Environmental tolerance of an invasive riparian tree and its potential for continued spread in the southwestern US

Questions: Exotic plant invasion may be aided by facilitation and broad tolerance of environmental conditions, yet these processes are poorly understood in species‐rich ecosystems such as riparian zones. In the southwestern United States (US) two plant species have invaded riparian zones: tamarisk (Tamarix ramosissima, T. chinensis, and their hybrids) and Russian olive (Elaeagnus angustifolia). We addressed the following questions: (1) is Russian olive able to tolerate drier and shadier conditions than cottonwood and tamarisk? (2) Can tamarisk and cottonwood facilitate Russian olive invasion? Location: Arid riparian zones, southwestern US. Methods: We analyzed riparian tree seedling requirements in a controlled experiment, performed empirical field studies, and analyzed stable oxygen isotopes to determine the water sources used by Russian olive. Results: Russian olive survival was significantly higher in dense shade and low moisture conditions than tamarisk and cottonwood. Field observations indicated Russian olive established where flooding cannot occur, and under dense canopies of tamarisk, cottonwood, and Russian olive. Tamarisk and native riparian plant species seedlings cannot establish in these dry, shaded habitats. Russian olive can rely on upper soil water until 15 years of age, before utilizing groundwater. Conclusions: We demonstrate that even though there is little evidence of facilitation by cottonwood and tamarisk, Russian olive is able to tolerate dense shade and low moisture conditions better than tamarisk and cottonwood. There is great potential for continued spread of Russian olive throughout the southwestern US because large areas of suitable habitat exist that are not yet inhabited by this species.     

Influences of the invasive tamarisk leaf beetle (<Emphasis Type="Italic">Diorhabda carinulata</Emphasis>) on avian diets along the Dolores River in Southwestern Colorado USA

The tamarisk leaf beetle (Diorhabda carinulata), introduced from Eurasia in 2001 as a biological control agent for the invasive plant Tamarix ramosissima, has spread widely throughout the western USA. With D. carinulata now very abundant, scientists and restoration managers have questioned what influence this introduced arthropod might have upon the avian component of riparian ecosystems. From 2009 through 2012 we studied the consequences of biological invasions of the introduced tamarisk shrub and tamarisk leaf beetles on the diets of native birds along the Dolores River in southwestern Colorado, USA. We examined avian foraging behavior, sampled the arthropod community, documented bird diets and the use of invasive tamarisk shrubs and tamarisk leaf beetles by birds. We documented D. carinulata abundance, on what plants the beetles occurred, and to what degree they were consumed by birds as compared to other arthropods. We hypothesized that if D. carinulata is an important new avian food source, birds should consume beetles at least in proportion to their abundance. We also hypothesized that birds should forage more in tamarisk in the late summer when tamarisk leaf beetle larvae are more abundant than in early summer, and that birds should select beetle-damaged tamarisk shrubs. We found that D. carinulata composed 24.0 percent (±?19.9–27.4%) and 35.4% biomass of all collected arthropods. From the gut contents of 188 birds (25 passerine species), only four species (n?=?11 birds) contained tamarisk leaf beetle parts. Although D. carinulata comprised one-quarter of total insect abundance, frequency of occurrence in bird gut contents was only 2.1% by abundance and 3.4% biomass. Birds used tamarisk shrubs for foraging in proportion to their availability, but foraging frequency did not increase during the late summer when more tamarisk leaf beetles were present and birds avoided beetle-damaged tamarisk shrubs. Despite D. carinulata being the most abundant arthropod in the environment, these invasive beetles were not frequently consumed by birds and seem not to provide a significant contribution to avian diets.     

