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5 December 2016

Gmelina arborea (candahar)

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

Abstract

This datasheet on Gmelina arborea covers Identity, Overview, Associated Diseases, Pests or Pathogens, Distribution, Dispersal, Diagnosis, Biology & Ecology, Environmental Requirements, Natural Enemies, Impacts, Uses, Management, Genetics and Breeding, Economics, Further Information.

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
Trade name
gamhar
Trade name
yamane

Pictures

Mature tree in Malim Nawar, Perak, Malayisia.
Roadside tree
Mature tree in Malim Nawar, Perak, Malayisia.
Lai Hoe Ang
In Bidor, west coast Peninsular Malaysia, loss of foliage begins in December and is completed in March.
Leaf-fall
In Bidor, west coast Peninsular Malaysia, loss of foliage begins in December and is completed in March.
Lai Hoe Ang
Sapling with relatively good stem form, in a secondary forest.
Sapling
Sapling with relatively good stem form, in a secondary forest.
Lai Hoe Ang
The cordate leaf blade with slender petiole showing the secondary vein distribution.
Leaves
The cordate leaf blade with slender petiole showing the secondary vein distribution.
Lai Hoe Ang
A pair of glands positioned at the base of a cordate leaf.
Glands at base of leaf
A pair of glands positioned at the base of a cordate leaf.
Lai Hoe Ang
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.
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
1. tree habit. 2. flowering twig. 3. flower. 4. fruits.
Line artwork
1. tree habit. 2. flowering twig. 3. flower. 4. fruits.
PROSEA Foundation

Overview

Importance

G. arborea produces an important and valuable creamy-white timber which is sometimes called white teak. It is a fast-growing species that demands good site conditions (Durant, 1941; Fox, 1967; Nanagas and Serna, 1970; Fasehun, 1975; Greaves, 1981; Webb et al., 1984; Luna, 1996). Native to Asia, the species has great plantation potential and is widely planted, especially in Africa. Large-scale plantations of G. arborea are found in Nigeria, Sierra Leone and Malawi (Streets, 1962). Compared with other high-quality timber species, G. arborea can be managed for relatively short rotations of 15 to 20 years (Fox, 1967). With recent developments in wood processing technology for utilizing smaller diameter logs, G. arborea plantations are often managed on a less than 15-year rotation; this is especially the case when fuelwood is the main end-product.
G. arborea is also valuable as a multipurpose tree (Tiwari et al, 2007) and is economically important in various agroforestry systems. Different parts of the tree are used in traditional medicine (Nayak et al., 2011).

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

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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).

Plant Type

Broadleaved
Perennial
Seed propagated
Tree
Vegetatively propagated
Woody

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).
Botanical Features

General

G. arborea is a medium-size tree which produces a clear, but frequently bent, bole. It reaches 30 m tall (e.g. in Myanmar) on favourable sites. However, it can become shrubby in drier environments, such as the Punjab and other parts of India. G. arborea has low, obscure buttresses and a yellowish-grey, smooth bole. The branches are numerous and spread to form a large shading crown. The corky bark is yellow inside and greyish-white outside when young, turning brown with age. The old bark then sheds, producing patches of paler bark on the bole.

Foliage

Leaves are simple, normally cordate, broad-ovate acuminate, about 10-25 cm long and 5-18 cm broad, denticulate, with about 3-5 primary veins originating from the base and 4-6 pairs of secondary veins. The petiole is about 7.5-12.0 cm long. A pair of shining glands is located at the intersection of the blade and petiole, along the lower half of the midrib.

Inflorescences, flowers and fruits

The flowers are numerous and arranged in a panicle about 30 cm long. Corolla brown-yellow, upper lip shortly bifid, larger than the lower. The fruit is a drupe with sweet flesh; fruit turning yellow while ripening, 2.0-2.5 cm long and with a bony endocarp (stone) in 1-4 cells, though it is rare for all 4 cells to be fertile and each fruit usually has only 2-3 seeds.

Phenology

G. arborea sheds its leaves from the end of January to March. The flowers and new leaves are seen together in April-May after the leaf shedding period. Flowering time varies according to location, individual tree, and even between inflorescences within trees. The fruits ripen from the end of April to June or July. Correlations of flowering time and fruit ripening with latitude or forest type have not been detected (Lauridsen, 1977). 

