Establishment Of Non-native Species Research Articles

Nutrients influence the multi-trophic impacts of an invasive species unaffected by native competitors or predators

Non-native species often lead to undesirable ecological and environmental impacts. Two hypotheses that predict establishment of non-native species are enemy release and biotic resistance. Support for these hypotheses in freshwater invasions is mixed. Experiments combined with field observations provide a complementary approach to understanding how interactions between native and non-native species lead to enemy release or biotic resistance. We tested experimentally whether these hypotheses provided insights into the invasion of the banded mystery snail (Viviparus georgianus), which has invaded the Great Lakes region and northeastern Unites States (US) from the southeastern US. Because freshwater systems vary widely in their nutrient concentrations due to natural and anthropogenic processes, we tested whether nutrient additions altered competitive and predatory interactions that regulate mechanisms of enemy release or biotic resistance. We evaluated the status of the mystery snail invasion in a 3-year field survey of Lake George (NY, US) to identify if field observations supported any experimental conclusions. The presence of the banded mystery snail led to a 14% and 27% reduction in biomass of a native competitor under low- and high-nutrient concentrations, respectively. The mystery snail also triggered a 29% biomass loss of a native snail predator, but only in low-nutrient concentrations. Field surveys indicated that the mystery snail dominated the snail community; of seven snail species, it comprised 77% of all snails. Results from the field surveys combined with experimental results indicate that neither competitors nor predators have likely suppressed the invasion of the banded mystery snail. This conclusion is consistent with competitive- and predatory-enemy release as we found no indication of biotic resistance via competition or predation from native species. Our results further highlight that the post-establishment impacts of invasive species are altered by the trophic state of freshwater ecosystems.

Exotic species dominate marinas between the two most populated regions in the southwestern Atlantic Ocean

Human occupation of coastal areas promotes the establishment of non-native species but information on bioinvasions is usually biased toward the Northern Hemisphere. We assessed non-native species' importance in sessile communities at six marinas along the most urbanized area of the Southwestern Atlantic coastline. We found 67 species, of which 19 are exotic. The most frequent species was the exotic polychaete Branchiomma luctuosum, while the most abundant was the exotic bryozoan Schizoporella errata that monopolized the substrata in three marinas. Along with S. errata, the exotic polychaete Hydroides elegans and ascidian Styela plicata dominated space in the three remaining marinas, while native species were in general rare. We show that communities associated with artificial substrata along this Brazilian urbanized area are dominated by exotic species and that using abundance data along with species identity can improve our understanding of the importance of exotic species for the dynamics of biological communities.

Effects of mastication of burned non-commercial Pinus pinaster Ait. trees on soil compaction and vegetation response

Mastication is commonly used as a fuel reduction treatment to minimize severe wildfires in fire-prone areas worldwide. Although mastication of non-commercial burned trees has become common practice in NW Spain in recent years, little is known about the possible effects on fuels, soil physical properties and vegetation recovery. In this study, sites in NW Spain (Arbo and Soutomaior) where non-commercial Pinus pinaster Ait. stands burned in August 2016 in NW Spain were selected for study. In both sites the fires scorched tree crowns and caused moderate soil burn severity. In January 2017, burned trees in 15 plots (10 in Arbo and 5 in Soutomaior) were masticated using a steel track tractor with a front mounted rotating toothed drum. In addition, five plots in Arbo were masticated using a walking excavator with a mulching head. In the 10 remaining plots (5 in each site) the trees were cut with a chainsaw and manually moved offsite. The fuel distribution categories were determined immediately after mastication. Soil penetration resistance and soil shear strength were measured immediately after wildfire, and also immediately, six months and one year after mastication. Species cover and pine seedling density were measured six months and one year after mastication. Percentage masticated material cover ranged from a mean of 60% (s.e. = 1.5) after mastication by tractor in Soutomaior to a mean of 81% (s.e. = 6.0) after mastication by walking excavator in Arbo. 64–75% of material was shredded after mastication. Mastication had no effect on soil shear strength or soil penetration resistance, and it also did not affect total vegetation cover. Species cover was not significantly affected by any of the environmental variables or masticated material cover. Mean pine seedling density was 0.2 seedlings m−2 in masticated and non-masticated plots, irrespective of the equipment used. Mastication did not favour the establishment of non-native species. Mastication after wildfire does not appear to be detrimental to soil conservation and vegetation recovery although further research is needed to confirm this.

