Factors affecting the efficacy of the leaf-spot fungus Mycosphaerella polygoni-cuspidati (Ascomycota): A potential classical biological control agent of the invasive alien weed Fallopia japonica (Polygonaceae) in the UK

Fallopia japonica, commonly known as Japanese knotweed, is an increasingly serious invasive alien weed in the UK and large parts of mainland Europe, as well as in North America. There is an urgent need to include classical biological control (CBC) into any integrated pest management strategy. The leaf-spot pathogen Mycosphaerella polygoni-cuspidati, a coevolved natural enemy of F. japonica present throughout its native Japanese range, is considered to have high potential as a CBC agent. In this study, the disease development of M. polygoni-cuspidati in the field and the optimum infection parameters under controlled conditions were investigated to elucidate the pathogen’s potential biocontrol efficacy against Japanese knotweed. Field observation in Japan showed that M. polygoni-cuspidati caused severe damage to its host plant. When sentinel knotweed plants from the UK were placed amongst naturally-infected field populations of F. japonica, disease incidence and severity were highest in July when monthly precipitation was also highest. In greenhouse inoculation tests, F. japonica was shown to be most susceptible at the young leaf stage (7–12 days after opening). Disease severity was highest after an initial dew period of 42–48 h, and severe defoliation followed inoculation at a temperature range of 15–25 °C. The optimum post-inoculation temperature after dew treatment for disease severity was 20–25 °C. In field inoculation tests, high disease incidence and severity indicate that the pathogen has the potential to control the plant effectively in the field. Humidity and temperature were shown to be the main factors influencing disease expression and lesion development of M. polygoni-cuspidati in a field situation. These results provide valuable information for any future use of M. polygoni-cuspidati as a CBC agent for management of Japanese knotweed in the UK.     

Barriers to hybridisation and their conservation implications for a highly threatened Australian fish species

Hybridisation and introgression are natural phenomena that may lead to the transfer of adaptive alleles from one species to another and increased species diversity. At the same time, hybridisation and subsequent introgression threaten many species world‐wide through the loss of genetic and species diversity. In Australia, introgressive hybridisation between native and alien species has not typically been considered a significant threat to native biodiversity because of the taxonomic distance between native and alien biota. However, many native fish have been introduced outside their natural range. Recently, four taxa in the genus Melanotaenia have been nationally listed as threatened due to introgressive hybridisation with introduced Melanotaenia splendida. We examined pre‐ and post‐zygotic barriers to hybridisation between M. splendida and one of these threatened taxa—Running River rainbowfish (RRR)—to assess the potential for hybridisation to occur. We used dichotomous mate choice experiments to examine pre‐zygotic barriers and mating experiments to examine post‐zygotic barriers. Size was not a significant predictor of the proportion of time subjects spent with a potential mate, nor was there any significant difference in the amount of time subjects spent with potential mates of their own or the opposite species. Eggs from hybrid pairings with female RRR had a slightly higher hatching rate than those from hybrid pairings with female M. splendida, but neither were significantly different from intraspecies crosses. We could not identify any definite barriers to hybridisation, demonstrating that the introduction of “native” fish species outside their natural range poses a higher risk of hybridisation than previously thought. We call for better education around the consequences of moving “native” fish and the development of rapid response plans to deal with recently established alien populations of Australian fish species in order to prevent future extinctions due to introgressive hybridisation.     

Extreme differences in population structure and genetic diversity for three invasive congeners: knotweeds in western North America

Japanese, giant, and the hybrid Bohemian knotweeds (Fallopia japonica, F. sachalinensis and F. × bohemica) have invaded the western USA and Canada, as well as other regions of the world. The distribution of these taxa in western North America, and their mode of invasion, is relatively unresolved. Using amplified fragment length polymorphisms of 858 plants from 131 populations from British Columbia to California to South Dakota, we determined that Bohemian knotweed was the most common taxon (71 % of all plants). This result is in contrast to earlier reports of F. × bohemica being uncommon or non-existent in the USA, and also differs from the European invasion where it is rarer. Japanese knotweed was monotypic, while giant knotweed and Bohemian knotweed were genetically diverse. Our genetic data suggest that Japanese knotweed in western North America spreads exclusively by vegetative reproduction. Giant knotweed populations were mostly monotypic, with most containing distinct genotypes, suggesting local spread by vegetative propagules, whereas Bohemian knotweed spreads by both seed and vegetative propagules, over both long and short distances. The high relative abundance and genetic diversity of Bohemian knotweed make it a priority for control in North America.     

