Clematis terniflora
(sweet autumn clematis)

Pictures

Picture	Title	Caption	Copyright
Title Habit Caption Clematis terniflora (sweet autumn clematis); invasive habit, showing the white flowers. The plant can climb woody vegetation to heights of >30m. USA. Copyright ©Leslie J. Mehrhoff/University of Connecticut/Bugwood.org - CC BY 3.0 US	Habit	Clematis terniflora (sweet autumn clematis); invasive habit, showing the white flowers. The plant can climb woody vegetation to heights of >30m. USA.	©Leslie J. Mehrhoff/University of Connecticut/Bugwood.org - CC BY 3.0 US
Title Habit Caption Clematis terniflora (sweet autumn clematis); flowering habit. USA. September 2003. Copyright ©Leslie J. Mehrhoff/University of Connecticut/Bugwood.org - CC BY 3.0 US	Habit	Clematis terniflora (sweet autumn clematis); flowering habit. USA. September 2003.	©Leslie J. Mehrhoff/University of Connecticut/Bugwood.org - CC BY 3.0 US
Title Habit Caption Clematis terniflora (sweet autumn clematis); flower and leaves. Omihachiman, Shiga prefecture, Japan. October 2013. Copyright ©Alpsdake/via wikipedia - CC BY-SA 4.0	Habit	Clematis terniflora (sweet autumn clematis); flower and leaves. Omihachiman, Shiga prefecture, Japan. October 2013.	©Alpsdake/via wikipedia - CC BY-SA 4.0
Title Flowering habit Caption Clematis terniflora (sweet autumn clematis); flowering habit. Kasugai, Aichi prefecture, Japan. August 2016. Copyright ©Alpsdake/via wikipedia - CC BY-SA 4.0	Flowering habit	Clematis terniflora (sweet autumn clematis); flowering habit. Kasugai, Aichi prefecture, Japan. August 2016.	©Alpsdake/via wikipedia - CC BY-SA 4.0
Title Flower Caption Clematis terniflora (sweet autumn clematis); close view of a flower. Mount Tado, Yoro Mountains, Kuwana, Mie prefecture, Japan. October 2013. Copyright ©Alpsdake/via wikipedia - CC BY-SA 4.0	Flower	Clematis terniflora (sweet autumn clematis); close view of a flower. Mount Tado, Yoro Mountains, Kuwana, Mie prefecture, Japan. October 2013.	©Alpsdake/via wikipedia - CC BY-SA 4.0
Title Habit Caption Clematis terniflora (sweet autumn clematis); habit, showing leaves. Mount Yokoyama, Nagahama, Shiga prefecture, Japan. August 2017. Copyright ©Alpsdake/via wikipedia - CC BY-SA 4.0	Habit	Clematis terniflora (sweet autumn clematis); habit, showing leaves. Mount Yokoyama, Nagahama, Shiga prefecture, Japan. August 2017.	©Alpsdake/via wikipedia - CC BY-SA 4.0
Title Fruits Caption Clematis terniflora (sweet autumn clematis); ripening fruits. USA. October 2006. Copyright ©Leslie J. Mehrhoff/University of Connecticut/Bugwood.org - CC BY 3.0 US	Fruits	Clematis terniflora (sweet autumn clematis); ripening fruits. USA. October 2006.	©Leslie J. Mehrhoff/University of Connecticut/Bugwood.org - CC BY 3.0 US
Title Fruits Caption Clematis terniflora (sweet autumn clematis); fruits. USA. October 2005. Copyright ©Leslie J. Mehrhoff/University of Connecticut/Bugwood.org - CC BY 3.0 US	Fruits	Clematis terniflora (sweet autumn clematis); fruits. USA. October 2005.	©Leslie J. Mehrhoff/University of Connecticut/Bugwood.org - CC BY 3.0 US
Title Young plants Caption Clematis terniflora (sweet autumn clematis); young plants. USA. December 2009. Copyright ©Leslie J. Mehrhoff/University of Connecticut/Bugwood.org - CC BY 3.0 US	Young plants	Clematis terniflora (sweet autumn clematis); young plants. USA. December 2009.	©Leslie J. Mehrhoff/University of Connecticut/Bugwood.org - CC BY 3.0 US

Identity

Preferred Scientific Name

• Clematis terniflora DC.

