Responses of the soil fungal communities to the co‐invasion of two invasive species with different cover classes

• Soil fungal communities play an important role in the successful invasion of non‐native species. It is common for two or more invasive plant species to co‐occur in invaded ecosystems.
• This study aimed to determine the effects of co‐invasion of two invasive species (Erigeron annuus and Solidago canadensis) with different cover classes on soil fungal communities using high‐throughput sequencing.
• Invasion of E. annuus and/or S. canadensis had positive effects on the sequence number, operational taxonomic unit (OTU) richness, Shannon diversity, abundance‐based cover estimator (ACE index) and Chao1 index of soil fungal communities, but negative effects on the Simpson index. Thus, invasion of E. annuus and/or S. canadensis could increase diversity and richness of soil fungal communities but decrease dominance of some members of these communities, in part to facilitate plant further invasion, because high soil microbial diversity could increase soil functions and plant nutrient acquisition. Some soil fungal species grow well, whereas others tend to extinction after non‐native plant invasion with increasing invasion degree and presumably time. The sequence number, OTU richness, Shannon diversity, ACE index and Chao1 index of soil fungal communities were higher under co‐invasion of E. annuus and S. canadensis than under independent invasion of either individual species.
• The co‐invasion of the two invasive species had a positive synergistic effect on diversity and abundance of soil fungal communities, partly to build a soil microenvironment to enhance competitiveness of the invaders. The changed diversity and community under co‐invasion could modify resource availability and niche differentiation within the soil fungal communities, mediated by differences in leaf litter quality and quantity, which can support different fungal/microbial species in the soil.

     

Understanding invasion history: genetic structure and diversity of two globally invasive plants and implications for their management

Aim Resolving the origin of invasive plant species is important for understanding the introduction histories of successful invaders and aiding strategies aimed at their management. This study aimed to infer the number and origin(s) of introduction for the globally invasive species, Macfadyena unguis‐cati and Jatropha gossypiifolia using molecular data. Location Native range: Neotropics; Invaded range: North America, Africa, Europe, Asia, Pacific Islands and Australia. Methods We used chloroplast microsatellites (cpSSRs) to elucidate the origin(s) of introduced populations and calculated the genetic diversity in native and introduced regions. Results Strong genetic structure was found within the native range of M. unguis‐cati, but no genetic structuring was evident in the native range of J. gossypiifolia. Overall, 27 haplotypes were found in the native range of M. unguis‐cati. Only four haplotypes were found in the introduced range, with more than 96% of introduced specimens matching a haplotype from Paraguay. In contrast, 15 haplotypes were found in the introduced range of J. gossypiifolia, with all invasive populations, except New Caledonia, comprising multiple haplotypes. Main conclusions These data show that two invasive plant species from the same native range have had vastly different introduction histories in their non‐native ranges. Invasive populations of M. unguis‐cati probably came from a single or few independent introductions, whereas most invasive J. gossypiifolia populations arose from multiple introductions or alternatively from a representative sample of genetic diversity from a panmictic native range. As introduced M. unguis‐cati populations are dominated by a single haplotype, locally adapted natural enemies should make the best control agents. However, invasive populations of J. gossypiifolia are genetically diverse and the selection of bio‐control agents will be considerably more complex.     

Ecological filtering of exotic plants in an Australian sub‐alpine environment

We investigated some of the factors influencing exotic invasion of native sub‐alpine plant communities at a site in southeast Australia. Structure, floristic composition and invasibility of the plant communities and attributes of the invasive species were studied. To determine the plant characteristics correlated with invasiveness, we distinguished between roadside invaders, native community invaders and non‐invasive exotic species, and compared these groups across a range of traits including functional group, taxonomic affinity, life history, mating system and morphology. Poa grasslands and Eucalyptus‐Poa woodlands contained the largest number of exotic species, although all communities studied appeared resilient to invasion by most species. Most community invaders were broad‐leaved herbs while roadside invaders contained both herbs and a range of grass species. Over the entire study area the richness and cover of native and exotic herbaceous species were positively related, but exotic herbs were more negatively related to cover of specific functional groups (e.g. trees) than native herbs. Compared with the overall pool of exotic species, those capable of invading native plant communities were disproportionately polycarpic, Asteracean and cross‐pollinating. Our data support the hypothesis that strong ecological filtering of exotic species generates an exotic assemblage containing few dominant species and which functionally converges on the native assemblage. These findings contrast with those observed in the majority of invaded natural systems. We conclude that the invasion of closed sub‐alpine communities must be viewed in terms of the unique attributes of the invading species, the structure and composition of the invaded communities and the strong extrinsic physical and climatic factors typical of the sub‐alpine environment.     

