Eucalyptus regnans

Family

Myrtaceae

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Chromosome number

2n = 22

Synonyms

Vernacular names

Mountain ash, Australian mountain ash, swamp gum, stringy gum, Victorian ash, white mountain ash (En).

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Origin and geographic distribution

Eucalyptus regnans is native to south-eastern Australia (Victoria and Tasmania), where it occurs in original forests and extensive plantations. It is widely planted in subtropical regions around the world and in the mid-altitude tropics, for instance in the United States, Chile, Spain, India and Sri Lanka. In New Zealand it is one of the primary plantation species. In Africa Eucalyptus regnans has been planted in Ethiopia, Kenya, Tanzania, Zimbabwe and South Africa.

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Uses

Eucalyptus regnans is the main Eucalyptus species used by the Australian pulp industry, and is a favoured pulpwood plantation species in southern Africa, New Zealand and Argentina. The timber (trade name: Tasmanian oak) is used for a wide range of purposes, including light and heavy construction, flooring, boat building, joinery, interior trim, furniture, agricultural implements, tool handles, kitchen utensils, poles, crates, matches, turnery, veneer, plywood, fibreboard, particleboard and wood wool. It is also used as fuelwood and for charcoal making. Waste material (leaves, branches, bark) from Eucalyptus regnans plantations can be collected and used as fuel. Planted strips of Eucalyptus regnans can be used as windbreaks. Bees make honey from its nectar. An essential oil can be extracted from the leaves, but the yield is so low that the species has little value as an essential-oil-producing tree.

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Production and international trade

Worldwide Eucalyptus plantations were estimated in 1995 at 14.6 million ha, of which 1.8 million ha were in Africa. Much of this was in South Africa. Very little of the world or African production of Eucalyptus wood and pulp comes from Eucalyptus regnans, but it is an important species in Tasmania and New Zealand.

Properties

Eucalyptus regnans is considered one of the best Eucalyptus species with respect to pulp yield and quality, for all types of mechanical and chemical pulping processes. A study in Australia on the effect of tree age on pulp quality indicates that pulp quality improves up to 20–60 years and remains fairly constant thereafter. The wood fibre cells are 0.5–1.0 mm long and 18–280 μm wide. The chemical composition of the fibres is: cellulose 49%, hemicelluloses 15%, lignin 28% and ash 0.4%.
The heartwood is straw-coloured to pale reddish brown, and often not clearly differentiated from the paler, up to 2.5 cm wide sapwood. The grain is usually straight, texture coarse. Growth rings are often prominent. Quarter-sawn surfaces have an attractive figure. The density is 510–720 kg/m³ at 12% moisture content, depending on age of the tree and growing conditions, with the wood of plantation-grown trees being less dense than that from natural stands. For instance, the density at 12% moisture content of wood from Australian natural forest is 650–720 kg/m³, while that recorded for wood from African plantations is 510–570 kg/m³. Due to the relatively low density of the wood, air-drying proceeds faster than in other Eucalyptus species, and the sapwood is easily treated with creosote or other chemical anti-rotting agents. The rates of shrinkage are high: from green to 12% moisture content 7% radial and 14% tangential. Common problems with the wood include growth tensions, and extreme shrinkage and collapse at drying, leading to bending and checking. Boards are usually quarter-sawn to reduce checking.
The wood is moderately strong, tough and hard. At 12% moisture content, the modulus of rupture is 110 N/mm², modulus of elasticity 16,000 N/mm² and compression parallel to grain 63 N/mm². The wood is easy to work, with only moderate blunting of tools. It saws cleanly and sharply and moulds well. Planing with the grain gives a very smooth surface. It drills easily and cleanly, and holds nails well, though pre-drilling is recommended. The wood takes stains and finishes well and glues satisfactorily. It is fairly bendable, and peels and slices satisfactorily. The durability of the wood is low, and it is not suitable for outdoor or ground-contact applications. The wood is liable to attacks by marine borers and termites. The sapwood is susceptible to attack by Lyctus borers, and therefore milled timber is usually sold without sapwood. The heartwood is resistant to impregnation with preservatives, the sapwood is permeable.
The wood has an energy value of 19,700 kJ/kg at 12% moisture content, and charcoal made from it has an energy value of 28,000 kJ/kg. The wood burns well and leaves little ash, while the charcoal burns almost without smoke. Conversion of wood to charcoal using traditional methods leads to a loss of more than two thirds of its embodied energy. In Spain the energy value of waste material (leaves, bark, and branches) can reach 65,000 MJ/ha per year.
The essential oil extracted from the leaves and branchlets of Eucalyptus regnans is markedly different from that of other Eucalyptus species, because it contains almost no cineole. It is mainly composed of α-phellandrene, and also contains p-cymene, eudesmols and small amounts of the ester geranyl-acetate.

