Fallopia x bohemica

Pictures

Picture	Title	Caption	Copyright
Title Flowering plant Caption Fallopia x bohemica; male fertile F. x bohemica plant. Copyright ©John P. Bailey	Flowering plant	Fallopia x bohemica; male fertile F. x bohemica plant.	©John P. Bailey
Title Flowering plant Caption Fallopia x bohemica; male fertile F. x bohemica plant Copyright ©John P. Bailey	Flowering plant	Fallopia x bohemica; male fertile F. x bohemica plant	©John P. Bailey
Title Flowers and foliage Caption Fallopia x bohemica; close-up of flowers on a fertile male F. x bohemica plant. Copyright ©John P. Bailey	Flowers and foliage	Fallopia x bohemica; close-up of flowers on a fertile male F. x bohemica plant.	©John P. Bailey
Title Leaves Caption Comparison of Fallopia leaves; (a) Fallopia sachalinensis. (b) Fallopia x Bohemica. (c) Fallopia japonica var. japonica. Copyright ©John P. Bailey	Leaves	Comparison of Fallopia leaves; (a) Fallopia sachalinensis. (b) Fallopia x Bohemica. (c) Fallopia japonica var. japonica.	©John P. Bailey

Identity

Preferred Scientific Name

• Fallopia x bohemica (J. Chrtek & A. Chrtková) J. P. Bailey

Other Scientific Names

• Polygonum x bohemicum (J. Chrtek & A. Chrtková) P.F. Zika & A.L. Jacobson
• Reynoutria x bohemica J. Chrtek & A. Chrtková

International Common Names

• English: bohemian knotweed; hybrid knotweed

Local Common Names

• Poland: rdest posredni; rdestowiec posredni
• Sweden: hybridslide

Summary of Invasiveness

It is generally an under-recorded component of the knotweed populations. It occupies the same sorts of habitat as Japanese knotweed (F. japonica) does in its adventive range. There is some evidence that it is more difficult to control than its parents (Bimova et al., 2003) and that it can exploit a wider range of habitat than its parents (Bailey and Wisskirchen, 2006). Some clones cover very large areas.

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Plantae
•         Phylum: Spermatophyta
•             Subphylum: Angiospermae
•                 Class: Dicotyledonae
•                     Order: Polygonales
•                         Family: Polygonaceae
•                             Genus: Fallopia
•                                 Species: Fallopia x bohemica

Notes on Taxonomy and Nomenclature

Fallopia x bohemica (Chrtek & Chrtková) J.P. Bailey (Stace, 1989) is the hybrid between Fallopia japonica and Fallopia sachalinensis and occurs at three different ploidy levels: 2n=44; 66; and 88. Although most literature nowadays refers to it under the Fallopia combination, older papers refer to Reynoutria x bohemica Chrtek & Chrtková, and the Americans have even published the Polygonum combination; Polygonum x bohemicum (Zika and Jacobson, 2003). There are two reasons behind this generic plentitude. Firstly, Japanese knotweed [Fallopia japonica] was described twice independently, the first name Reynoutria japonica being lost for more than 100 years, and in the mean time it was described again as Polygonum cuspidatum (Bailey and Stace, 1992). However, many taxonomists now accord generic status to the former sections of the Linnaean genus Polygonum. Fallopia and Reynoutria were then the correct names for the genera containing the climbers and the Japanese knotweeds, respectively. In the 1980s, morphological and hybridization studies indicated that these two genera should be merged under Fallopia, the older name (Ronse Decraene and Akeroyd, 1988; Bailey and Stace, 1992). A recent molecular study by Galsasso et al. (2009) proposes the transfer back to Reynoutria and The Plant List (2013) gives Reynoutria x bohemica Chrtek & Chrková as the preferred name. 

Description

It has the same general growth form as F. japonica var. japonica, but the leaves are much larger and do not have the truncate bases so typical of F. japonica. Unlike F. japonica var. japonica, both sexes are found, and in the UK at least, the hermaphrodites seem to outnumber the male-sterile plants. Clones extend by rhizome growth and may occupy considerable areas.

Inflorescences are similar to those of either of the parents. Florets are 1-2.5 mm long and functionally unisexual but with each male or female flower possessing the complementary but vestigial, organs of the other sex. Each floret has 5 petals and 8 stamens (Gillespie and Faithfull, 2004).

Plant Type

Distribution

Bailey et al. (1996) give detailed information about the distribution in the British Isles with six figure grid references; more up to date data on a 10 km square basis is to be found in Preston et al. (2002).

