Sandalwood Biblio - Cropwatch

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Cropwatch’s Sandalwood Bibliography. 

v1.13 June 2010. 

Note: in this bibliography, articles on sandalwood are arranged by relevance to 

geographical origin rather than being arranged species-by-species. More 

information on the ecological status of individual Santalum species & general 

notes are available on Cropwatch’s Updated List of Threatened Aromatic Plants 

Used in the Aroma & Cosmetic Industries. Please note also that where chemical 

formulae for certain sandalwood constituents are illustrated, some are added 

from the Cropwatch natural chemicals structure library. 

Contents: Australian Sandalwoods 

Biocidal properties 

Chemistry 

General articles 

Chinese Sandalwood 

East African Sandalwoods 

East Indian Sandalwood 

Biocidal propeties 

Contact dermatitis 

Cancer chemoprevention 

Chemistry 

General articles 

Indonesian Sandalwood 

Ogasawara Island Sandalwood 

Pacific Sandalwoods 

Cook Isllands 

Fiji 

Hawaii 

Marquesas Islands 

New Caledonia 

Tonga 

Vanuatu 

General Pacific Region 

Papua New Guinea Sandalwood 

Sri Lankan Sandalwood 

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Thai Sandalwood 

Timorese Sandalwood 

Unclassified Articles on Sandalwood 

Australian Sandalwoods. 

Cropwatch comments: Australia has two commercially important Sandalwood 

spp. (Santalum album & S. spicatum), and a number of other Sandalwood spp. of 

more minor economic importance (such as S. acuminatum, S. lanceolatum, and 

S. murrayanum). 

Biocidal Properties of Australian Sandalwoods 

Ritchie S.A., Williams C.R. & Montgomery B.L. (2006) "Field evaluation of New Mountain 

Sandalwood Mosquito Sticks and New Mountain Sandalwood Botanical Repellent against 

mosquitoes in North Queensland, Australia." J Am Mosq Control Assoc. 22(1), 158-60. Abstract. 

The mosquito repellent efficacy of New Mountain Sandalwood Mosquito Sticks (containing 0.5% 

w/w essential oils) and New Mountain Sandalwood Botanical Repellent (containing soybean and 

geranium oils) was assessed. Tests were conducted in the field with 4 volunteers in a wooded 

area near Cairns, North Queensland, Australia. Predominant biting species were Verrallina 

funerea and Ve. lineata. A pair of burning Mosquito Sticks immediately upwind of the subject 

(acting as an area repellent) provided a 73.1% mean reduction in mosquito landing and probing 

over the 3-h test period. The Botanical Repellent and a DEET-based control were both 100% 

effective in preventing mosquito probing for 3 h. These data are consistent with other studies of 

area repellents in that such products provide significant protection from mosquito bites, albeit 

inferior to the protection provided by topically applied repellents. 

Spafford H., Jardine A., Carver S., Tarala K., Van Wees M. & Weinstein P. (2007) "Laboratory 

determination of efficacy of a Santalum spicatum extract for mosquito control." J Am Mosq 

Control Assoc. 23(3), 304-11. Abstract. The activity of QN50, a sequiterpene alcohol derived from 

Australian sandalwood (Santalum spicatum), was tested for its effectiveness against larvae of 2 

mosquito species (Culex molestus and Aedes camptorhynchus [Diptera: Culicidael), nymphs of 2 

species of water boatmen (Micronecta robusta and Agraptocorixa [Hemiptera: Corixidae]), 

immature Daphnia sp. (Crustacea), and mosquito eggs (Cx. molestus). In a series of laboratory 

bioassays, field-collected mosquito larvae, eggs, and immature corixids and daphnids were 

placed in beakers with either QN50, methoprene or source water only (control). The mosquito 

larvae exposed to QN50 had reduced survivorship and average longevity relative to the control 

and to methoprene at most concentrations used in this study. The hatching rate of mosquito eggs 

was unaffected by methoprene or QN50. Corixid nymphs and daphnids experienced high 

mortality in both methoprene and QN50 relative to the control, but there was no difference in the 

effect between the compounds. The results of this preliminary study suggest that further research 

into the mode of action and efficacy of QN50 as a potential alternative to methoprene for 

mosquito abatement is warranted. 

Chemistry of Australian Sandalwoods 

Adams D.R., Bhatnagar S.P. & Cookson R.C. (1975) “Sesquiterpenes of Santalum spicatum” 

Phytochemistry 14(5-6), 1459-1460. 

Birch A.J., Moslyn K.M.C. & Penfold A.R. (1953) "The sesquiterpene alcohols of Eucarya spicata 

Sprague & Summ." Aust. J. Chem 6, 391-394. Cropwatch comments: Eucarya spicata Sprague 

& Summ. Is the outdated botanical name for Santalum spicata R.Br. 

Birch A.J., Chamberlain K.B., Moore B.P. & Powell V.H. (1970) “Termite attractants in Santalum 

spicatum.” Australian Journal of Chemistry 23(11), 2337-2341. Abstract. The oil of Santalum 

spicatum (R.Br.) A.DC. has been fractionated to yield 10-cis- (1) and 10-trans-2,6,10- 

trimethyldodeca-2,6,10-triene (2). These compounds have been synthesized by reduction of a 

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mixture of cis- and trans-farnesyl acetate. Although not identical with the trail pheromone of 

Nasutitermes they have similar specific trail activities, the former being the more active. 

Brand J., Kimber P. & Streatfield J. (2006). “Preliminary analysis of Indian sandalwood (Santalum 

album L.) oil from a 14-year-old plantation at Kununurra, Western Australia.” Sandalwood 

Research Newsletter 21. 

Braun N.A., Meier M. & Pickenhagen W. (2003) "Isolation & chiral GC analysis of beta-bisabolols 

- trace constituents from the essential oil of Santalum album L. (Santalaceae). J. Essent. Oil Res. 

15(1), 63-65. 

Braun N.A., Mieir M., Schmaus G., Holsher B. & Pickenhagen W (2003) "Enantioselectivity in 

odor perception: synthesis and olfactory properties of iso-beta-bisabolol, a new natural product." 

Helv Chim Acta 86(7), 2698-2708. Abstract. The odorous trace constituent iso--bisabolol (4) was 

isolated from East Indian and Western Australian sandalwood oil and synthesized by using the 

(E/Z)-triene 12 (iso--bisabolene) as a key intermediate. Only one of four stereoisomeric forms of 

4, (6R,7R)-4a, is odor active, having a strong floral, muguet-like, very pleasant scent. 

Braun N.A., Meier M., Kohlenberg B., Valder C. & Neugebauer M. (2003) “Santalum spicatum 

(R.Br.) A. DC. (Santalaceae) – nor-helifolenal and acorenol isomers: isolation & biogenic 

considerations.” J. Essen. Oil. Res. 15, 381-386. 

Braun N.A. & Spitzner D. (2007) “Synthesis and natural occurrence of (Z/E)-β-and γ-curcumen- 

12-ol." ARKIVOC (vii) 273-279. Abstract: (Z/E)-β-Curcumen-12-ol (Z/E)-(1) was synthesized via 

Birch reduction of acid 6 starting from α-curcumene (5). An olefin isomerization of 1 is the key 

step in the synthesis of (Z/E)-γ-curcumen-12-ol (Z/E)-(2). Sesquiterpene alcohol (E)-1 was found 

for the first time in nature as a minor constituent of different Santalum species by using the 

synthetic sample as reference. 

Bristow M., Taylor D. & Robson K. (2002) "Queensland Sandalwood (Santalum lanceolatum): 

regeneration following harvesting." Sandalwood Research Newsletter 2002. Abstract. In 1994, a 

trial, funded by Queensland Department of Primary Industries Forestry, was established near 

Hughenden investigating regeneration of natural stands of Queensland sandalwood from two 

harvestingmethods, vis, stump cutting vs. stump pulling. Merchantable size trees in five, one 

hectare plots wereharvested by the respective methods and vegetative regeneration was 

recorded over the successive fiveyear period. Overall indications are that retaining sandalwood 

stumps is unlikely to result in a greateramount or more successful coppice regeneration following 

harvesting than stump pulling, and that it may well result in less successful cpppice regeneration.. 

Data from the trial suggests that the propor-tion of pulled stumps that produce coppice is higher 

than the coppice produced through the cut stumpmethod, and these are more likely to survive. 

Concerns about the impact of stump pulling on soil prop-erties and erosion are unwarranted as 

the number of sandalwood removed from any area is relatively few and the area of soil disturbed 

during the operation is very small. 

Bristow M. (2004) "Review of Agroforestry in Tropical Savanna Regions of Northern Australia." A 

Report for the RIRDC/Land & Water Australia/FWPRDC/MDBC Joint Venture Agroforestry 

Program Mar 2004. "# 2.4 Ord River early sandalwood plantation projects." 

Brophy J.J., Fookes C.J.R. & Lassak E.V. (1991) “Constituents of Santalum spicatum (R. Br.) A. 

DC. Wood oil.” J. Essen. Record Res 3, 381-385. 

Jones G.P., Rao K.S., Tucker D.J., Richardson B.J., Barnes A. & Rivett D.E. (1995) 

“Antimicrobial activity of Santalum acuminatum (quandong) kernels.” International Journal 

Pharmacognosy 33, 120-123. 

Lawrence B.M. (2006) “Australian Sandalwood oil” in “Progress in Essential Oils” Perf & Flavorist 

31 (Sept 2005), 62-65. 

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Lawrence B.M. (2009) “Australian Sandalwood oil” in “Progress in Essential Oils” Perf & Flavorist 

34 (May 2009), 56. 

Liu Y.D., Longmore R.B. & Kailis S.G. (1995) “A comparison of kernel compositions of 

sandalwood (Santalum spicatum) seeds from different Western Australian locations. Mulga 

Research Centre Journal 12, 15-21. 

Liu Y.D., Longmore R.B., Fox J.E.D. (1996) “Separation & identification of ximenynic acid isomers 

in the seed oil of Santalum spicatum R. Br. as their 4,4-dimethyloxazoline derivatives.” Journal of 

the Americ. Oil Chemists Soc. 73(12), 1729-1731. 

Liu Y.D., Longmore R.B. & Kailis S.G. (1997) "Proximate and fatty acid composition changes in 

developing sandalwood (Santalum spicatum) seeds." Journal of the Science of Food and 

Agriculture 75(1), 27-30, 

Liu Y.D., Longmore R.B., Boddy M.R. & Fox J.E.D. (1997) “Separation & identification of 

triximenynin from Santalum spicatum R. Br.” Journal of the Americ. Oil Chemists Soc. 74(10), 

1269-1272. 

Loveys B.R., Tyerman S.D. & Loveys B.R. (2001) "Transfer of photosynthate and naturally 

occurring insecticidal compounds from host plants to the root hemiparasite Santalum acuminatum 

(Santalaceae)." Australian J of Botany 49(1), 9-16. Abstract. Plant hemiparasites obtain a wide 

range of primary compounds from their host plants, including carbon, water and ions. In this 

paper, we examine the transfer of carbon from the host plant Myoporum parvifolium and the 

movement of an insecticidal compound from the host Melia azedarach to the root hemiparasite 

Santalum acuminatum (R.Br) (quandong). By using 14 C we determined that glucose was moving 

from the M. parvifolium host to the parasite while the carbon fixed by quandongs was found to be 

mostly in mannitol. Mannitol occurred in fruit, leaf, stem and root tissue and also in xylem sap. We 

also provide evidence from direct infusion electrospray mass spectrometry (DIEMS) that 

quandong fruit from trees growing near Melia azedarach (L.) contain an insecticidal compound. 

This was supported by results from a bioassay in which apple moth (Epiphyas postvittana Wal 

ker) larvae suffered higher mortality when fed only on quandong fruit that was growing nearM. 

azedarach than those fed on quandong fruit from trees growing away from M. azedarach. 

Loveys B.R., Sedgley M. & Simpson R.F. (1984) “Identification and quantitative analysis of methyl 

benzoate in quandong (Santalum acuminatum) kernels. Food Technology Australia. 36, 280-289 

Moretta P., Ghisalbert E.L., Piggott M.J & Trengove R.D. (1998) “Extraction of oil from Santalum 

spicatum by supercritical fluid extraction.” ACIAR Proceedings Series 84, 83-85. Abstract. Steam 

distillation, solvent extraction, supercritical fluid extraction (SCCO2) and liquid CO2 extraction 

were used to obtain the volatile oil from Western Australian Sandalwood (Santalum spicatum (R. 

Br.) A. DC.). Supercritical fluid extraction afforded the highest yields of extractable material and 

total volatiles. The percentages of five sesquiterpene alcohols, epi--bisabolol (1), (Z)--santalol (2), 

2(E), 6(E)-farnesol (3), (Z)--santalol (4) and (Z)-nuciferol (5), were highest in the steam distillate. 

The variations in the relative amounts of these sesquiterpenes in the essential oil recovered by 

SCCO2 extraction of different sections of a single tree have been investigated.Cropwatch 

comments: According to ISO 9235, the supercritical fluid extraction of aromatic material 

produces an extract; it cannot be termed an essential oil. 

Moretta P. et al. (2001). "Longitudinal variation in the yield and composition of sandalwood oil 

from Santalum spicatum." Sandalwood Research Newsletter 14, 5-7. 

Marongiu B., Piras A., Porcedda S. & Tuveri E. (2006) "Extraction of Santalum album and 

Boswellia carterii Birdw. volatile oil by supercritical carbon dioxide: influence of some process 

parameters." Flavour and Fragrance Journal 21(4), 718 - 724 Abstract. Wood of Santalum 

album and resin of Boswellia carterii Birdw. were used to obtain their volatile oils by means of 

supercritical fluid extraction with carbon dioxide. Different extraction conditions were tested: 90 

bar, 45 °C; 120 bar, 60 °C; and 120 bar, 45 °C. On both matrices, a good process performance 

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was obtained working at 120 bar and 45 °C (density of CO2 = 0.658 g cm-3) in the extraction 

vessel, at 20 bar and 15 °C in the separator and at CO2 flow of 1.5 kg/h. At these conditions the 

higher yields were obtained: 1.9% for S. album and 6.5% for B. carterii. The main compounds 

contained in the sandalwood volatile oil were: -santalol (46.1%), -santalol (20.4%), epi--santalol 

(6.8%) and trans--bergamotol (5.4%). In the corresponding HD essential oil the -santalol and - 

santalol contents were lower: 35.0% and 14.0%, respectively. The volatile oil of B. carterii were 

made up of incensole acetate (32.0%), octanol acetate (25.1%), incensole (17.8%) and 

phyllocladene (7.7%). The percentage of the main constituents in the oil obtained by HD was 

quite different. It contained larger amounts of octanol acetate (45.2%) and phyllocladene (13.2%) 

and lower amounts of incesole (6.1%) and incensole acetate (13.0%). 

Penfold A. R. (1928) “Chemistry of West Australian Sandalwood oil.” J. Proc. Royal Soc. NSW. 

62, 60-71. 

Penfold A. R. (1932) “Chemistry of West Australian Sandalwood oil II.” J. Proc. Royal Soc. NSW. 

66, 240-247. 

Piggott M.J., Ghisalberti E.L. & Trengove R.D. (1997) “West Australian sandalwood oil: extraction 

by different techniques and variations of the major components in different sections of the same 

tree.” Fl. & Frag. J. 12, 43-46. 

Shellie R., Marriott P. & Morrison P. (2004) " Comprehensive two-dimensional gas 

chromatography with flame-ionization and time-of-flight mass spectrometry detection: qualitative 

& quantitative analysis of West Australian sandalwood oil" J Chromatog Sci. 42(8), 417-422. 

Abstract: The use of gas chromatography (GC)-mass spectrometry (MS), GC-time-of-flight MS 

(TOFMS), comprehensive two-dimensional GC (GCxGC)-flame ionization detection (FID), and 

GCxGC-TOFMS is discussed for the characterization of the eight important representative 

components, including Z-alpha-santalol, epi-alpha-bisabolol, Z-alpha-trans-bergamotol, epi-betasantalol, 

Z-beta-santalol, E,E-farnesol, Z-nuciferol, and Z-lanceol, in the oil of West Australian 

sandalwood (Santalum spicatum). Single-column GC-MS lacks the resolving power to separate 

all of the listed components as pure peaks and allow precise analytical measurement of individual 

component abundances. With enhanced peak resolution capabilities in GCxGC, these 

components are sufficiently well resolved to be quantitated using flame ionization detection, 

following initial characterization of components by using GCxGC-TOFMS. 

Srikrishna A. & Babu R.R. (2007) "Total syntheses of (±)--acorenol, β-acorenol, -epi-acorenol and 

β-epi-acorenol via an Ireland ester Claisen rearrangement and RCM reaction sequence." 

Tetrahedron Letters 48(39), 6916-6919. Abstract. Total syntheses of (±)-- and β-acorenols and 

(±)-- and β-epi-acorenols, spiro[4.5]decane sesquiterpenes, isolated from the western Australian 

sandalwood oil, have been accomplished employing a combination of Ireland ester Claisen 

rearrangement and RCM reactions for an efficient construction of the spiro[4.5]decane present in 

acoranes. 

Graphical abstract. 

Valder C., Neugebauer M., Meier M., Kohlenberg B., Hammerschmidt F.-J., Braun NA (2003) 

“Western Australian sandalwood oil – new constituents of Santalum spicatum (R.Br) A DC. 

(Santalaceae)” J. Essent. Oil Res. 15(3), 178-186. Abstract. Commercial Australian sandalwood 

oil produced from Santalum spicatum (R. Br.) A. DC. roots was analyzed using GC and GC/MS. 

Seventy constituents were identified: four monoterpenes, 64 sesquiterpenes and two others. Four 

compounds (Z)-beta-curcumen-12-ol, (Z)-12-hydroxysesquicineole, 6,10-epoxybisabol-2-en-12-ol 

andnor-helifolen12-al were found to our knowledge for the first time in nature and were 

5

characterized using^sup 1^H-,^sup 13^C-NM MR, GC/FTIR and GC/MS analyses. Cropwatch 

comments: The authors show lower concentration of cis- alpha santalol & cis-beta santalol, 

higher conc of (Z) trans-alpha bergamotol & epi-beta-santalol in the oils of S. spicatum compared 

with S. album. Regarding the bisabolols, the main isomer in S. spicatum is 6R, 7S-epi-betabisabolol 

whereas in S album it is 6R, 7S-beta-bisabolol. The oils should therefore be regarded 

as different 

Valder C., Neugebauer M., Meier M., Kohlenberg B., Hammerschmidt F.-J., Braun N.A. (2003a) 

“Santalum spicatum (R.Br.) A DC. (Santalaceae) – nor-helifolenal & acorenol Isomers: Isolation 

and biogenic considerations” J. Essent. Oil Res. 15, 381-386. 

Wedding B.B., White R.D., Grauf S., Tilse B., & Gadek P.A. (2009) “Near infrared spectroscopy 

as a rapid, non-invasive method for sandalwood oil determination.” Papers from the 11th Society 

for the Advancement of Breeding Research in Asia & Oceania Congress (SABRAO) In: 11th 

Society for the Advancement of Breeding Research in Asia & Oceania Congress (SABRAO), 10- 

14 August 2009, Cairns, QLD, Australia. Abstract. Fourier Transform (FT) - near infra-red 

spectroscopy (NIRS) was investigated as a non-invasive technique for predicting -santalol 

content in sandalwood chip samples. The correlation between the NIR spectral data and the a- 

santalol content from the GC-MS analysis was very high (R2 = 0.93). The feasibility study 

indicates that it is possible to use FT-NIRS to predict -santalol content in sandalwood chip 

samples. The technique of utilising NIRS technology for sandalwood quality and quantity 

determination needs to be further developed to be utilised as a tool for commercial applications. 

Australian Sandalwoods - General 

Anon (1919) “Western Australian sandalwood oil. An official statement on the products.” Perff 

Essen Oil Record 10(7), 194-195. 

Anon (1998) Australian Sandalwood oil. A new century – a new alternative. Mount Romance 

Australia Pty Ltd. Albany, Western Australia. 

Anon (2000) “Qld: Five fined for sandalwood harvesting” AAP General News Perth (Australia) 

Dec 12th WA: sandalwood claims would be dealt with if true: Abstract Court story alleging that 

West Australian government officials were exporting sandalwood to dealers in Taiwan who had 

offered state officials bribes or prostitutes. 

Anon (2002) AAP General News (Australia) Nov 18 (2002). Abstract: Five people fined in the 

Cairns Magistrate Court for illegal harvesting of the protected sandalwood plant, the Queensland 

EPA reportedly said. 

Anon (2002) “A Crop in Crisis” – a part of “A calming influence” Soap, Perfumery & Cosmetics 

(Oct 2002) p42-3. 

Anon (2006) "Big expansion for sandalwood plantation." ABC Newsonline Abstract 19th June 

2006. An Indian sandalwood plantation in the Ord Valley is undergoing its biggest expansion in 

seven years. Tropical Forestry Services is planting a further 235 hectares of the exotic hardwood, 

increasing its total plantation to more than 800 hectares. The company plans to harvest its first 

crop in 2012, banking on continuing strong demand from Asia, Europe and the United States. 

Chief executive Tom Cullity says the company is planning processing facilities at Kununurra to 

produce sandalwood oil which is used for perfumes and cosmetics. "Oil is from the hardwood. 

Over $100,000 Australian for a tonne of hardwood. The sandalwood oil that is distilled from the 

hardwood is very valuable and it's used in a lot of perfumes and cosmetics," he said. The other 

major grower of indian sandalwood in the Ord, ITC Limited, has now planted 750 hectares, 

owned by investors.Its first harvest is planned for 2014. 

Anon (2007) “W.A. Sandalwood set to dominate world trade.” ABC News 11/12/2007. Abstract. 

The head of one of the world's leading fragrance companies believes the Ord Valley in Western 

Australia will overtake India, as the major producer of Indian sandalwood. The Ord has the only 

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commerical crop of Indian sandalwood trees in the world. With a global shortage, oil from the 

processed timber is currently worth around $US1800 per kilogram. Georges Ferrando, from 

Albert Vieille, says with a processing plant due to be built in Kununurra next year, the region will 

become a world leader within five years. "India is number one in supplying sandalwood oil, but I 

think very, very quickly, Kununurra will become the supplier number one in the world," he says. 

Anon (2007) “Sandalwood oil – Smells like success.” RIRDC Press release 27.01.08 - see 

http://www.rirdc.gov.au/pub/media_releases/23jan07.html 

Anon (2008) “Event Notes: Sustainable Indian Sandalwood in Australia.” P&F Now June 25 th 

2008. Cropwatch comments: Sad to see P & F act as an advertising agent for TFS via their 

obedient reproduction of TFS promo material & sympathetic coverage of the recent Sandalwood 

conference at the Kimberly Grande Hotel in Kununurra Western Australia. Cropwatch has 

received opinions from conference attendees which give a more independent account, and that is 

what we should expect in Perfumer & Flavourist features. 

Anon (2008) “Givaudan enters ethical sustainability partnership for sandalwood oil.” The 

Givaudanian 05 Feb 2008 – see http://www.givaudan.com/vgn-exttemplating/v/index.jspvgnextoid=17889631fd5e7110VgnVCM1000004a53410aRCRD&cpsextcu 

rrchannel=1 Cropwatch comments: We believe that linking to Mount Romance, with its history 

of involvement in animal-products, was a major mistake by the Givaudin management. We are 

also told Givaudin are actively sourcing “more than 190 pure & natural raw materials for 

fragrances. “ 

Applegate G.B, Davis A. & Annable P.A (1990) “Managing sandalwood for conservation in N. 

Queensland, Australia” in Proc of the symposium on sandalwood in the Pacific: April 9-11, 1990, 

Honolulu, Hawai/technical co-ordinators: Lawrence Hamilton, C. Eugene Conrad. pub: 

Symposium on Sandalwood Conservation (1st: 1991: Honolulu, Hawaii). Abstract.: Santalum 

lanceolatum, the commercial species of sandalwood harvested in Queensland, was worth $4.2 

million in export earnings in 1988. The ecology of the species in natural forests is summarized, 

and information on seedling regeneration and coppice and root suckering strategies is provided. 

Stand characteristics and size class distribution in two different environments in northwest 

Queensland are provided. It is important to manage the resource for conservation. The harvesting 

guidelines, pricing criteria, and procedures are discussed along with information on heartwood 

recovery and moisture content of harvested sandalwood. Future research should be undertaken 

to monitor stand dynamics, growth rates, and the effects of land use practices on the distribution, 

growth, and dynamics of sandalwood in natural stands. 

Applegate G.B. & McKinnell F.H. (1993) “The Management & Conservation Status of Santalum 

species occurring in Australia.” In McKinnell F.H. ed. Sandalwood in the Pacific Region. 

Symposium 2nd June 1991 at XVII Pacific Science Congress, Honolulu, ACIAR Proceedings No. 

49, 5-12. 

Barrats D.R., Wijesuriya S.R. & Fox J.E.D. (1985) “Observations on foliar nutrient content of 

sandalwood (Santalum spicatum R. Br. DC.) Mulga Research Centre Journal 8, 81-91. 

Barrats D.R. (1987) “Initial observations on flowering and fruiting in Santalum spicatum (R. Br.) A. 

DC the Western Australian sandalwood.” Mulga Research Centre Journal, Australia 4, 61-65. 

Barrats D.R. (1987) “Germination & planting out techniques for the Western Australian 

sandalwood Santalum spicatum.” Mulga Research Centre Journal, Australia 9, 31-32. 

Barrett D R (1987) Initial observations on flowering and fruiting in Santalum spicatum (R. Br. ) A. 

DC. – the Western Australian sandalwood. Mulga Research Centre Journal 9:33–37. 

Bentley D. (1997) “Field grafting of Quandong. Acuminatum” Summer 1997 pp2-3. (Newsletter of 

the Australian Quandong Industry Association). 

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Bird K. (2008) "Lush secures supply of sustainable sandalwood." CosmeticsDesign-Europe 

21.02.2008. Cropwatch comments. Further move illustrating rising tendency of natural aromatic 

ingredient users to by-pass essential oil traders and sign contracts directly with producers. In this 

case the report notes the deal is to buy Indian sandalwood from the Australian TFS Corporation, 

which expects sandalwood oil to be available from its'plantations by 2011. For full story - see 

http://www.cosmeticsdesign-europe.com/news/ng.aspid=83433-lush-tfs-sandlewood 

Bird K. (2008) "Fragrance house sources sustainable ingredients." CosmeticsDesign-Europe 

07.02.2008. Cropwatch comments. Givaudin announced a partnership with Mount Romance, 

according to the article, and we are also informed that Givaudin claim to be the first fragrance 

house using an aboriginal source of wood, since we are told that the sandalwood is harvested by 

aboriginal communities in SW Australia, and inspected by the independent indigenous 

certification body, the Songman Circle of Wisdom. Full details can be seen at 

http://www.cosmeticsdesign-europe.com/news/ng.aspn=83107-givaudan-fragrance-naturalehtical. 

Mount Romance’s involvement with emu oil was quite well known (5,000 litres claimed to 

have been produced in 1997), as is Stephen Birkbeck’s (MD at Mount Romance) previous trackrecord 

in crocodile & turtle farming. Given this animal-product-exploitation scenario, the “ethical 

sustainability relationship” between Givaudin & Mount Romance woulf have probably ring hollow 

with many ecology-concious consumers & vegetarians, at least. Interestingly, the farm gate value 

of the emu-farming industry was put at $6-8 million/y (CoAS 2003), compared with a valuation of 

(only) $12m for the whole of the Australian tea tree oil industry. Opposition to emu farming in 

Australia by the Australian Royal Society for the Prevention of Cruelty to Animals, can be viewed 

at http://www.rspca.org.au/pdf/B_policystatements.pdf Further comments. Aveda also have an 

agreement with Mount Romance for supply steam-distilled Sandalwood oil (instead, apparently, 

of the solvent extract initially marketed by Mount Romance as ‘oil’) More details, as well as their 

involvement with the Ingenous Communities of Mardu Peoples of Kuktabubba for harvested 

sandalwood can be seen at http://aveda.aveda.com/protect/we/sandalwood.asp. 

Bolt C. (2001) “Tax scheme controversy fells plantation timber company” The Financial Review 

31 July 2001 p12. 

Bradfield A.E., Francis E.M., Penfold A.R. & Simonsen J.L. (1936) "Lanceol, a sesquiterpene– 

alcohol from the oil of Santalum lanceolatum. Part I." J. Chem. Soc., 1936, 1619 - 1625, 

Brand J.E. & Jones P.J. (year) “The influence of landforms on sandalwood (Santalum spicatum 

(R.Br) A.DC.) size structure & density in the North East Goldfields, Western Australia.” Rangeland 

Journal 24(2), 219-226. 

Brand J.E. (1993) “Preliminary observations on ecotypic variations in Santalum spicatum. 2. 

Genotypic variation.” Mulga Research Centre Journal 11, 13-19. 

Brand, J.E. (1994). “Genotypic variation in Santalum album.” Sandalwood Research Newsletter 2, 

2–4. 

Brand J.E. (1999) “Ecology of sandalwood (Santalum spicatum) near Paynes Find & Menzies, 

Western Australia: size structure & dry-sided stems” Rangeland Journal 21(2), 220-228. 

Abstract. Population size structure of sandalwood (Santalum spicatum) was studied on four 

pastoral leases near Paynes Find and Menzies, in semi-arid Western Australia. Stem diameter, 

height, height to crown and the orientation of dry-sided stems were recorded for 1017 individual 

sandalwood. Populations of S. spicatum at Paynes Find contained only mature trees, indicating 

no successful recruitment for at least 30 years. In contrast, populations of S. spicatum at Menzies 

had a high proportion of seedlings and saplings. Crown measurements of mature S. spicatum 

trees indicated high grazing intensity at Paynes Find: mean height to crown at Paynes Find (147- 

148 cm) was significantly higher than Menzies (92-94 cm). Dry-side percentage differed 

significantly between directional faces, consistent with sun damage. Highest mean dry-side 

percentages were on stem sides facing the sun between midday and late afternoon: west, northwest, 

south-west and north. This directional pattern was the same between pastoral leases, and 

8

there was no interaction between pastoral lease and dry-side direction. Mean percentage of 

mature trees with a dry-sided stem was also significantly higher at Paynes Find (76-82%) than at 

Menzies (42-46%). Significantly less foliage low to the ground on mature trees at Paynes Find 

may have exposed the stems to more sun damage. Land systems did not significantly influence 

dry-side direction on Burnerbinmah or Goongarrie. No S. spicatum seedlings or saplings had a 

dry-sided stem. 

Brand J.E., Ryan P.C. & Williams M.R. (1999) “Establishment and growth of sandalwood 

(Santalum spicatum) in South-Western Australia: the Northampton pilot trial.” Australian Forestry 

62(1), 33-37. 

Brand J. & Jones P. (1999). "Grow-ing sandalwood (Santalum spicatum) on farmland in Western 

Australia." Sandalwood Information Sheet No1. Conservation and Land Management (Perth WA) 

Brand J.E., Crombie D.S. & Mitchell M.D. (2000) “Establishment and growth of sandalwood 

(Santalum spicatum) in South-Western Australia: the influence of host species.” Australian 

Forestry 63(1), 60-65. 

Brand J.E., Fox J.E.D. & Moretta P. (2001). “Review of research into sandalwood (Santalum 

spicatum) tree farm systems in south-western Australia.” In Conference Proceedings: Forests in a 

Chang-ing Landscape: 16th Common-wealth Forestry Conferencejointly with the 19 th Biennial 

Con-ference of the Institute of Foresters of Australia, Fremantle, Western Australia, 18-25 April, 

2001 Promaco Conventions, Perth, pp 527-535 

Brand J.E. (2002) “Review of the Influence of Acacia species on establishment of sandalwood 

(Santalum spicatum) in Western Australia “ Conservation Science Western Australia 4(3), 125- 

129. Rangeland Journal 21(2), 220-228. 

Brand J.E., Robinson N & Archibald R.D. (2003) “Establishment & growth of sandalwood 

(Santalum spicatum) in South-Western Australia: Acacia host trials.” Australian Forestry 66(4), 

294-299. Abstract. The influence of four different Acacia species (Acacia acuminata, A. saligna, 

A. microbotrya and A. hemiteles) on the establishment and growth of sandalwood (Santalum 

spicatum) was examined at two sites in the wheatbelt, Western Australia, Australia. The host 

seedlings were planted in June 1998, and four S. spicatum seeds were planted adjacent to each 

host at age 1 year (May 1999). Direct sowing S. spicatum near 1-year-old host seedlings again 

proved to be a successful establishment technique, with 81-91% germination per spot, at both 

sites. At age 3 years, the survival of S. spicatum near A. saligna (94%) and A. acuminata (81%) 

was significantly greater than near A. hemiteles (45%). At the same age, the mean stem diameter 

of S. spicatum growing near A. saligna was 53 mm, significantly greater than near A. acuminata 

(33 mm), A. microbotrya (20 mm) and A. hemiteles (11 mm). Growth was superior at the 

Dandaragan site, with S. spicatum near A. saligna having a mean stem diameter of 59 mm and a 

mean height of 2.3 m. At the host age of 4 years, the mean height of A. microbotrya (4.3 m) was 

significantly greater than A. saligna (3.3 m), A. acuminata (3.2 m) and A. hemiteles (1.1 m). 

Between host ages of 1 and 4 years, the mean survival of A. saligna dropped by 27%, 

significantly more than the other host species (2.5-10%). Mean potassium and phosphorus 

concentrations in the foliage of S. spicatum were significantly higher near A. saligna than near A. 

hemiteles. The mean potassium:calcium ratio was highest near A. microbotrya (2.2-3.7) at both 

sites. Stem water potentials in S. spicatum were significantly lower near A. microbotrya (-2.9 

MPa) than near A. hemiteles (-2.2 MPa) at Dandaragan. There were no significant differences 

between S. spicatum stem water potentials at Narrogin. 

Brand J., Jones P & Donovan O. (2004). "Current growth rates and predicted yields of 

Sandalwood (Santalum spictum) grown in plantations in south-western Australia." Sandalwood 

Research Newsletter 19, 4-7 Abstract. Aromatic timber from S. spicatum is a valuable commodity, 

and this species hasthe potential to provide an income to farmers in the medium annual rainfall 

(400-600 mm) regions of the wheatbelt. Since 1987, S. spicatum plantations have 

beensuccessfully established in the wheatbelt, by direct seeding near 1-2 year old host seedlings, 

9

especially Acacia acuminata. This establishment technique has been very effective, with over 80 

% survival per spot, and mean stem diameters (at 150 mm above the ground) increasing at 10-12 

mm yr-1near A. acuminata. Allowing two years to establish both A. acuminata and S. spicatum, 

and then a mean stem diameter growth of only 7 mm yr-1for 18 years, the S. spicatum are 

expected to reach commercial size (127 mm) at plantation age 20 years. At this age, the 

expected yields are approximately 4.4 tonnes ha-1, with a net return ofover AU $14,000 ha-1. 

The sandalwood trees are also producing 60-170 kg ha- 1of seeds at age only 4-6 years. The 

value of the seeds may also provide a supple-mentary income to the sandalwood growers, while 

they are waiting for the trees to reach commercial size. Cropwatch comments: The authors 

state that core samples taken from 10-year old trees produced oil containing 16.7 to 21.1% α- & 

β-santalols, “which are the compounds that produce the distinct sandalwood fragrance” 

referencing Adams et al. (1975), The authors take no account of the effect on the odour profile of 

other major components found in the oil, such as the presence of 17.8% to 20.5% farnesol, a 

sesquiterpene alcohol recently identified as a sensitiser by IFRA and the subject of a recent 

SCCP Opinion. 

Braun N.A. & Meier M. (2004) “Western Australian & East Indian sandalwood oil – a comparison” 

Euro Cosmetics 12(1), 22-29. 

Bristow, M. et al. 2000. "Queensland sandalwood (Santalum lanceolatum): regeneration following 

harvesting." Sandalwood Research Newsletter 11, 4-8. 

Burfield T. & Wildwood C. (2004) “Cropwatch 2: Australian Sandalwood Oil: a tale of Spin & 

Hype” at http://www.cropwatch.org/cropwatch2.htm & www.users.globalnet.co.uk/~nodice/ 

Byrne M., McDonald B. & Brand J. (2003) “Phylogeography & divergence in the chloroplast 

genome of Western Australian sandalwood (Santalum spicatum) Heredity 91(4), 389-395. 

Abstract. Western Australian sandalwood (Santalum spicatum) is widespread throughout Western 

Australia across the semiarid and arid regions. The diversity and phylogeographic patterns within 

the chloroplast genome of S. spicatum were investigated using restriction fragment length 

polymorphism analysis of 23 populations. The chloroplast diversity was structured into two main 

clades that were geographically separated, one centred in the southern (semiarid region) and the 

other in the northern (arid) region. Fragmentation due to climatic instability was identified as the 

most likely influence on the differentiation of the lineages. The lineage in the arid region showed a 

greater level of differentiation than that in the southern region, suggesting a higher level of gene 

flow or a more recent range expansion of sandalwood in the southern region. The 

phylogeographic pattern in the chloroplast genome is congruent with that detected in the nuclear 

genome, which identified different genetic influences between the regions and also suggested a 

more recent expansion of sandalwood in the southern region. 

Byrne M., McDonald B., Broadhurst L. & Brand J. (2003) “Regional genetic differentiation in 

Western Australian sandalwood (Santalum spicatum) as revealed by nuclear RFLP analysis.” 

Theoretical & Applied Genetics 107(7), 1208-1214. Abstract. Western Australian sandalwood, 

Santalum spicatum, is widespread in the semi-arid and arid regions of Western Australia, and 

there is some morphological variation suggestive of two ecotypes. The level and structuring of 

genetic diversity within the species was investigated using anonymous nuclear RFLP loci. 

Santalum spicatum showed moderate levels of genetic diversity compared to other Australian 

tree species. The northern populations in the arid region showed greater levels of diversity and 

less population differentiation than the southern populations in the semi-arid region due to 

differences in the distribution of rare alleles. Equilibrium between drift and gene flow in the 

northern populations indicated that they have been established for a long period of time with 

stable conditions conducive to gene flow. In contrast, the southern populations showed a 

relationship between drift and gene flow indicative of a pattern of fragmentation and isolation 

where drift has greater effect than gene flow. The different patterns of diversity suggest that the 

ecotypes in the two regions have been subject to differences in the relative influences of drift and 

gene flow during their evolutionary history. 

10

CALM (2001) (Dept of Conservation & Land Management 2001) “New sandalwood contracts for 

station owners” Media Release 17th July Perth: Dept of Conservation & Land Management. 

Choupechoux R. (1931) Contribution a l'etude de l'essence de santal Australie. pub. Lyon, Boso 

& Riou 1931. 

Clarke M. (2006) “Australia’s Sandalwood Industry: an overview & analysis of research needs”. 

