Datasheet

Enhanced

8 May 2017

Morinda citrifolia (Indian mulberry)

Author: J Rojas-SandovalAuthors Info & Affiliations

Publication: CABI Compendium

34854

https://doi.org/10.1079/cabicompendium.34854

Datasheet Types: Invasive species, Tree, Host plant, Crop

Abstract

This datasheet on Morinda citrifolia covers Identity, Overview, Associated Diseases, Pests or Pathogens, Distribution, Dispersal, Biology & Ecology, Environmental Requirements, Natural Enemies, Impacts, Uses, Prevention/Control, Management, Genetics and Breeding, Food Quality, Food Safety, Economics, Further Information.

Identity

Preferred Scientific Name: Morinda citrifolia L.
Preferred Common Name: Indian mulberry
Other Scientific Names: Belicea hoffimannioides Lundell; Morinda aspera Wight & Arn.; Morinda asperula Standl.; Morinda bracteata Roxb.; Morinda chachuca Buch.-Ham.; Morinda elliptica (Hook.f.) Ridl.; Morinda ligulata Blanco; Morinda litoralis Blanco; Morinda macrophylla Desf.; Morinda mudia Buch.-Ham.; Morinda multiflora Roxb.; Morinda nodosa Buch.-Ham.; Morinda quadrangularis G.Don; Morinda stenophylla Spreng.; Morinda teysmanniana Miq.; Morinda tomentosa B.Heyne ex Roth; Morinda zollingeriana Miq.; Platanocephalus orientalis Crantz; Samama citrifolia (L.) Kuntze; Sarcocephalus leichhardtii F.Muell.
International Common Names: English
canary-wood
cheesefruit
great morinda; Spanish
mora de la India
morinda
noni
ruibarbo Caribe (Mexico); French
bois douleur
morinde; Chinese
hai bin mu ba ji
Local Common Names: Australia
canary wood
rotten cheesefruit; Brazil
pau-azeitona; Cambodia
nhoër srôk
nhoër thôm'; Cuba
árbol de queso
mora de la India
noni; Dominican Republic
bagá
buñuela
fruta del diablo
manzana de puerto rico
manzanilla
nigua
piña de puerco
piñecla
piñuela; Haiti
bois bouleur
fromagier; Indonesia
bengkudu
cangkudu
mengkudu; Jamaica
hog apple; Laos
nhoo baanz; Lesser Antilles
bilimbi
feuille froide
jumbie breadfruit
pain killer
pomme de singe
pomme-macaque
rhubarbe caraïbe; Malaysia
mengkudu besar
mengkudu jantan; Philippines
apatot
bankoro
tumbong-aso; Puerto Rico
gardenia hedionda
noni; Thailand
yo ban
EPPO code: MOJCI (Morinda citrifolia)

Pictures

Habit
Morinda citrifolia (Indian mulberry); habit. Waianapanapa, Maui, Hawaii, USA. April 2006.
©Forest & Kim Starr - CC BY 4.0

Habit
Morinda citrifolia (Indian mulberry); habit. LaPerouse, Maui, Hawaii, USA. December 2004.
©Forest & Kim Starr - CC BY 4.0

Branches and foliage
Morinda citrifolia (Indian mulberry); branches and foliage. Waianapanapa State Park Hana, Maui, Hawaii, USA. March 2007.
©Forest & Kim Starr - CC BY 4.0

Leaves
Morinda citrifolia (Indian mulberry); leaves and flowers. Hoolawa Farms, Maui, Hawaii, USA. November 2006.
©Forest & Kim Starr- 2006 - CC BY 4.0

Fruiting habit.
Morinda citrifolia (Indian mulberry); fruiting habit. Waihee Coastal Preserve, Maui, Hawaii, USA. June 2016.
©Forest & Kim Starr - CC BY 4.0

Flowers, fruits and foliage
Morinda citrifolia (Indian mulberry); flowers, fruits and foliage. Waihee Coastal Preserve, Maui, Hawaii, USA. June 2016.
©Forest & Kim Starr - CC BY 4.0

Flowers and fruit
Morinda citrifolia (Indian mulberry); flowers and unripe fruit. Waianapanapa, Maui, Hawaii, USA. April 2006.
©Forest & Kim Starr - CC BY 4.0

Flowers and fruit
Morinda citrifolia (Indian mulberry); flowers and ripening fruit. LaPerouse, Maui, Hawaii, USA. April 2004.
©Forest & Kim Starr - CC BY 4.0

Flowers
Morinda citrifolia (Indian mulberry); flowers. Waihee Coastal Preserve, Maui, Hawaii, USA. June 2016.
©Forest & Kim Starr - CC BY 4.0

Ripe fruit
Morinda citrifolia (Indian mulberry); ripe fruit, showing seed and pulp. Waianapanapa State Park Hana, Maui, Hawaii, USA. March 2007.
©Forest & Kim Starr - CC BY 4.0

Seeds and pulp
Morinda citrifolia (Indian mulberry); seeds and pulp. Waianapanapa State Park Hana, Maui, Hawaii, USA. March 2007.
©Forest & Kim Starr - CC BY 4.0

