Aleurites moluccanus (candlenut tree)

Pictures

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Picture

Title

Caption

Copyright

Tree habit

Aleurites moluccanus (kukui, kukui nut); tree habit. Hana Highway, Maui., Hawaii. June 18, 2009

©Forest & Kim Starr Images-2009. CC-BY-3.0

Typical habit in a grove

Aleurites moluccanus (kukui, kukui nut); typical yhabit in a grove. Honomanu Hana Highway, Maui, Hawaii, June 18, 2009

©Forest & Kim Starr Images-2009. CC-BY-3.0

Trunk and bark

Aleurites moluccanus (kukui, kukui nut); trunk. Iao, Maui, Hawaii. April 30, 2010

©Forest & Kim Starr Images-2010. CC-BY-3.0

Foliage

Aleurites moluccanus (kukui, kukui nut); foliage. Kula Botanical Garden, Maui. March 07, 2011

©Forest & Kim Starr Images-2011 - CC-BY-3.0

Fruits and foliage

Aleurites moluccanus (kukui, kukui nut); Fruits and foliage. Wahinepee, Maui, Hawaii. August 03, 2002

©Forest & Kim Starr Images-2002 - CC-BY-3.0

Flowers

Aleurites moluccanus (kukui, kukui nut); flowers at Keopuolani Park, Maui. July 03, 2006

©Forest & Kim Starr Images-2006 - CC-BY-3.0

Fruits as beach flotsam

Aleurites moluccanus (kukui, kukui nut); beach flotsam at Kanaha Beach, Maui. March 23, 2004

©Forest & Kim Starr Images-2004 - CC-BY-3.0

Mature Tree

Aleurites moluccanus (kukui, kukui nut); mature tree habit.

©Rafael T. Cadiz

Wildlings

Aleurites moluccanus (kukui, kukui nut); wildlings.

©Rafael T. Cadiz

Wildling

Aleurites moluccanus (kukui, kukui nut); wildling.

©Rafael T. Cadiz

Identity

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Preferred Scientific Name

Aleurites moluccanus (L.) Willd.

Preferred Common Name

candlenut tree

Other Scientific Names

Aleurites ambinux Pers
Aleurites angustifolius Vieill
Aleurites commutata Geiseler
Aleurites cordifolia Steud
Aleurites cordifolius (Gaertn.) Steud.
Aleurites erratica O. Deg., I. Deg. & K. Hummel
Aleurites integrifolia Vieill. ex Guillaumin
Aleurites integrifolius Vieill.
Aleurites javanicus Gand.
Aleurites lanceolatus Blanco
Aleurites lobatus Blanco
Aleurites moluccana (L.) Willd.
Aleurites moluccanus var. aulanii O.Deg. & I.Deg.
Aleurites moluccanus var. floccosa Airy Shaw
Aleurites moluccanus var. katoi O.Deg., I.Deg. & Stone
Aleurites moluccanus var. remyi (Sherff) Stone
Aleurites moluccanus var. serotina O.Deg. & Sherff
Aleurites moluccanus var. serotinus O. Deg. & Sherff
Aleurites remyi Sherff
Aleurites trilobus J. R. Forst. & G. Forst.
Camirium cordifolium Gaertn
Camirium moluccanum (L.) Ktze.
Camirium oleosum Reinw. ex Blume
Camirium oleosum Reinw. ex Müll. Arg.
Dryandra oleifera Lam.
Jatropha moluccana L.
Juglans camirium Lour
Mallotus moluccanus (L.) Müll.Arg.
Manihot moluccana (L.) Crantz
Ricinus dicoccus Roxb.
Rottlera moluccana (L.) Scheff.
Telopea perspicua Sol. ex Seem.

International Common Names

English: balucanat; Belgaum walnut; candle nut tree; candleberry; candlenut; candlenut oil tree; Candle-nut tree; Indian walnut; kukui nut; lumbangtree; Otaheite walnut; varnish tree; varnishtree
Spanish: Arbol de la cera; arbol llorón; avellano; avellano criollo; camirio; nogal de la India; nuez
French: aleurites; bancoulier; Bancoulier des Moluques; noisette; noix; noyer; noyer de bancoul; noyer des Indes; noyer des Moluques; noyer des Moluques
Chinese: shi li
Portuguese: calumbàn; lumbàn; noz da India

Local Common Names

Brazil: nogueira-de-igape; noz-da-índia; saboneteira
Cook Islands: tuitui
Cuba: árbol de la luz; nogal prieto; nogel de jardín
Fiji: lauci; nggerenggere; sikeci; sikeli; sikethi; toto; tuitui; tutui; waiwai
French Polynesia: ama; rama; tahii; tahiri; ti'a'iri; tiaiti; tuitui; tutu'i
Germany: Kerzennussbaum; Lichtnussbaum
Greece/Crete: lerit
Guam: lumbang
Indonesia: anoi; berau; bontalo dudulaa; boyau; buwa kare; kamere; kamiri; kembiri; kemili; kemiling; keminting; kemiri; kemwiri; kereh; komere; kumiri; lana; madang ijo; mi; midi; miri; muncang; nena; nyenga; pidekan; saketa; tanoan; tenu; wiau
Indonesia/Sumatra: kemiri
Italy: Albero della vernice
Micronesia, Federated states of: raguar; sakan; shakan
Niue: tuitui
Palau: sakan
Philippines: biau; kalumban; kami; kapili; lumbang; rumbang
Portugal: calumban; noz da India
Puerto Rico: nogal; nuez de la India; palo de nuez
Samoa: lama
Spain: arbol lloron; avellano; avellano criollo; nogal de la India; nuez
Tonga: tuitui
USA/Hawaii: kuikui; kukui
Vanuatu: kandeltri
Wallis and Futuna Islands: tuitui

EPPO code

ALEMO (Aleurites moluccana)

Trade name

tung

Summary of Invasiveness

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A. moluccanus, native to the Indo-Malaya region, is a distinctive medium-sized tree, recognisable by its silvery green leaves. It has been intentionally introduced, and is subsequently, widespread throughout the tropics. The species has occasionally escaped from cultivation into the wild, most prominently in Hawaii, but also in the Dominican Republic, Florida and sites in New South Wales, Queensland and Christmas Island (University of Queensland, 2011). It is reported to be invasive on American Samoa, the Cook Islands, Brazil, Ecuador, Paraguay, Dominican Republic, Federated States of Micronesia, Fiji, French Polynesia, New Caledonia, Niue, Pitcairn, Samoa, Tonga, Singapore and Mayotte and to be a pest in parts of Australia, including Christmas Island (Kairo et al., 2003; University of Queensland, 2011; PIER, 2013).