Publicn no 06/131- For RIRDC – see http://www.rirdc.gov.au/reports/EOI/06-131.pdf 

Crossland T. (1982). “Germination of sandalwood seed.” Mulga Research Centre Report, Curtin 

University, Perth. 5:13-16. 

Crossland T. (1982) “Response to fertiliser treatment by seedlings of sandalwood, Santalum 

spicatum (R.Br) DC.” Annual Report, Mulga Research Centre Australia 5, 13-16. 

Crossland T. (date) ”Preliminary investigations into germination and establishment of 

sandalwood, Santalum spicatum (R. Br.) DC. Annual Report, Mulga Research Centre Australia 4, 

61-65. 

Done C , Kimber P. & Underwood R. (2008) ““Development of the Indian Sandalwood industry on 

the Ord river irrigation area” Sandalwood Conference 2008 at The Kimberley Grande, 

Kununurra, W. Australia 13-15 May 2008. 

Donovan R.J. (undated) A history of the sandalwood industry of Western Australia Battye Library, 

Perth, Australia 

Duus J. E. (1987). “Harvesting of Sandal-wood from Crown Lands in Queen-sland.” 

(Unpublished). 

Fergeus J. (undated pamphlet) “Australian sandalwood aromatic review” Australian Botanical 

Products Pty Ltd. 

Flanagan F. & Barrett D.R. (1993) “Sandalwood nuts as food.” Mulga Research Centre Journal 

11, 21-26. 

Forest Products Commission WA Media Release (27 March 2006) “Preliminary oil results from a 

14-year-old Indian sandalwood plantation at Kununurra, WA.” 

Forest Products Commission WA Media Release (16 th 

plantations established in Carnavon.” 

May 2006) “First Indian sandalwood 

Fox J.E.D. & Brand J.E. (1993) “Preliminary observations on ecotypic variations in Santalum 

spicatum. 1. Phenotypic variation.” Mulga Research Centre Journal 11, 1-12. 

Fox J.E.D. & Wijesunya S.R. (1985) “Sandalwood planting with property owners” Mulga Research 

Centre Journal 8, 3340. 

Fox J E D & Brand J E (1993). “Preliminary observations on ecotypic variation in Santalum 

spicatum. 1. Phenotypic variation.” Mulga Research Centre Journal 11:1–12. 

Fox J.E.D. (1997) "Why is Santalum spicatum common near granite rocks" J. Royal Soc. of 

Western Australia 80, 209-220. Abstract. Sandford Rocks Nature Reserve is dominated by a 

large granite outcrop. This reserve is notably well-endowed with trees of the root parasite 

sandalwood (Santalum spicatum). These are comparatively common in and among granite 

exposures. Trees attain 4 m in height and 20 cm basal diameter on favourable sites but are small 

gnarled shrubs in rock fissures. Fruiting ability differs considerably between trees. Despite 

apparently high densities of rabbits, continuous regeneration appears to have occurred, but only 

in the vicinity of parent trees. The reserve contains a number of distinct vegetation associations 

that are soil determined. Although sandalwood is common near exposed granite it is rarely found 

11

in association with Eucalyptus stands. It is suggested that the water-shedding properties of the 

granite exposures are less important to sustaining sandalwood than the presence of preferentially 

parasitised host species. 

George, A. S. (1984). “Santalum.” in Flora of Australia, vol. 22. Bureau of Flora and Fauna. 

Australian Government Publishing Service. Canberra, Australia. 

George, A.S 1996) “Sandalwoods and quandongs of Australia.” Australian Plants 13, 318-319 

Gearon V. (2000) at 

http://www.essentiallyoils.com/Newsletters/October_2000_Newsletter/october_2000_newsletter.h 

tml 

Gearon V. (2002) Aromatherapy Today 24, Dec 2002 p21 

Gowda D. (2008) “Detergents“Decline in the Supply of Natural Sandalwood oil: deforestation, 

adulteration and synthetics.” Sandalwood Conference 2008 at The Kimberley Grande, 

Kununurra, W. Australia 13-15 May 2008. Cropwatch Comments: Gowda maintains that in spite 

of the official figures, the current (2008) annual production of sandalwood is 3,000 - 4,000 tons 

and for sandalwood oil is 120-150 tons, of which 80 tons/annum of sandalwood oil is consumed 

by the domestic market. Gowda is employed by Karnataka Soaps & Detergents Ltd. (KSDL) once 

the largest producers of sandalwood oil E.I. Gowda informs us that sandalwood oil distillation 

commenced in 1916 in Mysore, and 2 years later the essential oil was incorporated into 

sandalwood soap by KSDL. Gowda lists polyethylene glycols, African sandalwood oil, castor oil 

and coconut oil amongst the adulterants of E.I. Sandalwod oil. 

Grant, W.J.R. & Buttrose, M.S. (1978) "Domestication of the quandong, Santalum 

acuminatum.”" Australian Plants 9, 316-318 

Harbaugh D. (2007) "A taxonomic revision of Australian northern sandalwood (Santalum 

lanceolatum, Santalaceae)." Australian Systematic Botany 20(5) 409–416. Abstract. A previously 

published molecular phylogenetic analysis of the sandalwood genus, Santalum L. (Santalaceae), 

identified that the Australian endemic northern sandalwood, S. lanceolatum R.Br., is not 

monophyletic and contains a distinct, yet cryptic, lineage within it as currently circumscribed. This 

study examines nuclear ribosomal gene sequences of additional specimens from across its 

geographic range, and 30 morphological characters, in order to revise the taxonomy of S. 

lanceolatum sensu lat. (s.l.) and the segregate species that should bear the name S. leptocladum 

Gand. Santalum lanceolatum sensu stricto (s.s.) is distributed in the humid to subhumid regions 

of northern Australia north of 20°S latitude, whereas S. leptocladum occurs in the arid and 

temperate regions of central and southern Australia. Putative interspecific hybrids were 

discovered in two localities, and may represent either natural or human-mediated hybridisation. 

The results of this study have major economic and conservation implications because S. 

lanceolatum s.s., which is known to have higher levels of fragrance compounds than S. 

leptocladum, has a much more restricted range than previously thought. 

Harbaugh D.T. (2008) "Polyploid and Hybrid Origins of Pacific Island Sandalwoods (Santalum, 

Santalaceae) Inferred from Low-Copy Nuclear and Flow Cytometry Data." Int. J Plant Sci. 

169(5), 677–685. Abstract. It has been argued that polyploids are better adapted than diploids for 

long-distance dispersal to and establishment on oceanic islands. To address this issue in a 

molecular phylogenetic framework, the extensive history of auto- and allopolyploidization in 

Santalum (Santalaceae), the sandalwood genus, was studied by sequencing the low-copy 

nuclear gene waxy and investigating the ploidy level of all 16 species. Ploidy level was estimated 

by measuring the C value (total amount of DNA per nucleus) using flow cytometry and calibrating 

it by known chromosome numbers and new root-tip chromosome counts of several taxa. Results 

indicate four ploidy levels in Santalum: diploid (n=10), tetraploid (n=20), hexaploid (n=30), and 

octoploid (n=40). The waxy phylogeny suggests that at least six independent polyploid events 

occurred in the history of Santalum: two allopolyploid events between distantly related species 

12

and four putatively autopolyploid events. An additional hybrid event between two tetraploid 

Hawaiian clades evidently produced the tetraploid species S. boninense, endemic to the Bonin 

Islands. By finding more than twice as many long-distance island colonizations from polyploid as 

from diploid ancestors, this study provides novel evidence for the role of polyploidy in plant 

colonization throughout the Pacific Islands. 

Henschke I. (2000) “Sandalwood brings sweet smell of success.” Landline. Australian 

Broadcasting Corp at http://www.abc.net.au/landline/stories/s206172.html 

Herbert D A (1925) “The root parasitism of Western Australian Santalaceae.” Journal and 

Proceedings of the Royal Society of Western Australia 11, 127–149. 

Heuberger E., Gearon V., Birbeck S., Buchbauer G. (2002) “The essential oil of Australian 

sandalwood (Santalum spicatum) – effects of different samples on human physiology & 

subjective evaluation,” 33rd ISEO, Sept 2002, Lisbon, Portugal. 

Hobman, F.R. (1991) “The SA Dept of Agriculture evaluation programme for quandongs..” In 

Quandongs, a viable opportunity. Minnipa Research Centre, Oct. 18, 1991. Dept of Agriculture, 

South Australia 

Hood J.R., Cavanagh H.M. & Wilkinson J.M. (2004) "Effect of essential oil concentration on the 

pH of nutrient and Iso-sensitest broth." Phytother Res. 18(11), 947-9. Abstract. The role of pH on 

the antimicrobial activity of essential oils has not been well studied. The effect of four essential 

oils: Backhousia citriodora, Melaleuca alternifolia, Lavandula angustifolia and Santalum spicatum 

(0.1% to 10%) on the pH of two commonly used media, nutrient broth and Iso-sensitest broth, 

was therefore undertaken. Small (less than 0.5 pH units) but statistically significant differences 

between the pH of the two media followed the addition of M. alternifolia, L. angustifolia and S. 

spicatum essential oil. In general the effect on pH was greatest at higher concentrations and the 

fall in pH was greatest in the nutrient broth. The addition of B. citriodora essential oil to nutrient 

broth resulted in a fall in pH from 7.29 +/- 0.02 (no oil) to 5.2 +/- 0.03 (10% oil). This effect was 

not observed in the Iso-sensitest broth. 

Hudson (2008) “Kununurra could become world's biggest producer of Indian Sandalwood”. 

http://www.abc.net.au/rural/wa/content/2006/s2244847.htm Cropwatch comments: Hopefully 

the 75 or so delegates (presumably manily shareholders) were impressed by reports of 3,000 + 

ha of sandalwood under cultivation. No party-pooper mentioned the fact that due to the high cost 

of sandalwood oil, the hard-pressed perfumery trade has mainly switched to cheap sandalwood 

synthetics on cost grounds. 

Ilah A. et al. (2002). “Somatic embryo irregularities in in vitro cloning of sandal (Santalum album 

L.).” Sandalwood Research Newsletter 15, 2-3. 

Jain S.H., Angandi V.G. & Shankaranarayana, K.H. (2003) "Edaphic, environmental and genetic 

factors associatedwith growth and adaptability of Sandal (Santalum album L.) in provenances." 

Sandalwood Research Newsletter 17, 4-5. Abstract. Sandal tree grows under different edaphic 

and eco climatic conditions. Consid-ering large genetic distance between provenances, it is 

concluded that under di-verse locality factors sandal adapts very well in terms of growth, 

heartwood and oil content. 

Jones G.P., Tucker D.J., Rivett D.E. & Sedgley M." (1985). “The nutritional potential of the 

quandong (Santalum acuminatum) kernel.” Journal Plant Foods 6, 239-246. 

Jones G.P., Birkett A., Sanigorski A., Sinclair A.J., Hooper P.T., Watson T.& Rieger V. (1994) 

"The effect of feeding quandong (Santalum acuminatum) oil to rats on tissue lipids, hepatic 

ctochrome P450 and tissue histology." Food and Chemical Toxicology 32, 521-525 

Jones P. (1999) “'Growing Sandalwood (Santalum spicatum) on farmland in Western Australia.' 

Forest Products Commission Information Sheet Issue 1, May 1999. 

13

Jones P. (2002) “Estimating Returns on Plantation Grown Sandalwood (Santalum spicatum)” 

Forest Products Commission Sandalwood Information Sheet Issue 3, July 2002 

Jyothi P.V., Atluri J.B. & Subba R.C.(1991). "Pollination ecology of Santalum album 

(Santalaceae)." Tropical Ecology 32, 98-104. Abstract. Santalum album L. is obligately 

xenogamous and blooms thrice a year at Visakhapatnam (17 degree 42'N-82 degree 18'E). The 

flowers anthese between 0430 and 1930 hr and soon after the anthers dehisce. The pollen grains 

are viable for 16 hr and the stigma is receptive for 48 hr. Nectar production begins 24 hr after 

anthesis and continues through flower life. Ants, bees, flies, butterflies and wasps forage at the 

flowers; only the last two groups serve as pollinators, wasps being the "major". 

Kauber K. “Australian sandalwood oil – acute oral toxicity and acute dermal toxicity”, Scantox, 

Denmark 2000 (unpublished). Cropwatch comments: Cropwatch previously asked Scantox to 

release details of this study, allegedly funded by Mount Romance, which was said to include 

animal testing experiments. Scantox politely declined to release this confidential data. Later all 

references to this funded research were removed from Mount Romance’s internet presence, 

perhaps because many perfumery companies will not accept perfumery materials manufactured 

by companies who have tested their products on animals. However some aromatherapy oil 

traders (apparently customers of Mount Romance) failed to erase the data as promptly, which is 

where we initially learned of the Scantox studies. 

Kealley I.G. (1991) “The management of Sandalwood” Dept of Conservation & Land 

Management, W. Australian Wildlife Management Program No 8, 3-9.. 

Keenan R (1996) “Santalum lanceolatum in Queensland.” Sandalwood Research Newsletter - 

Issue 1.Department of Conservation and LandManagement, Kununurra, Western Australia. 

Kerr J. (2000) “Essential Oil Profile – Australian Sandalwood Oil” Aromatherapy Today 15, 8-12. 

Kerr J. (2002) “Editorial Comment” Aromatherapy Today 24 Dec 2002 p32-33. 

Lethbridge B. (1998) “Germinating bitter quandong.” Acuminatum Autumn 1998, p 4 

Lethbridge B. (1998) "Root rot, rootstock and phosphorous acid." Acuminatum, Winter 1998, p 4" 

Lethbridge, B. (1999) “Host plants I - Melaleucas. "Acuminatum, Autumn 1999, p 4. 

Lethbridge, B. (2001). “Grafting compatibility of quandong, Santalum acuminatum.”. Sandalwood 

Research Newsletter 12, 2. 

Lethbridge B. & Randell B. (2003) "Genetic and agronomic improve-ment of Quandong." RIRDC 

Pub-lication No. 03/110. 

Lethbridge B. (2004) "Do Our Own Research (DOOR) quandong production." RIRDC publication 

No. W04 / 111 

Lethbridge B.(2004) "Native Foods : Quandong." In The New Crop Industries Handbook. Edited 

by Salvin S., Bourke M. Byrne A. Rural Industries Research and Development Corporation. 

Lethbridgw B. (2005) "Field grafting of Quandong (Santalum acuminatum)." Sandalwood 

Research Newsletter 20. April 2005. 

Luong T.M. (2002) "Competitive effects within and between Santalum album and pot host 

Alternanthera dentata." Sandalwood Research Newsletter 16. Abstract. The growth of Santalum 

album seedlings and the preferred pot host Alternanthera dentata under nursery conditions is the 

first important step in establishing this species in planta-tions. A 19 week pot trial was conducted 

in a glasshouse at Curtin University of Technol-ogy, Perth, Western Australia. The aim was to test 

whether an increase in host density im-proved growth of sandalwood seedlings. S. album 

14

seedlings had a tendency to grow betterat lower densities of A. dentata (one or two hosts per 

pot), compared with higher densities (three or four hosts per pot). Seedlings with two hosts had 

greater heights, dry root andshoot weights and leaf area, while seedlings with one pot host had 

more leaves. There wereno clear trends between number of haustorial connections made as host 

density increased.As host density increased, the leaf area, root and shoot weights of A. dentata 

declined. Bothparasite and host were more affected by competition, however the host was more 

affected byintraspecific competition, indicated by large competitive responses to each other. S. 

album seedlings had less effect or response to density of A. dentata after 19 weeks, perhaps due 

tonot being limited by the same resources as the host at this early establishment phase 

Loveys B.R. & Jusaitis M. (1994) “Stimulation of germination of quandong (Santalum 

acuminatum) and other native plant seeds. Australian Journal of Botany. 42, 563-574. 

Liu Y.D. & Longmore R.B. (1997) “Dietary sandalwood seed oil modifies fatty acid composition of 

mouse adipose tissue, brain & liver.” Lipids 32(9), 965-969. Abstract. Sandalwood (Santalum 

spicatum) seed oil, which occurs to about 50% of the weight of the seed kernels, contains 30- 

35% of total fatty acids (FA) as ximenynic acid (XMYA). This study was designed to obtain basic 

information on changes in tissue FA composition and on the metabolic fate of XMYA in mice fed a 

sandalwood seed oil (SWSO)-enriched diet. Female mice were randomly divided into three 

groups, each receiving different semisynthetic diets containing 5.2% (w/w) fat (standard 

laboratory diet), 15% canola oil, or 15% SWSO for 8 wk. The effects of SWSO as a dietary fat on 

the FA composition of adipose tissue, brain, and liver lipids were determined by analyses of FA 

methyl ester derivatives of extracted total lipid. The FA compositions of the liver and adipose 

tissue were markedly altered by the dietary fats, and mice fed on a SWSO-enriched diet were 

found to contain XMYA but only in low concentration (0.3-3%) in these tissues; XMYA was not 

detected in brain. Oleic acid was suggested to be a principal XMYA biotransformation product. 

The results were interpreted to suggest that the metabolism of XMYA may involve both 

biohydrogenation and oxidation reactions. 

Liu Y.D., Longmore R.B. & Kailis S.G. (1997) “Proximate & fatty acid composition changes in 

developing sandalwood (Santalum spicatum) seeds.” J. Science of Food & Agriculture 75(1), 27- 

30. Abstract. Changes in the proximate composition of developing seeds of sandalwood 

(Santalum spicatum R Br) were quantified. The developing fruits were collected regularly over a 

period of 5 months commencing 14 days after flower opening. Rapid deposition of seed lipid 

began at about 91 days after flowering (DAF) at a level of 4 g kg-1 and continued to about 396 g 

kg-1 at 147 DAF. Protein and ash contents displayed similar trends to that of lipid with a 

corresponding decrease in moisture content. Fatty acid analysis of the seed oil demonstrated 

marked changes in composition during seed development. In particular, major increases in oleic 

and ximenynic acids were noted with corresponding decreases in the other fatty acids. 

Lonergan O.W. (1990) “Historical review of sandalwood (Santalum spicatum). Research in 

Australia”. Perth: Research Bulletin No 4 Dept of Conservation & Land Management Dec 1990 

p28. 

McKinnell F.H. (1990) “Status of management & silvicultural research on sandalwood in W. 

Australia & Indonesia” In Hamilton L. & Conrad C.E. ed. Proceedings of the Symposium on 

Sandalwood in the Pacific; April 9-11 1990 Tech. Rep PSW-122, Pacific Research Station, Forest 

Service, UJS Dept of Agric, Honolulu, 19-25. Abstract. The current status of the conservation and 

management of Santalum spicatum in Western Australia and S. album in East Indonesia is 

outlined. Natural and artificial regeneration techniques for both species in selected areas are 

discussed. The present Australian Centre for International Agricultural Research program on S. 

album in Nasa Tenggara Timur is described in relation to the management needs of the species 

in that province. In S. spicatum, research on silviculture is essentially complete, and interest is 

now focused on the marketability of the kernels for human consumption. 

15

Maslin B.R., Byrne M., Coates D., Broadhurst L. et al. (1999) "The Acacia acuminata (Jam) 

group: an analysis of variation to aid Sandalwood (Santalum spicatum) " Report to the 

Sandalwood Business Unit, Department of Environment & Conservation, Australia (1999). 

Misra U. (2009) "How India's Sandalwood Oil trade got hijacked." Business 6th Aug 2009. 

Cropwatch comments: The article comments on how India went from sandalwood oil exporter to 

importer, with a look at Australia's 20,000 ha sandalwood plantation profect. Tim Coakley, 

executive chairman of the Wescorp Group of Companies is quotes as estimating that 15 to 17 

tons/annum of oil of Santalum spicatum is currently produced in Australia and that exporting of oil 

of Santalum album would start in 3-7 years. 

Mullholland J. (1994) “An investigation of the harvesting processing and export of Western 

Australia sandalwood (Santalum spicatum).” - Abstract of Masters Thesis University of Australia; 

summary available at the website of Institute of Foresters of Australia 

http://www.ifa.unimelb.edu.au/abstracts/master/1994/mulholland1994.htm 

Murphy M.T., Garkakalis M.J. & Hardy G.E.S.J. (2005) “Seed catching by woylies Bettongia 

penicillata can increase sandalwood Santalum spicatum regeneration in Western Australia” 

Austral. Ecology 30(7), 747-755. 

Murphy M. & Mark Garkaklis M. (2003) "Hopping Into A Bright Future- The Woylie Sandalwood 

Story." Sandalwood Research Newsletter 18. 

Murthy S.G. (1985) “Sandalwood: case study of a resource in decline.” Garden 16-19. 

Oates A. (1989) The Story of Sandalwood The Museum of the Golfields Kalgoorlie 

Owen L.N. (1949) "Lanceol, a sesquiterpene alcohol from the oil of Santalum lanceolatum. Part II. 

Some observations on the degradation product." J. Chem. Soc., 1949, 1582 - 1586, 

Possingham J. (1986) "Selection for abetter Quandong." Australian Horticulture February 1986 

pp55-59 

Radomiljac A. (2000) see: http://users.bigpond.net.au/sellwood/kimsoc/pasttalk00.htm 

Radomiljac A.M. (1998). “The influence of pot host species, seedling age and supplementary 

nursery nutrition on Santalum album (Linn.) plantation establishment within the Ord River 

Irrigation Area, Western Australia.” Forest Ecology and Management 102(2–3), 193–201. 

Abstract. A factorial experiment investigated the effect of six pot host species treatments 

(Alternanthera nana, Sesbania formosa, Atalaya hemiglauca, Acacia hemignosta, Crotalaria 

retusa and no pot host), two Santalum album seedling age treatments (24 and 17 weeks at field 

establishment) and a supplementary nursery nutrition treatment (2×100 ml 5% Ca Wuxal®) on 

Sa. album survival and growth 287 days after field establishment. Significant variation exists 

between pot host species in increasing Sa. album survival and growth. Al. nana and Se. formosa 

pot host species significantly increased Sa. album survival, height and diameter. Sa. album 

survival, height and diameter was significantly better with supplementary nursery nutrition. Sa. 

album survival and height was significantly greater and pot host species survival was significantly 

poorer with older Sa. album seedlings. Older seedlings and supplementary nursery nutrition gave 

higher levels of Sa. album field survival and growth when parasitised to poor pot host species but 

not when parasitised to satisfactory pot host species. 

Raychaudhuri S.P. & Varma A. (1980) “Sandal spike” Review of Plant Pathology 59(3), 99-107. 

Razikari H. (1996) “An assessment of the commercial potential of quandong (Santalum 

acuminatum) varieties in Broken Hill. Thesis, University of Western Sydney, Hawkesbury. 

RIRDC (undated) see: http://www.rirdc.gov.au/champions/MtRomanceAustralia.html. 

16

Robson K. (2003). “Sandalwood species x host interaction trial in North Queensland.” Pacific 

Islands Forest and Trees, 3/2003, pp. 14-16. 

Robson K. (2004) “Experiences with Sandalwood in plantations in the South Pacific & N. 

Queensland.” In: Prospects for high value hardwood in the “dry” tropics of N. Australia. 

Proceedings of a workshop held in Mareeba, N. Queensland, Australia 19-21st Oct 2004. pub. 

Private Forestry North Queensland Association Inc. N. Queensland. Abstract. Sandalwood is an 

important commercial industry in the south western Pacific. A number of sandalwood species 

occur across the south western Pacific, Santalum austrocaledonicum in New Caledonia and 

Vanuatu, and Santalum yasi in the Fiji Islands and Tonga. Communities do the majority of 

sandalwood plantings, manage and harvest existing stands. There is a growing interest among 

villagers, other smallscale growers and Governments to expand the scale of planting in both 

countries. The most common type of planting is garden plantings of sandalwood by villagers. 

However, large investors and Governments now starting to invest in plantations across the south 

western Pacific. 

Rugkhla A. & Jones M.G.K. (1998) “Somatic embryogenesis & platelet formation in Santalum 

album & S. spicatum.” J of Exptl. Botany 49(320), 563-571. Abstract. A reproducible system for 

somatic embryogenesis and plantlet formation of sandalwood has been developed. A high 

frequency (100%) of somatic embryos were induced directly from various explants in MS 

(Murashige and Skoog, 1962) medium with thidiazuron (1 or 2 M) or indirectly in medium 

containing 2,4-D plus thidiazuron. Within 8 weeks, white globular somatic embryos or friable 

embryogenic tissue developed on cultured explants. In S. album the globular somatic embryos 

were transferred to MS medium supplemented with IAA (6 M) and kinetin (1 and M) where they 

developed further, multiplied and maintained friable embryogenic tissue. After 15-30 d, mature 

somatic embryos (1-2 mm) with well-developed cotyledons were separated and subcultured on to 

medium containing GA3 (6 M) for germination. Once germinated, elongated somatic embryos 

(10-20 mm long) grew further in MS supplemented with lower GA3 (3 M). In S. spicatum, the 

addition of casein hydrolysate and coconut milk was necessary for plantlet development from 

somatic embryos. From histological studies, it appeared that primary somatic embryos arose from 

single cells or had a multicellular origin from the epidermis or cortical parenchyma. Secondary 

somatic embryos and friable embryogenic tissue differentiated from groups of proembryogenic 

cells from a superficial layer of the primary somatic 

Rugkhla A., McComb J.A. & Jones M.G.K. (1997) “Intra- & inter-specific pollination of Santalum 

spicatum & S. album.” Australian J of Botany 45(6), 1083-1095. Abstract. The flower morphology, 

receptivity and sexual compatibility between genotypes and species were determined in Western 

Australian sandalwood (Santalum spicatum) and Indian sandalwood (S. album). The results 

showed that the stigma of both species became receptive at anthesis and reached a peak at 3 or 

4 days after anthesis. Pollen tubes took 2 days to grow to the ovary when pollinated at anthesis, 

and 1 day when pollinated 2 or 3 days after anthesis. The egg apparatus matured at least 2 days 

after pollination and varied between genotypes. Fertilisation occurred 2 or 3 days following cross 

pollination. Although 10–40% of ovules were fertilised following intra-specific crosses of both 

species, the average initial fruit set was much lower: 4% in S. spicatum and 19% in S. album. 

Most immature fruit (75–80%) abscised following intra-specific pollination. The number of pollen 

tubes that grew in styles after self-and inter-specific pollination was lower than that for intraspecific 

pollination. Following self and inter-specific pollination, growth of pollen tubes was 

arrested in the style, ovary and around the embryo sac; a few penetrated the embryo sac. Initial 

fruit set was low and developing fruit abscised prematurely. The results indicated that pre- and 

post-fertilisation mechanisms control self-incompatibility and inter-specific incompatibility between 

the sandalwood species. 

Ryan, P.C. & Brand, J.E. 2002. Techniques to improve sandalwood (Santalum spicatum) 

regeneration at Shark Bay, Western Australia: stem coppice and direct seeding. Sandalwood 

Research Newsletter 15, 4-7. 

17

Samson, Basil (1980): The camp at Wallaby Cross. Canberra: Australian Institute of Aboriginal 

Studies; 199-202. 

Sawyer (1892) through Applegate Graham B, Davis Allan G.W. & Annable. Peter A. (1990) 

“Managing Sandalwood for Conservation in N. Queensland, Australia” in Proc of the Symposium 

on Sandalwood in the Pacific: April 9-11, 1990, Honolulu, Hawai/technical co-ordinators: 

Lawrence Hamilton, C. Eugene Conrad. pub: Symposium on Sandalwood Conservation (1st: 

1991: Honolulu, Hawaii). 

Sedgley M. (1982). “Preliminary assessment of an orchard of quandong seedling trees.” J. Aust. 

Inst. Agr. Sci. 48(l):52-56. 

Sedgley M (1982) "Floral anatomy andpollen-tube growth in the Quan-dong (Santalum 

acuminatum (R Br) a Dc)." Australian Journal of Botany 30, 601-609..Abstract. Floral anatomy 

and pollen tube growth in the quandong were studied using light and scanning electron 

microscopy. The flowers had four perianth lobes and four stamens whose anthers dehisced by 

longitudinal slits. The pollen became caught in long unicellular hairs adjacent to the anthers. The 

central disc secreted nectar through raised stomata. The stigma papilla cells had a cuticle with a 

rough surface overlying thick PAS-positive walls. The half-inferior ovary normally contained two 

ovules. The embryo sac extended beyond the ovule at the micropylar end and into the placenta at 

the chalazal end. Half of the ovaries observed at both anthesis and 4 days following anthesis had 

no embryo sacs and the other half had one embryo sac. Occasional ovaries had two embryo sacs 

and some underdeveloped embryo sacs were observed that did not extend beyond the ovule or 

into the placenta. Pollen tubes had reached the ovary by 1 day following pollination and the 

stigma was receptive for 8 days following anthesis. Only half of the pistils had pollen tubes in the 

ovary. Unpollinated flowers had no pollen tube growth in the pistil. 

Sen-Sarma P.K. (1982) “Insect vectors of sandal spike disease” European J of Forest Pathology 

12(4/5), 297-299. 

Shea S.R., Radmomiljac A.M., Brand & Jones P. (1998) “An overview of sandalwood and the 

development of sandal in Farm Forestry in W. Australia”. ACIAR Proceedings 84, 9-18. 

Sidheswaran P. & Ganguli S. (1997) “Sandalwood oil substitutes – a review” Supplement to 

Cultivation & Utilisation of Aromatic Plants 123-139. 

Statham P. (1988). “The Australian sandalwood trade, small but significant.” Working Paper No. 

100. Canberra, Australia. Department of Economic History, The Australian National University. 36 

Statham P. (1990) “The sandalwood Industry in Australia: A history” in Proc of the Symposium on 

Sandalwood in the Pacific: April 9-11, 1990, Honolulu, Hawai/technical co-ordinators: Lawrence 

Hamilton, C. Eugene Conrad. Pub: Symposium on Sandalwood Conservation (1st: 1991: 

Honolulu, Hawaii). p26. Abstract. From its inception in 1805, when it contributed to Sydney 

merchant incomes from whaling ventures, until today, when it earns several million dollars in 

export revenue, the sandalwood industry has played a small but significant part in Australia's 

economic development. The history of the industry falls into three major stages: first is the offshore 

exploitation of the wood from Sydney, from 1805 to the 1840's and beyond; second is the 

free exploitation of Australian grown sandalwood from 1844 to 1929; and finally the period of 

government controlled exploitation from 1929 to the present. 

Struthers R., Lamont B.B., Fox J.E.D., Wijesuriya S. & Crossland T. “Mineral nutrition of 

sandalwood (Santalum spicatum).” J. of Exptl. Botany 37(182), 1274-1284. 

Surate I.K. (1994) “The effect of hostplants on the growth of sandalwood seedlings (Santalum 

album Linn).” In: Sandalwood Research Newsletter Issue 3. Department of Conservationand 

Land Management, Kununurra, Western Australia. 

18

Talbot L. (1983) "Wooden gold. Early days of the sandalwood industry." Forest Focus 30, 21-31. 

pub W. Austr. Forest Dept, Perth. 

Taylor D., Swift S. & Collins S. (2000) “Testing growth & survival of four sandalwood species in 

Queensland” Sandalwood Research Letter 10, 6-8. Abstract. Sandalwood production in 

Queensland has been based on harvesting from naturally occurring Santalum lanceolatum, 

principally from relatively remote areas in northern Queensland between Hughenden and 

Normanton. Santalum lanceolatum has a relatively low oil yield in comparison to other 

sandalwood species and a consequent lower market value. With declining amounts of natural 

sandalwood available for harvest and an increasing market the potential exists for sandalwood 

production from plantations. To date, little work has been done in Queensland on production of 

Sandalwood in plantations. This report details anexperiment established in late 1999 to 

investigate the growth and survival of four sandalwood species, viz; Santalum album, S. 

austrocaledonicum, S. yasi and S. macgregorii on two sites in Queensland 

Tennakoon K.U. & Cameron D.C. (2006) "The anatomy of Santalum album (Sandalwood) 

haustoria. Can. J. Bot. 84(10), 1608-1616. Abstract. Structural attributes of Santalum album L. 

(Sandalwood) haustoria have been long overlooked in the literature. This is surprising since 

successful haustorial formation is key to the survival of individuals of this ecologically and 

economically important plant. We investigated the morphology of haustoria formed by S. album 

attached to one of its principal hosts Tithonia diversifolia (Hemsley) A. Gray. The bell-shaped 

mature haustoria were composed of a peripheral hyaline body and a centrally located penetration 

peg. The parasite penetration peg can penetrate the host by means of direct pressure and the 

secretion of cell-wall-degrading enzymes when forming a successful graft union. The latter 

mechanism is supported by this study as we observed no evidence of collapsed host cells as the 

result of parasite applied pressure. Upon reaching the xylem tissue of the host root, the 

penetration peg formed a thin ellipsoidal disc and the host–parasite interface was almost entirely 

composed of parenchymatous tissue. Luminal continuities were absent between the xylem 

conducting tissues of the partners, thus suggesting mass flow of solutes is unlikely to occur in this 

association. High densities of contact parenchyma were found at the host–parasite interface; thus 

it is probable that these are the principal structures formed by the parasite that facilitate the 

acquisition of host-derived xylem resources. This study therefore concludes that haustorial 

anatomy of S. album supports cross membrane (potentially selective) uptake of host-derived 

solutes as opposed to mass flow via vascular continuity. 

Tennakoon K.U. & Pate J.S. (1997) “Biological and physiological aspects of the Santalum 

acuminatum (quandong) and its hosts in Western Australia.” Sandalwood Research Newsletter 6: 

1-2 

Tennakoon K.U., Pate J.S. & Arthur D. (1997) “Ecophysiological aspects of the woody root 

hemiparasite Santalum acuminatum and its common hosts in South Western Australia.” Annals of 

Botany. 80: 254-256 

Tennakoon K.U., Pate J.S. & Stewart, G.R. (1997) “Haustorium-related uptake and metabolism of 

host xylem solutes by the root hemiparasitic shrub Santalum acuminatum. Annals of Botany. 80: 

257-264 

Tonts M. (2001) Sandalwood Market Study (Draft Report) Perth: Dept of Agriculture. 

Tonts M. & Selwood J. (2002) “Niche Markets, Regional Diversification and the Reinvention of 

Western Australia’s Sandalwood Industry” Tijdschrift voor Economische en Sociale Geografie 

94(5), 564-575. Abstract. Diversification and niche marketing have become very important 

economic strategies for many rural small businesses, farmers and communities. As part of these 

strategies, new opportunities often emerge for traditional products and industries. In the case of 

Western Australia, this has contributed to the revitalisation of the sandalwood industry. While 

sandalwood has been exported from Western Australia for more than 150 years, for much of the 

second half of the twentieth century it was of little economic significance. In recent years, 

19

however, the industry has become increasingly entrepreneurial, successfully marketing its 

products into niche markets in the global economy. For farmers and communities in rural areas, 

the revitalisation of the sandalwood industry has also provided opportunities for economic 

diversification and a profitable way of tackling land degradation. 

Trueman S., Warburton C., James E., Fripp Y. &. Wallace H. (2001) “Clonality in remnant 

populations of Santalum lanceolatum.” Sandalwood Research Newsletter 14, 1–4. Abstract. 

Santalum lanceolatum, the northern sandalwood or plumbush, was very heavily harvested in 

Victoria and New South Wales in the late 1800s. Clearing, fire and grazing have also contributed 

to the species’ decline. Only seven populations remain in Victoria, where we studied the five 

southernmost populations of the species. Since exclusion of grazing animals, the remnant 

populations have been reproducing asexually by root suckers. However, we observed little or no 

fruit production in the populations, and allozyme and RAPD analyses suggested that sexual 

reproduction had not been contributing to recruitment. Each population appeared to exist as a 

unique single clone composed of numerous ramets of a single genet. Therefore, conservation of 

the species in Victoria may require protection of all remnant populations, and possibly the 

establishment of new populations. 

Vernes T. & Robson K. (2002). “Indian sandalwood industry in Australia.” Sandalwood Research 

Newsletter 16, 1-4. 

Warburton C.L. James E.A., Fripp Y.J., Trueman S.J. & Wallace H.M. (2000) "Clonality and 

sexual reproductive failure in remnant populations of Santalum lanceolatum (Santalaceae)." 

Biological Conservation 96(1), 45-54 Abstract. Habitat fragmentation can have important 

conservation consequences for clonal plant species that possess self-incompatibility 

mechanisms, as lack of genetic variability within remnant populations may result in sexual 

reproductive failure. Allozymes and RAPDs were used in this study to determine the extent of 

clonality in remnant Victorian populations of the northern sandalwood, Santalum lanceolatum 

(Santalaceae), a species that has been heavily wild-harvested. S. lanceolatum can reproduce 

asexually by root suckers, and each population was identified as a unique single clone composed 

of numerous ramets of a single genet. Examination of pollination and fruit set indicated that little 

or no sexual reproduction was occurring in the remnants, due to pollen sterility in one population 

and self-incompatibility or pistil dysfunction in others. Clonality, genetic isolation and sexual 

reproductive failure indicate that preservation of each population, and possibly the establishment 

of new ones, should be objectives of the conservation strategy for the S. lanceolatum remnants. 

Warburton C.L. (2001) "Clonality in remnant populations of Santalum lanceolatum" Sandalwood 

Research Newsletter: 14, 1-4. Abstract. Santalum lanceolatum, the northern sandalwood or 

plumbush, was very heavily harvested in Victoria and New South Wales in the late 1800s. 

Clearing, fire and grazing have also contributed to the species’ decline. Only seven populations 

remain in Victoria, where we studied the five southernmost populations of the species. Since 

exclusion of grazing animals, the remnant populations have been reproducing asexually by root 

suckers. However, we observed little or no fruit production in the populations, and allozyme and 

RAPD analyses suggested that sexual reproduction had not been contributing to recruitment. 

Each population appeared to exist as a unique single clone composed of numerous ramets of a 

single genet. Therefore, conservation of the species in Victoria may require protection of all 

remnant populations, and possibly the establishment of new populations. 

Wharton G. (1985). “Antiquarians and sandalwood-getters: the establishment of the Cape York 

Collection at Weipa.” In: Proceedings of the North Australian Mine Rehabilitation Workshop, No 9 

Weipa, 1985. 

Wijesuriya S.R. & Fox J.E.D. (1985) “Growth and nutrient concentration of sandalwood seedlings 

grown in different potting mixtures.” Mulga Research Centre Journal 8, 33-40. 

