European and Mediterranean Plant Protection Organization Organisation Européenne et Méditerranéenne pour la Protection des Plantes 07-13386 DRAFT Data sheets on invasive alien plants Fiches informatives sur les plantes exotiques envahissantes Polygonum perfoliatum Identity Scientific name: Polygonum perfoliatum (L.) L. Synonyms: Ampelogonum perfoliatum (L.) Roberty & Vautier, Ampelygonum perfoliatum (L.) Roberty & Vautier, Chylocalyx perfoliatus (L.) Hassk. Ex Miq., Echinocaulon perfoliatum (L.) Meisn. Ex Hassk, Fagopyrum perfoliatum (L.) Raf., Persicaria perfoliata (L.) H. Gross, Tracaulon perfoliatum (L.) Greene, Truellum perfoliatum (L.) Soják Taxonomic position: Polygonaceae Protologue: Systema naturae edition 10 p. 1006, published 1759 (The International Plant Names Index website) Common names: Asiatic tearthumb, devil's-tail tearthumb, mile-a-minute-vine, mile-a-minuteweed, minuteweed, tearthumb (English), knöterich, durchwachsener (German), ishimikawa (Japanese), gang ban gui (Chinese). EPPO code: POLPF Phytosanitary categorization: EPPO A2/352 Geographical distribution EPPO Region: Russia (Siberia, native), Turkey (alien, status unknown). Asia (native): Temperate: China (Anhui, Fujian, possibly eastern Gansu, Guangdong, Guangxi, Guizhou, Hainan, Hebei, Heilongjiang, Henan, Hubei, Hunan, Jiangsu, Jiangxi, Jilin, Liaoning, Neimenggu (Inner Mongolia, eastern part), southern Shaanxi, Shandong, Sichuan, Xizhang (Tibet, unconfirmed), Yunnan, Zhejiang), Japan, Republic of Korea, Russian Federation (Far East), Taiwan. Tropical: Bangladesh, Bhutan, India, Indonesia, Nepal, Thailand, Vietnam, Malaysia, Myanmar (Burma), Philippines. Oceania: Papua New Guinea. North America (alien): USA (Connecticut, Delaware, District of Columbia, Maryland, New Jersey, New York, North-Carolina, Ohio, Oregon, Pennsylvania, Rhode Island, Virginia, West Virginia, Wisconsin). Note: the plant has been eradicated from New Zealand. There is also a single record for Canada (British Columbia), but no additional records since 1954. In Turkey, the plant is present on the northern face of the Kaçkar range of mountains in North Eastern Turkey, in Rize, district Ardeşen, near Firtina Deresi (Black Sea region). 1 In the USA, the plant has been recorded in Mississippi, but herbaria curatorial staff in Mississippi stated that publications indicating its occurrence in this State are erroneous. History of introduction and spread The first record of P. perfoliatum in North America is from Portland, Oregon (1890) where it was believed to have arrived in ship ballasts (Stahl, 2002). This population was apparently short-lived as the plant disappeared (Hickman and Hickman, 1977). In 1937, the plant was found in Prince George County (Maryland), in the Glenn Dale Introduction Garden from a site planted with Meliosa seeds from China. According to Moul (1948) Dr. Joseph Ewan of the USDA Plant Industry Station in Beltsville, Maryland, writes that "Polygonum perfoliatum did appear at the Glenn Dale Introduction Garden, Maryland, at a site where Meliosa seed from Nanking, China, had been planted. The Meliosa failed to grow but the Polygonum perfoliatum appeared”. This population was eradicated by routine weeding practices (Moul, 1948). The genus Meliosa does not exist and may have been misspelled in which case it should read “Meliosma” (EPPO Panel on Invasive Alien Species). The discovery of plants from the Gable Nursery in Stewartstown, Pennsylvania, in 1946 is considered the first establishment site in the United States (Moul, 1948). Moul (1948) talked with someone at the Gable Nursery in Stewartstown, York County, Pennsylvania, and states "the plant made its first appearance in the nursery about ten years ago, when some holly seeds (Ilex spp.) sent from Japan were planted and it came up with the holly". On the other hand, according to Okay (1999) seeds of P. perfoliatum were probably unintentionally transported to this location in rhododendron nursery stock imported from Eastern Asia in the 1930s (Okay, 1999). Whatever the pathway of introduction (Ilex seeds or rhododendron stock), the owner of the nursery was impressed by the beauty of the fruit and reproduced it (Moul, 1948); subsequent efforts to eradicate it failed. It was also speculated that P. perfoliatum could have been introduced into the Gable Nursery in Stewartstown, Pennsylvania with rhododendron stock from the Glenn Dale Introduction Garden (Reed, 1979). Hickman and Hickman (1977) also reported its establishment on the campus of Swarthmore College in Southern Pennsylvania and believed that it was spread primarily by attachment to rhododendron plants purchased from the Gable Nursery. The distribution of P. perfoliatum has radiated from the York County site into neighbouring states. In the past 55 years, the range for this plant in the United States has extended up to approximately 500 km in several directions from the York County site (Mountain, 1995, Okay 1997). It was found in Connecticut in 2000, and although its mean of introduction is unclear, it had apparently spread for a few years before it was reported. It has also been reported in Rhode Island, where it was found growing in a rhododendron/azalea nursery (IPANE Website). Morphology Plant type Polygonum perfoliatum is an herbaceous trailing vine. It is an annual in temperate climates (Mountain, 1989), but could behave as a perennial in tropical climates such as in Florida (Olivier, 1994). Description P. perfoliatum has a stem that can grow up to 6 m in length and as much as 15 cm per day. Roots are few in number, fibrous, weak and do not penetrate the soil deeply (Moul, 1948). A characteristic cup-shaped ocrea (or bract) surrounds the stem at the base of the petiole; those of the upper leaves are conspicuously expanded. Stems, petioles and veins on the underside of the foliage are armed with curved, retrorse barbs. The petioles are long and perfoliate. The thin, jointed, highly branched stems are green to reddish-green in colour. The leaves are pale green, thin and glabrous. They are 2-8 cm wide and shaped like an Euclidian equilateral (equal-sided) triangle and alternate along the delicate stems (this leaf shape gives it one of its common names, devil's tail). 2 Two to four flowers are borne on racemes 1-2 cm in length that emerge from the ocrea. These inconspicuous white or pale red flowers, which become blue, at fruiting measure 3-5 mm. They are deep, 5 lobed and bear 3 styles and 8 stamens, and often remain closed. Green, berry-like fruits, 5 mm in diameter, are produced in June (in New England, USA) and become a pale, metallic blue colour as they ripen. Each fruit contains a shiny, black or reddish-black, nearly round achene 2 mm in diameter which weighs 27.4 mg on average (Van Clef and Stiles, 2001). P. perfoliatum produces fruit continuously until the first frost, when the plant begins to die back. Dead plants in winter are reddish-brown to tan in colour, often forming brittle mats (IPANE Website). Similarities to other species Polygonum arifolium, P. sagittatum and P. scandens var. cristatum have ocreas, but they are not entirely perfoliate (surrounding the stems) such as P. perfoliatum. Calystegia sepium has no ocrea. Fruits are berry-like in P. perfoliatum, while they are lenticular achenes in P. arifolium, P. arifolium and P. scandens var. cristatum and a capsule in Calystegia sepium (IPANE Website). Biology and ecology General P. perfoliatum is a very tender annual, withering with a slight frost, and reproducing successfully until the first frost. Germination occurs in early to mid-March the following year and continues through April in regions of North-Eastern United States (Kumar and DiTommaso, 2005). The plant reproduces sexually; vegetative propagation has never been reported. It is primarily a self-pollinating plant (supported by its inconspicuous, closed flowers and lack of a detectable scent), with occasional out crossing. Fruits and viable seeds are produced without assistance from pollinators. The self-compatibility of this species contributes to its successful dispersal because single plants once established in a new habitat can produce new populations without the need for cross-pollination from neighbouring conspecies (Okay, 1997). Until the first frost, the plant can grow up to 6 m long (15 cm per day), bearing about 50-100 seeds (Stahl, 2002). The species may produce a small peak of production of fruits in July (which may safeguard production in years of severe drought) and a large peak of production ripening from mid-September to early November (in regions of North-Eastern United States), which coincides with major bird migration. Seed production can reach 66 seeds/m² during the peak of production. Seed dormancy and germination of the plant is essential for predicting its potential range of distribution. Seed germination after 1, 2, and 3 years buried in forest soils was 75%, 25%, and 33%, respectively, which indicates that the species forms a long-term seed bank (Van Clef and Stiles, 2001). It senesces after the first frost in late October to early November (in regions of North-Eastern United States). Van Clef (2001) found that seed longevity, seedling growth advantages provided by larger seeds, and bird dispersal contribute most to increased invasiveness of the species in North America. Habitats In its native range, P. perfoliatum occurs in moist areas at elevations of 80-2300 m. It can be found along rivers and roadsides in Eastern China; along valley streams and in thickets in Northern China; mountain thickets, forest margins and stream banks at elevations of 200-1300 m in the Qinling Mountains and Loess Plateau areas of North-Western China; ditches, stream banks and wasteland in Central and Southern China; hillside thickets at 2100 m in Southern Tibet; and grassy slopes, forest margins, roadsides and