Euonymus japonicus
(Japanese spindle tree)

Pictures

Picture	Title	Caption	Copyright
Title Habit Caption Euonymus japonicus (Japanese spindle tree); habit, planted as an ornamental. Bupyung, Korea. June 2008. Copyright ©Dalgial/via wikipedia - CC BY-SA 3.0	Habit	Euonymus japonicus (Japanese spindle tree); habit, planted as an ornamental. Bupyung, Korea. June 2008.	©Dalgial/via wikipedia - CC BY-SA 3.0
Title Habit Caption Euonymus japonicus (Japanese spindle tree); habit, planted as an ornamental. Incheon, Korea. February 2008. Copyright ©Dalgial/via wikipedia - CC BY-SA 3.0	Habit	Euonymus japonicus (Japanese spindle tree); habit, planted as an ornamental. Incheon, Korea. February 2008.	©Dalgial/via wikipedia - CC BY-SA 3.0
Title Foliage Caption Euonymus japonicus (Japanese spindle tree); foliage, showing old and young leaves. Bupyung, Korea. June 2008. Copyright ©Dalgial/via wikipedia - CC BY-SA 3.0	Foliage	Euonymus japonicus (Japanese spindle tree); foliage, showing old and young leaves. Bupyung, Korea. June 2008.	©Dalgial/via wikipedia - CC BY-SA 3.0
Title Foliage Caption Euonymus japonicus (Japanese spindle tree); variegated foliage. The Botanical Garden of the University of Latvia. April 2015. Copyright ©Aleksandrs Balodis (AfroBrazilian)/via wikipedia - CC BY-SA 3.0	Foliage	Euonymus japonicus (Japanese spindle tree); variegated foliage. The Botanical Garden of the University of Latvia. April 2015.	©Aleksandrs Balodis (AfroBrazilian)/via wikipedia - CC BY-SA 3.0
Title Flowers Caption Euonymus japonicus (Japanese spindle tree); flowers. Hockenheim, Germany. August 2015. Copyright Public Domain - Released by AnRo0002/via wikipedia - CC0 1.0 Universal (CC0 1.0)	Flowers	Euonymus japonicus (Japanese spindle tree); flowers. Hockenheim, Germany. August 2015.	Public Domain - Released by AnRo0002/via wikipedia - CC0 1.0 Universal (CC0 1.0)
Title Fruits Caption Euonymus japonicus (Japanese spindle tree); fruits. Hockenheim, Germany. December 2016. Copyright Public Domain - Released by AnRo0002/via wikipedia - CC0 1.0 Universal (CC0 1.0)	Fruits	Euonymus japonicus (Japanese spindle tree); fruits. Hockenheim, Germany. December 2016.	Public Domain - Released by AnRo0002/via wikipedia - CC0 1.0 Universal (CC0 1.0)
Title Fruits Caption Euonymus japonicus (Japanese spindle tree); fruits and foliage. Aizu area, Fukushima prefecture, Japan. January 2010. Copyright Public Domain - Released by Qwert1234 /via wikipedia - CC0	Fruits	Euonymus japonicus (Japanese spindle tree); fruits and foliage. Aizu area, Fukushima prefecture, Japan. January 2010.	Public Domain - Released by Qwert1234 /via wikipedia - CC0

Identity

Preferred Scientific Name

• Euonymus japonicus Thunb.

