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Fallopia

Fallopia is a genus of approximately 12 species of flowering plants in the buckwheat family, Polygonaceae, comprising mostly herbaceous perennials or twining vines native to the temperate regions of the Northern Hemisphere. These plants are characterized by their fibrous or woody roots, often forming extensive rhizomes, and stems that range from erect and robust to scandent and climbing, typically glabrous or sparsely pubescent. Leaves are alternate, petiolate, and broadly ovate to triangular, with distinctive ocreae—papery sheaths at the nodes—that persist or decay with age. Flowers are small, bisexual or unisexual, arranged in racemes or panicles, with five tepals that become winged or accrescent in fruit, enclosing triangular achenes. The genus is distributed primarily across , , and parts of , with many species introduced and naturalized in other temperate zones worldwide, including , , and . Fallopia species often occupy disturbed habitats such as roadsides, riverbanks, and waste areas, thriving in a wide range of soils and exhibiting high ecological . occurs through and extensive clonal propagation via rhizomes and stem fragments, enabling rapid spread and resilience to environmental stresses. Several Fallopia species are notable for their ecological and economic impacts, particularly the invasive knotweed complex. Japanese knotweed (F. japonica), native to , is a highly aggressive that forms dense monocultures, displacing native flora, altering soil properties, and causing structural damage in introduced regions like and . Its hybrid with giant knotweed (F. sachalinensis), known as Bohemian knotweed (F. × bohemica), combines parental traits to enhance invasiveness, including increased fertility and adaptability. In contrast, species like silver lace vine (F. baldschuanica) are valued ornamentals for their profuse flowering and vigorous habit, though they can also become weedy in suitable climates. Other widespread species include black bindweed (F. convolvulus), an annual weed of arable lands, and climbing false buckwheat (F. scandens), native to .

Description

Morphology

Fallopia species are herbaceous vines or shrubs in the family, characterized by twining or scandent stems that can extend up to 10 meters or more in length, often featuring swollen nodes that contribute to their common name of knotweeds. These stems are typically glabrous or pubescent, woody at the base in perennial forms, and branched from near the ground, enabling both climbing and supportive growth. The leaves are alternate, , and petiolate, with blades ranging from ovate to lanceolate or broadly ovate to triangular, typically 5–15 cm long and 3–10 cm wide, with entire or slightly wavy margins. At each , distinctive ocreae—persistent or , chartaceous sheath-like structures—enclose the , a hallmark feature of the . These ocreae are often 5–20 mm long, with truncate to oblique apices, providing structural support and protection at the leaf bases. Flowers are small, bisexual or unisexual, measuring 2–4 in , with a campanulate that is pale green, white, or pinkish and glabrous. They occur in axillary racemes or panicles, 1–5 per ocreate fascicle, featuring five dimorphic tepals where the outer three are larger, often winged or keeled. The fruits are achenes, trigonous (3-gonous), 2–4 long, brown to black, glabrous, and typically enclosed or slightly exserted by the persistent . Root systems are fibrous or woody, with extensive rhizomes in many species that enable vegetative propagation; in some, such as , rhizomes penetrate up to 3 meters deep and extend horizontally up to 7 meters. Growth habits vary across the genus, with some species forming erect shrubs and others vigorous climbing vines, exemplified by the scandent versus more upright forms.