Preferred Common Name

• sweet autumn clematis

Other Scientific Names

• Clematis dioscoreifolia H. Lév. & Vaniot
• Clematis dioscoreifolia var. robusta (Carrière)
• Clematis flammula var. robusta (Carrière)
• Clematis maximowicziana Franch. & Sav.
• Clematis paniculata Thunb.
• Clematis recta subsp. paniculata (Thunb.) Kuntze
• Clematis terniflora var. robusta (Carrière) Tamura

International Common Names

• English: fall clematis; Japanese clematis; leatherleaf clematis; sweet autumn virginsbower; yam-leaved clematis
• French: clématite à panicules
• Chinese: yuan zhui tie xian lian
• German: rispenblütige waldrebe

Local Common Names

• Japan: sennin-sô
• Sweden: stor vippklematis

Summary of Invasiveness

Clematis terniflora is a perennial woody vine, native to Asia and introduced to North America as an ornamental. It can self-seed, and has escaped cultivation and naturalized in many parts of the USA. It is reported to be invasive in a number of eastern states. It grows in forest margins, scrub, grassy areas on hills and slopes, and in disturbed areas such as roadsides, thickets and urban green spaces. Seeds are widely dispersed by wind. It grows rapidly, forming dense clumps that outcompete and cover young native trees, shrubs and herbs at ground level and suppress seed germination. It can also climb to nearly 10 m, smothering trees and pulling down telephone poles. C. terniflora is difficult to control. Removal by hand can help encourage the growth of native species, but is unlikely to eliminate C. terniflora entirely due to root re-sprouting and prolific seed production. Some herbicides have proven effective in controlling the spread of this species; however, repeated applications are necessary.

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Plantae
•         Phylum: Spermatophyta
•             Subphylum: Angiospermae
•                 Class: Dicotyledonae
•                     Order: Ranunculales
•                         Family: Ranunculaceae
•                             Genus: Clematis
•                                 Species: Clematis terniflora

Notes on Taxonomy and Nomenclature

The name C. paniculata was incorrectly used for this species by Thunberg in 1794 (Flora of North America Editorial Committee, 2018). C. paniculata is a separate species, native to New Zealand (Missouri Botanical Garden, 2018).

Some authors have recognized two or more varieties of this species, correlated with their distribution in Asia. However these have now been reduced to synonymy (Flora of North America Editorial Committee, 2018).

Description

C. terniflora is a climbing, semi-evergreen, woody vine (Swearingen and Bargeron, 2016). Stems are 3-6 m, climbing with tendril-like petioles and leaf rachises (Flora of North America Editorial Committee, 2018). Leaves are shiny, green and leathery (Missouri Botanical Garden, 2018), and are opposite, compound, with 3-5 leaflets of 5-7.5 cm and margins entire (Swearingen and Bargeron, 2016). Leaflets are ovate or broadly lanceolate to narrowly deltate (Flora of North America Editorial Committee, 2018). Inflorescences are axillary 3-12-flowered cymes (or compound cymes or paniculate with cymose subunits) (Flora of North America Editorial Committee, 2018). Flowers are 1.4-3.0 cm in diameter (Flora of China Editorial Committee, 2018), bisexual often with some unisexual flowers in the same inflorescence, with pedicels of 1-3.5 cm (Flora of North America Editorial Committee, 2018). Flowers are fragrant with four slender white petal-like sepals (Swearingen and Bargeron, 2016) that are obovate-oblong and measure 5-15 x 2-6 mm (Flora of China Editorial Committee, 2018). Flowers have up to 50 stamens and 5-10 unicarpellate pistils (Flora of North America Editorial Committee, 2018). The ovary is superior (Burnham, 2013). Seeds are enclosed in flattened achenes, production is prolific, and seed heads have long, silvery-grey, feather-like hairs (Swearingen and Bargeron, 2016). Each achene has a plume attached (Mahr, 2017), and this helps with wind dispersal (Burnham, 2013). The mature bark is light brown and shreds longitudinally (Burnham, 2013).