Preferred Common Name

• Japanese spindle tree

Other Scientific Names

• Elaeodendron javanicum Turcz.
• Euonymus carrierei Dippel
• Euonymus fortunei var. alticola Hand.-Mazz.
• Euonymus pulchellus Dippel
• Euonymus repens Carrière
• Euonymus sinensis Carrière
• Masakia japonica (Thunb.) Nakai

International Common Names

• English: evergreen euonymus; evergreen spindle; spindle tree
• French: fusain du Japon

Local Common Names

• China: dong qing wei mao
• France: bonnet de prêtre
• Germany: Spindelstrauch, Japanischer
• Italy: evonimo del Giappone
• Japan: masaki
• Malaysia: belimbing hutan; belungkas; kemuning ayer
• Netherlands: kardinaalsmuts, Japanse
• Philippines: malasangki
• Portugal: evônimo-do-Japão
• Sweden: japansk benved
• Thailand: khao kwang; kraduuk kai

EPPO code

• EUOJA (Euonymus japonicus)

Summary of Invasiveness

Euonymus japonicus is native in Japan (and perhaps China and Korea) but has been widely introduced for cultivation as an ornamental or hedge plant. It has become naturalized in other Asian countries, a number of states in the USA, and in some European countries including the UK. It is of concern for its potential to be invasive. E. fortunei, which is closely related, is already proving invasive in some regions. In New Zealand, E. japonicus is cited as a ‘weed of concern on conservation lands’ (Invasive Plants Database, 2017). In Australia, it has been ranked as an environmental weed and declared weed (regulated) ‘with the potential to have serious impact' (Randall, 2007). E. japonicus was included in the Life IAP-Risk list of 37 potentially invasive species to the EU (Tanner, 2017), but was subsequently removed due to a lack of adequate information (Life IAP-Risk, 2016). As the seeds are dispersed by birds it has the potential to readily spread outside cultivation.

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Plantae
•         Phylum: Spermatophyta
•             Subphylum: Angiospermae
•                 Class: Dicotyledonae
•                     Order: Celastrales
•                         Family: Celastraceae
•                             Genus: Euonymus
•                                 Species: Euonymus japonicus

Notes on Taxonomy and Nomenclature

There are a number of synonyms listed for Euonymus japonicus but it is not believed that any of these are commonly used. Confusion could be caused by the use of E. fortunei var. alticola to refer to E. japonicus due to similarity with E. fortunei, which is also an invasive species. There are a range of recognized subspecies or varieties for E. japonicus, many developed as ornamentals. Some hybrids have also been developed from E. japonicus and related species (e.g. E. kiautschovicus and E. fortunei) that show greater cold hardiness (RenJun, 2013).

Description

E. japonicus grows as an evergreen shrub or small tree that can reach up to 3 m, sometimes dwarfed. Branches are grey-green to grey-brown, terete glabrous and sturdy. Twigs are green to light green, glabrous, and not evidently striate, especially when fresh. The petiole is 3-10 mm long. Leaf blades are leathery or thickly leathery, ovate, obovate, orbicular-ovate or long ovate, measuring (3-)5-10(-12) x (2-)3-5(-5.5) cm, base orbicular or semiorbicular, margin crenulate distally, nearly entire proximally, apex orbicular or semiorbicular; lateral veins 6-8 pairs, slightly visible or unclear, especially when dry. Leaves are often variegated in cultivated varieties. Cymes usually axillary, sometimes terminal, many branched with many flowers; peduncle up to 8 cm, sub-branches 2-4 cm; pedicel 4-7 mm. Flowers are 4-merous, 5-6 mm in diameter; sepals nearly orbicular; petals greenish white or yellowish green, sometimes cream, nearly orbicular. Capsule globose or subglobose, brown or yellow-brown to red-brown, 6-9(-12) mm in diameter, 4-lobed. Seeds 2 per locule, 5 x 3.5 mm, dark brown, globose; aril orange-red (Flora of China Editorial Committee, 2017).

Plant Type

Distribution

According to most sources, E. japonicus, is only native to Japan but some sources, e.g. World Agroforestry Centre (2017), indicate that it is also native to China and Korea. E. japonicus has been widely introduced for cultivation as an ornamental or hedge plant. This is well documented in many countries in Europe (DAISIE, 2017; Flora Europaea, 2017; Online Atlas of the British and Irish Flora, 2017), USA (USDA-NRCS, 2017), South and South East Asia (Flora of China Editorial Committee, 2017; USDA-ARS, 2017), Australia (Randall, 2007) and New Zealand (GBIF, 2017), and locally in South America (Bolivia) (Missouri Botanical Garden, 2017), and may well occur in more countries than are indicated in this datasheet. It has naturalized in some parts of the USA (USDA-NRCS, 2017) and Europe (DAISIE, 2017), and elsewhere (GBIF, 2017).