Distribution Table

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

History of Introduction and Spread

Initially this species was thought to be part of the range of variation of Fallopia japonica var. japonica, and it was only with the application of cytology that it became apparent that there was no morphological variation in the UK F. japonica var. japonica (subsequently shown to be a male-sterile clone) and that the morphologically intermediate plants were actually the hybrid. It was only discovered in the 1980s, therefore its earlier history can only be established by a study of dated herbarium specimens (Bailey and Conolly, 2000). The fact that all F. japonica var. japonica were male-sterile, meant that once male-fertile plants of Fallopia sachalinensis reached Europe, hybridization was inevitable. Any seed collected from Japanese knotweed would thus be of hybrid origin. Botanic gardens around the world were distributing this hybrid seed as F. japonica. Evidence from herbarium material shows that this hybrid arose not long after the arrival of F. sachalinensis in the late nineteenth century (Bailey and Conolly, 2000), whereas the earliest date for a plant established in the wild is currently 1954 for County Durham, UK.

Risk of Introduction

The main method of reproduction is by vegetative propagation so there is a danger of river-borne fragments of rhizome crossing country borders. Also there are some reports of short, sea-borne transfer of viable rhizome fragments of Fallopia japonica from island to island in the west of Scotland (Beerling et al., 1994; Hayward, 2002) and there is nothing to suggest that F. x bohemica is any less adept in this ability so this possibility cannot be excluded. Richards et al. (2008) describe the presence of F. japonica and F. x bohemica in salt marsh habitats in the USA and performed experiments that indicated that both taxa exhibited considerable plasticity in their response to exposure to salt. There is little risk of inadvertent transport to continents not already occupied.

Habitat

It occurs in habitats similar to those occupied by Fallopia japonica; principally in riparian and ruderal habitats, but also in salt marshes (Richards et al., 2008).

Habitat List

Hosts/Species Affected

Not usually a problem in cultivated ground.

Biology and Ecology

Genetics

Climate

Notes on Natural Enemies

No specific research has been carried out on F. x bohemica because it is rarely found in Japan, but much of what holds for Fallopia japonica might be expected to apply to the hybrid. F. japonica in Japan is attacked by a suite of natural enemies, both arthropod and fungal, not present in its adventive range. To date, 186 species of arthropod and around 40 species of fungus have been recorded from the plant in its native range of Japan (R Shaw, CABI, UK, personal communication, 2008). As a result of this attack, it is not able to compete with local flora as effectively as it does in the introduced range and does not normally reach the same massive size. Of these natural enemies, some exert significant damage such as the chrysomelid beetle Gallerucida nigromaculata, which is described as having potential as a biocontrol agent by Zwoelfer (1973). This is now thought to be Gallerucida bifasciata and not adequately specific. Recent observations by CABI Bioscience show fungal pathogens, as well as arthropods, are significant controlling factors in Japan, across the climatic range of the plant. This includes a number of obligate biotrophic fungi, as well as an ubiquitous leafspot pathogen. In its introduced range, F. japonica is attacked by the green dock beetle, Gastrophysa viridula, but this is only when its normal Rumex host has been consumed and beetle populations are elevated.

Pathway Causes

Pathway Vectors

Impact Summary

Economic Impact

The economic impact of this species is similar to that of Fallopia japonica in that it is a problem for developers of land infested with F. x bohemica, although it is not entirely clear whether this taxon is actually covered by the letter of the law that proscribes Japanese knotweed (1981 Wildlife and Countryside Act). It is unsightly in amenity areas and a threat to biodiversity along water courses, where, like F. japonica, it is able to spread vegetatively. Control costs will equal, if not exceed, the costs for F. japonica, as there is some evidence (Bimova et al. 2003) that the hybrid is more difficult to eradicate than either parent.

For F. japonica the estimated annual control costs for one county council in Wales, UK, in 1994 was £300,000 (approximately US $600,000). The budget needed to control the 64 ha knotweed infestation in the city and county of Swansea was estimated to be £5.79 million in 1998 (Shaw et al., 2001). To control F. japonica on a national scale in the UK would cost an extrapolated £1.56 billion (approximately US $3 billion) were it to be attempted, as reported by the UK Department of Environment, Food and Rural Affairs in its recent non-native species policy review. An accepted estimate of control costs is £10,000 per hectare for a 3-year spraying regime, with two sprays per year, but this is probably an underestimate if revegetation costs are taken into account. Its presence can add around 10% to the costs of a development project, especially if soil is considered contaminated and subject to additional removal fees. Indeed, a spraying programme on a development site is estimated to be £27.19 per m² (approximately US $54 per m²), and including finance costs this almost doubles to £50.88 per m² (approximately US $100 per m²) if soil has to be removed and clean soil imported and compacted (Child and Wade, 2000).

It is present by roads and railways, and stem debris may impede water flow in smaller water courses.

Environmental Impact

Impact on Habitats

Threatened Species

Social Impact

As mentioned in the Economic impact section, it may be considered unsightly and intrusive in amenity and other public spaces. In the UK, the constant barrage of publicity in the press (much of it ill-considered i.e. ‘can grow through concrete’) has given members of the public the impression that this is some sort of indestructible Triffid, and that once spotted in a garden it won’t be long before it is found growing up between the floorboards.