Potted plants
Morinda citrifolia (Indian mulberry); in pots, for sale. Hoolawa Farms, Maui, Hawaii, USA. November 2006.
©Forest & Kim Starr - CC BY 4.0

Overview

Morinda citrifolia is a small, evergreen tree or shrub of approximately 3–10 m at maturity, cultivated widely throughout the tropics, mainly for its fruits. It is a significant source of traditional medicines, dyes and food for indigenous societies and a popular herbal dietary supplement. It is native from South-east Asia (Indonesia) to Australia. It is often raised from seed, but it is possible to air layer or raise plants from cuttings. There has been little formal breeding work or evaluation of genetic resources. The fruits are harvested by hand, usually before fully ripe when they smell of rancid cheese. The fruit is primarily used, but all parts of the plant including seeds, leaves, bark and roots are used in traditional medicine. It is used as a tonic, but also for complaints involving digestive, intestinal, respiratory and immune systems. The juice is in great demand and other products include a fruit powder.

Summary of Invasiveness

M. citrifolia is a shrub or small tree that in recent years has attained significant economic importance worldwide due to the great variety of health and cosmetic products made from its leaves and fruits. Consequently, it has been extensively introduced in cultivation and can be found cultivated and naturalized across tropical and subtropical regions of the world (Groenendijk, 1991; Nelson, 2006; Orwa et al., 2009; USDA-ARS, 2017). Currently, M. citrifolia is listed as invasive in Cuba, Costa Rica, the Dominican Republic and Haiti (Kairo et al., 2003; Chacon and Saborio, 2012, Oviedo Prieto et al., 2012).

M. citrifolia is a species that can tolerate and thrive in very harsh conditions. It is well adapted to a wide range of environmental conditions and soil types. It can grow in infertile, acidic, and alkaline soils, and in areas with climates ranging from very dry to very wet. It is also tolerant to fire, waterlogging, wind, shaded conditions (>80% shade) and salt spray (Francis, 2004; Nelson, 2006; PROTA, 2017). For instance, M. citrifolia is one of the first plants to colonize harsh waste areas or lava flows on islands across the Pacific region and is also one of the few species that can thrive beneath the canopy of the allelopathic tree Casuarina equisetifolia (Nelson, 2006). Additionally, M. citrifolia has a deep taproot and an extensive and aggressive root system and once established it is very persistent and difficult to eradicate. Seeds have a distinct air chamber and can retain viability even after floating in water for months, facilitating the wide distribution and occurrence of this species on many seashores worldwide (Nelson, 2006).

Taxonomic Tree

Notes on Taxonomy and Nomenclature

Rubiaceae is a family of flowering plants comprising 611 genera and 13,150 species of herbs, shrubs, trees and lianas distributed worldwide but largely tropical, especially diverse in Madagascar and the Andes (Stevens, 2012). The genus Morinda includes about 100 species of climbing shrubs, erect shrubs, or small trees (The Plant List, 2013).

There are forms of M. citrifolia previously recognized as botanical varieties (M. citrifolia var. citrifolia, M. citrifolia var. bracteata and M. citrifolia var. elliptica) and one cultivar (M. citrifolia cultivar Potteri) (Nelson, 2006; The Plant List, 2013), but now regarded as synonyms. The former var. bracteata is a small fruited variety with conspicuous bracts subtending the fruit. It is found in Indonesia and other parts of the region between the Indian and Pacific oceans. It is cultivated in some locations. The former var. potteri is an ornamental, small-fruited, narrow-leafed type with green-and-white leaf variegation, which is distributed throughout the Pacific (Janick and Paull, 2008).

A proposal was been made to conserve the name Morinda citrifolia with a conserved type (Razafimandimbison et al., 2011). If this proposal had been rejected the name would have been applied to the species known as M. coreia and the species presently known as M. citrifolia would have to be renamed M. nodosa, which would cause confusion in this widely cultivated species.

Plant Type

Seed propagated

Tree

Vegetatively propagated

Broadleaved

Perennial

Shrub

Description

Morinda citrifolia is a small tree or large evergreen shrub approximately 3–10 m in height at maturity and 15 cm or more in stem diameter. The plant sometimes finds support on other plants as a liana. The sapwood is soft and yellow-brown and the bark relatively smooth to slightly rough and grey or light brown. The light green, four-angled twigs have opposite, pinnately veined, glossy leaves attached by stout petioles, 1.5–2 cm long. Stipules are connate or distinct, 10–12 mm long, the apex entire or two- to three-lobed. The membranous, glabrous leaf blades range from elliptic to elliptic-ovate and range in size from 20 to 45 cm long and 7–25 cm wide. The tubular flowers are perfect, with about 75–90 in ovoid to globose heads. Peduncles are 10–30 mm long; the calyx a truncated rim. The corolla is white, five lobed, with the tube greenish white, 7–9 mm long and lobes oblong-deltate, approximately 7 mm long. There are five stamens, scarcely exserted and the style is about 15 mm long. Fruit (a syncarp) are yellowish white and fleshy, 5–14 cm long, about 3–7.5 cm in diameter, soft and fetid when ripe. Seeds are brown, about 4–9 mm long and have a distinct air chamber. The plant has a rooting habit similar to citrus and coffee, with an extensive lateral root system and a deep taproot (Janick and Paull, 2008).