Woodall G.S. & Robinson C.J. (2002) "Same day plantation establishment of the root 

hemiparasite sandalwood (Santalum spicatum (R Br) A DC: Santalaceae) and hosts." J Royal 

20

Soc of Western Australia 85, 37-42. Abstract. Interest and investment in a plantation sandalwood 

(Santalum spicatum (R Br) A DC) industry in southern Western Australia has been steadily 

growing over the last few years. Current plantation establishment involves planting host seedlings 

in year one and then direct sowing of untreated seeds of the parasitic sandalwood in year two or 

three. An innovative establishment technique in which host seedlings of Acacia acuminata Benth 

and partially germinated sandalwood seeds are planted on the same day was compared to the 

current establishment methods. The study showed that sandalwood and host establishment in 

one season is achievable and that it was three times more successful than the most widely used 

and promoted technique at present. Results also indicated that water availability influenced the 

germination, summer survival and growth of sandalwood. The use of small seedling hosts on 

well-watered, cleared land results in a higher rate of sandalwood establishment and growth. 

Woodall G.S. & Robinson C.J. (2002) “Direct seeding Acacias of different form & function as 

hosts for Sandalwood (Santalum spicatum).” Conservation Science Western Australia 4(3), 130- 

134. 

Woodall G.S. & Robinson C.J. (2003) “Natural diversity of Santalum spicatum host species in 

south-coast river systems and their incorporation into profitable and biodiverse revegetation” 

Australian Journal of Botany 51(6), 741 –753. 

Woodall G.S. (2004) “Cracking the woody endocarp of Santalum spicatum nuts by wetting and 

rapid drying improves germination” Australian J. of Botany 52(2), 163-169. 

Chinese Sandalwood (Santalum album). 

Chen F. (1999) "Cuttage of Santalum album." Zhong Yao Cai 22(3), 109-111. Abstract: The 

effects of cuttage times, miaternal plant ages, hormones and mediums on the taking root of a 

cutting were studied in 1991-1996. The results showed that the sprouts of germinating and 

growing 20-30 days from the cut back of maternal plant as cuttings, the rate of the taking root get 

to about 70%; the suitable cuttage time was in June to August; the proper medium was river 

sands, but the effects of hormones were not obvious. 

Chen Z.-X. & Lin L. (2001) “Influences of various extraction methods on content & chemical 

components of volatile oil of Santalum album.” Guangzhou Zhongyiyao Daxue Xuebo 18(2), 174- 

177. 

Gao Z., Wu Y., Dong Z. & Wu W. (2004) "Habit & control of pests in Santalum album." Zhong Yao 

Cai 27(8), 549-51. Abstract: The habit of 5 species pests from South China Botanical Garden was 

reported in this paper, they are Delias aglaia Linni, Zenzera coffeae Nietner, Parlatoria pergandii 

Comstock, Scarab (grub), Agrotis ypsilon Rottemberg. Their control methods were presented. 

Ma G.H.., Bunn E., Zhang J.-F., Wu G.-J. (2006) "[Evidence of dichogamy in Santalum album L.]" 

J Integrative Plant Biology 48(3),300-306. Abstract. Flowering, fruit set, embryological 

development, and pollination trials were investigated in Santalum album L. Each ovary may have 

three to four ovules. Microsporogenesis and megasporogenesis in the same flower were 

synchronized at the earlier stages of flower development. However, at anthesis, when pollen was 

mature, the magaspore had developed only to the stage of a one- to two-nucleus embryo sac. As 

the eight-nucleus embryo sac developed, some mamelon cells began to undergo programmed 

cell death, forming holes into which the eight-nucleus embryo sacs extended, becoming "N" or 

"S" shaped. The development from a two-nucleus embryo sac to a matured eight-nucleus embryo 

sac lasted up to 10 d. Fruit-set from open pollination was less than 2%. The endosperm develops 

prior to division of the zygotic embryo and one to three embryos and endosperms were formed in 

the same fruit. A mature seed usually germinates to produce one seedling; however, two and 

three seedlings from one seed were also observed, albeit at a low frequency. Pollination trials 

showed that no seed sets when inflorescences were covered with a bag; however, artificial 

pollination could improve fruit set. Our pollination trials and embryological studies proved that the 

flower of S. album is dichogamous and fruit set has high heterozygosity. 

21

Ma G.-H., YueMin H., JingFeng Z., FuLian C (2005) "Study on semi-parasitism of sandalwood 

seedlings." Journal of Tropical and Subtropical Botany 13(3),233-238. Abstract. Semi-parasitism 

of sandalwood (Santalum album) seedlings was studied on the basis of the propagation of the 

different host plant species. Sandalwood plants can grow normally without host plant during its 

seed germination and early seedling stage. However, the subsequent growth needs roots of the 

host plant. Results indicated that the host plant species had a significant impact on the growth of 

sandalwood seedlings and their root haustoria as exhibited by the differences in haustorium's 

number, size and adhesiveness. Host plant species such as Hibiscus rosa-sinensis and 

Phyllanthus reticulatus were found as good host plants for the growth of sandalwood seedlings. 

Sandalwood roots lack root hairs, but its vessels were well developed, which are suitable for 

absorption of water and nutrients from the host's roots. The semi-parasitism of sandalwood on 

Hibiscus roots was also investigated. 

Ma G-H. & Bunn E. (2007) "Embryology and pollination trials support dichogamy in Santalum 

album L." Sandalwood Research Newsletter 23 (Oct 2007) Abstract. Embryo development and 

pollination trials were studied in Santalum album L. The formation of the male (microspore) and 

female (megaspore) tissues in the same flower were synchronized during the early stages of 

flower-bud development. How-ever, at anthesis when pollen was mature, the megaspore had 

developed only to the stage of a 1-2 nucleate embryo sac. The development from 2-nucleate 

embryo sac to matured 8-nucleate embryo sac lasted up to 10 days. These results indicate 

thatthe flower of S. album is dichogamous where the pollen matures before the embryo sac. 

Following fertilisation of the ovule the endosperm developed prior to division of the zygotic 

embryo, and 1-3 embryos and endosperms were formed in the samefruit. Seed-set resulting from 

open pollination was less than 3%. No seed set was observed when inflorescences were 

coveredwith a bag; however artificial pollination increased fruit set to14%. Mature seed usually 

germinated to produce one seed-ling, but two- and three-seedlings from one seed were also 

observed at low frequency 

Li Y. (1997) "Preliminary studies on grafting of Santalum album." Zhong Yao Cai 20(11), 543-545. 

Abstract: With the purpose of propagating high production Clone of Santalum album, the best 

season and practical method of grafting, and the selection of shoots for scion are studied. The 

preliminary results show: The best season for grafting in Guangzhou District occurs from June to 

October, when the daily mean temperature is over 25 degrees C, the side graft is recommedable; 

the scion from 1-5-year old young trees is much in favor for grafting than that from adult trees. In 

the right condition, side grafting of Santalum album has had up to 80 percent success rate. 

Lin L., Wei M., Xiao S., Xu X., Hu Z., Qiu J., Cai Y., Lu A., &Yuan L. (2000) "[The influence of 

external stimulation on content and quality of volatile oil in Lignun Santali albi]" Zhong Yao Cai. 

23(3), 152-4. Abstract. The authors analyzed the quality of Ligmum Santali Albi formed by the 

external stimulation of hormone and windburn by GC-MS-DS. The results showed that the 

content of volatile oil is 2.34% in the heart wood formed in 10 years tree age of Santalum album 

(SA) after 2 years stimulation continuously with a definite concentration of hormone, which is near 

to the 25 years tree age of SA in the same place. The GC-MS analysis showed that the content of 

santalol and other chemical components in volatile oil are similar to the 25 years tree age of SA. It 

is indicated that a definite concentration of hormone stimulated the SA may shorten the formation 

of the heart wood. The heart wood can be also formed by the broken branches after 2 years 

windburn, but its content of volatile oil is only 1/2 of the heart wood formed by hormone 

stimulation. 

Wei M, Lin L, Qiu JY, Chai YW, Lu AN, Yuan L, Liao HF, Xiao SE. (2000) "[Wind-damage effects 

on quality of heartwood of Lignum Santali Albi]" Zhongguo Zhong Yao Za Zhi 25(12), 710-3. 

Abstract. OBJECTIVE: To evaluate the wind-damage effects on quality of heartwood of Lignum 

Santali Albi. METHOD: GC-MS, TLC and pharmacodynamic test. RESULTS: The content of 

volatile oil from heartwood of Wind-damaged Lignum Santali Albi is 1.42%; the content of various 

components in the oil and the chromatography of different extracts are similar to those of 

reference drug and 25 years old trees. CONCLUSION: Wind-damage should accelerate the 

formation of heartwood of Lignum Santali Albi without influence on its quality. 

22

Yu J.G., Cong P.Z., Lin J.T., Fang H.J. (1988) "Studies on the chemical constituents of Chinese 

sandalwood oil & preliminary structures of five novel compounds". Yao Xue Xue Bao 23(11), 868- 

872. 

Yu, J. G., Cong P.Z., et al. (1993). “Studies on the structure of alpha-trans-bergamotenol from 

Chinese sandalwood oil.” Acta Pharmaceutica Sinica 28(11), 840-844. 

Zhu L.-J. Li Y.-H. Li B-L. Lu B.-Y & Xia N.H. (1993) Aromatic plants & essential constituents p60. 

South China Institute of Botany, Chinese Academy of Sciences, Hai Feng Publish Co, distributed 

by Peace Book Co. Ltd. Hong Kong, China (1993). 

East African Sandalwood. 

Cropwatch comments: ‘East African sandalwood’ includes Osyris spp. such as 

O. lanceolata & O. tenuifolia). 

Kamau P. & Wabuyele E. (2007) "Report on in-situ sustainable harvesting method for Osyris 

lanceolata (East African sandalwood) in Mbeere District, Kenya." pub: National Museum of 

Kenya, Niarobi. See http://idl-bnc.idrc.ca/dspace/handle/10625/42137 

H 

Koross K. (2008) "Kenya: Sandalwood Ban Proves Hard to Enforce." The Nation (Niaobi) 27th 

June 2008. Cropwatch comments: Story about 7 tons of sandalwood being impounded on 

Wednesday at Salawa Division in the Baringo Disatrict. Villagers from Baringo & East Pokot 

districts sell sandalwood to dealers in spite of the trade ban in 2007. The wood finds a ready 

market in China. The article goes on to speculate about corrupt officials & security officers being 

involved in the illegal trade as well as prominent individuals and politicians. 

Kreipl A. Th. & König W.A. (2004) “Sesquiterpenes from the East African sandalwood Osyris 

tenuifolia” Phytochem 65(14), 2045-2049. Abstract: The essential oil of the east African 

sandalwood Osyris tenuifolia was investigated by chromatographic and spectroscopic methods. 

Beside several already known sesquiterpenes four new compounds could be isolated by 

preparative gas chromatography and their structures investigated by mass spectroscopy and 

NMR techniques. Two of the new compounds – tenuifolene (17) and ar-tenuifolene (15) – show a 

new sesquiterpene backbone. 2,(7Z,10Z)-Bisabolatrien-13-ol (23) and the cyclic ether 

lanceoloxide (21) belong to the bisabolanes. 

Graphical Abstract: The essential oil of East African sandalwood Osyris tenuifolia was 

investigated by NMR, Mass spectrometry and chemical correlations. Four new sesquiterpenes 

including 15 and 17 with a new skeleton were identified. 

Mwang’ingo P.L. & Mwihomeke S.T. (1997) “Some highlight on a research program into 

cultivation of Osyris lanceolata (African sandalwood).” In: Mbwambo, L.R., Mwang’ingo, P.L., 

Masayanyika S.W and Isango, J.A (eds.). Proceedings of the Second Workshop on Setting 

Forestry Research Needs and Priorities. 18-22 August 1997 Moshi Tanzania. TAFORI, Morogoro, 

Tanzania. pp 82-84. 

Mwang'ingo P.L. (2002) "Ecology and silviculture of Osyris lanceolata (African Sandalwood) An 

aromatic tree of Tanzania." - see http://opensigle.inist.fr/handle/10068/365227 

Mwang'ingo P.L.., Teklehaimanot Z., Hall J.B. & Lulanda L.L. (2003) "African Sandalwood (Osyris 

lanceolata): resource assessment & quality variation among poulations in Tanzania: research 

note." Southern Hemisphere Forestry Journal 199, 77-88. Abstract. African sandalwood (Osyris 

lanceolata) populations occurring in Tanzania were assessed to determine the current resource 

status and ascertain variation in quality existing among them. This will provide a guide in the 

selection of populations where conservation efforts and improvement programmes can be 

concentrated. The resource status was assessed through estimation of the species' density per 

unit area and measurements of tree dimensions. Quality variation was assessed by determining 

the amount of oil extracted from a given amount of wood and the proportion composition of 

23

santalol, a prime determinant of sandalwood oil quality. The study revealed that populations 

supporting O. lanceolata in Tanzania occur mostly in arid to semiarid areas with the majority 

being on stony and rocky soils. However, big sized trees are supported in humid climates, being 

favoured by relatively low soil pH and reasonable amounts of soil nitrogen. Tree density ranged 

from 38 individuals to 76 per hectare. The mean tree height was 3, 8 m (2, 1 to 6, 5 m) while the 

mean diameter was 5, 7 cm (3, 6 cm to 8, 6 cm). The best quality and quantity of oil came from 

populations of relatively arid climates compared to those of humid climates. Populations differed 

significantly in both yield and quality. The highest yield obtained was 8, 45 ± 0, 54% from Gubali 

population while the highest santalol content (32, 2 ± 1, 2%) was from Bereko populations. Within 

trees, quantity and quality of oil was higher in wood portions close to the ground in both the root 

and shoot system. The amount decreased toward the root and shoot tip. The root and the shoot 

system were similar in quality and quantity of oil. The observed harvesting selectivity is thus 

primarily influenced by quality differences among populations while the large dimension and 

density differences among populations seem to be secondary. Inclusion of the root systems 

during harvesting is also a matter of maximizing the raw material to be collected rather than 

differences between the two portions. The exact factors controlling wood quality in the species 

have however remained uncertain. Probably, genetic factors alone or in combination with the 

environmental factors play a significant role. 

Mwang’ingo P.L., Teklehaimanot Z., Hall J.B, Zilihona J.E. (2007) "Sex distribution, reproductive 

biology and regeneration In the dioecious species Osyris lanceolata (African Sandalwood) In 

Tanzania." Tanzania Journal of Forestry and Nature Conservation 76, 118-133. Abstract. Sex 

distribution, reproductive biology and regeneration of African Sandalwood (Osyris lanceolata) 

were assessed in six natural populations of Tanzania between January 1999 and February 2001. 

The aim was to acquire basic information required for efficient management, conservation and 

sustainable utilization of the species. The study had four objectives: to assess the spatial 

distribution of male and female trees in O. lanceolata supporting stands and whether this has any 

significance in influencing the reproductive success; to document the phenological events 

occurring between flower initiation and fruit ripening; to examine the reproductive success of 

various stages through pollination experiment; and to assess the regeneration mode and potential 

of the species. The study revealed that, the distribution of male and female trees in most 

populations was random with no evidence of sex clustering. It takes 104 days from flowering until 

when 25% of fruit initiated become ripe. About 75% of the initiated fruits become ripe in 163 days. 

This study has also demonstrated absence of agamospermy behaviour in O. lanceolata. A limited 

reproductive success was noted however, due to either low level of pollen production or limited 

pollinators' movement. Assisted pollination significantly increased the reproductive success of the 

species. The tree regenerates through seeds, rootstocks and coppice. Of the total regenerating 

plants assessed at sapling stage, 61% had originated from rootstock or coppice while 39% came 

from seed source. It is concluded that, recruitment of the species relies mainly on rootstock or 

coppice source although the importance of seeds cannot be ignored. Thus uprooting of the 

species as a mode of harvesting has to be discouraged since the practice is likely to severely limit 

the recruitment rate. 

Mwang’ingo P.L, Teklehaimanot Z., Maliondo S.M. & Msanga H.P. (2004). "Storage and presowing 

treatment of recalcitrant seeds of Africa Sandalwood (Osyris lanceolata)." Seed Science 

and Technology, 32, 547-560. Abstract. The best seed conditions and environment in which 

seeds of Osyris lanceolata could be stored to prolong their life span were investigated at Iringa 

Tree Seed Centre, Tanzania, by varying the storage moisture content of seeds and storage 

temperatures. The study also investigated the effectiveness of various seed pre-sowing 

treatments in enhancing germination and early seedling growth. Seeds stored at 3-5°C, after 

being dried to moisture content of 20% retained viability longer than those stored at other 

conditions. By the end of the 36th week, the viability was 60% with 0.5% being as an estimated 

rate of viability loss per week. Temperatures below 3°C and over 13°C decreased rapidly the life 

span of seeds. Moisture content below 15% and over 25% were also noted to be lethal. Thus 

seeds of O. lanceolata could be stored at least for short-term supply, although their life span 

generally remains short, suggesting the need for further research to find out other better storage 

24

conditions. The seed coat covering the embryo plays a significant role in limiting germination by 

restricting gas and water entry. It also acts as a mechanical barrier to embryo growth. Complete 

removal of the seed coat and soaking in hot water enhanced seed germination (66.5% and 

57.5%, respectively), shortened the time of seed to commence germination and promoted early 

seedling growth and are thus recommended for adoption. Nevertheless, the highest germination 

(66.5%) attained in this study is still unsatisfactory, suggesting the existence of other types of 

dormancies. This calls for further investigation to identify the dormancies and the means of 

resolving them. The possible existence of chemical dormancies, which was not dealt with in the 

present study, be given a priority in future research. 

Mwang'ingo P.L., Teklehaimanot Z., Lulandala L.L. & Mwihomeke S.T. (2005). "Host plants of 

Osyris lanceolata (African Sandalwood) and their influence on its early growth performance in 

Tanzania." Southern African Forestry Journal 203(1), 55-66. Abstract. Identification of the host 

plants of the hemi-parasitic African sandalwood (Osyris lanceolata) and the influence of some on 

its early growth performance was investigated at Image, Nundu, Sao Hill and Iringa in the 

southern highlands of Tanzania. The aim was to identify host plants that support the growth of O. 

lanceolata, and to evaluate the potential of some in promoting its early growth under artificial 

establishment. The results revealed that O. lanceolata parasitises a wide range of hosts although 

some were preferred. The preferred hosts were Rhus natalensis, Dodonaea viscosa, Tecomaria 

capensis, Catha edulis, Apodytes dimidiata, Brachystegia spiciformis, Maytenus acuminatus and 

Aphloia theiformis. Of the preferred hosts, Brachytegia spiciformis, Rhus natalensis and 

Casuarina equisetifolia promoted most effectively the early growth of O. lanceolata in terms of 

height, diameter and overall root and shoot biomass. Possibly the light crown of these host 

species and the nitrogen fixing ability of C. equisetifolia played a significant role in conferring this 

advantage. The species are thus recommended as appropriate host plants when raising O. 

lanceolata seedlings for planting. However, a decision on whether these hosts will support the 

growth of O. lanceolata at a later stage is subject to further experimentation as they may only be 

serving as initial or intermediate hosts as reported in a related species Santalum album. 

Mwang’ingo P. L. Teklehaimanot Z., Lulandala L. L. & Maliondo S. M. (2006) "Propagating Osyris 

lanceolata (African sandalwood) through air layering: Its potential and limitation in Tanzania." 

Southern African Forestry Journal 207, 7-14. Synopsis. Propagation of African sandalwood 

(Osyris lanceolata) by air layering (marcotting) was investigated at Sao Hill, Tanzania, aiming at 

providing an alternative propagation technique to the use of seeds or cuttings that germinate or 

root poorly. Air layers were initiated on the young shoots (1 – 2 years old) of mature O. lanceolata 

trees growing at Sao Hill catchment Forest. After root initiation, which took 8 weeks, they were 

detached from the parents, potted in polyethylene tubes and reared at the nursery for a further 

three months. The factors assessed in this experiment were the effect of time at which air layers 

were initiated (i.e. February, June, September and December); and the influence of IBA as 

rooting promoter at three concentrations (50, 100 and 150 ppm). From the data collected it was 

observed that rooting success of up to 80% can be achieved from air layers, making this 

propagation technique a viable alternative to seedlings or cutting propagation. Rooting success 

was influenced by both the season and application of rooting hormone with optimal rooting being 

achieved during June and September with the addition of IBA at a rate of 50 ppm. The 

significance increase in rootability of air layers during June and September may be linked to the 

advantage of the dry season in Tanzania where reduction of plant development activities such as 

budding, leafing and flowering in the dormant dry season might have reduced resource 

competition and thus promoting the observed rooting. 

Mwang’ingo P.L., Teklehaimanot Z., Hall J.B., Zilihona L.E. (2007) "Sex distribution, reproductive 

biology and regenerationiIn the dioecious species Osyris lanceolata (African Sandalwood) nn 

Tanzania." Tanzania Journal of Forestry and Nature Conservation 76, 118-144. Abstract. Sex 

distribution, reproductive biology and regeneration of African Sandalwood (Osyris lanceolata) 

were assessed in six natural populations of Tanzania between January 1999 and February 2001. 

The aim was to acquire basic information required for efficient management, conservation and 

sustainable utilization of the species. The study had four objectives: to assess the spatial 

25

distribution of male and female trees in O. lanceolata supporting stands and whether this has any 

significance in influencing the reproductive success; to document the phenological events 

occurring between flower initiation and fruit ripening; to examine the reproductive success of 

various stages through pollination experiment; and to assess the regeneration mode and potential 

of the species. The study revealed that, the distribution of male and female trees in most 

populations was random with no evidence of sex clustering. It takes 104 days from flowering until 

when 25% of fruit initiated become ripe. About 75% of the initiated fruits become ripe in 163 days. 

This study has also demonstrated absence of agamospermy behaviour in O. lanceolata. A limited 

reproductive success was noted however, due to either low level of pollen production or limited 

pollinators\' movement. Assisted pollination significantly increased the reproductive success of 

the species. The tree regenerates through seeds, rootstocks and coppice. Of the total 

regenerating plants assessed at sapling stage, 61% had originated from rootstock or coppice 

while 39% came from seed source. It is concluded that, recruitment of the species relies mainly 

on rootstock or coppice source although the importance of seeds cannot be ignored. Thus 

uprooting of the species as a mode of harvesting has to be discouraged since the practice is 

likely to severely limit the recruitment rate. 

Srikrishnaa A. & Beeraiah B. (2005) "First synthesis of (′)-tenuifolene and ar-tenuifolene." Indian J 

of Chemistry Sect B. 44(8), 1641-1643. Abstract. First total synthesis of the sesquiterpenes (′)- 

tenuifolene and (′)-ar-tenuifolene, isolated from the essential oil of the East African sandalwood 

tree Osyris tenuifolia, has been accomplished. 

Teklehaimanot Z., Mwang ingo P. L., Mugasha A. G. & Ruffo, C. K. (2004) "Influence of the origin 

of stem cutting, season of collection and auxin application on the vegetative propagation of 

African Sandalwood (Osyris lanceolata) in Tanzania." Southern African Forestry Journal 201, 13- 

24. Abstract. An investigation into the possibility of propagating O. lanceolata through stem 

cutting was carried out at Tanzania Tree Seed Agency, Iringa Zone, Tanzania. The aim was to 

test the potential of stem cuttings in providing an alternative/supplement to the use of seeds that 

are constrained with germination and storage problems. Three treatments were investigated on 

the rooting success and subsequent nursery performance of the cuttings: the effect of season at 

which cuttings are collected i.e. December, February, June and September; the effect of origin of 

stem cutting in a shoot, i.e. basal and terminal portions; and the effect of different levels of IBA as 

root promoters, i.e. 0, 50, 100 and 150 ppm. The results revealed that stem cuttings collected 

from the sprouting stumps have a potential to be used in propagating O. lanceolata. Season at 

which cuttings are collected; origin of the stem cuttings in a shoot and application of auxins 

influenced the rooting success. Stem cuttings collected in September, originating from the basal 

portion had the best rooting (43.8 + 3.9%). This is possibly related to the high levels of stored 

food in the plant after undergoing active photosynthesis during the rain season, November-May. 

Auxin application in interaction with the season at which cuttings were collected enhanced the 

number of cuttings that rooted, the number of roots formed (13 + 0.4), the length (14 + 0.3 cm) 

and biomass of roots (6.95 + 3.9 g) produced. The concentration to be applied for effective 

rooting depended on the season at which cuttings were collected. Of the origin of stem cuttings, 

basal portions had better rooting than the terminal portion. The high nutrition status and low 

nitrogen content of basal portions may play a role in enhancing their performance. Thus when 

raising O. lanceolata from stem cuttings, best rooting is obtained from those raised between June 

and September using cuttings from the basal origin of the juvenile shoots. Application of IBA 

between 50 and 100 ppm further enhances rooting success. 

Wells R. (2006) "On the scent: Rhona Wells investigates sandalwood poaching, the ugly 

downside of the luxurious natural perfumery raw material trade" Soap, Perfumery & Cosmetics 

Feb 2006 79(2), 31. Cropwatch comments: Informative one-page article on the Tanzanian 

situation where sandalwood logs are smuggled to India for distillation to produce sandalwood oil. 

Yeboah E.M.O., Majinda R.R.T., Kadziola A. & Muller A. (2010) "Dihydro-β-agarofuran 

Sesquiterpenes and Pentacyclic Triterpenoids from the Root Bark of Osyris lanceolata." J Nat 

26

Products May 2010. Abstract. 

Three new dihydro-β-agarofuran polyesters, 1α,9β-difuranoyloxy-2-oxodihydro-β-agarofuran (1), 

1α,9β-difuranoyloxy-2-oxo-3-enedihydro-β-agarofuran (2), and 1α,9β-difuranoyloxydihydro-βagarofuran 

(3), have been isolated from the CHCl3 extract of the root bark of Osyris lanceolata, 

together with two known pentacylic triterpenoids, 4 and 5. Compounds 1−5 did not scavenge the 

DPPH radical within 30 min of reaction time. All five compounds displayed antifungal activity 

against Candida albicans. Compounds 1, 3, 4, and 5 showed antibacterial activity against the 

Gram-positive Bacillus subtilis and Staphylococcus aureus and Gram-negative Escherichia coli 

and Pseudomonas aeruginosa, with 4 and 5 being the most active. Compound 2 displayed weak 

antibacterial activity only against Escherichia coli. 

East Indian Sandalwood (Santalum album). 

Biocidal properties – E.I. Sandalwood oil. 

Amer A. & Mehlhorn H. (2006) "Larvicidal effects of various essential oils against Aedes, 

Anopheles, and Culex larvae (Diptera, Culicidae)." Parasitol Res. 99(4), 466-72. Abstract. 

Mosquitoes in the larval stage are attractive targets for pesticides because mosquitoes breed in 

water, and thus, it is easy to deal with them in this habitat. The use of conventional pesticides in 

the water sources, however, introduces many risks to people and/or the environment. Natural 

pesticides, especially those derived from plants, are more promising in this aspect. Aromatic 

plants and their essential oils are very important sources of many compounds that are used in 

different respects. In this study, the oils of 41 plants were evaluated for their effects against thirdinstar 

larvae of Aedes aegypti, Anopheles stephensi and Culex quinquefasciatus. At first, the oils 

were surveyed against A. aegypti using a 50-ppm solution. Thirteen oils from 41 plants (camphor, 

thyme, amyris, lemon, cedarwood, frankincense, dill, myrtle, juniper, black pepper, verbena, 

helichrysum and sandalwood) induced 100% mortality after 24 h, or even after shorter periods. 

The best oils were tested against third-instar larvae of the three mosquito species in 

concentrations of 1, 10, 50, 100 and 500 ppm. The lethal concentration 50 values of these oils 

ranged between 1 and 101.3 ppm against A. aegypti, between 9.7 and 101.4 ppm for A. 

stephensi and between 1 and 50.2 ppm for C. quinquefasciatus. 

Benencia F. & Courreges M.C. (1999) “Antiviral activity of sandalwood oil against herpes simplex 

viruses-1 and -2.” Phytomed. 6(2), 119-23. Abstract: Sandalwood oil, the essential oil of Santalum 

album L., was tested for in vitro antiviral activity against Herpes simplex viruses-1 and -2. It was 

found that the replication of these viruses was inhibited in the presence of the oil. This effect was 

dose-dependent and more pronounced against HSV-1. A slight diminution of the effect was 

observed at higher multiplicity of infections. The oil was not virucidal and showed no cytotoxicity 

at the concentrations tested. 

Courreges B.F. (1999). "Antiviral activity of sandalwood oil against Herpes simplex viruses- 1 and 

2." Phytomedicine 6, 119-123. 

Jirovetz L., Buchbauer G., Dednkova Z., Stoyanova A., Murgov I., Gearon V., Birkbeck S., 

Schmidt E., Gelssler M. (2006) "Comparative study on the antimicrobial activities of different 

sandalwood essential oils of various origin." Flavour and Fragrance Journal 21(3), 465 - 468. 

Abstract. In total, eight samples of different sandalwoods [Amyris balsamifera L., Santalum album 

L. and Santalum spicatum (R.Br.) A.DC.] and a mixture of - and -santalols, as well as eugenol as 

reference compound, were tested by an agar dilution and agar diffusion method for their 

antimicrobial activities against the yeast Candida albicans, the Gram-positive bacterium 

27

Staphylococcus aureus and the Gram-negative bacteria Escherichia coli, Pseudomonas 

aeruginosa and Klebsiella pneumoniae. The main compounds of each essential oil were 

investigated by gas chromatographic-spectroscopic (GC-FID and GC-MS) and -olfactory methods 

to obtain information about the inßuence of these volatiles on the observed antimicrobial effects. 

For the santalol mixture, as well as for one S. album and one S. spicatum sample with moderate 

concentrations of santalols, antimicrobial activity was found against all the strains used. The A. 

balsamifera sample, containing only a small quantity of -santalol and nearly no -santalol, showed 

high effects only against Klebsiella pneumoniae, while against the other strains weak or no 

activity was observed. Therefore, santalols in medium and/or high concentrations in sandalwood 

oils show a significant influence on antimicrobial potential in such natural products. 

Ochi T., Shibata H., Higuti T., Kodama K.H., Kusumi T., Takaishi Y "Anti-Heliobacter pylori 

compounds from Santalum album."J. Nat Products 68(6), 819-824. Abstract: Six new 

sesquiterpenes, (Z)-2-beta-hydroxy-14-hydro-beta-santalol (1), (Z)-2alpha-hydroxy-albumol (2), 

2R-(Z)-campherene-2,13-diol (3), (Z)-campherene-2beta,13-diol (4), (Z)-7-hydroxynuciferol (5), 

and (Z)-1beta-hydroxy-2-hydrolanceol (6), together with five known compounds, (Z)-alphasantalol 

(7), (Z)-beta-santalol (8), (Z)-lanceol (9), alpha-santaldiol (10), and beta-santaldiol (11), 

were isolated from Santalum album, by using bioassay-guided fractionation for Helicobacter 

pylori. The structures were determined by extensive NMR studies. The absolute configuration of 

compound 3 was determined by a modified Mosher method. The crude extracts as well as the 

isolated compounds showed antibacterial activity against H. pylori. Especially, compounds 7 and 

8 have strong anti-H. pylori activities against a clarithromycin-resistant strain (TS281) as well as 

other strains. 

Radhakrishnan A.N. & Giri K.V. (1954) "The isolation of allohydroxy-l-proline from sandal 

(Santalum album L.)." Biochem J. 58(1), 57-61. 

Shankaranarayana K.H., Shivaramakrishnan V.R., Ayya K.S. & Sen P.K. (1979) “Isolation of a 

compound from the bark of sandal and its activity against some lipidopterous & coleopterous 

insects.” J. Entomol. Res 3, 116-118. 

Shankaranarayana K.H., Ayyar K.S. & Rao G.S.K. (1980) "Insect growth inhibitor from the bark of 

Santalum album." Phytochemistry 19(6), 1239-1240. 

Schnitzler P., Koch C. & Reichling J. (2007) "Susceptibility of drug-resistant clinical herpes 

simplex virus type 1 strains to essential oils of ginger, thyme, hyssop, and sandalwood." 

Antimicrob Agents Chemother. 51(5):1859-62. Abstract. Acyclovir-resistant clinical isolates of 

herpes simplex virus type 1 (HSV-1) were analyzed in vitro for their susceptibilities to essential 

oils of ginger, thyme, hyssop, and sandalwood. All essential oils exhibited high levels of virucidal 

activity against acyclovir-sensitive strain KOS and acyclovir-resistant HSV-1 clinical isolates and 

reduced plaque formation significantly. 

Zhu J., Zeng X., O'Neal M., Schultz G., Tucker B., Coats J., Bartholomay L. & Xue RD. (2008) 

"Mosquito larvicidal activity of botanical-based mosquito repellents." J Am Mosq Control Assoc. 

24(1),161-8. Abstract. The larvicidal activity of 4 plant essential oils--innamon oil, lemon 

eucalyptus oil, sandalwood oil, and turmeric oil--previously reported as insect repellents was 

evaluated in the laboratory against 4th instars of Aedes albopictus, Ae. aegypti, and Culex 

pipiens. Sandalwood oil appeared to be the most effective of the larvicides, killing larvae of all 3 

mosquito species in relatively short times. The values of LT50 and LT90 at the application dosage 

(0.2 mg/ml) were 1.06 +/- 0.11 and 3.24 +/- 0.14 h for Ae. aegypti, 1.82 +/- 0.06 and 3.33 +/- 0.48 

h for Ae. albopictus, and 1.55 +/- 0.07 and 3.91 +/- 0.44 h for Cx. pipiens, respectively. Chemical 

compositions of these essential oils were also studied, and the lavicidal activity of their major 

ingredient compounds was compared with that of each of the essential oils. The acute toxicity of 

the 4 essential oils to fathead minnows was also evaluated. The safe use of these natural plant 

essential oils in future applications of mosquito control was discussed. 

Contact Dermatitis – E.I. Sandalwood oil. 

28

An S., Lee A.Y., Lee C.S., Kim D.W., Hahm J.H., Kim K.J., Moon K.C., Won Y.H., Ro Y.S., Eun 

H.C. (2005) "Fragrance contact dermatitis in Korea: a joint study." Contact Dermatitis 53(6) , 320- 

323. Abstract: The purpose of this study is to determine the frequency of responses to selected 

fragrances in patients with suspected fragrance allergy and to evaluate the risk factors. 9 

dermatology departments of university hospitals have participated in this study for the past 1 

year. To determine allergic response to fragrances, 18 additional fragrances in addition to the 

Korean standard and a commercial fragrance series were patch-tested in patients with suspecting 

cosmetic contact dermatitis. Over 80% of the patients were women, and the most common site 

was the face. Cinnamic alcohol and sandalwood oil (Santalum album L.) showed high 

frequencies of positive responses. Of the specific fragrances, ebanol, alpha-isomethyl-ionone 

(methyl ionone-gamma) and Lyral (hydroxyisohexyl 3-cyclohexane carboxdaldehyde) showed 

high positive responses. We compared the results obtained during this study with those of other 

studies and concluded that including additional fragrance allergens may be useful for the 

detection of fragrance allergy. 

Viardot-Helmer A., Merk HF, Hausen BM (2008) “[Delayed hypersensitivity to East Indian 

rosewood.]. Hautarzt. 59(6):465-466. 

Sharma R., Bajaj A.K. & Singh K.G. (1987) “Sandalwood dermatitis” Int. J. Dermatol 26(9), 597. 

Cropwatch comments: A short report about a man who had been appying Santalum album 

paste to his forehead daily for eight years. He presented with a well defined, hyperpigmented, 

erythematous plaque, with a mild surrounding zone of erythma. Patch tests proved positive to 

sandalwood, and the lesion disappeared after the application of a corticosteroid cream. 

Tewary M, Ahmed I. (2006) "Bindi dermatitis to 'chandan' bindi." Contact Dermatitis. 55(6), 372-4. 

Abstract. Bindi (meaning dot in Sanskrit) is a mark worn by most Indian women on their forehead 

for religious and social purposes. Traditionally it was worn by only Hindu women to signify their 

marital status. Nowadays, it is a huge fashion accessory, being worn in different sizes, shapes, 

designs and colours. The variety includes sequined designs, motifs dusted with gold and silver 

powder, studded with beads, or even surrounded by glittering gems. Stick-on and liquid ranges 

are both equally in demand. We report a case of bindi dermatitis with 'chandan' (sandalwood) 

bindi. To our knowledge this is the first report of contact allergic dermatitis to 'chandan' 

(sandalwood) bindi. 

Cancer Chemoprevention – E.I. Sandalwood oil. 

Arasada B.L., Bommareddy A., Zhang X., Bremmon K. & Dwivedi C. (2008) "Effects of alphasantalol 

on proapoptotic caspases and p53 expression in UVB irradiated mouse skin." Anticancer 

Res. 28(1A), 129-32. Abstract. BACKGROUND: Cancer chemoprevention by naturally occurring 

agents, especially phytochemicals, minerals and vitamins has shown promising results against 

various malignancies in a number of studies both under in vitro and in vivo conditions. One such 

phytochemical, alpha-santalol, a major component of sandalwood oil, is effective in preventing 

skin cancer in both chemically and UVB-induced skin cancer development in CD-1, SENCAR and 

SKH-1 mice; however, the mechanism of its efficacy is not fully understood. The objective of the 

present investigation was to study the effects of alpha-santalol on apoptosis proteins and p53 in 

UVB-induced skin tumor development in SKH-1 mice to elucidate the mechanism of action. 

MATERIALS AND METHODS: Female SKH-1 mice were divided into two groups: Group 1, which 

served as control received topical application of acetone (0.1 ml) one hour before UVB treatment; 

Group 2 received alpha-santalol (0.1 ml, 5% w/v in acetone, topical) one hour prior to UVB 

treatment. UVB-induced promotion was continued for 30 weeks. RESULTS: Pre-treatment with 

alpha-santalol one hour prior to UVB exposure significantly (p < 0.05) reduced tumor incidence 

and multiplicity, and resulted in a significant (p < 0.05) increase in apoptosis proteins, caspase-3 

and -8 levels and tumor suppressor protein, p53. CONCLUSION: These results suggest that 

alpha-santalol prevents skin cancer development by inducing proapoptotic proteins via an 

extrinsic pathway and increasing p53. 

29

Banerjee S., Ecavade A. & Rao A.R. (1993) “Modulatory influence of sandalwood oil on mouse 

hepatic glutathione S-transferase activity and acid soluble sulpydryl level” Cancer Lett 68(2), 

105-9. Abstract: The effect of the oil from the wood of Santalum album on glutathione S- 

transferase (GST) activity and acid soluble sulphydryl (SH) levels in the liver of adult male Swiss 

albino mice was investigated. Oral feeding by gavage to mice each day with 5 and 15 microliters 

sandalwood oil for 10 and 20 days exhibited an increase in GST activity in time- and doseresponsive 

manners. Feeding a dose of 5 microliters sandalwood oil for 10 and 20 days caused, 

respectively, a 1.80-fold (P < 0.001) and 1.93-fold (P < 0.001) increase in GST enzyme activity, 

while feeding a dose of 15 microliters of the oil per day for 10 and 20 days induced, respectively, 

4.73-fold (P < 0.001) and 6.10-fold (P < 0.001) increases in the enzyme's activity. In addition, 

there were 1.59-fold (P < 0.001) and 1.57 (P < 0.001) increases in acid-soluble SH levels in the 

hepatic tissue of the mice following feeding of the oil at the dose levels of 5 and 15 microliters for 

10 days. Furthermore, mice fed on a diet containing 1% 2(3)-butyl-4-hydroxyanisole (positive 

control) also showed an increase in hepatic GST activity and SH levels. Enhancement of GST 

activity and acid-soluble SH levels are suggestive of a possible chemopreventive action of 

sandalwood oil on carcinogenesis through a blocking mechanism. 