Taxonomic Tree

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Domain: Eukaryota
Kingdom: Plantae
Phylum: Spermatophyta
Subphylum: Angiospermae
Class: Dicotyledonae
Order: Euphorbiales
Family: Euphorbiaceae
Genus: Aleurites
Species: Aleurites moluccanus

Notes on Taxonomy and Nomenclature

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Aleurites moluccanus, the 'candlenut tree', is a culturally significant species of tree in the genus Aleurites of the family Euphorbiaceae. The genus contains two species: A. moluccanus (L.) Willd and Aleurites rockinghamensis (Baill.) P.I.Forst. The economically important tung oil tree, A. fordii, which was widely planted in South America and elsewhere during the twentieth century (Arce and Paull, 2008) is now recognized in the genus Vernicia.

The species was described by Carl Linnaeus in his 1753 work Species Plantarum as two species in separate genera: Croton moluccanus and Jatropha moluccana. The two names are now considered synonyms, with the name in Croton being disregarded (Stuppy et al., 1999). When the genus Aleurites was erected by naturalists Johann Reinhold Forster and Georg Forster, they nominated Aleurites triloba to be the type species. This is a junior subjective synonym of A. moluccanus. A range of other synonyms exist for A. moluccanus, as well as several nomina nuda.

The name Aleurites derives from a Greek word meaning floury, in reference to the underside of the leaves. A. moluccanus is morphologically variable, and a number of taxa have been recognised as various ranks, from varieties to separate species. Although none were accepted as valid by Stuppy et al. (1999), listed varieties of the species may include A. moluccanus var. aulanii, A. moluccanus var. floccose, A. moluccanus var. katoi, A. moluccanus var. remyi, A. moluccanus var. serotina and A. moluccanus var. serotinus. Due to its cultural significance and broad geographical range, A. moluccanus is known by a great variety of vernacular names. In English it is generally known as the 'candlenut tree' or 'Indian walnut', or by its Hawaiian name, kukui.

Description

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A. moluccanus is a medium-sized tree, reaching 20 m tall, with a diameter at breast height of up to 90cm. It has a large, spreading crown, and often has irregular branches, frequently spreading wide or drooping downwards to ground level. The tree is distinctive from a distance (Scott and Thomas, 2000) due to the silvery–green colour of the leaves. This pale colour is caused by a thick covering of stellate hairs, which often diminishes as the leaf ages.

The leaves are very distinctively shaped but also quite variable in gross morphology. Young leaves and leaves on lower branches are often three-lobed or five-lobed, while older leaves and those on higher branches, tend to be a simpler triangular or oval shape. They are typically 10–20cm long with wavy margins, and are arranged alternately. Where the leaf stalk joins the blade of the leaf, there are a pair of glands that produce a sweet secretion.

Flowers are in terminal cymes, each about 10–15cm long. Each female flower is surrounded by several smaller male flowers. The female (pistillate) flowers are up to 13mm long, with five separate creamy–white petals. The male (staminate) flowers are similar to the pistillate flowers, but are longer and thinner. The fruit are hard, round drupes, about 5–6 cm long and 5–7 cm wide. The seeds are contained within a tough black shell that resembles that of a walnut; de-husked seeds number approximately 100–120 per kilogram (Elevitch and Manner, 2006).

Plant Type

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Broadleaved
Perennial
Seed propagated
Tree
Woody

Distribution

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A. moluccanus was introduced aboriginally throughout the Pacific Islands and is now a common tree of the Pacific. To the Polynesians it is an important cultural tree, and they transported it across much of South-east Asia and Oceania in prehistoric times as a so-called “canoe plant”. This ancient transportation makes it difficult to distinguish between the natural range of the species and the areas where it has been introduced by man. Therefore, there is some debate as to the limits of its true native range. Its native distribution is, however, reported as centred on the Indo-Malaya region, and probably extends from Myanmar and China throughout the Malay Archipelago, and as far east as French Polynesia (PIER, 2013).

As an introduced species, A. moluccanus is now widespread across the tropics, being found in Bangladesh, Kenya, Uganda, Madagascar, the Comoros, South Africa, Brazil and many islands in the Caribbean (Elevitch and Manner, 2006; Oyen, 2007), and has even been planted on isolated oceanic islands such as Pitcairn and St Helena (Varnham, 2006). In Brazil, it is abundant in southern and south-eastern states, from São Paulo to Rio Grande do Sul (Quintão et al., 2011). In Hawaii, A. moluccanus is widespread, and has been designated as the official state tree (Little and Skolmen, 2003).

Distribution Table

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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.