Ecosystem response to removal of exotic riparian shrubs and a transition to upland vegetation

Understanding plant community change over time is essential for managing important ecosystems such as riparian areas. This study analyzed historic vegetation using soil seed banks and the effects of riparian shrub removal treatments and channel incision on ecosystem and plant community dynamics in Canyon de Chelly National Monument, Arizona. We focused on how seeds, nutrients, and ground water influence the floristic composition of post-treatment vegetation and addressed three questions: (1) How does pre-treatment soil seed bank composition reflect post-treatment vegetation composition? (2) How does shrub removal affect post-treatment riparian vegetation composition, seed rain inputs, and ground water dynamics? and (3) Is available soil nitrogen increased near dead Russian olive plants following removal and does this influence post-treatment vegetation? We analyzed seed bank composition across the study area, analyzed differences in vegetation, ground water levels, and seed rain between control, cut-stump and whole-plant removal areas, and compared soil nitrogen and vegetation near removed Russian olive to areas lacking Russian olive. The soil seed bank contained more riparian plants, more native and fewer exotic plants than the extant vegetation. Both shrub removal methods decreased exotic plant cover, decreased tamarisk and Russian olive seed inputs, and increased native plant cover after 2 years. Neither method increased ground water levels. Soil near dead Russian olive trees indicated a short-term increase in soil nitrogen following plant removal but did not influence vegetation composition compared to areas without Russian olive. Following tamarisk and Russian olive removal, our study sites were colonized by upland plant species. Many western North American rivers have tamarisk and Russian olive on floodplains abandoned by channel incision, river regulation or both. Our results are widely applicable to sites where drying has occurred and vegetation establishment following shrub removal is likely to be by upland species.     

Riparian bird density decline in response to biocontrol of <Emphasis Type="Italic">Tamarix</Emphasis> from riparian ecosystems along the Dolores River in SW Colorado,USA

Biocontrol of invasive tamarisk (Tamarix spp.) in the arid Southwest using the introduced tamarisk beetle (Diorhabda elongata) has been hypothesized to negatively affect some breeding bird species, but no studies to date have documented the effects of beetle-induced defoliation on riparian bird abundance. We assessed the effects of tamarisk defoliation by monitoring defoliation rates, changes in vegetation composition, and changes in density of six obligate riparian breeding bird species at two sites along the Dolores River in Colorado following the arrival of tamarisk beetles. We conducted bird point counts from 2010 to 2014 and modeled bird density as a function of native vegetation density and extent of defoliation using hierarchical distance sampling. Maximum annual defoliation decreased throughout the study period, peaking at 32–37% in 2009–2010 and dropping to 0.5–15% from 2011–2014. Stem density of both tamarisk and native plants declined throughout the study period until 2014. Density of all bird species declined throughout most of the study, with Song Sparrow disappearing from the study sites after 2011. Blue Grosbeak, Yellow-breasted Chat, and Yellow Warbler densities were negatively related to defoliation in the previous year, while Lazuli Bunting exhibited a positive relationship with defoliation. These findings corroborate earlier predictions of species expected to be sensitive to defoliation as a result of nest site selection. Tamarisk defoliation thus had short-term negative impacts on riparian bird species; active restoration may be needed to encourage the regrowth of native riparian vegetation, which in the longer-term may result in increased riparian bird density.     

Environmental and ecological factors influencing the spillover of the non-native kelp, <Emphasis Type="Italic">Undaria pinnatifida</Emphasis>, from marinas into natural rocky reef communities

The non-native kelp, Undaria pinnatifida, is considered one of the world’s worst invasive species. The northeast Atlantic is a hotspot of Undaria invasion, yet there is limited knowledge on its invasion dynamics. In the UK its distribution is strongly associated with artificial structures, primarily marina and harbour pontoons, with relatively few records of Undaria on natural substrates. Here, the southwest UK is used as a case region, to explicitly link Undaria distribution-abundance patterns in artificial marina habitats with those in natural rocky reef habitats. Using a mixture of in situ recording and video survey techniques, Undaria was found at all thirteen marina sites surveyed; but in only 17 of 35 rocky reef sites, all of which were in 2 of the 5 larger systems surveyed (Plymouth Sound and Torbay). The distribution-abundance patterns of Undaria at reef sites were analysed using zero-inflated models. The probability of finding Undaria on rocky reef increased with increasing proximity to marinas with high abundances of Undaria. Total propagule pressure from marinas also increased the probability of occurrence, and was positively related to Undaria abundance and cover at reef sites. Increases in the cover of native kelps, Laminaria spp., and wave exposure at reef sites were linked to a reduced probability of Undaria occurrence, and lower abundance and cover. Identifying high risk areas, natural boundaries and factors affecting the spread and abundance of non-native species in natural habitats is key to future management prioritisation. Where Undaria is confined to artificial substrates management may be deemed a low priority. However, the results of this study suggest that controlling the abundance and propagule pressure in artificial habitats may limit, to some extent, the spillover of Undaria into natural rocky reef habitats, where it has the potential to interact with and influence native communities.     