The potential role of polyploidy and hybridisation in the further evolution of the highly invasive <Emphasis Type="Italic">Fallopia</Emphasis> taxa in Europe

Japanese knotweed s.l. comprises Fallopia japonica, F. sachalinensis, F. × bohemica and any F2s or backcrosses. The parental taxa were introduced from the East to the West as garden ornamentals in the nineteenth century, and soon spread beyond the confines of the garden to become widespread and persistent weeds. Since only female F. japonica var. japonica was introduced, its impressive spread has occurred solely by vegetative means. However, the initial lack of genetic variability has been complemented by an extensive series of hybridisations in the adventive range. We examine the history, spread, reproductive biology and ecological impact of these species in the West. The role and importance of polyploidy and hybridisation in their invasion of the West is discussed, as are the implications of these factors for the potential further evolution of the group.     

Geographically extensive hybridization between the forest trees American butternut and Japanese walnut

We investigate the question of naturally occurring interspecific hybrids between two forest trees: the native North American butternut (Juglans cinerea L.) and the introduced Japanese walnut (Juglans ailantifolia Carrière). Using nuclear and chloroplast DNA markers, we provide evidence for 29 F₁ and 22 advanced generation hybrids in seven locations across the eastern and southern range of the native species. Two locations show extensive admixture (95% J. ailantifolia and hybrids) while other locations show limited admixture. Hybridization appears to be asymmetrical with 90.9 per cent of hybrids having J. ailantifolia as the maternal parent. This is, to our knowledge, the first genetic data supporting natural hybridization between these species. The long-term outcome of introgression could include loss of native diversity, but could also include transfer of useful traits from the introduced species.     

Evidence for introgressive hybridization of wild common quail (Coturnix coturnix) by domesticated Japanese quail (Coturnix japonica) in France

Many cases of introgressive hybridization have been reported among birds, particularly following introduction to the natural environment of individuals belonging to non-native similar taxa. This appears to be the case for common quail (Coturnix coturnix) in France where wild populations artificially come into contact with domesticated Japanese quail (Coturnix japonica) raised for meat and egg production but sometimes released for hunting purposes. In order to highlight the possible existence of gene flows between both taxa, a comparison of nuclear (25 microsatellite loci) and mitochondrial (sequencing and RFLP) DNA polymorphisms was performed on 375 common quails (from France, Spain and Morocco) and 140 Japanese quails (from France and Japan). Genetic diversity was assessed, and analyses (Factorial Correspondence Analysis, Bayesian admixture) of molecular polymorphisms revealed clear differentiation between the two taxa, making it possible to detect for hybrids among quails sampled in the wild. Eight birds expected to be common quail were found to be two pure Japanese quail, one probable backcross to C. japonica, three F1/F2 hybrids, and two probable backcrosses to Coturnix coturnix. These results show that Japanese quails were released and suggest that the two taxa hybridize in the wild. They confirm the urgent need for preventing the release of pure Japanese or hybrid quails to preserve the genetic integrity of C. coturnix. The tools developed for this study should be useful for accurate monitoring of wild quail populations within the framework of avifauna management programs.     

Leaf Decomposition and Stream Macroinvertebrate Colonisation of Japanese Knotweed,an Invasive Plant Species

Japanese knotweed (Fallopia japonica Houtt. Ronse Decrane ) is a highly invasive exotic plant that forms monocultures in riparian areas, effectively reducing plant diversity. This change in riparian plant composition alters the allocthonous input of leaf litter into adjacent streams. A field experiment was completed to understand how leaf decomposition and macroinvertebrate colonisation associated with the incorporation of exotic leaf litter. Leaf packs of Japanese knotweed, native alder (Alnus incana L.), native cottonwood (Populus trichocarpa Torr . and Gray ), and two additional mixed pack types (alder and cottonwood; alder, cottonwood, and Japanese knotweed) were placed into a 50 m stream reach in Clear Creek, Idaho, and removed over a three‐month period. Leaf decomposition and macroinvertebrate assemblages were similar between leaf types, despite differences in nitrogen and phosphorus content. The diversity of leaf types within a given leaf pack also had no effect on leaf decomposition or macroinvertebrate dynamics. These findings suggest that allochthonous inputs of Japanese knotweed fulfill a detrital function similar to that of native leaf litter. (© 2008 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