Responses of a native beetle to novel exotic plant species with varying invasion history

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Giant invasive Heracleum persicum: Friend or foe of plant diversity?

The impact of invasion on diversity varies widely and remains elusive. Despite the considerable attempts to understand mechanisms of biological invasion, it is largely unknown whether some communities’ characteristics promote biological invasion, or whether some inherent characteristics of invaders enable them to invade other communities. Our aims were to assess the impact of one of the massive plant invaders of Scandinavia on vascular plant species diversity, disentangle attributes of invasible and noninvasible communities, and evaluate the relationship between invasibility and genetic diversity of a dominant invader. We studied 56 pairs of Heracleum persicum Desf. ex Fisch.‐invaded and noninvaded plots from 12 locations in northern Norway. There was lower native cover, evenness, taxonomic diversity, native biomass, and species richness in the invaded plots than in the noninvaded plots. The invaded plots had nearly two native species fewer than the noninvaded plots on average. Within the invaded plots, cover of H. persicum had a strong negative effect on the native cover, evenness, and native biomass, and a positive association with the height of the native plants. Plant communities containing only native species appeared more invasible than those that included exotic species, particularly H. persicum. Genetic diversity of H. persicum was positively correlated with invasibility but not with community diversity. The invasion of a plant community by H. persicum exerts consistent negative pressure on vascular plant diversity. The lack of positive correlation between impacts and genetic diversity of H. persicum indicates that even a small founder population may cause high impact. We highlight community stability or saturation as an important determinant of invasibility. While the invasion by H. persicum may decrease susceptibility of a plant community to further invasion, it severely reduces the abundance of native species and makes them more vulnerable to competitive exclusion.     

Soil modification by invasive plants: effects on native and invasive species of mixed-grass prairies

Invasive plants are capable of modifying attributes of soil to facilitate further invasion by conspecifics and other invasive species. We assessed this capability in three important plant invaders of grasslands in the Great Plains region of North America: leafy spurge (Euphorbia esula), smooth brome (Bromus inermis) and crested wheatgrass (Agropyron cristatum). In a glasshouse, these three invasives or a group of native species were grown separately through three cycles of growth and soil conditioning in both steam-pasteurized and non-pasteurized soils, after which we assessed seedling growth in these soils. Two of the three invasive species, Bromus and Agropyron, exhibited significant self-facilitation via soil modification. Bromus and Agropyron also had significant facilitative effects on other invasives via soil modification, while Euphorbia had significant antagonistic effects on the other invasives. Both Agropyron and Euphorbia consistently suppressed growth of two of three native forbs, while three native grasses were generally less affected. Almost all intra- and interspecific effects of invasive soil conditioning were dependent upon presence of soil biota from field sites where these species were successful invaders. Overall, these results suggest that that invasive modification of soil microbiota can facilitate plant invasion directly or via ‘cross-facilitation’ of other invasive species, and moreover has potential to impede restoration of native communities after removal of an invasive species. However, certain native species that are relatively insensitive to altered soil biota (as we observed in the case of the forb Linum lewisii and the native grasses), may be valuable as ‘nurse’species in restoration efforts.     