Risk and Impact Factors

• Proved invasive outside its native range
• Highly adaptable to different environments
• Is a habitat generalist
• Long lived
• Fast growing
• Reproduces asexually
• Has high genetic variability

• Damaged ecosystem services
• Ecosystem change/ habitat alteration
• Infrastructure damage
• Modification of hydrology
• Modification of successional patterns
• Monoculture formation
• Negatively impacts tourism
• Reduced amenity values
• Reduced native biodiversity
• Threat to/ loss of endangered species
• Threat to/ loss of native species

• Allelopathic
• Competition - monopolizing resources
• Competition - shading
• Hybridization
• Rapid growth

• Difficult/costly to control

Uses

Economic Value

Uses List

Animal feed, fodder, forage

• Forage

Environmental

• Erosion control or dune stabilization

Fuels

• Biofuels

Human food and beverage

• Emergency (famine) food

Materials

• Chemicals
• Pesticide

Diagnosis

It is most likely to be mistaken for either of its parents Fallopia japonica and Fallopia sachalinensis. Identification is on the basis of leaf shape and the characteristic intermediate leaf hairs found on the lower epidermis of the leaves. For both of these characters, it is important that only large fully-expanded stem leaves are examined. The male-fertile clones are particularly easy to distinguish from F. japonica var. japonica. Clear characters for distinguishing the three taxa may be found in Bailey et al. (1996, 2008), Zika and Jacobson (2003) and Botta-Dukát and Balogh (2008).

Rapid Response

It is possible to eradicate knotweed if a new infestation of rhizome is spotted quickly and the resultant plants pulled or treated before the roots have become well established.

Eradication

There has been little success with eradication policies for F. japonica, but given the smaller population size, it is perhaps a realistic and worthwhile aspiration; although there is some evidence that it is more difficult to eradicate (Bimova et al., 2003), and contains greater reserves of genetic diversity than the clone of F. japonica var. japonica. Also see Child (1999), and Child and Wade (2000).

Detection and Inspection

The UK Environment Agency have produced a Code of Practice, and the Cornwall and Devon Knotweed Forum have produced an excellent guide, which has advice on identifying the plant in the field at various stages of the season.

Similarities to Other Species/Conditions

As a hybrid between Fallopia japonica and Fallopia sachalinensis it obviously shares many similarities with these plants. It should be noted that the tetraploid F. x bohemica is a cross between the dwarf montane F. japonica var. compacta and F. sachalinensis. The standard 6x and the rarer 8x F. x bohemica are both crosses between the F. japonica var. japonica clone and F. sachalinensis. The tetraploid and octoploid contain half japonica and half sachalinensis chromosomes, whereas the 6x is 66% japonica and 33% sachalinensis. This is reflected in their morphology; although they cannot currently be distinguished morphologically. It is also possible that the montane var. compacta component of the 4x may pre-adapt it to different habitats, because in the wild it is found up to 2500 m.a.s.l. (Bailey, 2003).

Prevention and Control

Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Prevention

F. japonica appears on the UK Wildlife and Countryside Act (1981) and as such it is illegal to cause the plant to grow in the wild. It is listed as a noxious weed in many states and provinces of North America and appears on many weed lists around the world. F. x bohemica is not such a well-recognised taxon, but the same conditions apply to its treatment.

SPS measures

Vehicles should be inspected when moving from infected sites to new ones.

Control

Due to a large and persistent rhizome system, F. japonica is highly resistant to control efforts (Ainsworth et al., 2002), and the same also applies to F. x bohemica. The effectiveness of control and eradication interventions has recently been reviewed thoroughly by Kabat et al. (2006), who included 65 articles in their meta-analysis. Six categories of intervention were included, none of which could eradicate Japanese knotweed or its hybrid in the short term. Cutting treatments alone were not found to result in significant decreases in knotweed abundance. However, statistically-significant reductions in abundance can be achieved by short-term application of: glyphosate; imazapyr; imazapyr and glyphosate; cutting followed by filling stems with glyphosate; and cutting followed by spraying with glyphosate (Kabat et al., 2006). However, these authors were still unable to conclude long-term efficacy for any control measure.

Physical/mechanical control

Mechanical control of F. japonica is difficult, but continual mowing will reduce the resources of the extensive rhizome system if carried out throughout the growing season. Glasshouse trials have shown that repeated cutting at least every 4 weeks and at least 7 weeks prior to senescence, can be effective (Seiger and Merchant, 1997). Pulling up plants complete with root systems can eliminate small stands and is appropriate for local eradication in sensitive areas, but only if carried out continually over a number of years (Baker, 1988). However, digging up roots is even more challenging because they can extend to a depth of 2 m, and 7 m away from the crown, and despite the best efforts, this normally results in an increased stem density. This may be useful for integrated control. With its Fallopia sachalinensis component, F. x bohemica may be slightly more palatable for stock. Certainly F. sachalinensis was promoted as fodder for livestock in the early 1900s (Bailey and Conolly, 2000).