Reproduction

Fallopia species exhibit both sexual and asexual reproductive strategies, with the latter often dominating in invasive populations. Fallopia have flowers that are typically bisexual, though some are unisexual and the plants may be dioecious (e.g., F. japonica) or exhibit mixtures, with like Fallopia convolvulus bearing bisexual flowers. Flowers are typically small and arranged in racemose or paniculate inflorescences, pollinated primarily by , with wind playing a minor role in some cases. Sexual reproduction occurs through seed production following pollination, but seed viability is generally low, particularly in introduced ranges where hybridization between species like Fallopia japonica and Fallopia sachalinense produces sterile or low-fertility hybrids such as Fallopia × bohemica. For instance, pure F. japonica clones rarely produce viable seeds outside their native range due to the absence of compatible pollinators or genetic constraints, while F. convolvulus achieves higher seed output as an species. Flowering generally takes place from late summer to autumn, with seed maturation varying by species—F. convolvulus produces abundant seeds from July onward, whereas hybrids yield few viable ones. Asexual reproduction is the primary mode of propagation and spread, occurring through rhizome fragmentation and adventitious rooting at stem nodes, enabling regeneration from fragments as small as 1-2 cm or weighing 7 g. This clonal strategy maintains high genetic uniformity in invasive populations, often consisting of a single widespread , such as the dominant F. japonica clone in and , which facilitates rapid establishment and limits sexual recombination.

Taxonomy

Etymology and history

The genus Fallopia was established in 1763 by the French botanist Michel Adanson, who named it in honor of the Italian anatomist and botanist Gabriello Fallopio (1523–1562), known for his contributions to anatomy, including the description of the Fallopian tubes. Early classifications placed species now recognized in Fallopia under the broader genus Polygonum, as described by Carl Linnaeus in Species Plantarum (1753), due to shared morphological traits such as ocreae and inflorescences within the Polygonaceae family. Although Adanson's genus was proposed, it was not immediately adopted, and many taxa were alternatively assigned to Reynoutria or retained in Polygonum through the 18th and much of the 19th centuries. Botanical explorations in Asia during the 19th century, led by collectors such as the Russian botanist Carl Johann Maximowicz (1827–1891), who traveled extensively in Japan and surrounding regions from 1860 onward, significantly advanced the documentation of Fallopia diversity and highlighted distinctions in habit, chromosome numbers, and floral structures that supported segregating the genus from Polygonum. By the early , the invasive potential of certain Fallopia species became evident after their introduction to and as ornamental plants in the late , prompting studies on their ecological impacts and hybridization. Post-2000 taxonomic revisions, driven by molecular phylogenetic analyses using and sequence data, addressed longstanding debates over the genus's and confirmed its close monophyletic relationship with genera like Reynoutria and within , leading to refined boundaries and synonymy resolutions. Taxonomic treatments vary; for example, POWO recognizes 12 species and places knotweeds in Reynoutria, while FNA and WFO include them in Fallopia with 12–16 species.

Classification and phylogeny

Fallopia belongs to the family , subfamily Polygonoideae, and tribe Polygoneae. Within this tribe, the genus is most closely related to Reynoutria and , forming a distinct in subtribe Reynoutriinae. Phylogenetic analyses using nuclear (ITS) sequences and markers such as matK, rbcL, and rps16 have resolved the evolutionary relationships within this group. These studies indicate that Fallopia is not monophyletic, with of nested within it, rendering the current circumscription paraphyletic. Basal divergences occur among Asian , reflecting the genus's primary center of in temperate . The genus originated in temperate , where its species exhibit adaptations such as climbing habits via twining stems and extensive rhizomatous growth, facilitating colonization of diverse habitats. Key synapomorphies supporting a broader Fallopia including include persistent ocreae, articulate pedicels, and trigonous achenes. Fallopia was segregated from the polyphyletic s.l. based on morphological traits like the , often climbing habit and specific ocrea structure. Molecular evidence from ITS and chloroplast genes confirms its separation from s.s. and related groups, though ongoing debates concern the exact boundaries with and Reynoutria.