Distribution

M. citrifolia is native to tropical and subtropical Asia and Australia and now has a pantropical distribution (Govaerts, 2017; USDA-ARS, 2017), occurring roughly between latitudes 19°N and S. The Indo-Pacific distribution includes Eastern Polynesia (e.g. Hawaii, the Line Islands, Marquesas, Society Islands, Australs, Tuamotus, Pitcairn and the Cook islands), Melanesia (e.g. Fiji, Vanuatu, New Guinea, New Caledonia and the Solomon Islands), Western Polynesia (e.g. Samoa, Tonga, Niue, ‘Uvea/Futuna, Rotuma and Tuvalu) and Micronesia (e.g. Pohnpei, Guam, Chuuk, Palau, the Marshall Islands and the Northern Marianas), Indonesia, Australia and South-east Asia. The species has also become naturalized on the open shores of Central and South America (from Mexico to Panama, Venezuela and Surinam) and on many islands of the West Indies, the Bahamas, Bermuda, the Florida Keys and parts of Africa (Janick and Paull, 2008).

A study of nucleotide sequence data by Razafimandimbison et al. (2010) suggests a Micronesian origin for M. citrifolia. Large fruited var. citrifolia may have been present in the Pacific before the arrival of Micronesian and Polynesian ancestors from South East Asia.

Distribution Map

Distribution Table

History of Introduction and Spread

M. citrifolia may have been distributed by man and carried westwards into the Indian Ocean by sea currents, reaching the Seychelles, and similarly into the Pacific between 30°N and 30°S latitude, reaching the Marquesas and Easter Island. It is present throughout South-East Asia both wild and cultivated (Groenendijk, 1991). In Hawaii, it was introduced in 1941 from Fiji and now it can be found planted and naturalized (Little and Skolmen, 2003).

M. citrifolia was widely spread across the Pacific by native peoples beginning 2000 years ago (Smith, 2002), and later in the late 1700s by Europeans exploring the islands of the South Pacific. European explorers noticed that this species was widely used by the native people across Pacific islands and for example, Captain Cook in his journals mentioned the use of “noni” for food and medicinally by island natives (Francis, 2004; Nelson, 2006).

Risk of Introduction

The likelihood of new introductions of M. citrifolia is very high. Worldwide, this species has attained significant economic importance due to the great variety of health and cosmetic products made from its leaves and fruits (Nelson, 2006; USDA-ARS, 2017). Consequently, it has been extensively introduced and in recent years many countries in Asia, Africa and America have approved its cultivation.

Means of Movement and Dispersal

M. citrifolia spreads by seeds (Francis, 2004; Orwa et al., 2009). In cultivation, it can also be propagated vegetatively by cuttings made from stems and roots (Nelson, 2006).

Natural Dispersal (Non-Biotic)

Seeds can be dispersed floating in water. Seeds have a distinct air chamber and can retain viability even after months floating in water (e.g., ocean currents and/or streams and rivers), until their deposition on suitable substrates (Nelson, 2006).

Vector Transmission (Biotic)

Seeds are dispersed by bats, birds, and other mammals (Francis, 2004; Orwa et al., 2009).

Intentional Introduction

M. citrifolia has been widely introduced across tropical and subtropical regions of the world mainly for its fruits and medicinal uses (Nelson, 2006; Govaerts, 2017; USDA-ARS, 2017).

Pathway Causes

Pathway cause	Notes	Long distance	Local	References
Crop production (pathway cause)	Cultivated for its fruits and leaves	Yes	Yes	Orwa et al. (2009)
Disturbance (pathway cause)	Naturalized in disturbed areas near villages	Yes	Yes	Nelson (2006)
Food (pathway cause)	Fruits/ Leaves consumed as vegetable	Yes	Yes	Nelson (2006)
Forage (pathway cause)	Fruits used to feed pigs	Yes	Yes	Francis (2004)
Habitat restoration and improvement (pathway cause)	Planted for control coastal erosion		Yes	Francis (2004)
Hedges and windbreaks (pathway cause)	Windbreak and shade tree	Yes	Yes	Francis (2004)
Horticulture (pathway cause)		Yes	Yes	Francis (2004)
Internet sales (pathway cause)	Seeds and fruits sold online	Yes	Yes
Medicinal use (pathway cause)	Widely used for medicinal purposes	Yes	Yes	Orwa et al. (2009)
Ornamental purposes (pathway cause)		Yes	Yes	Nelson (2006)
People foraging (pathway cause)	Fruits/ Leaves consumed as vegetable	Yes	Yes	Nelson (2006)

Pathway Vectors

Pathway vector	Notes	Long distance	Local	References
Floating vegetation and debris (pathway vector)	Seeds can float in water for months	Yes	Yes	Nelson (2006)
Water (pathway vector)	Seeds floating in water	Yes	Yes	Nelson (2006)
Host and vector organisms (pathway vector)	Seeds are dispersed by birds, bats, and other mammals	Yes	Yes	Nelson (2006)

Habitat

M. citrifolia can be found growing in a wide range of habitats (e.g., from very dry to wet sites) including coastal and littoral forests, disturbed forests, dry to mesic forests, deciduous forests, xerophytic habitats, grasslands, open areas near shoreline, abandoned pastures, and coconut plantations. It is also common in home gardens, backyards, disturbed sites around villages, and along streams near cultivation (Nelson, 2006).