Last updated: 17 Dec 2021

Continent/Country/Region	Distribution	Origin	First Reported	Invasive	Planted	Reference	Notes
Africa
Comoros	Present	Introduced				Oyen (2007)
Congo, Democratic Republic of the	Present	Introduced			Planted	Oyen (2007)
Gabon	Present	Introduced				ORSTOM (1988)
Kenya	Present	Introduced			Planted	Krisnawati et al. (2011)
Madagascar	Present	Introduced				Oyen (2007)
Mayotte	Present	Introduced		Invasive		PIER (2013) Seebens et al. (2017)
Rwanda	Present	Introduced				Govaerts (2014)
South Africa	Present	Introduced			Planted	Oyen (2007)	KwaZulu-Natal and Mpumalanga
Tanzania	Present	Introduced				Oyen (2007)
Uganda	Present	Introduced			Planted	Oyen (2007)
Asia
Bangladesh	Present	Introduced				Krisnawati et al. (2011)
Brunei	Present	Native				Krisnawati et al. (2011)
Cambodia	Present	Native				Krisnawati et al. (2011) USDA-ARS (2012)
China	Present	Native			Planted	Oyen (2007)
-Fujian	Present	Native				Flora of China Editorial Committee (2014)
-Guangdong	Present	Native				Flora of China Editorial Committee (2014)
-Guangxi	Present	Native				Flora of China Editorial Committee (2014)
-Hainan	Present	Native				Flora of China Editorial Committee (2014)
-Yunnan	Present	Native				Flora of China Editorial Committee (2014)
Hong Kong	Present	Introduced				PIER (2013)
India	Present	Native			Planted	Oyen (2007)
-Assam	Present	Native				USDA-ARS (2012)
-Karnataka	Present	Native				USDA-ARS (2012)
-Kerala	Present	Native				USDA-ARS (2012)
-Maharashtra	Present	Native				USDA-ARS (2012)
-Odisha	Present	Native				USDA-ARS (2012)
-Tamil Nadu	Present	Native				USDA-ARS (2012)
-West Bengal	Present	Native				USDA-ARS (2012)
Indonesia	Present	Native			Planted	Krisnawati et al. (2011) USDA-ARS (2012) Sartiami and Mound (2013) Govaerts (2014)
-Java	Present					Sartiami and Mound (2013)
-Sulawesi	Present	Native				Govaerts (2014)
Japan	Present	Introduced				Krisnawati et al. (2011)
Laos	Present	Native				Krisnawati et al. (2011)
Malaysia	Present	Native			Planted	Krisnawati et al. (2011) USDA-ARS (2012)
-Peninsular Malaysia	Present	Native				Govaerts (2014)
-Sabah	Present	Native				Govaerts (2014)
-Sarawak	Present	Native				Govaerts (2014)
Maldives	Present	Introduced		Invasive		PIER (2013)	Malè Atoll
Myanmar	Present	Native				Krisnawati et al. (2011) USDA-ARS (2012)
Philippines	Present	Native			Planted	USDA-ARS (2012)
Singapore	Present	Introduced		Invasive		PIER (2013)
Sri Lanka	Present	Native				USDA-ARS (2012)
Taiwan	Present	Native			Planted	USDA-ARS (2012)
Thailand	Present	Native				Krisnawati et al. (2011) USDA-ARS (2012)
Vietnam	Present	Native				Krisnawati et al. (2011) USDA-ARS (2012)
North America
Antigua and Barbuda	Present	Introduced				Krisnawati et al. (2011)
Bahamas	Present	Introduced				Krisnawati et al. (2011)
Barbados	Present	Introduced				Krisnawati et al. (2011)
Costa Rica	Present	Introduced				Burger and Huft (1995)
Cuba	Present	Introduced		Invasive	Planted	Krisnawati et al. (2011)	Naturalized
Dominica	Present	Introduced				ICRAF (2012)
Dominican Republic	Present	Introduced		Invasive		Krisnawati et al. (2011)
Grenada	Present	Introduced				Krisnawati et al. (2011)
Guadeloupe	Present	Introduced				Krisnawati et al. (2011)
Haiti	Present	Introduced				Krisnawati et al. (2011)
Honduras	Present	Introduced				Molina (1975)
Jamaica	Present	Introduced			Planted	Krisnawati et al. (2011)
Martinique	Present	Introduced				Krisnawati et al. (2011)
Mexico	Present	Introduced				Pérez et al. (2005)
Montserrat	Present	Introduced				Krisnawati et al. (2011)
Netherlands Antilles	Present	Introduced				Krisnawati et al. (2011)
Puerto Rico	Present	Introduced				Krisnawati et al. (2011)
Saint Kitts and Nevis	Present	Introduced				Krisnawati et al. (2011)
Saint Lucia	Present	Introduced				Krisnawati et al. (2011)
Saint Vincent and the Grenadines	Present	Introduced				Krisnawati et al. (2011)
Trinidad and Tobago	Present	Introduced				Krisnawati et al. (2011)
U.S. Virgin Islands	Present	Introduced				Krisnawati et al. (2011)
United States	Present	Introduced				Krisnawati et al. (2011)
-Florida	Present	Introduced				USDA-ARS (2012)
-Hawaii	Present, Widespread	Introduced			Planted	Oyen (2007)
Oceania
American Samoa	Present	Introduced		Invasive		Space and Flynn (2000)	Reported as an aboriginal introduction and an invasive species on Tau Island, Tutuila Island
Australia	Present	Native				USDA-ARS (2012) Seebens et al. (2017)
-New South Wales	Present					University of Queensland (2011)	Escaped cultivation and recognised as a weed
-Queensland	Present	Native				USDA-ARS (2012)
Christmas Island	Present	Introduced		Invasive		Govaerts (2014)
Cook Islands	Present			Invasive		Space and Flynn (2002) Krisnawati et al. (2011) PIER (2013)	Present on many islands but reported as aboriginal introductions and invasive on Mangaia Island, Ma‘uke Island, Miti'aro Island and Rarotonga Island
Federated States of Micronesia	Present			Invasive		Smith (1981) PIER (2013)	Possibly introduced to some parts of Pohnpei Island where it is invasive. Native to Yap (Waqab) Island
Fiji	Present			Invasive		Smith (1981) Krisnawati et al. (2011) PIER (2013)	Present on many islands but reported as introduced and invasive on Kandavu Island, Koro Island, Lakemba Island, Ovalau Island, Taveuni Island, Vanua Levu Island and Viti Levu Island. Most reported as aboriginal introductions
French Polynesia	Present			Invasive		Krisnawati et al. (2011) PIER (2013)	Reported as native to some islands but introduced and invasive on many. Mainly cited as aboriginal introductions
Guam	Present	Introduced				PIER (2013)	Aboriginal introduction. Cultivated
Kiribati	Present	Native				Krisnawati et al. (2011)
Marshall Islands	Present	Native				Krisnawati et al. (2011)
New Caledonia	Present	Native		Invasive		MacKee (1994) Krisnawati et al. (2011) PIER (2013)	Aboriginal introduction
New Zealand	Present	Native				Oyen (2007)
-Kermadec Islands	Present	Introduced	1836			Seebens et al. (2017)
Niue	Present	Introduced		Invasive		PIER (2013)	Aboriginal introduction
Norfolk Island	Present	Native				Krisnawati et al. (2011)
Palau	Present					PIER (2013)
Papua New Guinea	Present	Native			Planted	Krisnawati et al. (2011) USDA-ARS (2012)
Pitcairn	Present	Introduced		Invasive		PIER (2013)
Samoa	Present	Introduced		Invasive		Space and Flynn (2002) Krisnawati et al. (2011) PIER (2013)	Savai‘i Island. Aboriginal introduction
Solomon Islands	Present	Native				Krisnawati et al. (2011)
Tonga	Present	Introduced		Invasive		PIER (2013) Krisnawati et al. (2011)	Reported as aboriginal introduction and an invasive species on Ha‘apai Islands, Kao Island, Lifuka Island, Tofua Island, ‘Eua Island, Tongatapu Island, Late Island, Niuafo‘ou Island, Tafahi Island, Vava‘u Island
Vanuatu	Present	Native				Krisnawati et al. (2011)
Wallis and Futuna	Present	Introduced				PIER (2013)	Cultivated
South America
Brazil	Present	Introduced				Krisnawati et al. (2011)
-Ceara	Present	Introduced		Invasive		I3N-Brasil (2014)
-Parana	Present	Introduced				Quintão et al. (2011)
-Rio Grande do Sul	Present	Introduced				Quintão et al. (2011)
-Santa Catarina	Present	Introduced				Quintão et al. (2011)
-Sao Paulo	Present	Introduced				Quintão et al. (2011)
Ecuador	Present					CABI (Undated)	Present based on regional distribution.
-Galapagos Islands	Present	Introduced		Invasive		PIER (2013)	San Cristóbal Island
Paraguay	Present	Introduced		Invasive		IABIN (2014)