Description

Evergreen, very large tree up to 105 m tall; bole straight, up to 300 cm in diameter; bark rough and fibrous for the lower 15 m of the trunk, smooth and white or grey-green above, decorticating in long strips; crown open. Leaves alternate, simple and entire; stipules absent; petiole channelled, 8–25 mm long; blade lanceolate, 9–23 cm × 1.5–5 cm, slightly oblique or tapering at the base, acuminate at the apex, glossy, concolorous, green, venation oblique. Inflorescence a simple, umbel-like dichasium, paired in leaf axils, 9–15-flowered; peduncle angular, 4–14 mm long. Flowers bisexual, regular, white; pedicel 2–7 mm long; flower buds obovoid, 4–7 mm × 2–4 mm, divided into an obconical hypanthium (lower part) c. 3 mm × 3–4 mm, and a conical operculum (upper part) 2–3 mm × 3–4 mm, shed at anthesis; stamens numerous; ovary inferior, usually 3-celled. Fruit an obconical to pear-shaped capsule 5–9 mm × 4–7 mm, opening with 3 valves. Seeds pyramidal, 1.5–3 mm long, brown. Seedling with epigeal germination.

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Other botanical information

Eucalyptus comprises about 800 species, endemic to Australia, except for about 10 species in the eastern part of South-East Asia. Many Eucalyptus species are cultivated outside their natural distribution area, in tropical, subtropical and temperate regions, because of their rapid growth and adaptation to a wide range of ecological conditions. Eucalyptus is divided into several subgenera (7–10, depending on the author), which are subdivided into many sections and series. The results of phylogenetic studies within Eucalyptus suggest that the genus is polyphyletic, hence not of a single evolutionary origin, and consequently it has been proposed to divide the genus into several distinct genera. This has not yet been done, mainly because of the nomenclatural whirlpool this would bring about. Eucalyptus species hybridize easily, which adds to the taxonomic complexity.

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Growth and development

Eucalyptus regnans seedlings first send down a strong taproot, and later produce lateral roots. Growth is initially slow, but increases rapidly in the second year. In the sapling stage trees have narrow, conical crowns 9–12 m high, lateral roots spread far and send down sinker roots. In the pole stage the tree is c. 30 m tall, with a conical crown, the sinker roots become branched and the taproot dies back. In the spar stage the crown spreads out and the tree reaches almost full height while the lateral and sinker roots become very well developed. In the mature stage the crown opens and starts to die back, and buttresses are present at the base of the tree. Overmaturity is characterized by trunk epicormic growth and progressive die-back of the central areas of crown and roots. Tree ages of 350–400 years have been recorded.
Flowering is protandrous and the flowers are pollinated by insects or birds. Both outcrossing and self-pollination occur. In controlled-pollination experiments seedlings produced after self-pollination showed reduced growth compared to those raised from open-pollinated seed. The time from flowering to seed maturation is 8–10 months. Unlike many other Eucalyptus species, Eucalyptus regnans does not regenerate from lignotubers. This means that mature trees are sensitive to intense fires, and that coppice harvests are usually not possible after the first harvest.
Eucalyptus regnans is the tallest hardwood species in the world. In Kenya at 2400 m altitude 2-year-old trees with a height of 9 m have been recorded, and 35-year-old trees with a height of 80 m. In a small plantation at 3000 m altitude 30-years-old trees were up to 80 m tall. Growth measurements are available from Tanzania as well. In a stand in the Shagayu Forest Reserve (1900 m altitude, average annual rainfall 1200 mm) trees had a mean height of 26.3 m and a mean diameter of 22.2 cm at age 8, and a mean height of 34.6 m and a mean diameter of 28.2 cm at age 12. In the Lushoto Arboretum (1500 m altitude, average annual temperature 18°C, average annual rainfall 1070 mm) 10-year-old trees were 32.0 m tall and 32.0 cm in diameter. In Olmotonyi (1850 m altitude) the average height of 13-year old trees was 29.8 m, and the average diameter 24.7 cm. In the Kigogo Arboretum (2000 m altitude, average annual rainfall 1800 mm) 32-year-old trees were 44–49 m tall, with a diameter of 65–89 cm.