Biology and Ecology

Genetics

The chromosome number reported for M. citrifolia is n = 22 (Philip and Mathew, 1988). Within this species, there is great morphological variation for fruits and leaves with no clear subpopulations bearing unique traits (Nelson, 2006).

Reproductive Biology

Flowers in M. citrifolia are hermaphroditic. Anthesis is diurnal and flowers are visited and probably pollinated by bees, but plants can self-pollinate (Orwa et al., 2009; PROTA, 2017).

Physiology and Phenology

M. citrifolia has a moderate growth rate (0.75–1.5 m/year), slowing as the tree reaches maturity. However, juvenile plants may grow 1.2–1.5 m in just 6 months. Plants begin flowering and producing fruit in the first year after transplanting. Flowers and fruit are produced throughout the year. Unripe fruit are light green, turning whitish yellow when ripe. Fruit when harvested at the ‘hard white’ stage turn soft and translucent yellow within a few days. In Hawaii flowers on individual M. citrifolia trees are produced over a span of several weeks (or more) as fruit expand in size (Janick and Paull, 2008). Ripe fruit are distributed by several animals including fruit bats.

The seeds have an air chambers and float, facilitating their dispersal via oceanic currents (Macpherson et al., 2007). Dispersal of seeds in native habitats is probably by birds, rats, bats and other mammals. Seeds may remain viable for at least 6 months. Germination time is 3-9 weeks after sowing. Seed germination can be rapid and uniform (~20 days) in full sun to partial shade and mean temperature of approximately 38°C (Groenendijk, 1991; Nelson, 2006; Orwa et al., 2009; PROTA, 2017).

Qualitative and compositional changes were determined in harvested fruits and ripening went through three stages: no significant softening, significant softening and dramatic softening (Cárdenas-Coronel et al., 2016). Ripening was accompanied by accumulation of acidity and soluble solids, and the activity of pectinases and hemicellulases promoted the differential disassembly of cell wall polymers resulting in fruit softening.

Longevity

M. citrifolia is a perennial species and plants may live for 25 to 50 years, probably longer (Nelson, 2006).

Associations

A symbiotic association in the native range with the ant species Oecophylla smaragdina has been described. The plant provides the ants with food and leaves for nesting in exchange for protection from insect predators (Tan, 2001; Francis, 2004). M. citrifolia is the principal larval host of the hawk moth, Macroglossum hirundo vitiensis in Fiji (Nelson, 2006).

Environmental Requirements

M. citrifolia grows in a very wide range of environments and soils and has an unusual ability to survive in harsh conditions such as coral atolls or basaltic lava flows. The species grows from sea level to about 1500 m, depending on latitude and environment. It prefers 20–35°C, although it can tolerate a minimum temperature of about 5°C.

M. citrifolia is a salt and salt-spray tolerant plant and is most competitive where many other plants cannot grow, such as on coral beach sands, volcanic lava flows, in brackish tide pools or on the slopes of very steep gulches. It grows very well on rocky soils, but may not compete well with grasses or other weeds in deep, silty soils. It can tolerate a wide range of precipitation patterns up to 3000 mm/year, including summer, winter, bimodal and uniform. Mature, cultivated M. citrifolia can easily withstand drought of 6 months or more. Wild M. citrifolia plants growing in arid conditions can spend their entire lives in conditions of perpetual drought. It also tolerates waterlogging, salty soils and salt spray.

M. citrifolia can grow well under a wide range of light intensities, from full sun to over 80% shade. It can regenerate after fire by sprouting new foliage from roots or stems. The plant tolerates shallow, sodic and infertile soils (Janick and Paull, 2008). It is tolerant of windy locations, but yields and growth are retarded. The brittle woody branches are easily broken by high winds, but are easily regenerated following storms (Macpherson et al., 2007).

Climate

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

Latitude/Altitude Ranges

Latitude North (°N)	Latitude South (°S)	Altitude lower (m)	Altitude upper (m)
30	30	0	800

Air Temperature

Parameter	Lower limit (°C)	Upper limit (°C)
Mean annual temperature	20	25
Mean maximum temperature of hottest month	32	38
Mean minimum temperature of coldest month	5	18

Rainfall

Parameter	Lower limit	Upper limit	Description
Dry season duration	0	4	number of consecutive months with <40 mm rainfall
Mean annual rainfall	250 mm	4000 mm	mm; lower/upper limits