Biological control

Much work has been done on F. japonica at CABI Biosciences (Shaw et al., 2009), but this was mainly based on the European clone of F. japonica, and the aim was to obtain agents specific to that clone. The F. japonica programme has been underway, on behalf of UK and North American sponsors, since May 2003, with two candidate agents, namely a Mycosphaerella leafspot and the psyllid Aphalara itadori. Both of these agents have undergone extensive host range testing and have good potential as biological control agents. Given the difficulty faced by property developers, there would appear to be a market for a mycoherbicide, although registration costs are hindering this approach. As a newly arisen hybrid it is doubtful that any specialist predators actually exist for F. x bohemica and it will be a matter of selecting agents that are restricted to F. japonica and F. sachalinensis.

Chemical control

For F. japonica, chemicals that are permitted on or near water are normally restricted as will be the potential for full control. Child and Wade (2000) recommended five herbicides for F. japonica control, to be applied as foliar sprays. Triclopyr and imazapyr can be applied to young, actively-growing shoots when grasslands need to be protected; glyphosate is suitable during active growth periods when leaves are fully expanded, although larger plants may need to be sprayed using a telescopic/long lance sprayer; picloram can also be used as a soil drench due to its persistence, but not where planting is required within 2 years; and 2,4-D amine is used during the active growing period and as a selective translocated herbicide to be used in grassland, amenity areas and forest situations, although this may depend on which formulation is used in which country. Of the five herbicides, only glyphosate and 2,4-D amine can be used near water. In general, cutting and removing dead stems at the end of the season, prior to a spraying regime the following season, is advisable to aid access. F. japonica is a very resilient plant and unless extremely toxic chemicals are appropriate, repeated well-timed applications should be anticipated, and follow-up spot treatments of any regrowth will often be required.

Stem injection of various herbicides is a relatively modern phenomenon and can produce very good results in some conditions, but concerns remain over the amount of chemical that is actually applied per hectare exceeding statutory maxima. Hagen and Dunwiddie (2008) discovered that using glyphosate, through the injection method, results in the short-term dieback of injected stems. However, drawbacks to its use in certain scenarios should be considered when developing an integrated management plan for knotweed control.

Much of the above will also apply to F. x bohemica, although the larger leaf area and the fact that chemical control is more difficult with F. x bohemica (Bimova et al., 2003), may point to the need for some experimentation with the dosage used and the times and frequency of application.

IPM

For F. japonica the use of a combination of mechanical and chemical techniques has proved effective, i.e. cutting and a follow-up spray of new growth, but it is necessary to apply the chemical more than once a season (de Waal, 1995). There are two basic methods: to cut plants to 5 cm height and immediately apply a 25% solution of glyphosate or triclopyr to the cut stems; or cut or mow infestations when the plants reach the early bud stage in the late spring or summer and treat the regrowth in the autumn with glyphosate or triclopyr. If deep digging is used to effectively increase the above ground:below ground biomass ratio, then subsequent chemical application can reduce the time required to achieve effective control (Child et al., 1998). Another herbicide strategy is an integrated strategy with mowing or cutting.

Similar approaches for F. x bohemica should be attempted in order to assess their effectiveness.

Monitoring and Surveillance

There are various GIS surveys on-going in the UK, the first being in Swansea, followed by Cornwall and Devon. Professor Denis Murphy and Daniel Jones at the University of Glamorgan are currently working in this area. Whilst it is highly unlikely that the system would ever be able to discriminate between the different knotweed taxa, it is nevertheless a powerful tool for charting the distribution and extent of this group.

Mitigation

Rapid eradication of F. x bohemica is only possible if rhizome growth has not been too extensive.

Ecosystem Restoration

F. japonica is able to hyper-accumulate heavy metals, including copper, zinc and cadmium, more effectively than other angiosperm species as has been demonstrated in Japan (Nishizono et al., 1989) and Croatia (Hulina and Dumija, 1999). However, the abilities of F. x bohemica in this direction remain untested.
 

Gaps in Knowledge/Research Needs

Do the Wildlife and Countryside Act and subsequent Duty of Care regulations actually apply to the hybrid F. x bohemica and if not the appropriate legislative changes should be made?

References

Adolphi K, 1999. First record of Fallopia x bohemica (Chrtek & Chrtkova) J. Bailey in the Netherlands. (De eerste vondst van Fallopia x bohemica (Chrtek and Chrtkova) J. Bailey in Nederland.) Gorteria, 25:140-142.

Bailey J, 2003. Japanese Knotweed s.l. at home and abroad. In: Plant invasions: ecological threats and management solutions [ed. by Child, L.\Brock, J. H.\Brundu, G.\Prach, K.\Pyse?k, K.\Wade, P. M.\Williamson, M.]. Leiden, Netherlands: Backhuys Publishers, 183-196.

Bailey JP, 1994. Reproductive biology and fertility of Fallopia japonica (Japanese knotweed) and its hybrids in the British Isles. In: Ecology and management of invasive riverside plants [ed. by Waal, L.C. de\Child, L.E.\Wade, P.M.\Brock, J.H.]. Chichester, UK: John Wiley & Sons Ltd, 141-158.