Species list

The genus Fallopia consists of approximately 12–16 accepted species (depending on taxonomic treatment), predominantly climbing annuals or perennials native to temperate regions of and , with the type species being . These species are characterized by twining stems, often with ocreae (sheath-like structures at nodes), and small flowers in racemes or panicles. Below is a catalog of accepted species, with brief distinguishing traits highlighted for key examples; some taxa are debated or treated in related genera in certain floras. Hybrids are listed separately.
  • Fallopia aubertii (L.Henry) Holub: A vigorous woody climber with panicles of small white flowers; often cultivated as an ornamental (sometimes synonymous with F. baldschuanica).
  • Fallopia baldschuanica (Regel) Holub: Known as Bukhara fleeceflower or Russian , this deciduous climber features arching stems up to 12 m long and abundant creamy-white flowers in long panicles during late summer; native to .
  • Fallopia cilinodis (Michx.) Holub: A climber with fringed ocreae and lanceolate leaves; found in eastern .
  • Fallopia convolvulus (L.) Á.Löve: The black bindweed, an annual twiner with arrow-shaped leaves and greenish-white flowers; a common weed in arable fields across and introduced elsewhere.
  • Fallopia cristata (Engelm. & A.Gray) Holub: Features crested ocreae and ovate leaves; native to central .
  • Fallopia cynanchoides (Hemsl.) Haraldson: A rare Asian species with slender stems and small, clustered flowers.
  • Fallopia dentatoalata (F.Schmidt) Holub: An annual climber with toothed leaves and dentate ocreae; distributed in .
  • Fallopia denticulata (C.C.Huang) Holub: Characterized by finely denticulate leaf margins; endemic to .
  • Fallopia dumetorum (L.) Holub: The thicket bindweed, a climbing annual with heart-shaped leaves and winged fruits; native to .
  • Fallopia japonica (Houtt.) Ronse Decr.: knotweed, a rhizomatous with hollow, bamboo-like stems up to 3 m tall and broad, heart-shaped leaves; notorious for its invasiveness and rapid spread via rhizomes (sometimes placed in Reynoutria).
  • Fallopia multiflora (Thunb.) Haraldson: Features reddish stems and dense panicles of small white flowers; used in in Asia (sometimes placed in Pleuropterus).
  • Fallopia scandens (L.) Holub: Climbing false , a with ovoid-orbicular leaves and greenish flowers; native to .
  • Fallopia sieboldii (Miq.) Miyabe: A species with slender, twining stems and inconspicuous flowers.
  • Fallopia tenuicaulis (Nakai) Holub: A herb native to , with slender stems and small flowers; known from temperate forests.
Notable hybrids include Fallopia × conollyana J.P.Bailey (a sterile hybrid with robust growth and variegated stems; rare) and Fallopia × convolvuloides (Brügger) Holub (a hybrid climber resembling F. convolvulus but with intermediate floral traits). Fallopia × bohemica (formerly in Fallopia but now often classified separately or in Reynoutria) is noted in phylogenetic studies but treated as a stable in some floras.