Rainfall Regime

Summer

Winter

Bimodal

Uniform

Soil Tolerances

Special soil tolerances > infertile

Special soil tolerances > shallow

Special soil tolerances > saline

Special soil tolerances > sodic

Soil reaction > acid

Soil reaction > alkaline

Soil texture > heavy

Soil texture > light

Soil texture > medium

Soil texture

Soil reaction > neutral

Soil drainage > free

Soil drainage > seasonally waterlogged

Notes on Pests

When grown in the Pacific it is susceptible to attack and damage by a range of insects, such as aphids (e.g. the melon aphid, Aphis gossypii), scales (e.g. the green scale, Coccus viridis), weevils, leaf miners, whiteflies (e.g. the Kirkaldy whitefly, Dialeurodes kirkaldyi), caterpillars (e.g. croton caterpillar, Achaea janata), thrips (e.g. the greenhouse thrips, Heliothrips haemorroidalis), ants and mites. The most severe damage in Hawaii, USA, is associated with whiteflies, whereas in Micronesia a species of leaf miner is damaging. In general, insect damage may be more severe in relatively dry or low rainfall locations or in full-sun plantings where Morinda citrifolia is cultivated as an expansive monocrop. Most diseases cause relatively minor damage. The biggest threat to cultivation in the Pacific is root-knot nematodes (Meloidogyne spp.), the cause of root knot disease. The roots are very susceptible to attack by these nematodes and farm failure due to the disease is not uncommon in Hawaii. Attack by the nematodes severely stunts plant growth and allows other opportunistic pathogens, such as the fungus Athelia rolfsii, to infect the roots. Root knot is best controlled through avoidance of infection in the nursery. In Hawaii, there is also a very severe Phytophthora disease known as black flag, caused by an unidentified Phytophthora species. Infection results in extensive blight of leaves, stems and fruit. The disease may be controlled with foliar applications of phosphoric acid fertilizers (Janick and Paull, 2008).

List of Pests

Notes on Natural Enemies

Nelson (2006) reports that M. citrifolia is susceptible to attack and damage by a range of insects, diseases, and pests, including:

•

Aphis gosypii (aphids)

•

Coccus viridis (green scale)

•

Dialeurodes kirkaldyi (whitefly)

•

Achaea janata

•

Heliothrips haemorroidalis (thrips)

•

Colletotrichum spp. (leaf spot disease)

•

Phytophthora spp. (fungi)

•

Athelia rolfsii

•

Meloidogyne spp. (nematodes)

M. citrifolia is also susceptible to infection by coastline parasitic plants such as Cuscuta spp. and Cassytha filiformis (Nelson, 2006).

Natural enemies

Natural enemy	Type	Life stages	Specificity
Aphis gossypii (cotton aphid)		Whole plant	not specific
Coccus viridis (soft green scale)		Whole plant	not specific
Achaea janata (castor semilooper)	Herbivore	Whole plant	not specific
Heliothrips haemorrhoidalis (black tea thrips)		Whole plant	not specific
Athelia rolfsii (sclerotium rot)	Pathogen	Whole plant	not specific
Dialeurodes kirkaldyi		Whole plant	not specific
Cassytha filiformis (love-vine)	Parasite	Whole plant	not specific

Impact: Environmental

M. citrifolia is an invasive species with the capability to survive harsh environmental conditions and extended periods of drought. It is also recognized for its ability of persist and to disperse and colonize new areas without a specific biological dispersal agent, such as humans, rodents, and birds. Once naturalized, this species grows forming dense canopies that inhibit the establishment of native vegetation. It also has a very aggressive and extensive root system that outcompetes native plants for resources such as water and nutrients. On some islands across the Pacific region (i.e., Micronesia) M. citrifolia is considered a weed in some agroforestry or diversified farming settings (Nelson, 2006).

Risk and Impact Factors

Invasiveness

Proved invasive outside its native range

Has a broad native range

Abundant in its native range

Highly adaptable to different environments

Is a habitat generalist

Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc

Pioneering in disturbed areas

Tolerant of shade

Highly mobile locally

Benefits from human association (i.e. it is a human commensal)

Long lived

Fast growing

Gregarious

Reproduces asexually

Impact outcomes

Altered trophic level

Damaged ecosystem services

Ecosystem change/ habitat alteration

Modification of nutrient regime

Modification of successional patterns

Monoculture formation

Reduced amenity values

Reduced native biodiversity

Threat to/ loss of native species

Impact mechanisms

Competition - monopolizing resources

Competition - shading

Rapid growth

Rooting

Likelihood of entry/control

Highly likely to be transported internationally deliberately

Difficult/costly to control

Uses

M. citrifolia is harvested from both plantations and from the wild. This species is one of the most important botanical remedies and food supplements traded on the international market. Its fruits, leaves, fruit juice, and extracts are sold worldwide. The fruit and leaves are edible raw or cooked. Unripe fruit are cooked in curries and ripe fruit are consumed with salt in Myanmar. The ripe fruit has the smell of putrid cheese. Cooked fruit is mixed with coconut in Nauru. Very young leaves are cooked as vegetables (containing 4–6% protein) and eaten with rice in Java and Thailand (Janick and Paull, 2008).

M. citrifolia is often planted as a windbreak, to provide support for pepper vines, and as a shade tree in coffee plantations (Tan, 2001). It is sometime planted as an ornamental, but this practice is not very common due to the strong and sometimes offensive odour of ripened fruits and because the fallen fruits attract many flies and other insects.