Bailey JP; Bímová K; Mandák B, 2008. Asexual spread versus sexual reproduction and evolution in Japanese Knotweed s.l. - the "Battle of the Clones"? Biological Invasions, 11:1189-1203.

Bailey JP; Child LE; Conolly AP, 1996. A survey of the distribution of Fallopia x bohemica (Chrtek & Chrtkova) J.Bailey (Polygonaceae) in the British Isles. Watsonia, 21:187-198.

Bailey JP; Conolly AP, 2000. Prize-winners to pariahs - a history of Japanese knotweed s.l. (Polygonaceae) in the British Isles. Watsonia, 23(1):93-110.

Bailey JP; Stace CA, 1992. Chromosome number, morphology, pairing, and DNA values of species and hybrids in the genus Fallopia (Polygonaceae). Plant Systematics and Evolution, 180(1/2):29-52.

Bailey JP; Wisskirchen R, 2006. The distribution and origins of Fallopia x bohemica (Polygonaceae) in Europe. Nord. J. Bot, 24:173-200.

Balogh L, 1998. Exomorphological observations in support of the presence of the hybrid species Fallopia x bohemica in Hungary. (Külsõ alaktani megfigyelések a Fallopia x bohemica (Chrtek & Chrtková) J. Bailey (F. japonica x F. sachalinensis) hibridfaj magyarországi jelenlétének alátámasztásához.) Kitaibelia (Debrecen), 3(2):255-256.

Balogh L, 2003. Species of the genus Fallopia sectio Reynoutria in Hungary - a land of the hybrid Fallopia x bohemica? In: Invasive Plants in Natural and Managed Systems: Linking Science and Management and 7th International Conference on the Ecology and Management of Alien Plant Invasions, Wyndham Bonaventure Resort, Ft. Lauderdale, Forida, USA, November 3-7, 2003. 7.

Beerling DJ; Bailey JP; Conolly AP, 1994. Biological Flora of the British Isles No. 183 Fallopia japonica. Journal of Ecology, 82(4):959-979.

Bímová K; Mandák B; Pysek P, 2003. Experimental study of vegetative regeneration in four invasive Reynoutria taxa (Polygonaceae). Plant Ecology, 166(1):1-11.

Botta-Dukát Z; Balogh L, 2008. The most important invasive plants in Hungary [ed. by Botta-Dukát, Z.\Balogh, L.]. Vácrátót, Hungary: HAS Institute of Ecology and Botany, 255 pp.

Callaghan TV; Lawson GJ; Scott R, 1984. Natural vegetation as a renewable energy resource in Great Britain. Cumbria, UK: Merlewood.

Child L; Wade M, 2000. The Japanese knotweed manual: the management and control of an invasive alien weed. The Japanese knotweed manual: the management and control of an invasive alien weed, xi + 123 pp.

Child LE, 1999. Vegetative regeneration and distribution of Fallopia japonica and Fallopia × bohemica: Implications for control and management., UK: Loughbrough University.

Chrtek J; Chrtkova A, 1983. [English title not available]. (Reynoutria x bohemica, novy krizenec z celedi Rdesnovitych.) Casopis Narodniho Muzea, 152:120.

Chrtek J; Chrtkova A, 1985. [English title not available]. (Krizenec Reynoutria x bohemica v Pruhonickem parku.) Ziva, 33:136-137.

Conolly AP, 1998. Fallopia x bohemica: a new record from Australia? BSBI News, 78:88.

Conolly AP, 2001. Japanese Knotweeds in Australia. BSBI News, 87:74-75.

Decraene LPR; Akeroyd JR, 1988. Generic limits in Polygonum and related genera (Polygonaceae) on the basis of floral characters. Botanical Journal of the Linnean Society, 98(4):321-371.

eFloras, 2008. Missouri Botanical Garden & Harvard University Herbaria. St Louis, USA: Missouri Botanical Garden. http://www.efloras.org

Eliás P, 2001. [English title not available]. (Dva naturalizovaNé neofyty [Sarcoca esculenta (van Houtte) skalicky a Fallopia x bohemica (Chrtek et Chrtková) J.P.Bailey v meste trnave.) Acta Fac. Paed. Univ. Tyrnaviensis, Ser. B, 5:13-21.

EPPO, 2014. PQR database. Paris, France: European and Mediterranean Plant Protection Organization. http://www.eppo.int/DATABASES/pqr/pqr.htm

Fehér A, 1999. Reconstruction of the spread of invasive species of Fallopia genus in Pozitavska pahorkatina region. In: Invasions and invasive organisms [ed. by Elias, P.]. Bratislava-Nitra, Slovakia: SNK Scope & Sekos, 96-103.

Fojcik B; Tokarska-Guzik B, 2000. Reynoutria x bohemica (Polygonaceae): a new taxon to the Polish flora. (Reynoutria x bohemica (Polygonaceae): nowy takson we florze Polski.) Fragm. Flor. Geobot. Polon, 7:63-71.