History of Introduction and Spread

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This species has been widely introduced to South and Central America, the Caribbean, Japan and Africa. In the tropical oceanic islands of the Pacific, A. moluccanus was intentionally introduced as a useful plant by the first Polynesian migrants at least 1000 years ago and is now widely naturalized (Larrue et al., 2010). In the West Indies, this species first appears in herbarium collections made in Puerto Rico in 1885, in Cuba in 1889 and in Guadeloupe in 1892 (US National Herbarium).

Risk of Introduction

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The large seeds of A. moluccanus are poorly suited to long-distance dispersal. Although it is capable of surviving in new areas outside its native range, its rate of unaided spread is generally low, and it “is rarely considered a harmful invasive or pest species” (Elevitch and Manner, 2006). It is, however, widely planted across a very large geographical area, which significantly increases the potential for it to escape into the wild. It is reported to be invasive in the Dominican Republic (Kairo et al., 2003) and to be a pest in parts of Australia, including Christmas Island (University of Queensland, 2011).

Habitat List

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Category	Sub-Category	Habitat	Presence	Status
Terrestrial
Terrestrial	Managed	Managed forests, plantations and orchards	Present, no further details	Productive/non-natural
Terrestrial	Managed	Disturbed areas	Present, no further details	Productive/non-natural
Terrestrial	Managed	Urban / peri-urban areas	Present, no further details	Productive/non-natural
Terrestrial	Natural / Semi-natural	Natural forests	Present, no further details	Natural
Terrestrial	Natural / Semi-natural	Natural forests	Present, no further details	Productive/non-natural
Terrestrial	Natural / Semi-natural	Riverbanks	Present, no further details	Natural
Terrestrial	Natural / Semi-natural	Riverbanks	Present, no further details	Productive/non-natural

Biology and Ecology

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Physiology and Phenology

There is only weak seasonality to flowering and fruiting in A. moluccanus, with plants above 3–4 years old producing flowers and fruit continuously, and with all stages of the fruiting cycle often present on a single tree at the same time. The flowers are, however, more abundant in the spring. It is estimated that individual trees of A. moluccanus can live for 40–60 years (Elevitch and Manner, 2006). There are few pests of A. moluccanus, none which are serious (Krisnawati et al., 2011).

Associations

The tree species associated with A. moluccanus vary across its natural and introduced range. In the Cook Islands, they include the native species Elaeocarpus floridanus and Hernandia moerenhoutiana and the introduced species Cocos nucifera, Morinda citrifolia, Hibiscus tiliaceus and Psidium guajava; in French Polynesia, they include H. tiliaceus and Rhus taitensis on forested slopes, and a wider range of native species on rocky slopes. In Hawaii, the native species associated with A. moluccanus include species of Diospyros and Pisonia, and introduced species include Syzygium cumini, Schinus terebinthifolia and Eucalyptus species (Elevitch and Manner, 2006).

Environmental Requirements

A. moluccanus grows chiefly on well-drained, moist, slightly acidic soils. It can reach elevations of 1200 m (3900 ft.) close to the Equator, but only reaches 700 m (2300 ft.) in Hawaii. It typically grows in mesic environments, but is apparently unaffected by varying patterns of rainfall, especially since it often grows close to streams. It generally grows in areas with a mean annual rainfall range of 640-4290 mm and a mean of 1940 mm (Elevitch and Manner, 2006). However, it can tolerate as little as 200 mm annual rainfall in Indonesia and once fully grown, A. moluccanus tolerates drought reasonably well (Krisnawati et al., 2011). It grows in areas with a mean annual temperature of 19–27°C and is estimated to tolerate temperatures no colder than 8°C (Elevitch and Manner, 2006).