Plant Type

Distribution

C. terniflora is native to China, Japan, Korea, Mongolia, Taiwan and parts of Russia (Flora of China Editorial Committee, 2018; USDA-ARS, 2018). It has escaped cultivation and is naturalized in many parts of the USA, particularly in the East and Midwest (Missouri Botanical Garden, 2018). It is reported as invasive in Alabama, Delaware, Florida, Georgia, South Carolina and Tennessee in the USA (Alabama Invasive Species Council, 2007; SC-EPPC, 2011; FLEPPC, 2017; GA-EPPC, 2018; TE-EPPC, 2018; Delaware Invasive Species Council, 2019). It is also introduced in Canada, Indonesia and Nepal (Roskov et al., 2018).

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

C. terniflora was introduced to the USA as an ornamental plant in 1877 (Mahr, 2017; SC-EPPC, undated) via seeds sent to an Arboretum in Boston (Mahr, 2017).

Introductions

Risk of Introduction

C. terniflora is still widely available as an ornamental, so it is likely that it will continue to spread into wild areas from gardens, where it is grown.

Habitat

In the USA, C. terniflora is found along roadsides and in thickets, forest edges, rights of ways and urban green space, especially near creeks (Swearingen and Bargeron, 2016; Flora of North America Editorial Committee, 2018).

In China, C. terniflora is found in forest margins, scrub on slopes, grassy areas on hills, among rocks in coastal areas at sea level to 800 m (Flora of China Editorial Committee, 2018).

Habitat List

Biology and Ecology

Genetics

C. terniflora is tetraploid (2n = 4x = 32) (Wang et al., 2017).

Reproductive Biology

When C. terniflora was grown under natural photoperiods of at least 14 h, flowering was favoured by day/night temperatures of 24/19˚C and 30/25˚C (Suzuki and Hori, 1991). At temperatures of 17/12˚C, shoot growth was delayed and flower buds did not develop (Suzuki and Hori, 1991). Flower buds did not develop under short photoperiods of 8 h or 12 h (Suzuki and Hori, 1991). Spraying plants with GA3 decreased flower numbers and resulted in over 50% of the flowers having degenerated anthers and undeveloped pistils (Suzuki and Hori, 1991).  

C. terniflora is a perennial plant that reproduces both vegetatively and by seed (Global Invasive Species Database, 2010). Seeds (achenes) are small and brown and germinate within 1-9 months (Global Invasive Species Database, 2010). C. terniflora is self-pollinating and a high proportion of its seed is viable (Skibo, 2006).

Physiology and Phenology

With support, C. terniflora can grow to over 9 m high (Global Invasive Species Database, 2010). Where it is unable to climb, it will sprawl across the ground forming dense, 15-30 cm tall and 3 m wide, clumps (Global Invasive Species Database, 2010; Missouri Botanical Garden, 2018).

Flowering occurs from July to September (Flora of North America Editorial Committee, 2018), and flowers attract bees and flies (Burnham, 2013).

Environmental Requirements

C. terniflora prefers full sun or partial shade, where its leaves can be in the sun and its roots in the shade (Floridata, 2015; Missouri Botanical Garden, 2018). It can grow in light (sandy), medium (loamy), or heavy (clay) soils that are fairly well drained. It also tolerates a variety of pH, including highly alkaline settings (Global Invasive Species Database, 2010).

In the USA, C. terniflora grows to elevations of 1000 m, and in China, it has been recorded up to 800 m above sea level (Flora of China Editorial Committee, 2018; Flora of North America Editorial Committee, 2018).