Elevated dominance of extrafloral nectary-bearing plants is associated with increased abundances of an invasive ant and reduced native ant richness

Aim Invasive ants can have substantial and detrimental effects on co‐occurring community members, especially other ants. However, the ecological factors that promote both their population growth and their negative influences remain elusive. Opportunistic associations between invasive ants and extrafloral nectary (EFN)‐bearing plants are common and may fuel population expansion and subsequent impacts of invasive ants on native communities. We examined three predictions of this hypothesis, compared ant assemblages between invaded and uninvaded sites and assessed the extent of this species in Samoa. Location The Samoan Archipelago (six islands and 35 sites). Methods We surveyed abundances of the invasive ant Anoplolepis gracilipes, other ant species and EFN‐bearing plants. Results Anoplolepis gracilipes was significantly more widely distributed in 2006 than in 1962, suggesting that the invasion of A. gracilipes in Samoa has progressed. Furthermore, (non‐A. gracilipes) ant assemblages differed significantly between invaded and uninvaded sites. Anoplolepis gracilipes workers were found more frequently at nectaries than other plant parts, suggesting that nectar resources were important to this species. There was a strong, positive relationship between the dominance of EFN‐bearing plants in the community and A. gracilipes abundance on plants, a relationship that co‐occurring ants did not display. High abundances of A. gracilipes at sites dominated by EFN‐bearing plants were associated with low species richness of native plant‐visiting ant species. Anoplolepis gracilipes did not display any significant relationships with the diversity of other non‐native ants. Main conclusions Together, these data suggest that EFN‐bearing plants may promote negative impacts of A. gracilipes on co‐occurring ants across broad spatial scales. This study underscores the potential importance of positive interactions in the dynamics of species invasions. Furthermore, they suggest that conservation managers may benefit from explicit considerations of potential positive interactions in predicting the identities of problematic invaders or the outcomes of species invasions.     

Differential invasion success in aquatic invasive species: the role of within- and among-population genetic diversity

Despite a well-developed theoretical basis for the role of genetic diversity in the colonization process, contemporary investigations of genetic diversity in biological invasions have downplayed its importance. Observed reductions in genetic diversity have been argued to have a limited effect on the success of establishment and impact based on empirical studies; however, those studies rarely include assessment of failed or comparatively less-successful biological invasions. We address this gap by comparing genetic diversity at microsatellite loci for taxonomically and geographically paired aquatic invasive species. Our four species pairs contain one highly successful and one less-successful invasive species (Gobies: Neogobius melanostomus, Proterorhinus semilunaris; waterfleas: Bythotrephes longimanus, Cercopagis pengoi; oysters: Crassostrea gigas, Crassostrea virginica; tunicates: Bortylloides violaceous, Botryllus schlosseri). We genotyped 2717 individuals across all species from multiple locations in multiple years and explicitly test whether genetic diversity is lower for less-successful biological invaders within each species pair. We demonstrate that, for gobies and tunicates, reduced allelic diversity is associated with lower success of invasion. We also found that less-successful invasive species tend to have greater divergence among populations. This suggests that intraspecific hybridization may be acting to convert among-population variation to within-population variation for highly successful invasive species and buffering any loss of diversity. While our findings highlight the species-specific nature of the effects of genetic diversity on invasion success, they do support the use of genetic diversity information in the management of current species invasions and in the risk assessment of potential future invaders.     

There is more than one ‘killer shrimp’: trophic positions and predatory abilities of invasive amphipods of Ponto‐Caspian origin

1. Biological invasions are regarded as one of the greatest threats to biological diversity. One of the macroinvertebrate groups with the largest number of invasive species in fresh water are gammarid amphipods. Their omnivorous (including predatory) feeding behaviour may facilitate their spread and establishment in new areas. 2. Dikerogammarus villosus, the ‘killer shrimp’, is a well‐known example of a Ponto‐Caspian gammarid that is a very effective predator and successful coloniser in Europe. There are, however, other invasive Ponto‐Caspian amphipods, which have spread successfully in Northern, Central and Western Europe. Our aim here was to test whether two of such invaders (Pontogammarus robustoides and Dikerogammarus haemobaphes) are also more predacious than a native species (Gammarus fossarum). 3. Stable isotope analysis (δ¹⁵N and δ¹³C) of Ponto‐Caspian amphipods coexisting in a reservoir demonstrated that the trophic positions of P. robustoides and D. haemobaphes were similar to that of D. villosus. Echinogammarus ischnus and Chelicorophium curvispinum occupied the lowest position in the food web, while the native Gammarus fossarum (collected from another waterbody) had an intermediate trophic position. 4. Stomach content analysis of P. robustoides, D. haemobaphes and G. fossarum collected in the field, as well as laboratory feeding experiments, was used to compare diet and feeding preferences among the two invasive and one native species. All three species were omnivorous and predacious. However, the two invasive species (P. robustoides and D. haemobaphes) were more effective predators than G. fossarum and showed a clear preference for animal prey and tissue. 5. Pontogammarus robustoides and D. haemobaphes may, like D. villosus, also be called ‘killer shrimps’ and could have a similar impact as invaders of European freshwater and brackish waterbodies.     