Dwivedi C. & Abu-Ghazaleh A. (1997) "Chemopreventive effects of sandalwood oil on skin 

papillomas in mice." Eur J Cancer Prev. 6(4), 399-401. Abstract. The essential oil, emulsion or 

paste of sandalwood (Santalum album L) has been used in India as an ayurvedic medicinal agent 

for the treatment of inflammatory and eruptive skin diseases. In this investigation, the 

chemopreventive effects of sandalwood oil (5% in acetone, w/v) on 7,12- 

dimethylbenz(a)anthracene-(DMBA)-initiated and 12-O-tetradecanoyl phorbol-13-acetate(TPA)- 

promoted skin papillomas, and TPA-induced ornithine decarboxylase (ODC) activity in CD1 mice 

were studied. Sandalwood oil treatment significantly decreased papilloma incidence by 67%, 

multiplicity by 96%, and TPA-induced ODC activity by 70%. This oil could be an effective 

chemopreventive agent against skin cancer. 

Dwivedi C., Guan X., Harmsen W.L., Voss A.L., Goetz-Parten D.E., Koopman E.M., Johnson 

K.M., Valluri H.B. & Matthees D.P. (2003) " Chemopreventive effects of alpha-santalol on skin 

tumor development in CD-1 and SENCAR mice." Cancer Epidemiol Biomarkers Prev. 12(2), 151- 

6. Abstract. Studies from our laboratory have indicated skin cancer chemopreventive effectsof 

sandalwood oil in CD-1 mice. The purpose of this investigation was to study the skin cancer 

chemopreventive effects of alpha-santalol, a principal component of sandalwood oil in CD-1 and 

SENCAR mice. alpha-Santalol was isolated from sandalwood oil by distillation under vacuum and 

characterized by nuclear magnetic resonance and gas chromatography-mass spectrometry. 

Chemopreventive effects of alpha-santalol were determined during initiation and promotion phase 

in female CD-1 and SENCAR mice. Carcinogenesis was initiated with 7,12- 

dimethylbenz(a)anthracene and promoted with 12-O-tetradecanoylphorbol-13-acetate (TPA). The 

effects of alpha-santalol treatment on TPA-induced epidermal ornithine decarboxylase (ODC) 

activity and (3)H-thymidine incorporation in epidermal DNA of CD-1 and SENCAR mice were also 

investigated. alpha-Santalol treatment during promotion phase delayed the papilloma 

development by 2 weeks in both CD-1 and SENCAR strains of mice. alpha-Santalol treatment 

during promotion phase significantly (P < 0.05) decreased the papilloma incidence and multiplicity 

when compared with control and treatment during initiation phase during 20 weeks of promotion 

in both CD-1 and SENCAR strains of mice. alpha-Santalol treatment resulted in a significant (P < 

0.05) inhibition in TPA-induced ODC activity and incorporation of (3)H-thymidine in DNA in the 

epidermis of both strains of mice. alpha-Santalol significantly prevents papilloma development 

during promotion phase of 7,12-dimethylbenz(a)anthracene-TPA carcinogenesis protocol in both 

CD-1 and SENCAR mice, possibly by inhibiting TPA-induced ODC activity and DNA synthesis. 

alpha-Santalol could be an effective chemopreventive agent for skin cancer. Additional 

experimental and clinical studies are needed to investigate the chemopreventive effect of alphasantalol 

in skin cancer. 

Dwivedi C., Maydew E.R., Hora J.J., Ramaeker D.M. & Guan X. (2005) "Chemopreventive effects 

of various concentrations of alpha-santalol on skin cancer development in CD-1 mice." Eur J 

30

Cancer Prev. 14(5), 473-6. Abstract. Previous studies from this laboratory have indicated that 

alpha-santalol (5%) provides chemopreventive effects in 7,12-dimethylbenz[a]anthracene 

(DMBA)-initiated and 12-O-tetradecanoylphorbol-13-acetate (TPA)-promoted skin cancer in CD-1 

and SENCAR mice. Skin cancer development is associated with increased ornithine 

decarboxylase (ODC) activity, DNA synthesis and rapid proliferation of epidermal cells. The 

purpose of this investigation was to determine the effects of various concentrations (1.25% and 

2.5%) of alpha-santalol on DMBA-initiated and TPA-promoted skin cancer development, TPAinduced 

ODC activity, and DNA synthesis in CD-1 mice. alpha-Santalol treatment at both 

concentrations (1.25% and 2.5%) prevented the skin cancer development. alpha-Santalol 

treatment (1.25% and 2.5%) resulted in a significant decrease in the TPA-induced ODC activity 

and incorporation of [3H]thymidine in DNA in the epidermis of CD-1 mice. There was no 

significant difference in the effects of 1.25% and 2.5% alpha-santalol on tumour incidence, 

multiplicity, epidermal TPA-induced ODC activity, or DNA synthesis in CD-1 mice. 

Dwivedi C., Valluri H.B., Guan X. & Agarwal R. (2006) "Chemopreventive effects of alpha-santalol 

on ultraviolet B radiation-induced skin tumor development in SKH-1 hairless mice." 

Carcinogenesis. 27(9),1917-22. Abstract. Recent studies from our laboratory have shown the 

chemopreventive effects of alpha-santalol against 7,12-dimethylbenzanthracene (DMBA) initiated 

and 12-O-tetradecanoylphorbol-13-acetate (TPA) promoted skin tumor development in mice. The 

objective of the present investigation was to study the effects of alpha-santalol on ultraviolet B 

(UVB) radiation-induced skin tumor development and UVB-caused increase in epidermal 

ornithine decarboxylase (ODC) activity in female hairless SKH-1 mice. For the tumor studies, 180 

mice were divided into three groups of 60 mice each, and each group was divided into two 

subgroups of 30 mice. The first subgroup served as control and was treated topically on the 

dorsal skin with acetone. The second subgroup served as experimental and was treated topically 

on the dorsal skin with alpha-santalol (5%, w/v in acetone). The tumorigenesis in the first group 

was initiated with UVB radiation and promoted with TPA; in the second group it was initiated with 

DMBA and promoted with UVB radiation; and in the third group it was both initiated and promoted 

with UVB radiation. In each case, the study was terminated at 30 weeks. Topical application of 

alpha-santalol significantly (P

treatment of cells with caspase-8 or -9 inhibitor, pan caspase inhibitor or cycloheximide totally 

blocked alpha-santalol-caused caspase-3 activity and cleavage, but only partially reversed 

apoptotic cell death. This suggests involvement of both caspase-dependent and -independent 

pathways, at least under caspase inhibiting conditions, in alpha-santalol-caused apoptosis. 

Together, this study for the first time identifies the apoptotic effect of alpha-santalol, and defines 

the mechanism of apoptotic cascade activated by this agent in A431 cells, which might be 

contributing to its overall cancer preventive efficacy in mouse skin cancer models. 

Kim T.H., Ho H., Takayasu T., Tokuda H., Machiguchi M. & T. (2006) "New antitumor 

sesquiterpenoids from Santalum album of Indian origin." Tetrahedron 62 (29), 6981-6989. 

Abstract. Three new campherenane-type (1, 4, 7) and three new santalane-type (9, 11, 12) 

sesquiterpenoids, and two aromatic glycosides (21, 22) together with 12 known metabolites 

including β-santalols (14, 18), (E)-,β-santalals (15, 19), β-santaldiols (16, 20), -santalenoic acid 

(17), and vanillic acid 4-O-neohesperidoside were isolated from Santalum album chips of Indian 

origin. The structures of the new compounds, including absolute configurations, were elucidated 

by 1D- and 2D-NMR spectroscopic and chemical methods. The antitumor promoting activity of 

these isolates along with several neolignans previously isolated from the same source was 

evaluated for both in vitro Epstein–Barr virus early antigen (EBV-EA) activation and in vivo twostage 

carcinogenesis assays. Among them, compound 1 exhibited a potent inhibitory effect on 

EBV-EA activation, and also strongly suppressed two-stage carcinogenesis on mouse skin. 

Graphical abstract. 

Kim T.K., Ito H., Hayashi K., Hasegawa T., Machiguchi T. & Yoshida T. (2005) "Aromatic 

Constituents from the Heartwood of Santalum album L." Chem. Phar. Bull. 53(6), 641-644. 

Abstract. A phytochemical investigation of the polar constituents in the heartwood of Indian 

Santalum album L. resulted in the isolation of three new neolignans (1—3) and a new aromatic 

ester (4), along with 14 known components. The structures of the new compounds (1—4) were 

established using spectroscopic methods. 

Matsuo Y. & Mimaki Y. (2010) "Lignans from Santalum album and their cytotoxic activities." 

Chem Pharm Bull 58(4):587-90. Abstract. A new neolignan, (7R,8R)-5-O-demethylbilagrewin (1), 

together with four known lignans (2-5), were isolated from the heartwood of Santalum album 

(Santalaceae). The structure of 1 was determined by analysis of extensive spectroscopic data. 

The isolated compounds and derivatives were evaluated for their cytotoxic activities against HL- 

60 human promyelocytic leukemia cells and A549 human lung adenocarcinoma cells. 

Compounds 1 and 2 exhibited cytotoxicity against HL-60 cells with IC(50) values of 1.5+/-0.02 

and 4.3+/-0.13 microM, and against A549 cells with IC(50) values of 13.6+/-0.32 and 19.9+/-1.27 

microM, respectively. The aldehyde group of 1 and 2 was revealed to be a structural requirement 

for the appearance of cytotoxicity in this type of lignans. These tumor cell deaths were shown to 

be mediated through induction of apoptosis. 

Palep S. & Lebwohl M. (2007) "Inhibitory effects of alpha- and beta-santalol on UVB-induced 

mouse skin carcinogenesis." Journal of the American Academy of Dermatology 56(2) Suppl. 2, 

pAB36. 

Chemistry of E.I. Sandalwood. 

Anonis D.P. (1998) “Sandalwood & sandalwood compounds” Perf. & Flav. 23(5), 19-24. 

Bajgrowicz J.A. & Frater G. (2000) "Chiral recognition of sandalwood odourants." Enantiomer 

5(3-4), 225-234. Abstract: Looking for more efficient sandalwood oil smelling compounds, new 

32

campholenic aldehyde derivatives with rigidifying cyclopropane rings were prepared. For some of 

them, having the lowest odor threshold ever measured for this type of odorants and a very 

appreciated scent, close to that of the scarce natural sandalwood oils, pure stereoisomers were 

obtained and their olfactory properties were evaluated. Thus acquired structure-odor relationship 

data, together with consolidated and completed previous knowledge on structurally different 

sandalwood-smelling compounds, allowed to propose new models of the sandalwood 

olfactophore. 

Beyer A., Wolschann P., Becker A., Pranka E. & Buchbauer G. (1988) "Conformational 

calculations in odiferous molecules of sandalwood." Montash. Chem. 119, 711. 

Beyer A., Wolschann P., Becker A., Pranka E. & Buchbauer G. (1988) "Conformational 

calculations in sandalwood odour molecules" Flav. Frag. J. 3, 173. 

Bhati A. (1962) “Studies in the Sandalwood oil Series. 111. Chain Effect on Terpene 

Transformations.” J of Organic Chemistry Dec 1962 p4485. Abstract. Thc carboxyl chain of some 

moleciiles has been found to be responsible for causing rearrangements and controlling their 

course, This chain effect, which opcrates during reactions involving carbonium ions, is illustrated 

with examples from Sandalwood oil chemistry. 

Bohlmann F. & Zedro C. (1968) “Isolierung von (-)-a-santalal aus Piqueria Trinerva” Tetrahedr. 

Lett. 1533. 

Braun N.A., Meier M., Schmaus G., Hölscher B. & Pickenhagen (date) “Enantioselectivity in 

odour perception: synthesis & olfactory properties of iso-b-bisabolol, a new natural product” Helv. 

Chim. Acta 86, 2698-2708. 

Briggs C.H. (1915). “Some notes on Sandalwood, its assay, yield of oil, and changes in the oil 

during distillation.” J of Industrial. & Engineering. Chemistry 8(5), 428. 

Brocke C., Eh M. & Finke A.(2008) "Recent developments in the chemistry of sandalwood 

odorants." Chem Biodivers 5(6), 1000-10. Abstract. Natural sandalwood oil, a unique and 

valuable ingredient in fine perfumery, has been the focus of scientific interest for many years. Due 

to its scarcity and its high price, the search for novel synthetic raw materials imitating the 

characteristic odor profile of sandalwood oil is as challenging as ever. In this context, the 

preparation of the novel sandalwood odorants 26, 33, and 39 will be discussed, including their 

sensory properties and structure-odor relationship. 

Brunke E.-J. & Hammerschmidt F.-J. (1980) “New Constituents of East Indian Sandalwood oil”. 

Proceedings of VIII Congress Intl. Des Huiles Essentielles Oct 1980 Pub. 1982 Fedarom. 

Brunke E-J. & Rojahn W. (1980) “Sandalwood Oil” Dragoco Report 5/1980, 127-135. 

Brunke E-J (1983) “Woody Aroma Chemicals” Dragoco Report 6/1983 p146 

Brunke E-J. & Hammerschmidt F.-J. (1988) “Constituents of East Indian sandalwood oil – an 

eighty year old stabilty test” Dragoco Report 4/1988 pp107-113. 

Brunke E.J. & Schmaus (1995) “New active odour constituents in Sandalwood Oil: part 2: 

Isolation, structural elucidation and synthesis of nor-a-trans-bergamotenone” Dragoco Report 

6/1995 p245-257. 

Brunke, E. J., Vollhardt J., et al. (1995). “Cyclosantalal and epicyclosantalal-new sesquiterpene 

aldehydes from East Indian sandalwood oil.” Flavour and Fragrance Journal 10(3), 211-219 

Brunke E-J, Fahlbusch K-G, Schmaus G & Volhart (1997) “The chemistry of sandalwood odour – 

a review of the last 10 years”. In Rivista Ital. EPOS (Actes des 15emes Journeés Internationales 

Huilles Essentielles; Digne-les-Baines, France 5-7th Sept 1996 special issue 01/97) pp49-83. 

33

Brunke E.J. & Tumbrink L. (1986) "First total synthesis of spirosantalol." Progress in Essential Oil 

Research pp321-327. 

Brunke E.-J. Hammerschmidt F.-J. & Struwe H. (1980) "(+)-epi-Santalol isolierung aus 

sandelholzol und partialsynthese aus (+) -alpha-santalol. Tetrahedron Lett. (1980) 2405. 

Brunke E.-J. & Klein E. (1982) "Chemistry of sandalwood fragrance" In Fragrance Chemistry. The 

Science of Smell, Academic Press NY p397. 

Buchbauer G., Stappen I., Pretterklieber C & Wolschann P. (2004) “Structure–activity 

relationships of sandalwood odorants: synthesis and odor of tricyclo β-santalol” Eur J Med Chem 

39(12), 1039-1046. Abstract: In a series of structure–odor relationship investigations the 

synthesis of a new tricyclic β-santalol derivative is described. The product of a multi-step 

synthesis appears in an olfactive evaluation more or less odorless may be slightly creamy but 

definitely with no sandalwood odor. This modification with a bulky aliphatic bridge in the 

neighbourhood of the quaternary C3-atom demonstrated the sensitivity of sandalwood odor on 

the structure of β-santalol analogues. 

Buchbauer G., Winiwarter S. & Wolschann P. (1992) "Surface comparisons of some odour 

molecules: conformational calculations on sandalwood odour V.” J. Comput. Aided Mol. Des. 

6(6), 583-592. Abstract: Molecular surface comparison seems to be a very suitable tool for the 

investigation of small differences between biologically active and inactive compounds of the same 

structural type. A fast method for such comparisons, based on volume matching followed by the 

estimation of comparable surface dots, is presented and applied on a few selected sandalwood 

odour molecules. 

Demole E., Demole C. & Enggist P. (1976) “A chemical investigation of volatile constituents of 

volatile constituents of East Indian sandalwood oil (Santalum album L.)” Helv. Chim. Acta 59, 

737. Abstract. Distillation foreruns from East Indian sandalwood oil (Santalum album L.), 

representing 5-8% of the oil, have been investigated using fractional distillation, preparative 

column chromatography, gas liquid chromatography (GLC.), and chemical treatments. This 

allowed the isolation and characterization by their spectral data of 46 compounds. 32 of them 

were newly identified sandalwood oil constituents including 4 novel substances: santalone (2), 4- 

methylcyclohexa-1,3-dien-1-yl methyl ketone (4), 5,6-dimethyl-5-norbornen-exo-2-ol (7), and (E)- 

5-(2,3-dimethyl-3-nortricyclyl)-pent-3-en-2-one (20). The other constituents identified were 1- 

furfuryl-pyrrole (10) and 10 phenols accompanied by 17 terpene and sesquiterpene derivatives. 

Endo-2,endo-3-dimethyl-norbornan-exo-2-ol (6), an -santenol (z), precursor, was present in the 

last group of constituents. The compounds 2, 4, 6, 7, 20 have been synthesized as well as 

another novel constituent, endo-2-mythyl-3-methylidene-norbornan-exo-2-ol (5). 

Dimoglo A.S., Beda A.A., Shvets N.M., Gorbachov M.Yu., Kheifits L.A. & Aulchenko S. (1995) 

"Investigation of the relationship between sandalwood odour & chemical structure: electron 

topological approach.” New J of Chemistry 19(2), 149-154. 

Hatt H.H. & Schoenfeld R. (1956) “Some seed fats of the Santalaceae family.” J. Sci Food Agric 

7(2), 130-133. Cropwatch comments. The drying oil from the hard-shelled seeds (50-60% fixed 

oil) contains 30-35% santalbic acid and 1% stearolic acid. Tthese acetylenic compounds inhibit 

lipoenzymes in experimental animals. 

CH 3 

(CH 2 

) 5 

CH C 

H 

(CH 2 

) 7 

COOH 

santalbic acid 

Hayashi K., Haseegawa T., Machiguchi T. & Yoshida T. (2005) “"Isolation and structure of a new 

aroma constituent from Indian Sandalwood, Santalum album L." Nippon Kagakkai Koen Yokoshu 

85(2), 863. Abstract. This work presents the isolation and structural elucidation of a new aroma 

compound in the major component from Indian sandalwood tree, Santalum album L., not from 

sandalwood oil obtained through steam distillation. We have found that the compound has a 

34

novel hemiacetal structure and has sandalwood odor stronger than those of.alpha.- and.beta.- 

santalols (the major components of sandalwood oil). (author abst.) 

Heissler D. & Riehl J.-J. (1980) “Synthesis with benzenesulfenyl chloride. On the structure of a 

C 12 H 18 hydrocarbon from East Indian sandalwood oil” Tetrahedron Letters 21(49), 4711-4714. 

Abstract: The tetracyclic hydrocarbon was synthesized by means of the electrophilic addition of 

benzenesulfenyl chloride to an appropriately substituted methylenenorbornene. The synthetic 

methodology used to prepare this letter compound includes a mild enol ether hydrolysis with 

acidic silica gel. 

Hopkins C.Y. & Chisholm M.J. (1969) "Fatty acid composition of some Santalaceae seed oils." 

Phytochem. 8, 161-165. 

Howes M.-J. R., Simmonds M.S.J. & Kite G.C. (2003) “Evaluation of the quality of sandalwood 

essential oils by gas chromatography–mass spectrometry” Journal of Chromatography A, 

1028(2), 307-312. Abstract: Trade and historic oils from ‘sandalwoods’, labelled as Amyris 

balsamifera, Eremophila mitchelli, Fusanus acuminatus (= Santalum acuminatum), Santalum 

album, S. austrocaledonicum, S. latifolium, S. spicatum and S. yasi, were assessed using gas 

chromatography–mass spectrometry (GC–MS). Using GC–MS, none of the oils assessed 

complied with the internationally recognised standard of a 90% santalol content, and only about 

half of the trade sandalwood oils met with recent International Organisation for Standardisation 

standards. The majority of trade oils, reportedly from S. album, contained approximately 50–70% 

santalols (Z-α and Z-β). Thus, the internationally recognised specification (90% santalols) for S. 

album requires re-evaluation by more efficient analysis methods. In view of the issues associated 

with the quality of sandalwood oils being traded, specifications of ≥43% Z-α-santalol and ≥18% Z- 

β-santalol for S. album oil estimated by GC–MS are suggested. GC–MS are recommended as it 

assists with authentication and quality control issues associated with sandalwood oils. 

Cropwatch comments: The authors seem confused. The ‘90% santalols figure’ is largely a relict 

of the past from when santalols in sandalwood oil were estimated by wet chemical methods – 

either by the acetylation method e.g. by EOA Determination 1B as set out in EOA Spec. No 103, 

or by wide-bore GC. This result is inaccurate and non-comparable to the superior information 

revealed by modern high performance capillary GC/MS determinations. The latter can break 

down the identity of a number of santalol isomers within sandalwood oil, and can help identify 

other sesquiterpene alcohols which might have previously have been included with the total 

santatols figure by the wet chemical procedure. Thus, by high performance capillary gas 

chromatography, a different story unfolds, and some 16 years previously, Verghese et al. (1990a) 

established that in Sandalwood oil E.I. the normal range is as follows: α-santalol 40-45% and β- 

santalol 17-27% [Lawrence (1991) q.v.]. The ISO standard ISO 3518 (2002) for sandalwood oil is 

surely taken by most workers as the current standard for the commodity and sets the limits on the 

Z-α-santalol content to 41-55% and the Z-β-santalol content to 16-24%. Although smaller 

amounts of other santalols are present in Sandalwood oil e.g. epi-β-santalol, Cropwatch does not 

accept that the situation for steam distilled sandalwood oils is quite as Howes et al. present it to 

be. Sandalwood extracts – via the benzene (etc.) extraction of sandalwood powder to produce 

sandalwood concrete, followed by methanolic extraction to produce a so-called ‘oil’ – can 

however produce high santalol containing sandalwood commodities in higher yield than steam 

distillation, which are sometimes traded as ‘oils’ or mixed in with the normal oil. Co-distillation 

technology with high boiling solvents (which are subsequently removed) can also produce high 

santalol containing sandalwood ‘oils’. East African sandalwood oil and certain fractions of 

Australian sandalwood oil have also frequently been added as adulterants to traded East Indian 

sandalwood oils. The analyst should be aware therefore that not everything offered as 

sandalwood oil is as necessarily ‘100% derived from the named botanical source’ – but this is 

hardly news to any experienced essential oil analyst! 

Jie M. S. F. L. K., Pasha M.K. et al. (1996). “Ultrasound-assisted synthesis of santalbic acid and a 

study of triacylglycerol species in Santalum album (Linn.) seed oil.” Lipids 31(10), 1083-1089. 

35

Jirovetz L. et al. (1988). “Differentiation of double bond isomers of sesquiterpene alcohols in East 

Indian sandalwood oil by means of GC-MS and GC-FTIR: Dihydrosantalols.” Spectroscopy 6(5- 

6), 283-294. 

John M.D., Paul T.M. & Jaiswal P.K. (1991) “Detection of adulteration of polyethylene glycol in oil 

of sandalwood” Indian Perfumer 35(4), 186-187.. 

Kim T.H., Ito H., Hayashi K., Hasegawa T., Machguchi T., & Yoshida T. (2005) "Aromatic 

constituents from the heartwood of Santalum album." Chem Bull Pharm (Tokyo) 53(6), 641-646. 

Abstract: A phytochemical investigation of the polar constituents in the heartwood of Indian 

Santalum album L. resulted in the isolation of three new neolignans (1-3) and a new aromatic 

ester (4), along with 14 known components. The structures of the new compounds (1-4) were 

established using spectroscopic methods. 

Kim T.H., Ito H., Hatano T., Haswegawa T., Akiba A., Machiguchi T., Yoshida T. (2005) 

"Bisabolane & santalane-type sequiterpemoids from Santalum album of East Indian origin" J. Nat 

Products 68(12), 1805-1808. Abstract: Six new bisabolane-type (1-3) and santalane-type (4-6) 

sesquiterpenoids, together with (+)-alpha-nuciferol, (+)-citronellol, and geraniol, were isolated 

from the heartwood of Santalum album of Indian origin. Their structures, including two bisabolol 

diastereomers (1, 2), were established on the basis of spectroscopic data interpretation. 

Kovatcheva A., Buchbauer G., Golbraikh A. & Wolschann P. (2003) “QSAR modeling of alphacampholenic 

derivatives with sandalwood odor.” J Chem Inf Comput Sci. 43(1), 259-66. Abstract. 

Three-dimensional quantitative structure-activity relationship (3D-QSAR) models were developed 

for a series of 44 synthetic alpha-campholenic derivatives with sandalwood odor. These 

compounds have complex stereochemistry as they contain up to five chiral atoms. To address 

stereospecificity of odor intensity, a 3D-QSAR method was developed, which does not require 

spatial alignment of molecules. In this method, compounds are represented as derivatives of 

several common structural templates with several substituents, which are numbered according to 

their relative spatial positions in the molecule. Both wholistic and substituent descriptors 

calculated with the TSAR software were used as independent variables. Based on published 

experimental data of sandalwood odor intensities, two discrete scales of the odor intensity with 

equal or unequal intervals between the threshold values were developed. The data set was 

divided into a training set of 38 compounds and a test set of six compounds. To build QSAR 

models, a stepwise multiple linear regression method was used. The best model was obtained 

using the unequal scale of odor intensity: for the training set, the leave one out cross-validated 

R(2) (q(2)) was 0.80, the correlation coefficient R between actual and predicted odor intensities 

was 0.93, and the correlation coefficient for the test set was 0.95. The QSAR models developed 

in this study contribute to the better understanding of structural, electronic, and lipophilic 

properties responsible for sandalwood odor. Furthermore, the QSAR approach reported herein 

can be applied to other data sets that include compounds with complex stereochemistry. 

Kretschmar H.C., Barneis Z.J. & Erman W.F. (1970) “The isolation & synthesis of a novel 

tetracyclic ether from East Indian sandalwood oil. A facile intramolecular Prins reaction.” 

Tetrahedron Letters 11(1), 37-40. 

Kuttan R. & Radhakrishnan A.N. (1972) "Studies on the biosynthesis of sym-homospermidine in 

sandal (Santalum album L.)." Biochem J. 127(1), 61-67. Abstract. The biosynthesis of the newly 

isolated polyamine, sym-homospermidine (NH2–[CH2]4–NH–[CH2]4 –NH2), was studied by 

using radioactive amino acids. Arginine was the most effective precursor, being about 10 times as 

active as ornithine. Unlabelled agmatine and putrescine markedly inhibited the incorporation of 

[14C]arginine into homospermidine. Similarly the incorporation of ornithine was inhibited by 

unlabelled arginine and putrescine. γ-Aminobutyraldehyde, the oxidation product of putrescine, 

was considered to be one of the intermediates in the biosynthesis of homospermidine. The 

biosynthesis may involve a Schiff-base formation of putrescine with γ-aminobutyraldehyde and 

subsequent reduction. A limited synthesis of spermidine also takes place under these conditions. 

36

Lawrence B. M. (1991) “Progress in Essential Oils: Sandalwood Oil” Perf & Flav. 16(6), 50-52. 

Lawrence B. M. (1981) “Progress in Essential Oils: Sandalwood Oil” Perf & Flav 6(5), 32-34. 

Lawrence B. M. (1976) “Progress in Essential Oils: Sandalwood Oil” Perf & Flav. 1(5),14. 

Lawrence B. M. (1976) “Progress in Essential Oils: Sandalwood Oil” Perf & Flav.1(1),5. 

Manjarrez A., Rios T. & Guzman A. (1954) “The stereochemistry of l-lanceol and the synthesis of 

its racemate” Tetrahedron 20, 333-339. 

Mookerjee B.D., Trenkle B.W. & Wilson R.A. (1992) “New insights in the three most important 

natural gragrance products: Wood, amber & musk..” In: Proceedings of 12 th International 

Congress of flavours, fragrances & essential oils. Oct 4-8 1992. Eds: H. Woldich & G. Buchbauer 

pp234-262. Austrian Assocn. Flav. Frag. Industry, Vienna, Austria (1992). 

Mörgenthaler J.M. & Spitzner (2004) “Ring-closing olefin metathesis reactions: synthesis of iso-bbisabolol” 

Tetrahedron Letters 45, 1171-1172. 

Muratore A., Clinet J.C. & Duñach E. (2010) "Synthesis of new exo- and endo-3,8-dihydro-betasantalols 

and other norbornyl-derived alcohols." Chem Biodivers. 7(3),623-38. Abstract. Several 

new and differently functionalized cis-2,3-dimethylnorbornane derivatives presenting diverse sidechain 

lengths were prepared, the structures of which are related to the natural fragrance betasantalol. 

In particular, exo- and endo-3,8-dihydro-beta-santalols, with either (E) or (Z) C==C-bond 

configuration on the side chain, were synthesized in seven steps and 21-24% overall yields. 

Several other exo- and endo-norbornyl alcohols with shorter side chains were also prepared in 

high yields. The olfactory evaluation indicated woody, sandalwood, as well as fruity notes for 

some of the derivatives. 

Nikiforov A., Jirovetz C., Buchbauer G. & Raverdino V. (1988) “GC-FTIR and GC-MS in odour 

analysis of essential oils. Mikrochim Acta (Vienna) (11), 193-198. 

Ohloff G. (1994) Scent & Fragrances Springer-Verlag p175-178. 

Pasha, M. K. & Ahmad F. (1993). “Synthesis of oxygenated fatty acid esters from santalbic acid 

ester.” Lipids 28(11), 1027-1031. 

Ruzicka L. & Thomann G. (1935) "Polyterpene und polyterpenoide XCIII. Uber die konstitution 

des beta-santalols und des beta-santalenes. Helv. Chim. Acta 18, 355. 

Sato K., Miyamoto O., Inoue S. & Honda K. (1980) "Stereospecific synthesis of beta-santalol" 

Paper presented at Proceedings the VIII Congress International des Huiles Essentielles, Cannes, 

Grasse, Oct. 1980 pub Fedarom 1982 

Schmaus G., Meier M., Braun N.A., Hölscher B. & Pickenhagen (2001) “Iso-β-bisabolol as a 

fragrance & aroma substance” WO 03/011802.Schmincke K.-H. (1985) “Sandalwood Oil” in 

Flavour & Fragrances of Plant Origin: (Non-Wood Forest Products 1) Food & Agricultural 

Organisation of the United Nation (FAO) Rome. 

Semmler F.W. (1908) "Zur Kenntnis der Bestandtelle atherische Ole. (Weitere Mitteilungen uber 

die Santole C15H24O und ihre Derivative). Ber. Dtsche. Chem. Ges. 41, 1488. 

Semmler F.W. (1910) Zur Kenntnis der Bestandtelle atherische Ole. (Konstitution der alpha- 

Santalol und alpha-Santalen-Reihe: Die Konstitution der Seasquiterpenalkohole und 

Seaquiterpene). Ber. Dtsche. Chem. Ges. 43, 1893. 

37

Shankaranarayana K.H., Angadi V.G., Rajeevalochan A.N.,. Theagarajan K.S., Sarma C.R. & 

Rangas-wamy C.R. (1997). “A rapid method ofestimating essential oil content inheartwood of 

Santalum album Linn.” Current Science 72(4), 241-242. 

Shankaranarayana K.H., Rajeevalochan G., Rajeevalochan A.N. & Angadi V.G. (2005) "Fragrant 

oils from exhausted sandalwood powder and sandal sapwood." J. Sci.Ind. Res. 64, 965-966. 

Shukla B.V., Ravindra M., Shukla S.V., Lahire L. & Singh D.P. (1999) “Qualitative assessment of 

sandalwood oil using gas chromatography” PAFAI J. 13, 41-43. 

Sidheswaran P. & Ganguli S. (1997) "Sandalwood oil substitutes - A review" Supplement to 

Cultivation & Utilisation of Aromatic Plants CIMAP (1997). 

Spreitzer, H., Roesslhuber I, et al. (1990). “Structure/odor relationships of beta-santalol 

analogues: E-homo-beta-santalol and E-dehydrohomo-beta-santalol.” Monatshefte Fuer Chemie 

121(2-3), 195-202. 

Stappena I., Höfinghoff J., Friedl S., Pammer C., Wolschann P. & Buchbauer G. (2008) 

"Structure–activity relationships of sandalwood odorants: Total synthesis and fragrance 

properties of cyclopropano-β-santalol." European Journal of Medicinal Chemistry 43(7),1525- 

1529. Abstract. The synthesis and odor properties of cyclopropano-β-santalol, a new santalol 

analogue, are described. The exocyclic double bond of the original molecule, β-santalol, is 

replaced by a cyclopropane ring. Despite the analogies in the binding properties between the 

double bond and cyclopropane this change in the bulky hydrophobic part of the molecule leads to 

the complete loss of the characteristic sandalwood odor: in an olfactory evaluation the (Z)-product 

appears spicy and sweet, the (E)-isomer woody, but neither of them exhibits the typical 

sandalwood character. 

Graphical abstract: 

Verghese J., Sunny T.P. & Balkrishnan K.V. (1990) “-(-)-α-santalol & (-)-β-santalol concentration, 

a new quality determinant of East Indian sandalwood oil.” Flavours & Fragrances J. 5, 223-226 

Walker G.T. (1968) "The chemistry of the oil of sandalwood." Perf & Essen Oil Record 59(11), 

778-781. 

Wang Z. & Hong X. (1991) "Comparitive GC analysis of essential oil in imported sandalwood" 

Zongguo Zong Yuo Za Zhi 16(1), 40-43, 64. Abstract: The GC-fingerprint spectra of essential oils 

in imported sandalwood are established by the new technique of GC-relative retention value 

fingerprint spectrum (GC-FPS). According to the GC-FPS of samples, their chromatographic 

peaks, overlap ratio of peaks and eight strong peaks are studied comparatively. 

Witteveen J.G. & van der Weerdt A.J.A. (1987) "Structure-odour relationships of some new 

synthetic sandalwood aroma chemicals - Synthesis and olefactive properties in a series of bicyclo 

[4.4.0] decan-3-ols. Rec Trauvase Chim., Pays-Bas 106, 29 

General Articles – E.I. Sandalwood. 

Adkoli N.S. & Kushalappa K.A. (eds)(1977) Proceedings of All India Sandal Seminar, Bangalore, 

7-8 February, 1977. Myforest, Special Issue. pub Karnataka Forest Dept, Bangalore. 

Ananthapadmanabha H.S. (2000) “The present status of sandalwood in India & Australia” PAFAI 

Journal 2, 33-36. 

Ananthapadmanabha H.S. (2000) “Sandalwood and its marketing trend.” My Forest 36, 147-151.. 

38

Ananthapadmanabha H.S., Nagaveni H.C. & Rai S.N. (1988). “Dormancy principles in 

sandalwood seeds (Santalum album) Linn. Myforest 24(l):22-24. 

Ananthapadmanabha H.S., Rangaswamy C.R.; Sarma C.R.., Nagaveni H.C., Jain S.H.. 

Venkatesan K.R. & Krishanappa H.P. (1984). “Host requirements of sandal (Santalum album 

L.).”. Indian Forester 110 (3). 

Angadi V.G. & Anathapadmanabha H.S. (1988). “Variations in isoenzyme pattern associated with 

spike disease in sandal.” Indian Journal of Forestry 11(1), 37-38. 

Angadi V.G., Ramalakshmi S., Jain S.H., Rangaswamy C.R. & Theagarajan K.S. (1999). 

"Allozymicvariation in the seed tissue of Sandal (Santalum album L.) population of different 

provenances." (unpublished). 

Angadi V.G., Jain S.H., Rajeevalochan A.N., Ravikumar G. & Shankaran-yana K.H. (2002) "A 

note on peroxidase reagents to distinguish between highand low yielders of Sandal (Santalum 

album) in the field." Sandalwood Research Newsletter 2002. 

Anil V.S., Harmon A.C.& Rao K.S. (2000) "Spatio-temporal accumulation and activity of calciumdependent 

protein kinases during embryogenesis, seed development, and germination in 

sandalwood." Plant Physiol. 122(4), 1035-43. Abstract. Western-blot analysis and protein kinase 

assays identified two Ca(2+)-dependent protein kinases (CDPKs) of 55 to 60 kD in soluble 

protein extracts of embryogenic cultures of sandalwood (Santalum album L.). However, these 

sandalwood CDPKs (swCDPKs) were absent in plantlets regenerated from somatic embryos. 

swCDPKs exhibited differential expression (monitored at the level of the protein) and activity in 

different developmental stages. Zygotic embryos, seedlings, and endosperm showed high 

accumulation of swCDPK, but the enzyme was not detected in the soluble proteins of shoots and 

flowers. swCDPK exhibited a temporal pattern of expression in endosperm, showing high 

accumulation and activity in mature fruit and germinating stages; the enzyme was localized 

strongly in the storage bodies of the endosperm cells. The study also reports for the first time to 

our knowledge a post-translational inhibition/inactivation of swCDPK in zygotic embryos during 

seed dormancy and early stages of germination. The temporal expression of swCDPK during 

somatic/zygotic embryogenesis, seed maturation, and germination suggests involvement of the 

enzyme in these developmental processes. 

Anil, V. S. &. Rao K..S. (2000). “Calcium-mediated signaling during sandalwood somatic 

embryogenesis. Role for exogenous calcium as second messenger.” Plant Physiology August 

2000 123(4), 1301-1311. Abstract. Calcium-dependent protein kinase (CDPK) is expressed in 

sandalwood (Santalum album L.) seeds under developmental regulation, and it is localized with 

spherical storage organelles in the endosperm [Anil et al. (2000) Plant Physiol. 122: 1035]. This 

study identifies these storage organelles as oil bodies. A 55 kDa protein associated with isolated 

oil bodies, showed Ca(2+)-dependent autophosphorylation and also cross-reacted with antisoybean 

CDPK. The CDPK activity detected in the oil body-protein fraction was calmodulinindependent 

and sensitive to W7 (N-(6-aminohexyl)-5-chloro-1-naphthalene sulfonamide) 

inhibition. Differences in Michaelis Menton kinetics, rate of histone phosphorylation and sensitivity 

to W7 inhibition between a soluble CDPK from embryos and the oil body-associated CDPK of 

endosperm suggest that these are tissue-specific isozymes. The association of CDPK with oil 

bodies of endosperm was found to show a temporal pattern during seed development. CDPK 

protein and activity, and the in vivo phosphorylation of Ser and Thr residues were detected 

strongly in the oil bodies of endosperm from maturing seed. Since oil body formation occurs 

during seed maturation, the observations indicate that CDPK and Ca(2+) may have a regulatory 

role during oil accumulation/oil body biogenesis. The detection of CDPK-protein and activity in oil 

bodies of groundnut, sesame, cotton, sunflower, soybean and safflower suggests the ubiquity of 

the association of CDPKs with oil bodies. 

