Gmelina arborea
(candahar)

Pictures

Picture	Title	Caption	Copyright
Title Roadside tree Caption Mature tree in Malim Nawar, Perak, Malayisia. Copyright Lai Hoe Ang	Roadside tree	Mature tree in Malim Nawar, Perak, Malayisia.	Lai Hoe Ang
Title Leaf-fall Caption In Bidor, west coast Peninsular Malaysia, loss of foliage begins in December and is completed in March. Copyright Lai Hoe Ang	Leaf-fall	In Bidor, west coast Peninsular Malaysia, loss of foliage begins in December and is completed in March.	Lai Hoe Ang
Title Sapling Caption Sapling with relatively good stem form, in a secondary forest. Copyright Lai Hoe Ang	Sapling	Sapling with relatively good stem form, in a secondary forest.	Lai Hoe Ang
Title Leaves Caption The cordate leaf blade with slender petiole showing the secondary vein distribution. Copyright Lai Hoe Ang	Leaves	The cordate leaf blade with slender petiole showing the secondary vein distribution.	Lai Hoe Ang
Title Glands at base of leaf Caption A pair of glands positioned at the base of a cordate leaf. Copyright Lai Hoe Ang	Glands at base of leaf	A pair of glands positioned at the base of a cordate leaf.	Lai Hoe Ang
Title Fruit and aborted flowers Caption Green fruits (drupe) with aborted flowers of cordate leaf blade of Gmelina arborea growing at Bidor. Perak in December 1997. The diameter of the coin is 2.4cm. Copyright Lai Hoe Ang	Fruit and aborted flowers	Green fruits (drupe) with aborted flowers of cordate leaf blade of Gmelina arborea growing at Bidor. Perak in December 1997. The diameter of the coin is 2.4cm.	Lai Hoe Ang
Title Line artwork Caption 1. tree habit. 2. flowering twig. 3. flower. 4. fruits. Copyright PROSEA Foundation	Line artwork	1. tree habit. 2. flowering twig. 3. flower. 4. fruits.	PROSEA Foundation

Identity

Preferred Scientific Name

• Gmelina arborea Roxb.

Preferred Common Name

• candahar

Variety

• Gmelina arborea var. arborea
• Gmelina arborea var. canescens Haines
• Gmelina arborea var. glaucescens Haines

Other Scientific Names

• Gmelina sinuata Link

International Common Names

• English: gamhar; gmelina; goomar teak; Kashmir tree; Malay beechwood; Malay bush-beech; snapdragon tree; white beech; white teak
• Spanish: ciruela Malaya; jobo de Africa; melina
• French: peuplier d’ Afrique; yemane
• Chinese: yun nan shi zi
• Portuguese: árvore-boca-de-leão; guemelina; guimelina

Local Common Names

• Bangladesh: gamar
• Brazil: gmelinea
• Costa Rica: melina
• Cuba: Álamo blanco
• Dominican Republic: ciruela de Malaya
• India: badraparmi; ban; bhodroporrni; gamari; gambari; gambhar; ghambari; ghandari; gomari; gumadi; gumartek; gumbar; gumhar; gummadi; kasmari; kasmiri-mara; khambhari; kumbil; kumhar; kummadi; perungumpil; sewan; shewan; shivan; shivani; shriparmi; umi-thekku
• India/Assam: gomari
• Myanmar: yamane
• Nepal: gamari; gambari; gumhari; khamari; yemane
• Philippines: yemane
• Thailand: so-maeo

EPPO code

• GMEAR (Gmelina arborea)

Trade name

• gamari
• gamhar
• yamane

Summary of Invasiveness

Due to its rapid growth rate, G. arborea is a tree which has been widely used in reforestation programs in tropical and subtropical regions of the world and as a source of commercial timber and cellulose (Dvorak, 2004; Rojas-Rodríguez et al., 2004; Silva et al., 2005; USDA-ARS, 2016). This species produces large numbers of fertile fruits that are easily dispersed by birds and bats, spreading seedlings quite far from the parent tree (Orwa et al., 2009). In this way, G. arborea has escaped from plantations and entered wild habitats where it is now replacing native trees and becoming invasive (IUCN, 2013). Currently, it is listed as invasive in Costa Rica, the Dominican Republic, Ghana, Australia and the Cook Islands (Chacón and Saborío, 2012; Mir, 2012; IUCN, 2013; PIER, 2016; Weeds of Australia, 2016). It is also separately reported as invasive in Malawi, Tanzania and Zambia.