Climate

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Climate	Status	Description
Af - Tropical rainforest climate	Preferred	> 60mm precipitation per month
Am - Tropical monsoon climate	Preferred	Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))
Aw - Tropical wet and dry savanna climate	Preferred	< 60mm precipitation driest month (in winter) and < (100 - [total annual precipitation{mm}/25])

Latitude/Altitude Ranges

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Latitude North (°N)	Latitude South (°S)	Altitude Lower (m)	Altitude Upper (m)
20-25	27-32	0	700

Air Temperature

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Parameter	Lower limit	Upper limit
Absolute minimum temperature (ºC)	8
Mean annual temperature (ºC)		27
Mean maximum temperature of hottest month (ºC)	26	30
Mean minimum temperature of coldest month (ºC)	8	13

Rainfall

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Parameter	Lower limit	Upper limit	Description
Dry season duration	3	5	number of consecutive months with <40 mm rainfall
Mean annual rainfall	640	4290	mm; lower/upper limits

Soil Tolerances

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Soil drainage

free

Soil reaction

acid
alkaline
neutral

Soil texture

heavy
light
medium

Special soil tolerances

infertile
shallow

Impact Summary

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Category	Impact
Cultural/amenity	Positive
Environment (generally)	Negative

Economic Impact

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A. moluccanus provides a number of useful products, although few of economic significance (see Uses).

Environmental Impact

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The direct impact of A. moluccanus on native ecosystems is not well known. Simply by virtue of its abundance, it must reduce the abundances of native species, but few specific interactions have been documented. On Christmas Island, its spread into the natural forest vegetation causes attrition of the forest edges and interrupts the natural succession (University of Queensland, 2011).

Threatened Species

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Threatened Species	Conservation Status	Where Threatened	Mechanism	References
Abutilon sandwicense (greenflower Indian mallow)	CR (IUCN red list: Critically endangered); USA ESA listing as endangered species	Hawaii	Competition - monopolizing resources	US Fish and Wildlife Service (1998)
Isodendrion longifolium (longleaf isodendrion)	EN (IUCN red list: Endangered); USA ESA listing as threatened species	Hawaii	Competition - monopolizing resources	US Fish and Wildlife Service (2011b)
Peucedanum sandwicense (makou)	NatureServe; USA ESA listing as threatened species	Hawaii	Competition - shading	US Fish and Wildlife Service (2011a)
Pritchardia napaliensis	CR (IUCN red list: Critically endangered); USA ESA listing as endangered species	Hawaii	Competition - monopolizing resources	US Fish and Wildlife Service (2010a)
Schiedea hookeri (sprawling schiedea)	CR (IUCN red list: Critically endangered); USA ESA listing as endangered species	Hawaii	Competition - monopolizing resources; Ecosystem change / habitat alteration	US Fish and Wildlife Service (2011c)
Schiedea kealiae (Waianae Range schiedea)	CR (IUCN red list: Critically endangered); USA ESA listing as endangered species	Hawaii	Competition - monopolizing resources	US Fish and Wildlife Service (2010b)
Stenogyne kanehoana (Oahu stenogyne)	CR (IUCN red list: Critically endangered); USA ESA listing as endangered species	Hawaii	Competition - monopolizing resources	US Fish and Wildlife Service (1997)
Urera kaalae	CR (IUCN red list: Critically endangered); USA ESA listing as endangered species	Hawaii	Ecosystem change / habitat alteration; Pest and disease transmission	US Fish and Wildlife Service (2011d)
Wikstroemia villosa.	CR (IUCN red list: Critically endangered)	Hawaii	Competition (unspecified)	US Fish and Wildlife Service (2012a)
Zanthoxylum oahuense (Oahu pricklyash)	VU (IUCN red list: Vulnerable); USA ESA listing as endangered species	Hawaii	Competition (unspecified)	US Fish and Wildlife Service (2012a)

Risk and Impact Factors

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Invasiveness

Proved invasive outside its native range
Has a broad native range
Abundant in its native range
Highly adaptable to different environments
Pioneering in disturbed areas
Tolerant of shade
Benefits from human association (i.e. it is a human commensal)
Long lived
Fast growing
Has high reproductive potential
Has propagules that can remain viable for more than one year
Has high genetic variability

Impact outcomes

Altered trophic level
Ecosystem change/ habitat alteration
Modification of nutrient regime
Modification of successional patterns
Reduced native biodiversity
Soil accretion
Threat to/ loss of native species

Impact mechanisms

Competition - monopolizing resources
Competition - shading
Competition - smothering
Poisoning
Rapid growth

Likelihood of entry/control

Highly likely to be transported internationally deliberately

Uses

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Social Benefit

A. moluccanus has a vast number of uses, and almost every part of the plant can be used for some purpose. The living plant is used as an ornamental tree, or as a living fence or windbreak. The bark is used to make an infusion that preserves fishing nets. The wood is not resistant to rot but can be used as an effective substrate for growing mushrooms, as low-quality fuel, or for making floats or short-lived canoes. The sap has been used to waterproof cloth. The leaves have been used in poultices, or in leis (garlands), either with or without the flowers (Elevitch and Manner, 2006).

Elevitch and Manner (2006) however, highlight that the majority of uses of A. moluccanus derive from their seeds. They are edible in small quantities when cooked, but are otherwise generally toxic. However, there is a variety growing in Vanuatu that appears to lack the toxic effect. The empty seed shells can be used to make jewellery or as part of a lei, or can be burned to produce soot that can be used in tattooing and dyeing. An oil can be extracted from the seeds which is used for protecting cotton bolls from insect attack, as a laxative or general folk remedy, for preserving surfboards, for waterproofing paper, in making varnishes and paints, or which can be burnt for illumination. This traditional use explains the tree’s name in Polynesian languages, which comes from a root meaning ‘light’.

Economic Value

The tree’s chief commercial product is probably a soap substitute produced from the seed oil. Once the oil has been harvested, the remaining seed cake can be used for animal fodder or as fertiliser.

Uses List

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Animal feed, fodder, forage

Fodder/animal feed

Environmental

Agroforestry
Amenity
Boundary, barrier or support
Windbreak

Fuels

Fuelwood

General

Ornamental
Sociocultural value

Human food and beverage

Fruits
Oil/fat
Seeds
Spices and culinary herbs

Materials

Bark products
Cosmetics
Dyestuffs
Essential oils
Lipids
Oils
Poisonous to mammals
Resins
Wood/timber

Medicinal, pharmaceutical

Source of medicine/pharmaceutical
Traditional/folklore

Wood Products

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Boats

Sawn or hewn building timbers

For light construction

Woodware

Industrial and domestic woodware
Matches

Similarities to Other Species/Conditions

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The related species Aleurites trisperma resembles A. moluccanus, but has entire (unlobed) leaves, and its three-sided fruits have prominent ridges (Elevitch and Manner, 2006).