Many alien invasive plants disperse against the direction of stream flow in riparian areas

Propagule pressure plays an important role in the invasion of alien plants into riparian areas. In this study, we focused on propagule pressure from both neighboring riparian areas and anthropogenic land-use areas because propagules are likely to originate from both sources. We tested the effects of whether neighboring units contained the alien plant species, focusing on the direction of invasion by alien plant species into the focal unit, and how much anthropogenic land was contained within the unit, focusing on both farmland and urbanized areas, on alien plant occurrences in Hyogo Prefecture, Japan. We modeled the occurrence of 10 alien plants using generalized linear models to evaluate species invasions by both propagules from both neighboring units and anthropogenic land within a unit. We also investigated the biological and ecological plant attributes that are likely related to invasion success, such as seed dispersal methods, seed size, and clonality, and tested the relationships between the model results and each species’ attributes. Results showed that the occurrence of an affected neighboring unit was positively associated with the occurrence of all 10 alien plants. Note that two alien invasive species were influenced by upstream flow direction, six species by downstream flow direction, and in two species, propagule supply was not distinguished by direction. In short, the dominant direction of dispersal was against the stream current, while dispersal in the downstream direction was less common. Species attributes were associated with these directions of dispersal. In addition, anthropogenic land was positively associated with the occurrence of most alien plants, although this effect was weaker than the neighbor unit effects. These results indicate that alien plants spreading into riparian areas do not always follow the natural flow regime; rather, they spread against the flow regime in some cases. We discuss an ecological explanation for these results and provide perspectives for future river management of alien plants that invade the riparian zone.     

Distribution and Abundance of Saltcedar and Russian Olive in the Western United States

Over the past century, two introduced Eurasian trees, saltcedar (Tamarix spp.) and Russian olive (Elaeagnus angustifolia) have become wide spread on western United States of American (U.S.) rivers. This paper reviews the literature on the following five key areas related to their distribution and abundance in the western United States: (1) the history of introduction, planting, and spread of saltcedar and Russian olive; (2) their current distribution; (3) their current abundance; (4) factors controlling their current distribution and abundance; and (5) models that have been developed to predict their future distribution and abundance. Saltcedar and Russian olive are now the third and fourth most frequently occurring woody riparian plants and the second and fifth most abundant species (out of 42 native and non-native species) along rivers in the western United States. Currently there is not a precise estimate of the areas that these species occupy in the entire West. Climatic variables are important determinants of their distribution and abundance. For example, saltcedar is limited by its sensitivity to hard freezes, whereas Russian olive appears to have a chilling requirement for bud break and seed germination, and can presumably survive colder winter temperatures. Either species can be dominant, co-dominant or sub-dominant relative to native species on a given river system. A number of environmental factors such as water availability, soil salinity, degree of streamflow regulation, and fire frequency can influence the abundance of these species relative to native species. Numerous studies suggest that both species have spread on western rivers primarily through a replacement process, whereby stress-tolerant species have moved into expanded niches that are no longer suitable for mesic native pioneer species. Better maps of current distribution and rigorous monitoring of distributional changes though time can help to resolve differences in predictions of potential future spread. An adequate understanding does not yet exist of what fraction of western riparian zones is resistant to dominance by either of these species, what fraction is at risk and could benefit from intervention, and what fraction has been altered to the point that saltcedar or Russian olive are most likely to thrive.     