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).

Review of Natural Distribution

G. arborea is indigenous to India, Pakistan, Bangladesh, Myanmar, Sri Lanka, Thailand, Laos, Cambodia, Vietnam, and Yunnan and Guangxi provinces in China (Troup, 1921; Moldenke, 1971; Greaves, 1981; Gupta, 1993; Luna, 1996). It is found in deciduous forest and moist deciduous forest, but also sometimes occurs in evergreen and Shorea robusta forests. It is seldom found above 1200 m altitude in the Himalayan region (Troup, 1921), but it has been observed above 1500 m in the Sri Lankan moist forests (Greaves, 1981).

Location of Introductions

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).

Distribution Map

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Distribution Table

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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

Introduced toIntroduced fromYearReasonsIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
BrazilAsia1960s YesNo 
Costa RicaAsia1966 YesNo 

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).

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

Pathway causeNotesLong distanceLocalReferences
Disturbance (pathway cause)Naturalized in disturbed sites and along roadsidesYesYes
Escape from confinement or garden escape (pathway cause)Fruits –escaped from plantations and colonizing wild habitatsYesYes
Forestry (pathway cause)Plantation timber productionYesYes
Habitat restoration and improvement (pathway cause)Often used in large-scale reforestation programsYesYes
Horticulture (pathway cause)Often planted as shade tree in cacao and coffee plantationsYesYes
Medicinal use (pathway cause)Used in traditional Asian medicine Yes
Ornamental purposes (pathway cause)Planted as ornamental and shade tree in parks and avenuesYesYes

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).

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

CategorySub categoryHabitatPresenceStatus
Terrestrial    
TerrestrialTerrestrial – ManagedManaged forests, plantations and orchardsPrincipal habitatProductive/non-natural
TerrestrialTerrestrial – ManagedDisturbed areasPresent, no further detailsHarmful (pest or invasive)
TerrestrialTerrestrial – ManagedDisturbed areasPresent, no further detailsNatural
TerrestrialTerrestrial – ManagedDisturbed areasPresent, no further detailsProductive/non-natural
TerrestrialTerrestrial – ManagedRail / roadsidesPresent, no further detailsHarmful (pest or invasive)
TerrestrialTerrestrial – ManagedRail / roadsidesPresent, no further detailsNatural
TerrestrialTerrestrial – ManagedRail / roadsidesPresent, no further detailsProductive/non-natural
TerrestrialTerrestrial ‑ Natural / Semi-naturalNatural forestsPresent, no further detailsHarmful (pest or invasive)
TerrestrialTerrestrial ‑ Natural / Semi-naturalNatural forestsPresent, no further detailsNatural
TerrestrialTerrestrial ‑ Natural / Semi-naturalNatural forestsPresent, no further detailsProductive/non-natural

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

G. arborea grows best in the tropics, especially in low-lying areas. However, it is found naturally in subtropical climates up to 1200 m altitude in India (Greaves, 1981). The absolute maximum temperature varies from 38°C to 48°C. 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%. Average monthly temperatures generally between 18 and 35°C for the coolest and warmest months, respectively. Average annual rainfall generally between 900 and 2300 mm (Troup, 1921; Greaves, 1981; Webb et al., 1984).

A modified description of climatic requirements (see climatic data table of this data sheet) was prepared by CSIRO (see Booth and Jovanovic, 2000).

Soil and Physiography

G. arborea grows well on a range of soil types, particularly on sandy-loam soils with high site quality and good drainage. It is also found on shallow and rocky dry sites, where as a result of drought it can remain stunted, maintaining a shrubby habit. G. arborea does not grow on waterlogged sites or on peat soils.