Prevention and Control

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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.

Where required, populations of A. moluccanus can be controlled by felling and treating the stumps with glyphosate to prevent the tree from re-sprouting. Seedlings can be sprayed with Roundup [glyphosate] plus Pulse in clean water and saplings and trees either cut stump treated or stem injected with Roundup in clean water or basal bark treated with Garlon 600 [triclopyr] in diesel oil (Swarbrick, 1997).

The tree is a conspicuous and distinctive species that is unlikely to be overlooked or misidentified.

Gaps in Knowledge/Research Needs

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The ecological impact of A. moluccanus on native habitats requires further research. Although it is generally considered a beneficial tree, it may have detrimental effects that have not yet come to light. Compared to the relative depth of study afforded to the species in Hawaii, little is known about its spread in other areas, such as the Dominican Republic and many other countries, where it is reported to be invasive.

References

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Aguilar RH, 1917. A comparison of linseed oil and lumbang oil as paint vehicles. Philipinne Journal of Science, 12(5):235

Alambat CF, 1971. Pointers on How to Grow Lumbang Trees. Forests and Farms, Philipinnes, 4-20

Arce RKM, Paull RE, 2008. Aleurites moluccana. In: The encyclopedia of fruit and nuts [ed. by Janick, J. \Paull, R.]. Wallingford, UK: CAB International, 363-364 pp

Bartolome A, Logarita E, 1981. Methods of Stratification on Pili and Lumbang Seeds in nursery production. BS Thesis (Unpublished). Lambunao, Iloilo, Philippines: INCA

Browne FG, 1968. Pests and diseases of forest plantation trees: an annotated list of the principal species occurring in the British Commonwealth. Clarendon Press, Oxford University Press, Oxford

Burger WC, Huft MJ, 1995. Family 113. Euphorbiaceae. Fieldiana, 36:1-169. [Flora Costaricensis.]

Chong KY, Tan HTW, Corlett RT, 2009. A checklist of the total vascular plant flora of Singapore: native, naturalised and cultivated species. Singapore: Raffles Museum of Biodiversity Research, National University of Singapore, 273 pp. http://lkcnhm.nus.edu.sg/nus/pdf/PUBLICATION/LKCNH%20Museum%20Books/LKCNHM%20Books/flora_of_singapore_tc.pdf

Dayan MP, Constantino CH, 1990. Lumbang (Aleurites moluccana (L) Willd.). RISE Compilation (Research Information Series on Ecosystems). College, Laguna, Philippines: ERDB, DENR

Eakle TW, Garcia AS, 1977. Hastening the germination of lumbang (Aleurites moluccana (L.) Willd.) seeds. Sylvatrop, Philipinne Forest Research Journal, 2(4):291-295; 7 ref

Elevitch CR, Manner HI, 2006. Aleurites moluccana (kukui). In: Traditional trees of Pacific islands: their culture, environment and use [ed. by Elevitch, C. R.]. Holualoa, Hawaii, USA: Permanent Agriculture Resources, 41-55 pp. http://www.agroforestry.net/tti/Aleurites-kukui.pdf

Flora of China Editorial Committee, 2014. 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

Gordon DR, Onderdonk DA, Fox AM, Stocker RK, Gantz C, 2008. Predicting invasive plants in Florida using the Australian weed risk assessment. Invasive Plant Science and Management, 1(2):178-195. http://www.wssa.net

Govaerts R, 2014. Family Euphorbiaceae - World Checklist of Euphorbiaceae. London, UK: Royal Botanic Gardens, Kew. http://apps.kew.org/wcsp/

Guzman E de, Umali RM, Sotalbo ED, 1986. Guide to Philippine flora and fauna. Natural Resources Management Center, Ministry of Natural Resources and University of the Philippine. Manila, Philippines: JMC Press Inc

I3N-Brasil, 2014. Base de dados nacional de espécies exóticas invasora (National database of exotic invasive species). Florianópolis - SC, Brazil: I3N Brasil, Instituto Hórus de Desenvolvimento e Conservação Ambiental. http://i3n.institutohorus.org.br

IABIN, 2014. Inter-American Biodiversity Information Network (IABIN). Red de Informacion sobre especies invasoras ([English title not available]). http://www.oas.org/en/sedi/dsd/iabin/

ICRAF, 2012. World Agroforestry Centre: AgroForestryTree Database. Nairobi, Kenya: International Centre for Research in Agroforestry (ICRAF). http://www.worldagroforestrycentre.org

Jacalne DV, 1955. Summary of some germination studies in the Divsion of Forestry. Investigation. Philipinne Journal of Forestry, 11(3 and 4)

Kairo M, Ali B, Cheesman O, Haysom K, Murphy S, 2003. Invasive species threats in the Caribbean region. Report to the Nature Conservancy. Curepe, Trinidad and Tobago: CAB International, 132 pp. http://www.issg.org/database/species/reference_files/Kairo%20et%20al,%202003.pdf

Krisnawati H, Kallio MH, Kanninen M, 2011. Aleurites moluccana (L.) Willd.: ecology, silviculture and productivity. Jakarta, Indonesia: Center for International Forestry Research (CIFOR), vi + 9 pp. http://www.cifor.org/nc/online-library/browse/view-publication/publication/3393.html

Larrue S, Meyer JY, Chiron T, 2010. Anthropogenic vegetation contributions to Polynesia's social heritage: the legacy of candlenut tree (Aleurites moluccana) forests and Bamboo (Schizostachyum glaucifolium) groves on the island of Tahiti. Economic Botany, 64(4):329-339. http://www.springerlink.com/content/4407k53815h75744/

Little, EL, Skolmen RG, 2003. Common forest trees of Hawaii (native and introduced), Agriculture Handbook 679. Honolulu, HI: University of Hawaii at Manoa

Mabberley DJ, 1997. The plant-book: a portable dictionary of the vascular plants. Ed. 2: xvi + 858 pp. Cambridge, UK: Cambridge University Press