Impacts of the invader giant reed (<Emphasis Type="Italic">Arundo donax</Emphasis>) on riparian habitats and ground arthropod communities

Riparian areas have experienced long-term anthropogenic impacts including the effects of plant introductions. In this study, 27 plots were surveyed across three Mediterranean rivers in north-eastern Spain to explore the effects of the invader giant reed (Arundo donax) on riparian habitat features and the diversity, trophic structure, body size, and abundances of epigeal and hypogeal arthropods in riparian areas. Using pitfall traps and Berlese funnels, this study detected a significant increase in collembola abundance and a decrease in the abundance, body size and diversity of macro-arthropods at order and family levels in invaded plots compared to native stands. Invaded and un-invaded areas also differed in the taxonomical structure of arthropod assemblies but not in trophic guild proportions. However, the fact that arthropods were smaller in A. donax soils, together with the absence of particular taxa within each trophic guild or even an entire trophic group (parasitoids), suggests that food-web alterations in invaded areas cannot be discarded. Habitat features also differed between invaded and un-invaded areas with the poorest herbaceous understory and the largest leaf litter deposition and soil carbon stock observed in A. donax plots. The type of vegetation in riparian areas followed by the total native plant species richness were identified as major causal factors to changes in the abundance, diversity and composition of macro-arthropods. However, our analyses also showed that some alterations related to A. donax invasion were inconsistent across rivers, suggesting that A. donax effects may be context dependent. In conclusion, this study highlights an impoverishment of native flora and arthropod fauna in A. donax soils, and suggests major changes in riparian food webs if A. donax displaces native riparian vegetation.     

Similar dietary but different numerical responses to nonnative tamarisk (<Emphasis Type="Italic">Tamarix</Emphasis> spp.) by two native warblers

Native species can have a range of responses to nonnative introductions, from negative to positive, and understanding how and why native species respond differently to nonnatives remains an important management challenge. Based on differences and similarities in ecology and behavior, we predicted how abundance and diet of two native warblers, Lucy’s warbler (Oreothlypis luciae) and yellow warbler (Setophaga petechia), would differ in habitats with different amounts of nonnative tamarisk trees and the three nonnative insects obligately dependent on tamarisk (Tamarix spp.). Specifically, we predicted that Lucy’s warblers would have similar densities across sites, yellow warbler densities would be inversely related to tamarisk cover, and both warblers, being generalist insectivores, would incorporate tamarisk biocontrol insects in their diet. Based on point counts and fecal samples at six sites along the Virgin River in the southwestern United States, we found that yellow warblers decreased in abundance with increasing tamarisk cover, while Lucy’s warbler abundance did not and that diet of the two warblers did not differ, with both species exhibiting strong selection for the nonnative tamarisk weevil (Coniatus splendidulus) and weak to no selection for the nonnative tamarisk leafhopper (Opsius stactogalus). Both warblers showed negative selection for the tamarisk beetle (Diorhabda carinulata) and its larvae, even when those insects were 10–100 times more abundant during outbreaks. Although both warblers exploited the novel food resources offered by tamarisk, with those insects contributing half or more of total prey biomass, Lucy’s warblers were better able to maintain densities in tamarisk habitats. We hypothesize this was due to the Lucy’s warbler’s ability to exploit a broader array of habitats surrounding tamarisk sites and its cavity nesting habit that buffers its nests from the higher temperatures and lower humidity of tamarisk-dominated habitat. Our results suggest that predictions based on detailed knowledge of the form and function of native and nonnative species can be used to predict native bird response to nonnatives.     

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.     

Urban riparian systems function as corridors for both native and invasive plant species

Riparian areas are often the only green areas left in urban and suburban landscapes, providing opportunities for conservation and connectivity of both aquatic and terrestrial organisms. While city planners and land managers often tout the importance of riparian networks for these uses, it is not well established if urban riparian plant communities are actually functioning as connected assemblages. Furthermore, urban riparian zones are well known to be highly invaded by non-native plant species and may be functioning to increase the spread of non-native species across the landscape. Here we examine connectivity of plant assemblages in riparian networks within an extensively urbanized landscape. We sampled riparian plant communities at 13 sites along three second-order streams of the Rahway River watershed, New Jersey. We also characterized propagule dispersal at each site by sampling litter packs on the river banks five times between March–October 2011 and identifying germinants from litter packs after cold stratification. Species turnover of both riparian and litter vegetation was more strongly associated with flow distance, particularly for native species, indicating that riverine systems are important for promoting connectivity of native plant assemblages in urban landscapes. However, non-native germinants significantly dominated propagule dispersal along the stream reaches, particularly early in the growing season, suggesting spread utilizing the river system and preemption may be an important mechanism for invasion success in this system. Our data show that management of invasive species should be planned and implemented at the watershed scale to reduce spread via the river system.     