Vegetation Types

mixed forests
moist forests
rain forests
secondary forests

Climate

Climate typeDescriptionPreferred or toleratedRemarks
Af - Tropical rainforest climate> 60mm precipitation per monthPreferred 
Am - Tropical monsoon climateTropical 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)
254011200

Air Temperature

ParameterLower limit (°C)Upper limit (°C)
Absolute minimum temperature5 
Mean annual temperature2128
Mean maximum temperature of hottest month2440
Mean minimum temperature of coldest month1424

Rainfall

ParameterLower limitUpper limitDescription
Dry season duration16number of consecutive months with <40 mm rainfall
Mean annual rainfall7504500mm; lower/upper limits

Rainfall Regime

Summer

Soil Tolerances

Soil texture > light
Soil texture > medium
Soil reaction > acid
Soil reaction > neutral
Soil drainage > free
Soil texture > heavy
Soil texture

Soil Types

alluvial soils
ferralsols
granite soils
red soils
subtropical soils
tropical soils
ultisols

Notes on Pests

Severe pest and disease problems are frequent in plantations within the natural distribution of G. arborea and have even led to the failure of plantations (Homfray, 1937; Allsop, 1945). Insect pests that are common in G. arborea plantations within the natural distribution area are stem borers, leaf defoliators and leaf or shoot cutters. Stem borers include Acalolepta cervina (Beeson, 1961) in India and Myanmar; leaf defoliators include larvae of Calopepla leayana in Assam, Bengal, Meghalaya and Myanmar, and nymphs and adults of Tingis beesoni in India and Myanmar (Greaves, 1981; Sarma and Rahman, 2010). 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).
Although G. arborea planted outside its natural range suffers similar pest problems, on the whole these are less serious. Leaf defoliators on seedlings and in mature stands are very common. In Nigeria alone, Roberts (1969) noted that defoliators such 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 Latin America, leaf cutting ants are a major problem for stem quality and growth of plantations (Greaves, 1981). In Malaysia, the beehole borer Duomitus ceramica is a damaging insect pest (Ahmad Said, 1989). Chey (1996) reported that Coptotermes curvignathus, a termite pest that severely damages G. arborea plantations in Malaysia, can be controlled by the application of the termiticide chlorpyrifos.
Fungal attacks are less prevalent, but can cause damage to G. arborea stands. Bagchee (1952) recorded a leaf spot leading to defoliation caused by a Gnomonia sp., while in Kerala (India) various Coniella species were identified as causing foliar diseases (Mohanan et al., 2010). A Poria species attacks G. arborea in India and Bangladesh during waterlogging (Bagchee, 1952, 1953). A powdery mildew, Phyllactinia suffulta var. gmelinae, attacks leaves but does not seriously affect the whole tree. Fusarium wilt is a problem for seedlings, but can be suppressed by applying various combinations of biocontrol agents, organic matter and biofertilizers (Singh et al., 2003).
Outside the natural distribution of G. arborea, fungal diseases are mainly root diseases found in Africa and Latin America. Root diseases of nursery seedlings are normally caused by Gibberella fujikuroi in Gambia and Sclerotium rolfsii [Athelia rolfsii] in Sierra Leone, Gambia and Nigeria (Gibson, 1975). Thanatephorus cucumeris, Chaetophoma sp., Polyporus and Armillaria mellea are also causal agents of root diseases in Nigeria and Côte d'Ivoire (Gibson, 1975). In Amazonia, Ceratocystis fimbriata, which is an important pathogen of rubber (Hevea brasiliensis), mango (Mangifera indica), coffee and cocoa, also infects G. arborea; the vectors of the fungus are insects of the genera Scolytus and Platypus (Muchovej et al., 1978). In Malaysia, diseases such as leaf spot, collar-rot and wilt have been identified, caused respectively by the fungi Colletotrichum gloeosporioides [Glomerella cingulate] (leaf-spot disease), Rhizoctonia solani (collar-rot disease) and Pythium spp. (wilting disease); treatments to overcome these diseases are prescribed by Maziah and Norani (1988). Stem rot and anthracnose are common seedling diseases in Malaysia; these can be controlled (Lee and Goh, 1989).
In Zaria (Nigeria), individual trees showed severe infestation by mistletoe (Tapinanthus sp.) (Nwanosike, 2005).

List of Pests

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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).