MacKee HS, 1994. Catalogue of introduced and cultivated plants in New Caledonia. (Catalogue des plantes introduites et cultivées en Nouvelle-Calédonie.) Paris, France: Muséum National d'Histoire Naturelle, unpaginated

Molina RA, 1975. Enumeration of the plants of Honduras. (Enumeración de las plantas de Honduras) Ceiba, 19(1):1-118

ORSTOM, 1988. List of vascular plants of Gabon

Oyen LPA, 2007. Aleuritia moluccensis (L.) Willd. In: Vegetable oils. Plant Resources of Tropical Africa, 14 [ed. by Vossen, H. A. M. van der \Mkamilo, G.]. Wageningen, Netherlands: PROTA Foundation

Pérez J, Sousa Sánchez LAM, Hanan-Alipi AM, Chiang Cabrera F, Tenorio PL, 2005. Terrestrial vegetation. (Vegetación terrestre.) In: Biodiversidad [ed. by Bueno, J. \Álvarez, F. \Santiago, S.]. Tabasco, Mexico: CONABIO-UNAM, 65-110

PIER, 2013. Pacific Islands Ecosystems at Risk. Honolulu, Hawaii, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html

Quintão NLM, Meyre-Silva C, Silva GF, Antonialli CS, Rocha LW, Lucinda-Silva RM, Malheiros A, Souza MM, Cechinel Filho V, Bresolin TMB, 2011. Aleurites moluccana (L.) Willd. leaves: mechanical antinociceptive properties of a standardized dried extract and its chemical markers. Evidence-based Complementary and Alternative Medicine, 2011:Article ID 179890. http://www.hindawi.com/journals/ecam/2011/179890/

Scott S, Thomas C, 2000. Candlenut (kukui) (Aleurites moluccana). In: Poisonous plants of paradise: first aid and medical treatment of injuries from Hawaii's plant [ed. by Scott, S. \Thomas, C.]. Honolulu, Hawaii, USA: University of Hawaii Press, 25-29 pp

Smith AC, 1981. Flora Vitiensis Nova: A new flora of Fiji. Lawai, Kauai, Hawaii. National Tropical Botanical Garden, 2:290

Space J, Flynn T, 2002. Report to the Government of the Cook Islands on Invasive Plant Species of Environmental Concern. Honolulu, USA: USDA Forest Service, Pacific Southwest Research Station, Institute of Pacific Islands Forestry, 146

Space JC, Flynn T, 2000. Observations on invasive plant species in American Samoa. USDA Forest Service, Honolulu, 51

Space JC, Flynn T, 2001. Report to the Kingdom of Tonga on invasive plant species of environmental concern. Hawaii, USA: USDA Forest Service, Pacific Southwest Research Station, Institute of Pacific Islands Forestry, 78 pp

Space JC, Flynn T, 2002a. Report to the Government of Samoa on invasive plant species of environmental concern. Honolulu, USA: USDA Forest Service, 83 pp

Stuppy W, Welzen PCvan, Klinratana P, Posa MCT, 1999. Revision of the genera Aleurites, Reutealis and Vernicia (Euphorbiaceae). Blumea, 44(1):73-98

Swarbrick JT, 1997. Environmental weeds and exotic plants on Christmas Island, Indian Ocean. Report to Parks Australia. J.T. Swarbrick, Weed Science Consultancy, 131 pp

University of Queensland, 2011. Candle nut, Aleurites moluccana. Weeds of Australia: Biosecurity Queensland Edition. Queensland, Australia: University of Queensland. http://keyserver.lucidcentral.org/weeds/data/03030800-0b07-490a-8d04-0605030c0f01/media/Html/Aleurites_moluccana.htm

US Fish and Wildlife Service, 1997. Recovery Plan for the Maui Plant Cluster. In: Recovery Plan for the Maui Plant Cluster : US Fish and Wildlife Service.130 pp. + appendices.

US Fish and Wildlife Service, 1998. Recovery Plan for Oahu Plants. In: Recovery Plan for Oahu Plants : US Fish and Wildlife Service.207 pp. + appendices.

US Fish and Wildlife Service, 2010. 5-Year Review, Short Form Summary: Species Reviewed: Pritchardia napaliensis (loulu palm). In: 5-Year Review, Short Form Summary: Species Reviewed: Pritchardia napaliensis (loulu palm) : US Fish and Wildlife Service.10 pp.

US Fish and Wildlife Service, 2010. 5-Year Review, Short Form Summary: Species Reviewed: Schiedea kealiae (ma oli oli). In: 5-Year Review, Short Form Summary: Species Reviewed: Schiedea kealiae (ma oli oli) : US Fish and Wildlife Service.6 pp.

US Fish and Wildlife Service, 2011. Endangered and Threatened Wildlife and Plants; Listing 23 Species on Oahu as Endangered and Designating Critical Habitat for 124 Species. In: Federal Register , 76(148) : US Fish and Wildlife Service.46362-46593. http://www.gpo.gov/fdsys/pkg/FR-2011-08-02/pdf/2011-17162.pdf

US Fish and Wildlife Service, 2011. Isodendrion longifolium (aupaka). 5-Year Review: Summary and Evaluation. In: Isodendrion longifolium (aupaka). 5-Year Review: Summary and Evaluation : US Fish and Wildlife Service.28 pp. http://ecos.fws.gov/docs/five_year_review/doc3809.pdf

US Fish and Wildlife Service, 2011. Schiedea hookeri (no common name). 5-Year Review: Summary and Evaluation. In: Schiedea hookeri (no common name). 5-Year Review: Summary and Evaluation : US Fish and Wildlife Service.20 pp.

US Fish and Wildlife Service, 2011. Urera kaalae (opuhe). 5-Year Review: Summary and Evaluation. In: Urera kaalae (opuhe). 5-Year Review: Summary and Evaluation : US Fish and Wildlife Service.19 pp.