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Plantae
•         Phylum: Spermatophyta
•             Subphylum: Angiospermae
•                 Class: Dicotyledonae
•                     Order: Lamiales
•                         Family: Lamiaceae
•                             Genus: Gmelina
•                                 Species: Gmelina arborea

Notes on Taxonomy and Nomenclature

The family Lamiaceae comprises around 236 genera and 7173 species distributed worldwide (Stevens, 2012). The genus Gmelina includes 38 species of trees occurring in tropical Asia, Australia and New Guinea (Flora of China Editorial Committee, 2016). Gmelina is related to the large genus Vitex, which also includes several timber trees. Vitex can be distinguished by its more narrowly funnel-shaped corolla and less fleshy drupe (Flora of China Editorial Committee, 2016; PROTA, 2016).

The genus Gmelina, here assigned to the family Lamiaceae, is sometimes placed in the Verbenaceae. It was first described by Roxburgh in 1832 and later in a more complete form by Hooker in 1885 (Brandis, 1924). G. arborea var. canescens and var. glaucescens (Haines 1910, cited by Greaves, 1981) are differentiated from the type variety by their lamina features, i.e. stellately-hairy beneath (type), subcoriaceous and grey-pubescent with simple hairs (var. canescens) and glaucous and glabrous beneath (var. glaucescens). The genus name Gmelina was named by Carl Linnaeus in honour of the German botanist Johann Georg Gmelin. The specific name means treelike, from the Latin ‘arbor’ (tree).

Description

Trees about 15 m tall; bark grayish brown; branchlets, petioles, and inflorescences densely yellow-brown tomentose. Branchlets slightly 4-angled when young, becoming terete, lenticellate, leaf scars prominent. Petiole terete, 3.5-10 cm; leaf blade broadly ovate, 8-19 X 4.5-15 cm, thickly papery, base broadly cuneate to subcordate, apex acuminate; veins 3-5 pairs, abaxially prominent. Inflorescences terminal, narrow thyrses; peduncle 15-30 cm. Calyx 3-5 mm, with several black discoid gland patches; teeth 5, sharply triangular. Corolla yellow, 3-4 cm, 2-lipped, sparsely glandular; lower lip 3-lobed, outside yellowish brown puberulent, inside glabrous; upper lip entire or slightly 2-cleft. Ovary glabrous, glandular. Stigma unequally 2-cleft. Drupes yellow when ripe and black when dry, ellipsoid to obovoid-ellipsoid, 1.5-2 cm (Flora of China Editorial Committee, 2016).

Plant Type

Distribution

G. arborea is native to Asia, from Pakistan to Sri Lanka, Myanmar, Thailand, Vietnam and southern China (Troup, 1921; Moldenke, 1977; Greaves, 1981; Gupta, 1993; Luna, 1996). This species has been extensively planted as a fast-growing tree in tropical areas of Africa, Asia, Australia, America, the West Indies, and on several islands in the Pacific Ocean (PROTA, 2016).

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

G. arborea was first introduced from Myanmar as a fast-growing tree species into forest plantations of Peninsular and East Malaysia. It was also introduced to other ASEAN countries, such as the Philippines and Indonesia. By the 1960s, the FAO encouraged the utilization of G. arborea for reforestation projects and commercial purposes across the tropics (FAO, 1981). By that time, G. arborea was featured as a promising species due to its ease and inexpensive establishment, rapid early growth, quick return on investment, and for its wood characteristics including high durability, good yield and high quality pulp. So, during the following 30-35 years, small and large-scale plantations of G. arborea were established through Southeastern Asia, Australia, West Africa, and South and Central America (Lauridsen and Kjaer, 2002).

In Brazil, G. arborea was first introduced from Asia in the 1960s during a reforestation project located in the Amazon basin (Silva et al., 2005).

In Central America, the first plantation of G. arborea was established in 1966 in Costa Rica (in the Caribbean coast of Siquirres) using seeds introduced from more than 20 sites across its native distribution in Asia (Rojas-Rodríguez et al., 2004). Later, during the 1980s-1990s, the establishment of G. arborea plantations was promoted by local and international agencies in Panama, Guatemala, El Salvador, Nicaragua and Honduras. The total area of G. arborea plantations in Central America has been estimated at 52,000 ha, with biggest plantations located in Costa Rica and Guatemala (Rojas-Rodríguez et al., 2004).