Ecology

Eucalyptus regnans prefers a cool climate. It grows in areas with a mean annual temperature of 7–20°C, a mean maximum temperature of the warmest month of 17–29°C, and a mean minimum temperature of the coldest month of –2 to +10°C. The average annual rainfall is 650–2000 mm, with a dry season of 0–2 months. It is intolerant of drought and severe cold (below –7°C). In its natural distribution area it occurs from sea level up to 1100 m altitude, while in tropical countries it usually grows at 2000–3200 m altitude. In provenance trials with 18 Eucalyptus species at 0–2800 m altitude in the wet tropics of Antioquia, Colombia, Eucalyptus regnans was the best performer at 2400 m altitude, and one of the three best at 2800 m altitude. It requires a moderately fertile, well-drained soil, and is sensitive to salinity.
Like many other Eucalyptus species, Eucalyptus regnans is very intolerant of competition for light and soil resources, which means that seedlings do not grow well under a pre-existing canopy. For this reason natural forests of Eucalyptus regnans are said to be self-thinning, i.e. the tallest, most dominant individuals shade out smaller individuals, which die. In native forests in south-eastern Australia Eucalyptus regnans grows in pure stands or as the canopy species, with an understory of Acacia melanoxylon R.Br. or Acacia dealbata Link.
Eucalyptus regnans has allelopathic effects on understory plants, especially on its own seedlings growing near mature trees or in fields in which mature trees have grown. These effects are alternately believed to be due to a buildup of waxy lipids in the soil from decaying Eucalyptus regnans leaves, or to a rhizosphere fungus, Cylindrocarpon destructans. The resulting difficulty of re-planting plantations of Eucalyptus regnans after harvest is a likely reason for the preference of Eucalyptus globulus Labill. in many plantations. However, seedling growth of Eucalyptus regnans is much stimulated when soil is exposed to fire or sterilization, which may be due to reduction or breakdown of inhibiting fungi or compounds in the soil. Hence natural stands are usually even-aged and can grow for centuries until they are stricken by a catastrophic fire, which kills all standing trees and stimulates massive seed germination. There is high mortality among these new seedlings and saplings, eventually leaving once again a mature, even-aged forest, which grows until the next periodic fire strikes.
The absence of lignotubers may be considered an advantage in terms of the ecological profile of Eucalyptus regnans outside of its native range. Eucalyptus species are often feared as invasive plants, and the lignotubers of many species make eucalyptus stands hard to eradicate once established. This risk is not present with Eucalyptus regnans, though mature trees are prodigious producers of seed.

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Propagation and planting

Eucalyptus regnans is usually propagated by seed. The 1000-seed weight is 5.5–12.5 g. Seed can be stored for several years in airtight containers under dry (5–8% moisture content) and cool (3–5°C) conditions. Cold and moist stratification of the seed at 3–5°C for 3 weeks is recommended to break dormancy and promote even germination. Germination is optimal at 15–20°C and usually takes 10–20 days. Seedlings raised in nurseries can be planted out after 8–9 months. In New Zealand Eucalyptus regnans seedlings are often grown in nursery beds for a year before bare-rooted seedlings are planted out in the field. Growing in polyethylene bags in the nursery is also possible. In-vitro propagation with microcuttings obtained from node-derived shoot cultures is possible.
Planting distances in the field usually range from 2 m × 2 m (2500 trees/ha) to 3 m × 3 m (1100 trees/ha). Planting for high-quality, large sawlogs must allow more space for each tree, and must be accompanied by pruning and thinning regimes. Closer spacing and higher densities can be used if the intended end product is pulp, fuel, or small posts, though rows of 3 m are necessary for mechanization, and a spacing of 3 m × 3 m is necessary for mechanized weeding in two directions. Another common strategy is to burn logging debris after harvest of a stand of Eucalyptus regnans and broadcast a large amount of seed (about 1 kg/ha) on the ash bed, eventually thinning the many trees that germinate and grow.