Anil V.S. & Rao K.S. (2001) "Purification and characterization of a Ca(2+)-dependent protein 

kinase from sandalwood (Santalum album L.): evidence for Ca(2+)-induced conformational 

39

changes." Phytochemistry 58(2), 203-12. Abstract. An early development-specific soluble 55 kDa 

Ca(2+)-dependent protein kinase has been purified to homogeneity from sandalwood somatic 

embryos and biochemically characterized. The purified enzyme, swCDPK, resolved into a single 

band on 10% polyacrylamide gels, both under denaturing and non-denaturing conditions. 

swCDPK activity was strictly dependent on Ca(2+), K(0.5) (apparent binding constant) for Ca(2+)- 

activation of substrate phosphorylation activity being 0.7 microM and for autophosphorylation 

activity approximately 50 nM. Ca(2+)-dependence for activation, CaM-independence, inhibition by 

CaM-antagonist (IC(50) for W7=6 microM, for W5=46 microM) and cross-reaction with polyclonal 

antibodies directed against the CaM-like domain of soybean CDPK, confirmed the presence of an 

endogenous CaM-like domain in the purified enzyme. Kinetic studies revealed a K(m) value of 1.3 

mg/ml for histone III-S and a V(max) value of 0.1 nmol min(-1) mg(-1). The enzyme exhibited high 

specificity for ATP with a K(m) value of 10 nM. Titration with calcium resulted in the enhancement 

of intrinsic emission fluorescence of swCDPK and a shift in the lambda(max) emission from 

tryptophan residues. A reduction in the efficiency of non-radiative energy transfer from tyrosine to 

tryptophan residues was also observed. These are taken as evidence for the occurrence of 

Ca(2+)-induced conformational change in swCDPK. The emission spectral properties of swCDPK 

in conjunction with Ca(2+) levels required for autophosphorylation and substrate phosphorylation 

help understand mode of Ca(2+) activation of this enzyme. 

Anil V.S., Harmon A.C. & Rao K.S. (2003) "Temporal association of Ca(2+)-dependent protein 

kinase with oil bodies during seed development in Santalum album L.: its biochemical 

characterization and significance." Plant Cell Physiol. 44(4),367-76. Abstract. Calcium-dependent 

protein kinase (CDPK) is expressed in sandalwood (Santalum album L.) seeds under 

developmental regulation, and it is localized with spherical storage organelles in the endosperm 

[Anil et al. (2000) Plant Physiol. 122: 1035]. This study identifies these storage organelles as oil 

bodies. A 55 kDa protein associated with isolated oil bodies, showed Ca(2+)-dependent 

autophosphorylation and also cross-reacted with anti-soybean CDPK. The CDPK activity 

detected in the oil body-protein fraction was calmodulin-independent and sensitive to W7 (N-(6- 

aminohexyl)-5-chloro-1-naphthalene sulfonamide) inhibition. Differences in Michaelis Menton 

kinetics, rate of histone phosphorylation and sensitivity to W7 inhibition between a soluble CDPK 

from embryos and the oil body-associated CDPK of endosperm suggest that these are tissuespecific 

isozymes. The association of CDPK with oil bodies of endosperm was found to show a 

temporal pattern during seed development. CDPK protein and activity, and the in vivo 

phosphorylation of Ser and Thr residues were detected strongly in the oil bodies of endosperm 

from maturing seed. Since oil body formation occurs during seed maturation, the observations 

indicate that CDPK and Ca(2+) may have a regulatory role during oil accumulation/oil body 

biogenesis. The detection of CDPK-protein and activity in oil bodies of groundnut, sesame, 

cotton, sunflower, soybean and safflower suggests the ubiquity of the association of CDPKs with 

oil bodies. 

Anon (1999) “Tree farming: More on sandalwood cultivation: Economics of sandalwood 

plantations.” Agric. & Industry Survey, Mar/April pp30-32. 

Anon (1997) “Sandalwood oil crop suffers burn.” HerbalGram 41, 54. 

Anon (2004) “Squads to check Sandalwood smuggling” The Hindu 14.05.04 at: 

www.hinduonnet.com/thehindu/2002/05/ 14/stories/2002051403200400.htm 

Anon (2007) "Company News: Karnataka Soaps launches turmeric soap in Andhra." Focus on 

Surfacants 12 (Dec 2007), p7. Quote: “Karnataka Soaps also plans to increase its production of 

sandalwood soaps from 7200 tonnes in 2005-2006 to 8000 tonnes in 2007-2008.” 

Anon (2009) "Forest Dept to promote sandalwood cultivation." The Hindu 17th July 2009. 

Synopsis: Karnataka forest department has decided to promote sandalwood cultivation in the 

district by distributing around 50,000 seedlings this year.... 

40

Bagchi S. K. & Veerendra H.C.S. “Variation and relationship in developmental growth phases of 

Santalum album after pruning.” Indian Forester 117(12), 1053-1058. 

Bagchi S.K. & Sharma V.P. (1989) "Biometrical studies on seed characters of 

aaqswdeftghjukiolp L." Silvae Genetica 38(3-4), 152-153. Abstract. Les 

caractéristiques (longueur, largeur, poids) de graines de Santalum album, résoltées sur 10 arbres 

phénotypiquement supérieurs pour différentes localités. La variation entre les individus est 

significative au niveau de probabilité 0,001. Les caractéristiques de la graine sont fortement 

héritables et toutes les caractéristiques sont significativement corrélées. 

Bagchi S.K. & Kulkarni H.D. (1987). "A note on seedling abnormality frequency in the half-sib 

progenies of Santalum album." Indian Forest 113, 650-651. 

Balachandran N. & Kichenamourthy S. (2007) "Profile of natural stands of Santalum album L. in 

the Pondicherry region, India." Sandalwood Research Newsletter 22. Abstract. The enumeration 

of sandal and its habitats, soil type, girth, height, status of re-production and associated plants 

were studied within the Pondicherry region in India. A total of 463 mature sandal trees (girth at 

breast height (gbh) ≥ 10 cm) and 360 sapling trees (

Barrett, D. R. & Fox J.E.D. (1994). “Early growth of Santalum album in relation to shade.” 

Australian Journal of Botany 42(1), 83-93. 

Barrett D R & Fox J E D (1995) “Geographical distribution of Santalaceae and botanical 

characteristics of species in the genus Santalum.” In: Sandalwood Seed Nursery and Plantation 

Technology (eds L Gjerum, J E D Fox & L Ehrhart). FAO, Suva, Fiji. RAS/92/361. Field Document 

8, 3–23. 

Barrett D.R. & Fox J.E.D. (1997). “Santalum album: Kernel composition, morphological and 

nutrient characteristics of pre-parasitic seedlings under various nutrient regimes.” Annals of 

Botany 79(1), 59-66. 

Bapat V. A., Iyer R.K., et al. (1996). “Effect of cyanobacterial extract on somatic embryogenesis in 

tissue cultures of sandalwood (Santalum album).” Journal of Medicinal & Aromatic Plant Sciences 

18(1), 10-14. 

Bapat V.A. & Rao P.S. (1992). “Biotechnological approaches for sandalwood (Santalum album L.) 

micropropagation.” Indian Forester 118(1), 48-54. 

Bapat V.A. & Rao P.S. (1988). “Sandalwood plantlets from synthetic seeds.” Plant Cell Reports 

7(6), 434-436. 

Bapat V.A. & Rao P.S. (1984). “Regulatory factors for in vitro multiplication of sandalwood tree 

(Santalum album): 1. Shoot bud regeneration and somatic embryogenesis in hypocotyl cultures.” 

Proceedings Of The Indian Academy Of Sciences Plant Sciences 93(1), 19-28. 

Bapat V.A., Fulzele D.P., et al. (1990). “Production of sandalwood somatic embryos in 

bioreactors.” Current Science 59(15), 746-748. 

Baruah A. D. (1999).” The economics of sandal-oil.” The Indian Forester 125 (6), 640-643. 

Bhaskar, V. (1992). “”Pollination biology and fertilization in Santalum album L. (Santalaceae).” 

Flora Jena 187(1-2), 73-78. 

Bhaskar V. & Swami Rao N. (1999) “Vegetative and physico-anatomical traits and their relation 

heartwood content in small leaved and large leaved forms of sandal tree (Santalum album L.).” 

PAFAI Journal 1, 33-38. 

Bhatnagar S.P. (1965) Studies in Angiospermic Parasites (No 2) – Santalum album – The 

sandalwood tree. Pub. National Botanical Gardens, Lucknow, India 1965, 

Bhatnagar S.P. (1959). “Some observa-tions on the post-fertilization devel-opment of the embryo 

sac of Santalum.” Phytomorphology 9, 87-91. 

Bhattacharya A. & Lakshmi S.G. (1999). “Isolation and characterization of PR1 homolog from the 

genomic DNA of sandalwood (Santalum album L.).” Current Science 77(7), 958-961. Abstract. 

Genomic library was constructed using nuclear DNA prepared from tender leaves of sandalwood. 

Subsequently, screening with heterologous probes we could isolate the PR1 genomic hemolog, 

Restriction mapping and hybridization experiments were carried out to obtain the coding region 

for PR1 gene. A 750 bp EcoRI fragment thus obtained was subcloned to yield pSaPR1, which 

was compared with the related sequences. Southern hybridization with genomic DNA digests was 

carried out to check its genomic organization. The induction of this gene was observed in the 

somatic embryos treated with salicylic acid, thereby implying its possible involvement during 

systemic acquired resistance. 

Bhattacharya, A. & Sita G.L. (1998). “cDNA cloning and characterization of a proline (or 

hydroxyproline)-rich protein from Santalum album L.” Current Science 75(7), 697-701. 

42

Bieri S., Monastyrskaia K. & Schilling B. (2004) "Olfactory receptor neuron profiling using 

sandalwood odourants" Chem Senses 29(6), 483-487. Abstract: The mammalian olfactory 

system can discriminate between volatile molecules with subtle differences in their molecular 

structures. Efforts in synthetic chemistry have delivered a myriad of smelling compounds of 

different qualities as well as many molecules with very similar olfactive properties. One important 

class of molecules in the fragrance industry are sandalwood odorants. Sandalwood oil and four 

synthetic sandalwood molecules were selected to study the activation profile of endogenous 

olfactory receptors when exposed to compounds from the same odorant family. Dissociated rat 

olfactory receptor neurons were exposed to the sandalwood molecules and the receptor 

activation studied by monitoring fluxes in the internal calcium concentration. Olfactory receptor 

neurons were identified that were specifically stimulated by sandalwood compounds. These 

neurons expressed olfactory receptors that can discriminate between sandalwood odorants with 

slight differences in their molecular structures. This is the first study in which an important class of 

perfume compounds was analyzed for its ability to activate endogenous olfactory receptors in 

olfactory receptor neurons. 

Bock J. (2003) “Sandalwood oil’s effect on the autonomic nervous system” Original Internist 

3/1/2003. Abstract: The hypothesis is sandalwood oil causes a decrease in sympathetic tone as 

accessed by patients with Heart Rate Variation (HRV), blood pressure (systolic, diastolic, and 

pulse pressure) and pulse rate. 

Brand, J.E. 1994. “Genotypic variation in Santalum album.” Sandalwood Research Newsletter 2, 

2-4. 

Brummitt, R.K. (1992). “Santalaceae.” pp 659–660. In: Brummitt, R.K. Vascular plant Families 

and genera. Royal Botanic Gardens, Kew, UK. 

Burdock GA & Carabin IG (2008) "Safety assessment of sandalwood oil (Santalum album L.)." 

Food Chem Toxicol. 46(2), 421-32. Abstract. Sandalwood (Santalum album L.) is a fragrant wood 

from which oil is derived for use in food and cosmetics. Sandalwood oil is used in the food 

industry as a flavor ingredient with a daily consumption of 0.0074 mg/kg. Over 100 constituents 

have been identified in sandalwood oil with the major constituent being alpha-santalol. 

Sandalwood oil and its major constituent have low acute oral and dermal toxicity in laboratory 

animals. Sandalwood oil was not mutagenic in spore Rec assay and was found to have 

anticarcinogenic, antiviral and bactericidal activity. Occasional cases of irritation or sensitization 

reactions to sandalwood oil in humans are reported in the literature. Although the available 

information on toxicity of sandalwood oil is limited, it has a long history of oral use without any 

reported adverse effects and is considered safe at present use levels. 

Castro J.M., Linares-Palomino P.J., Salido S., Altarejos J., Nogueras & Sanchez A. (2005) 

"Enantiospecific synthesis, separation & olfactory evaluation of all diastereomers of a 

homologuee of the sandalwood odournt Polysantol." Tetrahedron 61(47), 11192-11203. Abstract. 

The four stereoisomers of (5E)-4,4-dimethyl-6-(2′,2′,3′-trimethylcyclopent-3′-en-1′-yl)-hex-5-en-3- 

ol, a homologue of the valuable sandalwood-type odorant Polysantol®, were enantiospecifically 

synthesized from (+)- and (−)-α-pinene, through (−)- and (+)-campholenic aldehyde, by aldol 

condensation with 3-pentanone, deconjugative α-methylation and reduction. The mixtures of 

epimeric alcohols obtained after reduction were separated by means of derivatization with (−)- 

(1S)-camphanic chloride. The enantiomerically pure final products were evaluated 

organoleptically. 

Chabra S. K. & Rao A.R. (1993). “Postnatal modulation of xenobiotic metabolizing enzymes in 

liver of mouse pups following translactational exposure to sandalwood oil.” Nutrition Research 

13(10), 1191-1202. 

Chana J.S. (1994) “Sandalwood Production.” Internat. J. Aromatherapy 6(4), 11-13. 

43

Chandrashekharaiah A.M. & Dahgar V.M. (1998) “The effect of sandalwood availability on the 

craftsman community.” In: Sandal & its products. ACIAR Proceedings (84) eds. ACIAR 

Proceedings (84) eds. A.M. Radomiljac & R.H. Aanathapadmanabha, Welbourn R.M. and 

Satyanarayana Rao, Publication – Australian Centre for International Agricultural Research, 

Canberra 19-21. (1998). 

Chandrashekharaiah A.M. (1971) “Sandal tree”. My Forest 8, 21-25. 

Chapuis C. (2004) "In the quest for a virtual pseudo receptor for sandalwood-like odorants. Part I: 

The empirical approach." Chem Biodivers.1(7):980-1021. Abstract. Based on similarities between 

naturally occurring (-)-(Z)-β-- or (+)-(Z)-α-santalol ((-)- 1 or (+)-2, resp.) and the reversed (E)- 

configured synthetic derivatives from campholenal (7a), a simple model A was developed. 

Besides reconciliation of this stereochemical aspect, this initial model also tentatively explained 

the enantiodiscriminations as well as the large spectra of distances separating the OH function 

from the lipophilic quaternary center(s) reported for different classes of substrates. Evolution, 

modifications, and refinement of this imperfect model allied with the research for alternative 

possibilities are illustrated, along with a historical guideline, in the light of olfactively challenging 

synthetic seco-substructures as well as literature reports. Despite evolution of the inadequate 

model A and a plausible interpretation of the lipophilic part, the topological positions of the OH 

function and its vicinal alkyl substituent could nevertheless not be fully ascertained by this 

approach. This apparently inconclusive empirical concept prompted us to turn our attention 

towards a computerized methodology, which will constitute the second and third part of this study. 

Chawla G. (2008) ““The demise of India’s supply: resorting to substitutes to meet demand.” 

Sandalwood Conference 2008 at The Kimberley Grande, Kununurra, W. Australia 13-15 May 

2008. Cropwatch Comments: Chawla illustrates the demise of EI sandalwood’s decline via the 

annual production figure for sandalwwod sales in Tamilnadu, which peaked at 2330.5 tons in 

1999-2000, against just 14 tons in 2007-8. Also of interest were facts about the selection of 79 or 

more sandalwwod trees maintained at the IWST germplasm bank in Gottipura and about the 

clonal seed orchards from these trees maintained at Nalla Nallal and Jarakabande and at the 

Andhra Pradesh Forest Department Research Center, BIOTRIM, at Tirupathi. . 

Christenson P.A., Secord N. & Willis B.J. (1981) Phytochemistry 20(5), 1139-1141. Abstract: An 

analysis of East Indian sandalwood oil (Santalum album) has resulted in the isolation and 

identification of trans-β-santalol and epi-cis-β- santalol. 

Choudhuri J.C.B. (1963) "Sandalwood tree & its diseases." Indian Forester 89(7), 456-462. 

Choueiri A. (2008) “Sustainable ingredients in the fragrance industry and the use of Indian 

Sandalwood in L’Oreal products”. Sandalwood Conference 2008 at The Kimberley Grande, 

Kununurra, W. Australia 13-15 May 2008 Cropwatch Comments: Drawing on data from 

Edwards M. Fragrances of the World, Choueiri (Head of Lancome UK) includes the point that of 

106 current fragrances listing sandalwood, only 36 detail Indian sandalwood, and of those, only 

16 detail Mysore sandalwood (the rest we assume use sandalwood from other sources). Of these 

sixteen current fragrances allegedly employing Mysore sandalwood, four are supplied by IFF, two 

by Givaudin (Quest), one by Firmenich and one by Symrise. The present situation of shortage 

seems a far cry from the original launching of three fragrances containing authentic sandalwood 

by the antique pharmacy of Santa Maria Novella, Florence in 1828. 

Das S., Das S., Pal S., Mujib A., Sahoo S. S., Dey S., Ponde N. R. & Dasgupta S.(1999). “A 

novel process for rapidmass propagation of Santalum album L. in liquid media and bioreactors.” 

In: Giberti, G. (Ed) Proc. WOCMAP-2. Acta Hort. 502(ISHS), 281-288. 

Das S., Ray S., Dey S. & Dasgupta S. (2001). "Optimisation of sucrose, inorganic nitrogen and 

abscisic acid levels for Santalum album L. somatic embryoproduc tion in suspension culture.” 

Process Biochemistry (details) 

44

Desai V. B., Hiremath R.D., et al. (1991). “On the pharmacological screening of HESP and 

sandalwood oils.” Indian Perfumer 35(2), 69-70. 

Desai V.B. & Shankaranarayana K.H. (1990) “On utilization aspects of sandal seed oil.” Res 

Industry 35, 232-233. 

Dey S. (2002) “Mass cloning of Santalum album L. through somatic embryogenesis: scale up in 

bioreactor.” Sandalwood Research Newsletter (Australia), 13, 1-3. Abstract. Santalum album L., 

the East Indian Sandalwood, enjoys acceptance worldwide because of the uniquefragrance in it’s 

oil and wood. The natural propagation of this important plant faces twin threats –spike disease 

and poaching. Regeneration by silvicultural methods being insufficient to meet the demand, 

several biotechnological routes of propagation has been tried. Somatic embryogenesis offers 

highest clonal propagation efficiency. Scale up in air-lift bioreactor improves embryo quality, 

saves laboratory space and minimizes incubation time as well as production cost. 

Dijkstra J. & Hiruki C. (1974) “A histochemical study on sandal (Santalum album) affected with 

spike disease and its diagnostic value.” Netherlands J. of Plant Pathology 80(2), 37-47. 

Dijkstra J. & Lee P.E. (1972) “Transmission by dodder of sandal spike disease and the 

accompanying mycoplasma-like organisms via Vinca rosea.” Netherlands J. of Plant Pathology 

78(5), 218-224. 

Doran J.C., Thomson L.A.C. &. Brophy J.J.(2002). “Sandalwood.” Paper to Regional Workshop 

on Sandalwood Research, Development and Extension in the Pacific Islands and Asia. Noumea, 

New Caledonia, 7–11 October 2002. 

Erligmann A. (2001) “Sandalwood oils” Int. J. of Aromatherapy 11(4), 186-192. 

European Patent EP1059086 “Use of sandal wood oil or constituents of sandal wood oil for the 

prevention and treatment of warts, skin blemishes and other viral-induced tumors.” Abstract The 

present invention provides a method for the prevention and treatment of viral-induced tumors, 

more specifically, human warts. The method uses sandalwood oil and/or derivatives from the 

sandalwood oil to prepare medicaments for the prevention and treatment of viral-induced tumors 

(i.e., warts caused by the human papillomavirus (HPV)) in humans. The method of the invention 

comprises the topical administration of the sandalwood oil or a composition derived therefrom to 

the human epidermis and/or to the genital tract as needed. The present invention is also 

concerned with a unique antiviral composition useful for topical application. The antiviral 

composition according to this invention is also effective against other DNA viruses such as the 

DNA pox virus that causes Molluscum contagiosum and may be effective against other DNA 

viruses such as AIDS virus and RNA viruses. The sandalwood oil compositions are also effective 

against genital warts and HPV of the genital tract and will prevent cancer of the skin and cervix. 

Sandalwood oil or a constituent of sandalwood oil, is also effective in preventing dryness of the 

skin, rashes and flakiness associated with seborrheic dermatitis, psoriasis and allergic or 

eczematous rashes of the skin. This oil or constituent is also effective in the treatment of acne 

lesions of the face and the body and in the eradication of pustular acne lesions caused by 

Staphylococcal acne and Streptococcal bacterial infections. 

Fernandes P. C., Bapat V.A. et al. (1992). “In vivo germination of encapsulated somatic embryos 

of Santalum album L. (Sandalwood).” Indian Journal of Experimental Biology 30(9), 839-841. 

Fernandes, P., Bapat V.A., et al. (1994). “Effect of crushed seed homogenate on germination of 

synthetic seeds of Santalum album L.” Indian Journal of Experimental Biology 32(11), 840-841. 

Fernandes P. C., Bapat V.A., et al. (1994). “Investigations on the development of somatic seeds 

of Santalum album L. (Sandalwood).” Proceedings of the National Academy of Sciences India 

Section B Biological Sciences 64(1), 1-8. 

45

Florento A. (1997) “Sandalwood oil faces trouble as crop is destroyed by fire” Chemical Marketing 

Reporter March 31, 1997. 

Fox J.E.D.. (2000) “Sandalwood: the royal tree” Biologist London 47(1), 31-34. Abstract. 

Sandalwood is the most valuable tree in the world. As with gold, platinum and diamonds, it owes 

its value to a demand based on ritual, fashion and scarcity. It is the stuff of mystery and intrigue, 

and fortunes can still be made from it. 

Fragrance raw materials monographs (1974): “Sandalwood oil, East Indian.” Food & Cosmetics 

Toxicology 12(7-8), 989-990. 

Gjerum L., Fox, J.E.D. & Ehrhart L., (eds) (1995) Sandalwood Seed Nursery and Plantation 

Technology FAO, Suva. RAS/92/2361. Field Document No.8. 

Gleason J. (2009) “Comparing notes: Formulating with coumarin, sandalwood & ethyl linalool: an 

extended conversation with fine fragrance perfumers Kevin Verspoor & Pierre Gueros.”” Perf & 

Flav 34, April 2008, pp26-29. Cropwatch comments: In spite of the promising title, one short 

section on sandalwood tells you littlethat you probably didn’t know already, namely that 

sandalwood oil E.I. can be used in almost every type of perfume, making it full and long-lasting, 

and it is difficult to overdose.”. 

Griffith W (1836). "On the ovulum of Santalum album." Transactions ofthe Linnean Society of 

London (Botany) 18, 59-70. 

Haffner D. (1993). “Determining heartwood formation within Santalum album and Santalum 

spicatum.” Sandalwood Research Newsletter 1, 4–5. 

Hart H.H. & Schoemfelt F. (1956) “Some seed fats of Santalaceae & Oleaceae.” Aust J Chem 12, 

190. 

Haque M.H. & Haque A.U. (date) United States Patent 6406706; EP Patent 1,059,086, 2000 

"Use of α- and β-santalols major constituents of sandal wood oil, in the treatment of warts, skin 

blemishes and other viral-induced tumors." Abstract. The present invention provides a method for 

the treatment of viral-induced tumors in mammals, more specifically, human warts. The method 

uses α- and β-santalols, or mixtures or derivatives thereof, to prepare medicaments for the 

treatment of viral-induced tumors i.e., warts caused by the human papillomavirus (HPV) in 

humans. The method of the invention comprises the topical administration of α- and β-santalols, 

or mixtures or derivatives thereof, in a composition derived therefrom, to the human epidermis, as 

needed. The present invention is also concerned with a unique antiviral composition useful for 

topical application. The antiviral composition according to this invention is also effective against 

other DNA viruses such as the DNA pox virus that causes Molluscum contagiosum and may be 

effective against other DNA viruses such as AIDS virus and RNA viruses. The α- and β-santalols 

composition, or mixtures or derivatives thereof, may also be effective in the treatment of genital 

warts and HPV of the genital tract and in the treatment of cancer of the skin and cervix. The α- 

and β-santalols, or mixtures or derivatives thereof, may also be effective in the prevention of 

dryness of the skin, rashes and flakiness associated with seborrheic dermatitis, psoriasis and 

allergic or eczematous rashes of the skin. The α- and β-santalols, or mixtures or derivatives 

thereof, may also be effective in the treatment of acne lesions of the face and the body and in the 

eradication of pustular acne lesions caused by staphylococcal acne and streptococcal bacterial 

infections. 

Henfrey A (1856). "On the develop-ment of Santalum album." Transac-tions of the Linnean 

Society of Lon-don (Botany) 22, 69-79. 

Heuberger E., Hongratanaworakit T. & Buchbauer G. (2001) “Biological properties of the 

essential oil of East Indian sandalwood (Santalum album L.) and its main compounds alpha- and 

beta-santalol.” Oral presentation 4éme Symposium Internat. D’Aromatherapie Scientifique, March 

2001, Grasse, France. 

46

Heuberger E., Hongratanaworakit T., Buchbauer G. (2001) “Die Wirkung von Sandelholzöl auf 

das autonome Nervensystem und die subjective Befindlichkeit.” Vortrag 3, Internat. Primavera 

Life-Kongress Oct 2001. 

Heuberger E, Hongratanaworakit T, Buchbauer G. (2006) "East Indian Sandalwood and alphasantalol 

odor increase physiological and self-rated arousal in humans. Planta Med. 72(9), 792- 

800. Abstract. In Ayurvedic medicine, East Indian Sandalwood is an important remedy for the 

treatment of both somatic and mental disorders. In this investigation, the effects of inhalation of 

East Indian Sandalwood essential oil and its main compound, alpha-santalol, on human 

physiological parameters (blood oxygen saturation, respiration rate, eye-blink rate, pulse rate, 

skin conductance, skin temperature, surface electromyogram, and blood pressure) and selfratings 

of arousal (alertness, attentiveness, calmness, mood, relaxation and vigor) were studied 

in healthy volunteers. Compared to either an odorless placebo or alpha-santalol, Sandalwood oil 

elevated pulse rate, skin conductance level, and systolic blood pressure. alpha-Santalol, 

however, elicited higher ratings of attentiveness and mood than did Sandalwood oil or the 

placebo. Correlation analyses revealed that these effects are mainly due to perceived odor 

quality. The results suggest a relation between differences in perceived odor quality and 

differences in arousal level. 

Hill R., Harne R.W. & Nayar R.M. (1969) “Mycoplasma-like bodies associated with sandal spike” 

Nature 224, 1121-1122. 

Hongratanaworakit T. Dissertation, Effects of fractions of sandalwood oil on human physiological 

parameters by inhalation and massage. Vienna, Austria. June 2001. (through Bock J. (2003). 

Hongratanaworakit T., Heuberger E., Buchbauer G. (2000). “Effects of sandalwood oil & alphasantalol 

on humans I: Inhalation.” Poster presentation, 31st ISEO, Sept 2000, Hamburg, 

Germany. 

Hongratanaworakit T, Heuberger E. & Buchbauer G. (2004) “Evaluation of the effects of East 

Indian sandalwood oil and alpha-santalol on humans after transdermal absorption.” Planta Med. 

70(1),3-7. Abstract. The aim of the study was to investigate the effects of East Indian sandalwood 

oil (Santalum album, Santalaceae) and alpha-santalol on physiological parameters as well as on 

mental and emotional conditions in healthy human subjects after transdermal absorption. In order 

to exclude any olfactory stimulation, the inhalation of the fragrances was prevented by breathing 

masks. Eight physiological parameters, i. e., blood oxygen saturation, blood pressure, breathing 

rate, eye-blink rate, pulse rate, skin conductance, skin temperature, and surface electromyogram 

were recorded. Subjective mental and emotional condition was assessed by means of rating 

scales. While alpha-santalol caused significant physiological changes which are interpreted in 

terms of a relaxing/sedative effect, sandalwood oil provoked physiological deactivation but 

behavioral activation. These findings are likely to represent an uncoupling of physiological and 

behavioral arousal processes by sandalwood oil. 

Husain M., Ponnuswamy A.M. & Ponnuswamy P.K. (1982) “An innovation in the vegetative 

propagation of sandal (Santalum album L.).” Indian J. Forest. 5, 1-7. 

ISO 3518 Oil of Sandalwood (Santalum album L.) 2nd edn 2002. pub. ISO. Geneva 2002. 

Iyengar G.S. (1937). "Life-history of Santalum album L." Journal of Indian Botanical Society 16, 

175-195 

Iyengar A.V.V. (1968). "The East Indian sandalwood oil." Indian Forester 57, 57-68 

Iyengar A.V.V. (1972) “Control of sandal spike” Current Science 41(9), 318-319. 

Iyenger A.V.V. (1972) "Some aspects of sandal spike disease" J. Scient. Ind. Res. 31, 331-342. 

47

Jain S.H. Angadi V.G., Shankaranarayana K.H. & Ravikumar G. (2003) "Relationship between 

girth and percentage of oil in trees of sandal (Santalum album L.) provenances." Sandalwood 

Research Newsletter 18. Abstract. In three provenance areas of sandal viz. Bangalore, Thangli 

(Karnataka) and Maryoor (Kerala), studies have been made in respect of GBH and oil. It was 

observed that percentage of oil remains nearly constant at 4 % after 80 cm girth and that rise in 

oil percentage beyond 80 cm girth was found to be just marginal. 

Jain S.H., Angadi V.G., Ravikumar G., Thegrajan K.S. & Shankaranarayana (1999) “Studies on 

cultivation & chemical utilisation of sandal (Santalum album L.).” PAFAI Journal 1, 49-53. 

Jain S.H., Angadi V.G., Rajeevalochan A.N., Shankaranarayana K.H., Theagarajan K.S. & 

Rangaswamy C.R. (1998) "Identification of provenances of Sandal in India for genetic 

conservation" ACIAR Proceedings, No.84, 1998, 117-120. 

Jain S.H. Augundi V.G. Rajeevalochan K.H., Shankaranarayana K.H., Theagarajan K.S. & 

Rangaswamy C.R (1992) “Identification of provenances of sandal in India for genetic 

conservation.” In: Sandal & Its Products ACIAR Proceedings 84, 117-120. . 

Jain S.H. & Rangaswamy C.R. (1998). "Soil properties and their relationship to the growth of 

Sandal (Santalum album L) in three study areas in Karnataka." My Forest 24, 141-146. 

Jirovetz L, Buchbauer G, & Jager W. (1992) “Analysis of fragrance compounds in blood samples 

of mice by gas chromatography, mass spectrometry, GC/FTIR and GC/AES after inhalation of 

sandalwood oil.” Biomed. Chromatography 6, 133-134. Abstract. After inhalation experiments with 

sandalwood oil and the pure fragrance compounds coumarin and alpha-terpineol, substances 

were detected and measured in the blood samples of test animals (mice) using gas 

chromatography/mass spectrometry (GC/MS) (MID) in connection with GC/FTIR (SWC), GC/AES 

(carbon and oxygen trace) and flame ionization detection/gas chromatography. Using tiglinic acid 

benzyl ester as the internal standard the following concentrations in serum could be found: alphasantalol 

6.1 ng/mL, beta-santalol 5.3 ng/mL and alpha-santalene 0.5 ng/mL. In separate 

inhalation experiments with coumarin and with alpha-terpineol the corresponding concentrations 

were 7.7 ng/mL and 6.9 ng/mL, respectively. 

Jones C.G., Ghisalberti E.L., Plummer J.A. & Barbour E.L. (2006) "Quantitative co-occurrence of 

sesquiterpenes; a tool for elucidating their biosynthesis in Indian sandalwood, Santalum album." 

Phytochemistry. 67(22), 2463-8. Abstract. A chemotaxonomic approach was used to investigate 

biosynthetic relationships between heartwood sesquiterpenes in Indian sandalwood, Santalum 

album L. Strong, linear relationships exist between four structural classes of sesquiterpenes; 

alpha- and beta-santalenes and bergamotene; gamma- and beta-curcumene; beta-bisabolene 

and alpha-bisabolol and four unidentified sesquiterpenes. All samples within the heartwood 

yielded the same co-occurrence patterns, however wood from young trees tended to be more 

variable. It is proposed that the biosynthesis of each structural class of sesquiterpene in 

sandalwood oil is linked through common carbocation intermediates. Lack of co-occurrence 

between each structural class suggests that four separate cyclase enzymes may be operative. 

The biosynthesis of sandalwood oil sesquiterpenes is discussed with respect to these cooccurrence 

patterns. Extractable oil yield was correlated to heartwood content of each wood core 

and the oil composition did not vary significantly throughout the tree. 

Jones C.G., Keeling C.I., Ghisalberti E.L., Barbour E.L., Plummer J.A., Bohlmann J.(2008) 

"Isolation of cDNAs and functional characterisation of two multi-product terpene synthase 

enzymes from sandalwood, Santalum album L." Arch Biochem Biophys. 2008 May 27. Abstract. 

Sandalwood, Santalum album (Santalaceae) is a small hemi-parasitic tropical tree of great 

economic value. Sandalwood timber contains resins and essential oils, particularly the santalols, 

santalenes and dozens of other minor sesquiterpenoids. These sesquiterpenoids provide the 

unique sandalwood fragrance. The research described in this paper set out to identify genes 

involved in essential oil biosynthesis, particularly terpene synthases (TPS) in S. album, with the 

long-term aim of better understanding heartwood oil production. Degenerate TPS primers 

48

amplified two genomic TPS fragments from S. album, one of which enabled the isolation of two 

TPS cDNAs, SamonoTPS1 (1731bp) and SasesquiTPS1 (1680bp). Both translated protein 

sequences shared highest similarity with known TPS from grapevine (Vitis vinifera). Heterologous 

expression in Escherichia coli produced catalytically active proteins. SamonoTPS1 was identified 

as a monoterpene synthase which produced a mixture of (+)-alpha-terpineol and (-)-limonene, 

along with small quantities of linalool, myrcene, (-)-alpha-pinene, (+)-sabinene and geraniol when 

assayed with geranyl diphosphate. Sesquiterpene synthase SasesquiTPS1 produced the 

monocyclic sesquiterpene alcohol germacrene D-4-ol and helminthogermacrene, when incubated 

with farnesyl diphosphate. Also present were alpha-bulnesene, gamma-muurolene, alpha- and 

beta-selinenes, as well as several other minor bicyclic compounds. Although these 

sesquiterpenes are present in only minute quantities in the distilled sandalwood oil, the genes 

and their encoded enzymes described here represent the first TPS isolated and characterised 

from a member of the Santalaceae plant family and they may enable the future discovery of 

additional TPS genes in sandalwood. 

Jones C.G (2008) The best of Santalum album: Essential oil composition, biosynthesis and 

genetic diversity in the Australian tropical sandalwood collection. PhD thesis for Univ W Australia 

June 2008. Abstract. An investigation into the heartwood and essential oil content of Australian 

plantation sandalwood, Santalum album was undertaken. Genetic diversity of 233 S. album five 

S. austrocaledonicum and fifteen S. macgregorii trees growing in the Forest Products Arboretum, 

Kununurra WA, was assessed using nuclear and chloroplast RFLPs. Nuclear genetic diversity of 

the S. album collection was very low with expected and observed heterozygosity levels of 0.047. 

This was lower than the results previously reported in the bliterature for trees in India, however a 

different technique was used. Based on allelic patterns, the collection was able to be catagorised 

into 19 genotypes: each representing some shared genetic origin. Some groups were highly 

redundant, with 56 trees being represented, whilst others were populated by just one tree. The 

essential oil yield and heartwood contents of trees from these genetic groups were compared. 

Yields were highly variable both within and between groups of trees which share a common 

genetic history, suggesting a significant environmental component was contributing to the 

observed phenotype despite identical soil and climatic oconditions. 

Ancestral lineages were tested using chloroplast RFLP’s, although a lack of shared mutations 

between species made this difficult. Only one S. album tree from S. Timor was resolved using 

nuclear RFLPs, with the other trees being grouped with material sourced from India. There was 

no resolution of Indian S. album from Timorese using chloroplast RFLPs, however one S. album 

tree grown from Indian seed possessed a single unique mutation. The low genetic diversity of the 

Ausatralian S. album collection is likely to be a combination of incomplete seed sourcing and 

highly restricted gene flow during the evolution of the species. Combined with information 

gathered on the phylogeny of the genus by other researchers, S album is postulated to have 

arisen from an over-sea dispersal out of northern Australia or Papua New Guinea 3 to 5 million 

years ago. 

Essential oil yield & composition was assessed for 100 S. album trees growing in the collection, 

ranging in years from 8 to 17 years. Oil content of the heartwood ranged from 30mg.g -1 to 

60mg.g -1 and the transition zone 36mg.g -1 to 90mg.g -1 . Sapwood contained almost no 

sesquiterpene oils. Despite the highly variable total yields, the chemical profile of the oil did not 

vary, suggesting there was limited genetic diversity within this region of the genome. Strong 

positive correlations existed between sesquiterpenoids in the essential oil of S. album. This was 

particularly evident in the santalenes and α-bergamotene, although trends were also seen in the 

curcumenes, bisabolene and bisabolol, and an unidentified group of sesquiterpenes. Cooccurrence 

patternes indicate shared chemical intermediates from which compounds are 

partitioned, resulting in multiple product formation. To further test the multiple product hypothesis 

two terpene synthase (TPS) genes werte isolated from S. album leaf and wood cDNA. These 

were cloned and the enzymes were heterologously expressed in Escherichia coli. One enzyme 

sasesquiTPS1 converted the universal sesquiterpene precursor farnesyl diphosphate into 

germacrene D-4-ol, helminthogermacrene, α-bulnescene, γ-muurolene and α- andβ-selinene. 