Implications of non-native species for mutualistic network resistance and resilience.

Resilience theory aims to understand and predict ecosystem state changes resulting from disturbances. Non-native species are ubiquitous in ecological communities and integrated into many described ecological interaction networks, including mutualisms. By altering the fitness landscape and rewiring species interactions, such network invasion may carry important implications for ecosystem resistance and resilience under continued environmental change. Here, I hypothesize that the tendency of established non-native species to be generalists may make them more likely than natives to occupy central network roles and may link them to the resistance and resilience of the overall network. I use a quantitative research synthesis of 58 empirical pollination and seed dispersal networks, along with extinction simulations, to examine the roles of known non-natives in networks. I show that non-native species in networks enhance network redundancy and may thereby bolster the ecological resistance or functional persistence of ecosystems in the face of disturbance. At the same time, non-natives are unlikely to partner with specialist natives, thus failing to support the resilience of native species assemblages. Non-natives significantly exceed natives in network centrality, normalized degree, and Pollination Service Index. Networks containing non-natives exhibit lower connectance, more links on average, and higher generality and vulnerability than networks lacking non-natives. As environmental change progresses, specialists are particularly likely to be impacted, reducing species diversity in many communities and network types. This work implies that functional diversity may be retained but taxonomic diversity decline as non-native species become established in networks worldwide.

Low litter cover, high light availability and rock cover favour the establishment of Ailanthus altissima in forests in southern Switzerland

Future forest composition is uncertain in many areas due to climate change. The spread of non-native species adds to these uncertainties, particularly in forests recently colonised by novel tree species. To anticipate future forest composition, and thus the provision of ecosystem services, a thorough understanding of the factors influencing the establishment of non-native tree species is essential. We studied the presence and abundance of regeneration of Ailanthusaltissima (Mill.) Swingle in 89 plots on a regular grid in three sites in southern Switzerland to determine the main drivers of its establishment. All sites are located in abandoned, i.e. currently unmanaged stands of Castaneasativa Mill. Propagule pressure is caused by single mature A.altissima that colonised the area ca 40 years ago. We found high rock cover, low litter cover and high light availability to be the most important predictors for the presence of A.altissima regeneration, whereas its abundance was positively influenced by high light availability, low litter cover and high browsing on regeneration of competing species. However, the presence models performed much better than the abundance models. Interestingly, the most important factors favouring the establishment of A.altissima in recently undisturbed sites were found to be similar in a nearby site after a severe forest fire, which suggests a similar establishment strategy after a disturbance as in recently undisturbed forests. Based on our results we expect a further expansion of the species in lowland forests currently dominated by C.sativa, likely controlled primarily by light availability.

Native and alien flower visitors differ in partner fidelity and network integration

Globalisation persistently fuels the establishment of non-native species outside their natural ranges. While alien plants have been intensively studied, little is known about alien flower visitors, and especially, how they integrate into natural communities. Here, we focus on mutualistic networks from five Galápagos islands to quantify whether alien and native flower visitors differ consistently in their pairwise interactions. We find that (1) alien flower visitors have more interaction partners and larger species strengths (i.e. plants are more connected to alien insects), (2) native insects tend to have higher partner fidelity as they deviate more from random partner utilisation, and iii) the difference between native and alien flower visitors in network integration intensifies with island degradation. Thus, native and alien flower visitors are not interchangeable, and alien establishment might have yet unforeseen consequences for the pairwise dynamics between plants and flower visitors on the Galápagos - especially on the heavily disturbed islands.

Phenology in a warming world: differences between native and non‐native plant species

Phenology is a harbinger of climate change, with many species advancing flowering in response to rising temperatures. However, there is tremendous variation among species in phenological response to warming, and any phenological differences between native and non-native species may influence invasion outcomes under global warming. We simulated global warming in the field and found that non-native species flowered earlier and were more phenologically plastic to temperature than natives, which did not accelerate flowering in response to warming. Non-native species' flowering also became more synchronous with other community members under warming. Earlier flowering was associated with greater geographic spread of non-native species, implicating phenology as a potential trait associated with the successful establishment of non-native species across large geographic regions. Such phenological differences in both timing and plasticity between native and non-natives are hypothesised to promote invasion success and population persistence, potentially benefiting non-native over native species under climate change.