Domination of invasive Nostocales (Cyanoprokaryota) at 52°N latitude

Cylindrospermopsis raciborskii and Sphaerospermopsis aphanizomenoides are increasingly widespread cyanobacterial species, considered invasive. However, they have never been found to dominate the phytoplankton in the northern part of the invaded area. The aim of our study was to expand the understanding of the invasion process in Nostocales, by answering the question which environmental conditions enabled the first ever noted bloom incident of C. raciborskii and S. aphanizomenoides in the northern part of their current occurrence ranges. We examined the population dynamics of the two invasive cyanobacteria and the abiotic conditions in which their blooms developed in a reservoir at 52°N. We also examined the phytoplankton community diversity and composition before and during the invasive species blooms. As shown by redundancy analysis, the competitive advantage of S. aphanizomenoides depended strongly on high water temperature and high concentration of phosphates, and was positively related to other summer conditions (low water transparency, high ammonium nitrogen concentration). C. raciborskii biomass was not related to temperature and phosphates, but we argue that high water temperature in the first half of July enabled abundant akinete germination, and pulsed phosphorus availability synergized to bring about a later bloom. Both the invasive species co‐occurred with the native cyanobacterium Aphanizomenon gracile, but C. raciborskii bloomed after the native species domination period. The results of the study corroborate the high competitiveness of S. aphanizomenoides in hot, nutrient‐rich conditions, and the adaptation of the invaders to the environment in the invaded regions, inter alia by the overwintering mechanism of numerous akinete differentiation. Our results also contradict the assumption that the phytoplankton diversity index would decrease after colonization of freshwater bodies by invasive cyanoprokaryotes.     

Low genetic diversity of the successful invasive African clawed frog Xenopus laevis (Pipidae) in Chile

In Africa, the genus Xenopus presents cryptic species and diverse hybrids between species. It has been assumed that the invasive populations of this genus correspond to X. laevis and that they are derived from the subspecies that inhabits the Mediterranean Cape region of South Africa. In part, this is supported by the successful establishment of this species in several Mediterranean regions of the world. In Mediterranean Chile, Xenopus has invaded an area of about 21,000 km², with scarce attention to genetic aspects underlying its invasion. Using mitochondrial DNA sequences we determined that Xenopus laevis laevis from the Cape region of South Africa is the subspecies that invaded Chile. The analysis indicated that the invaders have low genetic diversity (only two haplotypes, compared to 10 in two localities of their native range), and that probably the invasion in Chile occurred only once. Landscape genetics revealed that factors such as aridity and elevation have determined the spread of the species, both from the ecological and genetic points of view. Our results show that the invasion of the African clawed frog in Chile has been successful for at least 30 years, in spite of low genetic variability, few events of introduction, low propagule pressure, and bottlenecks in the founding population.     

Native jewelweed,but not other native species,displays post‐invasion trait divergence

Invasive exotic plants reduce the diversity of native communities by displacing native species. According to the coexistence theory, native plants are able to coexist with invaders only when their fitness is not significantly smaller than that of the exotics or when they occupy a different niche. It has therefore been hypothesized that the survival of some native species at invaded sites is due to post‐invasion evolutionary changes in fitness and/or niche traits. In common garden experiments, we tested whether plants from invaded sites of two native species, Impatiens noli‐tangere and Galeopsis speciosa, outperform conspecifics from non‐invaded sites when grown in competition with the invader (Impatiens parviflora). We further examined whether the expected superior performance of the plants from the invaded sites is due to changes in the plant size (fitness proxy) and/or changes in the germination phenology and phenotypic plasticity (niche proxies). Invasion history did not influence the performance of any native species when grown with the exotic competitor. In I. noli‐tangere, however, we found significant trait divergence with regard to plant size, germination phenology and phenotypic plasticity. In the absence of a competitor, plants of I. noli‐tangere from invaded sites were larger than plants from non‐invaded sites. The former plants germinated earlier than inexperienced conspecifics or an exotic congener. Invasion experience was also associated with increased phenotypic plasticity and an improved shade‐avoidance syndrome. Although these changes indicate fitness and niche differentiation of I. noli‐tangere at invaded sites, future research should examine more closely the adaptive value of these changes and their genetic basis.     