Climate

Latitude/Altitude Ranges

Soil Tolerances

Soil drainage

• free

Soil reaction

• acid
• alkaline
• neutral
• very alkaline

Soil texture

• heavy
• light
• medium

Natural enemies

Means of Movement and Dispersal

Natural Dispersal

Seeds have long feathery plumes and are widely dispersed by the wind (Skibo, 2006; Global Invasive Species Database, 2010).

Vector Transmission (biotic)

Seeds of C. terniflora are spread by wildlife and human activity (Skibo, 2006).

Accidental Introduction

C. terniflora can self-seed and has escaped cultivation and naturalized in many parts of the USA (Missouri Botanical Garden, 2018).

Intentional Introduction

C. terniflora has been introduced outside its native range as an ornamental plant (Mahr, 2017).

Pathway Causes

Pathway Vectors

Impact Summary

Environmental Impact

C. terniflora can aggressively self-seed and invade roadsides, thickets and wood edges near creeks (SC-EPPC, undated). The presence of C. terniflora degrades the quality of natural habitats by decreasing the light, space, moisture and nutrients available to other species (SC-EPPC, undated). It also inhibits the growth of legumes (Burnham 2013; SC-EPPC, undated).

Impact: Biodiversity

C. terniflora grows rapidly and forms dense clumps that smother and shade slower growing trees, shrubs and herbs, and also suppress seed bank germination (Skibo, 2006; Global Invasive Species Database, 2010; Missouri Botanical Garden, 2018).

Social Impact

C. terniflora is capable of pulling down telephone poles (Global Invasive Species Database, 2010).

Risk and Impact Factors

• Proved invasive outside its native range
• Fast growing
• Has high reproductive potential

• Ecosystem change/ habitat alteration
• Reduced native biodiversity
• Threat to/ loss of native species

• Competition - monopolizing resources
• Competition - shading
• Competition - smothering
• Competition - strangling
• Rapid growth

Uses

Economic Value

C. terniflora is traded as an ornamental and used for landscaping in southeastern USA (Langeland and Meisenburg, 2009).   

Social Benefit

C. terniflora has been used in traditional Chinese medicine to treat tonsillitis, rheumatoid arthritis and prostatitis, and investigations have found it to contain anti-inflammatory and antinociceptive compounds (Liu et al., 2015). In Korea, C. terniflora is used to treat dysentery, neuralgia and gynecological problems (Burnham, 2013). It is sold in Chinese and Korean markets (Han et al., 2013).

Some Clematis species are mildly poisonous; however young shoots and buds of C. ternifolia can be eaten if carefully boiled, roasted or pickled (PFAF, 2012).

Uses List

General

• Botanical garden/zoo

Medicinal, pharmaceutical

• Traditional/folklore

Ornamental

• garden plant

Similarities to Other Species/Conditions

The species C. virginiana, native to the USA, is very similar to C. terniflora, although it tends to have toothed leaf margins and is not as prone to self-seeding and spreading as C. terniflora (Swearingen and Bargeron, 2016). The floristic changes in the New York metropolitan region were monitored between two time periods (1901-1950 and 1951-2000) (Clemants and Moore, 2005). It was found that C. terniflora was spreading in the area, whilst the native C. virginiana was declining (Clemants and Moore, 2005).

C. terniflora is easily distinguished from C. occidentalis during flowering as the latter has purple, solitary flowers (Burnham, 2013). 

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

It has been suggested that one way to limit the spread of C. terniflora, is to encourage people to plant alternative species, such as native vines, in their gardens instead (Skibo, 2006).

Control

Physical/Mechanical Control

In a study conducted over six growing seasons, removal of non-native plants, including C. terniflora, either by hand, using chemicals, or a combination of both, led to an increase in native plant cover and richness, and a decline in the abundance of non-native species (Farmer et al., 2016). However, C. terniflora was still a significant community component even after removal treatments, suggesting that it is difficult to control (Farmer et al., 2016).