All parts of the plant have traditional uses. The roots and bark are used for dyes and medicines; the trunks are used for firewood and tools; the leaves are used as leafy vegetables and for wrapping foods, in medicines and poultices; and the fruit are used as famine food, in juices and for topical and internal medicines. The root bark produces a red dye (morindin) and the plant was widely grown in Java in the late 19th century for this purpose before synthetic dyes became available. The dye is still used in high quality batik (Janick and Paull, 2008). The fruit pulp is used to clean hair, iron and steel, while the wood is occasionally used for poles and plant supports (Nguyen and Nguyen, 2003).

Economic Value

The worldwide market for products of M. citrifolia was an estimated US$400 million in 2002 and projected to reach up to US$ 1.3 billion in 2015 (Potterat and Hamburger, 2007; Assi et al., 2017). M. citrifolia contains phytochemicals that own antibacterial, antiviral, antifungal, antitumor, anthelminthic, analgesic, hypotensive, anti-inflammatory and immune enhancing effects that have attracted industries to employ it as a part of various products and for wide applications such as a natural source of medicines and chemical reagents as well as a green insecticide (Assi et al., 2017).

Social Benefit

M. citrifolia has been used for centuries for its curative properties to treat varieties of illnesses. The purported modern application of M. citrifolia as a complementary alternative medicine spans a vast array of maladies including high blood pressure, diabetes, beri-beri, asthma, coughs, fevers, centipede bites, sores, headaches, pneumonia, diarrhoea, pain, arthritis, depression, cancer, AIDS, skin parasites, skin and stomach ulcers, arteriosclerosis and senility. Some of these uses can be accounted for by the presence of a number of physiologically active chemicals including anthraquinones, alkaloids, scopoletin, glycosides, polysaccharides, asperuloide and organic acids such as caprioc, caprylic and ursolic acids.

While the plant enjoys an outstanding anecdotal reputation regarding these maladies and shows promise as an anti-cancer agent in mice experiments, many purported benefits of using M. citrifolia topically or internally have yet to be supported by published data from peer-reviewed clinical trials involving humans. Nevertheless, M. citrifolia has attained significant economic importance worldwide through a variety of health and cosmetic products made from its leaves and fruit. These products, including fruit juice and powders derived from fruit or foliage, are some of the most important botanical remedies and food supplements currently traded on the international market (Nguyen and Nguyen, 2003; Janick and Paull, 2008).

The ethnomedicinal and pharmaceutical properties are reviewed by Macpherson et al., 2007, Assi et al. (2017), Ali et al. (2016) and Torres et al. (2017). Gu et al. (2018) demonstrated that leaf extracts enhanced osteogenic differentiation in human cell lines. Pandy and Vijeepallam (2017) showed that methanolic fruit extracts, specifically the scopoletin and rutin components, alleviated symptoms of schizophrenia in mice.

Environmental Services

M. citrifolia has been planted in coastal areas for erosion control (Francis, 2004). On islands across the Pacific region, M. citrifolia is also used in traditional subsistence farming, intercropped with breadfruit, kava, papaya, mango, coconut, and bananas (Nelson, 2006).

Uses List

Ornamental

Ornamental > Christmas tree

Ornamental > Cut flower

Ornamental > garden plant

Ornamental > Potted plant

Ornamental > Propagation material

Ornamental > Seed trade

Materials > Dye/tanning

Medicinal, pharmaceutical > Traditional/folklore

Human food and beverage > Fruits

Human food and beverage > Vegetable

Materials > Wood/timber

Environmental > Agroforestry

Environmental > Amenity

Environmental > Erosion control or dune stabilization

Environmental > Windbreak

Materials > Chemicals

Materials > Cosmetics

Medicinal, pharmaceutical > Source of medicine/pharmaceutical

Fuels > Fuelwood

Human food and beverage > Beverage base

Human food and beverage > Emergency (famine) food

Human food and beverage > Honey/honey flora

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.

M. citrifolia has deep taproot and an extensive and aggressive root system and once established it is very persistent and difficult to eradicate (Nelson, 2006). There is no information available for the chemical control of M. citrifolia.

Silviculture Characteristics

Tolerates > drought

Tolerates > wind

Tolerates > waterlogging

Cultivation

Optimal growth of M. citrifolia cultivation is obtained on well drained soils, but sites infested with rootknot nematodes (Meloidogyne spp.) should be avoided. Plants do well in rocky soils, probably due to the absence of nematodes in these soils. Sites with full sun or partial shade are preferred (Nelson, 2003; Macpherson et al., 2007). In Hawaiian plantations, plant spacing ranges from 4 to 6 m (Janick and Paull, 2008) although Macpherson et al. (2007) notes that the recommended plant spacing in Hawaii is 3 to 4.6 m to give densities of 473–1111 plants/ha. Higher densities are thought to cause pest and disease problems.