Environmentally‐contingent behaviour of invasive plants as drivers or passengers

Invasive plants are often regarded as drivers that actively reduce diversity and alter ecosystem processes such as succession. Alternatively, invaders may be passengers that simply colonize openings produced by anthropogenic disturbance and are present only temporarily. Here we test whether the behaviour of invasive species as drivers or passengers is contingent on disturbance and nutrient availability. We created twelve experimental environments (three levels of annual disturbance × four levels of nitrogen availability) for 18 years in a grassland at the northern edge of the North American Great Plains. Out of 19 invasive species initially present, two perennials (Bromus inermis, a grass, and Cirsium arvense, a forb) acted as drivers, maintaining or increasing dominance, maintaining low species richness, and forming an invader‐ dominated successional sequence. Behaviour as drivers was environmentally contingent: Bromus behaved as a driver only in less disturbed environments, and the tendency of Cirsium to behave as a driver increased significantly with both disturbance and nitrogen availability. Most invasive species (90%), however, consistently behaved as passengers, disappearing or becoming rare. The importance of disturbance and fertility for starting invasions is well‐known, but our study shows that these factors also contribute to the behaviour of some invaders as drivers. The emergence of drivers and invader‐dominated successional sequences suggests that, as rates of invasion, disturbance and eutrophication continue to increase with human activity, invasive species that act as drivers may form low‐diversity communities that persist for decades.     

Contrasting patterns of genetic diversity and spatial structure in an invasive symbiont-host association

Do host invaders and their associated symbiont co-invaders have different genetic responses to the same invasion process? To answer this question, we compared genetic patterns of native and exotic populations of an invasive symbiont-host association. This is an approach applied by very few studies, of which most are based on parasites with complex life cycles. We used the mitochondrial genetic marker cytochrome oxidase subunit I (COI) to investigate a non-parasitic freshwater ectosymbiont with direct life-cycle, low host specificity and well-documented invasion history. The study system was the crayfish Procambarus clarkii and its commensal ostracod Ankylocythere sinuosa, sampled in native (N American) and exotic (European) ranges. Results of analyses indicated: (1) higher genetic diversity in the symbiont than its host; (2) genetic diversity loss in the exotic range for both species, but less pronounced in the symbiont; (3) native populations genetically structured in space, with stronger patterns in the symbiont and (4) loss of spatial genetic structure in the exotic range in both species. The combination of historical, demographic and genetic data supports a higher genetic diversity of source populations and a higher propagule size that allowed the symbiont to overcome founder effects better than its host co-invader. Thus, the symbiont might be endowed with a higher adaptive potential to new hosts or off-host environmental pressures expected in the invasive range. We highlight the usefulness of this relatively unexplored kind of symbiont-host systems in the invasion context to test important ecological and evolutionary questions.     

Unravelling the global invasion routes of a worldwide invader,the red swamp crayfish (Procambarus clarkii)

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Genotypic diversity enhances invasive ability of Spartina alterniflora

Although genetic diversity is very important for alien species, which have to cope with new environments, little is known about the role that genetic diversity plays in their invasive success. In this study, we set up a manipulation experiment including three levels of genotypic diversity to test whether genotypic diversity can enhance the invasive ability of alien species, in our case the invasive Spartina alterniflora in China, and to infer the underlying mechanisms. There was no significant relationship between genotypic diversity and parameters of performance in the first year; however, from the summer of the second year onwards, genotypic diversity enhanced four of the six parameters of performance. After two growing seasons, there were significant positive relationships between genotypic diversity and maximum spread distance, patch size, shoot number per patch, and aboveground biomass. Moreover, abundance of the native dominant species Scirpus mariqueter was marginally significantly decreased with genotypic diversity of S. alterniflora, suggesting that enhanced invasive ability of S. alterniflora may have depressed the growth of the native species. There was no significant difference in most measures of performance among six genotypes, but we observed a transgressive over performance in four measures in multiple‐genotype patches. At the end of the experiment, there were significant nonadditive effects of genotypic diversity according to Monte Carlo permutations, in six‐genotype, but not three‐genotype plots. Our results indicated that both additive and nonadditive effects played roles in the positive relationship between genetic diversity and invasion success, and nonadditive effects were stronger as duration increased.     