In tropical Africa, G. arborea is planted in many countries, and large-scale plantations can be found in Senegal, Gambia, Sierra Leone, Ivory Coast, Mali, Burkina Faso, Ghana, Nigeria, Cameroon and Malawi. The total area of G. arborea plantations in Africa has been estimated at 130,000 ha (PROTA, 2016).

In Asia and the Pacific region, large plantations have been established in Malaysia, the Philippines, Singapore, and Solomon and Fiji Islands. There is an estimation of 50,000 ha of G. arborea plantations in Asia and the Pacific (Lauridsen and Kjaer, 2002).

Introductions

Risk of Introduction

The likelihood of further introductions of G. arborea is very high. At the present, approximately 700,000 ha of G. arborea have been established in plantations in Africa, Southeast Asia, the South Pacific, and Central and South America. It is expected that planting areas will expand to 800,000 ha by 2020 (Dvorak, 2004). Although it is widely introduced, a risk assessment for Hawaii gave it a low risk score of 2 for the danger of becoming a problem (PIER, 2016).

Habitat

G. arborea occurs in a wide range of forest habitats, including tropical rainforests, evergreen forests, submontane thickets, very moist forests, deciduous forests and dry forests (Nair, 2001). It also occurs in low alluvial savannah and woodland (Orwa et al., 2009). In China, it grows in open forests along roadsides and near farmhouses at elevations below 1500 m (Flora of China Editorial Committee, 2016). In Central America, plantations of G. arborea can be found in areas with climate varying from tropical dry to tropical moist and wet climate (Rojas-Rodríguez et al., 2004).

Habitat List

Biology and Ecology

Genetics

The chromosome number reported for G. arborea varies from 2n = 36, 2n = 38 to 2n = 40 (Chatha and Bir, 1986; PROTA, 2016). In provenance trials, plants of G. arborea originating from plantations often performed better than plants originating from forests within its natural distribution. This can be due to: (i) a positive selection during thinning in the plantations, (ii) a result of lower inbreeding in the plantations, or (iii) a positive response and adaptation to local conditions (because the plantings often represent local ‘landraces’; Lauridsen and Kjaer, 2002).

Reproductive Biology

G. arborea produces large, bisexual, and zygomorphic nectariferous flowers. The breeding system involves both self- and cross-pollination, but most of the self-pollinated flowers are aborted after two weeks of growth. Flowers are often visited and pollinated by bees (especially Apis and Xylocopa bees) and passerine birds (Orwa et al., 2009). Fruit-set under natural conditions is low (Solomon and Purnachandra, 2006).

Physiology and Phenology

G. arborea starts producing flowers and fruits when trees are 6-8 years old, however, trees in plantations can flower when they are 3-4 years old (Rojas-Rodríguez et al., 2004; Orwa et al., 2009). Under favourable conditions G. arborea is capable of reaching an annual increment of 20–25 m³/ha with impressive exceptions of over 30 m³/ha. On poor sandy soils a yield of only 84 m³/ha after 12 years was reported, whereas on very favorable soils a production of 304 m³/ha after 10 years can be reached (PROTA, 2016).

In China, G. arborea has been recorded flowering from April to May and fruiting from May to July (Flora of China Editorial Committee, 2016). In India, it flowers from February to March and fruits ripen from the end of April to June (Orwa et al., 2009). In Central America, trees often produce flowers from December to March (Rojas-Rodríguez et al., 2004).

Longevity

G. arborea is a perennial fast-growing tree (Sanon et al., 2006; Flora of China Editorial Committee, 2016). It has also been classified as a long-lived pioneer species (PROTA, 2016).

Environmental Requirements

G. arborea grows in both dry and moist areas at elevations from sea level to 1500 m and annual rainfall from 750 mm to 4500 mm (Flora of China Editorial Committee, 2016; PROTA, 2016). It can tolerate a dry season of up to 6–7 months (Duke, 1983), although the optimum climate for the species is in areas with a short dry period of 3-5 months and an average relative humidity of about 40%. It grows best in climates with mean annual temperature ranging from 21°C to 28°C (Moya and Tomazello, 2008; Orwa et al., 2009), and with average monthly temperatures generally between 18 and 35°C for the coolest and warmest months, respectively.