The other enzyme samonoTPS1 converted geranyl diphosphate into (+)-α-terpineol, and (-)- 

limonene, with small amounts of sabinene, linalool, α-terpinolene, myrcene and geraniol. These 

49

epresent the first TPS genes to be isolated from sandalwood and will enable further elucidation 

of the oil biosynthesis genes. 

This thesis `completes a three-pronged approach to understanding the underlying causes of oil 

yield variance in S. album. As a species for which so little is known, the research presented here 

provides a major leap forward for tree improvement, breeding and silviculture. Hence the best of 

sandalwood research is presented. 

Jones C. (date) "Indian Sandalwood: Genetic and oil diversity and bio-chemistry of the 

Australian germplasm collection." Sandalwood Research Letter 

Jyothi, P. V., J. B. Atluri, et al. (1991). “Pollination ecology of Santalum album (Santalaceae).” 

Tropical Ecology 32(1), 98-104. 

Kababick J. P. (1996). “Evaluation on incense purity using simultaneous steam distillationextraction 

and HRGC analysis.” Journal of High Resolution Chromatography 19(4), 241-242. 

Kaikini N.S. (1969) “The Indian sandal wood.” My Forest 25-40 (1969). 

Kapoor M.L. (1981) “A technique for chromosome count from developing leaf buds of induced 

tetraploids & diploid Santalum album L.” Indian Forester 107, 290-300. 

Kawanishi Y., Nin K. & Toyoda K. (2004) “The influence on the autonomic nervous system of a 

preference for the smell of sandalwood” Aroma Research 5(4), 382-385. 

Khan M.M.., Faroqui A.A., Vasundhara M. & Srinisappa K.N. (1999) “Clonal propagation of 

sandalwood (Santalum album Linn.) PAFAI Journal 1, 20-24. 

Kulkami H.D. & Srithmathi R.A. (1981) “Polyembryony in the genus Santalum L.” Indian Forester 

107(11), 704-706. 

Kushalappa K.A. (1999) “Trade liberalization in sandalwood.” Indian Forester 125(9), 891-894. 

Madhi A. (1986) "The biology of S. album seed." Biotrop. Tech. Bull. 1(1), 1-9. 

Mathur, N.K. 1979. An annotated bibliography of spike disease of sandal (Santalum album Linn.) 

Forestry Research Institute. Dehra Dun. 74 p. 

Mayar R. (1988) “Cultivation Information, Exploitation & Protection of Santalum album.” Advances 

in Forestry research in India Vol II p117-152. 

Meera C., Nageswara Rao M., Ganeshaiah K.N., Uma Shankaar R. (2000) “Conservation of 

sanadal genetic resources I. Extraction patterns and threats to sandal resources in Karnataka.” 

My Forest 36(2), 125-132. 

Merlin M. & Van Ravenswaay D. “The history of human impact on the genus Santalum in 

Hawai’i.” Paper presented at the Symposium on Sandalwood in the Pacific, 1990. Abstract. 

Adaptive radiation of Santalum in the Hawaiian archipelago has provided these remote islands 

with a number of endemic species and varieties. The prehistoric Polynesian inhabitants of Hawai‘i 

utilized the sandalwood trees for many of the same traditional purposes as their South Pacific 

ancestors who had developed ethnobotanical relationships with Santalum. The ancient Hawaiians 

probably reduced the number and geographical distribution of sandal-wood trees 

significantly through their extensive cutting and burning, especially in the dry forest regions. 

Nevertheless, vast numbers of the fragrant trees still existed in Hawai‘i at the time of Western 

contact in 1778. Within a century after this contact, the extensive trade in sandalwood produced a 

massive decline in the Hawaiian species of Santalum. Although cultivation attempts during this 

cen-tury with both introduced and native sandalwood species have had limited success in 

Hawai‘i, there is renewed interest in developing a sustainable forest industry based on the 

production of sandalwood for export trade. Biologists in general, however, have cautioned against 

50

large-scale harvesting of the remain-ing Santalum trees, suggesting that more research be 

undertaken first to deter-mine the distribution and vigor of the remaining species. 

Mitchell J.F.M. (1941) Sandalwood problems, factors factors affecting heartwood and oil content 

in sandalwood. Proc 5 th Silvi Conf, Dehra Dun 1941. 

Mookherjee B., Kamath V., Patel R. & Shuster E. (of International Flavours & Fragrances Ltd.) 

(1976) US Patent 4,000.050. 

Mookherjee B., Kamath V. & Shuster E. (of International Flavours & Fragrances Ltd.) (1977) US 

Patent 4,014,823. 

Morris E.T. (1983) “The Fascinating History of Sandalwood” Dragoco Report 44-51.(1983). 

Mujib A. (2005) “In vitro regeneration of sandal (Santalum album L.) from leaves.” Turk J. Bot. 29, 

63-67. 

Nageswara-Rao M., Ganeshaiah K.N & Uma Shankaar R. (2008) “Fading fragrance.” Decan 

Herald May 6 th, 2008. . 

Nageswara-Rao M., Padmini S., Ganeshaiah K.N., Shaanker R.U. & Soneji J.R. (2008) “Materials 

Study: Indian sandalwood crisis.” Perfumer & Flavorist 33 (Oct 2008), 38-43. Cropwatch 

comments:: Excellent article on the decline in the supply of East Indian sandalwood:due to overexploitation 

of the dwindling resource. The authour notes most of the former primary centres for 

sandalwood carving have now closed down and the sandalwood oil factory at Shimoga is now 

closed, just leaving the Mysore factory in the Karnataka district. The author also comments on the 

loss of sandalwood genetic resources.and remarks on the fact that the loss of natural genetic 

variability has been a major handicap in formulating conservation plans. Suggestions are made 

for a way forward however. 

Nageswara Rao M., Ganeshaiah K.N., & Shaanker R.U. (2007) “Assessing threats and mapping 

sandal (Santalum album) resources in peninsula India. Identification of genetic hot-spot for in-situ 

conservation.” Conservation Genet 8, 925-935. 

Nageswara Rao M. (2007) Mapping genetic diversity of sandal (Santalum album L.) genetic 

resources in peninsular India using biochemical and molecular markers. Lessons for in-situ 

conservation. PhD Thesis, Forest Research Institute (FRI) ICFRE Dehra Duhn, India 2004. 

Nageswara Rao M., Ganeshaiah K.N., & Shnkar R.U. (2001) “Mapping genetic diversity of sandal 

(Santalum album L.) in South India. Lessons for in-situ conservation of sandalwood resources.” 

In:: Forest Genetic Resources Status, Threats and Conservation Strategies.eds. R Uman 

Shankaar, KN Ganeshaiah & KS Bawa. Oxford & IBH Pakistan Publications, New Delhi 49-67 

(2001). 

Nageswara Rao M., Padmini S. Ganeshaiah K.N. & Uma Shankaar R. “Sandal genetic resources 

of South India. Threats and conservation approaches.” In: National symposium on role of plant 

tissue culture in biodiversity, conservation and economic development, 63, Kuri-Katarmal. Almora 

U.P. (1999). 

Nagaveni H.C. & Srimathi R.A. (1980) "Use of giberellic acid to assist germination of sandal 

seeds." Indian Forester 106(11), 792-799. 

Nagaveni H.C. & Srimathi R.H. (1981) “Studies on germination of sandal seeds Santalum album 

L. Linn. II” Indian Forester 106, 792-799. 

Nagaveni. H.C. & Vijayalakshmi G. (2003) Sandalwood Research Newsletter 18. "Growth 

performance of sandal (Santalum album L.) with different host species." Abstract. Santalum 

album L (Sandal plant) is a partial root parasite on several host plants.It shows a preference to 

51

certain host species and grows well. In the presentstudy, Pongamia pinnata and Casuarina 

equisetifolia supported the sandal plants, yielding robust growth, where as some hosts like 

Artocarpus integrifolia, Acacia auriculiformis and Swietenia mahogany hindered the growth of 

sandal. Understanding the dynamics of parasitism may help in raising successful multi species 

plantations of sandal along with other valuable timber species. 

Narayana K.H.S. & Parthasarathi K. (1986) "HESP - a new essential oil from the acid hydrolysis 

of spent sandal heartwood." Perf. Flav. 10(6), 60-1. 

Naqvi A.A., Singh A.K. & Mandal S. (1999) “Quality evaluation of sandalwood (Santalum album) 

oil grown in northern parts of India. Indian Perfumer 43, 67-69. 

Nasi, R., and Y. Ehrhart. (1996). “Le Santal, un parfum de prospérité. 1ère partie–une longue 

histoire.” Bois et Forêts des Tropiques 247, 5–19. 

Nasi R. & Ehrhart Y. (1996) “Sandalwood, a perfume of prosperity” Part 2 Plantations.” Bois-et- 

Forets-des-Tropiques 248, 5-20. 

Nayar R. (1981) “Interrelations between mycoplasma-like organisms in spiked sandal (Santalum 

album L.) & infected collateral hosts” Eur. J. For. Pathol. 2, 29-32. 

Nayar, R. (1984). “Investigations with sandalwood (Santalum album) Mycoplasma and toxins.” 

European Journal Of Forest Pathology 14(1), 59-64. 

Nayar R. (1988) "Cultivation, improvement, exploitation & protection of Santalum album Linn." 

Advances in Forestry Research in India 2, 117-151. 

Ngo K-S. & Brown G. D. (2000) "Autoxidation of alpha-santalene." Journal of Chemical Research 

2000 (2). 68-70(3) Abstract. Fifteen compounds (2 - 11) have been isolated from the 

spontaneous slow autoxidation of the tri-substituted double bond in the side-chain of the tricyclic 

sesquiterpene -santalene; most of these compounds have also been reported as natural 

products. 

Ohmori A., Shinomiya K., Utsu Y., Tokunaga S., Hasegawa Y. & Kamei C. (2007) "[Effect of 

santalol on the sleep-wake cycle in sleep-disturbed rats]" Nihon Shinkei Seishin Yakurigaku 

Zasshi. 27(4),167-71. Abstract. Sandalwood oil is widely used in aromatherapy for alleviating 

various symptoms. Santalol, a major component of sandalwood oil, has been reported to have 

central nervous system depressant effects such as sedation. In the present study, we 

investigated the effect of santalol on the sleep-wake cycle in sleep-disturbed rats. When inhaled 

at a concentration of 5 X 10(-2) ppm, santalol caused a significant decrease in total waking time 

and an increase in total non-rapid eye movement (NREM) sleep time. In order to clarify the 

mechanism of action, olfactory hypofunction was caused in rats by intranasal application of 5% 

zinc sulfate solution, and thereafter the effects of inhalation of fragrances were evaluated. In this 

study, it was found that the impairment of the olfactory system showed no significant effect on the 

changes in sleep parameters induced by santalol. This result suggests that santalol may act via 

the circulatory system rather than the olfactory system. That is, santalol is thought to be absorbed 

into the blood through the respiratory mucosa, and then exert its action. From these results, it is 

concluded that santalol may be useful in patients having difficulty maintaining sleep without being 

affected by individual differences in perfume-related preference. 

Okugawa H., Ueda R., Matsumoto K., Kawanishi K. & Kato A. (1995). “Effect of a-santalol and b- 

santalol from sandalwood on the central nervous system in mice.” Phytomed 2 (2), 119-126. 

Padmanabha A. (200 “Indian Sandalwood –the history, the uses and the future of supply” 

Sandalwood Conference 2008 at The Kimberley Grande, Kununurra, W. Australia 13-15 May 

2008 

Parry E.J. Sandalwood Oil Pub. Govt. of Mysore (date unknown). 

52

Parthasarathi K., Rangaswamy C.R. & Anguah V.C. (1985) "Leaf peroxidase, malate 

dehydrogenase & ertrase isoenzyme pattern in ten sandalwood (Santalum album) types showing 

variation in leasf pattern." Indian Forester 111, 441-449. 

Parthasarathi K., Rangaswamy C.R., Jayaraman K & Rao P.S (1973) “Studies on sandal spike. 

Part X. Deoxyribonuclease & ribonuclease activities and nucleic acid levels in sandal (Santalum 

album Linn.) affected by spike mycoplasma.” Proceedings of the Indian Academy of Sciences 

77(3), 131-136. 

Parthasarathi K. et al. (1973) "Hosts of sandalwood." Current Science 43(1), 20. 

Parthasarathi K., Gupta S.K. & Rao P.S. (1974). "Differential response in the cationexchange 

capacity of the hostplants on parasitization on sandal (Santalum album)." Curr.Sci. 43, 20 

Parthasarathi K. & Venkatesan K.R. (1982) "Sandal spike disease" Current Science 51(5), 225- 

230. 

Parthasarathi K., Angadi V.G.,Shankaranarayana K.H. & Ra-jeevalochan, A.N. (1986). "Peroxidase 

isoenzyme activity in the livingbask tissue as a marker for the oil-bearing capacity in 

Sandal." Current Science 55, 831-34. 

Parthasarathi, K; Rai, S.N. (1989). “Physiology, chemistry and utilization of sandal (Santalum 

album Linn.).” My Forest 25(2): 181-219. 

Prakash NA, Farooqi AA & Vasundhara M. (1999) “Studies on root parasitism and nutrition of 

sandalwood (Santalum album L.) PAFAI Journal 1, 25-29. 

Preen C. (2005) "Update on Sandalwood Essential Oil" Aromatherapy Regulation News Summer 

2005 Newsletter 2(2), 4. Aug 2005. Cropwatch comments: Further proof, if it were needed, that 

some Aromatherapy (AT) organisation officials have been 'in denial' about the role of 

aromatherapy as a consumer market affecting the serious demise of sandalwood species - here 

Preen attempted to shift the blame to the perfumery trade. Further, Preen argued that Santalum 

album is not actually endangered (the IUCN Red List 

http://www.iucnredlist.org/search/details.php/31852/all classifies it as vulnerable), wrings her 

hands a little about sandalwood smuggling, and pledges faith in the much-criticised sandalwood 

replanting schemes to eventually solve the problem 30-40 years hence. Cropwatch currently 

observes that little, if any, Sandalwood oil East Indian is currently available on the spot market, 

and what there is, is practically always adulterated - it is of note that the AT professional 

organisations have no formal analytical standards for sandalwood oil used in aromatherapy to 

determine whether or not this is the case. Further, the carbon footprint of sandalwood oil is 

particularly unacceptable wrt climate change concerns, with excessive energy consumption 

occurring as a result of long distillation times. Therefore by continuing to defend the use of E.I. 

sandalwood oil in AT, one can only conclude that any ecological interests have been 

inappropriately sold out to the commercial interests of AT essential oil traders, who anyway have 

an unhealthy & unseen influence on the policy of many AT professional organisations. Further, 

aromatherapists are likely to be indirectly supporting gangland by consuming sandalwood oil, 

most of which is either smuggled with or without the help of corrupt officials or otherwise illegally 

produced. This was a completely wrong-headed & misleadfing article (in Cropwatch's humble 

opinion, of course).. 

Radomiljac, A. M., J. A. McComb, et al. (1998). “Xylem transfer of organic solutes in Santalum 

album L. (Indian sandalwood) in association with legume and non-legume hosts.” Annals of 

Botany 82(5), 675-682. 

Radomiljac, A.M., McComb J.A., Shea S.R. (1998) “Field establishment of Santalum album – the 

effect of the time of the introduction of a pot host (Alternanthia nana R.Br.)” Forest Ecology & 

Management 111 (2-3), 107-118. Abstract: Field establishment of the root hemi-parasite 

53

Santalum album L. under large-scale plantation conditions, until recently, has been largely 

unsuccessful. In this experiment, the growth of S. album seedlings grown with the herbaceous pot 

host Alternanthera nana R. Br. for 134, 109, 84, 60 and 35 days in a nursery container prior to 

field establishment was examined after 11, 16 and 23 weeks in the field. S. album survival and 

growth was greater, and root:shoot ratio was lower for the 23 weeks for S. album seedlings grown 

with A. nana compared with seedlings grown without a host. Seedlings grown with A. nana for 

134 days in the nursery prior to field establishment had greater stem diameter, height and root, 

shoot and total plant dry weight (DW) over the 23 weeks in the field than all other treatments. 

Seedlings grown with A. nana for 109 days in the nursery prior to field establishment had greater 

field survival than all other treatments. A. nana survival in the field remained high when grown 

with S. album for 134 and 109 days in the nursery prior to field establishment whereas survival 

within remaining treatments declined significantly and A. nana growth was significantly less. S. 

album grown with A. nana for 134 days in the nursery prior to field establishment had a lower 

root:shoot ratio than all other treatments at all assessments. A strong negative linear relationship 

exists between S. album root:shoot ratio and A. nana DW, whereas a positive linear relationship 

exists between S. album DW and A. nana DW. Foliar phosphorus and sodium concentrations for 

S. album were lower and foliar potassium concentration higher when seedlings were grown with 

A. nana for 134 days in the nursery prior to field establishment compared with the remaining 

treatments at the 16-week assessment. The period of the S. album:-A. nana association in the 

nursery significantly influenced S. album survival and growth following field planting. 

Radomiljac A. M. & McComb J.A. et al. (1999). “Intermediate host influences on the root hemiparasite 

Santalum album L. biomass partitioning.” Forest Ecology and Management 113(2-3), 

143-153. Abstract. Santalum album L. seedlings parasitised on the N2-fixing woody hosts 

Sesbania formosa (F. Muell.) N. Burb., Acacia trachycarpa E. Pritzel and A. ampliceps Maslin and 

the non-N2-fixing woody host Eucalyptus camaldulensis Dehnh. were grown for 38 weeks in 25 l 

nursery containers. S. album growth was greater and root : shoot ratio lower for S. album 

seedlings grown with N2-fixing hosts compared with seedlings grown with E. camaldulensis or 

with no host. Seedlings grown with S. formosa had a greater stem diameter, height, leaf area, 

root and shoot dry weight (DW) than all other treatments. S. album grown with S. formosa and A. 

ampliceps had a lower root : shoot ratio than all other treatments at all assessments. The root : 

shoot ratio of unattached S. album increased exponentially over the 38 week period. Seedling 

growth declined for all treatments between the 33 and 38 week harvests, except for those 

seedlings attached to A. trachycarpa. A strong positive linear relationship was shown between S. 

album leaf area and shoot DW irrespective of host species. No relationship was found between S. 

album shoot DW and root : shoot ratio with host shoot DW. The combined E. camaldulensis and 

S. album root system supported a smaller S. album shoot biomass compared with the S. album 

shoot biomass supported by the combined root systems of the N2-fixing hosts and S. album. 

Compared with all other host species the A. trachycarpa root system was more efficient in 

supporting its own shoot biomass and the total biomass of S. album. Host use efficiency (S. 

album shoot DW/host shoot DW) values of S. formosa and A. trachycarpa were greater than the 

host use efficiency values of A. ampliceps and E. camaldulensis. Values for unparasitised S. 

formosa leaf area, shoot and root DW and root : shoot ratio were greater than those for 

parasitised plants. 

Radomiljac A. M. & McComb J.A. et al. (1999). “Gas exchange and water relations of the root 

hemi-parasite Santalum album L. in association with legume and non-legume hosts.” Annals of 

Botany 83(3), 215-224. 

Radomiljac A.M., Ananthapadmanabho H.S., Welbourne R.M & Satyanarayan Rao, K.(eds), 

(1999). “Sandal and its Products”. Proceedings of an International Seminar held on 18-19 

December 1997 organised by the Institute of Wood Science and Technology (ICFRE) and 

Karnataka State Forest Department, Bangalore India. Canberra: ACIAR Proceedings No 84. 

Radomiljac, A.M., McComb, J.A. & Pate, J.S. (1999). “Organic solute transport and assimilation in 

Santalum album L. (Indian sandalwood): intermediate host partnerships involving beneficial and 

non-beneficial hosts.” Annals of Botany 83, 215-224. 

54

Rai V.R. & McComb J. (2002) “Direct somatic embryogenesis from mature embryos of 

sandalwood.” Plant Cell Tiss. Org. Culture 69, 65-70. 

Rai S.N. & Sarma C.R. (1990) "Depleting sandalwood production and rising prices. Indian 

Forester 116, 348-355. 

Ral S N (1990) “Status and cultivation of sandalwood in India.” In: Proceedings of the Symposium 

on Sandalwood in the Pacific (eds L Hamilton. & C E Conrad). General Technical Report PSW – 

122. USDA Forest Service, Berkeley, 66–71. Abstract: Sandalwood (Santalum album) has been 

part of Indian culture and heritage for thousands of years, and was one of the first items traded 

with other countries. The heartwood yields fragrant oil, which is used mainly in the perfume 

industry but also has medicinal properties. The wood is used for carving and manufacturing 

incense. Generally S. album is found in the dry deciduous forests of Deccan Plateau, mostly in 

the states of Karnataka and Tamil Nadu, The evergreen tree regenerates naturally when 

conditions are favorable and has been spreading in its distribution. Lack of understanding of the 

dynamics of hemiparasitism by sandalwood has caused failure of pure plantations in the past; 

haustorial connections with its hosts supply sandalwood with nitrogen, phos-phorus, and 

potassium. Plantable seedlings can now be raised in the nursery in 6-8 months with the 

protection of a nematicide and fungicide. Several tech-niques for planting seeds directly in the 

field have also been developed. A tree that is growing well can put on an annual increment of 1 

kg per year. The sandalwood resource in India is currently threatened by four factors: fire, 

browsing by livestock, spike (little leaf) disease, and smuggling. 

Rao M.R. (1911) “Host plants of the sandal tree.” Indian Forest Records. 2(4), 159-207. 

Rao P.S. & Ozias-Akins P. (1985) "Plant regeneration through somatic embryogenesis in 

protoplast cultures of sandalwood (Santalum album L.)." Protoplasma 124(1-2), 80-86. Abstract. 

Protoplasts were isolated from embryogenic cell suspension cultures derived from proliferating 

shoot segments of a 20-year-old sandalwood tree (Santalum album Linn.). Under appropriate 

conditions, isolated protoplasts divided in liquid culture medium and produced embryogenic cell 

aggregates and globular embryos. Plating of cell aggregates on a fresh medium facilitated the 

differentiation of somatic emryos which further developed into plantlets. 

Ramanathan C. (1997). “Indian Sandalwood Trade.” In: TED Case Studies: Sandalwood Case. 

www.american.edu/projects/mandala/TED/sandalwood.htm 

Rangaswamy N. S. & Rao P.S.(1963). "Experimental studies on Santalum album L.: 

Establishment of tissue culture of endosperm." Phytomorphology 13, 450-454. 

Rangaswamy, C.R., Jain S.H. & Parthasarathy K. (1986). "Soil proper-ties of some Sandal 

bearing areas." Van Vigyan 24 (3&4), 61-68. 

Rangaswamy K.T. & Jayarajan R. (1998) “The distribution of spike disease in the forests of Tamil 

Nadu.” Current Research University of Agricultural Sciences Bangalore 27(4), 76-77. 

Rao P.S., Chrungoo N.K., et al. (1996). “Characterization of somatic embryogenesis in 

sandalwood (Santalum album L.). In vitro cellular and developmental biology” Plant 32(3),123- 

128. 

Rao P.S. (1987) “Clonal multiplication of plants of economic value: sandalwood, mulberry and oil 

palm.”. In Proceedings of Workshop on Increasing Crop Productivity, Bombay, 20-21 June, 1986. 

New Delhi: Oxford and IBH. pp. 225-229. 

Rao P.S. & Sharma C.L. (1986) “Relationship between height & diameter increment of sandal 

Santalum album L.” Van Vigyan 24, 105-138. 

55

Rao P.S. & Ozias-Atkins P. (1985) "Plant regeneration through somatic embryogenesis in 

protoplast cultures of sandalwood.” Protoplasma 124(1-2), 80-86. 

Rao P.S., Bapat V.A., et al. (1984). “Regulatory factors for in vitro multiplication of sandalwood 

tree (Santalum album): 2. Plant regeneration in nodal and internodal stem explants and 

occurrence of somaclonal variations in tissue culture raised plants.” Proceedings of the Indian 

National Science Academy Part B: Biological Sciences 50(2), 196-202. 

Rao P.S. & Srimati R..A. (1977) “Vegetative propagation of sandal (Santalum album L.). Current 

Science 46, 276. . 

Remadevi, O. K., Muthukrishnan R., et al. (1997). “Epidemic outbreak of lac insect, Kerria lacca 

(Kerr.), on Santalum album (Sandal) and its control.” Indian Forester 123(2), 143-147. 

Ross M.S. (1983). Bibliography on Sandalwood, Santalum album. Unpublished - copy held at 

University of Oxford, UK (although see Ross (1985). Cropwatch comments: A unique 

bibliography of nearly 800 references & papers, many of them reviewed (& much of the subject 

matter concerning splike disease) drawn from references within the former Commonwealth 

Forestry Institute at Oxford, and from other sources 

Ross M.S. (1985). Annotated bibliography on sandalwood, Santalum album, and its uses. Bos- 

Document 2. Dutch Forestry Development Cooperation, Wageningen, The Netherlands. 135pp 

Roychoudhuri S.P. & Verma A. (1980) “Sandalwood spike.” PAFAI Journal 1, 25-29. 

Sahai A.. & Shivanna K.R. (1984). “Seed germination, seedling growth and haustorial induction in 

Santalum album, a semi-root parasite.” Proceedings ff the Indian Academy of Sciences: Plant 

Sciences 93(5), 571-580. 

Sanjaya B.M., Thrilok S.R. & Vittal R.R. (2006) "Micropropagation of an endangered Indian 

sandalwood (Santalum album L.). J. of Forest Research 11(3), 203-209. Abstract Santalum 

album is known as East Indian sandalwood. It is the most economically important tree harvested 

for heartwood oil, and India is among the chief exporters of sandalwood and its products. Multiple 

shoots were induced from nodal shoot segments derived from a 50- to 60-year-old candidate plus 

tree (CPT) on Murashige and Skoog (MS) medium supplemented with 0.53 µM α- 

naphthaleneacetic acid (NAA) and 11.09 µM 6-benzylaminopurine (BA). In vitro differentiated 

shoots were multiplied on MS medium with 0.53 µM NAA, 4.44 µM BA, and additives: 

283.93 µM ascorbic acid, 118.10 µM citric acid, 104.04 µM cystine, 342.24 µM glutamine, and 

10% (v/v) coconut milk. New shoots were harvested repeatedly for up to three subculture 

passages on fresh medium at 4-week intervals. Microshoots treated with 98.4 µM indole-3- 

butyric acid (IBA) for 48 h produced roots on growth-regulator-free, quarter-strength MS basal 

salts medium with vitamin B5 and 2% sucrose. In vitro root induction was achieved from 

microshoots pulsed with 1230 µM IBA for 30 min in soilrite rooting medium. The percentage of 

rooting in soilrite was higher than that for agar medium, and in vitro raised plants were 

established in the field and showed normal growth. 

Sanjaya B.M. Anathapadmanabha & Rai V.S. (2003) “In vitro and in vivo micrografting of 

Santalum album shoot tips.” J. Trop. Forest Sci. 15, 234-236. 

Scartezzini P. & Speroni E. (2000). “Review on some plants of Indian traditional medicine with 

antioxidant activity.” J. of Ethnopharm. 71(1-2), 23-43. 

Scheffel M. (1990) “Sandalwood: Current Interest and Activity by the Hawaii Division of Forestry 

and Wildlife.” Proceedings of the Symposium in the Pacific 1990 Abstract: The State of Hawaii 

Department of Land & Natural Resources (DLNR) protects native species growing on State land, 

but has no official program funding for growing sandalwood. Part of the DLNR, the Division of 

Forestry and Wildlife forest and nursery managers maintain exuberant activity in attempting to 

56

establish their nursery stock of sandalwood in the field out of personal interest. Nursery and 

planting techniques are described. 

Scott J. (1871) "Root parasitism by sandalwood." J. Agric. Hort. Soc. India 2,287. 

Sekhar A. R. C & Vinaya Rai R. S. (2000) “Production and export of selected non-wood forest 

products in India in 2005.” Journal of Tropical Forest Products 6 (1) 1-11 

Sen Sarma P.K. (1982) “Sandalwood – its cultivation & utilisation.” In Cultivation and Utilisation of 

Medicinal & Aromatic Plants (eds C.K. Atal & B.M. Kapur). Regional Research Laboratory, 

Jammi–Tawi pp 395-405. 

Setzer W.N. (2009) "Essential oils & anxiolytic aromatherapy." Nat Prod Commun. 4(9), 1305-16. 

Abstract. A number of essential oils are currently in use as aromatherapy agents to relieve 

anxiety, stress, and depression. Popular anxiolytic oils include lavender (Lavandula angustifolia), 

rose (Rosa damascena), orange (Citrus sinensis), bergamot (Citrus aurantium), lemon (Citrus 

limon), sandalwood (Santalum album), clary sage (Salvia sclarea), Roman chamomile (Anthemis 

nobilis), and rose-scented geranium (Pelargonium spp.). This review discusses the chemical 

constituents and CNS effects of these aromatherapeutic essential oils, as well as recent studies 

on additional essential oils with anxiolytic activities. 

Shankaranarayana K.H., Ravikumar G. Rajeevalochan A.N. , Theagarajan K.S. & Ramaswamy 

C.R. (1998) “Content & composition of oil from central and transition zones.” In: Sandal & Its 

Products. ACAIR Proceedings (84) eds A.M. Radomiljac H.S. Aanathapadmanabba, Wellburn R 

& Satyanarayana Rao. Publication Australian Centre for International Agricultural Research, 

Canberra 86-88 (1998). 

Shankaranarayana, K.H., Ravikumar G., Rajeevalochan A.N., Theagarajan K.S. & Rangaswamy 

C.R. (1998). “Content and composition of oil from the central and transition zones of sandalwood 

discs. ACIAR Proceedings 84, 86–88. 

Shankaranarayana, K. H., Angadi V.G., et al. (1997). “A rapid method of estimating essential oil 

content in heartwood of Santalum album Linn.” Current Science 72(4), 241-242. 

Shankaranarayana K.H. & Parthasarathi K. (1987) “On th content and composition of oil from 

heartwood at different levels in sandal.” Indian-Parfum Kanpur Essential Oil Association of India 

28, 138-141. 

Shankaranarayana K.H. & Parthasarathi K. (1984). “Compositional differences in sandal 

(Santalum album) oils from young and mature trees and in the sandal oils undergoing color 

change on standing.” Indian Perfumer 28(3-4), 138-141. 

Shankaranarayana, K. H. & Venkatesan KR..(1982) “Chemical aspects of sandalwood oil.” In: 

Cultivation and utilization of aromatic plants. CK Atakal & RPL Kapoor (CSIR) Jammu 406-411. 

Sheen J. & Stevens J. (2001) “Self-perceived effects of Sandalwood” Intl J. of Aromatherapy 

11(4), 213-219. Abstract: Eight female participants used the essential oil of Santalum album, East 

Indian Sandalwood, as a perfume daily for a week. Their self-perceived effects were analyzed for 

common experiences, using the grounded theory method. Four categories of the experience were 

developed into an initial theory of the effects of sandalwood. It was found that sandalwood did 

have self-perceived effects, which varied with initial psychological state and emporal factors. The 

observed self-perceived effects of calming, ability to manage and well being have limited corelation 

with claimed therapeutic effects. A further self-perceived effect, uplifting, was observed 

such that further investigation is required. This study is a demonstration of the initial steps of a 

holistic research model that would allow for aromatherapy, essential oils, their therapeutic effects 

and the experience of their use to be researched. Thus a sound scientific knowledge base for the 

profession of aromatherapy, relevant to its practice can be developed.Cropwatch comments: 

Recommended reading on self-perceived therapeutic effects of sandalwood! 

57

Shieh J. C., Lin T.S. et al. (1990). “Essential oil yield and component variation from Santalum 

album wood of different age in Taiwan.” Bulletin Of Taiwan Forestry Research Institute: New 

Series 5(1), 45-52. 

Shineberg D. (1967) They came for Sandalwood: A study of Sandalwood Trade in The SW 

Pacific 1830-1865.”Melborn University Press 1967. 

Shiri, V. & K. S. Rao K.S. (1998). “Introduction and expression of marker genes in sandalwood 

(Santalum album L.) following Agrobacterium-mediated transformation. Plant Science 131(1), 53- 

63. 

Shankaranarayana K.H. & Parthasarathi K. (1984) "Compositional differences in sandal oils from 

young and mature trees and in the sandal oils undergoing colour change on standing." Indian 

Perfumer 28(3/4), 138-141. 

Shankaranarayana, K. H. & K. Parthasarathi K.(1984) "Synthetic sandalwood aroma chemicals." 

Perf. & Flav. 9(1), 17-20. 

Shankaranarayana, K. H. & Kamala B.S. (1989) "Fragrant products from less obvious 

sandalwood oil" Perf. & Flav. 14(1), 19-21. 

Sindhuveerendra, H. C., S. Ramalakshmi, et al. (1999). Variation in seed characteristics in 

provenances of sandal (Santalum album L.). Indian Forester 125(3), 308-312. 

Singh P. (1911) “Memorandum on the oil value of sandalwoods from Madras. Forest Bull. 6.. 

Sita G.L. (1991) "Tissue cultured sandalwood." Current Science 61(12), 794-795. 

Sindhuveerendra HC & Sujatha M. (1989). "Pollination studies in Santalum album L." Current 

Science 58, 629-630. 

Sreenivasan V.V., Shivaramakrishnana C.R., Rangaaswamy H.S., Anathapadmanabba H.S. & 

Shankaranarayan K.H. (1992) Sandal ICFRE Dehra Dun, India 1992. 

Srimati E.A. Venkateshan K.R. & Kulkarni H.D. (1995) “Guidelines for selection & establishment 

of seedstands, seed production areas, plus trees & clonal seed orchards for Santalum album L.). 

In: Recent Advances in Research and Management of Sandal (Santalum album L.) in India. eds 

R.A. Srimati, K.R. Venkateshan & H.D. Kulkarni Associated Press, New Delhi 1995 pp281-299. 

Srinivasa I.G. (1937). "Life history of Santalum album." Journal of Indian Botanical Sciences 

16,175-196 

Srivinivisan V.V. Sivaramakrishnan V.R., Rangaswamy C.R., Ananathpodmanabha MS & 

Shankaranarayana (1995) “Sandal (Santalum album)”. Indian Council of Forestry Research & 

Education, Dehradun. 

Starke J.C. (1967) "Photoallergy to sandalwood oil" Arch Dermatol 96(1), 62-63. Abstract. A case 

of photocontact dermatitis to commercial sandalwood oil, an ingredient in many men's toiletries, is 

described. Patient reacted on skin testing to a broad spectrum of ultraviolet light. Dermatitis was 

persistent in spite of the absence of the original sensitizer and minimal erythemal dose was 

lowered, as in other "persistent light reactors." 

Struthers R.; Lamong B.B.; Fox J.E.D.; Wejesuriya S.R. & Crossland T. (1986). “Mineral nutrition 

of sandalwood (Santalum spicatum). Journal of Experimental Biology 37(182), 1274-1284. 

Abstract. Acacia acuminata is a preferred host of the root hemiparasitic tree, Santalum spicatum 

(sandalwood). Comparison between nutrient content of adult trees of sandalwood and results for 

an earlier study of the mistletoe, Amyema preissii, on the same host species, A. acuminata, 

showed similar high levels of K and Na and low levels of Zn in both parasites compared with the 

58

host plants. Differences in K, Ca, N and Cu levels between parasitized and uninfected Acacias 

imply that the host plant contributes to the nutrition of sandalwood. The high K/Ca ratio in 

sandalwood confirms that K uptake in preference to Ca is a general feature of all categories of 

angiosperm parasites. 

Patterns of distribution of nutrients between various parts of sandalwood and A. acuminata 

depend on the type of nutrient, but levels are usually highest in leaves of both species and the 

haustoria. Although K, Ca and Na are much lower in the kernels than in vegetative parts of the 

parasite, only seedlings without supplementary Ca in a nutrient omission experiment failed to 

grow at all in the absence of hosts. Growth is not dependent on the level of K in the unattached 

plants but other evidence indicates it may have a role in water uptake in the attached plant. 

Calcium supply has a marked effect on internal Ca levels and growth of unattached plants. 

Compared with field plants, levels of Ca, and to a lesser extent Zn, were much higher in plants of 

the Ca/K treatment that produced greatest growth over 34 weeks. 

Haustorial formation is enhanced by the presence of A. acuminata roots. However, competition 

for nutrients, especially Ca, from co-planted A. acuminata seedlings results in suppression of 

growth of young sandalwood compared with their growth in the absence of the host species. 

Subbarao, N. S., Yadav D., et al. (1990). “Nodule haustoria and microbial features of Cajanus 

and Pongamia parasitized by sandal (sandal wood).” Plant And Soil 128(2), 249-256. 

Suma T.B. & Balasundaran M. (2003). "Isozyme variation in fiveprovenances of Santalum album 

in India." Australian Journal of Botany 51, 243-249. 

Sunil T. & Balasundaran, M (2001) "Purification of sandal spike phytoplasma for the production of 

polyclonal antibody." Current Science Online 80(12), 1489-1494. Abstract. Sandal (Santalum 

album. L), a semiroot parasitic tree is the source of the East Indian sandalwood and oil. Spike 

disease caused by phytoplasma is the major disease of sandal. The disease is noticed in all 

major sandal-growing states of India. 

Suriamidhardja S. (1978) "Problems on Sandalwood (Santalum album Linn.) silviculture & 

improving its production." In: Proc of 3rd Seminar on Volatile Oils, Bogor, Indonesia, July 1978. 

Bogor: Balai Penelitian Kimia 

Surendran, C., Partiban, K.T., Bhuvenaswaran, C. and Murugesh, M. (1998) . “Silvicultural 

strategies for augmentation of sandal regeneration.” In: Radomiljac A.M., Ananthapathmanabha 

H.S., Welbourn R.M., Stayanarayan K. (eds.) Sandal and its products. ACIAR Proceedings 

Volume 84. Arawang Communications, Canberra. pp. 69-73. 

Swami R.N. & Srinivasa R.V. (1980) “Accelerated germination of sandal seeds (Santalum album 

L.) Lal Bagh J. 25, 68-69. 

Thomas, S. & Balasundaran M. (1999). “Detection of sandal spike phytoplasma by polymerase 

chain reaction.” Current Science 76(12), 1574-1576. 

Uma Shankaar R., Ganeshaiah K.N. & Nageswara Rao M. & Aravind N.A. (2004) “Ecological 

consequences of forest-use from genes to ecosystem: a case study in the Bilgiri Ranganswamy 

Temple Wildlife Sanctuary, South India.” In: Conservation Soc. 2, 347-363. 

Uma Shankaar R., Ganeshaiah K.N., & Nageswara Rao M. and Ravikanth G. (2002) “Forest 

gene banks – a new integrated approach for the conservation of forest tree genetic resources.” 