Forman J; Kesseli RV, 2003. Sexual reproduction in the invasive species Fallopia japonica (Polygonaceae). American Journal of Botany, 90(4):586-592; 26 ref.

Fremsted E; Elven R, 1997. [English title not available]. (Fremmede planter i Norge. De store Fallopia-artene.) Blyttia, 55:3-14.

Galasso G; Banfi E; Mattia FDe; Grassi F; Sgorbati S; Labra M, 2009. Molecular phylogeny of Polygonum L. (Polygonoideae, Polygonaceae), focusing on European taxa: preliminary results and systematic considerations based on rbcL plastidial sequence data. Atti della Società italiana di scienze naturali e del Museo civico di storia naturale di Milano, 150(1):113-148.

Gammon MA; Grimsby JL; Tsirelson D; Kesseli R, 2007. Molecular and morphological evidence reveals introgressions in swarms of the invasive taxa Fallopia japonica, F. sachalinensis, and F. × bohemica (Polygonaceae) in the United States. American Journal of Botany, 94:948-956.

Gilbert OL, 1992. The ecology of an urban river. British Wildlife, 3:129-136.

Gillespie P; Faithfull I, 2004. Knotweed: state prohibited weed. Knotweed: state prohibited weed. Victoria, Australia: Department of Primary Industries. http://www.dpi.vic.gov.au/dpi/nreninf.nsf/LinkView/643CD116260B0928CA256E9000824464663C5274163D2336CA256E8D001EFC6C

Hayward CM, 2002. Fallopia japonica on the Isle of Arran. BSBI News, 89:44-46.

Hollingsworth ML; Bailey JP; Hollingsworth PM; Ferris C, 1999. Chloroplast DNA variation and hybridisation between invasive populations of Japanese Knotweed and Giant Knotweed (Fallopia, Polygonaceae). Bot. J. Lin. Soc, 129:139-154.

Hollingsworth ML; Hollingsworth PM; Jenkins GI; Bailey JP; Ferris C, 1998. The use of molecular markers to study patterns of genotypic diversity in some invasive alien Fallopia spp. (Polygonaceae). Molecular Ecology, 7(12):1681-1691.

Jalas J; Suominen J, 1979. Atlas Florae Europeae Vol. 4 Polygonaceae. Helsinki, Finland: Societas Biologica Fennica Vanamo, 71 pp.

Jonsell B; Karlsson T, 2000. Flora Nordica, volume 1: Lycopdiaceae - Polygonaceae. Stockholm, Sweden: Swedish Royal Academy of Sciences, 274 pp.

Keil P; Alberternst B, 1995. [English title not available]. (Reynoutria x bohemica Chrtek & Chrtkova im westlichen Ruhrgebiet.) Natur und Heimat (Münster), 55:85-88.

Kim JY; Park CW, 2000. Morphological and Chromosomal variation in Fallopia section Reynoutria (Polygonaceae) in Korea. Brittonia, 52:34-48.

Mandák B; Pysek P; Bímová K, 2004. History of the invasion and distribution of Reynoutria taxa in the Czech Republic: a hybrid spreading faster than its parents. Preslia, 76(1):15-64.

Meerts P; Tiebre MS, 2007. [English title not available]. (Fallopia x bohemica est beaucoup plus repandu que F. sachalinensis dans la region de Bruxelles.) Dumortiera, 92:22-24.

Padula M; Lastrucci L; Fiorini G; Galasso G; Zoccola A; Quilghini G, 2008. [English title not available]. (Prime segnalazioni di Reynoutria x bohemica Chrtek & Chrtkova (Polygonaceae) per l'Italia e analisi dell distribuzione del genere Reynoutria Houtt.) Atti Soc. It. Sci. nat. Museo civ. Stor. Nat Milano, 149:77-108.

Pashley CH; Bailey JP; Ferris C, 2003. Further evidence of the role of Dolgellau, Wales, in the production and dispersal of Japanese Knotweed s.l. In: Plant invasions: ecological threats and management solutions [ed. by Child, L.\Brock, J. H.\Brundu, G.\Prach, K.\Pyse?k, K.\Wade, P. M.\Williamson, M.]. Leiden, Netherlands: Backhuys Publishers, 197-211.

Preston CD; Pearman DA; Dines TD, 2002. New atlas of the British and Irish flora. An atlas of the vascular plants of Britain, Ireland, the Isle of Man and the Channel Islands. Oxford, UK: Oxford University Press, xi + 910 pp.

Pysek P; Brock JH; Bímová K; Mandák B; Jarosík V; Koukolíková I; Pergl J; Stepánek J, 2003. Vegetative regeneration in invasive Reynoutria (Polygonaceae) taxa: the determinant of invasibility at the genotype level. American Journal of Botany, 90(10):1487-1495.