The maintenance of hybrids by parasitism in a freshwater snail

Hybrids have often been labelled evolutionary dead-ends due to their lower fertility and viability. However, there is growing awareness that hybridisation between different species may play a constructive role in animal evolution as a means to create variability. Thus, hybridisation and introgression may contribute to adaptive evolution, for example with regards to natural antagonists (parasites, predators, competitors) and adaptation to local environmental conditions. Here we investigated whether parasite intensity contributes to the continuous recreation of hybrids in 74 natural populations of Melanopsis, a complex of freshwater snails with three species. We also examined, under laboratory conditions, whether hybrids and their parental taxa differ in their tolerance of low and high temperatures and salinity levels. Infections were consistently less prevalent in males than in females, and lower in snails from deeper habitats. Infection prevalence in hybrids was significantly lower than in the parental taxa. Low hybrid infection rates could not be explained by sediment type, snail density or geographic distribution of the sampling sites. Interestingly, infected hybrid snails did not show signs of parasite-induced gigantism, whereas all parental taxa did. We found that hybrids mostly coped with extreme temperatures and salinity levels as well as their parental taxa did. Taken together, our results suggest that Melanopsis hybrids perform better in the presence of parasites and environmental stress. This may explain the widespread and long-term occurrence of Melanopsis hybrids as evidenced by paleontological and biogeographic data. Hybridisation may be an adaptive host strategy, reducing infection rates and resisting gigantism.     

Fallopia japonica, an increasingly intractable weed problem in the UK: Can fungi help cut through this Gordian knot?

A new approach to the management of one of the UK's alien superweeds, Japanese knotweed (Fallopia japonica), is currently being investigated. The classical biological control strategy is based on the premise that those neophytes which become invasive and problematic are depauperate in natural enemies, both coevolved and new encounter species. Surveys have confirmed the absence of any significant natural enemy pressure in the UK, and the presence of an extensive guild of specialist arthropod and fungal natural enemies in Japan. The results of these surveys and the follow-up studies in the UK with selected fungi are discussed.     

Reciprocally beneficial interactions between introduced plants and ants are induced by the presence of a third introduced species

Interspecific interactions play an important role in the success of introduced species. For example, the ‘enemy release’ hypothesis posits that introduced species become invasive because they escape top–down regulation by natural enemies while the ‘invasional meltdown’ hypothesis posits that invasions may be facilitated by synergistic interactions between introduced species. Here, we explore how facilitation and enemy release interact to moderate the potential effect of a large category of positive interactions – protection mutualisms. We use the interactions between an introduced plant (Japanese knotweed Fallopia japonica), an introduced herbivore (Japanese beetle Popillia japonica), an introduced ant (European red ant Myrmica rubra), and native ants and herbivores in riparian zones of the northeastern United States as a model system. Japanese knotweed produces sugary extrafloral nectar that is attractive to ants, and we show that both sugar reward production and ant attendance increase when plants experience a level of leaf damage that is typical in the plants’ native range. Using manipulative experiments at six sites, we demonstrate low levels of ant patrolling, little effect of ants on herbivory rates, and low herbivore pressure during midsummer. Herbivory rates and the capacity of ants to protect plants (as evidenced by effects of ant exclusion) increased significantly when plants were exposed to introduced Japanese beetles that attack plants in the late summer. Beetles were also associated with greater on‐plant foraging by ants, and among‐plant differences in ant‐foraging were correlated with the magnitude of damage inflicted on plants by the beetles. Last, we found that sites occupied by introduced M. rubra ants almost invariably included Japanese knotweed. Thus, underlying variation in the spatiotemporal distribution of the introduced herbivore influences the provision of benefits to the introduced plant and to the introduced ant. More specifically, the presence of the introduced herbivore converts an otherwise weak interaction between two introduced species into a reciprocally beneficial mutualism. Because the prospects for facilitation are linked to the prospects for enemy release in protection mutualisms, species introductions can have complex effects on existing species interactions, between both native and introduced species.     