Synonyms and former classifications

The genus Fallopia Adans. has a complex nomenclatural history, with its name originally proposed in 1763 but largely unused until emended by Holub in 1971 to encompass certain twining and climbing species previously placed in the polyphyletic L. A key generic synonym is Bilderdykia Dumort. (1827), which included some of the same taxa and was later reduced to synonymy under Fallopia based on shared morphological traits such as ocrea structure and type. Many species now in Fallopia were reclassified from Polygonum or Reynoutria Houtt. during 20th-century revisions, driven by detailed studies of floral morphology, achene characteristics, and later DNA analyses that highlighted polyphyletic groupings in broader Polygonaceae classifications. For instance, Japanese knotweed, widely recognized as Fallopia japonica (Houtt.) Ronse Decr. (1988), was first described as Reynoutria japonica Houtt. (1777) and later as Polygonum cuspidatum Siebold & Zucc. (1846) or Polygonum sieboldii de Vriese ex Reinw. (1848); the transfer to Fallopia resolved its distinct perianth and twining habit from core Polygonum species. Similar reclassifications apply to other prominent taxa. Giant knotweed, F. sachalinensis (F. Schmidt) Ronse Decr. (1988), was originally Polygonum sachalinense F. Schmidt (1859), with the move to Fallopia supported by chromosome studies and molecular data indicating closer affinity to section Reynoutria within the genus. Russian vine, F. baldschuanica (Regel) Holub (1978), derives from Polygonum baldschuanicum Regel (1884), reclassified due to its scandent growth and ocreal features distinguishing it from herbaceous Polygonum. Hybrids have also contributed to nomenclatural shifts, often misclassified as distinct species owing to fertile intermediates. Bohemian knotweed, F. × bohemica (Chrtek & Chrtková) J.P. (2003), was initially named Reynoutria × bohemica Chrtek & Chrtková (1987) as the hybrid of F. japonica and F. sachalinensis; DNA evidence confirmed its hybrid origin and placement in Fallopia section Reynoutria. Some species formerly included in Fallopia have been excluded in recent phylogenies. For example, Fallopia cilinerve (Ohwi) Holub (1978) was transferred to Aconogonon Reichenb. as A. cilinerve (Ohwi) S. Sarkissian (2003) based on nuclear and chloroplast DNA sequences showing its alignment with the Aconogonon clade rather than Fallopia's twining group. Additional transfers from Fallopia or related segregates to Bistorta L. or Aconogonon occurred in the late 20th century, refining genus boundaries through integrative morphology and molecular systematics. The following table summarizes major species-level synonyms and reclassifications:
Current NameMajor Synonym(s)Original Author(s) and Year
F. japonicaReynoutria japonica, Polygonum cuspidatumHoutt. (1777), Siebold & Zucc. (1846)
F. sachalinensisPolygonum sachalinenseF. Schmidt (1859)
F. baldschuanicaPolygonum baldschuanicumRegel (1884)
F. × bohemicaReynoutria × bohemicaChrtek & Chrtková (1987)
F. aubertiiPolygonum aubertiiL'Hér. ex Loisel. (1828)
F. convolvulusPolygonum convolvulusL. (1753)
F. scandensPolygonum scandensL. (1753)
F. multifloraPolygonum multiflorumThunb. (1784)
These changes, particularly from the onward, reflect advances in resolving the tribe Polygoneae's phylogeny, prioritizing monophyletic genera over historical convenience.

Distribution and habitat

Native ranges

The genus Fallopia is predominantly native to temperate and subtropical regions of , including , , and , where the majority of its approximately 12 occur. Some species extend into and , reflecting a broader temperate distribution. For example, F. japonica is endemic to eastern Asia, specifically , , and eastern , with related taxa in . F. baldschuanica originates from , spanning the , western , , , , , and . In contrast, F. convolvulus, an annual species, is native across from (e.g., to the ) to northern and into parts of temperate (e.g., North-Central and ), with post-glacial expansions contributing to its wide range. In , F. scandens is native, occurring from and the (e.g., to ) eastward, often in moist woodlands and floodplains. Endemism patterns are particularly high in the Sino-Himalayan region, where several Fallopia species exhibit adaptations to diverse elevations from lowlands to montane zones up to 3000 m.

Introduced ranges

Fallopia species, particularly F. japonica, F. sachalinensis, and their hybrid F. × bohemica, have been introduced to regions outside their native East Asian ranges primarily through human-mediated activities in the 19th century. These plants were valued as ornamentals for their attractive foliage and were first brought to Europe around the mid-1800s; for instance, F. japonica was imported to the United Kingdom in 1849 via the Royal Botanic Gardens at Kew from specimens collected in Japan. In North America, introductions occurred slightly later, with F. japonica arriving in the United States by the late 19th century, documented as early as 1890 in coastal areas like Massachusetts. Today, Fallopia species are established across temperate regions globally, including much of (from the to central and eastern countries), (throughout the and southern ), , and . Smaller populations have been reported in parts of , such as , and southern , though these remain less extensive than in northern temperate zones. The genus thrives in disturbed environments like roadsides, riverbanks, and urban edges, where it forms dense monocultures. Initial introductions occurred mainly via the horticultural trade, with plants propagated and sold by nurseries for gardens and . Subsequent spread has been facilitated by accidental transport of rhizome fragments in soil from ships, construction materials, and flood debris, as well as deliberate movement for . Rhizomes enable rapid vegetative propagation, allowing the plants to colonize new areas along transportation corridors and waterways at rates exceeding 10 meters per year in favorable conditions. Establishment has been highly successful in human-altered habitats, where competition from native vegetation is reduced; for example, F. japonica now occupies an estimated area exceeding 10,000 km² across , particularly in riparian zones and abandoned industrial sites. is influencing distribution patterns, with models predicting potential southward expansion in the as warmer temperatures extend suitable habitats beyond current temperate limits.