Richards CL; Walls RL; Bailey JP; Parameswaran R; George T; Pigliucci M, 2008. Plasticity in salt tolerance traits allows for invasion of novel habitat by Japanese knotweed s. l. (Fallopia japonica and F. × bohemica, Polygonaceae). American Journal of Botany, 95(8):931-942. http://www.amjbot.org/

Schmitz J; Strank KJ, 1985. [English title not available]. (Die drei Reynoutria-Sippen (Polygonaceae) des Aachener Stadtwaldes.) Gött. Flor. Rundbr, 19:17-25.

Schnitzler A; Bailey J, 2008. Polymorphisme génétique et plasticité phénotypique deux atouts pour la dispersion des Renouées Asiatiques? Rev. Ecol, 63:209-217.

Schnitzler A; Bailey J; Hansen CN, 2007. Genotypic and phenotypic variation in a Fallopia x bohemica population in north-eastern France. In: Plant Invasions: Human perception, ecological impacts and management [ed. by Tokarska-Guzik, B. \Brock, J. H. \Brundu, G. \Child, L. \Daehler, C. C. \Pysek, P.]. Leiden, The Netherlands: Backhuys Publishers, 133-144.

Shaw RH; Child LE; Evans HC; Bailey JP, 2001. The Biological Control Programme for Japanese Knotweed (Fallopia japonica) in the UK and USA. A report of the Phase 1. Oxfordshire, UK: CABI, 105 pp.

Stace CA, 1989. New combinations in the British & Irish flora. Watsonia, 17:442-444.

Stace CA, 2001. New Flora of the British Isles. Cambridge, UK: Cambridge University Press, 1165 pp.

The Plant List, 2013. The Plant List: a working list of all plant species. Version 1.1. London, UK: Royal Botanic Gardens, Kew. http://www.theplantlist.org

Tiébré MS; Bizoux JP; Hardy OJ; Bailey JP; Mahy G, 2007. Hybridisation and morphogenetic variation in the invasive alien Fallopia (Polygonaceae) Complex in Belgium (Western Europe). American Journal of Botany, 94(11):1900-1910.

USDA-NRCS, 2008. The PLANTS Database. Baton Rouge, USA: National Plant Data Center. http://plants.usda.gov/

Zika PF; Jacobson AL, 2003. An overlooked hybrid Japanese knotweed (Polygonum cuspidatum × sachalinense; Polygonaceae) in North America. Rhodora, 105(922):143-152.

Distribution References

Adolphi K, 1999. First record of Fallopia x bohemica (Chrtek & Chrtkova) J. Bailey in the Netherlands. (De eerste vondst van Fallopia x bohemica (Chrtek and Chrtkova) J. Bailey in Nederland). Gorteria. 140-142.

Anon, 2000. Flora Nordica, volume 1: Lycopdiaceae - Polygonaceae. [ed. by Jonsell B, Karlsson T]. Stockholm, Sweden: Swedish Royal Academy of Sciences. 274 pp.

Anon, 2002. New atlas of the British and Irish flora. An atlas of the vascular plants of Britain, Ireland, the Isle of Man and the Channel Islands. [ed. by Preston C D, Pearman D A, Dines T D]. Oxford, UK: Oxford University Press. xi + 910 pp.

Bailey J P, Child L E, Conolly A P, 1996. A survey of the distribution of Fallopia x bohemica (Chrtek & Chrtkova) J. Bailey (Polygonaceae) in the British Isles. Watsonia. 187-198.

Bailey J P, Wisskirchen R, 2006. The distribution and origins of Fallopia x bohemica (Polygonaceae) in Europe. Nordic Journal of Botany. 173-200. DOI:10.1111/j.1756-1051.2004.tb00832.x

Bailey J, 2003. Japanese Knotweed s.l. at home and abroad. In: Plant invasions: ecological threats and management solutions. [ed. by Child L, Brock J H, Brundu G, Prach K, Pysĕk K, Wade P M, Williamson M]. Leiden, Netherlands: Backhuys Publishers. 183-196.

Balogh L, 1998. Exomorphological observations in support of the presence of the hybrid species Fallopia x bohemica in Hungary. (Külsõ alaktani megfigyelések a Fallopia x bohemica (Chrtek & Chrtková) J. Bailey (F. japonica x F. sachalinensis) hibridfaj magyarországi jelenlétének alátámasztásához). Kitaibelia (Debrecen). 3 (2), 255-256.

Balogh L, 2003. Species of the genus Fallopia sectio Reynoutria in Hungary - a land of the hybrid Fallopia x bohemica? In: Invasive Plants in Natural and Managed Systems: Linking Science and Management and 7th International Conference on the Ecology and Management of Alien Plant Invasions, Wyndham Bonaventure Resort, Ft. Lauderdale, Forida [Invasive Plants in Natural and Managed Systems: Linking Science and Management and 7th International Conference on the Ecology and Management of Alien Plant Invasions, Wyndham Bonaventure Resort, Ft. Lauderdale, Forida], 7.