river banks at 500-2100 m in Yunnan, South-Western China (Zheng et al., 2005). In its alien range, P. perfoliatum invades a wide range of habitats, mainly open and disturbed ones; edges of pastures, edges of woods, early successional forests, abandoned fields, roadsides, railroad, nurseries, wood-piles, clearings and ditches. It thrives where forests are clear-cut (Oliver, 1996). The 3 species has been reported to grow in areas where other invasive plants such as Pueraria lobata have been killed by herbicides (Wu et al., 2002). It is also found in freshwater habitats such as stream banks and moist thickets. Environmental requirements Although light and soil moisture are preferable for successful colonization by this species, it is tolerant to shade and dryness. It will tolerate shade for a part of the day but needs a good percentage (63-100%) of the available light. Weak growth can occur in low light (16% of ambient) under greenhouse conditions (Van Clef, 2001). It can reach areas of higher light intensity by attaching to and climbing over other plants with its recurved barbs and has been observed climbing the trunks of trees up to 4 m high. It can survive in areas with relatively low soil moisture, but demonstrates a preference for high soil moisture. P. perfoliatum generally grows in areas with an abundance of leaf litter on the soil surface (Okay, 1999), but has also been found in extremely wet environments with poor soil structure. Johnson (1996) reported that seeds of P. perfoliatum germinate over a wide range of temperatures from 5°C to 20°C and require at least 6 weeks of moist stratification at 2°C. Okay (1997) determined that a stratification period of at least 8 weeks at 5°C was required for breaking seed dormancy. At pH 3.5, however, stratification was not required, although germination levels were significantly lower (16.7%) than at pH 7.5 with cold treatment (46.7%). Although stratification requirements for germination make P. perfoliatum an unlikely candidate to cause serious problems in Florida (Okay, 1997), germination could occur under acidic conditions in cypress swamps of Florida where the pH can range from 2.8 to 3.9 (Kumar and DiTommaso, 2005). The ability of the plant seeds to germinate at relatively cold temperatures provides it a competitive advantage over other annual and perennial weeds that germinate at higher soil temperatures later in the spring (Kumar and DiTommaso, 2005). Climatic and vegetational categorization P. perfoliatum is considered a temperate species with subtropical tendencies and therefore has the potential to invade those portions of the contiguous United States that have the appropriate climate to provide a minimal eight week cold vernalization period. It is considered that its current geographical distribution in the USA only represents 20% of its possible range (Okay 1999). The plant is hardy in zones 6 and 7, meaning that average annual minimum temperature are between -18°C and -12°C (Okay, 1997). Pathways for movement Many seeds fall to the ground beneath parent plants, which may allow local site dominance over time for this annual species (Van Clef, 2001). The small, palatable, bright and attractive fruits are welladapted for dispersal by birds (Reifner, 1982). The seeds remain viable after passage through avian digestive tracts (Okay, 1997). Birds are probably the primary long-distance dispersal agents of P. perfoliatum (Okay, 1999). Transport of seeds over short distances by ant species has been observed because they are attracted by the elaiosome at the top of the seeds which is a source of food (Okay, 1999). These seed-carrying ants may play an important role in the survival and germination of the seeds of the plant. Other animals observed eating P. perfoliatum fruits are chipmunks, squirrel and deer, and passage through their gut is not thought to alter germinability of the seeds (Toni DiTommaso, pers. com.). Water is also an important mode of long distance dispersal for P. perfoliatum as the fruits are buoyant and well adapted for water-borne dispersal (Reed, 1979). The long vines frequently hang over waterways, allowing fruits that detach to be carried away in the water current. Its fruits can remain buoyant for 7-9 days. During storm events the potential spread of this plant is greatly increased throughout watersheds (Okay, 1999). The drupes may also be spread by railroad cars, mowing equipment or shoes (Wikimanual of Gardening Website; Cusick and Ort, 1987). 