Patterns of floral resource use by two dominant ant species in a biodiversity hotspot

Ecological dominance in ants is often fuelled by carbohydrate intake. Most studies have focused on the importance of invasive ant mutualistic associations with trophobionts whereas few studies have investigated the importance of floral nectar on invasion success. In this study, utilisation of temporarily available floral nectar by the invasive Argentine ant, Linepithema humile, was compared to that of the dominant native ant, Anoplolepis custodiens, within the Cape Floristic Region (CFR), a biodiversity hotspot. The effect of these two focal ant species on species composition and abundance of ground foraging ants as well as floral arthropod visitors in inflorescences of Proteacea species was assessed. Foraging activity, and trophic ecology inferred from the abundance of natural stable isotopes of Carbon (δ¹³C) and Nitrogen (δ¹⁵N), and the ratio of Carbon to Nitrogen (C:N) were compared between the two ant species during three flowering periods. Linepithema humile significantly reduced the abundance and species diversity of both above-ground and floral arthropod species abundance and composition. Linepithema humile increased its foraging activity with increasing nectar availability, switching its diet to a more herbivorous one. Anoplolepis custodiens did not respond as effectively to increasing floral nectar or negatively impact floral arthropod visitors. This study showed that the availability of floral nectar and ability of L. humile to more effectively utilise this temporarily available resource than native ants, can contribute significantly to the further spread and persistence of L. humile in natural environments in the CFR.     

Colonization potential of an endangered riparian shrub species

Riparian areas and their plant communities are threatened due to human exploitation and habitat loss. Conservation of riparian vegetation requires knowledge on limiting factors in the biology of species preventing its spread along suitable areas. It needs to be assessed if an endangered species is trapped in an extinction vortex or whether it can recover from its current bottleneck situation by management measurements. We investigate the recovery potential of an endangered riparian shrub species of European rivers, the German tamarisk, Myricaria germanica, by combining field and lab experiments on seed production, germination and wind dispersal with a modelling approach on species distribution. While the seed potential is high, wind-mediated dispersal is average, with a majority of seeds falling next to the mother shrub. The modelled dispersal kernel shows highest goodness-of-fit with a polynomic function. Including this kernel in a model on the future distribution of the species based on identification of suitable habitat, limited spread to new areas in Switzerland after 20 and 50 dispersal events is predicted. Given the current limited distribution of the German tamarisk in Switzerland, conservation efforts are required to allow for the formation of new riparian habitat. Additionally, connectivity along river networks has to be enhanced to help the species to escape the extinction vortex it is trapped in.     

Biodiversity influences invasion success of a facultative epiphytic seaweed in a marine forest

The biotic resistance hypothesis predicts that more diverse communities should have greater resistance to invasions than species-poor communities. However for facultative and obligate epiphytic invaders a high native species richness, abundance and community complexity might provide more resources for the invader to thrive to. We conducted surveys across space and time to test for the influence of native algal species abundance and richness on the abundance of the invasive facultative epiphytic filamentous alga Lophocladia lallemandii in a Mediterranean Cystoseira balearica seaweed forest. By removing different functional groups of algae, we also tested whether these relationships were dependent on the complexity and abundance of the native algal community. When invasion was first detected, Lophocladia abundance was positively related to species richness, but the correlation became negative after two years of invasion. Similarly, a negative relationship was also observed across sites. The removal experiment revealed that more complex native communities were more heavily invaded, where also a positive relationship was found between native algal richness and Lophocladia, independently of the native algal abundance. Our observational and experimental data show that, at early stages of invasion, species-rich seaweed forests are not more resistant to invasion than species-poor communities. Higher richness of native algal species may increase resource availability (i.e. substrate) for invader establishment, thus facilitating invasion. After the initial invasion stage, native species richness decreases with time since invasion, suggesting negative impacts of invasive species on native biodiversity.     