Water temperature as a hindrance, but not limiting factor for the survival of warm water invasive crayfish introduced in cold periods

The success of non-native species establishment depends on various abiotic and biotic factors that determine the outcome of an introduction event. Limiting temperature ranges have been studied for various non-native species; however, such previous assessments of species-specific temperature thresholds may be inadequate. Because several non-native crayfish species prefer warmer water temperatures, introductions were generally assumed to occur during preferable, warmer periods. However, despite the generality, traditionally considered ‘warm-water’ species are gradually appearing in new habitats, which were previously considered too cold for successful establishment. Newly discovered overwintering abilities of these species are likely related to the winter stratification in lentic ecosystems, which maintain tolerable conditions. To understand better the survivability of two such non-native species, red swamp crayfish Procambarus clarkii and marbled crayfish Procambarus virginalis individuals were abruptly subjected to a thermic shock which lowered the water temperature from 20 °C (room temperature) to 6 °C, 4 °C and 2 °C, thus mimicking the release by pet owners during various phases of winter. The survival rate and foraging activity were monitored for up to 98 days. Procambarus clarkii showed a considerable higher survival rate at low temperatures (4 °C, 2 °C) compared to that of P. virginalis with neither sex nor size differences evident. Our findings reveal the ability of warm water invaders to withstand a shock during introduction at low temperature periods without acclimation. Considering these newly discovered shifts in physiological limitations, particularly for the red swamp crayfish, this may indicate a higher threat for areas with colder conditions.

Antarctica: The final frontier for marine biological invasions.

Antarctica is experiencing significant ecological and environmental change, which may facilitate the establishment of non‐native marine species. Non‐native marine species will interact with other anthropogenic stressors affecting Antarctic ecosystems, such as climate change (warming, ocean acidification) and pollution, with irreversible ramifications for biodiversity and ecosystem services. We review current knowledge of non‐native marine species in the Antarctic region, the physical and physiological factors that resist establishment of non‐native marine species, changes to resistance under climate change, the role of legislation in limiting marine introductions, and the effect of increasing human activity on vectors and pathways of introduction. Evidence of non‐native marine species is limited: just four marine non‐native and one cryptogenic species that were likely introduced anthropogenically have been reported freely living in Antarctic or sub‐Antarctic waters, but no established populations have been reported; an additional six species have been observed in pathways to Antarctica that are potentially at risk of becoming invasive. We present estimates of the intensity of ship activity across fishing, tourism and research sectors: there may be approximately 180 vessels and 500+ voyages in Antarctic waters annually. However, these estimates are necessarily speculative because relevant data are scarce. To facilitate well‐informed policy and management, we make recommendations for future research into the likelihood of marine biological invasions in the Antarctic region.

Modelling the establishment, spread and distribution shifts of pests

During the last century increasing human trade networks have interacted with other global changes – such as land use and climate change – to accelerate the range expansion of pest species worldwide. In this special issue, we present a series of articles that model the risk of establishment and spread of non-native harmful species, as well as shifts in the distribution of native pest species. We also provide a state-of the art review of the available literature on species distribution models for pests, weeds, and pathogens. Recent advances in modelling approaches have allowed us to significantly improve our abilities to predict changes in species distribution, and these tools have been widely used to enhance biosecurity planning and pest management. We expect that this special issue contributes toward and integration of the scientific knowledge and the development of novel approaches to model pest species distribution, as well as to better understand the factors that influence their expansion in the context of a rapidly changing world.