4 The plant’s seeds may be planted inadvertently with other plants while being transported in nursery stock (e.g. Rhododendron stock) (IPANE, 2001). The plant is believed to have been introduced into Staten Island (New York State) around 2000 with nursery material from New Jersey, Delaware, Maryland or Pennsylvania. Uses and benefits The plant has “no apparent redeeming economic or social value” (Stevens, 1994). Nevertheless, in its native range, it is used as an herbal medicine (Yang and Kim, 1993). The fruits are edible and of 40 wild fruits studied in Nepal, it had the highest sodium and potassium content (Bajracharya, 1980). Impact Effects on plants Because P. perfoliatum can smother tree seedlings, this weed has a negative effect on forest regeneration and commercial forest areas (Christmas tree farms). In revegetation areas of Virginia, additional costs for site preparation, weed management, and labour to replant tree saplings overtopped by this weed have been estimated to range from $60-500 per ha (Stanosz and Jackson 1991). At harvested sites in Pennsylvania, Mc Cormick and Hartwig (1995) observed mortality of regenerating tree saplings including Pinus taeda because of the dense canopy formed by this weed. P. perfoliatum is thought to have the potential to be a problem to nurseries, orchards and to the ornamental shrub industry that are not regularly tilled as a cultivation practice (Lehtonen, 1994). Most nurseries in the mid-Atlantic U.S.A are aware of P. perfoliatum, where it is considered 'an emerging threat'. Its range is limited, thus most of the industry outside its range is unaware of this species and does not perceive it as widespread or serious. This weed is not common in container nurseries and may only become a concern in nurseries under very low management systems (0-1 maintenance operations per year), such as Christmas tree farms (R Bates, pers. com., 2007). Infestation of apple trees can cause some defoliation (Moul, 1948). Even if P. perfoliatum has been observed growing on the edges of corn and soybean fields in Delaware (Lehtonen, 1994), it does not appear to be a threat in agricultural production probably due to continuous tillage and herbicide use (Kumar and DiTommaso, 2005). Nevertheless, it can easily become a pest to gardeners and landscapers, destroying ornamental plants and landscaped yards (Invasive Alien plants of Virginia Website). It is considered an important weed especially in less intensively managed agricultural lands in Eastern Asia by Kasahara (1954), but Mountain (1989) suggests that in Korea and Japan, plant scientists consider it a weed with little or no agricultural significance. The species is a widely distributed native plant in Japan, but not listed as invasive either in orchards or in nurseries (S Kurokawa, pers. com. 2007). In Siberia, where it is also native, the plant is not recorded as being a threat (A. Orlinski, pers. com., 2007). Environmental and social impact P. perfoliatum is a threat to ecosystems as it has the ability to outgrow other species (Oliver, 1996). It is known to grow rapidly, scrambling over shrubs and other vegetation, blocking the foliage of covered plants from available light and reducing their ability to photosynthesize, which stresses and weakens them, the shade killing grasses and wildflowers (Stevens, 1994). IPANE (2001) states that trees and other native plants could suffer mechanical damage due to the weight of this plant. Okay (1997) observed decreases in native plant species diversity in areas colonized by P. perfoliatum. Lehtonen (1994) reports that the invasive Lonicera japonica has been displaced by P. perfoliatum and that Sambucus canadensis and Rubus spp. were overgrown and killed by the competition. Replacement of existing vegetation will deprive native animals of habitat and food (Olivier, 1994). 5 Dense thickets of the sharp-spined plants are unpleasant for people (Binion, 2005) and can restrict the movement of wildlife in natural areas (Okay, 1997). Summary of invasiveness The plant has negative economic impacts by smothering tree seedlings in tree farms and forest and by outcompeting native vegetation in freshwater ecosystems. Control This species can form a long-term seed bank, which must be suppressed for any management technique used (ISSG Database website, Van Clef, pers. comm., 2004). Cultural and mechanical control Control efforts for this plant should focus on eliminating or reducing seed output (Olivier & Coile, 1994), limiting the creation of gaps or openings in existing vegetation (Okay, 1999) and to maintain vegetative community stability. Cultural methods can be utilized to create conditions which are not favourable to the establishment of P. perfoliatum. Maintaining broad vegetative buffers along streams and forest edges should help to shade out and prevent establishment of this weed. This should also help to reduce the dispersal of fruits by water. Physical removal is not recommended after fruit production which begins in July (in the mid-Atlantic region of the USA) because it will help dispersal of the species (Van Clef, pers. comm 2004). Hand pulling of seedlings is best done before the recurved barbs on the stem and leaves harden but may be done afterwards with the help of thick gloves. The vines can be balled up in piles that can be left to dehydrate for several days before