After transplanting, balanced NPK fertilizers (13:13:13 or 16:16:16) are used to promote vegetative growth. Later, growers in Hawaii may switch to unbalanced fertilizers (such as 10:20:10 or 10:20:20) containing more potassium to stimulate flower production and to supply the fruit’s needs. Seedlings are given controlled release formulations while older plants are given rapidly available granular formulations. These are applied at the drip line where water drips from the edge of the leaf canopy (Nelson, 2003). Integrated nutrient management for the production of M. citrifolia in India was studied by Bhoomika et al. (2017). In trials they noted that optimum growth and yields were obtained using applications of 50:225:50 kg/ha NPK fertilizer with 50% or 100% of the recommended phosphorus supplied as bonemeal. Most farms in the Pacific do not use supplemental irrigation due to the plant’s drought tolerance. The lower branches tend to droop when bearing fruit and contact the ground. These branches are pruned away where needed to allow other farm operations such as weed control and fertilization (Janick and Paull, 2008). Young plants less than 3 years old may be pruned back after first fruiting to promote bushiness as trees can grow to 10 m in height (Nelson, 2003).

M. citrifolia is suitable for intercropping with cashew (Kumara et al., 2016) and coconut (Bhalerao et al., 2016). Khandekar et al. (2015) found that intercropping increases the profitability of coconut cultivation.

Harvesting

Plants often begin to bear fruit around 9 to 12 months after planting but, as fruits are small and few, farmers forgo this harvest and prune back the branches instead. Fruiting occurs all year round, but seasonal trends may be affected by the weather and by management. According to Janick and Paull (2008), fruit are harvested throughout the year, usually once a week. Fruit may be picked and used at any stage of development, depending on the product being made. For most juice operations, fruit are picked when they are at the ‘hard white’ stage (i.e. when almost all the green colour has turned to a whitish yellow, but before they turn soft and fall from the tree). In Hawaii harvesting is undertaken 2–3 times per month, and fruits are harvested manually. Fruits are placed in baskets, bags or placed in bins for transport to processing. They tend not to bruise or damage easily and do not need refrigeration after harvest (Nelson, 2003).

Yields within farms increase each year as the plants grow, as the plants may not reach maximum size for 10 years or more. Fruit yields depend on many environmental and cultural factors. In Hawaii, yields of 80,000 kg/ha may be reasonably expected, with some farms reporting much greater yields (Janick and Paull, 2008). Macpherson et al. (2007) noted that yields in Hawaii are 114–227 kg/plant/year depending on spacing and fertilizers, but yields may exceed these figures.

Postharvest Treatment

Fruit should be washed before they soften. Softened fruit are placed into large juice collection containers where the fluids seep out naturally and separate from the pulp, or are placed into a bladder press to squeeze out the fresh juice (Janick and Paull, 2008). However, fruits destined for juice production are normally held at ambient temperatures for one to several days to ripen fully. For processing for powders or minimally processed fruit, fruits are generally processed immediately before they are fully ripe. Unripe fruits are easier to process mechanically (Nelson, 2003).

Wall et al. (2015) noted that the fruits have a non-climacteric respiratory pattern (34 mg CO₂/kg/hour) with no detectable ethylene production. López-Vázquez et al. (2013) evaluated fruit stored at 0, 5, 15 and a control temperature of 28°C. Fruits stored at 0 and 5°C were severely damaged by cold, but storage at 15°C increased postharvest life to 12 days. However, weight loss was 13.4%.

Fruit juice is processed and produced by a variety of methods and is available fermented and unfermented or fresh squeezed or drip extracted. It may be bottled with or without pasteurization. So-called "traditional" juice is drip extracted and fermented or aged for at least 2 months. For the production of traditional juice, ripening fruits are washed, placed in fermentation vessels for 2 months or longer and the juice drips from the pulp and gradually ferments. Air is excluded and the juice is run off and filtered and bottled. This juice has a dark brown appearance similar to soy sauce with a sour taste. Juice recovery rates are 40–50% of the original fruit weight. Sometimes the pulp may be pressed to release the remaining juice. Occasionally the juice is pasteurized, altering the flavour, but the low pH of the juice makes pasteurization unnecessary. If an unfermented product is required, juice is drawn off every couple of days. Unfermented juice is lighter coloured and sweeter. The juice may be mixed with sugar or other fruit juices to improve the palatability. Fruit juice may be evaporated to produce a powder which can be used in various products, including reconstituted juices (Nelson, 2003).

Genetic Resources and Breeding

A chromosome count of 2n = 22 is recorded by the Chromosome Counts Database (2018).

There is a high degree of variability in fruit and leaf morphology, but little selection has so far been undertaken. Singh (2010) and Waki et al. (2008) have published descriptor lists for M. citrifolia.

RAPD markers were used to evaluate 13 Brazilian accessions and revealed that genetic diversity was low. Genetic distance ranged from 0.04 to 0.49 (Bordallo et al., 2017). RAPD and SSR markers were used by Patel et al. (2014) to assess 13 Indian genotypes. Genotypes clustered into one cluster, suggesting low variability. Wu and Lan (2014) used ISSR markers to characterise 13 accessions and two clusters were obtained with small fruited accessions clustered together.

Lee et al. (2013) carried out Agrobacterium-mediated genetic transformation using direct exposure of seeds which were clipped and incubated in the inoculation medium. A transformation efficiency of 96.8% was obtained with 2 hours co-cultivation.