Ice-free area expansion compounds the non-native species threat to Antarctic terrestrial biodiversity

Warming across ice-covered regions will result in changes to both the physical and climatic environment, revealing new ice-free habitat and new climatically suitable habitats for non-native species establishment. Recent studies have independently quantified each of these aspects in Antarctica, where ice-free areas form crucial habitat for the majority of terrestrial biodiversity. Here we synthesise projections of Antarctic ice-free area expansion, recent spatial predictions of non-native species risk, and the frequency of human activities to quantify how these facets of anthropogenic change may interact now and in the future. Under a high-emissions future climate scenario, over a quarter of ice-free area and over 80% of the ~14 thousand km2 of newly uncovered ice-free area could be vulnerable to invasion by one or more of the modelled non-native species by the end of the century. Ice-free areas identified as vulnerable to non-native species establishment were significantly closer to human activity than unsuitable areas were. Furthermore, almost half of the new vulnerable ice-free area is within 20 km of a site of current human activity. The Antarctic Peninsula, where human activity is heavily concentrated, will be at particular risk. The implications of this for conservation values of Antarctica and the management efforts required to mitigate against it are in need of urgent consideration.

Assessment of Topsoil Salvage and Seed Augmentation in the Restoration of Coastal Sage Scrub on Santa Catalina Island, California

The Catalina Island Conservancy manages over 22,000 acres of coastal (California) sage scrub (CSS) habitat off the coast of Southern California, including much of the infrastructure, roads, and recreation associated with habitat disturbance. Restoration trials were initiated to establish best management practices for roadside and disturbed site revegetation under local site conditions. Site treatments compared excavated native subsoil and surface soil (±20 cm) reapplication practices with and without seed augmentation. Evaluation of native and nonnative germinants indicated that salvaged surface soil alone may not be sufficient to restore native cover. Seeding improved native cover and reduced establishment of nonnative species on unirrigated sites. Evaluation of water potential of seedlings indicated that high-frequency irrigation may favor establishment of nonnative annual grasses. Most compelling for land managers is the indication that CSS habitats may contain a significant nonnative seed bank in spite of the composition of aboveground vegetation, such that disturbance may facilitate habitat-type conversion without seed augmentation or weed control.

The role of biotic factors during plant establishment in novel communities assessed with an agent-based simulation model.

BackgroundEstablishment success of non-native species is not only influenced by environmental conditions, but also by interactions with local competitors and enemies. The magnitude of these biotic interactions is mediated by species traits that reflect competitive strength or defence mechanisms. Our aim was to investigate the importance of species traits for successful establishment of non-native species in a native community exhibiting biotic resistance in the form of competition and herbivory.MethodsWe developed a trait-based, individual-based simulation model tracking the survival of non-native plants in a native community. In the model, non-native plants are characterized by high or low values of competition and defence traits. Model scenarios included variation of initial number of non-natives, intensity of competitive interaction, density of herbivores and density as well as mixture of the native community.ResultsTraits related to competition had a much greater impact on survival of non-native species than traits related to defence. Survival rates of strong competitors never fell below 50% while survival of weak competitors averaged at about 10%. Weak competitors were also much more susceptible to competitive pressures such as community density, composition and competition intensity. Strong competitors responded negatively to changes in competition intensity, but hardly to composition or density of the native community. High initial numbers of non-native individuals decreased survival rate of strong competitors, but increased the survival rate of weak competitors. Survival under herbivore attack was only slightly higher for plants with high defensive ability than for those with low defensive ability. Surprisingly, though, herbivory increased survival of species classified as weak competitors.DiscussionHigh survival rates of strong non-native competitors relate to a higher probability of successful establishment than for weak competitors. However, the reduced survival of strong competitors at high initial numbers indicates a self-thinning effect, probably mediated by a strongly competitive milieu. For weak competitors, our model emphasizes positive effects of high propagule pressure known from field studies. General effects of herbivory or defence abilities on survival were not supported by our model. However, the positive effect of herbivory on survival of weak competitors indicated side effects of herbivory, such as weakening resident competitors. This might play an important role for establishment of non-natives in a new community.

Import volumes and biosecurity interventions shape the arrival rate of fungal pathogens.