Crayfish co-introduced symbiotic ostracod found on native crab in Japan: The first record of epibiont ostracod found a new host

Ankylocythere sinuosa (Rioja, 1942), a symbiotic ostracod native to North America, was found from the Japanese mitten crab Eriocheir japonica (De Haan, 1835), a species native to Japan, collected from a pond in Shizuoka City, Shizuoka Prefecture, central Japan. Introduced North American crayfish Procambarus clarkii (Girard, 1852), which is a host of A. sinuosa in their native range, inhabits ponds sympatrically with Japanese mitten crabs, and it is thought that the ostracods transferred from the exotic crayfish to the native crabs. In recent years, along with the artificial transportation of crayfish around the world, their symbiotic ostracods also have been found on the body surfaces of exotic crayfish in Europe and Japan. However, no studies have confirmed the infestation of exotic ostracods on native crustaceans in the field. A wide range of developmental stages of A. sinuosa from juveniles to adults were found in Japanese mitten crabs, and mating individuals were also found. This strongly suggests that they can reproduce on the body surface of Japanese mitten crabs. In the future, it will be necessary to strengthen measures against alien species to prevent these exotic symbionts from infestating native ecosystems, and we also need to investigate the exact impact of this symbiont on Japanese mitten crabs.     

Recombinational and mutational analyses of subspecific differentiation in Oryza sativa L.

Recombinational and mutational analyses were performed in an attempt to elucidate the nature of the genic differences and the ancestral relationship between the indica and japonica subspecies of Oryza sativa L. by taking representative varieties from each group. The absence of cytological aberrations at meiosis in indica×japonica hybrids suggest that genetic differences are predominantly genic rather than chromosomal. The pattern of variation induced in the key characters, and the frequency and spectra of mutations obtained in each group, failed to support the hypothesis that indica has given rise to japonica, and suggested that the two subspecies have originated from a common ancestor.     

Gallerucida bifasciata (Coleoptera: Chrysomelidae), a potential biological control agent for Japanese knotweed (Fallopia japonica)

Host specificity of foreign natural enemies are becoming more and more critical in classical biological control programs, as concerns about potential risk from introduced biocontrol agents have been increasing recently. Understanding the insect's fundamental and ecological host ranges is the first step in determining the potential for introduction of an insect to control invasive plants. Japanese knotweed, Fallopia japonica (Houttuyn) Ronse Decraene (Polygonaceae) is an invasive weed in the United States and Europe. A leaf beetle, Gallerucida bifasciata (Coleoptera: Chrysomelidae) is an important natural enemy attacking this plant in Asia. However, its host range records were ambiguous. This study examined the beetle's host specificity through a set of choice and no-choice tests in the laboratory and field in its native China. Gallerucida bifasciata larvae were able to complete development on seven of 87 plant species in larval development tests, while adults fed and oviposited on 10 plants in no-choice tests. Multiple choice tests showed adults strongly preferred Fallopia japonica, Persicaria perfoliata (L.) H. Gross and Polygonum multiflorum Thunb over all other plants. Open field tests and field surveys further revealed that these three species were in its ecological host range. The results of this study suggest that G. bifasciata is a potential promising agent for control of Japanese knotweed in the United States and Europe, although additional host specificity tests and risk assessment should be completed.     

Green Frogs Show Reduced Foraging Success in Habitats Invaded by Japanese knotweed

Habitat loss is causing amphibian population declines worldwide, so there is increased attention to forces that degrade remaining habitats. Terrestrial habitats surrounding wetlands are critical foraging areas for temperate anurans. We investigated plant community changes in two old fields invaded by Japanese knotweed (Fallopia japonica) and the foraging success of Green frogs (Rana clamitans) in invaded and non-invaded portions of those fields. Within each field, vegetation data were recorded in quadrats located along two transects bisecting the invasion fronts. We placed frogs in ‘foraging buckets’ along transects and measured their change in mass over a 38 h period. There were significant changes in vegetation structure and composition associated with Japanese knotweed invasion. Diverse assemblages of native plants that covered non-invaded plots were absent from areas invaded by Japanese knotweed. There was also a significant change in vegetation architecture between invaded and non-invaded habitats. Change in frog mass declined significantly along transects, with most frogs in non-invaded plots gaining mass and no frogs in invaded plots gaining mass. Most frogs from non-invaded plots but only two from invaded plots defecated shortly after removal from foraging buckets (verification of recent feeding). We hypothesize that Japanese knotweed invasions degrade terrestrial habitat quality for frogs by indirectly reducing arthropod abundance. Nonnative plant invasions may be another factor contributing to amphibian population declines.     