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

E. japonicus was introduced to cultivation in the UK in 1804 and has been known in the wild on the Isles of Scilly since 1897 (Online Atlas of the British and Irish Flora, 2017).

Introductions

Risk of Introduction

There is a high risk of introduction, due to its popularity as an ornamental plant. The likelihood of it becoming invasive when introduced is far less certain. As the seeds are dispersed by birds it has the potential to readily spread outside cultivation.

Habitat

In the UK it has naturalized in woodland, on sea-cliffs and roadsides (Online Atlas of the British and Irish Flora, 2017; PFAF, 2017). In China it occurs from sea level up to 1400 m.

Habitat List

Biology and Ecology

Genetics

The chromosome number of E. japonicus is 2n = 32 (Missouri Botanical Garden, 2017).

Reproductive Biology

E. japonicus reproduces by seed. Flowering occurs in early summer and flowers are pollinated by bees and flies. Seed is dormant when shed and requires stratification. Stored seeds require 3 months of cold stratification, and can take up to 18 months to germinate (PFAF, 2017). Increasing the period of cold stratification can improve the rate of seed germination to 65% after 60 days germination. Soaking the seeds in gibberellin can further enhance germination. Optimum germination occurs at about 25^°C (Xue, 2011).

For commercial purposes propagation can be obtained from cuttings (Tawfik, 2001).

Physiology and Phenology

E. japonicus is an evergreen shrub with a periodic growth pattern consisting of several growth flushes per year. In each flush, 9-11 nodes are added to the stem length. Nodes formed at the beginning of a growth flush are short and bear scale-like leaves. In the middle of the flush, the nodes are longer and become shorter again toward the end of the flush (Zieslin and Pines, 1987).

Longevity

Trees may live for up to 150 years (Urban Forest Ecosystems Institute, 2017).

Environmental Requirements

E. japonicus tolerates temperatures as low as -10°C (PFAF, 2017) and is rated by PFAF (2017) as being tolerant to both UK and USA hardiness zones 7 (down to -20°C). It also has a wide tolerance to different soil types and acidity levels and some tolerance of salinity (Urban Forest Ecosystems Institute, 2017). It does not tolerate deep shade but has medium shade tolerance (Lin ShuYan, 2007). Plants are very tolerant of maritime exposure, succeeding even when grown on the seashore but they can be killed by cold drying winds (Useful Tropical Plants, 2017).

When grown as an ornamental, it tolerates light, medium and heavy (clay) soils, but prefers well-drained soil. It also tolerates acid, neutral and basic (alkaline) soils. The plant prefers night temperatures of 4-12°C and day temperatures of 19°C or lower.

Climate

Latitude/Altitude Ranges

Air Temperature

Rainfall

Rainfall Regime

Soil Tolerances

Soil drainage

• free

Soil reaction

• acid
• alkaline
• neutral

Soil texture

• heavy
• light
• medium

Natural enemies

Notes on Natural Enemies

A wide range of pests and pathogens have been recorded on E. japonicus, scale insects, thrips and powdery mildew are perhaps the most common causes of damage.

Major host of:

Colletotrichum boninense; Erysiphe euonymi-japonici; Luperomorpha xanthodera; Parthenolecanium persicae (peach scale); Senecio vulgaris.

Minor host of:

Alfalfa mosaic virus (alfalfa yellow spot); Armillaria luteobubalina (armillaria root rot); Ceroplastes floridensis (soft scale); Ceroplastes japonicus (tortoise wax scale); Cydalima perspectalis (box tree moth); Epiphyas postvittana (light brown apple moth); Exomala orientalis (oriental beetle); Metcalfa pruinosa (frosted moth-bug); Pratylenchus penetrans (nematode, northern root lesion).