Ecology

Growth habits

Fallopia species are primarily herbaceous or semi-woody s characterized by rapid vegetative growth and seasonal dieback, with new stems emerging from rhizomes or crowns in and aboveground parts senescing in autumn. In the , leaf-out typically occurs from mid- ( to May), driven by increasing temperatures, allowing shoots to reach heights of 2–3 m or more by midsummer under optimal conditions. Growth rates can be exceptionally fast, with Japanese () extending 5–10 cm per day during peak growth, enabling quick canopy formation and resource capture. By late summer, flowering commences in many species, followed by around to October, when leaves wilt and stems die back with the onset of , leaving persistent canes that provide winter structure. Rhizomes enter dormancy during winter, storing carbohydrates to fuel the next season's growth, a strategy that supports their across temperate climates. These plants exhibit broad tolerance to environmental stressors, thriving in diverse conditions despite a preference for moist, nitrogen-rich substrates. Fallopia species, particularly F. japonica, adapt to poor, compacted, or urban soils, including those with low fertility or high compaction, and can penetrate substrates up to 3 m deep with their rhizomes. tolerance spans acidic to neutral ranges, from 4.5 to 7.5, allowing establishment in naturally acidic or industrially disturbed sites. Regarding , they perform best in full sun but tolerate partial shade, with shade-intolerant forms like F. japonica competing effectively in canopy gaps by rapid upward growth or, in vining species such as silver lace vine (F. baldschuanica), by climbing supports to access sunlight. This flexibility in light regimes supports their of disturbed edges, riverbanks, and open woodlands. A key in Fallopia is high , enabling morphological adjustments to local conditions without genetic change. For instance, plants in shaded or colder environments often develop shorter statures and reduced leaf sizes to optimize capture and allocation, while those in open, nutrient-rich sites produce taller, more robust stems. This extends to architecture, where extension rates and depths vary by resistance and moisture, averaging 1.4–2 m horizontally in mature stands but capable of up to 4 m or more. Such responsiveness enhances survival and spread in heterogeneous habitats, from coastal marshes to inland areas, underscoring their as invasive perennials.

Interactions with other organisms

Fallopia species engage in pollination interactions primarily with , including bees and flies, which visit the small, greenish-white flowers to facilitate transfer. For instance, in , observations confirm visitation by various hymenopterans and dipterans, contributing to its reproductive success as an entomophilous species. Many Fallopia taxa, such as F. japonica, display strong , preventing self-fertilization and necessitating cross- by these pollinators for seed production. Herbivory on Fallopia is predominantly by feeders, with few native herbivores reported in introduced ranges. Deer, for example, browse leaves and stems of F. japonica, though damage levels are typically lower compared to native congeners, supporting the enemy release hypothesis for invasives. In its native Asian range, the psyllid Aphalara itadori acts as a herbivore, feeding on leaves and stems; this has been introduced in and as a classical biological to reduce biomass and growth of invasive Fallopia species like F. japonica. Fallopia plants are susceptible to several pathogens, including fungal rusts and eriophyid mites, though disease impacts are often minimal in invasive populations due to reduced pathogen pressure. The rust fungus polygoni-amphibii var. tovariae infects F. japonica in its native range, causing leaf spots and reduced vigor, and has been evaluated for biocontrol potential in invaded areas. Viral infections are less documented but occur, with low overall disease prevalence contributing to the resilience of invasives like F. japonica in non-native habitats. In terms of competitive interactions, Fallopia species exert allelopathic effects through root exudates containing that inhibit and of neighboring , facilitating dominance in mixed communities. This chemical interference, combined with rapid clonal , enables Fallopia to form dense monocultures that displace native and reduce local in riparian and disturbed habitats. Mutualistic relationships in Fallopia include associations with arbuscular mycorrhizal fungi (AMF), which colonize to enhance and uptake, particularly in nutrient-poor soils. Although AMF communities in invasive F. are species-poor compared to natives, these symbioses still support improved nutrient acquisition and in challenging environments.