Management

Weeding between rows (possibly mechanical) and hand weeding near seedlings is very important in the first year of growth, but becomes less critical after canopy closure. Young plants are commonly, but not universally, fertilized. In Tasmania fertilization with N greatly augmented growth, but recommended quantities vary with site conditions. High fertilizer rates risk acidifying the soil. Pruning is often considered to be unnecessary for Eucalyptus species, as they are largely self-pruning. However, allowing trees to self-prune by dropping branches can decrease wood quality by leaving knots and entrance points for decay. This is not a problem in plantations for pulp or fuel, but it can decrease the value of timber. Pruning of young branches can greatly improve timber quality by limiting knots to the innermost wood of the trunk. Pruning of young branches should coincide with canopy closure, as pruning before this point can slow down growth. The coppicing ability of Eucalyptus regnans is poor due to the absence of lignotubers.

Diseases and pests

Phytophthora fallax is a fungal crown dieback disease of Eucalyptus regnans that occurs episodically in New Zealand, with individual trees showing differing degrees of susceptibility. Symptoms include leaf spots, petiole infection and branchlet lesions. Mycosphaerella cryptica is a leaf disease that can result in heavy defoliation and dieback. Certain provenances of Eucalyptus regnans are more susceptible than others, which implies that there is potential for breeding for resistance to the disease. No fungicides are known to give good control. Some fungi mainly affect seeds and seedlings, such as Penicillium spp., Fusarium spp., Phytophthora spp. and Botrytis cinerea, but they are not widely reported on Eucalyptus regnans, and good phytosanitary practices suffice to avoid them.
Eucalyptus regnans is one of the Eucalyptus species susceptible to the eucalyptus snout beetle (Gonipterus scutellatus), which occurs on all continents and is known to be present in Kenya, Uganda, Zimbabwe, South Africa, Mauritius and Madagascar. The eucalyptus snout beetle can be controlled with the hymenopterous parasite Anaphoides nitens. The longicorn beetle Phoracantha semipunctata is a borer that attacks young trees, stressed trees, and stumps of recently harvested trees. It has been recorded from Zimbabwe and South Africa. Insecticides are not effective, so it must be controlled by the removal and burning of infected material. The principal pest in Eucalyptus regnans plantations in Tasmania is Chrysophtharta bimaculata, a leaf eating beetle. It is controlled with an integrated approach involving insecticides and natural predators and parasites, together with strict monitoring of population numbers and breeding of more resistant trees. In Africa termites are a risk in Eucalyptus nurseries and young plantations up to 3–4 years old.

Harvesting

For densely planted pulp or fuelwood plantations, harvesting occurs 8–15 years after planting, while plantations for large sawlogs are harvested when the plantation is 30–40 years old and has been thinned at age 10–12. The bark can be removed before the tree is felled by ringing the bark and pulling it upward in strips. However, this is not usually necessary, as rough bark only occurs at the base of Eucalyptus regnans, and this bark is usable in the paper pulping process. For larger trees debarking may need to occur after felling and sectioning of the trunk. Two men working with chainsaws can cut 300–500 trees a day in a well-ordered plantation. Clear-cutting of areas to be replanted to seedlings of Eucalyptus regnans is recommended, as standing trees are shown to have a strong negative effect on seedling growth.

Yield

Under good conditions and with fertilization, mean annual volume increments of Eucalyptus regnans are 25–30 m³/ha. In a 42-year-old plantation in Kenya mean annual volume increments of 56 m³/ha have been recorded.

Handling after harvest

Felled trunks may be sectioned into shorter lengths in the field to ease transport, and leaves, branches, and other waste may be localized in a few piles to facilitate circulation of people and vehicles. Pruning of branches after felling is usually unnecessary, as little branching occurs in dense plantations. On larger plantations with trees going to an off-site sawmill or pulping mill, trees or sections are dragged or carried to a road by manpower, animals, or tractors. Once near the road, the trees are picked up by a truck and taken to the plant. Trunks or sections can be left on the ground to air dry over months in areas where termites and rotting are not a problem. Air-drying on a paved lot reduces the risks of biological attack of the wood, and drying under a roof slows drying by reducing exposure to direct sun. This reduces the amount of tangential checking that is to some extent inevitable during the drying of round logs.