Predation and avoidance behaviour in aphid‐ladybird interactions of native and invasive ladybirds in Europe

1. While detrimental effects of invasive predators on native species are well documented, we often lack a mechanistic understanding of the invasion success. Lack of prey avoidance behaviour can lead to higher consumption rates by invasive predators compared to native predators. This competitive advantage is expected to contribute to the invasion success of non‐native predators.
2. We compared aphid consumption and cue avoidance behaviour of aphids between four native ladybird species (Coccinella septempunctata, Adalia bipunctata, Propylea quatuordecimpunctata, and Hippodamia variegata) and the invasive Asian ladybird Harmonia axyridis.
3. The invasive H. axyridis and the native C. septempunctata consumed more aphids than the three smaller native ladybird species. In line with our expectations, aphids avoided leaves bearing cues of most native ladybird species but not of the invasive H. axyridis.
4. Our results indicate that body size rather than ladybird origin determined aphid predation rates. The lack of aphid avoidance behaviour towards cues of H. axyridis indicates that they were not able to recognise the chemical cues of the invasive predator.
5. Relatively large body size and the absence of cue avoidance in aphids might benefit the invasive H. axyridis, particularly in comparison to smaller native ladybird species. The absence of avoidance behaviour in aphids might lead to even higher predation rates of H. axyridis under more natural conditions.

     

Genetic patterns across multiple introductions of the globally invasive crab genus Carcinus

The European green crab Carcinus maenas is one of the world's most successful aquatic invaders, having established populations on every continent with temperate shores. Here we describe patterns of genetic diversity across both the native and introduced ranges of C. maenas and its sister species, C. aestuarii, including all known non‐native populations. The global data set includes sequences from the mitochondrial cytochrome c oxidase subunit I gene, as well as multilocus genotype data from nine polymorphic nuclear microsatellite loci. Combined phylogeographic and population genetic analyses clarify the global colonization history of C. maenas, providing evidence of multiple invasions to Atlantic North America and South Africa, secondary invasions to the northeastern Pacific, Tasmania, and Argentina, and a strong likelihood of C. maenas × C. aestuarii hybrids in South Africa and Japan. Successful C. maenas invasions vary broadly in the degree to which they retain genetic diversity, although populations with the least variation typically derive from secondary invasions or from introductions that occurred more than 100 years ago.     

The making of a rapid plant invader: genetic diversity and differentiation in the native and invaded range of Senecio inaequidens

To become invasive, exotic species have to succeed in the consecutive phases of introduction, naturalization, and invasion. Each of these phases leaves traces in genetic structure, which may affect the species’ success in subsequent phases. We examined this interplay of genetic structure and invasion dynamics in the South African Ragwort (Senecio inaequidens), one of Europe’s fastest plant invaders. We used AFLP and microsatellite markers to analyze 19 native African and 32 invasive European populations. In combination with historic data, we distinguished invasion routes and traced them back to the native source areas. This revealed that different introduction sites had markedly different success in the three invasion phases. Notably, an observed lag‐phase in Northern Germany was evidently not terminated by factors increasing the invasiveness of the resident population but by invasive spread from another introduction centre. The lineage invading Central Europe was introduced to sites in which winters are more benign than in the native source region. Subsequently, this lineage spread into areas in which winter temperatures match the native climate more closely. Genetic diversity clearly increases with population age in Europe and less clearly decreases with spread rate up to population establishment. This indicates that gene flow along well‐connected invasion routes counteracted losses of genetic diversity during rapid spread. In summary, this study suggests that multiple introductions, environmental preadaptation and high gene flow along invasion routes contributed to the success of this rapid invader. More generally, it demonstrates the benefit of combining genetic, historical, and climatic data for understanding biological invasions.