Wild host of:

Aphis fabae (black bean aphid).

Host of:

Aphis fabae cirsiiacanthoidis; Fusarium oxysporum (basal rot); Pryeria sinica (pellucid, zygaenid (Japan)); Unaspis euonymi (euonymus scale).

Pryeria sinica (Euonymus leaf notcher) is newly recorded in the USA (University of Maryland Extension, 2017). E. japonicus is noted to be susceptible to scales, spider mites and thrips, oak root rot, powdery mildew and root rot (Urban Forest Ecosystems Institute, 2017).

In California, E. japonicus can be heavily infested by the light brown apple moth, Epiphyas postvittana, an exotic introduction from Australia (Wang et al., 2012).

Unaspis euonymi is found in the centre of Brussels, Belgium, and in Belgrade, Serbia, causing severe leaf loss and dieback (Graora, 2007; Malumphy, 2016)

Heavy infestations of Eutetranychus orientalis have been recorded in Shandong, China (Sun et al., 1996).

Rhizoctonia solani [Thanatephorus cucumeris] can be a serious problem in seedling nurseries of E. japonicus (Lee et al., 1996).

Paratylenchus dianthus was found to cause severe leaf drop of E. japonicus in Chongqing, China (Liu, 1995).

It may also be a host of the sugar beet fly, Pegomya mixta [Pegomya cunicularia] (PFAF, 2017).

Means of Movement and Dispersal

Vector Transmission (biotic)

A wide range of bird species disperse the seeds of E. japonicus (Orwa et al., 2009; Forest and Bird, 2017).

Accidental Introduction

Accidental introduction is extremely unlikely.

Intentional Introduction

E. japonicus has been widely introduced deliberately for horticultural purposes, as an ornamental or hedge plant. It is available from a wide range of companies and intentional introduction is therefore highly likely.

Pathway Causes

Pathway Vectors

Impact Summary

Impact: Biodiversity

There are no specific examples of problems, but of concern for its potential invasiveness in several countries.

Social Impact

All parts of E. japonicus are reported to be poisonous when ingested, causing vomiting, diarrhoea, weakness, chills, coma and convulsions. Seeds and arils are particularly poisonous. Otherwise, large amounts of foliage may be needed to cause acute poisoning. Though the toxic principle is not clearly defined, it is ‘believed to be a glycoside’ (North Carolina State University, 2017). In Belgium, E. japonicus has been responsible for causing sickness and death in sheep (Baert et al., 2005).

Poison Delivery Mode: ingestion

Symptoms: vomiting, diarrhoea, weakness, chills, coma and convulsions.

Toxic Principle: unidentified, possibly a glycoside, peptide or sesquiterpene alkaloid.

Risk and Impact Factors

• Benefits from human association (i.e. it is a human commensal)
• Long lived
• Fast growing
• Has propagules that can remain viable for more than one year

• Negatively impacts human health
• Reduced native biodiversity

• Poisoning
• Rapid growth

• Highly likely to be transported internationally deliberately

Uses

Economic Value

E. japonicus is primarily used as a hardy, versatile ornamental garden plant, being available in a wide variety of foliage colours and plant shapes, available from many internet suppliers across the world. It is also valued as a hedge plant and in Korea it is of interest for sound proofing (Kim et al., 1989).

E. japonicus is also described as an important forest tree species in Shandong, China (Sun et al., 1996).

Roots and stems yield up to 7% gutta-percha, a non-elastic rubber used as an electrical insulator and in making plastics (Useful Tropical Plants, 2017). It is cultivated for this purpose in Russia and in Spain (PFAF, 2017).