Human interactions

Economic uses

Fallopia species have several economic applications, primarily in ornamental , traditional medicine, and limited food uses. Several species, notably Fallopia baldschuanica (synonym F. aubertii), are valued as ornamental climbers for their rapid growth and attractive foliage and flowers. Introduced to and in the late , these vines provide quick coverage for fences, arbors, trellises, and walls, often reaching 10-15 meters in a single season. The roots of Fallopia multiflora, known as He Shou Wu in , are harvested for their purported health benefits, including anti-aging effects, liver tonification, and kidney support. These properties are attributed to bioactive stilbenes such as 2,3,5,4'-tetrahydroxystilbene-2-O-β-D-glucoside and glucosides, which exhibit and activities. Commercial cultivation of F. multiflora is primarily concentrated in and other parts of for pharmaceutical and herbal supplement production. Young shoots of are consumed in Japan as "itadori," prepared similarly to or in savory or sweet dishes due to their tart flavor. Historically in , the plant has served as for , including and goats, providing nutritious from its shoots and foliage.

Invasiveness and control

Several within the Fallopia, particularly F. japonica (Japanese ), F. sachalinensis (giant ), and the F. × bohemica (Bohemian ), are recognized as highly invasive outside their native ranges and are listed as noxious weeds in multiple regions. , these are designated as noxious weeds under federal and state regulations, with restrictions on their sale, transport, and cultivation in states such as and . In the , they appear on the list of of Union concern under Regulation (EU) No 1143/2014, prohibiting their trade and intentional introduction. In the , F. japonica is specifically protected under Schedule 9 of the , making it illegal to plant or cause the to grow in the wild. Similar protections extend to F. sachalinensis and F. × bohemica due to their invasive potential and hybridization risks. These knotweeds exert significant ecological and economic impacts in invaded areas. They reduce native , with significant declines in density, herbaceous cover, and populations in affected riparian zones. Structurally, the expansive systems can infiltrate cracks in buildings, pavements, and flood defenses, leading to costly repairs. Economically, management and damage mitigation for F. japonica alone cost the approximately £246.5 million annually as of 2023, encompassing property devaluation, remediation, and repairs. Spread primarily occurs vegetatively through rhizome fragments, which can regenerate into new from pieces as small as 0.7 grams containing a ; these fragments are dispersed by machinery, movement, flooding, or activities. s remain viable in for over 20 years, enabling long-term persistence and re-infestation even after apparent removal. Control strategies for Fallopia species integrate mechanical, chemical, and biological approaches, often requiring multi-year efforts for eradication. Mechanical methods include excavation of s to depths of at least 2 meters or repeated cutting to exhaust underground reserves, though these risk fragment dispersal if not managed carefully. Chemical control typically involves foliar or stem-injection applications of , with eradication timelines of 2–5 years depending on size and follow-up treatments; rates of 4–6 liters per in late summer or autumn maximize efficacy. Biological agents, such as the psyllid Aphalara itadori (previously trialed as a host-specific ), have been released in the UK since 2010 and in the since 2020 to suppress through feeding , though and remain under . Emerging non-chemical methods as of 2025 include thermo-electric treatments applying 3000–5000 volts to destroy s. Prevention focuses on regulatory measures and habitat restoration. Bans on the sale, transport, and planting of Fallopia species have been implemented since the 2010s in countries including the , member states, and several states, alongside requirements for in property transactions. Post-removal sites are restored by planting competitive native species to prevent re-establishment and support recovery. As of 2025, refined (IPM) protocols combining timing with biological releases offer sustainable, site-specific control.