Major Cultivars

A grower in Queensland, Australia, has registered a selection (cv. 'Allright') with large (>300 g) pink fruits which are firmer in texture. The juice is apparently less cheesy and ripe fruits are sweeter with high fructose levels ( Macpherson et al., 2007 ; Aurait Supreme Pty Ltd, 2008).

Propagation

This species is usually propagated by seeds and/or cuttings. Fully ripe fruit are crushed by hand and the seeds and pulp allowed to fall into a pot or container filled with a growth medium. The layer of pulp and seeds is then covered with a shallow layer of growth medium and the pot is placed in a warm location and watered daily. Without scarification seeds take several months or a year to germinate. Germination may be reduced to a month by applying heat. Seeds can tolerate 38°C or even higher, and heat may be supplied by warming cables in the greenhouse (Nelson, 2003). In Hawaii, fruit contain up to 260 seeds each. The seeds (air dried) weigh from 0.21 to 0.25 g/seed, 10 kg of fruit yielding about 250 g of air dried seeds. The seedcoat is very fibrous, tough and relatively impermeable to water. Unscarified seeds may require 60 days or longer for germination, depending on temperature. Scarified seeds can germinate in 3–4 weeks. The seed remains viable for about 6 months (Janick and Paull, 2008). Various techniques were used to break seed dormancy by Elakkuvan and Manivannan (2010). Highest germination rates (94%) were obtained within 7–12 days when nicked seeds were soaked in hot water (45°C) for 24 hours followed by gibberellic acid (1000 ppm) treatment for 24 hours before germination.

Stem cuttings root easily in 3–4 weeks in an inert growth medium such as vermiculite or volcanic cinders. Woody vertical stems from vigorous plants are selected. Root sprouts are also produced which can be dug up and planted. Air layering of branches is also possible (Janick and Paull, 2008).

Huang et al. (2017) used leaves as explants for tissue culture. Two routes for in vitro regeneration were established: induction of callus followed by induction of roots and buds, and induction of adventitious roots followed by induction of buds. Callus was induced on modified MS medium containing 0.1 mg/litre benzyladenine + 2 mg/litre 2,4-D. Leaf callus formed roots and shoots on MS medium containing 1 or 2 mg/litre benzyladenine + 0.4 mg/litre NAA. Roots and bud regeneration directly from leaf explants on MS medium containing 1 mg/litre benzyladenine + 0.4 mg/litre NAA. A micropropagation protocol utilizing nodal explants was reported by Shekhawat et al. (2015). MS medium containing 4 mg/litre benzyladenine allowed the regeneration of 4.6 shoots per explant. Rooting was achieved on half-strength MS medium containing 1 mg/litre IBA producing 44.3 roots per shoot.

Rootstocks

No rootstocks are used or have been reported in the cultivation of M. citrifolia ( Janick and Paull, 2008 ).

Nutritional Value

Among the nutritional characteristics of M. citrifolia, the levels of protein (4–6% in leaves) and ascorbic acid (24–258 mg/100 g dried fruit, 65 mg/100 ml pure juice) are perhaps most notable (Janick and Paull, 2008). Studies on the nutritional value of noni fruit and its products are reviewed by Motshakeri and Ghazali (2015).

Rosalizan et al. (2010) reported the proximate composition (per 100 g) of noni fruit during ripening. At full maturity (15 weeks after fruit set) moisture contents were 88.1 g, protein content was 0.56 g, fat 0.08 g, dietary fibre 9.63 g, carbohydrate 0.60 g, and ash 1.03 g. Retinol composition decreased from a high of 57 µg RE at week 11 to 4.67 µg RE at full maturity in week 15. Ascorbic acid content increased to 6.82 mg at maturity. The mineral contents of the fruit did not differ significantly during maturation. At maturity fruits (per 100 g) contained 3.52 mg sodium, 38.08 mg calcium, 369.93 mg potassium, 0.46 mg iron, 19.57 mg phosphorus, 17.40 mg magnesium and 0.26 mg zinc.

Phytosanitary Issues/Food Safety

There have been several cases of hepatotoxicity caused by drinking M. citrifolia juice (Yüce et al., 2006) but a safety review revealed no adverse health effects from M. citrifolia dietary supplements, even at high doses (West et al., 2006). Intake of 750 ml of juice per day is considered safe. The presence of anthraquinones was thought to be the causal agent in hepatotoxicity and some individuals are particularly sensitive. A European Food Safety Authority (EFSA) panel has concluded that fruit puree and concentrate are safe for consumption (EFSA, 2009).

Production and Trade

Although there are no worldwide cultivation statistics, the number of M. citrifolia farms has increased dramatically in the Pacific and worldwide since the early 1990s to meet the growing market demand for products derived from the plant (Janick and Paull, 2008). The largest markets are North America, Europe, Japan, Mexico, Asia and Australia with the worldwide market for these products estimated at US$400 million (Macpherson et al., 2007).

Links to Websites

Name	URL	Comment
GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gateway	https://doi.org/10.5061/dryad.m93f6	Data source for updated system data added to species habitat list.

References

Chromosome Counts Database, 2018. http://ccdb.tau.ac.il/home/

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