US Fish and Wildlife Service, 2012. Endangered and Threatened Wildlife and Plants; Endangered Status for 23 Species on Oahu and Designation of Critical Habitat for 124 Species; Final Rule. In: Federal Register , 77(181) : US Fish and Wildlife Service.57648-57862. https://www.gpo.gov/fdsys/pkg/FR-2012-09-18/pdf/2012-19561.pdf

USDA-ARS, 2012. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysearch.aspx

Varnham K, 2006. Non-native species in UK Overseas Territories: a review. JNCC Report, 372. Peterborough, UK: Joint Nature Conservation Committee. http://www.jncc.gov.uk/page-3660

Weidelt HJ, Agpaoa A, Endangan D, Festins S, Gumayagay J, Hoenninger T, Seeber G, Unkel K, eds, 1975. Manual of reforestation and erosion control for the Philippines. Schriftenreihe der GTZ, Deutsche Gesellschaft fur Technische Zusammenarbeit, German Federal Republic, No. 22, 569 pp

Distribution References

Burger W, Huft M, 1995. Flora Costaricensis. Family No. 113 Euphorbiaceae. In: Fieldiana, Botany, v + 169 pp.

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

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

Flora of China Editorial Committee, 2014. 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

Govaerts R, 2014. Family Euphorbiaceae - World Checklist of Euphorbiaceae., London, UK: Royal Botanic Gardens, Kew. http://apps.kew.org/wcsp/

I3N-Brasil, 2014. National database of exotic invasive species. (Base de dados nacional de espécies exóticas invasora)., Florianópolis - SC, Brazil: I3N Brasil, Instituto Hórus de Desenvolvimento e Conservação Ambiental. http://i3n.institutohorus.org.br

IABIN, 2014. [English title not available]. (Red de Informacion sobre especies invasoras)., Inter-American Biodiversity Information Network (IABIN). http://www.oas.org/en/sedi/dsd/iabin/

ICRAF, 2012. World Agroforestry Centre: AgroForestryTree Database., Nairobi, Kenya: International Centre for Research in Agroforestry (ICRAF). http://www.worldagroforestrycentre.org

Krisnawati H, Kallio M H, Kanninen M, 2011. Aleurites moluccana (L.) Willd.: ecology, silviculture and productivity. In: Aleurites moluccana (L.) Willd.: ecology, silviculture and productivity, Jakarta, Indonesia: Center for International Forestry Research (CIFOR). vi + 9 pp. http://www.cifor.org/nc/online-library/browse/view-publication/publication/3393.html

MacKee H S, 1994. Catalogue des plantes introduites et cultivées en Nouvelle-Calédonie. Paris, France: Muséum National d'Histoire Naturelle. unpaginated.

Molina R A, 1975. Enumeration of the plants of Honduras. (Enumeración de las plantas de Honduras). Ceiba. 19 (1), 1-118.

ORSTOM, 1988. List of vascular plants of Gabon.,

Oyen LPA, 2007. (Aleuritia moluccensis (L.) Willd). In: Vegetable oils. Plant Resources of Tropical Africa, 14 [ed. by Vossen HAM, van der Mkamilo G]. Wageningen, Netherlands: PROTA Foundation.

Pérez J, Sousa Sánchez LAM, Hanan-Alipi AM, Chiang Cabrera F, Tenorio PL, 2005. Terrestrial vegetation. (Vegetación terrestre). In: Biodiversidad, [ed. by Bueno J, Álvarez F, Santiago S]. Tabasco, Mexico: CONABIO-UNAM. 65-110.

PIER, 2013. Pacific Islands Ecosystems at Risk., Honolulu, Hawaii, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html

Quintão N L M, Meyre-Silva C, Silva G F, Antonialli C S, Rocha L W, Lucinda-Silva R M, Malheiros A, Souza M M, Cechinel Filho V, Bresolin T M B, 2011. Aleurites moluccana (L.) Willd. leaves: mechanical antinociceptive properties of a standardized dried extract and its chemical markers. Evidence-based Complementary and Alternative Medicine. Article ID 179890. http://www.hindawi.com/journals/ecam/2011/179890/ DOI:10.1155/2011/179890

Sartiami D, Mound L A, 2013. Identification of the terebrantian thrips (Insecta, Thysanoptera) associated with cultivated plants in Java, Indonesia. ZooKeys. 1-21. http://www.pensoft.net/journals/zookeys/article/5455/identification-of-the-terebrantian-thrips-insecta-thysanoptera-associated-with-cultivated-plants-in-java-indonesia

Seebens H, Blackburn T M, Dyer E E, Genovesi P, Hulme P E, Jeschke J M, Pagad S, Pyšek P, Winter M, Arianoutsou M, Bacher S, Blasius B, Brundu G, Capinha C, Celesti-Grapow L, Dawson W, Dullinger S, Fuentes N, Jäger H, Kartesz J, Kenis M, Kreft H, Kühn I, Lenzner B, Liebhold A, Mosena A (et al), 2017. No saturation in the accumulation of alien species worldwide. Nature Communications. 8 (2), 14435. http://www.nature.com/articles/ncomms14435

Smith AC, 1981. Flora Vitiensis Nova: A new flora of Fiji. Lawai, Kauai, Hawaii., 2 National Tropical Botanical Garden. 290.

Space J, Flynn T, 2002. Report to the Government of the Cook Islands on Invasive Plant Species of Environmental Concern., Honolulu, USA: USDA Forest Service, Pacific Southwest Research Station, Institute of Pacific Islands Forestry. 146.

Space JC, Flynn T, 2000. Observations on invasive plant species in American Samoa., Honolulu, USDA Forest Service. 51.

University of Queensland, 2011. Candle nut, Aleurites moluccana. In: Weeds of Australia: Biosecurity Queensland Edition, Queensland, Australia: University of Queensland. http://keyserver.lucidcentral.org/weeds/data/03030800-0b07-490a-8d04-0605030c0f01/media/Html/Aleurites_moluccana.htm

USDA-ARS, 2012. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysimple.aspx

Contributors

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18/06/14 Updated by:

Julissa Rojas-Sandoval, Department of Botany-Smithsonian NMNH, Washington DC, USA

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

03/01/13 Original text by:

Christopher J. Dixon, University of Oxford, Department of Plant Sciences, South Parks Road, Oxford, OX1 3RB, UK

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Aleurites moluccanus (candlenut tree)

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