Natural enemies

Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Armillaria mellea (armillaria root rot)Pathogen
Other/All Stages
not specific   
Ceratocystis fimbriata (Ceratocystis blight)Pathogen
Other/All Stages
not specific   
Gibberella fujikuroi (bakanae disease of rice)Pathogen
Seedlings
not specific   
Athelia rolfsii (sclerotium rot)Pathogen
Seedlings
not specific   
Globisporangium splendens (blast of oil palm)Pathogen
Seedlings
not specific   
Fusarium oxysporum (basal rot)Pathogen
Seedlings
not specific   
Thanetophorus cucumerisPathogen
Seedlings
not specific   
Craspedonta leayanaHerbivore
Leaves
to species   
Acalolepta cervina (coffee longhorn)Herbivore
Other/All Stages
not specific   

Impact Summary

CategoryImpact
Economic/livelihoodPositive and negative
Environment (generally)Positive and negative
Human healthPositive

Impact: Environmental

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

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
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 successional patterns
Monoculture formation
Reduced native biodiversity
Threat to/ loss of native species

Impact mechanisms

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

Likelihood of entry/control

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: Wood Uses

G. arborea produces a highly-prized cream-coloured timber known for its easy working and good finishing properties. Untreated G. arborea timber is not very durable and had slightly more than one year life in a graveyard test, but it is readily treated with creosote (Lee, 1964). It has good working properties, including for peeling, gluing and seasoning, etc. (Wong and Khoo, 1980). Details on sawing, woodworking, peeling, gluing and kiln drying of G. arborea have been extensively recorded (Peason and Brown, 1932; Thomas, 1939; Lee, 1964; Lee et al., 1974; Lopez, 1977; Wong and Khoo, 1980). The suitability of various types and concentrations of wood preservatives was determined by Lee (1964). When the timber of G. arborea is adequately protected, it is excellent for house posts, boat decks, furniture, boxes, musical instruments, cars, railway carriages, matches, curing tobacco, building, panels and general carpentry work. It has also been used to manufacture plywood (Lamb, 1968; Limaye, 1934; Wong and Khoo, 1980; Greaves, 1981) and wood wool cement board with high resistance to subterranean termite attack (Garcia et al., 2012). G. arborea can easily be pulped and produces paper for various uses, including wrapping, writing and printing. Pulping methods are described in Peh (1964). Pulp yield is generally high (Phillips et al. 1978; Villaneura et al., 1970). G. arborea wood can be pyrolysed to produce bio-oil and bio-char (Okoroigwe et al., 2012).
G. arborea in Malaysia had superior strength properties when compared with G. arborea grown in Myanmar and India (Wong and Khoo, 1980).

Uses: Non-Wood Uses

Various parts of the tree are used in traditional medicine, in both its native range and introduced range, e.g. Nigeria (Idu and Onyibe, 2007). 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.
Bark extracts have also been used to control crop pests (Mbonu, 2010; Oparaeke et al., 2010), while wood ash was equally as effective as commercial products in controlling nematodes (Meloidogyne spp.) in cowpea crops (Ononuju and Nzenwa, 2011).
Leaves have potential as feed for goats (Abdu et al., 2012).

Uses: Land Uses

G. arborea is much used in agroforestry systems involving intercropping with arable crops, coffee or fruit trees (Arteaga Martínez and Castelán Lorenzo, 2008; Swamy et al., 2008; Vanlalhluna and Sahoo, 2009, 2010; Dhillon et al., 2011; Miole et al., 2011), where it has been shown to give better growth and yield than in sole plantations (Banerjee and Dhara, 2010). In intercrops with maize, a high pruning intensity and/or widely spaced tree rows (10 m or more) are recommended to reduce the suppressive effect of G. arborea on the crop, thus increasing maize yields and farm income (Bertomeu et al., 2011, 2012). G. arborea is recommended for homestead forests in Bangladesh in agroforestry situations with fruit trees (Motiur et al., 2005).
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).