G. arborea grows on many soil types, including dry sandy soils and heavily leached acidic soils with pH ranging from 5 to 8 (Duke, 1983; Orwa et al., 2009). When established under poor soil conditions, trees often remain stunted or become little more than a shrub. This species prefers well-drained soils and does not tolerate waterlogged sites or peat soils. It has a high light requirement (PROTA, 2016).

Climate

Latitude/Altitude Ranges

Air Temperature

Rainfall

Rainfall Regime

Soil Tolerances

Soil drainage

• free

Soil reaction

• acid
• neutral

Soil texture

• heavy
• light
• medium

Natural enemies

Notes on Natural Enemies

In Brazil, the fungus Parapyricularia brasiliensis was observed on leaf spot of G. arborea (Silva et al., 2005).

Serious fungal infestation has been observed in G. arborea plantations. Armillaria mellea, Ceratocystis fimbriata, Gnomonia spp., and Poria rhizomorpha [Antrodia rhizomorpha] are some of the fungi that cause serious damage to plantations. In Africa, fungal diseases are mainly root diseases of nursery seedlings, caused by Gibberella fujikuroi and Sclerotium rolfsii [Athelia rolfsii] (Duke, 1983; Nair, 2001; PROTA, 2016). Other nursery pathogens include: Pythium splendens [Globisporangium splendens], which causes wilting in 1-2 month old seedlings; Fusarium oxysporum, a fungus causing high seedling mortality, and Rhizoctonia solani [Thanetophorus cucumeris], a root-collar disease (Orwa et al., 2009).

Defoliators are very common, both on seedlings and mature trees. Leaf-cutting ants (Atta species) cause severe defoliation. The insect Craspedonta leayana has been reported causing serious defoliation in Bangladesh (Baksha, 1997) and India (Rishi and Barthakur, 2016). In India, Thailand and Malaysia the borer Acalolepta cervina has killed or severely damaged whole plantations (Nair, 2001; Orwa et al., 2009). Dieback is caused by shoot cutters such as the larvae of Alcidodes gmelinae in Assam, Bengal and Myanmar (Greaves, 1981). Alcidodes ludificator is also a serious pest in nursery and young plantations in NE India (Senthilkumar and Barthakur, 2008). In Nigeria, Roberts (1969) noted that defoliators such as Empacamenta calabrica, Apophylia nigricollis, Zonoceros variegatus and Achaea lienardi (skeletonizer) are common in plantations. In the Philippines, defoliators such as Chrysodeixis chalcites, Acherontia lachesis, Ozola minor and Attacus spp. are common in nurseries and plantations (Lapis and Bautista, 1977; Lapis and Genil, 1979).

In Zaria (Nigeria), individual trees showed severe infestation by mistletoe (Tapinanthus sp.) (Nwanosike, 2005).

Means of Movement and Dispersal

Vector Transmission (Biotic)

G. arborea spreads by seeds. Birds and bats attracted by the smell of fruits are the main seed dispersal agents (Orwa et al., 2009).

Intentional Introduction

G. arborea has been actively introduced into many tropical and subtropical regions to be used as a forestry and ornamental tree. Worldwide estimates suggest that by 2004 plantations of G. arborea covered an area of about 700,000 ha (Dvorak, 2004).

Pathway Causes

Impact Summary

Environmental Impact

G. arborea produces many fertile fruits that are easily dispersed by birds and bats (Orwa et al., 2009), spreading seedlings quite far from the parent tree. In this way, it has escaped from plantations and entered wild habitats where it is now replacing native trees and becoming invasive (IUCN, 2013). It is listed as invasive in Costa Rica, Dominican Republic, Ghana, Australia and Cook Islands (Chacón and Saborío, 2012; Mir, 2012; IUCN, 2013; PIER, 2016; Weeds of Australia, 2016). In Australia, it is regarded as an environmental weed in the Northern Territory and as a potential environmental weed in Queensland. This species is naturalizing from settlements in the tropical savannas of the Northern Territory. It is also listed as a high priority weed in Aboriginal lands in the Northern Land Council area, and is of particular concern in Maningrida, in Arnhem Land (Weeds of Australia, 2016). It has become naturalized in many African countries, where it may be somewhat invasive and in Ghana it is spreading across the Mole National Park (IUCN, 2013; PROTA, 2016).