Does the invasive species Reynoutria japonica have an impact on soil and flora in urban wastelands?

Invasive plants are recognised as a major threat to biodiversity. Although they are well-established in natural areas, the supposed negative impacts of invasive plants upon communities and ecosystems have so far been poorly investigated in urban areas, where invasions are a main issue for ecologists and for urban planners and managers. We propose to assess the effects of an invasive species along an invasion gradient in a typical urban habitat. We focused on the Japanese knotweed (Reynoutria japonica Houtt.), a widespread invasive species in Europe and North America. We considered eight urban wastelands invaded by this species in the heart of the Greater Paris Area, France. On each site, we ran four transects from the centre of the Japanese knotweed patch towards the uninvaded peripheral vegetation. We recorded the flora using the line intercept method, and several soil parameters (thickness of A horizon, abundance of earthworm casts, topsoil Munsell value, pH) every metre along each transect. The A horizon was thicker and the topsoil darker under R. japonica canopy. Thus, this invasive plant species seemed to influence soil organic matter pool. However, our results also steadily showed that R. japonica locally excluded and/or severely reduced the cover of many plant species through competition. Our study clarified the local effects of R. japonica: an influence on the soil organic matter, and a severe negative impact on wasteland plant communities. We suggest implications in both conservation and restoration ecology.     

Alien molluscan species established along the Italian shores: an update,with discussions on some?Mediterranean “alien species” categories

The state of knowledge of the alien marine Mollusca in Italy is reviewed and updated. Littorina saxatilis (Olivi, 1792), Polycera hedgpethi Er. Marcus, 1964 and Haminoea japonica Pilsbry, 1895are here considered as established on the basis of published and unpublished data, and recent records of the latter considerably expand its known Mediterranean range to the Tyrrhenian Sea. COI sequences obtained indicate that a comprehensive survey of additional European localities is needed to elucidate the dispersal pathways of Haminoea japonica.Recent records and interpretation of several molluscan taxa as alien are discussed both in light of new Mediterranean (published and unpublished) records and of four categories previously excluded from alien species lists. Within this framework, ten taxa are no longer considered as alien species, or their records from Italy are refuted. Furthermore, Trochocochlea castriotae Bellini, 1903 is considered a new synonym for Gibbula albida (Gmelin, 1791). Data provided here leave unchanged as 35 the number of alien molluscan taxa recorded from Italy as well as the percentage of the most plausible vectors of introduction, but raise to 22 the number of established species along the Italian shores during the 2005–2010 period, and backdate to 1792 the first introduction of an alien molluscan species (Littorina saxatilis) to the Italian shores.     

Unexpected genetic diversity of Fallopia japonica from Central Europe revealed after AFLP analysis

Recently much attention has been paid to genetic aspects of invasive success in Japanese knotweed s.l. One hypothesis to explain the invasive spread of these species is a multiple introduction, which leads to a higher level of genetic diversity in the invaded range. Fallopia japonica is considered to be genetically uniform in Europe, introduced as a single female clone. However, there is some evidence suggesting that invasion history and dynamics differ between Western and Central-Eastern Europe. We used AFLP markers to characterize genetic diversity of three Fallopia taxa that occur in Poland: F. japonica, F. sachalinensis and their hybrid Fallopia × bohemica, growing in so-called ‘homogeneous’ populations, consisting of one taxon and ‘heterogeneous’ populations, composed of the three taxa cohabiting together. No polymorphism, resp. an insignificantly low variability was observed in the ‘homogeneous’ populations. In the ‘heterogeneous’ stands polymorphism was detected within each taxa, with one exception that concerns individuals of F. sachalinensis from a riparian habitat. The highest level of polymorphism was found among individuals of F. × bohemica. The most striking result of our study is the observation of polymorphism between individuals of F. japonica. The AFLP data also showed that F. × bohemica is most diverse when occurring in a heterogeneous configuration with F. japonica and F. sachalinensis in the same habitat. Our results are the first evidence of genetic diversity in F. japonica populations in Central Europe and can implicate the possibility of its multiple introduction in this region or the existence of sexual reproduction of this species.     