Uses List

Ornamental
Ornamental > Christmas tree
Ornamental > Cut flower
Ornamental > garden plant
Ornamental > Potted plant
Ornamental > Propagation material
Ornamental > Seed trade
Environmental > Agroforestry
Environmental > Erosion control or dune stabilization
Environmental > Revegetation
Environmental > Shade and shelter
Materials > Carved material
Materials > Essential oils
Materials > Fibre
Materials > Miscellaneous materials
Materials > Wood/timber
Medicinal, pharmaceutical > Source of medicine/pharmaceutical
Medicinal, pharmaceutical > Traditional/folklore
Fuels > Charcoal
Fuels > Fuelwood
Environmental > Amenity
Materials > Dyestuffs
Animal feed, fodder, forage > Forage
Human food and beverage > Honey/honey flora

Wood Products

Boats
Charcoal
Containers > Boxes
Containers > Crates
Furniture
Other cellulose derivatives
Pulp > Short-fibre pulp
Railway sleepers
Sawn or hewn building timbers > Carpentry/joinery (exterior/interior)
Sawn or hewn building timbers > Exterior fittings
Sawn or hewn building timbers > Fences
Sawn or hewn building timbers > Flooring
Sawn or hewn building timbers > For light construction
Sawn or hewn building timbers > Wall panelling
Veneers
Wood extractives (including oil)
Wood hydrolysates
Wood residues
Wood-based materials > Plywood
Woodware > Matches
Woodware > Musical instruments

Silviculture Characteristics

G. arborea is rarely gregarious, although it grows well in pure plantations. It is a pioneer species and prefers full sunlight, although it can withstand partial shade; it usually colonizes gap openings in disturbed forests. It is moderately drought and frost tolerant, but severe drought and frost may kill seedlings. The shedding of leaves after drought is also common in this species. After severe drought damage to the main stem, G. arborea can still produce shoots which will grow vigorously in subsequent favourable periods (Troup, 1921; Greaves, 1981).  The permanent wilting point has been suggested as a criterion for assigning clones to sites differing in water availability (Mariño Macana and Rodríguez Melo, 2010).  The ability to coppice is an added advantage for G. arborea as a timber plantation species.
G. arborea can flourish on wasteland, roadsides and occasionally in patches of urban secondary forest.

Silviculture Characteristics

Tolerates > shade
Ability to > regenerate rapidly
Ability to > coppice
Ability to > pollard