Botta-Dukát Z, Balogh L, 2008. The most important invasive plants in Hungary. [ed. by Botta-Dukát Z, Balogh L]. Vácrátót, Hungary: HAS Institute of Ecology and Botany. 255 pp.

CABI, Undated. Compendium record. Wallingford, UK: CABI

CABI, Undated a. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI

CABI, Undated b. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI

Claeson S M, Bisson P A, 2013. Passive reestablishment of riparian vegetation following removal of invasive knotweed (Polygonum). Invasive Plant Science and Management. 6 (2), 208-218. DOI:10.1614/IPSM-D-12-00070.1

Conolly A P, 1998. Fallopia x bohemica: a new record from Australia? BSBI News. 88.

Conolly A P, 2001. Japanese Knotweeds in Australia. BSBI News. 74-75.

Eliás P, 2001. [English title not available]. (Dva naturalizovaNé neofyty [Sarcoca esculenta (van Houtte) skalicky a Fallopia x bohemica (Chrtek et Chrtková) J. P. Bailey v meste trnave). Acta Fac. Paed. Univ. Tyrnaviensis, Ser. B. 13-21.

EPPO, 2020. EPPO Global database. In: EPPO Global database, Paris, France: EPPO.

Fehér A, 1999. Reconstruction of the spread of invasive species of Fallopia genus in Pozitavska pahorkatina region. In: Invasions and invasive organisms, Vol. 2. [ed. by Elias P]. Bratislava-Nitra, Slovakia: SNK Scope & Sekos. 96-103.

Fojcik B, Tokarska-Guzik B, 2000. Reynoutria x bohemica (Polygonaceae): a new taxon to the Polish flora. (Reynoutria x bohemica (Polygonaceae): nowy takson we florze Polski). Fragm. Flor. Geobot. Polon. 63-71.

Fremsted E, Elven R, 1997. Foreign plants in Norway. The main Fallopia species. (Fremmede planter i Norge. De store Fallopia-artene). Blyttia. 3-14.

Gammon M A, Grimsby J L, Tsirelson D, Kesseli R, 2007. Molecular and morphological evidence reveals introgressions in swarms of the invasive taxa Fallopia japonica, F. sachalinensis, and F. × bohemica (Polygonaceae) in the United States. American Journal of Botany. 948-956.

Keil P, Alberternst B, 1995. Reynoutria x bohemica Chrtek & Chrtkova in the western Ruhr area. (Reynoutria x bohemica Chrtek & Chrtkova im westlichen Ruhrgebiet). Natur und Heimat (Münster). 85-88.

Mandák B, Pyšek P, Bímová K, 2004. History of the invasion and distribution of Reynoutria taxa in the Czech Republic: a hybrid spreading faster than its parents. Preslia. 76 (1), 15-64.

Meerts P, Tiebre M S, 2007. [English title not available]. (Fallopia x bohemica est beaucoup plus repandu que F. sachalinensis dans la region de Bruxelles). Dumortiera. 22-24.

Padula M, Lastrucci L, Fiorini G, Galasso G, Zoccola A, Quilghini G, 2008. [English title not available]. (Prime segnalazioni di Reynoutria x bohemica Chrtek & Chrtkova (Polygonaceae) per l'Italia e analisi dell distribuzione del genere Reynoutria Houtt.). Atti Soc. It. Sci. nat. Museo civ. Stor. Nat. Milano. 77-108.

Parepa M, Schaffner U, Bossdorf O, 2013. Help from under ground: soil biota facilitate knotweed invasion. Ecosphere. 4 (2), art31. DOI:10.1890/ES13-00011.1

Schnitzler A, Bailey J, 2008. [English title not available]. (Polymorphisme génétique et plasticité phénotypique deux atouts pour la dispersion des Renouées Asiatiques?). Rev. Ecol. 209-217.

Schnitzler A, Bailey J, Hansen C N, 2007. Genotypic and phenotypic variation in a Fallopia x bohemica population in north-eastern France. In: Plant Invasions: Human perception, ecological impacts and management. [ed. by Tokarska-Guzik B, Brock J H, Brundu G, Child L, Daehler C C, Pyšek P]. Leiden, Netherlands: Backhuys Publishers BV. 133-144.

Urgenson L S, Reichard S H, Halpern C B, 2012. Multiple competitive mechanisms underlie the effects of a strong invader on early- to late-seral tree seedlings. Journal of Ecology (Oxford). 100 (5), 1204-1215. DOI:10.1111/j.1365-2745.2012.01995.x

Urgenson L S, Reichard S H, Halpern C B, 2014. Habitat factors and species' traits influence riparian community recovery following removal of Bohemian knotweed (Polygonum × bohemicum). Northwest Science. 88 (3), 246-260. DOI:10.3955/046.088.0307

USA, USDA-NRCS, 2008. The PLANTS Database. In: The PLANTS Database, Baton Rouge, USA: National Plant Data Center. http://plants.usda.gov

Contributors

07/07/09 Original text by:

John Bailey, University of Leicester, UK

Distribution Maps