Gene flow between alien and native races of the holoparasitic angiosperm Orobanche minor (Orobanchaceae)

The holoparasitic angiosperm Orobanche minor parasitizes a diverse range of flowering plants from at least 16 orders in both the monocots and eudicots. However, populations of O. minor show host specificity at a local level. Our previous work identified the potential for host specificity to act as a catalyst for genetic divergence among populations of O. minor. Here we have extended this investigation by sampling populations from multiple hosts, across a broad geographic range. Sequence characterised amplified region (SCAR) data identified an exotic host-generalist lineage and a native host-specialist lineage of O. minor, suggesting genetic structure in this species is defined by both host specificity and geography. In addition, host-range overlap, discordant tree topologies, and cryptic morphology indicate the presence of gene flow between alien races and endemic populations. Therefore, repeated introductions of alien races of O. minor from disparate sources leading to introgression with native populations, and cryptic race formation, seem to have contributed to the taxonomic confusion associated with this species. We speculate that radiations associated with broad host range and divergent host ecologies may have promoted the unusually wide geographic distribution and diversification of this species. Finally, evidence of multiple shifts to exotic hosts, coupled with the predicted northward shift in climatic suitability, identify the potential for range expansion in alien races of O. minor, which may threaten nationally scarce native taxa with genetic assimilation. Our phylogenetic analysis provides a framework for identifying host races in Orobanche with a view to setting conservation priorities.     

Plant genotypic diversity reduces the rate of consumer resource utilization

While plant species diversity can reduce herbivore densities and herbivory, little is known regarding how plant genotypic diversity alters resource utilization by herbivores. Here, we show that an invasive folivore—the Japanese beetle (Popillia japonica)—increases 28 per cent in abundance, but consumes 24 per cent less foliage in genotypic polycultures compared with monocultures of the common evening primrose (Oenothera biennis). We found strong complementarity for reduced herbivore damage among plant genotypes growing in polycultures and a weak dominance effect of particularly resistant genotypes. Sequential feeding by P. japonica on different genotypes from polycultures resulted in reduced consumption compared with feeding on different plants of the same genotype from monocultures. Thus, diet mixing among plant genotypes reduced herbivore consumption efficiency. Despite positive complementarity driving an increase in fruit production in polycultures, we observed a trade-off between complementarity for increased plant productivity and resistance to herbivory, suggesting costs in the complementary use of resources by plant genotypes may manifest across trophic levels. These results elucidate mechanisms for how plant genotypic diversity simultaneously alters resource utilization by both producers and consumers, and show that population genotypic diversity can increase the resistance of a native plant to an invasive herbivore.     

Establishment of the biological control agent Aphalara itadori is limited by native predators and foliage age

The knotweed psyllid, Aphalara itadori, is a biological control agent for invasive knotweed species in North America and Europe. Initial releases were conducted in Canada in 2014 but establishment has been slow, seemingly as a result of low nymphal survival. We conducted two field experiments in Ontario, Canada, to explore the effects of native predators and the age of knotweed (Fallopia japonica) foliage on nymphal survival in A. itadori. Survival of A. itadori nymphs was significantly reduced on potted plants that were exposed to native predators in the field, compared to plants from which predators were excluded. The number of surviving nymphs was also significantly reduced on older F. japonica foliage, compared to recent regrowth after a summer cutting treatment. We discuss our findings in the context of biological invasion theory and emphasize the potential for increased overlap between the fields of invasion ecology and biological control. Finally, we advocate the use of A. itadori in combination with other control measures as part of an integrated pest management programme, rather than as a solitary measure. Specifically, we recommend that future releases of A. itadori be concentrated shortly after cutting or herbicide treatments in order to maximize the availability of young tender foliage.