Silviculture Practice

Artificial regeneration

G. arborea planting stock can consist of seedlings, stump plants, cuttings or coppice shoots. Good-quality seedlings usually derive from high-quality seeds. The abundance of fruiting can be improved by treating the mother trees with Ethrel (ethephon) and girdling (Wong, 1987). A total of 14,000 freshly depulped seeds weigh about 1 kg and a kilogram of fruits produces about 60 g of clean seed. Freshly-collected seeds have a germination rate of 90% if sown directly after harvesting (Troup, 1921). Seed stands 3- to 7-years-old can yield viable seeds throughout the year, but the peak fruit season only occurs in January-February (Chinte, 1971; Rogers, 1957; Pringle, 1950; Barnard, 1953; Greaves, 1981). Seeds can be harvested directly from the tree or collected from mature fruits on the ground (Barnard, 1953). The better quality seed is harvested from fruit on the ground, rather than from trees bearing immature fruits (Aminuddin and Zakaria, 1980). Seed processing before germination involves depulping, cleaning and storing (Barnard, 1953; Woessner and McNabb 1979; Greaves, 1981). The seeds collected should be depulped within two days after collection and sown immediately to obtain maximum germination (Troup, 1921; Ogunnika and Kadeba, 1993). In Malaysia, fruits need to be depulped within 24 hours after collection to avoid fermentation; details of the depulping process are given in Sim and Wong (1985). However, if depulping is done immediately and seeds are properly dried at 45°C for 24 hours to 8-10% moisture content, the treated seeds can be stored in sealed containers for two years without affecting viability (Woessner and McNabb, 1979). A methodology for assessing seed lot vigour was developed by Pandey and Sharma (2007).
Germination of G. arborea seeds occurs 7-15 days after sowing; the sowing medium should be kept wet and well-aerated throughout the period (Chinte, 1971; Homfray, 1937; Troup, 1921). Solarization of nursery seedbeds eliminates pests and pathogens and increases seedling height and collar diameter (Verma et al., 2010). It has been noted that there is a higher percentage of seeds germinating in full sunlight and that shading inhibits the process (Aminuddin and Ng, 1982). Large fruit stones weighing >1 g have a high germination rate (Woessner and McNabb, 1979). The optimum temperature for maximum total germination (about 78%), has been observed at 27°C, with low germination at 10-12°C (Ng and Hellum, 1985). Seeds at the yellow-brown stage of maturity with the best germinative traits after soaking in water for 48 hours are recommended for producing the most robust seedlings (Adebisi et al., 2011).
Seedlings should be transplanted to a good potting mixture (one part organic material, three parts sand and two parts topsoil), to provide good aeration and drainage. Potted seedlings are kept in the nursery until reaching 25-30 cm tall. Application of nitrogen, phosphorus and potassium (15:15:15) fertilizer at monthly intervals is necessary to ensure good growth of the potted seedlings. Root trainer size has a significant effect on growth (Durai, 2012). Darus (1984) provides a detailed description of the nursery techniques used for propagation of G. arborea by seed at FRIM, Malaysia.
Stump plants are a popular planting stock because of their relatively high survival rates in the field (Setten, 1953; Greaves 1981). The quality of stump plants is dependent on their seed quality. One-year-old seedlings growing on germination beds are kept for producing stump plants. Details on preparation of the germination bed for stump plant production are well documented (Pringle, 1950; Jackson, 1974). The germination bed should be slightly raised to about 40 cm and 1.2 m wide; four sowing lines are set out at 23 to 30 cm apart (Rajkhowa, 1964). Sowing must be timely so that the seedlings can obtain a collar diameter of 2.5 cm and height of 1 m before the planting season, but seedlings of smaller size should be rejected for stump plant production. Stumps 7- to 10-months-old have shown good survival in field planting (Zamora and Agpaoa, 1976). The seedlings will then be lifted, pruned back to about 5 cm of stem and 20 cm of root to produce stump plants; these are then tied in bundles and wrapped in wet sacks for transport to the planting site.
Vegetative cuttings have been successfully propagated. Vegetative materials used for cutting are sprouts from stock plants (Romero, 1966) and stems (Sabado and Valiente, 1972; Rahman, 1977; Zabala, 1977). Stem cuttings are preferred over other vegetative parts because they produce better planting stock (Itam et al., 1986). Plant growth regulators such as IBA improve the rooting of stem cuttings (Hamsawi and Srivastava, 1985). Cuttings of G. arborea have been successfully produced by treating stem cuttings with 60% NAA (Surendran and Seethalakshmi, 1987). Protocols for in vitro micropropagation from nodal tissue have also been developed (Gamboa and Abdelnour, 1999; Behera et al., 2008; Mishra and Shirin, 2009).
G. arborea coppice is a potential source of planting stock. Studies in Malaysia indicated that coppice shoots derived from a 6-year-old G. arborea stand had good potential to form the next plantation crop (Afzal-Ata and Muhamad, 1987).

Planting/sowing

Planting using stump plants or seedlings requires good site preparation. Establishment of G. arborea necessitates removal of existing vegetation and a clean site void of Imperata cylindrica (Durant, 1941; Setten, 1953). Burning of felled vegetation is recommended for improving site fertility (Durant, 1941; Setten, 1953), although it can encourage erosion on some soils. Planting must be done in the early rainy season to ensure good survival and establishment of transplanted seedlings (Greaves, 1981). Stump plants are popular for plantation establishment (McEwan, 1961) due to their drought tolerance, relatively low cost and ease of planting. Stump plants require shoot pruning after sprouting to maintain a single leader.
A high density of direct sowing (50 kg/ha) gives rise to successful establishment and good growth of seedlings (Greaves, 1981); seedlings can attain a height of 2.4-2.7 m six months after direct sowing. In the Philippines, direct sowing of seeds in a scratch line, covered with soil and weeded quarterly showed good survival (72%) and growth rate, with seedlings attaining 1.8 m in height one year after sowing (Chinte, 1971). Direct sowing of G. arborea is not encouraged on Imperata cylindrica-infested sites. Differences in survival rates on poorly- and well-burned sites where Imperata cylindrica thrive have been recorded, with a higher survival of seedlings on the latter (Dawkins, 1919).