Global trade and the movement of people accelerate biological invasions by spreading species worldwide. Biosecurity measures seek to allow trade and passenger movements while preventing incursions that could lead to the establishment of unwanted pests, pathogens, and weeds. However, few data exist to evaluate whether changes in trade volumes, passenger arrivals, and biosecurity measures have altered rates of establishment of nonnative species over time. This is particularly true for pathogens, which pose significant risks to animal and plant health and are consequently a major focus of biosecurity efforts but are difficult to detect. Here, we use a database of all known plant pathogen associations recorded in New Zealand to estimate the rate at which new fungal pathogens arrived and established on 131 economically important plant species over the last 133 years. We show that the annual arrival rate of new fungal pathogens increased from 1880 to about 1980 in parallel with increasing import trade volume but subsequently stabilised despite continued rapid growth in import trade and recent rapid increases in international passenger arrivals. Nevertheless, while pathogen arrival rates for crop and pasture species have declined in recent decades, arrival rates have increased for forestry and fruit tree species. These contrasting trends between production sectors reflect differences in biosecurity effort and suggest that targeted biosecurity can slow pathogen arrival and establishment despite increasing trade and international movement of people.

Influence of seed size on performance of non-native annual plant species in a novel community at two planting densities

Climate warming enables plant species to migrate to higher latitudes and altitudes. Within Europe, the Mediterranean harbours many species that might expand their ranges towards Western Europe. Small seed size may facilitate dispersal, however, it may impair establishment of the range-expanding plant species in the novel vegetation. In a greenhouse experiment, we examined effects of average seed size of Mediterranean plant species on their establishment in a mixed community of Western European plant species. Applying two levels of densities of the natives and a herbivory treatment, we tested how seed size is linked to response in plant growth and fitness in novel vegetation. While all non-native plant species showed a negative response to increased planting density, species with small seeds showed a less negative response. This effect persisted under herbivory. Our data suggest that small-seeded non-native plant species may tolerate competitive pressure from novel plant communities better than large-seeded species, so that small seed size may confer a higher probability of establishment of non-native species in novel communities.

Herbivory enhances the resistance of mangrove forest to cordgrass invasion

The biotic resistance hypothesis proposes that biotic interactions, such as competition and herbivory, resist the establishment and spread of non-native species. The relative and interactive role of competition and herbivory in resisting plant invasions, however, remains poorly understood. We investigated the interactive role of competition and herbivory (by the native rodent Rattus losea) in resisting Spartina alterniflora (cordgrass) invasions into mangrove forests. In southern China, although exotic cordgrass numerically dominates intertidal mudflats and open gaps in mangrove forests, intact forests appear to be highly resistant to cordgrass invasion. A field transplant and rodent exclusion experiment showed that while the impact of rodent grazing on cordgrass was weak on mangrove forest edges and open mudflats, rodent grazing strongly suppressed cordgrass in mangrove understory habitats. A greenhouse experiment confirmed a synergistic interaction between grazing and light availability (a proxy for mangrove shading and light competition) in suppressing cordgrass establishment, with the strongest impacts of grazing in low light conditions that likely weakened cordgrass to survive and resprout. When both were present, as in mangrove understory habitats, grazing and low light acted in concert to eliminate cordgrass establishment, resulting in resistance of mangrove forests to cordgrass invasion. Our results reveal that grazing by native herbivores can enhance the resistance of mangrove forests to cordgrass invasion in southern China, and suggest that investigating multifactor interactions may be critical to understanding community resistance to exotic invasions.

Propagules are not all equal: traits of vegetative fragments and disturbance regulate invasion success.

Invasion success is regulated by multiple factors. While the roles of disturbance and propagule pressure in regulating the establishment of non-native species are widely acknowledged, that of propagule morphology (a proxy for quality) is poorly known. By means of a multi-factorial field experiment, we tested how the number (5 vs. 10) and quality (intact, without fronds or without rhizoids) of fragments of the clonal invasive seaweed, Caulerpa cylindracea, influenced its ability to establish in patches of the native seagrass, Posidonia oceanica, exposed to different intensities of disturbance (0, 50, or 100% reduction in canopy cover). We hypothesized that the ability of fragments to establish would be greater for intact fragments (high quality) and reduced more by frond removal (low quality) than rhizoid removal (intermediate quality). At low propagule pressure or quality, fragment establishment was predicted to increase with increasing disturbance, whereas, at high propagule pressure or quality, it was predicted to be high regardless of disturbance intensity. Disturbance intensity, fragment number and quality had independent effects on C.cylindracea establishment success. Disturbance always facilitated fragment establishment. However, fragments retaining fronds, either intact or deprived of rhizoids, had higher establishment success than fragments deprived of fronds. Increasing propagule number had weak effects on the cover of C.cylindracea. Our results demonstrate that propagule traits enabling the acquisition of resources made available by disturbance can be more important than propagule number in determining the establishment and spread of clonal non-native plants. More generally, our study suggests that propagule quality is a key, yet underexplored, determinant of invasion success.