E. japonicus has apparently been used widely as a research tool in the study of photosynthesis, cold and drought-hardiness, salt-tolerance and other physiological reactions, especially in China. Some of this work relates to its use for ornamental and conservation purposes, but some may be because of its common availability.

Social Benefit

Decoctions from the bark are considered to be tonic, anti-rheumatic, anhidoritic and diuretic. Chinese women use the leaves to aid difficult childbirths (Duke and Ayensu, 1985 in: Useful Tropical Plants, 2017). American Indians use the occidental vicariads in gynaecological applications (Orwa et al., 2009).

In spite of indications of toxicity, there are reports of the leaves being edible (PFAF, 2017).

Environmental Services

E. japonicus is widely studied in China for its use in conservation and re-vegetation.

Uses List

Environmental

• Agroforestry
• Amenity
• Landscape improvement
• Revegetation
• Soil conservation

General

• Research model

Medicinal, pharmaceutical

• Source of medicine/pharmaceutical

Ornamental

• Propagation material

Similarities to Other Species/Conditions

E. japonicus is very similar to E. fortunei but is more erect, growing to 3 m tall, while the latter has a climbing or procumbent habit (Flora of China Editorial Committee, 2017). It is distinguished from most other species by its leathery leaves and smooth, rounded fruit, not lobed (Stace, 1997; Schulz, 2006).

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.

Physical/Mechanical Control

Established plants can be removed by cutting and by uprooting if small enough, but larger stumps may need to be covered with thick black polyethylene to exclude light (Forest and Bird, 2017).

Biological Control

E. japonicus is damaged by a wide range of insects, pathogens and nematodes but there are no reports of attempts at exploiting these for biological control.

Chemical Control

The growth of E. japonicus can be inhibited by the growth regulator paclobutrazol (Keever et al., 1990); also by uniconazole (Norcini and Knox, 1990); and by dikegulac (Yamazaki et al., 1979) but these would only limit the size of established plants. In New Zealand, recommendations include cutting and painting the stump with a 1-2 mm layer of metsulfuron gel; also spraying foliage with metsulfuron (Forest and Bird, 2017). Glyphosate is not included as a means of control in New Zealand but it is known to be mildly damaging to the foliage and GISD (2017) recommendations for control of the closely related E. fortunei include the application of glyphosate or triclopyr to freshly cut stems.

References

Baert, K., Croubels, S., Steurbaut, N., Boever, S. de, Vercauteren, G., Ducatelle, R., Verbeken, A., Backer, P. de, 2005. Two unusual cases of plant intoxication in small ruminants. Vlaams Diergeneeskundig Tijdschrift, 74(2), 149-153. HTTP://VDT.UGENT.BE

DAISIE, 2017. Delivering Alien Invasive Species Inventories for Europe. European Invasive Alien Species Gateway. www.europe-aliens.org/default.do

Dave's Garden, 2017. Dave's Garden. http://davesgarden.com/

Encyclopedia of Life, 2017. Encyclopedia of Life. http://eol.org/

EPPO, 2017. EPPO Global Database. Paris, France: European and Mediterranean Plant Protection Organization.https://gd.eppo.int/

Flora Europaea, 2017. Euonymus japonicus. http://193.62.154.38/cgi-bin/nph-readbtree.pl/feout?FAMILY_XREF=&GENUS_XREF=Euonymus&SPECIES_XREF=japonic*&TAXON_NAME_XREF=&RANK=

Flora of China Editorial Committee, 2017. 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

Forest and Bird, 2017. Weed Control Guide. http://www.forestandbird.org.nz/files/Weed_Guide_Final_small.pdf

GBIF, 2017. Global Biodiversity Information Facility. http://www.gbif.org/species

GISD, 2017. Global Invasive Species Database (GISD). http://www.iucngisd.org/gisd/

Graora, D., 2007. Biology and harmfulness of Unaspis euonymi (Comstock) in the area of Belgrade. Pesticidi i Fitomedicina, 22(3), 247-253.