Impact on habitats and biodiversity

G. arborea is an opportunist species and it has been classified as a long-lived pioneer. Thus, it has the potential to disrupt successional processes in areas where it is invading and outcompete native vegetation (IUCN, 2013; PROTA, 2016).

Risk and Impact Factors

• 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
• Highly mobile locally
• Benefits from human association (i.e. it is a human commensal)
• Long lived
• Fast growing
• Gregarious
• Reproduces asexually

• Altered trophic level
• Damaged ecosystem services
• Ecosystem change/ habitat alteration
• Modification of successional patterns
• Monoculture formation
• Reduced native biodiversity
• Threat to/ loss of native species

• Competition - monopolizing resources
• Competition - shading
• Pest and disease transmission
• Rapid growth

• Highly likely to be transported internationally deliberately

Uses

Economic Value

G. arborea is a fast-growing tree often grown in large-scale plantations to produce wood for light construction, crafts, decorative veneers, pulp, fuel, and charcoal. It produces high-quality wood, which is harvested for the manufacture of furniture, musical instruments and to make plywood, matches, agricultural implements and even artificial limbs. The wood also produces good quality pulp used in the manufacture of cardboard and paper. Flowers produce abundant nectar, which produces high-quality honey. The leaves are widely used as cattle fodder, and in silkworm culture. The wood ash and fruit yield very persistent yellow dyes (Orwa et al., 2009; PROTA, 2016).

Social Benefit

Roots, bark and seeds of G. arborea are used in traditional Asian medicine. In India G. arborea stem bark and roots are used in Ayurvedic preparations to treat a range of ailments (Shenoy and Yoganarasimhan, 2009; Yogesh and Veeranjaneyulu, 2010; Acharya et al., 2012). The fruit and bark are used to treat various conditions in Sri Lanka (Trimen, 1895; Wagman, 1982). Other vegetative parts (including young leaves) also have medicinal value (Burkhill, 1935). Chemicals that may have commercial potential for drug or chemical production (Greaves, 1981) include apigenin, luteolin and quercetagetin in leaves, gmelinoceryl alcohol in roots, and lignins from the wood.

It is also cultivated as an ornamental tree in gardens and avenues (Orwa et al., 2009).

Environmental Services

G. arborea is often used in reforestation programs (Moya and Tomazello, 2008). The species is also planted in agroforestry systems and as a shade tree on cassava, maize, coffee, and cacao plantations (Orwa et al., 2009; USDA-ARS, 2016). G. arborea has been shown to have some potential for the phytoremediation of crude oil polluted soils as long as the contamination level is below 5% (Agbogidi et al., 2007).

Rotenberg (2007) examines the use of plantations of G. arborea by native birds in Guatemala and assesses its suitability as bird habitat.

Uses List

Animal feed, fodder, forage

• Forage

Environmental

• Agroforestry
• Amenity
• Erosion control or dune stabilization
• Revegetation
• Shade and shelter

Fuels

• Charcoal
• Fuelwood

Human food and beverage

• Honey/honey flora

Materials

• Carved material
• Dyestuffs
• Essential oils
• Fibre
• Miscellaneous materials
• Wood/timber

Medicinal, pharmaceutical

• Source of medicine/pharmaceutical
• Traditional/folklore

Ornamental

• Christmas tree
• Cut flower
• garden plant
• Potted plant
• Propagation material
• Seed trade

Wood Products

Boats

Charcoal

Containers

• Boxes
• Crates

Furniture

Other cellulose derivatives

Pulp

• Short-fibre pulp

Railway sleepers

Sawn or hewn building timbers

• Carpentry/joinery (exterior/interior)
• Exterior fittings
• Fences
• Flooring
• For light construction
• Wall panelling

Veneers

Wood extractives (including oil)

Wood hydrolysates

Wood residues

Wood-based materials

• Plywood

Woodware

• Matches
• Musical instruments

Similarities to Other Species/Conditions

G. arborea looks similar to G. asiatica, but differs in having erect inflorescences and larger leaves than G. asiatica (Flora of China Editorial Committee, 2016).

References

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Links to Websites

Contributors

05/12/16 Original text by:

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

Distribution Maps