Spatial scales and the invasion paradox: a test using fish assemblages in a Neotropical floodplain

The invasion paradox refers to the existence of biotic resistance and biotic acceptance hypotheses. According to the biotic resistance hypothesis, the higher the richness present in a community, the more resistant it is against invaders. In contrast, the biotic acceptance hypothesis states that if the environment is suitable for a high richness of native species, it will be similarly suitable for establishment of non-native species. Previous studies on terrestrial plant communities considered the scale-dependent nature of native and non-native relationships: the relationship tends to be negative at small scales (biotic resistance) and positive at broader scales (biotic acceptance). We tested the hypothesized role of spatial scale on the relationship between non-native and native species richness using a spatially nested design and a long-term dataset (146 communities sampled during 13 years) of fish species richness from the upper Parana River floodplain, Brazil. Contrary to expectations, non-native fish species richness was positively correlated with native species richness at all spatial scales. Mobility of vertebrates, mode of invasion, and environmental disturbance may affect the role of spatial scale in potentially mediating relative importance of biotic acceptance versus resistance. In this context, the present study provides a unique contribution towards resolving the invasion paradox.

Functional distance and establishment of non-native species with complex life cycles

More than 80% of animals have complex life cycles and undergo distinct changes in ecology and morphology during development. The strength and type of factors regulating each life-stage may differ as an organism may occupy different niches during ontogeny. We examined the functional distance at larval and adult life-stages of two non-native anurans (Green Tree Frog [Hyla cinerea] and Bullfrog [Lithobates catesbeianus]) that have established in a Chihuahuan Desert anuran assemblage in Big Bend National Park. Both life stages of both non-native species occupied niche space outside of the native assemblage. At the larval stage, the ability of the tadpoles to utilize permanent aquatic habitats and coexist with predatory fishes differentiated the non-native species from the majority of the native species that are restricted to temporary pools. At the post-metamorphic life stage, each species appears to have established by exploiting unoccupied habitat and trophic niches in the recipient community. The arboreal habits of H. cinerea may enable it to utilize resources in microhabitats that are otherwise not used by native species because arboreal frogs are absent from this native assemblage. The large body size of post-metamorphic L. catesbeianus may enable it to utilize larger food resources that are otherwise unavailable to the smaller-bodied natives. Separate comparison of larval and adult functional traits between non-natives and the native community may help predict their potential establishment or invasion success as well as aid in the development of stage-specific control or eradication efforts.

How Widespread are Nonnative Species? Estimating Habitat Occupancy in an Ecological Reserve within a Megacity

Exotic species are fairly common within cities. However, urban areas also contain isolated fragments of original ecosystems, which may act as small ecological reserves. Unfortunately, these natural areas that are completely surrounded by urban structures are more vulnerable to human-induced disturbance and to the presence of nonnative species. Given the importance of these remnants of original ecosystems embedded within cities, efforts must be made to know if, once a nonnative species enters an urban reserve, it occupies the majority of the area or becomes restricted to some specific patches. This knowledge is important to determine the invasiveness and success of nonnative species, as well as the level of threat to the native biota. Additionally, it is important to understand which microhabitat conditions promote the presence and establishment of such nonnative species. Here, we used occupancy models to estimate the proportion of the total area that is occupied by exotic species of plants and animals within a natural reserve completely surrounded by urban areas. We also examined which environmental factors are associated with the presence of these exotic plants and animals. Our results revealed that occupancy of nonnative species is relatively high (>40%) within this urban reserve. However, we found an overall preference of nonnatives for specific patches of the reserve where human-made structures are present and where gardening activities take place. Some of the species that we studied deserve special attention because of their potential negative effects on native species.