Intra- and interspecific hybridization in invasive Siberian elm

Hybridization creates unique allele combinations which can facilitate the evolution of invasiveness. Frequent interspecific hybridization between the Siberian elm, Ulmus pumila, and native elm species has been detected in the Midwestern United States, Italy and Spain. However, Ulmus pumila also occurs in the western United States and Argentina, regions where no native elm species capable of hybridizing with it occurs. We examined whether inter- or intraspecific hybridization could be detected in these regions. Nuclear markers and the program STRUCTURE helped detect interspecific hybridization and determine the population genetic structure in both the native and the two non-native ranges. Chloroplast markers identified sources of introduction into these two non-native ranges. No significant interspecific hybridization was detected between U. pumila and U. rubra in the western United States or between U. pumila and U. minor in Argentina and vice versa. However, the genetic findings supported the presence of intraspecific hybridization and high levels of genetic diversity in both non-native ranges. The evidence presented for intraspecific hybridization in the current study, combined with reports of interspecific hybridization from previous studies, identifies elm as a genus where both inter- and intraspecific hybridization may occur and help maintain high levels of genetic diversity potentially associated with invasiveness.     

Utilization of invasive tamarisk by salt marsh consumers

Plant invasions of coastal wetlands are rapidly changing the structure and function of these systems globally. Alteration of litter dynamics represents one of the fundamental impacts of an invasive plant on salt marsh ecosystems. Tamarisk species (Tamarix spp.), which extensively invade terrestrial and riparian habitats, have been demonstrated to enter food webs in these ecosystems. However, the trophic impacts of the relatively new invasion of tamarisk into marine ecosystem have not been assessed. We evaluated the trophic consequences of invasion by tamarisk for detrital food chains in the Tijuana River National Estuarine Research Reserve salt marsh using litter dynamics techniques and stable isotope enrichment experiments. The observations of a short residence time for tamarisk combined with relatively low C:N values indicate that tamarisk is a relatively available and labile food source. With an isotopic (¹⁵N) enrichment of tamarisk, we demonstrated that numerous macroinvertebrate taxonomic and trophic groups, both within and on the sediment, utilized ¹⁵N derived from labeled tamarisk detritus. Infaunal invertebrate species that took up no or limited ¹⁵N from labeled tamarisk (A. californica, enchytraeid oligochaetes, coleoptera larvae) occurred in lower abundance in the tamarisk-invaded environment. In contrast, species that utilized significant ¹⁵N from the labeled tamarisk, such as psychodid insects, an exotic amphipod, and an oniscid isopod, either did not change or occurred in higher abundance. Our research supports the hypothesis that invasive species can alter the trophic structure of an environment through addition of detritus and can also potentially impact higher trophic levels by shifting dominance within the invertebrate community to species not widely consumed. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Temporal and spatial impact of <Emphasis Type="Italic">Spartina alterniflora</Emphasis> invasion on methanogens community in Chongming Island,China

Methane production by methanogens in wetland is recognized as a significant contributor to global warming. Spartina alterniflora (S. alterniflora), which is an invasion plant in China’s wetland, was reported to have enormous effects on methane production. But studies on shifts in the methanogen community in response to S. alterniflora invasion at temporal and spatial scales in the initial invasion years are rare. Sediments derived from the invasive species S. alterniflora and the native species Phragmites australis (P. australis) in pairwise sites and an invasion chronosequence patch (4 years) were analyzed to investigate the abundance and community structure of methanogens using quantitative real-time PCR (qPCR) and Denaturing gradient gel electrophoresis (DGGE) cloning of the methyl-coenzyme M reductase A (mcrA) gene. For the pairwise sites, the abundance of methanogens in S. alterniflora soils was lower than that of P. australis soils. For the chronosequence patch, the abundance and diversity of methanogens was highest in the soil subjected to two years invasion, in which we detected some rare groups including Methanocellales and Methanococcales. These results indicated a priming effect at the initial invasion stages of S. alterniflora for microorganisms in the soil, which was also supported by the diverse root exudates. The shifts of methanogen communities after S. alterniflora invasion were due to changes in pH, salinity and sulfate. The results indicate that root exudates from S. alterniflora have a priming effect on methanogens in the initial years after invasion, and the predominate methylotrophic groups (Methanosarcinales) may adapt to the availability of diverse substrates and reflects the potential for high methane production after invasion by S. alterniflora.     