Genetic resources and breeding

Breeding and provenance trial programs in Australia and New Zealand using progeny from hundreds of trees serve as a sort of in situ conservation of genetic diversity. In addition to this, natural and managed forests in south-eastern Australia and Tasmania preserve an estimated one third of the original natural genetic diversity of the species, as a large portion of them are the offspring of regeneration from original forest after fires or harvesting. Hence genetic erosion is not a serious threat, though the practice of replanting harvested forests with seedlings or direct seeding (common since 1960) amounts to a reduction of genetic diversity, since planted trees come from a limited number of accessions.

Breeding

The main (perhaps only) breeding programs for Eucalyptus regnans are in Australia and New Zealand, and focus on frost resistance and rapid growth.

Prospects

Eucalyptus regnans is not a major world plantation species, and Eucalyptus globulus is commonly used in the subtropical or high-altitude tropical environments well-suited to Eucalyptus regnans. Eucalyptus regnans is well-suited for the production of paper pulp, the most important product of industrial Eucalyptus globulus, but cannot substitute the latter as a source of eucalyptus oil. Little breeding work is done on Eucalyptus regnans, and hence most of the future world growth in eucalyptus planting and use will likely come from cosmopolitan species such as Eucalyptus camaldulensis Dehnh., Eucalyptus globulus and Eucalyptus grandis W.Hill ex Maiden. The unsuitability of Eucalyptus regnans wood for outdoor uses would somewhat limit its expansion in Africa. Another major issue is that Eucalyptus regnans does not possess lignotubers, and hence does not produce coppice regrowth. This is a disadvantage for intensive plantations because it means that trees must be replanted after every harvest, which, in combination with the poor growth of Eucalyptus regnans seedlings in soil from mature Eucalyptus regnans stands, may be the prime limitation to current and future use of Eucalyptus regnans in plantations.

Major references

• Ashton, D.H., 1975. The root and shoot development of Eucalyptus regnans F. Muell. Australian Journal of Botany 23(6): 867–887.
• Borota, J., 1991. Tropical forests: some African and Asian case studies of composition and structure. Developments in Agricultural and Managed-Forest Ecology No 22. Elsevier, Amsterdam, Netherlands. 274 pp.
• CPBR, 2006. EUCLID: Eucalypts of Australia. 3rd Edition. [Internet] Centre for Plant Biodiversity Research, Canberra, Australia. http://www.anbg.gov.au/ cpbr/cd-keys/euclid3/euclidsample/html/ Eucalyptus_regnans.htm. Accessed May 2009.
• Eldridge, K., Davidson, J., Harwood, C. & van Wijk, G., 1993. Eucalypt domestication and breeding. Clarendon Press, Oxford, United Kingdom. xix + 288 pp.
• FAO, 1958. Principal pests and diseases of eucalypts outside Australia. Unasylva 12(2): 77–79.
• Forest Products Commission, 2008. Timber Advisory Note No 11: Mountain Ash – Eucalyptus regnans. [Internet] Department of Conservation and Land Management, Como, Australia. http://www.agric.wa.gov.au/PC_92542.html?s=937741519. Accessed May 2009.
• Jacobs, M.R., 1981. Eucalypts for planting. 2nd Edition. FAO Forestry Series No 11. Food and Agriculture Organization of the United Nations, Rome, Italy. 677 pp.
• Keane, P.J., Kile, G.A., Podger, F.D. & Brown, B.N. (Editors), 2000. Diseases and pathogens of eucalypts. CSIRO Publishing, Collingwood, Australia. 565 pp.
• Montagu, K.D., Kearney, D.E. & Smith, R.G.B., 2003. The biology and silviculture of pruning planted eucalypts for clear wood production – a review. Forest Ecology and Management 179(1–3): 1–13.
• Verdcourt, B., 2001. Myrtaceae. In: Beentje, H.J. (Editor). Flora of Tropical East Africa. A.A. Balkema, Rotterdam, Netherlands. 89 pp.