Hagiladi, A., Raviv, M., Ran, I., Alzam, O., Raviv, A., Ben-Jaacov, J., 1981. Cultivation of Myrtus and Euonymus spp. in the mountainous regions of Israel. Hassadeh, 61(10), 1682-1685.

Invasive Plants Database, 2017. Greenery Technology and Landscape Planning. http://invasive.m-fuukei.jp/

Keever, G. J., Foster, W. J., Stephenson, J. C., 1990. Paclobutrazol inhibits growth of woody landscape plants. Journal of Environmental Horticulture, 8(1), 41-47.

Kim, Y. S., Chang, H. G., Kim, Y. H., 1989. Noise attenuation by landscape woody plants. Comparisons as a hedge species between Japanese spindle tree and oriental arbor-vitae. Journal of Korean Forestry Society, 78(1), 30-34.

Lee, Y. H., Huang, J. W., Leu, L. S., 1996. Integrated control of seedling blight of Euonymus japonicus. Plant Protection Bulletin (Taipei), 38(3), 261-273.

Life IAP-Risk, 2016. Life IAP-Risk Newsletter No 1. http://www.iap-risk.eu/media/files/LIFE_IAP_RISK_Newsletter_1_August_2016.pdf

Lin Shu-Yan, Zhang Qing-Feng, Chen Qi-Xu, 2007. Shade-tolerance of ten species of garden plants. Journal of Northeast Forestry University, 35(7), 32-34.

Liu, D., 1995. Biology and control test of Paratylenchus dianthus. Acta Phytopathologica Sinica, 25(3), 271-274.

Malumphy, C., 2016. Six non-native species of scale insect (Hemiptera: Coccoidea) new for Belgium. Entomologist's Monthly Magazine, 152(4), 245-251. http://www.pemberleybooks.com

Missouri Botanical Garden, 2017. Tropicos database. St. Louis, Missouri, USA: Missouri Botanical Garden.http://www.tropicos.org/

Norcini, J. G., Knox, G. W., 1990. Effect of pruning on the growth inhibiting activity of Sumagic (uniconazole). Journal of Environmental Horticulture, 8(4), 199-204.

North Carolina State University, 2017. Plants. https://plants.ces.ncsu.edu/plants/plant-list

Online Atlas of the British and Irish Flora, 2017. Online Atlas of the British and Irish Flora. http://www.brc.ac.uk/plantatlas/

Orwa C, Mutua A, Kindt R, Jamnadass R, Anthony S, 2009. Agroforestree Database: a tree reference and selection guide, version 4.0. http://www.worldagroforestry.org/output/agroforestree-database

PFAF, 2017. Plants for a Future Database. http://www.pfaf.org/USER/Default.aspx

Randall, R. P., 2007. The introduced flora of Australia and its weed status, CRC for Australian Weed Management.iv + 524 pp.

RenJun, 2013. The Cold Resistance of the Hybrid of Euonymus japonicus and Euonymus kiautschovicus. http://www.dissertationtopic.net/doc/2150204

Schulz B, 2006. Studies of fruit and seed characters of selected Euonymus species. Mitteilungen der Deutschen Dendrologischen Gesellschaft, 91, 127-145.

Stace, C., 1997. New flora of the British Isles, (Ed. 2) : Cambridge University Press.xxvii + 1130 pp.

Sun Xu-Gen, Liu Zhen-Yu, Zhou Cheng-Gang, Qiao Lu-Qin, 1996. Damage of Eutetranychus orientalis to Euonymus japonicus. Plant Protection, 22(5), 29-30.

Tanner R, 2017. Updates from EPPO. [12th meeting Bern Convention Group of Experts on Invasive Alien Species. Madeira, 2017-06-1/3], The European and Mediterranean, Plant Protection Organization. https://rm.coe.int/updates-from-eppo/1680730319

Tawfik, A. A., 2001. Fast and efficient mass-propagation of some ornamental plants using simple water air-flow medium. Acta Horticulturae, (No.548), 611-618.