Predicted range expansion of the invasive plant <Emphasis Type="Italic">Leucaena leucocephala</Emphasis> in the Hengchun peninsula,Taiwan

We integrated correlative (species distribution) and process-based (agent-based) modeling approaches—to predict future range expansion of an invasive plant (Leucaena leucocephala) in the lowland rainforest and tropical coastal forest ecosystems of the Hengchun peninsula, Taiwan. We simulated range expansion to the year 2027 using a spatially-explicit, agent-based model representing invasion rate as a function of habitat quality and propagule pressure. We developed an index of habitat quality by relating 1988 plant distribution data to geo-referenced data on climatic conditions, landscape features, and anthropogenic factors via logistic regression. We represented propagule pressure using a lognormal dispersal kernel. We evaluated model performance by simulating range expansion from 1988 to 2007 and comparing simulated distribution patterns to those observed in 2007. Results of logistic regression indicated that L. leucocephala is more likely to occur in warm, dry areas containing a higher percentage of natural landscape (forest or grassland), areas adjacent to the forest edge, and areas disturbed by human activities such as a main road or an abandoned sisal plantation. Model evaluation showed that the prediction performance is excellent with AUC > 0.9 and Mantel’s r = 0.77. Our results indicated that L. leucocephala will continue from the western portion toward the southern and central portions of Checheng township, and throughout the southern portion of Hengchun and Manjhou townships, with about 2500 ha of new area invaded within the next 20 years. Our predictions should allow managers to develop proactive management plans for the areas most likely to be invaded.     

Drivers of invasive tree and shrub natural regeneration in temperate forests

We assessed drivers of ecological success along resource availability gradients for three invasive woody species: Prunus serotina Ehrh., Quercus rubra L. and Robinia pseudoacacia L. We aimed to check how much of invasion success, measured by invader biomass, is explained by propagule pressure and plant community invasibility. Using 3 years of observations from 372 study plots (100 m² each) in temperate forests of Wielkopolski National Park (Poland) we investigated the hierarchy of predictors and partial dependencies using the random forest method. Our study indicated that propagule pressure explained more variance in success of invaders than invasibility—describing availability of resources and competitors in understory vegetation. We also found different responses of seedlings and saplings, connected with dependence on stored carbohydrates, which decreased seedling responses to resource availability gradients. However, resource availability (light and leaf litter predictors) had greater influence than predictors describing understory vegetation. Based on importance and response strength the species studied may be arranged by decreasing requirements for soil fertility and acidity: P. serotina?<?Q. rubra?<?R. pseudoacacia, whereas for light requirements and competition vulnerability the order is: P. serotina?>?Q. rubra?>?R. pseudoacacia. However, low light requirements of R. pseudoacacia may be biased by high proportion of sprouts supplied by parental trees. Results provide guidelines for effective management of invasive woody species in forest ecosystems and describe complex interactions between factors studied on ecological success of invaders.     

Freshwater invasibility level depends on the population age structure of the invading mussel species

Environmental conditions may affect invasibility and potentially prevent successful invasions of the freshwater bivalve Limnoperna fortunei Dunker, 1857. However, even though the larval stage may be considered an important species’ characteristic, the invasion processes of L. fortunei are mainly evaluated using only the adult stage. Therefore, the aim of this study was to identify what, and how, environmental filters might predict the likelihood of occurrence of each of the L. fortunei larval stages. Logistic regressions were applied using the larval stages of L. fortunei as a surrogate of population age structure and the environmental variables as the main filters that potentially predicted the likelihood of the occurrence of each larval stage of L. fortunei. The turbidity predicted the occurrence of the D-shaped larvae and straight-hinged veliger stages, while the umbonated-veliger and pediveliger stages were predicted by the pH and conductivity. Finally, the phytoplankton density (diatoms) predicted the occurrence of the umbonated-veliger, pediveliger and mainly the plantigrade stages. Our findings suggested that, during larvae development, from younger to older stages, the main environmental predictor of larvae occurrence shifted from abiotic to biotic variables, indicating that larval stages are indeed an important factor that helps in the evaluation of freshwater invasibility. These findings are particularly important for floodplain systems, where flood pulse dynamics may increase propagule pressure, leading to a successful spread among habitats. Then, even though environmental filters were important to predict successful invasions, we highlight that a successful invasion might depend on the population age structure of the invading species arriving in the new habitat. Thus, successful L. fortunei invasions in floodplain systems may depend on the interaction between invasiveness, invasibility and propagule pressure.