C. terniflora seedlings can be pulled by hand or mowed, and mature plants can be cut down by hand or mowed (Skibo, 2006). Plants are dormant in winter, making it easier to cut down larger plants at this time (Skibo, 2006).

Chemical Control

In Alachua County, Florida, C. terniflora is treated with foliar applications of the herbicides, glyphosate and pelargonic acid (Global Invasive Species Database, 2010). Glyphosate-based products or growth regulator herbicides such as 2,4-D and Dicamba may be used to control C. terniflora (Skibo, 2006).

In herbicide trials in Florida, imazapic 0.6 g ae/L, glyphosate as low as 3.6 g ae/L, triclopyr as low as 3.6 g ae/L, metsulfuron 0.07 g ai/L, and fluroxypyr as low as 1.8 g ae/L, resulted in 72-99% control of C. terniflora 90 days after application (Langeland and Meisenburg, 2009). It is suggested that repeat applications will be required to control this plant where it is invasive (Langeland and Meisenburg, 2009).

Gaps in Knowledge/Research Needs

There is a lack of information about the direct effects of C. terniflora on native species, such as C. virginiana. More information is also needed about the costs and timescales required to clear C. terniflora infestations and the potential for use of biological control agents. 

References

Alabama Invasive Plant Council, 2007. Invasive Plant List. https://www.se-eppc.org/alabama/2007plantlist.pdf

Bowen B, Johnson K, Franklin S, Call G, Webber M, 2002. Invasive exotic pest plants in Tennessee. Journal of the Tennessee Academy of Science, 77(2), 45-48. http://www.tennacadofsci.org/journal/articles/vol77/JTAS77-2-45.pdf

Burnham RJ, 2013. Climbers. Censusing lianas in mesic biomes of Eastern regions. http://climbers.lsa.umich.edu/

Clemants SE, Moore G, 2005. The changing flora of the New York metropolitan region. Urban Habitats, 3(1), 192-210.

Delaware Invasive Species Council, 2019. Invasive Plant List. https://www.invasive.org/species/list.cfm?id=90

Dong Yan, Sun YanMei, Xue XiaoFeng, 2016. Two new eriophyid mite species associated with Clematis terniflora var. mandshurica in China (Acari, Eriophyidae). ZooKeys, (No.621), 1-14. http://zookeys.pensoft.net/articles.php?id=9443&display_type=element&element_type=8&element_id=27896&element_name=

Farmer, S., Ward, J. R., Horton, J. L., Clarke, H. D., 2016. Southern Appalachian urban forest response to three invasive plant removal treatments. Management of Biological Invasions, 7(4), 329-342. http://www.reabic.net/journals/mbi/2016/4/MBI_2016_Farmer_etal.pdf doi: 10.3391/mbi.2016.7.4.03

FLEPPC, 2017. 2017 FLEPPC List of Invasive Plant Species. The Florida Exotic Plant Pest Council.http://bugwoodcloud.org/CDN/fleppc/plantlists/2017/2017FLEPPCLIST-TRIFOLD-FINALAPPROVEDBYKEN-SUBMITTEDTOALTA.pdf

Flora of China Editorial Committee, 2018. Flora of China. St. Louis, Missouri and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria.http://www.efloras.org/flora_page.aspx?flora_id=2

Flora of North America Editorial Committee, 2018. Flora of North America of Mexico. St. Louis, Missouri and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria.http://www.efloras.org/flora_page.aspx?flora_id=1

Floridata, 2015. Floridata Plant Encyclopedia. https://floridata.com/plantlist/

GA-EPPC, 2018. List of non-native invasive plants in Georgia. Georgia Exotic Plant Pest Council.https://www.gaeppc.org/list/

Global Invasive Species Database, 2010. Global Invasive Species Database (GISD). http://issg.org/database/welcome/

Han Yan, Kitaoka, F., Mano, M., Sasaki, Y., Mikage, M., 2013. ITS analysis of Clematis plants from East Asia and the botanical origin of Clematidis Radix sold in modern markets. Journal of Traditional Medicines, 30(3), 91-101. http://www.wakan-iyaku.gr.jp

