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.     

Relationships of the Galapagos flora

Joseph Dalton Hooker's pioneer 1847 paper on Galapagos plants and their relationships is a classic in the field of phytogeography. It was the first study of its kind to be published, comparing the islands' flora with island and continental floras elsewhere, hypothesizing on the dispersal mechanisms of the plants, and pointing out anomalies in the inter-island distributions of the native species. These are still three of the primary concerns of contemporary phytogeographers, and the present paper contrasts Hooker's findings with those of today. Despite the accumulation of a large amount of data since his time, many of Hooker's conclusions regarding Galapagos phytogeography remain valid.     

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.     

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.     

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 role of propagule pressure in explaining species invasions

Human-mediated species invasions are a significant component of current global environmental change. There is every indication that the rate at which locations are accumulating non-native species is accelerating as free trade and globalization advance. Thus, the need to incorporate predictive models in the assessment of invasion risk has become acute. However, finding elements of the invasion process that provide consistent explanatory power has proved elusive. Here, we propose propagule pressure as a key element to understanding why some introduced populations fail to establish whereas others succeed. In the process, we illustrate how the study of propagule pressure can provide an opportunity to tie together disparate research agendas within invasion ecology.     

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

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

An updated checklist of the vascular flora alien to Italy

An updated inventory of the vascular flora alien to Italy, providing details on the occurrence at regional level, is presented. The checklist includes 1597 species, subspecies, and hybrids, distributed in 725 genera and 152 families; 2 taxa are lycophytes, 11 ferns and fern allies, 33 gymnosperms, and 1551 angiosperms. 157 taxa are archaeophytes and 1440 neophytes. The alien taxa currently established in Italy are 791 (570 naturalized and 221 invasive), while 705 taxa are casual aliens, 4 are not assessed, 7 are of unknown regional distribution, 47 have not been confirmed in recent times, 3 are considered extinct or possibly extinct in the country, and 40 are doubtfully occurring in Italy. This checklist allows to establish an up-to-date number (9792) of taxa constituting the whole (native and alien) Italian flora.     

长春地区入侵植物种类组成及区系分析

为了解长春地区入侵植物分布规律及其影响因子,对长春地区入侵植物种类组成和植物区系进行了研究。结果表明:长春地区共有外来入侵植物85种,隶属于26科64属,其中4种为有害检疫杂草;菊科、苋科、禾本科、豆科和茄科构成了长春市入侵植物的主体,其中以菊科(18种)为最多,占总种数的21.18%;草本植物共81种,占95.29%,说明草本植物比木本植物对长春地区更具有入侵性;长春地区入侵植物区系成分复杂,入侵植物科的区系类型主要为世界分布;入侵属的分布区系类型是世界分布占优势,其次为北温带和泛热带分布。根据入侵现状,本文对长春地区防范外来植物入侵提出了建议。     

Residence time and exposure time of sinking phytoplankton in the euphotic layer

The residence time of a sinking particle in the euphotic layer is usually defined as the time taken by this particle to reach for the first time the bottom of the euphotic layer. According to this definition, the concept of residence time does not take into account the fact that many cells leaving the euphotic layer at some time can re-enter the euphotic layer at a later time. Therefore, the exposure time in the surface layer, i.e. the total time spent by the particles in the euphotic layer irrespective of their possible excursions outside the surface layer, is a more relevant concept to diagnose the effect of diffusion on the survival of phytoplankton cells sinking through the water column.While increasing the diffusion coefficient can induce both a decrease or an increase of the residence time, the exposure time in the euphotic layer increases monotonically with the diffusion coefficient, at least when the settling velocity does not increase with depth. Turbulence is therefore shown to increase the total time spent by phytoplankton cells in the euphotic layer.The generalization of the concept of exposure time to take into account the variations of the light intensity with depth or the functional response of phytoplankton cells to irradiance leads to the definition of the concepts of light exposure and effective light exposure. The former provides a measure of the total light energy received by the cells during their cycling through the water column while the latter diagnose the potential growth rate.The exposure time, the light exposure and the effective light exposure can all be computed as the solution of a differential problem that generalizes the adjoint approach introduced by Delhez et al. (2004) for the residence time. A general analytical solution of the 1D steady-state version of this equation is derived from which the properties of the different diagnostic tools can be obtained.     

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.     

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.     

The alien flora of Europe: a taxonomic and biogeographic review

Abstract. A geographic and taxonomic overview of the non-indigenous plant species of Europe, based on the ‘Flora Europaea’ is given. The flora of Europe includes 1568 species which have either expanded their ranges within Europe under human influence (naturalized European species) or are of non-European origin (exotic species). The latter group consists of 580 species (37%) which form a diverse group in terms of their taxonomic composition and geographic origin. The exotics are represented by 113 families, of which the Compositae, Rosaceae and Gramineae are most important. The ratio of species to families is low. Most exotic species in Europe originate from the Americas and Asia. Countries of southern Europe have a higher relative number of exotics in their non-native flora than northern ones. The species-range size distribution differs between naturalized European and exotic species; the latter are on average more widespread than the naturalized.     

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.     

Functional diversity changes in native and alien urban flora over three centuries

Biological Invasions - Alien species in urban areas have a large effect on overall species diversity. A suitable metric of flora’s response to environmental change is functional diversity...     

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.     

Whale Shark (Rhincodon typus) Seasonal Presence,Residence Time and Habitat Use at Darwin Island,Galapagos Marine Reserve

The life history of the whale shark (Rhincodon typus), including its reproductive ecology, still remains largely unknown. Here, we present results from the first whale shark population study around Darwin Island, Galapagos Marine Reserve. Following a diversified approach we characterized seasonal occurrence, population structure and size, and described habitat use of whale sharks based on fine scale movements around the island. Whale shark presence at Darwin Island was negatively correlated with Sea Surface Temperature (SST), with highest abundance corresponding to a cool season between July and December over six years of monitoring. From 2011 to 2013 we photo-identified 82 whale sharks ranging from 4 to 13.1 m Total Length (TL). Size distribution was bimodal, with a great majority (91.5%) of adult female individuals averaging 11.35 m±0.12 m (TL±SE), all but one showing signs of a potential pregnancy. Population dynamics models for apparently pregnant sharks estimated the presence of 3.76±0.90 (mean ± SE) sharks in the study area per day with an individual residence time of 2.09±0.51 (mean ± SE) days. Movement patterns analysis of four apparently pregnant individuals tracked with acoustic tags at Darwin Island revealed an intense use of Darwin''s Arch, where no feeding or specific behavior has been recorded, together with periodic excursions around the island''s vicinity. Sharks showed a preference for intermediate depths (20–30 m) with occasional dives mostly to mid-water, remaining the majority of their time at water temperatures between 24–25°C. All of our results point to Darwin Island as an important stopover in a migration, possibly with reproductive purposes, rather than an aggregation site. Current studies carried out in this area to investigate regional scale movement patterns may provide essential information about possible pupping grounds for this enigmatic species.