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Other references

• Baker, R.T. & Smith, H.G., 1920. A research on the eucalypts especially in regard to their essential oils. [Internet] Authority of the Government of the State of New South Wales, Sydney, Australia. http://chestofbooks.com/ health/aromatherapy/Eucalyptus-Essential-Oils/ 161-Eucalyptus-regnans.html. Accessed May 2009.
• Blomstedt, C., Cameron, J., Whiteman, P. & Chandler, S.F., 1991. Micropropagation of juvenile Eucalyptus regnans (mountain ash). Australian Journal of Botany 39(2): 179–186.
• Booth, T.H., Nix, H.A., Hutchinson, M.F. & Jovanovic, T., 1988. Niche analysis and tree species introduction. Forest Ecology and Management 23: 47–59.
• CAB International, 2010. Forestry Compendium. Eucalyptus regnans. [Internet] http://www.cabi.org/ fc/?compid=2&dsid=22833&loadmodule=datasheet&page=2147&site=163. Accessed April 2011.
• Coppen, J.J.W., 2002. Eucalyptus: the genus Eucalyptus. Medicinal and aromatic plants - industrial profiles, vol. 22. Taylor & Francis, London, United Kingdom. 450 pp.
• Dick, M.A., Dobbie, K., Cooke, D.E.L. & Brasier, C.M., 2006. Phytophthora captiosa sp. nov. and P. fallax sp. nov. causing crown dieback of Eucalyptus in New Zealand. Mycological Research 110(4): 393–404.
• Dignan, P., King, M., Saveneh, A. & Walters, M., 1998. The regeneration of Eucalyptus regnans F. Muell. under retained overwood: seedling growth and density. Forest Ecology and Management 102(1): 1–7.
• Ilvessalo-Pfäffli, M.-S., 1995. Fiber atlas. Identification of papermaking fibers. Springer Verlag, Berlin, Germany. 400 pp.
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• Phillips, C., 1993. Longicorn beetles 1. Forest Health Information Fact Sheets No 17. PIRSA Forestry, Mount Gambier, Australia. 3 pp.
• Phillips, C., 1994. Chrysomelid beetles – Chrysophtharta spp. and Paropsis spp. Forest Health Information Fact Sheets No 21. PIRSA Forestry, Mount Gambier, Australia. 6 pp.
• Raymond, C.A., 1995. Genetic variation in Eucalyptus regnans and Eucalyptus nitens for levels of observed defoliation caused by the Eucalyptus leaf beetle, Chrysophtharta bimaculata Olivier, in Tasmania. Forest Ecology and Management 72(1): 21–29.
• Restrepo U., G. & Atehortua V., L.M., 1989. Some results from species and provenance trials of Eucalyptus species in the department of Antioquia, Colombia. Forest Genetic Resources 17: 20–23.
• Ringrose, C. & Neilsen, W.A., 2005. Growth response of Eucalyptus regnans and soil changes following periodic fertilization. Soil Science Society of America Journal 69: 1806–1812.
• Timber Research Unit, 2007. Tasmanian oak: Eucalyptus delegatensis, E. obliqua & E. regnans. [Internet] Tasmanian Timber Promotion Board. http://www.tastimber.tas.gov.au/ species/pdfs/ TasoakV4.pdf. Accessed May 2009.
• Webb, D.B., Wood, P.J., Smith, J.P. & Henman, G.S., 1984. A guide to species selection for tropical and sub-tropical plantations. 2nd Edition. Tropical Forestry Papers No 15. Commonwealth Forestry Institute, University of Oxford, United Kingdom. 256 pp.
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Sources of illustration

• Costermans, L., 1983. Native trees and shrubs of south-eastern Australia. Revised edition. New Holland Publishers, Sydney, Australia. 424 pp.

Author(s)

•	G. Vaughan Museo Arqueológico de Tunja, UPTC, Avenida Central del Norte, Tunja, Boyacá, Colombia

Editors

•	M. Brink PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands
•	E.G. Achigan Dako PROTA Network Office Africa, World Agroforestry Centre (ICRAF), P.O. Box 30677-00100, Nairobi, Kenya

Photo editor

•	G.H. Schmelzer PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands

Correct citation of this article

Vaughan, G., 2011. Eucalyptus regnans F.Muell. [Internet] Record from PROTA4U. Brink, M. & Achigan-Dako, E.G. (Editors). PROTA (Plant Resources of Tropical Africa / Ressources végétales de l’Afrique tropicale), Wageningen, Netherlands. <http://www.prota4u.org/search.asp>. Accessed .

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