Spacing

Initial spacing of G. arborea stands varies according to management objective. For chip or pulpwood production, G. arborea can be planted at 1.2 x 1.2 m to 1.8 x 1.8 m. For sawlog production wider spacing is anticipated; an initial spacing of 3 x 3 m is recommended for sawlog production in Malaysia (Wong and Khoo, 1980; Greaves, 1981; Gupta, 1993; Luna, 1996). Spacing of 2.5 x 2.5 m ensures early suppression of weed growth and 4.5 x 4.5 m decreases the form quality of stands.

Tending

Weeding is a very important silvicultural practice for enhancing the survival and growth of G. arborea. Weeds must be removed prior to planting (see above) and an intensive weeding regime should be implemented until crown closure (Barnard, 1953; Mitchell, 1962). In Malaysia, studies have shown that planting G. arborea using the taungya system was successful where intensive weeding and early pruning were carried out by farmers (Durant, 1941). Weeding regimes differ depending on the severity of weed infestation. Three to four rounds of circular weeding are required to ensure a high survival of seedlings. In Colombia, 2 rounds of chemical weed control plus a high dose of NPK-Mg fertilizer gave the best growth response in young trees (Barrios et al., 2011).
Intensive care and frequent fertilizer application will help the establishment of G. arborea seedlings (Maun, 1977; Agpaoa and Zamora, 1976; Kamis and Ismail, 1986). Low growth rates for G. arborea have been noted on poorer soils and ridge site tops which are less fertile than foothills (Kamis and Ismail, 1986). The high nutrient demand of G. arborea is illustrated by its high accumulation of nitrogen, phosphorus and potassium (Helenda, 1993). Ogbonnaya (1994) and Ezema (1989) showed the effects that different nutrients have on G. arborea growth, while Raman et al. (2008) showed that high levels of mineral fertilizer and farmyard manure promote seedling growth and quality. Mendoza and Glori (1976) discuss the importance of quantity and timing of fertilizer application for successful development of G. arborea, while. Jackson (1973) showed that seedlings grow best with relatively small doses of urea. It was also shown that good soil water availability and high organic material content favour root system development (Mbakwe, 1989).
Thinning is usually undertaken only in plantations producing veneer and sawlog timber. Thinning intensity depends very much on the initial spacing and there is a trade-off with the weeding intensity required. Fox (1967) suggests that plantations with 2.1 x 2.1 m and 3.7 x 3.7 m spacing be thinned when the mean height reaches 6 and 9 m, respectively, usually within 3-4 years after planting, reducing the stock to 740 stems per hectare at 9 m and 125 stems per hectare at 15 m. In northern Brazil, growing stock control is also used to regulate the thinning regime, e.g. stocking of 900-950 stems per hectare and 800-1000 stems per hectare at five years after planting for good and poor sites, respectively (Kalish, 1979).

Silviculture Practice

Seed storage > orthodox
Vegetative propagation by > cuttings
Vegetative propagation by > stump plants
Vegetative propagation by > grafting
Vegetative propagation by > tissue culture
Stand establishment using > natural regeneration
Stand establishment using > direct sowing
Stand establishment using > planting stock
Stand establishment using > wildings