Langeland, K., Meisenburg, M., 2009. Herbicide evaluation to control Clematis terniflora invading natural areas in Gainesville, Florida. Invasive Plant Science and Management, 2(1), 70-73. http://www.wssa.net doi: 10.1614/IPSM-08-110.1

Liu, X. B., Yang, B. X., Zhang, L., Lu, Y. Z., Gong, M. H., Tian, J. K., 2015. An in vivo and in vitro assessment of the anti-inflammatory, antinociceptive, and immunomodulatory activities of Clematis terniflora DC. extract, participation of aurantiamide acetate. Journal of Ethnopharmacology, 169, 287-294. http://www.sciencedirect.com/science/article/pii/S0378874115002603

Mahr S, 2017. Sweet Autumn Clematis, Clematis terniflora. In: Master Gardener Program, Wisconsin, USA: University of Wisconsin Extension.https://wimastergardener.org/article/sweet-autumn-clematis-clematis-terniflora/

MIPN, 2018. Midwest Invasive Plant List. https://www.mipn.org/plantlist/

Missouri Botanical Garden, 2018. Missouri Botanical Garden Plant Finder. St. Louis, MO, USA: Missouri Botanical Garden.http://www.missouribotanicalgarden.org/plantfinder/plantfindersearch.aspx

PFAF, 2012. Plants For A Future Database. http://www.pfaf.org/USER/Default.aspx

Roskov Y, Abucay L, Orrell T, Nicolson D, Bailly N, Kirk PM, Bourgoin T, DeWalt RE, Decock W, De Wever A, Nieukerken E van, Zarucchi J, Penev L, 2018. Species 2000 & ITIS Catalogue of Life. Leiden, The Netherlands: Species 2000, Naturalis.www.catalogueoflife.org/col

SC-EPPC, 2004. South Carolina Exotic Pest Plant Council Non-Native Invasive Plant Species List – March, 2004. https://www.se-eppc.org/southcarolina/SCList.pdf

SC-EPPC, 2011. Invasive Plant Pest Species of South Carolina. Forestry Leaflet 28, July 2011. Clemson University Cooperative Extension Service. South Carolina Exotic Pest Plant Council.https://www.se-eppc.org/southcarolina/scinvasives.pdf

SC-EPPC, undated. Green Gone Bad. South Carolina Exotic Pest Plant Council.https://www.se-eppc.org/southcarolina/Publications/ClematisTernifoliaArticleSCNLA.pdf

Skibo AZ, 2006. Invasive Plant List. Planting for a livable Delaware. http://dnrec.delaware.gov/fw/dplap/information/Documents/DE%20Invasive%20Plant%20Booklet%20(DDA%20Plant%20Industries).pdf

Suzuki, S., Hori, Y., 1991. Effects of temperature, photoperiod and exogenous gibberellin on flowering of Clematis terniflora DC. Journal of the Japanese Society for Horticultural Science, 60(3), 643-650. doi: 10.2503/jjshs.60.643

Swearingen J, Bargeron C, 2016. Invasive Plant Atlas of the United States. University of Georgia Center for Invasive Species and Ecosystem Health.http://www.invasiveplantatlas.org/

TE-EPPC, 2009. Invasive Exotic Pest Plants in Tennessee – 2009. Tennessee Exotic Plant Pest Council.http://s3.amazonaws.com/tneppc2/uploads/619/original/tn-eppc-plant-list-ww-w09-final-1.pdf

Tredici, P. del, 2017. The introduction of Japanese plants into North America. Botanical Review, 83(3), 215-252. https://link.springer.com/article/10.1007/s12229-017-9184-3 doi: 10.1007/s12229-017-9184-3

USDA-ARS, 2018. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, USA: National Germplasm Resources Laboratory.http://www.ars-grin.gov/cgi-bin/npgs/html/tax_search.pl