disposal. The site must be rechecked at frequent intervals, and removal of new plants should continue until the seed germination period is complete, roughly early April until early July in the mid-Atlantic region of the United States. Repeated mowing or trimming of plants should prevent the plants from flowering and thus reduce or eliminate fruit and seed production. Heavy deposits of dead and decaying plant matter should be cleared to reduce the mulch available to seeds (Mountain, 1989). Old woodpiles should be removed or, where appropriate, eliminated using fire (Olivier, 1996). Chemical control Studies have shown that pre-emergence applications of herbicide are most effective in controlling P. perfoliatum with the herbicides sulfometuron methyl, hexazinone, imazapyr, atrazine, and imazethapyr being the most effective. glyphosate and imazapyr are best for post-emergence control (McCormick & Hartwig, 1995). Herbicidal soap helps burning back foliage of P. perfoliatum. Because these products do not have the systemic ability of herbicides like glyphosate (i.e., these products do not travel to the roots), they must be reapplied all season long to any regrowth. glyphosate formulations adapted to upland areas or to wetlands, applied at a low rate will probably be effective in killing the weed. Biological control A biological control program is currently undertaken in the USA (University of Delaware Website) and a number of potential biological control agents for P. perfoliatum have been identified in China. The weevil Rhinoncomimus latipes (Coleoptera: Curculionidae) is regarded as a very promising agent (J Hough-Goldstein, pers. com., 2007). It is host specific for P. perfoliatum (Ding et al. 2004). and should have minimal potential non-target effects if released in the USA (Colpetzer et al. 2004a, b). An environmental assessment has been performed for the field release of R. latipes (USDA, 2004). Regarding the release of R. latipes, this agent appears to have established and over wintered wherever it has been released, and has built up to high numbers and apparently caused plant mortality in a few places so far. It is still early to assess its effectiveness (since it has only been in the field since late 2004). 6 Other possibilities included the oligophagous leaf beetles Smaragdina nigrifons (Coleoptera: Eumolpidae), Gallerucida bifasciata (Coleoptera: Chrysomelidae) and Galerucella placida (Coleoptera: Chrysomelidae), and the geometrid moth Timandra griseata (Lepidoptera: Geometridiae). These all had an impact on the growth and reproduction of P. perfoliatum. The bug Cletus schmidti (Hemiptera: Coreidae) and the sawfly Allantus nigrocaerulleus (Hymenoptera: Tenthredinidae) were recommended for further trials of host specificity (Ding et al. 2004, Zheng et al., 2005). Timandra griseata was found to have too broad a host range to be considered for release in the USA., as it also fed on Fagopyrum esculentum and Fagopyrum tartaricum. Homorosoma chinensis (Coleoptera: Curculionidae) is possibly host-specific to P. perfoliatum, but further trials are needed with other potential host plants before its release in the USA. could be recommended (Price et al., 2003). Possibilities for eradication P. perfoliatum was eliminated or did not establish permanent populations in both sites of first introduction in the USA (Portland, Oregon, 1890 and Beltsville, Maryland, 1937) and in Canada (Pitt, Meadows, British Columbia, 1954). However, efforts to eradicate it were not successful when it was introduced in York County, Pennsylvania and spread with Rhododendron stock. In New Zealand, Lee et al. (2001) stated that P. perfoliatum has been eradicated from Auckland. Regulatory status The plant is listed as a noxious plant by the following states in USA: Alabama, Connecticut, Massachusetts, North Carolina, Ohio, Pennsylvania, South Carolina (USDA Website). In New Zealand, it is also listed as a Total Control Pest Plant throughout the Auckland region (Auckland Regional Council, 2006). Acknowledgements The following people provided information for this data sheet: E Snyder, Canadian Food Inspection Agency, Canada (snydere@inspection.gc.ca), J HoughGoldstein, University of Delaware, USA (jhough@udel.edu), RC Reardon, USDAFS, USA (rreardon@fs.fed.us), R Bates, Department of Horticulture, Penn State University, USA (rmb30@psu.edu), S Kurokawa, National Agriculture and Food Research Organization, Japan (shunji@affrc.go.jp), A DiTommaso, Department of Crop & Soil Sciences, Cornell University, USA (ad97@cornell.edu). 7 References Auckland Regional Council (2006) Proposed Auckland Regional Pest Management Strategy 20072012. Auckland regional Council. 182 p. http://www.arc.govt.nz/shadomx/apps/fms/fmsdownload.cfm?file_uuid=A53F0E5B-BCD4-1A249B45-1AAA5D56839E&siteName=arc Bajracharya D (1980) Nutritive values of Nepalese edible wild fruits. 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