University of Maryland Extension, 2017. Home & Garden Information Center. https://extension.umd.edu/hgic

Urban Forest Ecosystems Institute, 2017. SelecTree: A Tree Selection Guide. https://selectree.calpoly.edu/

USDA-ARS, 2017. 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, 2017. The PLANTS Database. Baton Rouge, USA: National Plant Data Center.http://plants.usda.gov/

Useful Tropical Plants, 2017. Useful Tropical Plants Database. http://tropical.theferns.info/

Wang, X. G., Levy, K., Mills, N. J., Daane, K. M., 2012. Light brown apple moth in California: a diversity of host plants and indigenous parasitoids. Environmental Entomology, 41(1), 81-90. http://esa.publisher.ingentaconnect.com/content/esa/envent/2012/00000041/00000001/art00009 doi: 10.1603/EN11160

World Agroforestry Centre, 2017. Euonymus japonicus. http://www.worldagroforestry.org/treedb/AFTPDFS/Euonymus_japonicus.PDF

Xue Hui-Wen, 2011. The Research on the Propagation Techniques of Euonymus Japonicus Cv. Microphyllus Butterscotch. Changsha, Hunan, China: http://www.dissertationtopic.net/doc/187462

Yamazaki, K., Takahashi, E., Okabe, M., 1979. Experiments on the practical use of dikegulac for ornamental trees. Effects on retarding hedgerow-tree growth, induction of flower buds in azaleas and abortion of fruit in Japanese holly. Bulletin of the Kanagawa Horticultural Experiment Station, (No. 26), 111-117.

Zieslin, N., Pines, N., 1987. Growth correlations in Euonymus japonicus Thunb. (Celastraceae). Israel Journal of Botany, 36(3), 113-123.

Distribution References

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

DAISIE, 2017. Delivering Alien Invasive Species Inventories for Europe. In: Delivering Alien Invasive Species Inventories for Europe. http://www.europe-aliens.org/

Encyclopedia of Life, 2017. Encyclopedia of Life., http://eol.org/

Flora Europaea, 2017. Euonymus japonicus., http://193.62.154.38/cgi-bin/nph-readbtree.pl/feout?FAMILY_XREF=&GENUS_XREF=Euonymus&SPECIES_XREF=japonic*&TAXON_NAME_XREF=&RANK=

Flora of China Editorial Committee, 2017. 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

GBIF, 2017. Global Biodiversity Information Facility. In: Global Biodiversity Information Facility. http://www.gbif.org/species

Hagiladi A, Raviv M, Ran I, Alzam O, Raviv A, Ben-Jaacov J, 1981. Cultivation of Myrtus and Euonymus spp. in the mountainous regions of Israel. Hassadeh. 61 (10), 1682-1685.

Missouri Botanical Garden, 2017. Tropicos database., St. Louis, Missouri, USA: Missouri Botanical Garden. http://www.tropicos.org/

Orwa C, Mutua A, Kindt R, Jamnadass R, Anthony S, 2009. Agroforestree Database: a tree reference and selection guide, version 4.0., http://www.worldagroforestry.org/output/agroforestree-database

PFAF, 2017. Plants For A Future Database., http://www.pfaf.org/USER/Default.aspx

Randall R P, 2007. The introduced flora of Australia and its weed status. [ed. by Randall R P]. Glen Osmond, Australia: CRC for Australian Weed Management. iv + 524 pp.

USDA-ARS, 2017. 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, 2017. The PLANTS Database. In: The PLANTS Database. Greensboro, North Carolina, USA: National Plant Data Team. https://plants.sc.egov.usda.gov

Links to Websites

Principal Source

Draft datasheet under review

Contributors

08/02/17 Original text by:

Chris Parker, Consultant, UK

Pedro Acevedo-Rodríguez, Department of Botany-Smithsonian NMNH, Washington DC, USA

Distribution Maps