In: Managing Plant Genetic Diversity eds JMM Eugels, ADH Brown & MT Jackson. CAIBI 

Publishing Nosworthy, Wallington, Oxon UK 229-235. 

Uma Shankaar R., Ganeshaiah K.N. & Nageswara Rao M. (2000) “Conservation of sandal 

genetic resources in India: problems and psospects. IN; International Conference on Science & 

Technology for Managing Plant Genetic Diversity in 21 st Century, Kuala LAmpur, Malaysia 2000. 

59

Uniyal, D. P., Thapliyal R.C. et al. (1985). “Vegetative propagation of sandal (Santalum album) 

by root cuttings.” Indian Forester 111(3), 145-148. 

Valluri J.V. (2009) "Bioreactor Production of Secondary Metabolites from Cell Cultures of 

Periwinkle and Sandalwood" In S. Mohan Jain & Praveen K. Saxena (eds.), Methods in Molecular 

Biology, Protocols for In Vitro Cultures and Secondary Metabolite Analysis of Aromatic and 

Medicinal Plants, vol. 547 pub Humana Press (Springer Science + Business Media). Abstract. A 

bench-top bioreactor allowing continuous extraction of secondary metabolites is designed for 

Catharanthus roseus L. (G.) Don (periwinkle) and Santalum album L. (sandalwood) plant cell 

suspensions. Periwinkle cell cultures are exposed to biotic elicitors (Aspergillus niger, crude 

chitin) and abiotic elicitors (mannitol, methyl jasmonate) to induce alkaloid production. Whereas 

most of the biotic elicitors are effective when added on day 15 of culture, the abiotic elicitors are 

effective when added on day 20. The use of trans-cinnamic acid, an inhibitor of phenylalanine 

ammonia lyase (PAL) activity, results in significant increase in the alkaloid production of 

periwinkle cell cultures. Exposure of the cells to mannitol-induced osmotic stress produced 

marked increment in the total alkaloid production. When biotic and abiotic stress treatments are 

applied sequentially, an additive effect in alkaloid accumulation is observed. Although no 

essential oils are detected, secondary metabolites in the form of phenolics are produced by the 

sandalwood cell cultures in the bioreactor environment. The use of morphologic modification such 

as organ cultures and transformed cultures is believed to be required for both production and 

storage of essential oil constituents in sandalwood. The present chapter demonstrates that 

periwinkle and sandalwood cell suspensions could be developed and successfully cultured in a 

modified air-lift bioreactor. The exploitation of variant cell strains and biotransformation of added 

precursors can certainly improve the use of periwinkle and sandalwood cell cultures for the 

bioproduction of desired compounds. 

Valluri, J. V., Treat W.J. et al. (1991). “Bioreactor culture of heterotrophic sandalwood (Santalum 

album L.) cell suspensions utilizing a cell-lift impeller.” Plant Cell Reports 10(6-7), 366-370. 

Valluri J.V., and D.B. Chambers (1992). "Protein synthesis in sandalwood callus cultures exposed 

to drought and salt stress." Plant Physiology and Biochemistry 99(5), 51 – 52. . 

Vaze Suresh (1999) “Sandal – introductory Note” PAFAI Journal 1, 17-19. 

Veakatesha Gowda V.S. Patil K.B. & Perumal I.R. (2006) “Forest based essential oils viz 

sandalwood oil production and future scenario.” Indian Perfumer 30, 45-50. 

Veakatesha Gowda V.S. Patil K.B. & Perumal I.R. (2006) “Forest based essential oils viz 

sandalwood oil production and future scenario.” PAFAI 8(1), 63-70. 

Veerendra, H. C. S. & Sarma C.R. (1990). “Variation studies in sandal (Santalum album L.): I. 

Time of emergence and seedling vigor.” Indian Forester 116(7), 568-571. 

Veerendra, H. C. S. & Padmanabha H.S.A. (1996). “The breeding system in sandal (Santalum 

album L.).” Silvae Genetica 45(4), 188-190. 

Venkata Rao, M.G. (1938). “The influ-ence of host plants on sandal andspike disease.” Ind. For. 

64(11) : 656-669. 

Venkatesha, M. G. & Gopinath K. (1994). “Description of immature stages of a species of gt 

lyptapanteles (Hymenoptera: Braconidae), a gregarious endoparasitoid of Amata passalis 

(Fabricius) (Lepidoptera: Arctiidae), a defoliator of sandalwood, Santalum album L. Insect 

Science and its Application 15(2), 161-165. 

Walker H. (1966). "The market for sandalwood oil." Tropical Product Institute, Ministry of 

Overseas Development. London. 

60

Wilson C.C. (1915) “Sandalwood (a) a parasite (b) susceptibility to fire (c) damage by borers (d) 

spike disease.” Science 188, 1081-1021. 

Yusuf, R. (1999). “Santalum album L.” pp 161–167 In: Oyen, L.P.A., and Nguyen Xuan Dund 

(eds.). Plant Resources of South-East Asia Vol 19. Essential-oil Plants. Prosea, Bogor, 

Indonesia. 

Indonesian Sandalwood (Santalum album L.). 

Anon (2003) “Police seize 13.6 tons of sandalwood” Jakarta Post 9.30.2003 KUPANG, East Nusa 

Tenggara: Abstract. 13,645 kilograms of sandalwood allegedly smuggled from neighboring East 

Timor was seized, supposedly being sent to sandalwood distilling firm PT Tropicana Oil. 

Husain A.M.M. (1983) “Report on the Rehabilitation of Sandalwood & the trade in Nasa Tenggara 

Timur Indonesia. World Bank PPIPD Project Report, West Timor. 

Omon, R. M. (1994). “The effects of N, P, K and NPK fertilizer on the growth of Cendana 

(Santalum album Linn.) in nursery of Latosol soil.” Buletin Penelitian Hutan (565), 55-64. Forest 

Research and Development Centre, Bogor 16001, Indonesia. 

Risseeuw P. (1950).”Sandelhout (Sandalwood).” In C.J.J. van Hall and C.van deKoppel, De 

Landbouw in de Indische Archipel (Agriculture in the Indonesian Archipelago). The Hague Vol 3 

pp686-705. 

Suriamihardia S. (1978) "[Problems on sandalwood (Santalum album Linn.) silviculture and 

improving its production]". pp. 115-125. In Proceedings of Third Seminar on Volatile Oils, Bogor, 

Indonesia, July, 1978. Bogor: Balai Penelitian Kimia. 

Walker H (1966) “The Market for Sandalwood Oil” TPI Rep. Trop. Prod. Inst. London (G22): 13. 

Widiadana, Rita A. (13.03.2002) “Illegal trade in endangered species on the rise in RI” Jakarta 

Post 13.03.2002. Abstract. Widiadana comments on the increasing trade in endangered species 

including trade in orchids and sandalwood. 

Yadav V.G. (1993) "Sandalwood: its origin, synthetic substitutes & structure-odour relationship." 

PAFAI Journal 15(4), 21-54. 

Yemris Fointuna (2004) “Myth about sandalwood broken” Jakarta Post 9.8.2004. Abstract. 

Sandalwood now only found on Timor and Sumba islands. According to the East Nusa Tenggara 

Statistics Office, the number of sandalwood trees is estimated to be only some 100,000 due to 

illegal logging. 

Ogasawara Island Sandalwood (Santalum boninensis). 

Maina S. L., Pray L. A. & Defilipps R. A. 1988. “A historical note on the endangered Santalum 

boninensis (Santalacaea) of the Ogasawara Islands: Early reports by Jahrasi Tuyoma.” Atoll 

Research Bulletin 319, 19-24. 

Tuyama, T. (1939). "On Santalum boninense, and the distribution of the species of Santalum." 

Jap. J. Bot. 15, 697-712. 

Pacific Sandalwoods. 

Cropwatch comments: Several Sandalwood spp are distributed throughout the 

Pacific including Santalum austrocaledonicum (Vanuatu & New Caledonia & 

Santalum yasi (Fiji). The Lush company of the UK publically own up to using 1 

ton per annum of New Caledonian sandalwood oil at 

http://www.lush.co.uk/Shop/FeatureDetail.aspxfdShopFeatureId=6888 

61

Cook Islands 

Sykes W.R. (1980).” Sandalwood in the Cook Islands.” Pacific Science 34(l), 77-82. 

Fiji (Santalum yasi, Santalum album). 

Bulai P.B. (1995). “Sandalwood in Fiji.” In L Gerum, JED Fox and Y Ehrhart (eds.) Sandalwood 

seed, nursery and plantation technology. Proceedings of a regional workshop for Pacific Island 

Countries; August 1-11, 1994; Noumea, New Caledonia. RAS/92/361. Field Document No. 8. 

UNDP/FAO South Pacific Forestry Development Programme, Suva, Fiji. Pp 167-172. 

Bulai P. & Nataniela V. (2002). “Research, development and extension of Sandalwood in Fiji: A 

new Beginning.” In Proceedings of SPC Regional Workshop On Sandalwood Research, 

Development And Extension In The Pacific Islands And Asia. 7-11 October 2002, Noumea, New 

Caledonia. 

Jiko L.R. (2000) “Status & current interest in Sandalwood in Fiji” Sandalwood Research 

Newsletter 10, 1-3. Abstract. Santalum yasi, the only sandalwood species in Fiji, earned 4.74 

million Fijian dollars in foreign exchange over the period 1987-90. This recent resurgence in the 

utilisation of sandalwood has created an interest among landowners in planting and restocking 

new areas with the species. Information is presented on early history, traditional uses, growing 

conditions, silviculture and marketing of yasi, together with some notes on current research 

directions. 

Silaitoga S. (2008) “Sandalwood rip-off.” Fiji Times 17 th Feb 2008. Cropwatch comments: Article 

describes how traders in the North are being offered 50c per kilo (down from $1) by local buyers 

for Sandalwood yasi. The traders are members of mataqali Nakorovatu of Nabavatu Village in 

Dreketi, and they want the Ministry of Forestry to intervene. 

Smith R.M. & Morris P.R. (1979). Composition of Fijian sandalwood oil (Santalum yasi). 

International Flavour and Food Additives 10(2), 57. 

Tabunakawai, K. & A. Chang, (1984) "Sandalwood resource of the Ono-i-LauIslands. Suva, Fiji." 

unpublished reportof the Forestry Department. 

Usumaki, J.T. (1981). Sandalwood Survey Report—Bua Province. Unpublished report of Forestry 

Department. Suva, Fiji. 

Hawaii (Santalum haleakalae - Maui only, Santalum freycinetianum, Santalum. 

paniculatum & Santalum ellipticum). 

Le Barron, Russell K. (1970).”Hawaii's sandalwood. Aloha Aina.” Department of Land and Natural 

Resources, State of Hawaii; 6-7. 

Cartwright B. (1935). “Extinction of trees soon followed the Sandalwood Rush—Hawaii’s unhappy 

first export trade.” Paradise of the Pacific, Honolulu. 

Harada-Stone, D. (1988). “Sandalwood logging defended, ripped at special Kona hearing.” 

Hawaii Tribune-Herald, September 30, 1, 10. 

Judd C.S. (1933). "The parasite habit of the sandalwood tree." Thrum, Hawaiian Annual, 

Honolulu: The Printshop Co. Ltd. 59th Issue; 81-88. 

Judd C.S. (1935). "Reviving the sandalwood industry." Paradise of the Pacific. April 1935 p 19. 

Judd C.S. (1936) “Growing Sandalwood in the territory of Hawaii.” Journal of Forestry Vol XXXIV (1) 

1936. 

Kepler A.K. (1985). “Sandalwood: Hawaii’s precious ‘iliahi.” Mauian 2(6), 6–11. 

Lydgate J.M. (1916). “Sandalwood days.” Thrum's Hawaiian Annual. 

62

Merlin M. & VanRavenswaay D. (1990) "The History of Human Impact on the Genus Santalum in 

Hawaii." Proceedings of the Symposium on Sandalwood in the Pacific April 9-11, 1990, Honolulu, 

Hawaii. Abstract: Adaptive radiation of Santalum in the Hawaiian archipelago has provided these 

remote islands with a number of endemic species and varieties. The prehistoric Polynesian 

inhabitants of Hawai‘i utilized the sandalwood trees for many of the same traditional purposes as 

their South Pacific ancestors who had developed ethnobotanical relationships with Santalum. The 

ancient Ha-waiians probably reduced the number and geographical distribution of sandal-wood 

trees significantly through their extensive cutting and burning, especially in the dry forest regions. 

Nevertheless, vast numbers of the fragrant trees still existed in Hawai‘i at the time of Western 

contact in 1778. Within a century after this contact, the extensive trade in sandalwood produced a 

massive decline in the Hawaiian species of Santalum. Although cultivation attempts during this 

cen-tury with both introduced and native sandalwood species have had limited success in 

Hawai‘i, there is renewed interest in developing a sustainable forest industry based on the 

production of sandalwood for export trade. Biologists in general, however, have cautioned against 

large-scale harvesting of the remain-ing Santalum trees, suggesting that more research be 

undertaken first to determine the distribution & vigor of the remaining species. 

Rock J.F. (1916) "The sandalwoods of Hawaii. “A revision of the Hawaiian species of the genus 

Santalum." Hawaii Board Agric. Forest. Bot. Bull. 3, 1–43. 

St. John, H. (1947). “The History, Present Distribution, and Abundance of Sandalwood on O‘ahu, 

Hawaiian Islands.” Hawaiian Plant Studies 14, 1(1): 5–20. 

Scheffel M. (1990) "Sandalwood: Current Interest and Activity by the Hawaii Division of Forestry 

and Wildlife." Proceedings of the Symposium on Sandalwood in the Pacific April 9-11, 1990, 

Honolulu, Hawaii. Abstract: The State of Hawaii Department of Land & Natural Resources 

(DLNR) protects native species growing on State land, but has no official program funding for 

growing sandalwood. Part of the DLNR, the Division of Forestry and Wildlife forest and nursery 

managers maintain exuberant activity in attempting to establish their nursery stock of sandalwood 

in the field out of personal interest. Nursery and planting techniques are described. 

Stemmermann L. (1980). “Vegetative anatomy of the Hawaiian species of Santalum 

(Santalaceae).” Pacific Science 34(l):55-75 

Stemmermann R.L. (1980). “Observations of the Genus Santalum (Santalaceae)” in Hawaii 

Pacific Science 34(1), 41-54. Abstract. Some of the taxonomic problems of species of Santalum 

in Hawai'i are resolved by proposing one new taxon and two combinations. These are based on 

observations of the genus in the field, herbarium studies, and anatomical differences found. 

Stemmermann L. (1990) "Distribution and Status of Sandalwood in Hawaii." Proceedings of the 

Symposium on Sandalwood in the Pacific April 9-11, 1990, Honolulu, Hawaii. Abstract. This 

paper attempts to summarize what is known of the distribution and status of sandalwoods in 

Hawai‘i. Four species of sandalwood are recog-nized as being endemic to the Hawaiian Islands, 

and one has been introduced. Ecological factors affecting the present and former distribution of 

Hawaiian sandalwoods are considered. 

TenBruggencate J. (1988). “Private sandalwood logging has state upset”. Hono-lulu Star- 

Bulletin/Advertiser, September 27, A3. 

U.S. Fish & Wildlife Service (1985). Endangered and threatened wildlife and plants: proposed 

endangered status for Santalum freycinetianum Guad. var. lanaiense Rock (Lanai sandalwood or 

‘iliahi). Fed. Reg. 50(44): 9086-9089. 

Wagner J.P. (1986). “The rape of the fragrant trees.” Honolulu Magazine, November, 97 ff. 

63

Wilkinson K.M. (2007) "Propagation Protocol for 'iliahi (Santalum freycinetianum).” Native Plants 

Journal 8(3),248-251. Abstract. 'Iliahi or Hawaiian sandalwood (Santalum freycinetianum 

Gaudich. [Santalaceae]) is a hemiparasitic plant that can be readily grown in the nursery, 

provided some general guidelines are followed. Seeds germinate best if scarified and sown fresh. 

Plants can be grown to outplanting size (20 cm [8.0 in] tall with stems 8 mm [0.3 in] in diameter) 

in just 8 to 12 mo using controlled release fertilizer. The best survival and growth occurs when 

sandalwood is grown with a companion plant. Keywords sandalwood, nursery host plant, 

Santalaceae, Hawai'i Nomenclature USDA NRCS (2007) Click for larger view Jack Jeffrey 

inspects Santalum paniculatum tree on Mauna Kea. Photo by Craig Elevitch [Begin Page 250] 

liahi or Hawaiian sandalwood (Santalum freycinetianum Gaudich. [Santalaceae]) is endemic to 

the Hawaiian islands of O'ahu, Kaua'i, Lana'i, Maui, and Moloka'i. It is found in dry, mesic, and 

wet forest, with rainfall of 50 to 380 cm (20 to 150 in) and at elevations of 250 to 950 m (820 to 

3120 ft). It is a hemiparasitic plant, meaning its roots attach to the root systems of other plants to. 

Marquesas Islands (Santalum insulare, Santalum marchionense). 

Butaud J-F, Raharivelomanana P, Bianchini J-P & Baron V. (2003) “A new chemotype of 

Sandalwood (Santalum insulare Bertero ex A DC.) from Marquesas Islands” J. Essen. Oil Res. 

15, 323-6. Abstract. Volatile constituents of sandalwood (S. insulare) concrete from the island of 

Nuku-Hiva in Marquesas Islands were studied using GC, GC-MS, HPLC and NMR. The 

investigation of nine main compounds showed important variations among sandalwood samples 

(from 3.5 to 53.2% for α-santalol and from trace to 29.3% for (Z)-nuciferol). Statistical analysis put 

in relief a geographical segregation between sandalwoods growing in dry area in Terre-Déserte 

(14.6% of α- and β-santalol, 17.1% of (Z)-nuciferol and 11.7% of 6,13-dihydroxybisabola-2,10- 

diene) and sandalwoods growing in wetter area of the other parts of the island (60.9% of α- and 

β-santalol, 1.2% of (Z)-nuciferol and 0.7% of 6,13-dihydroxybisabola-2,10-diene). The chemotype 

rich in (Z)-nuciferol of Terre-Déserte constitutes a rare and new chemotype, which is described 

for the first time. 

Verhaegen D. (2000) “AMI in the Marquesas Islands: Sandal Preservation.” L’Ami Ingrediénts 

Naturels No 26, 1-2. 

New Caledonia (Santalum austrocaledonicum). 

Alpha T., Raharivelomanana P., Blanchini J.-P., Faure R. & Cambon A. & Joncheray L. (1996) 

“Santalenes from Santalum austrocaledonicum.” Phytochemistry 41, 829-832. 

Alpha T., Raharivelomanana P., Blanchini J.-P., Faure R. & Cambon A. (1997) “A sequiterpenoid 

from Santalum austrocaledonicum var. austrocaledonicum.” Phytochemistry 46, 1237-1239. 

Abstract. A new sesquiterpenoid, campherene-2,13-diol, has been isolated and characterized 

from the heartwood of Santalum austrocaledonicum var austrocaledonicum. Its structure has 

been established by the use of 1D and 2D NMR spectral techniques and shown to contain the 

campherenane skeleton. 

Alpha T., Raharivelomanana P., Blanchini J.-P., Faure R. & Cambon A. (1997) “Bisabolane 

sesquiterpenoids from Santalum austrocaledonicum”. Phytochemistry 44, 1519-1552. Abstract. 

Two new sesquiterpenoids, 6,13-dihydroxybisabola-2,10-diene and 7,13-dihydroxybisabola-2,10- 

diene, were isolated, together with (E)-anceol, from the heartwood of Santalum 

austrocaledonicum var. austrocaledonicum. The compounds were characterized by one- and twodimensional 

NMR. 

64

Alpha T., Raharivelomanana P., Blanchini J.-P., Faure R. & Cambon A. (1997) “Identification de 

deux nouveaux dihydroxyles du bisabolene a partir de santal oceanien.” In Rivista Ital. EPPOS 

(Actes des 15emes Journeés Internationales Huiles Essentielles: Digne-les-Bains, France., 5,6& 

7 Sept 1996 Special Issue 01/97 pp 84-91. 

Azais, T. (1995). “Sandalwood management in the Southern Province of New Caledonia.”. pp. 

217–227. In: Gerum, Fox, and Ehrhart Gerum, L., J.E.D. Fox, and Y. Ehrhart (eds.). 1995. 

Sandalwood Seed, Nursery and Plantation Technology. Proceedings of a regional workshop for 

Pacific Island Countries, August 1–11, 1994, Noumea, New Caledonia. RAS/92/361. Field 

Document 8. UNDP/FAO South Pacific Forestry Development Programme, Suva, Fiji. 

Bottin L, Vaillant A, Sire P, Cardi C, Bouvet J M (2005) “Isolation and characterization of 

microsatellite loci in Santalum austrocaledonicum, Santalaceae”, Molecular Ecology Notes, 5(4), 

800-802. 

Bottin L., Isnard C., Godefroy C., Lagrange A., Butaud. & Raharivelomanana, Bianchini & Bouvet 

J.M. (2005). “Chemical variability of sandalwood in New-Caledonia.” Technical note CIRAD 20p + 

annexes. 

Bottin L., Verhaegen D., Tassin J., Olivieri I., Vallant A. & Bouvet J.M. (2005) “Genetic Diversity & 

Population Structure of an Insular tree, Santalum austrocaledonicum in New Caledonian 

archipelago.” Molecular Ecology 14(7), 1979-89. Abstract: We present a study of the genetic 

diversity and structure of a tropical tree in an insular system. Santalum austrocaledonicum is 

endemic to the archipelago of New Caledonia and is exploited for oil extraction from heartwood. A 

total of 431 individuals over 17 populations were analysed for eight polymorphic microsatellite 

loci. The number of alleles per locus ranged from 3 to 33 and the observed heterozygosity per 

population ranged from 0.01 in Mare to 0.74 in Ile des Pins. The genetic diversity was lowest in 

the most recent islands, the Loyautes, and highest in the oldest island, Grande Terre, as well as 

the nearby small Ile des Pins. Significant departures from panmixia were observed for some locipopulation 

combinations (per population FIS = 0-0.03 on Grande-Terre and Ile des Pins, and 0- 

0.67 on Loyautes). A strong genetic differentiation among all islands was observed (FST = 0.22), 

and the amount of differentiation increased with geographic distance in Iles Loyaute and in 

Grande Terre. At both population and island levels, island age and isolation seem to be the main 

factors influencing the amount of genetic diversity. In particular, populations from recent islands 

had large average FIS that could not be entirely explained by null alleles or a Wahlund effect. 

This result suggests that, at least in some populations, selfing occurred extensively. Conclusively, 

our results indicate a strong influence of insularity on the genetic diversity and structure of 

Santalum austrocaledonicum. 

Bottin L. (2006) Thesis: Ecole Nationale Superieure d’Agrnomie de Montpellier. Agro Montpellier: 

Déterminants de la variation moléculaire et phénotypique d'une espèce forestière en milieu 

insulaire: cas de Santalum austrocaledonicum en Nouvelle Calédonie. – see http://tel.archivesouvertes.fr/tel-00097974/en/ 

Abstract. Les îles océaniques constituent de véritables « 

laboratoires naturels » pour comprendre l'impact des forces évolutives sur la biodiversité. Les 

effets de dérive génétique et l'impact de la sélection naturelle apparaissent d'autant plus 

exacerbés que les îles sont isolées et soumises à de forts gradients environnementaux. Notre 

étude associe des marqueurs moléculaires neutres et des caractères liés à l'adaptation afin 

d'évaluer l'influence de ces différentes forces dans le contexte insulaire de Nouvelle-Calédonie 

sur l'espèce forestière Santalum austrocaledonicum. L'étude des microsatellites nucléaires et 

chloroplastiques montre une différenciation nette des populations des petites îles Loyauté et un 

isolement par la distance au sein de l'île la plus vaste, Grande Terre. En outre elle met en 

évidence un déficit en hérérozygotes au sein de certaines populations pouvant être attribué à une 

sous-structuration spatiale ou un régime de reproduction autogame. La variation de la taille des 

feuilles et des graines, caractères liés à l'adaptation, résulte des effets de dérive mais aussi de la 

sélection naturelle provoquée par des contrastes environnementaux notamment par des 

différences de pluviométrie. De même la composition chimique du bois de coeur, analysée par 

chromatographie, subirait, en plus de la dérive, une pression sélective exercée par le cortège 

65

d'insectes et de champignons phytophages. Cette étude exploratoire permet de dégager de 

nombreuses perspectives de recherche relevant des questions évolutives en milieu insulaire. Sur 

un plan opérationnel, elle permet de définir des unités de gestion de l'espèce associant 

caractères adaptatifs et variables moléculaires. 

Bottin L., Isnard C., Lagrange A. & Bouvet J.M. (2007) "Comparative molecular and 

phytochemical study of the tree species Santalum austrocaledonicum (Santalaceae) distributed in 

the New-Caledonian archipelago." Chem Biodivers. 4(7):1541-56. Abstract. We have tried to 

elucidate the origin of phytochemical variation in trees by studying concomitantly the chemical 

and microsatellite variations in Santalum austrocaledonicum. Eight natural populations were 

sampled in the New-Caledonian archipelago, a total of 157 individuals being analyzed. The main 

components, as revealed by gas chromatography (GC), were alpha- and beta-santalol (as in 

other sandalwood species), although the level of (Z)-lanceol was particularly high. Most of the 

chemical variation was observed within populations (83.7%). With microsatellites, the variation 

between populations was more pronounced (32% of the total variation). Although the chemical 

variation between populations was small, we investigated the effects of genetic drift and migration 

by comparing the chemical- and molecular-differentiation patterns. The poor congruence between 

neighbor-joining trees, confirmed by the non-significant Mantel test between the molecular and 

chemical distance matrices (R=0.26, P=0.12), showed that genetic drift and migration are not the 

main evolutionary forces acting on chemical differentiation between populations. We could not 

find any effect of soil and rainfall conditions neither. Although the impact of drift and migration 

cannot be discounted in rationalizing between-population differentiation, the low variation among 

populations could result from a stabilizing selection caused by the same phytopathogen charge 

across the natural range. 

Bottin L., Tassin J., Nasi R. & Bouvet J.-M. (2007) "Molecular, quantitative and abiotic variables 

for the delineation of evolutionary significant units: case of sandalwood (Santalum 

austrocaledonicum Vieillard) in New Caledonia." Conservation Genetics 8(1), 99-109. Abstract. 

Various approaches have been developed to define conservation units for plant and animal 

species. In this study we combined nuclear microsatellites (from a previous published study) and 

chloroplast microsatellites (assessed in the present study), leaf and seed morphology traits and 

abiotic variables (climate and soil) to define evolutionary significant units (ESU) of Santalum 

austrocaledonicum, a tree species growing in New Caledonia. Results for chloroplast 

microsatellites showed that the total population heterozygosity was␣high, (H cp = 0.84) but varied 

between islands. Differentiation was strong in the total population (F stcp = 0.66) but also within 

the main island Grande Terre (F stcp = 0.73) and within Iles Loyauté (F stcp = 0.52), highlighting 

a limited gene flow between populations. These results confirmed those obtained with nuclear 

microsatellites. The cluster analysis on molecular markers discriminated two main groups 

constituted by the populations of Grande Terre and the populations of Iles Loyauté. A principal 

component analysis of leaf and seed morphology traits singled out the populations of Iles Loyauté 

and the western populations of Grande Terre. Quantitative genetic analyses showed that the 

variation between populations was under genetic control (broad sense heritability close to 80%). 

A high correlation between rainfall and morphological traits suggested an impact of climate on 

this variation. The integration of these results allows to define two ESUs, one corresponding to 

Grande Terre and Ile des Pins and the other the Iles Loyauté archipelago. This study stresses the 

need to restore some populations of Grande Terre that are currently threatened by their small 

size. 

Braun N.A., Meier M. & Hammweschmidt F.-J. (2005) “New Caledonian sandalwood – a 

substitute for East Indian sandalwood oil” J. Essen Oil Res 17, 477-480. Abstract: Three 

qualities of New Caledonian sandalwood oil were analysed using GC and GC/MS. Eighty-four 

constituents were identified: 10 monoterpenes, 72 sesquiterpenes and two others. In addition b- 

bisabolol/epi-b-bisabolol isomers were isolated and characterised via chiral GC chromatography. 

Our results indicate that New Caledonian sandalwood oil is much closer related to East Indian 

sandalwood oil than its West Australian counterpart. Cropwatch comments: Arguably in 2005, 

the world production of sandalwood oil was approx. 50 tons/annum, set against a demand of 200 

66

tons/annum. How then can the authors maintain, bearing in mind New Caledonia’s very limited 

production capability (1-2 tons at most), that this oil can be a substitute for the ever-scarcer East 

Indian Sandalwood oil Furthermore, the authors assume that the GC analytical trace similarity 

(i.e. between E.I. sandalwood oil against New Caledon sandalwood oil) will make it an automatic 

pertfumery substitution choice, without performing detailed odour profiling trials, or by comparing 

perforemance in product In fact the authors own figures show considerable differences in 

compositiomn exist between New Caledonoium & E.I. sandalwood oils, especially in respect to 

the high (Z)-lanceol (9.1%) and high (Z)-trans-α-bergamotol (9.9%) figures. 

H 

H 

H 

HO 

z 

Z-lanceol 

HO 

H 

(Z)-trans-alpha-bergamotol 

Brennan P. & Merlin M (1993). “Biogeography and traditional use of Santalum in the Pacific 

Region”. pp. 30–38. In: McKinnell, F.H. (ed.). 1993. Sandalwood in the Pacific Region. 

Proceedings of a symposium held on 2 June 1991 at the XVII Pacific Science Congress, 

Honolulu, Hawaii. ACIAR Proceedings 49. ACIAR, Canberra, Australia. Abstract. Santalum has a 

disjunct known distribution among the islands of the Pacific Ocean. During the prehistoric 

period, Melanesian and Polynesian Islanders, who had access to native sandalwood trees and 

shrubs, utilised the aromatic heartwood for a variety of medicinal and other purposes. Some uses 

had significant social import, motivating trade of Santalum from Fiji to Tonga for status and 

aesthetic reasons. Pre-contact trade of sandalwood may also have occurred between other South 

Pacific Islands in Eastern Polynesia. The biogeography of Santalum spp. is described, and some 

aspects of the ancient and more recent history of the use of, and human environmental impact 

on, sandalwood species in the Pacific are reviewed. 

Bulai P. & Nataniela V. (2002).. “Research, development and extension of Sandalwood in Fiji - A 

new beginning.” Paper to Regional Workshop on Sandalwood Research, Development and 

Extension in the Pacific Islands and Asia. Noumea, New Caledonia, 7–11 October 2002. 

Chauvin J.P. & Ehrhart Y. (1998). “Germination of two provenances of Santalum 

austrocaledonicum var. austrocaledonicum.” ACIAR Proceedings 84: 113–116. 

Chauvin, J.P. (1990). “La production de plants de santal en Nouvelle Caledonie.” Bois et Forests 

des Tropiques N°218, 1-10. 

Cherrier, J-F (1993). “Sandalwood in New Caledonia”. In F.H. McKinnell (ed) Sandalwood in the 

Pacific Region. Proceedings of a symposium held on 2 June 1991 at the XVII Pacific Science 

Congress, Honolulu, Hawaii. Canberra: ACIAR Proceedings No.49. pp19-23. Abstract. Results of 

research on wood formation in Santalum austrocaledonicum in New Caledonia are discussed. 

There is high variability of heartwood content at any tree size. Trees also reach maturity at 

different heights and diameters, making predictive models of limited value. The best correlation of 

yield of heartwood is with sapwood width. The latter is positively correlated with recent growth 

rate. Sapwood is at a minimum and the proportion of heartwood is highest when the tree matures 

and growth rate is reduced. It is concluded that the management of sandalwood to maximise 

heartwood production is complex. 

Douheret J. (1981). “Le santal en Nouvelle Calédonie.” Nature calédonienne 11/1981. 

Ehrhart Y. (1996). “The status of the genus Santalum and Agathis in New Caledonia.” Paper at 

SPRIG (South Pacific Regional Initiative on Forest Genetic Resources) Meeting, Nadi, Fiji 2–4 

December 1996. Unpublished. 

67

Lawrence B.M. (2008) “Progress in Essential oils. New Caledonian Sandalwood oil.” Perf. & Flav. 

33 (Juy 2008) p 44. 

Veillon J.M. & Jaffré T. (1995) “Sandalwood (Santautm azrstrocaledonicum Vieillard) in New 

Caledonia: taxonomy, distribution, ecology.” In L Gerum, JED Fox and Y Ehrhart (eds.) 

Sandalwood seed, nursery and plantation technology. Proceedings of a regional workshop for 

Pacific Island Countries; August 1-11, 1994; Noumea, New Caledonia. RAS/92/361. Field 

Document No. 8. UNDP/FAO South Pacific Forestry Development Programme, Suva, Fiji. Pp. 25- 

36.Abstract. Sandalwood is represented in New Caledonia by a single species, 

S.austrocaledonicum, which is divided into three varieties showing different geographic 

distributions: the Nouméa area for the pifosulicm variety, the foot of ultramafic rock formations for 

the minutzim variety and the Loyalty Islands, the Isle of Pines and a few locations in New 

Caledonia for the austrocaledonicum variety. It is mainly limited to secondary vegetation stands 

with diverse flora, but can also be found in sclerophyll forest and in low altitude scrub, which 

could well be its original environment. It grows in extremely varied soils, with a pH of 4 to 8 or 

more, containing a variety of exchangeable bases (Ca, K, Na); these soils may be rich in 

magnesium, nickel and chrome. Its foliar mineral composition features relatively high 

concentrations of nitrogen and potassium, low to medium-level concentrations of phosphorus and 

highly variable levels of calcium. As in other Pacific islands, sandalwood, which is highly sought 

after for its wood and essential oils, has undergone intensive harvesting in New Caledonia. This 

overexploitation, along with the destruction of its original biotopes (sclerophyll forest and lowaltítude 

scrub) by both agricultural and grazing activities and fire, has contributed to the growing 

scarcity of this species through the disappearance of numerous stands.At the present time, the 

minufum variety, which may well be an edaphic scrubland variety, can only be found in a single 

location and must be considered as an endangered species. Urgent action is therefore necessary 

for its survival. 

Tonga (Santalum yasi). 

Erhart Y. (1997) “Technical Report on Sandalwood Workshop, Tonga 17–21 November 1997. 

CIRAD–Forêt/New-Caledonia 

Kaufusi S. (1995) “Status of the Sandalwood tree in Tonga.” In:: Gerum, L., J.E.D. Fox, and Y. 

Ehrhart (eds.). 1995. Sandalwood Seed, Nursery and Plantation Technology. Proceedings of a 

regional workshop for Pacific Island Countries, August 1–11, 1994, Noumea, New Caledonia. 

RAS/92/361. Field Document 8. UNDP/FAO South Pacific Forestry Development Programme, 

Suva, Fiji.Gerum, Fox, and Ehrhart (eds) , 

Kaufusi S, Harmani S, and Thomson L. (1999). “Sandalwood work on ‘Eua, Kingdom of Tonga.”. 

Sandalwood Research Newsletter. CALM, Kununnura, Western Australia. 

Siwatibau S., Bani C, & Kaloptap J. (1998). “SPRIG Rapid Rural Appraisal Survey of selected 

tree species in Vanuatu.” Report by Island Consulting to CSIRO Division of Forestry/SPRIG 

Project. 

Yuncker T, 1959. Plants of Tonga. B.P. Bishop Museum Bulletin, 220pp. 

Vanuatu (Santalum austrocaledonicum). 

Barrance A.J. (1989). “Controlled development of sandalwood in Vanuatu - a mid-term review of 

the five year moratorium on sandalwood cutting.” Vanuata Forest Service, June 1989. 

Barrance, A. 1989 (November). Research Trials: Results of trials up to 1989. Research report on 

file, Forestry Department, Port Vila, Vanuatu. 

Berry A. (2002). Vanuatu country report. In proceedings of SPC Regional Workshop On 

Sandalwood Research, Development And Extension In The Pacific Islands And Asia. 7-11 

October 2002, Noumea, New Caledonia. (2002). 

68

Bule L. & Daruhi G. (1990) “Status of sandalwood resources in Vanuatu.” Proceedings of the 

Symposium on Sandalwood in the Pacific April 9-11, 1990, Honolulu, Hawaii Abstract. On eight 

islands of Vanuatu archipelago, sandalwood stands have been heavily exploited since the late 

1800's. Because of the over-exploitation, which worried the Vanuatu Government, a moratorium 

was imposed in early 1987. The status of the valuable wood and the beginnings of research into 

one of the country's potential commodities are reviewed. 

Channel S. & Thompson L. (1999) “Development of a Sandalwood conservation strategy for 

Vanuatu. In Forest Genetic Resources No 27, 68-72. 

Daruhi, G. (1991). “Sandelwud blong Vanuatu. A bright future” Sandalwood paper for the XVII e 

P.S.A. Congress. Honolulu, Hawaii. Forestry Department, Vila. 

Ehrhart Y. 1998. “Oil composition of the sandalwood (Santalum austrocaledonicum) from 

Erromango and Aniwa Islands, Vanuatu.” Report for CIRAD–Foret, Nouvelle–Calédonie, 10 June 

1998. Unpublished. 

Neil P.E. (1986) “Sandalwood of Vanuatu” Forest Research Report 5/86, Vanuatu Forest Service 

(5/86): ii + 7. 

Page T., Tate H., Tungon J., Sam C., Dickinson C., Robson K., Southwell I., Russell M., Waycott 

M., Leakey R. (2004). "Evaluation of heartwood and oil characteristics in nine populations of 

Santalum austrocaledonicum from Vanuatu." Sandalwood Research Newsletter Abstract. 

Heartwood cores were collected from 222 trees across nine populations on six different islands 

from Vanuatu. Oil was ethanol extracted and oil concentration and the main constituents were 

determined for each of the cores sampled andanalysed on a tree-to-tree and site-by-site basis. 

Heartwood oil concentration and all major oil constituents exhibited significanttree-to-tree 

variation, within and between all populations. Each population had a range of trees with high and 

low concentrations of α- and β-santalol. The popula-tions from the two northern islands had a 

greater proportion of trees with high santalol content than the populations sampled from the 

southern islands 

Tacconi L. & Mele. L. (1995). Economic Aspects of Sandalwood Cultivation in Relation to the 

Erromango Kauri Protected Area. Vanuatu Forest Conservation Research Report 7. Department 

of Economics and Management, University College, University of New South Wales, Campbell, 

Canberra, Australia. 

Tacconi L. & Mele l. (1997). “Sandalwood Cultivation and the Establishment of the Erromango 

Kauri Protected Area, 71-85.” In: Tacconi, L. and Bennett, J. (Eds) Protected Area Assessment 

and Establishment in Vanuatu: a Socioeconomic Approach ACIAR Monograph 38, ACIAR, 

Canberra, 180p. 