USDA-NRCS, 2018. The PLANTS Database. Baton Rouge, USA: National Plant Data Center.http://plants.usda.gov/

Wang Na, Wang KuiLing, Liu QingHua, Liu QingChao, 2017. Karyotype analysis of seven wild Clematis species. Acta Prataculturae Sinica, 26(11), 123-130. http://cyxb.lzu.edu.cn/EN/volumn/current.shtml

Distribution References

Alabama Invasive Plant Council, 2007. Invasive Plant List., https://www.se-eppc.org/alabama/2007plantlist.pdf

Bowen B, Johnson K, Franklin S, Call G, Webber M, 2002. Invasive exotic pest plants in Tennessee. In: Journal of the Tennessee Academy of Science, 77 (2) 45-48. http://www.tennacadofsci.org/journal/articles/vol77/JTAS77-2-45.pdf

Burnham RJ, 2013. Climbers. Censusing lianas in mesic biomes of Eastern regions., http://climbers.lsa.umich.edu/

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

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

Clemants SE, Moore G, 2005. The changing flora of the New York metropolitan region. In: Urban Habitats, 3 (1) 192-210.

Delaware Invasive Species Council, 2019. Invasive Plant List., https://www.invasive.org/species/list.cfm?id=90

FLEPPC, 2017. 2017 FLEPPC List of Invasive Plant Species., The Florida Exotic Plant Pest Council. http://bugwoodcloud.org/CDN/fleppc/plantlists/2017/2017FLEPPCLIST-TRIFOLD-FINALAPPROVEDBYKEN-SUBMITTEDTOALTA.pdf

Flora of China Editorial Committee, 2018. Flora of China. In: Flora of China. St. Louis, Missouri and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria. http://www.efloras.org/flora_page.aspx?flora_id=2

GA-EPPC, 2018. List of non-native invasive plants in Georgia., Georgia Exotic Plant Pest Council. https://www.gaeppc.org/list/

Langeland K, Meisenburg M, 2009. Herbicide evaluation to control Clematis terniflora invading natural areas in Gainesville, Florida. Invasive Plant Science and Management. 2 (1), 70-73. http://www.wssa.net DOI:10.1614/IPSM-08-110.1

MIPN, 2018. Midwest Invasive Plant List., https://www.mipn.org/plantlist/

Roskov Y, Abucay L, Orrell T, Nicolson D, Bailly N, Kirk PM, Bourgoin T, DeWalt RE, Decock W, De Wever A, Nieukerken E van, Zarucchi J, Penev L, 2018. Species 2000 & ITIS Catalogue of Life., Leiden, The Netherlands: Species 2000, Naturalis. http://www.catalogueoflife.org/col

SC-EPPC, 2004. South Carolina Exotic Pest Plant Council Non-Native Invasive Plant Species List – March, 2004., https://www.se-eppc.org/southcarolina/SCList.pdf

SC-EPPC, 2011. Invasive Plant Pest Species of South Carolina. Forestry Leaflet 28, July 2011., Clemson University Cooperative Extension Service. South Carolina Exotic Pest Plant Council. https://www.se-eppc.org/southcarolina/scinvasives.pdf

Skibo AZ, 2006. Invasive Plant List. Planting for a livable Delaware., http://dnrec.delaware.gov/fw/dplap/information/Documents/DE%20Invasive%20Plant%20Booklet%20(DDA%20Plant%20Industries).pdf

Swearingen J, Bargeron C, 2016. Invasive Plant Atlas of the United States., University of Georgia Center for Invasive Species and Ecosystem Health. http://www.invasiveplantatlas.org/

USDA-ARS, 2018. Germplasm Resources Information Network (GRIN). Online Database. In: Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysimple.aspx

USDA-NRCS, 2018. The PLANTS Database. In: The PLANTS Database. Greensboro, North Carolina, USA: National Plant Data Team. https://plants.sc.egov.usda.gov

Links to Websites

Contributors

03/06/18 Original text by:

Vicki Cottrell, Consultant, UK

Distribution Maps