the presence of strong propagule pressure, variation in local abiotic conditions can explain differences in invasibility within a local environment, and intermediate levels of natural disturbance and stress offer the best conditions for spread of alien species. However, in our model system, propagule pressure is actually the main determinant of alien species occurrence and spread. We demonstrated that extending the information of environmental features measured in a subsample of vegetation plots through predictive modelling allows complex questions in invasion biology to be addressed without requiring disproportionate funding and sampling effort.     

Spatial and temporal variability in propagule limitation of California native grasses

Community composition and diversity arise from limitations in propagule supply (i.e. propagule or seed limitation) and propagule establishment after arriving at a site (i.e. establishment or microsite limitation). Recent meta‐analyses suggest that the degree of propagule limitation depends on local abiotic and biotic conditions, which in turn are likely to vary spatially and temporally. Nevertheless, seed addition studies testing propagule limitation are rarely replicated in multiple locations and years and often lack experimental manipulations of critical determinants of propagule limitation, such as the density and species richness of the propagule pool and disturbance and resource supply of recipient site. The invasion of California (USA) grasslands by exotic annual species from the Mediterranean region is unique in its scope (over 9 million ha) and persistence (more than 150 years). This invasion provides an exciting context in which to test the role of spatial and temporal variability in mediating propagule limitation and ultimately the potential for restoration. Here I present the results of native‐grass, seed‐addition trials conducted along a 500 km gradient in California in three consecutive years spanning a wide range of environmental conditions and initial conditions (e.g. rainfall, seeding density and disturbance). While native grasses were able to establish at many locations, per capita seed survival was low suggesting that the fate of these species is governed by interplay between propagule and establishment limitation. Establishment primarily varied spatially, with the most successful establishment occurring at fertile locations with low resident species richness. While recruitment was highly variable among years initially, there was no difference among seedling trials after three years.