Management

In its natural habitat, such as in Myanmar, it is common to find individual trees attaining a diameter of 1.4 m, with 15 m of clear bole and a top height of 30 m (Rodger, 1913). On good sites, it can easily attain an average top height of 25 m and diameter at breast height (dbh) of 20-30 cm at 15 years old.
G. arborea is very sensitive to site quality; studies show that it does not grow well in adverse conditions such as waterlogged sites (Durant, 1941), aluminium-contaminated sites (Fasehun, 1975) or sites affected by mild frost (Greaves, 1981). On good-quality sites, annual yields of G. arborea can range from 2 to 5 cubic metres per hectare. The highest value recorded in the Philippines is 12 cubic metres/hectare per year. Mean annual volume increments of as low as 0.9 cubic metres per hectare have been recorded on poor sites.
G. arborea responds well to increased CO²concentration, with a 30% increase in photosynthetic rate, higher growth rate and above-ground biomass production, and high carbon sequestration potential (Rasineni et al., 2011). 
The crown diameter of G. arborea is linearly related to dbh, with a curvilinear relation between crown diameter and total height (Freezaillah and Sandrasegaran, 1966). Bark thickness is related to dbh in a linear manner (Chinte, 1971a; 1971b); volume is linearly correlated with basal area (Greaves, 1981). The relation of stem volume to diameter is normally used to construct volume tables, e.g. for India (Chatuverdi, 1973; Sharma and Jain, 1977), Malaysia (Freezaillah and Sandrasegaran, 1966) and Nigeria (Okojie, 1976; Greaves, 1981; Akinnifesi and Akinsanmi, 1995).
The stem form quality of G. arborea is generally below the desirable standard. Its poor stem form reduces its popularity among sawlog timber planters (Barnard and Beveridge, 1957; Leggate, 1966; Tillman, 1975). Many studies provide interesting observations on the causes of poor stem quality in G. arborea. These are: the destruction of apical shoots by leaf cutting ants; stump plant stock produces several sprouts which form leaders; destruction of apical shoot by climbers; and poor genetic material. Poor form quality can be slightly improved by use of closer initial spacings. However, a proper system that allows intensive monitoring of growth must be in place; this ensures that severe competition, which reduces productivity and stand health, is avoided.
In plantation trials in Mexico, G. arborea has regularly outperformed other tree species; rates of annual increase in height and dbh were greater in G. arborea than in Tectona grandis and Acrocarpus fraxinifolius in the dry tropics of Michoacan (Muñoz Flores et al., 2009), and in a mixed plantation of eight tree species in Veracruz (López Ayala et al., 2010).
G. arborea has been successfully managed in Aragua (Venezuela) under small-scale forestry systems (Torres Lezama et al., 2010, 2011). In SE Bangladesh, coppice management has been traditionally practised by the Bwan tribal community (Mohiuddin and Alam, 2011). Managing plantations for fuelwood production, where good stem quality is not required, is discussed in Setten (1953). G. arborea is a promising species for pulpwood plantations in the tropics on a rotation of 7-8 years. G. arborea can be used as a nurse crop or nurse tree providing a more conducive environment for the establishment of climax species, such for as dipterocarps in Malaysia (Ang, 1994).

Genetic Resources and Breeding

G. arborea provenances have distinct variation in fruit size, seed and seedling characteristics, leaf size and petiole length (Arimah, 1979). Genetic variation for seed-related traits and response to in vitro culture has been observed in eight Indian provenances (Naik et al., 2009), while Wee et al. (2012) reported good levels of genetic diversity, highly polymorphic microsatellite loci and a deficiency of heterozygotes in 19 populations from India, China, Thailand and Myanmar. Genetic diversity and heterosis could be maximized by including selections from each geographic region in breeding programmes.
Presumably self-pollination is common for G. arborea, because large quantities of fruit are found on isolated trees. Self-pollination is supported further by the favourable structure of the flowers (Greaves, 1981). However, trees may also be insect pollinated.
Tewari (1995) reported that harvested seeds from a germplasm bank showed heterosis of 19.6-27.6% over their mid-parent for seed quality and 30% heterosis for seed germination. Lokmal (1994) suggests that a low intensity of mass selection followed by a high intensity of family selection may yield genetic gain for diameter and height growth in G. arborea. A trial of 70 clones from 10 sites in NE India at age 24 months showed significant differences for growth traits, indicating good potential for selection and improvement (Kumar, 2007). Occasionally, a super plus tree is encountered in G. arborea stands and can act as a source of genetic material for improvement (Greaves, 1981). Selection work in the Philippines has shown that at least 50 plus trees need to be included in any breeding programme, and selection intensity can be as low as one tree in 10,000 (Haque et al., 2001). Cooperative provenance/progeny tests involving the evaluation of 60 open-pollinated Thai families in 4 countries gave genetic gains and production gains in the best 30 trees using family and within family selection of 39% and 31% above the overall test means, respectively (Wijoyo, 2001). An account of propagation protocol evaluations and provenance/progeny trials being conducted by the Ecosystems Research and Development Bureau in the Philippines is given by Cadiz et al (2010).

Disadvantages

The tendency for development of poor stem form in G. arborea should be noted.

Links to Websites

NameURLComment
GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gatewayhttps://doi.org/10.5061/dryad.m93f6Data source for updated system data added to species habitat list.

References

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