Tate H., Sethy M. & Tungon J. (2004) "Grafting Sandalwood in Vanuatu." Sandalwood Research 

Newsletter . Abstract. Historically sandalwood plantings in Vanuatu have been established 

mainly by seed propagation and transplanted wildings. This method continues to be very 

important for village communities to grow sandalwood collected from their natu-ral sources. With 

increasing interest across the country in planting sandalwoodthe Department of Forests (DoF) is 

actively encouraging improved clonal seed orchards to keep up with demand. Clonal propagation 

of mature trees by cuttings has been difficult to achieve byconventional methods, but grafting has 

proven a viable alternative method. Thesuperior individuals identified within the current ACIAR 

sandalwood project are now being grafted using the methods developed in conjunction with 

SPRIG 

General Pacific Region 

Allen J.A. (2002).” Santalum freycinetianum Gaudich.” In Vozzo, J.A. (ed.). Tropical Tree Seed 

Manual. Agriculture Handbook 721. U.S. Forest Service, Washington, DC. 

69

Alpha T., Raharivelomanana P., Bianchini J.-P., Faure R., Cambon A., & Joncheray L. (1995) 

"alpha-Santaldiol & beta-santaldiol, two santalane sesquiterpenes from Santalum insulare." 

Phytochem. 41(3), 829-831. Abstract. Two new sesquiterpene alcohols, beta-santaldiol and 

alpha-santaldiol, have been isolated from the heartwood of Santalum insulare var. marchionese 

and, by means of two-dimensional NMR experiments, shown to have the beta- and alphasantalane 

skeleton respectively. 

Alpha, T. (1997). Etude des concrètes et des essences de santal d’origine océanienne. 

Elucidation de nouveaux sesquiterpenoïdes par la RMN multi-impulsionnelle et bidimensionnelle. 

Ph.D. Thesis. Université Française du Pacifique, Papeete, French Polynesia. 

Alpha, T., P. Raharivelomanana, J.-P. Bianchini, Y. Ehrhart & A. Cambon. (1997). “Etude de la 

composition chimique d’essences de santal d’origine du Pacific Sud.” pp 499–465 In: Revista 

Italiana EPPOS (Actes des 15èmes Journées Internationales Huiles Essentielles; Digne-les- 

Bains, France, 5, 6 & 7 Septembre 1996, Special issue 01/97. 

Applegate G.B. (1990). “Sandalwood in the Pacific: A state of knowledge. Synthesis and 

summary from the April 1990 symposium.” pp 1–11. In: Hamilton, L., and C.E. Conrad (eds.). 

1990. Proceedings of the Symposium on Sandalwood in the Pacific, April 9–11, 1990, Honolulu, 

Hawai‘i. General Technical Report PSW–122. Pacific Southwest Research 

Barrau, J. (1960). “Plantes utiles des îles du Pacifique–Le Santal.” Bulletin des études Pacifiques, 

July 1960. 

Brennan, P., and M. Merlin. (1993). "Biogeography and traditional use of Santalum in the Pacific 

Region". In: F. McKinnell (ed.). Proceedings of a symposium held on June 2, 1991 at the XVII 

Pacific Science Congress, Honolulu, Hawai‘i. ACIAR Proceedings 49. ACIAR, Canberra, 

Australia. 

Butaud J.F. & Tetuanui W. (2002). “Le Santal en Polynésie Française.” Proceedings of the 

Regional Workshop on Sandalwood Re-search, Development and Extension in the Pacific 

Islands andAsia, 7-11 October, 2002,Noumea, New Caledonia 

Butaud J.-F. (2004) "Santalum insulare (Bertero ex A. DC.): Distribution and ecology." 

Sandalwood Research Newsletter 19. July 2004. Abstract. The Eastern Polynesian sandalwood 

is one of the sixteen Santalum species of Asia and the Pacific. It is known under the name puahi 

in Marquesas Islands, ai in Cook Islands and ahi elsewhere in its geographical area. 

Overexploited during the beginning of the 19 th century, Polynesian sandalwood is still used for 

carvingor in powder mixed with coconut oil (monoi ahi or pani puahi) for cosmetic or medicinal 

purposes. 

Butaud J.F. (2003). “Autécologie et phytosociologie des Santals de Polynésie française.” 

Proceedingsof the Regional Workshop on San-dalwood Research, Developmentand Extension in 

the Pacific Is-lands and Asia, 7-11 October, 2002, Noumea, New Caledonia. 

Butaud J.-F., Rives F., Verhaegen D. * Bouvet J.-M. (2005) “Phylogeography of Eastern 

Polynesian sandalwood (Santalum insulare), an endangered tree species from the Pacific: a 

study based on chloroplast microsatellites.” Journal of Biogeograohy 32 (10) , 1763–1774. 

Abstract. Aim: Patterns of genetic variation within forest species are poorly documented in island 

ecosystems. The distribution of molecular variation for Santalum insulare, an endangered tree 

species endemic to the islands of eastern Polynesia, was analysed using chloroplast 

microsatellite markers. The aims were to quantify the genetic diversity; to assess the genetic 

structure; and to analyse the geographical distribution of the diversity within and between 

archipelagoes. The ultimate goal was to pre-define evolutionary significant units (ESUs) for 

conservation and restoration programmes of this species, which constitutes a natural resource on 

small, isolated islands. 

70

Location Eleven populations, each representative of one island, covering most of the natural 

occurrence of S. insulare were sampled: five populations from the Marquesas Archipelago; three 

from the Society Archipelago; and three from the Cook–Austral Archipelago. These South Pacific 

islands are known for their high degree of plant endemism, and for their human occupation by 

Polynesian migrations. The extensive exploitation of sandalwood by Europeans nearly 200 years 

ago for its fragrant heartwood, used overseas in incense, carving and essential oil production for 

perfume, has dramatically reduced the population size of this species. 

Methods We used chloroplast microsatellites, which provide useful information in 

phylogeographical forest tree analyses. They are maternally inherited in most angiosperms and 

present high polymorphism. Among the 499 individuals sampled, 345 were genotyped 

successfully. Classical models of population genetics were used to assess diversity parameters 

and phylogenetic relationships between populations. 

Results Four microsatellite primers showed 16 alleles and their combinations provided 17 

chlorotypes, of which four exhibited a frequency > 10% in the total population. The gene diversity 

index was high for the total population (He = 0.82) and varied among archipelagoes from He = 

0.40 to 0.67. Genetic structure is characterized by high levels of differentiation between 

archipelagoes (36% of total variation) and between islands, but differentiation between islands 

varied according to archipelago. The relationship between genetic and geographical distance 

confirms the low gene flow between archipelagoes. The minimum spanning tree of chlorotypes 

exhibits three clusters corresponding to the geographical distribution in the three main 

archipelagoes. 

Main conclusions The high level of diversity within the species was explained by an ancient 

presence on and around the hotspot traces currently occupied by young islands. Diversity in the 

species has enabled survival in a range of habitats. Relationships between islands show that the 

Cook–Austral chlorotype cluster constitutes a link between the Marquesas and the Society 

Islands. This can be explained by the evolution of the island systems over millions of years, and 

extinction of intermediary populations on the Tuamotu Islands following subsidence there. Based 

on the unrooted neighbour-joining tree and on the genetic structure, we propose four ESUs to 

guide the conservation and population restoration of Polynesian Sandalwood: the Society 

Archipelago; the Marquesas Archipelago; Raivavae Island; and Rapa Island. 

Butaud J.-F., Raharivelomanana P., Bianchini J.-P. Faurec R. & Gaydou E.M. (2006) "Leaf C- 

glycosylflavones from Santalum insulare (Santalaceae)" Biochemical Systematics and Ecology 

34(5), 433-435 

Butaud J.-F., Raharivelomanana P., Bianchini J.-P. & Gaydou E.M. (2008) "Santalum insulare 

Acetylenic Fatty Acid Seed Oils: Comparison within the Santalum Genus." J of American Oil 

Chemists Society 85(4), 353-356.. Abstract. The sandalwood kernels of Santalum insulare 

(Santalaceae) collected in French Polynesia give seed oils containing significant amounts of 

ximenynic acid, E-11-octadecen-9-oic acid (64–86%). Fatty acid (FA) identifications were 

performed by gas chromatography/mass spectrometry (GC/MS) of FA methyl esters. Among the 

other main eight identified fatty acids, oleic acid was found at a 7–28% level. The content in 

stearolic acid, octadec-9-ynoic acid, was low (0.7–3.0%). An inverse relationship was 

demonstrated between ximenynic acid and oleic acid using 20 seed oils. Results obtained have 

been compared to other previously published data on species belonging to the Santalum genus, 

using multivariate statistical analysis. The relative FA S. insulare composition, rich in ximenynic 

acid is in the same order of those given for S. album or S. obtusifolium. The other compared 

species (S. acuminatum, S. lanceolatum, S. spicatum and S. murrayanum) are richer in oleic acid 

(40–59%) with some little differences in linolenic content. 

Chauvin J.-P. & Erhart J. “Germination of two provenances of Santalum austrocaledonicum var. 

austrocaledonicum.” ACIAR-Proceedings Series 84, 113-116. 

Doran J., Thomson L., Brophy J., Goldsack B., Bulai P., Faka'osi & Mokosa T. (date) "Variation 

in heartwood oil composition of young sandalwood trees in the South Pacific (Santalum yasi, S. 

album and F1 hybrids in Fiji, and S. yasi in Tonga and Niue)." Abstract. This study was 

undertaken during 2003 as part of AusAID’s SPRIG (South Pacific Regional Initiative in Forest 

71

Genetic Re-sources) project. It had the primary aim of extending the knowledge base on the 

production of heartwood and heartwood oils in young Pacific Island sandalwoods, Santalum yasi, 

the introduced S. album, and the spontaneous F1 hybrid, S. album ×yasi. A solvent (pentane) 

extraction technique was used to determine heartwood oil chemistry, following verification 

againststeam distillation. The heartwood was obtained from trees by non-destructive coring. Ages 

of the trees sampled ranged between 5 years and more than 25 years. Many of them had not yet 

started to lay downheartwood at their base. For those that had, heartwood was restricted to the 

lower most cores i.e. 0.1m or 0.2m above groundor very occasionally extending to 0.3m in older 

trees. Tree-to-tree variation in oil quality in S. yasi, as determined by allow-able α-santalol and β- 

santalol levels in the International Standard (2002) for S. album, was substantial indicating a 

potentialof improvement through selection and breeding if genetic parameters are favourable. 

Trees in Fiji of the spontaneous F1 hy-brid, S. album × yasi, were very vigorous and the 

heartwood oil of two (out of three) of the 7-year-old trees with heartwoodwas of excellent quality. 

The results suggest that rotation lengths of 25 to 30 years for the Pacific sandalwoods may be 

more realistic than the 15 to 20 year rotation lengths suggested by some workers. 

Ehrhart Y. (1997). Technical Report on Sandalwood Workshop, Tonga 17–21 November 1997. 

CIRAD–Forêt/ New Caledonia, Pouembout. Unpublished. 

Ehrhart Y. (1998) "Descriptions of some sandal tree populations in the South West Pacific : 

consequences for the silviculture of these species and provenances." In : Radomiljac A.M. (ed.), 

Ananthapadmanabho H.S. (ed.), Welbourn R.M. (ed.), Satyanarayana Rao K. (ed.). Sandal and 

its products : proceedings of an international seminar. Canberra : ACIAR, p.105-112. Sandal and 

its Products, 1997-12-18/1997-12-19, (Bangalore, Inde). Abstract. Many of the islands of the 

South West Pacific that bear sandal have been visited and the stands described. Mostly the 

population is depleted, but some stands still exist. Depending on the status of the existing 

population, several possible management strategies are feasible. The aim is to rebuild stands 

which are as diverse as possible which will be able to be managed sustainably in a few decades. 

Some are presently managed with the objective of regular annual heartwood production with an 

increase of the stock. The observations reported here, especially those regarding shade intensity, 

can be used to improve the silviculture of the various provenances which differ markedly. Even 

aspects of seed storage differ, and this demande further investigation. New techniques, which 

differ significantly from those previously identified for the Ile des Pins provenance, are proposed. 

(Résumé d'auteur) 

Elevitch C.R. & K.M. Wilkinson K.M..(2003). “Propagation protocol for production of container 

Santalum freycinetianum Gaudich.” In: Native Plant Network.University of Idaho, College of 

Natural Resources, Forest Research Nursery, Moscow, Idaho.–see 

http://www.nativeplantnetwork.org 

Emeline L., Alexandre V, Jean-Francois B. et al. (2006) "Isolation & characterisation of 

microsatellite loci in Santalum insulare, Santalaceae." Molecular Ecology Notes, 2006. 

Felgelson J. (1990) "Sandalwood - the myth & the reality." Paper presented at the Symposium on 

Sandalwood in the Pacific, April 9-11, 1990, Honolulu, Hawai‘i. Abstract: Santalum paniculatum 

trade after more than a century was revived by the author in 1988. Revival of the trade has called 

attention to this resource, and the focus is now on management of this resource. A discussion 

about recent sandalwood logging and marketing activities in Hawai‘i is presented. The author also 

points out various anomalies that may be related to habitat and land use variations. The 

obligatory parasitic nature of this species is questioned and the coppicing tendency is confirmed. 

Criteria are suggested concerning the harvesting and sales that minimize fragmentation of forest 

areas. The concept of establishing a sandalwood research center and the cultivation of 

sandalwood in Hawaii is presented. 

Fosberg F.R. & Sachet M.H. “Santalum in Eastern Polynesia” Candollea 40, 459-470. 

72

Harbaugh D.T. & Bruce G. Baldwin B.G. (2007) American Journal of Botany 94, 1028-1040. 

"Phylogeny and biogeography of the sandalwoods (Santalum, Santalaceae): repeated dispersals 

throughout the Pacific." Abstract. Results of the first genus-wide phylogenetic analysis for 

Santalum (Santalaceae), using a combination of 18S–26S nuclear ribosomal (ITS, ETS) and 

chloroplast (3' trnK intron) DNA sequences, provide new perspectives on relationships and 

biogeographic patterns among the widespread and economically important sandalwoods. 

Congruent trees based on maximum parsimony, maximum likelihood, and Bayesian methods 

support an origin of Santalum in Australia and at least five putatively bird-mediated, long-distance 

dispersal events out of Australia, with two colonizations of Melanesia, two of the Hawaiian 

Islands, and one of the Juan Fernandez Islands. The phylogenetic data also provide the best 

available evidence for plant dispersal out of the Hawaiian Islands to the Bonin Islands and 

eastern Polynesia. Inability to reject rate constancy of Santalum ITS evolution and use of fossilbased 

calibrations yielded estimates for timing of speciation and colonization events in the 

Pacific, with dates of 1.0–1.5 million yr ago (Ma) and 0.4–0.6 Ma for onset of diversification of the 

two Hawaiian lineages. The results indicate that the previously recognized sections Polynesica, 

Santalum, and Solenantha, the widespread Australian species S. lanceolatum, and the Hawaiian 

species S. freycinetianum are not monophyletic and need taxonomic revision, which is currently 

being pursued 

Hirano R.T. (1990) “Propagation of Santalum, Sandalwood tree.” Proceedings of the Symposium 

on Sandalwood in the Pacific April 9-11, 1990, Honolulu, Hawaii Abstract. The history of the 

genus Santalum (sandalwood) in Hawaii is re-viewed, along with all the early reference regarding 

its botany and horticulture. This paper gives some seed germination and viability information on 

Santalum haleakalae Hbd. and S. paniculatum H. & A. both native to Hawaii and Santalum album 

L. native to Indonesia. Germination was shown to be highly variable: as early as 26 days after 

sowing for S. album, 75 days for S. paniculatum, and 155 days for S. haleakalae. Seed viability 

varied from 324 days in S. album, 387 days in S. haleakalae and 824 days in S. paniculatum. 

Germination percentages ranged from 38 percent to 77 percent. This study also showed that 

supplemental chelated iron is essential in the propagation of all the species tested. 

Lhuillier E., Butaud J.F. & Bouvet J.M. (2006) "Extensive clonality and strong differentiation in the 

insular pacific tree Santalum insulare: implications for its conservation." Ann Bot (Lond). 98(5), 

1061-72. Abstract. BACKGROUND AND AIMS: The impact of evolutionary forces on insular 

systems is particularly exacerbated by the remoteness of islands, strong founder effects, small 

population size and the influence of biotic and abiotic factors. Patterns of molecular diversity were 

analysed in an island system with Santalum insulare, a sandalwood species endemic to eastern 

Polynesia. The aims were to evaluate clonality and to study the genetic diversity and structure of 

this species, in order to understand the evolutionary process and to define a conservation 

strategy. METHODS: Eight nuclear microsatellites were used to investigate clonality, genetic 

variation and structure of the French Polynesian sandalwood populations found on ten islands 

distributed over three archipelagos. KEY RESULTS: It was found that 58 % of the 384 trees 

analysed were clones. The real size of the populations is thus dramatically reduced, with 

sometimes only one genet producing ramets by root suckering. The diversity parameters were 

low for islands (n(A) = 1.5-5.0; H(E) = 0.28-0.49). No departure from Hardy-Weinberg proportion 

was observed except within Tahiti island, where a significant excess of homozygotes was noted 

in the highland population. Genetic structure was characterized by high levels of differentiation 

between archipelagos (27 % of the total variation) and islands (F(ST) = 0.50). The neighbourjoining 

tree did not discriminate the three archipelagos but separated the Society archipelago 

from the other two. CONCLUSIONS: This study shows that clonality is a frequent phenomenon in 

S. insulare. The genetic diversity within populations is lower than the values assessed in species 

distributed on the mainland, as a consequence of insularity. But this can also be explained by the 

overexploitation of sandalwood. The differentiation between archipelagos and islands within 

archipelagos is very high because of the limited gene flow due to oceanic barriers. Delineation of 

evolutionary significant units and principles for population management are proposed based on 

this molecular analysis. 

73

McKinnell, F.H. (ed.). (1993). "Sandalwood in the Pacific Region". ACIAR Proceedings 49. 

ACIAR, Canberra, Australia. 

Merlin M.D., Thomson L.A.J. & Elevitch C.R. (2005). “Santalum ellipticum, S. freycinetianum, S. 

haleakalae, and S. paniculatum (Hawaiian sandalwood), ver. 3.1.” In: C.R. Elevitch (ed.). Species 

Profiles for Pacific Island Agroforestry. Permanent Agriculture Resources (PAR), Holualoa, 

Hawai‘i. http://www.traditionaltree.org. Cropwatch comments: Recommended article & excellent 

biblio! 

Raharivelomanana P., Bianchini J.-P., Faure R., Cambon A. & Azzaro M. (1994) Phytochem. 

35,1059. 

Raharivelomanana P., Faure R., Cambon A. & Azzaro M. (1993) Phytochem. 33,235. 

Skottsberg C. (1930) "Further notes on Pacific sandalwoods." Acta Horti Gothob. 5, 135–145. 

Skottsberg C. (1930) "The geographical distribution of the sandalwoods and its significance." 

Proc. 4th Pacific Sci. Congr., Java 3, 435–442. 

Thomson, L.A.J. (2005). “Santalum austrocaledonicum and S. yasi (sandalwood).” ver. 1.1. In: 

Elevitch, C.R. (ed.). Species Profiles for Pacific Island Agroforestry. Permanent Agriculture 

Resources (PAR), Hōlualoa, Hawai‘i.

Radomiljac, A. M. & Bosimbi, D. (1999) “Santalum macgregorii F. v. Mueller in Papua New 

Guinea.” Sandalwood Research Newsletter 8, 5. Department ofConservation & Land 

Management, West-ern Australia 

Thomson, L. & Bosimbi, D. (2000) “Santalum macgregorii – PNG sandalwood.” Un-published 

paper prepared for CSIRO/PNGForest Research Institute/ACIAR project entitled Domestication of 

papua NewGuinea’s Indigenous Forest Species. Australian Tree Seed Centre, CSIRO Forestry 

and Forest Products, Yarralumla, ACT. 

Vernes T. (2001) “Preliminary results from Santalum macgregorii ex situ conservation planting.” 

Sandalwood Research Newsletter 10, 6-8. 

Sri Lankan Sandalwood (Santalum album) 

Anon (2000) “Santalum album in Sri Lanka.” Parasitic Plants Newsletter No 38 (2000). 

Kathriarachchi H. S. & Tennakoon K. U. (1999). "A preliminary investigationon the biology of 

Santalum album (Sandalwood) in Sri Lanka." Proceedings of the Sri lanka Association forthe 

Advancement of Science, Annual Sessions 55, 147-148 

Tennakoon K.U., Ekanayake S.P. & Etampawala L. (2000). “An overview of Santalum album 

research in Sri Lanka.” International Sandalwood Research News Letter 11, (1-4). Abstract. This 

paper outlines the background and present status of Santalum album research in Sri Lanka. The 

cur-rent project is a detailed study undertaken to investigate the biology, ecology, siviculture and 

physiology of sandalwood in Sri Lanka. Assessments made during the pilot study and the 

experimental data collectedfrom the two established model nurseries of sandalwood will be used 

to provide training and know how tothe farmers and interested governmental and nongovernmental 

organisations to establish their own San-dal nurseries and subsequent sandalwood 

plantations. This project is funded by the Community Environ-ment Initiative Facility implemented 

by the Environment Action 1 project of the Ministry of Forestry and Environment. Sri Lanka under 

a World Bank fund. 

Wijesinghe S. (2009) "A ‘scented’ gesture towards a green world." Daily News 27th July 2009. 

Synopsis: Sri Lankan Environment and Natural Resources Minister, Patali Champika Ranawaka 

in combination with the patronage of Tharunyayata Hetak president Namal Rajapaksa announced 

a policy of Sandalwood (Santalum album) cultivation where the species will be planted in places 

of religious worship. Torchwood Ivestments plc has a progamme planting Sandalwood trees at 

22,400 religious locations in Sri Lanka. 

Thai Sandalwood (Santalum album). 

Anon (2004). “Police hunt for Thai sandalwood collectors” New Straits Times, Johor Baru. 

28.6.2004 

Timorese Sandalwood (Santalum album). 

Alongi D.M. & de Carvalho N.A. (date) "The effect of small-scale logging on stand 

charfacteristics & soil biogeochemistry in mangrove forests of Timor Leste." Forest Ecology and 

Management, 255 (3),1359-1366. Abstract. The impact of small-scale cutting of mangroves by 

family groups was examined in three high-salinity forests on the dry tropical, north coast of Timor 

Leste. Before logging, these forests were characterized by moderately dense stands (3633–9610 

stems ha−1) of Ceriops tagal, Rhizophora apiculata, Bruguiera gymnorrhiza, and Avicennia 

marina, with average basal areas of 13–34 m2 ha−1, total above-ground biomass of 51–221.5 t 

ha−1, canopy cover of 61–73%, and leaf area index (LAI) of 4.9–5.4 m2 leaf area m−2 ground 

area. Approximately 1 year after the start of harvesting, these forests experienced a 30–50% 

decline in live stems and a 46–86% loss of above-ground biomass with more canopy gaps 

between less dense, smaller trees. There was some evidence of selectivity of trees 5–15 cm dbh 

in size, interpreted as a trade-off between cutting trees small enough for women and children to 

carry but large enough to warrant cost/benefit of selling for firewood. Concentrations of most 

75

particulate nutrients increased in surface soils in the harvested stands, reflecting bark, leaves, 

twigs, and small branches discarded on the forest floor. Interstitial concentrations of dissolved 

sulfide, metals, and ammonium also increased due to enhanced soil desiccation (evidenced by 

increased salinity) and decline in solute uptake and O2 translocation to live roots. Rates of 

anaerobic soil metabolism (sulfate reduction) declined after the onset of cutting, attributed to the 

decline in live roots and their metabolic activities. These cutting operations, although small-scale, 

are unsustainable as these forests are likely to be slow-growing in such highly saline soils. A 

community-based approach to conservation and sustainable management of the remaining 

mangrove forests of Timor Leste is recommended.Cropwatch comments: Article mentions 

decline of sandalwood forests, once plentiful with white sandalwood up to 1915, through export of 

wood to China, Indonesia & Europe 

Badan PengembanganEkspor Nasional (BPEN). 1993. "Informasi perkembangan harga minyak 

Cendana (Sandalwood) tahun 1993." Pusat Informasi dan Analisa Pasar. Jakarta. 

Brand. J.E. (1993) Phenotype and genotype variation within Santalum album in West Timor. 

Thesis for degree of master of Science in Biology, Curtin University of Technology (unpublished). 

DepartmentKehutanan (1991). “Cendana (Santalum album L.).” Kupang. Bagian Proyek 

Perencanaan Pimbinaan dan Pengendalian Pembangunan Kehutanan Kantor Wilayah, 

Department Kehutanan. 

Effendi M. & Susila, I.W.W. (1994) "Genetics improvement of sandalwood (Santalum album L.) in 

Nusa Tenggara Timur" International Symposium on Asian Tropical Forest Management: 

proceedings, Samarinda, 13-15 Sep 1994 

Effendi M. I. & Rachmawati dan U.R.Fauzi (1995). “Identifikasi sumber benih cendana (Santalum 

album) di Nusa Tenggara Timur dan Yogyakarta.” Santalum 17. 20-27 

Fox, J.E.D. (1990). “Silviculture of Santalam album (sic) in Timor NTT. (Report for the period 

1988-1990). ACIAR /Australia-Indonesia Sandalwood Project. Curtin University Western 

Australia. 

Fox J.E.D., Brand J.E., Barrett D.R., Markhum E. (1995). “Genetic variation in Santalum album in 

Timor.” In Sandalwood Seed Nursery and Plantation Technology, (Eds. LGjerum, JED Fox, L 

Erhart) pp 93–110. (FAO: Suva, Fiji). 

Fox J E D, Doronila A I, Barrett D R & Surata I K (1996) “Desmanthus virgatus (L.) Willd. an 

efficient intermediate host for the parasitic species Santalum album L. in Timor, Indonesia.” 

Journal of Sustainable Forestry 3(4):13–23. 

Harisetijono & Sutarjo Suriamihardja (1993). “Sandalwood in Nusa Tenggara Timur.” In 

McKinnell, F.H. (ed) Sandalwood in the Pacific Region. Proceedings of a symposium held on 2 

June 1991 at the XVII Pacific Science Congress, Honolulu, Hawaii. Canberra ACIAR Proceedings 

No.49 pp39-43. 

Hamzah (1976). “Sifat Silvika dan Silivikultur Cendana (Santalum album L.) di Pulau Timor.” 

Laporan Lembaga Penelitian Hutan. Bogor 227. 

Husain A.M.M. (1983). Report on the Rehabilitation of Sandalwood and the Trade in Nusa 

Tenggara Timur Indonesia. PPIPD West Timor 

Kharisma & Sutarjo, S. (1988). “Effects of host plants on seedling growth of Cendana (Santalum 

album L.). Santalum 2,1-8 

Kushalapa K.A. (1998). “Trade liberalisation in Sandalwood.” In Radomiljac, A.M, 

Ananthapadmanabho, H.S, Welbourne, R.M, and Satyanarayan Rao, K. (eds), Sandal and its 

Products. Proceedings of an international seminar held on 18-19 December 1997 organised by 

76

the Institute of Wood Science and Technology (ICFRE) and Karnataka State Forest Department, 

Bangalore India. Canberra: ACIAR Proceedings No 84 24-26. 

McWilliam A. (2001) “Haumeni, not many: renewed plunder and mismanagement in the Timorese 

Sandalwood Industry” Resource Management in Asia Pacific Working Paper No 29 pub. 

Resource Management in Asia-Pacific Program, Division of Pacific and Asian History, Research 

School for Pacific and Asian Studies, The Australian National University, Canberra 2001. 

Marks S.V. (2002) “NTT sandalwood: roots of disaster.” Bulletin of Indonesian Economic Studies 

38(2), 223-240. Abstract. For decades the government of Nusa Tenggara Timur (NTT) province 

has exploited the sandalwood sector, to the detriment of the growers of the trees. Severe 

depletion of the stock of sandalwood in the province has been the result. This paper documents 

NTT policies toward the sector, which it argues have been both inefficient and inequitable, and 

offers a detailed approach for reform. It also examines the political economy of these policies, 

and argues that the case of sandalwood provides an example of the dangers of decentralisation 

of economic authority in the absence of local democracy. 

Messakh, M & Dewa A. (1999). “Dalam Hutanku ada cendana, tapi bukan milikku. [In my forest 

there is sandalwood but it does not belong to me].” Udik: Advokasi Newsletter (5) Kupang. August 

Messakh M.V. (1999). “Orang Timor Mencuri Cendana di Tanah Sendiri. Suatu tinjauan terhadap 

kebijakan Pemerintah Daerah NTT tentang komoditas cendana dan implikasi bagi kesejahteraan 

masyarakat lokal. [Timorese Steal Sandalwood from their Own Land: A study of NTT Government 

policy towards sandalwood and the implications for local community welfare].” Lokakarya 

Penulisan Pegelolaan Sumber Daya Alam untuk Rakyat. Lembaga Alam Tropika Indonesia 

(LATIN) [unpublished]. 

Nuningsih R., Mudita I .W., & Mella W. (1994). “Kajian Permudaan Cendana (Santalum album L) 

Secara vegetatif pada Habitat Alamiah di Timor Tengah Selatan NTT. [Study of vegetative root 

propagation of Sandalwood in natural habitats of South Central Timor].” Kupang. Universitas 

Nusa Cendana. 

Nuningsih R. (1996). “Kajian Perkembangan Sistem Perakaran Anakan Vegetatif Alami Cendana 

(Santalum album L.) pada Habitat Alaminya di Kabuapaten Timor Tengah Selatan. [Study of 

development of vegetative root sprouting of Sandalwood in natural habitats of South Central 

Timor]”. Kupang: Universitas Nusa Cendana. 

Ormeling F.J. (1955) "The Timor problem. A geographical interpretation of an undeveloped 

island." PhD thesis, University of Indonesia. J.B. Walters, Djakarta & Groningen (through Rohadi 

et al. (undated). 

Rahm Th. (1925).”Sandelhout op Timor.” Tectona Buitenzorg 18, 499-545. 

Rohadi D., Maryani R., Widyana M. & Azhar I. (undated) "Ch 12. A case study of the productionto-consumption 

system of sandalwood (Santalum album) in South Central Timor, Indonesia. See 

http://www.cifor.cgiar.org/publications/pdf_files/Books/NTFPAsia/Chapter12-Chapter16.PDF. 

Cropwatch comments: Recommended study of the Timorese sandalwood situation. 

Rohadi D., MAryani R., Belcher B., Perez M., & Widnyana M. (2000). "Can sandalwood in East 

Nusa Tenggara survive Lessons from the policy impact on resource sustainability." Sandalwood 

Research Newsletter Issue 10, 3-6. Abstract. This paper discusses the policy aspects of 

sandalwood in East Nusa Tenggara province, focusing primar-ily on the impacts of regional 

government regulations on the resource sustainability. The paper is basedon a field survey that 

was conducted during July-August 1999, as well as from various publications and official reports 

from the region 

Setiadi D & Komar T.E. (2001) "Current Sandalwood seed source in Timor Island." Sandalwood 

Research Newsletter 13. Abstract. Sandalwood (Santalum album Linn) is one of the native 

77

species to East Nusa Tenggara which hashigh economic value. Effort has been put to increase 

its productivity, especially through artificialplantation since its natural regeneration success is very 

low. Artificial regeneration is the onlyalternative to overcome the shortage of raw material for 

various wood-base industries as well as for the production of santalol. 

Steenis C.G.G.J van (1939). “The native country of sandalwood and teak: a plant geographical 

study.” Hendelingen 8e Nederland Indische. Natuurwetenschappelijke Congres, Sorabaja, pp408- 

418. 

Susila I. W. W. (1994). “Estimate of hard-wood yield and natural regeneration ofsandalwood 

(Santalumalbum) inAmanuban Selatan, Timor Tengah Selatan.” Santalum 15. 

Surata K. (1992). “Effect of host plants on growth of sandalwood (Santalum album) seedlings.” 

Santalum 9,1-10. 

Surata I. K., Sutrisno E. & Sinaga M. (1995) "Utilisation and conservation ofSandalwood in Nusa 

Tenggara Timur,Indonesia." In Cjerum L., Fox J. E. D. & Ehrhart Y. (Editors). Sandalwood seed 

nursery and plantation technology (Proceedings). RAS/92/361 Field Document No. 8. CIRAD, 

ACIAR and UNDP. 

Surata, K., Harisetijono & Sinaga, M., (1993). “Effect of intercropping system on sandalwood 

growth (Santalum album) “ Santalum 20, 17-24. 

Suriamihardja S & Susila I.W.W. (1993). “Strategi dan Upaya Pelestarian Potensi Cendana di 

Nusa Tenggara Timur [Strategies and Efforts for the Preservation of Sandalwood in NTT]” 

Savanna. Kupang: Balai Penelitian Kehutanan. 1-8. Suripto, 1992. Pemulihan Potensi Cendana 

di NTT. Makalah 

Susila I.W.W. & Ormeling, F.J. (1955) “The Timor Problem.”. J.B. Walters, Djakarta, Gröningen. 

994). "Estimate of hardwood yield & natural regeneration of sandalwood (Santalum album) in 

Amanuban Seletan, Timor Tengah Seltan. Santalum 15 

Susila I.W.W. (1994) “Estimation of heartwood yield & natural regeneration of sandalwood in 

Amanuban Selatan, Timor Tenga Selatan.” Santalum 15. 

Wright, A. (2001). “East Timor (Timor Timur) sandalwood plantation development: a feasibility 

study.” Sandalwood Research Newsletter 12, 5-6. 

Unclassified Articles. 

Denham R (1998) “Southern Sandalwood: an introduction” 

http://agspsrv38.agric.wa.gov.au/pls/portal30/docs/folder/ikmp/lwe/vegt/trees/f02798.pdf. 

McKinnell F.H. (1990) “Status of Management & Silviculture research on Sandalwood in W 

Australia and Indonesia” in Proc of the symposium on sandalwood in the Pacific : April 9-11, 

1990, Honolulu, Hawai/technical co-ordinators: Lawrence Hamilton, C. Eugene Conrad. Pub: 

Symposium on Sandalwood Conservation (1st: 1991: Honolulu, Hawaii). Abstract. The current 

status of the conservation and management of Santalum spicatum in Western Australia and S. 

album in East Indonesia is outlined. Natural and artificial regeneration techniques for both species 

in selected areas are discussed. The present Australian Centre for International Agricultural 

Research program on S. album in Nasa Tenggara Timur is described in relation to the 

management needs of the species in that province. In S. spicatum, research on silviculture is 

essentially complete, and interest is now focused on the marketability of the kernels for human 

consumption. 

Metcalf C.R. (1935) “The structure of some Sandalwoods and their substitutes and of some other 

little known scented woods.” Bulletin of Miscellaneous Information (Royal Gardens, Kew), 1935 – 

JSTOR. 

78

Nagaraja Rao (1939) J Ind Chem Soc Ind Division (1939) 2, 1. 

Naipawer R.E. (1988) "Synthetic sandalwood chemistry - a decade in review." Dev. Food Sci. 18, 

805-818. 

Neil, P. E. (1989). “Possible techniques for raising and planting sandalwood in Nepal.” Banko 

Janakari 2(3):1-6; 1989. 

Neil P.E. (1990) “Growing sandalwood in Nepal - Potential Silvicultural Methods and Research 

Priorities.” Proceedings of the Symposium on Sandalwood in the Pacific April 9-11, 1990, 

Honolulu, Hawaii Abstract. Interest in sandalwood has increased recently in Nepal as a result of a 

royal directive to plant it in the Eastern Development Region. The most suitable seed sources, 

seed acquisition, nursery techniques, direct sowing and plantation establishment methods are 

discussed here on the basis of results from elsewhere. Suggestions are made as to what 

research is most needed to assist with successful establishment of sandalwood in Nepal. The 

silvicultural methods discussed could well be of use to other countries that are interested in 

introducing and establishing sandalwood plantations. 

Surendran, C., Partiban, K.T., Bhuvenaswaran, C. & Murugesh, M. (1998). “Silvicultural 

strategies for augmentation of sandal regeneration.” In: Radomiljac AM, Ananthapathmanabha 

HS, Welbourn RM, Stayanarayan K. (eds.) Sandal and its products. ACIAR Proceedings Volume 

84. Arawang Communications, Canberra. pp. 69-73. 

Wang Z, Hong X. (1991) "]Comparative GC analysis of essential oil in imported sandalwood] 

Zhongguo Zhong Yao Za Zhi. 16(1), 40-3 Abstract. The GC-fingerprint spectra of essential oils in 

imported sandalwood are established by the new technique of GC-relative retention value 

fingerprint spectrum (GC-FPS). According to the GC-FPS of samples, their chromatographic 

peaks, overlap ratio of peaks and eight strong peaks are studied comparatively. 

Widiarti, A. (1991). “Site and heartwood formation of sandalwood.” Buletin Penelitian Hutan (534), 

1-14. 

Zhang D-Q., Ma Q-Z, Peng W-x & Liu Q-M. (2008) "Pyrolysis- GC/MS analysis of biomedical 

components of the pyrolyzate of Santalum album leaf treated by benzene/ethanol extraction." 

Bioinformatics and Biomedical Engineering 2nd Intl. Conference 16-18 May 2008, 1213-1218, 

Zhang Q., Tang Y., Wang R., Wang S., Fang M., Zhang Y. & Zheng X. (2009) "[Effect of 

Santalum album on tissue distribution of danshensu in rabbits by HPLC]" Zhongguo Zhong Yao 

Za Zhi 34(15), 1968-70. Abstract. OBJECTIVE: To investigate the influence of Shi herb 

(Santalum album, SA) to the tissue distribution of danshensu (DSS) which is the main 

hydrosoluble component of Jun herb (Salvia miltiorrhiza, SM) in rabbits by HPLC. METHOD: 

Rabbits were oral administrated decoction of SM and SM-SA, respectively. Perchloric acid (10%) 

was used to precipitate the tissue samples of rabbits heart, brain, liver, kidney, acetic ether was 

used to extracte supernatant, and the internal standard was p-hydroxybenzoic acid. The content 

of DSS of SM in tissues was detected. RESULT: The content of DSS reached the highest point 

close to 50 min in the mentioned tissues. Before and after co-administration, the sequences of 

average concentration of DSS in tissues were C(kidney) > C(heart) > C(brain) > C(liver) and 

C(kidney) > C(liver) > C(brain) > C(heart) respectively. Compared with SM administrated singly, 

the content of DSS in every tissues of co-administration was higher. CONCLUSION: In 

Danshenyin Formulae, SA can increase concentration of DSS in target tissues significantly, and 

therefore therapeutic effect of SM for cardiovascular disease is raised. 

79

Sandalwood Biblio - Cropwatch

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